Larger patches of diverse floral resources increase insect pollinator density,diversity, and their pollination of native wildflowers

Native wildflower plantings can be used to provide nutritional resources to support pollinating insects, yet the effects of planting size and bloom richness on the density, diversity, and function of these insects are not well understood. We established stands of twelve native flowering perennial plant species in replicated plots ranging in size from 1 to 100 m². These plots were sampled for insect pollinators, bloom richness, and seed production by three wildflower species. Honeybees, wild bees, and hoverflies all responded positively to increasing flower richness, whereas particular insect pollinator groups responded differently to the size of the flowering plant area. The density of honeybees and hoverflies was not affected by increasing flowering patch size, whereas in general, wild bees were observed at higher density and diversity in the 30 and 100 m² patches. Increasing wildflower patch size, and thus wild bee density, resulted in greater seed set in the sampled wildflowers. These results indicate that wild bees are sensitive to the area and richness of floral resources in patches, even at relatively small scales. Therefore, larger wildflower plantings with more diverse flower species mixes are more suitable for the conservation of wild pollinators and reproduction of sown species.     

Intraspecific variation in species interactions promotes the feasibility of mutualistic assemblages

Patterns of resource use observed at the species level emerge from the way individuals exploit the range of available resources. Hence, accounting for interindividual differences in resource use, such as pollinator use by plants, is essential to advance our understanding of community assembly and persistence. By using finely resolved data on plant–pollinator interactions, we evaluated how interindividual plant variation in pollinator use scales up to affect community structure and dynamics. All co-occurring plant species comprised specialists interacting with proper subsets of pollinators that visited generalists, and differences in interaction patterns were driven by among-individual trait variation. Furthermore, the nested structure and feasibility of plant–pollinator communities were maximised at higher levels of interindividual plant variation in traits and pollinator use. Our study sheds light on how pervasive properties of community structure arise from individual-level processes and contributes to elucidate the importance of preserving intraspecific variation in traits and resource use within populations.     

Functional flower traits and their diversity drive pollinator visitation

Recent studies have shown that the diversity of flowering plants can enhance pollinator richness and visitation frequency and thereby increase the resilience of pollination. It is assumed that flower traits explain these effects, but it is still unclear which flower traits are responsible, and knowing that, if pollinator richness and visitation frequency are more driven by mass‐ratio effects (mean trait values) or by trait diversity. Here, we analyse a three‐year data set of pollinator observations collected in a European grassland plant diversity experiment (The Jena experiment). The data entail comprehensive flower trait measurements, including reward traits (nectar and pollen amount), morphological traits (height, symmetry, area, colour spectra) and chemical traits (nectar‐amino acid and nectar‐sugar concentration). We test if pollinator species richness and visitation frequency of flower communities depend on overall functional diversity combining all flower traits within a community, single trait diversities (within trait variation) and community‐weighted means of the single traits, using Bayesian inference. Overall functional diversity did not affect pollinator species richness, but reduced visitation frequency. When looking at individual flower traits separately, we found that single trait diversity of flower reflectance and flower morphology were important predictors of pollinator visitation frequency. Moreover, independent of total flower abundance, community‐weighted means of flower height, area, reflectance, nectar‐sugar concentration and nectar‐amino acid concentration strongly affected both pollinator species richness and visitation frequency. Our results, challenge the idea that functional diversity always positively affects ecosystem functions. Nonetheless, we demonstrate that both single trait diversity and mass‐ratio effects of flower traits play an important role for diverse and frequent flower visits, which underlines the functionality of flower traits for pollination services.     

Volatility of network indices due to undersampling of intraspecific variation in plant–insect interactions

Appropriate sampling effort of interaction networks is necessary to extract robust indices describing the structure of species interactions. Here we show that time-invariant variation in the composition and diversity of interaction partners of plant individuals of the same species explains volatility in aggregate network statistics due to undersampling. Within a multi-species pollinator–plant interaction matrix, we replaced the interactions observed on multiple individuals of a single plant species (Sinapis arvensis, pooled interactions) with the plant–insect interactions observed on a single plant individual. In the resampling approach, we considered the interactions of 1 to 84 S. arvensis individuals in different combinations. For each resampled network, several commonly applied aggregated statistics were calculated to test how intraspecific variation affects the properties of a multi-species network. Our results showed that aggregate statistics are sensitive towards qualitative and quantitative intraspecific variation of flower–visitor interactions within a multi-species network, which may affect the ecological interpretation about the properties of a community. These findings challenge the robustness of commonly applied network indices, confirm the urge for a sufficient and representative sampling of interactions, and emphasize the significance of intraspecific variation in the context of communities and networks.     

Wild bees and natural enemies prefer similar flower species and respond to similar plant traits

Designing wildflower habitats to support beneficial insects providing pollination and pest control services is important for supporting sustainable crop production. It is often desirable to support both groups of insects, making the selection of resource plants for insect conservation programs more challenging. Moreover, the process of selecting resource plants is complicated by the array of possible options in each region, and the need to provide resources over the entire growing season. Identifying traits shared by resource plants that are attractive to both bees and natural enemies can reduce the need to evaluate new plants in each region, by providing a guide for the types of plants expected to be rewarding to these insects. Using insect visitation data collected from replicated common garden plantings of native wildflower and shrub species from the Great Lakes region of the United States, we found a high degree of correlation between the abundance of bees and natural enemies visiting native plant species. These results were used to identify a set of 15 plant species that can provide resources for these insects throughout the summer. Across all tested species, pollen quantity per flower and the week of bloom were positively correlated with some, but not all, taxonomic groupings of beneficial insects. In contrast, floral area was consistently positively associated with visitation of both natural enemies and wild bees. This trait is easy to document and can allow for efficient local testing of potential resource plants, providing a faster path to implementing insect conservation in working landscapes.     

Intraspecific variation in host plant traits mediates taxonomic and functional composition of local insect herbivore communities

1. Host plant phenotypic traits affect the structure of the associated consumer community and mediate species interactions. Intraspecific variation in host traits is well documented, although a functional understanding of variable traits that drive herbivore community response is lacking. We address this gap by modelling the trait-environment relationship using insect traits and host plant traits in a multilevel model. 2. We compare herbivore assemblages from the canopy of the phenotypically variable tree Metrosideros polymorpha on Hawai‘i Island. Multiple distinct varieties of M. polymorpha frequently co-occur, with variation in morphological traits. Using this system, we identify host and insect traits that underlie patterns of herbivore abundance and quantify the strength of host-insect trait interactions. 3. This work examines plant-insect interactions at a community scale, across 36 herbivore species in three orders. We find that co-occurring trees of varying phenotype support distinct communities. Leaf traits, including specific leaf area, trichome presence, and leaf nutrients, explain 46% of variation in insect communities. We find that feeding guild and nymphal life history are correlated with host plant traits, and we show that model predictions are improved by including the host and insect trait interaction. 4. This study demonstrates how insect herbivores traits influence community response to morphologically variable hosts. Environmental heterogeneity indirectly affected herbivore community structure via intraspecific variation in host plants, providing an important source of variation for maintaining diversity in the broader community.     

Phenotypic divergence and inter-specific trait correlation in a plant-pollinator/seed predator mutualism

Plant-pollinator interactions have been suggested as key drivers of morphological divergence and speciation of the involved taxa. These interactions can also promote sexual dimorphism in both the plant and pollinator, particularly if the pollinator is also a seed-eater and/or exerts different selection pressures on male and female plants. Here we tested the hypotheses that plant-pollinator interactions can be reflected in trait variation and sexual dimorphism in both organisms within and across populations. Across nine European populations, we examined intraspecific variation and sexual dimorphism in phenotypic traits potentially involved in the plant–insect interaction of the dioecious white campion Silene latifolia (Caryophyllaceae) and its specialist pollinator Hadena bicruris (Noctuidae). This interaction is expected to entail sex-specific selective pressures, as female moths lay eggs on female plants and the larvae predate on the seeds during their development. We compared divergence in phenotypic traits among populations and between sexes within populations, examined correlations between plant and pollinator traits, and between phenotypic distances and genetic distances among co-occurring populations for both plants and insects. We found key differences in phenotypic traits across populations of both the plant and moth, though only in the moth were these differences correlated with geographic distances. We also found evidence for sexual dimorphism in the plant but not in the pollinator. Evolution of floral sexual dimorphism in S. latifolia most likely results from the joint contribution of different selective forces, including biotic interactions with H. bicruris moths.     

Complex floral traits shape pollinator attraction to ornamental plants

Background and AimsOrnamental flowering plant species are often used in managed greenspaces to attract and support pollinator populations. In natural systems, selection by pollinators is hypothesized to result in convergent multimodal floral phenotypes that are more attractive to specific pollinator taxa. In contrast, ornamental cultivars are bred via artificial selection by humans, and exhibit diverse and distinct phenotypes. Despite their prevalence in managed habitats, the influence of cultivar phenotypic variation on plant attractiveness to pollinator taxa is not well resolved.MethodsWe used a combination of field and behavioural assays to evaluate how variation in floral visual, chemical and nutritional traits impacted overall attractiveness and visitation by pollinator taxonomic groups and bee species to 25 cultivars of five herbaceous perennial ornamental plant genera.Key resultsDespite significant phenotypic variation, cultivars tended to attract a broad range of pollinator species. Nonetheless, at the level of insect order (bee, fly, butterfly, beetle), attraction was generally modulated by traits consistent with the pollination syndrome hypothesis. At the level of bee species, the relative influence of traits on visitation varied across plant genera, with some floral phenotypes leading to a broadening of the visitor community, and others leading to exclusion of visitation by certain bee species.ConclusionsOur results demonstrate how pollinator choice is mediated by complex multimodal floral signals. Importantly, the traits that had the greatest and most consistent effect on regulating pollinator attraction were those that are commonly selected for in cultivar development. Though variation among cultivars in floral traits may limit the pollinator community by excluding certain species, it may also encourage interactions with generalist taxa to support pollinator diversity in managed landscapes.     

Leaf traits and performance vary with plant age and water availability in Artemisia californica

Background and AimsLeaf functional traits are strongly tied to growth strategies and ecological processes across species, but few efforts have linked intraspecific trait variation to performance across ontogenetic and environmental gradients. Plants are believed to shift towards more resource-conservative traits in stressful environments and as they age. However, uncertainty as to how intraspecific trait variation aligns with plant age and performance in the context of environmental variation may limit our ability to use traits to infer ecological processes at larger scales.MethodsWe measured leaf physiological and morphological traits, canopy volume and flowering effort for Artemisia californica (California sagebrush), a dominant shrub species in the coastal sage scrub community, under conditions of 50, 100 and 150 % ambient precipitation for 3 years.Key ResultsPlant age was a stronger driver of variation in traits and performance than water availability. Older plants demonstrated trait values consistent with a more conservative resource-use strategy, and trait values were less sensitive to drought. Several trait correlations were consistent across years and treatments; for example, plants with high photosynthetic rates tended to have high stomatal conductance, leaf nitrogen concentration and light-use efficiency. However, the trade-off between leaf construction and leaf nitrogen evident in older plants was absent for first-year plants. While few traits correlated with plant growth and flowering effort, we observed a positive correlation between leaf mass per area and performance in some groups of older plants.ConclusionsOverall, our results suggest that trait sensitivity to the environment is most visible during earlier stages of development, after which intraspecific trait variation and relationships may stabilize. While plant age plays a major role in intraspecific trait variation and sensitivity (and thus trait-based inferences), the direct influence of environment on growth and fecundity is just as critical to predicting plant performance in a changing environment.     

Functional trait differences and the outcome of community assembly: an experimental test with vernal pool annual plants

Functional trait‐based approaches have seen rapid development in community ecology and biogeography in recent years, as they promise to offer a better mechanistic and predictive understanding of community structure. However, several key challenges remain. First, while many studies have explored connections between functional traits and abiotic gradients, far fewer have directly tested the common assumption that functional trait differences influence interspecific interactions. Second, empirical studies often ignore intraspecific trait variation within communities, even though intraspecific variation has been known to have substantial impacts on community dynamics. Here we present an experiment designed to assess the role of functional trait differences in predicting the outcome of interspecific species interactions among a suite of California vernal pool annual plants. Eight species were grown in pairwise combinations in two levels of inundation in a greenhouse and functional traits were measured on all individuals. Nested models predicting focal plant performance were fit to the data. For seven of the eight species in the experiment, the best model included a functional trait difference term that was consistent with a competitive hierarchy, indicating that focal species tended to do better when they had larger leaf size, lower specific leaf area, and greater investment in lateral canopy spread than their neighbors. Models that included individually measured trait values generally performed better than models using species trait averages. We tested if the same trait measurements predicted tolerance of inundation (a feature of vernal pool habitats), and species depth distributions from extensive field surveys, though we did not find strong relationships. Our results suggest that functional traits can be used to make inferences about the outcome of interspecific interactions, and that greater predictive power can come from considering intraspecific variation in functional traits, particularly in low diversity communities.     

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。     

Interspecific plant functional variation prevails over intraspecific variation in driving spider beta diversity

1. Non-trophic interactions between plants and animals can affect community structure and species trait composition. However, it is unclear how changes in intra- and interspecific morphological traits of plant species affect non-trophic interactions at a metacommunity scale. Additionally, whether plant evolutionary history determines taxonomic and functional diversity of plant-dwelling predators is an open question. 2. To address these gaps, this study used a published dataset with spiders dwelling exclusively on bromeliads to investigate if: (i) intra- and interspecific variability in host plant morphological traits affects spider taxonomic and functional diversity; and (ii) bromeliad trait evolution determines present-day patterns of spider trait diversity. 3. Spider and bromeliad traits were measured, and a new statistical framework was used to quantify the response of spider beta diversity to intra- and interspecific variation in bromeliad traits. In addition, bromeliad traits were decomposed across its phylogenetic tree to check whether the current variation in morphological traits of bromeliads is a result of either ancestral or recent diversification. 4. Bromeliad intraspecific variation did not affect spiders, but leaf length variation between bromeliad species had a positive effect on spider functional beta diversity. Interestingly, the most ancestral split between two subfamilies explained most of the variation in bromeliad species, which suggests that spider functional diversity could represent an outcome of bromeliad evolutionary history. 5. Overall, the results of this study suggest that interactions between plants and organisms that do not feed directly on their tissues could be shaped by plant evolutionary history, which in turn suggests that non-trophic interactions can be maintained over time.     

cati: an R package using functional traits to detect and quantify multi‐level community assembly processes

Community ecologists are active in describing species by their functional traits, quantifying the functional structure of plant and animal assemblages and inferring community assembly processes with null‐model analyses of trait distribution and functional diversity indices. Intraspecific variation in traits and effects of spatial scale are potentially important in these analyses. Here, we introduce the R package cati (Community Assembly by Traits: Individuals and beyond) available on CRAN, for the analysis of community assembly with functional traits. cati builds on a recent approach to community assembly that explicitly incorporates individual differences in community assembly analyses and decomposes phenotypic variations across scales and organizational levels, based on three phenotypic variance ratios, termed the T‐statistics. More generally, the cati package 1) calculates a variety of single‐trait and multi‐trait indices from interspecific and intraspecific trait measures; 2) it partitions functional trait variation among spatial and taxonomic levels; 3) it implements a palette of flexible null models for detecting non‐random patterns of functional traits. These patterns can be used to draw inferences about hypotheses of community assembly such as environmental filtering and species interactions. The basic input for cati is a data frame in which columns are traits, rows are species or individuals, and entries are the measured trait values. The cati package can also incorporate a square distance matrix into analyses, which could include phylogenetic or genetic distances among individuals or species. Users select from a variety of functional trait metrics and analyze these relative to a null model that specifies trait distributions in a regional source pool.     

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.     

Plant reproductive strategies vary under low and high pollinator densities

Long‐term variation in the population density of honey bees Apis mellifera across landscapes has been shown to correlate with variation in the floral traits of plant populations in these landscapes, suggesting that variations in pollinator population density and foraging rates can drive floral trait evolution of their host plants. However, it remained to be determined whether this variation in plant traits is associated with adaptive variation in plant reproductive strategies under conditions of high and low pollinator densities. Here we conducted a reciprocal transplant experiment to examine how this variation in floral traits, under conditions of either high and low pollinator density, impacted seed production in the Tibetan lotus Saussurea nigrescens. In 2014 and 2015, we recorded the floral traits, pollinator visitation rates, and seed production of S. nigrescens populations grown in both home sites and foreign sites, where sites varied in honey bee population density. Our results demonstrated that the floral traits reflected those of their original population, regardless of their current location. However, seed production varied with both population origin and transplant site. Seed number was positively correlated with flower abundance in the pollinator‐rich sites, but with nectar production in the pollinator‐poor sites. Pollinator visitation rate was also positively correlated with flower number at pollinator‐rich sites, and with nectar volume at pollinator‐poor sites. Overall, the local genotype had higher seed production than nonlocal genotypes in home sites. However, when pollen is hand‐supplemented, plants from pollinator‐rich populations had higher seed production than plants from pollinator‐poor populations, regardless of whether they were transplanted to pollinator‐rich or ‐poor sites. These results suggest that the plant genotypic differences primarily drive variation in pollinator attraction, and this ultimately drives variation in seed: ovule ratio. Thus, our results suggest that flowering plant species use different reproductive strategies to respond to high or low pollinator densities.     

Augmenting flower trait diversity in wildflower strips to optimise the conservation of arthropod functional groups for multiple agroecosystem services

Sown wildflower strips are increasingly being established in Europe for enhancing arthropod conservation and the provision of ecosystem services, including biotic pollination and natural pest control. Here we use floral traits to identify different plant functional effect groups. Floral resources were provided in four experimental levels characterised by a cumulatively increasing flower trait diversity and vegetation stand complexity. The first level consisted of a bare control strip, whilst in each subsequent level three wildflower species with different functional traits were added (Level 0: control; Level 1: three Apiaceae species; Level 2: three Apiaceae and three Fabaceae species; Level 3: three Apiaceae, three Fabaceae species, and Centaurea jacea (Asteraceae), Fagopyrum esculentum (Polygonaceae), Sinapis alba (Brassicaceae)). Plots with sown wildflower strip mixtures were located adjacent to experimental plots of organically-managed tomato crop, which is attacked by multiple pests and partially relies on bees for fruit production, and hence dependent on the provision of pollination and pest control services. Results obtained here show that the inclusion of functionally diverse wildflower species was associated with an augmented availability of floral resources across time, and this increased the abundance of bees and anthocorids throughout the crop season. Several natural enemy groups, such as parasitoids, coccinelids and ground-dwelling predators, were not significantly enhanced by the inclusion of additional flower traits within the strips but the presence of flower resources was important to enhance their conservation in an arable cropping system.     

Spatial configuration and landscape context of wildflower areas determine their benefits to pollinator α- and β-diversity

Wildflower areas have become a staple tool within agro-environmental schemes (AES) to counteract pollinator declines. While their role in providing food resources to resident flower-visiting insects is unambiguous, the conservation effectiveness in a landscape context is less clear. Particularly, how multiple vs. single wildflower area utilization differs between simple and complex landscapes is understudied. We examined colonisation and community dynamics of wild bees and hoverflies in 33 newly established wildflower areas across a gradient of landscape complexity (amount of semi-natural habitat) and connectivity (presence of additional wildflower areas) for seven weeks during three consecutive years (one year during and two after establishment). We recorded more than 25% of the wild bee and hoverfly species of Hesse in an area of approx. 10 ha, substantiating the general benefit of wildflower areas to pollinators. While alpha-diversity increased with landscape complexity in isolated wildflower areas, the opposite pattern was observed for connected areas. The low alpha-diversity in complex landscapes indicated a dilution effect between connected sites. The inverse relationship between alpha and beta diversity among wildflower areas within landscapes suggests interspecific trade-offs between local resource conditions and landscape context. Accordingly, the establishment of multiple wildflower areas within AES is advisable to increase connectivity of suitable habitats in simple landscapes. Moreover, adjusting local conditions (plant diversity) to landscape context likely optimizes conservation effectiveness in modern agroecosystems.     

Can novel seed mixes provide a more diverse,abundant, earlier,and longer-lasting floral resource for bees than current mixes?

Several agri-environment schemes aim to improve pollinator diversity and abundance, including the sowing of wildflower areas. These seed mixes are often either low in floral diversity and target few pollinator species (mainly social bees), or high in floral diversity but with limited evidence of good establishment of the component species. In order to support a greater diversity of wild bees in farmland, we need more diverse seed mixes, containing species shown to support a wide diversity of insect pollinators, with good establishment and long flowering periods. Here we trialled two typical seed mixes, a low-diversity Fabaceae-heavy mix (FAB) and a more diverse wildflower mix (WF), against two novel wildflower mixes, one based on literature sources (LT), and one based on first-hand surveys of pollinator attraction to flowers growing on a wildflower farm (WB). Both new mixes were focussed on plants attractive to wild bee species. Replicated field plots were set up on two farms and monitored over three years. Our novel wildflower mixes had higher floral diversity and abundance than the FAB mix, and began flowering earlier, reaching their floral peak before the FAB mix, potentially providing forage for a broader range of pollinators or those with earlier flight seasons. The high floral abundance in LT and WB was driven by annuals in the first year, and then multiple perennials in the second and third year. We identified five perennials from four families (Daucus carota, Leucanthemum vulgare, Geranium pyrenaicum, Lotus corniculatus and Trifolium hybridum) that established well on both farms, are known to be attractive to a diversity of bee species, and thus could be considered as providing a more taxonomically diverse base for creating future mixes. However, the mixes provided few floral resources in April (needed by early-flying wild bees), and more research is required in this area.     

Size‐dependent and environment‐mediated shifts in leaf traits of a deciduous tree species in a subtropical forest

AimsUnderstanding the joint effects of plant development and environment on shifts of intraspecific leaf traits will advance the understandings of the causes of intraspecific trait variation. We address this question by focusing on a widespread species Clausena dunniana in a subtropical broad‐leaved forest.MethodsWe sampled 262 individuals of C. dunniana at two major topographic habitat types, the slope and hilltop, within the karst forests in Maolan Nature Reserve in southwestern China. We measured individual plant level leaf traits (i.e., specific leaf area (SLA), leaf area, leaf dry‐matter content (LDMC), and leaf thickness) that are associated with plant resource‐use strategies. We adopted a linear mixed‐effects model in which the plant size (i.e., the first principal component of plant basal diameter and plant height) and environmental factors (i.e., topographic habitat, canopy height, and rock‐bareness) were used as independent variables, to estimate their influences on the shifts of leaf traits.Key ResultsWe found that (1) plant size and the environmental factors independently drove the intraspecific leaf trait shifts of C. dunniana, of which plant size explained less variances than environmental factors. (2) With increasing plant size, C. dunniana individuals had increasingly smaller SLA but larger sized leaves. (3) The most influential environmental factor was topographic habitat; it drove the shifts of all the four traits examined. Clausena dunniana individuals on hilltops had leaf traits representing more conservative resource‐use strategies (e.g., smaller SLA, higher LDMC) than individuals on slopes. On top of that, local‐scale environmental factors further modified leaf trait shifts.ConclusionsPlant size and environment independently shaped the variations in intraspecific leaf traits of C. dunniana in the subtropical karst forest of Maolan. Compared with plant size, the environment played a more critical role in shaping intraspecific leaf trait variations, and potentially also the underlying individual‐level plant resource‐use strategies.     

Epigenetic variation predicts regional and local intraspecific functional diversity in a perennial herb

The ecological significance of epigenetic variation has been generally inferred from studies on model plants under artificial conditions, but the importance of epigenetic differences between individuals as a source of intraspecific diversity in natural plant populations remains essentially unknown. This study investigates the relationship between epigenetic variation and functional plant diversity by conducting epigenetic (methylation‐sensitive amplified fragment length polymorphisms, MSAP) and genetic (amplified fragment length polymorphisms, AFLP) marker–trait association analyses for 20 whole‐plant, leaf and regenerative functional traits in a large sample of wild‐growing plants of the perennial herb Helleborus foetidus from ten sampling sites in south‐eastern Spain. Plants differed widely in functional characteristics, and exhibited greater epigenetic than genetic diversity, as shown by per cent polymorphism of MSAP fragments (92%) or markers (69%) greatly exceeding that for AFLP ones (41%). After controlling for genetic structuring and possible cryptic relatedness, every functional trait considered exhibited a significant association with at least one AFLP or MSAP marker. A total of 27 MSAP (13.0% of total) and 12 AFLP (4.4%) markers were involved in significant associations, which explained on average 8.2% and 8.0% of trait variance, respectively. Individual MSAP markers were more likely to be associated with functional traits than AFLP markers. Between‐site differences in multivariate functional diversity were directly related to variation in multilocus epigenetic diversity after multilocus genetic diversity was statistically accounted for. Results suggest that epigenetic variation can be an important source of intraspecific functional diversity in H. foetidus, possibly endowing this species with the capacity to exploit a broad range of ecological conditions despite its modest genetic diversity.