Andrew Siefert

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.

Incorporating intraspecific variation in tests of trait-based community assembly

Environmental filtering and niche differentiation are processes proposed to drive community assembly, generating nonrandom patterns in community trait distributions. Despite the substantial intraspecific trait variation present in plant communities, most previous studies of trait-based community assembly have used species mean trait values and therefore not accounted for intraspecific variation. Using a null model approach, I tested for environmental filtering and niche differentiation acting on three key functional traits—vegetative height, specific leaf area (SLA), and leaf dry matter content (LDMC)—in old-field plant communities. I also examined how accounting for intraspecific variation at the among-plot and individual levels affected the detection of nonrandom assembly patterns. Tests using fixed species mean trait values provided evidence of environmental filtering acting on height and SLA and niche differentiation acting on SLA. Including plot-level intraspecific variation increased the strength of these patterns, indicating an important role of intraspecific variation in community assembly. Tests using individual trait data indicated strong environmental filtering acting on all traits, but provided no evidence of niche differentiation, although these signals may have been obscured by the effects of dispersal limitation and spatial aggregation of conspecific individuals. There was also strong evidence of nonrandom assembly of individuals within single species, with the strength of environmental filtering varying among species. This study demonstrates that, while analyses using fixed species mean trait values can provide insights into community assembly processes, accounting for intraspecific variation provides a more complete view of communities and the processes driving their assembly.

A global method for calculating plant CSR ecological strategies applied across biomes world‐wide

Summary nCompetitor, stress-tolerator, ruderal (CSR) theory is a prominent plant functional strategy scheme previously applied to local floras. Globally, the wide geographic and phylogenetic coverage of available values of leaf area (LA), leaf dry matter content (LDMC) and specific leaf area (SLA) (representing, respectively, interspecific variation in plant size and conservative vs. acquisitive resource economics) promises the general application of CSR strategies across biomes, including the tropical forests hosting a large proportion of Earths diversity. nWe used trait variation for 3068 tracheophytes (representing 198 families, six continents and 14 biomes) to create a globally calibrated CSR strategy calculator tool and investigate strategy–environment relationships across biomes world-wide. nDue to disparity in trait availability globally, co-inertia analysis was used to check correspondence between a ‘wide geographic coverage, few traits’ data set and a ‘restricted coverage, many traits’ subset of 371 species for which 14 whole-plant, flowering, seed and leaf traits (including leaf nitrogen content) were available. CSR strategy/environment relationships within biomes were investigated using fourth-corner and RLQ analyses to determine strategy/climate specializations. nStrong, significant concordance (RVxa0=xa00·597; Pxa0<xa00·0001) was evident between the 14 trait multivariate space and when only LA, LDMC and SLA were used. nBiomes such as tropical moist broadleaf forests exhibited strategy convergence (i.e. clustered around a CS/CSR median; C:S:Rxa0=xa043:42:15%), with CS-selection associated with warm, stable situations (lesser temperature seasonality), with greater annual precipitation and potential evapotranspiration. Other biomes were characterized by strategy divergence: for example, deserts varied between xeromorphic perennials such as Larrea divaricata, classified as S-selected (C:S:Rxa0=xa01:99:0%) and broadly R-selected annual herbs (e.g. Claytonia perfoliata; R/CR-selected; C:S:Rxa0=xa021:0:79%). Strategy convergence was evident for several growth habits (e.g. trees) but not others (forbs). nThe CSR strategies of vascular plants can now be compared quantitatively within and between biomes at the global scale. Through known linkages between underlying leaf traits and growth rates, herbivory and decomposition rates, this method and the strategy–environment relationships it elucidates will help to predict which kinds of species may assemble in response to changes in biogeochemical cycles, climate and land use.

Phenological variation of leaf functional traits within species

A basic assumption of the trait-based approach in plant ecology is that differences in functional trait values are greater between species than within species. We questioned this assumption by assessing (1) the relative extent of inter- and intraspecific leaf trait variation throughout a complete growing season (phenological variation) in a group of deciduous and evergreen woody species, and (2) whether species rankings based on leaf traits were maintained across the growing season. We analysed leaf mass per area (LMA) and leaf nutrient concentrations (C, N, P), including the C:N and N:P ratios. Intraspecific trait variation (ITV) due to phenology was significantly greater than interspecific variation for leaf N concentration on a mass basis (Nm; 68.90xa0%) and for the leaf C:N ratio (60.60xa0%), whereas interspecific variation was significantly higher than ITV for LMA (62.30xa0%) and for leaf C concentration on a mass (Cm) and area (Ca) basis (Cm 70.40xa0%; Ca 65.30xa0%). ITV was particularly low for LMA (<20xa0%). Species rankings were highly modified by phenology for a number of leaf traits (Pm, N:P ratio) but were relatively well conserved throughout the growing season for others (LMA, Nm). Patterns of ITV across the growing season differed significantly between deciduous and evergreen species for all traits except leaf P but did not vary between native and exotic species. Overall, our results show that intraspecific phenological variation in leaf traits may be similar to or greater than interspecific variation and that temporal patterns of ITV vary considerably among traits and species, especially for leaf nutrient concentrations, factors which can potentially affect quantitative interspecific relationships.

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 variation drives functional responses of old-field plant communities to nutrient enrichment

Environmental changes are expected to shift the distribution of functional trait values in plant communities through a combination of species turnover and intraspecific variation. The strength of these shifts may depend on the availability of individuals with trait values adapted to new environmental conditions, represented by the functional diversity (FD) of existing community residents or dispersal from the regional species pool. We conducted a 3-year nutrient- and seed-addition experiment in old-field plant communities to examine the contributions of species turnover and intraspecific variation to community trait shifts, focusing on four key plant functional traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We further examined the influence of initial FD and seed availability on the strength of these shifts. Community mean height, leaf area, and SLA increased in response to fertilization, and these shifts were driven almost entirely by intraspecific variation. The strength of intraspecific shifts in height and leaf area was positively related to initial intraspecific FD in these traits. Intraspecific trait responses to fertilization varied among species, with species of short stature displaying stronger shifts in SLA and LDMC but weaker shifts in leaf area. Trait shifts due to species turnover were generally weak and opposed intraspecific responses. Seed addition altered community taxonomic composition but had little effect on community trait shifts. These results highlight the importance of intraspecific variation for short-term community functional responses and demonstrate that the strength of these responses may be mediated by community FD.

Habitat filtering determines the functional niche occupancy of plant communities worldwide

How the patterns of niche occupancy vary from species-poor to species-rich communities is a fundamental question in ecology that has a central bearing on the processes that drive patterns of biodiversity. As species richness increases, habitat filtering should constrain the expansion of total niche volume, while limiting similarity should restrict the degree of niche overlap between species. Here, by explicitly incorporating intraspecific trait variability, we investigate the relationship between functional niche occupancy and species richness at the global scale. 2.We assembled 21 datasets worldwide, spanning tropical to temperate biomes and consisting of 313 plant communities representing different growth forms. We quantified three key niche occupancy components (the total functional volume, the functional overlap between species and the average functional volume per species) for each community, related each component to species richness, and compared each component to the null expectations. 3.As species richness increased, communities were more functionally diverse (an increase in total functional volume), and species overlapped more within the community (an increase in functional overlap) but did not more finely divide the functional space (no decline in average functional volume). Null model analyses provided evidence for habitat filtering (smaller total functional volume than expectation), but not for limiting similarity (larger functional overlap and larger average functional volume than expectation) as a process driving the pattern of functional niche occupancy. 4.Synthesis. Habitat filtering is a widespread process driving the pattern of functional niche occupancy across plant communities and coexisting species tend to be more functionally similar rather than more functionally specialized. Our results indicate that including intraspecific trait variability will contribute to a better understanding of the processes driving patterns of functional niche occupancy

Soil microbial communities alter conspecific and congeneric competition consistent with patterns of field coexistence in three Trifolium congeners

Coexistence and diversity in plant communities depend upon outcomes of plant competition. Competition and coexistence can be mediated by abiotic soil nutrient differences as well as by soil microbial communities. The latter effects occur through various mechanisms including negative plant–soil feedbacks, when plants foster the build‐up of specialized pathogenic microbes, which ultimately reduce conspecific, but not heterospecific, densities. Microbial mutualists can have generalized associations with host plants, and by associating with multiple species might affect coexistence by conferring different levels of benefit to hosts. We examined the effects of abiotic differences and soil microbial communities, including mutualistic nitrogen‐fixing rhizobial bacteria, on coexistence processes and asked whether these interactions inform patterns of co‐occurrence in natural communities. We measured plant–soil feedbacks in the greenhouse for three native Trifolium species that either highly co‐occurred or were spatially repulsed at our field site. Using size‐fractioned soil microbial inocula prepared from field‐collected soils, we explored the effect of soil microbes on nodulation and the outcome of competition (relative interaction intensity). We also examined the effects of soil origin (home vs. away soil) on the outcome of competition between species. Soil microbes had strong positive effects on plant growth and nodulation. Microbes in general reduced the strength of plant competition relative to competition in sterilized soil and altered the relative strength of interactions with conspecific vs. congeneric neighbours, which often occurred in ways predicted to enhance coexistence. In one pair of highly co‐occurring Trifolium, competition was strong in sterilized soils, but these species facilitated one other in the presence of the full microbial community. The net effect of these microbes in general reduced competition while also increasing performance over that in sterilized soil, suggesting a role for soil microbial mutualists in host coexistence. Synthesis. Our results demonstrate that interactions between plants and diverse soil communities can alter plant–plant interactions and plant–soil feedbacks. They may increase niche differences and equalize fitness differences in ways consistent with observed co‐occurrence in the field.

Greenhouse experiment neighborhood data

Individual biomass, trait, and neighborhood composition for Trifolium greenhouse experiment

Intraspecific trait variation and the leaf economics spectrum across resource gradients and levels of organization.

Understanding patterns of functional trait variation across environmental gradients offers an opportunity to increase inference in the mechanistic causes of plant community assembly. The leaf economics spectrum (LES) predicts global tradeoffs in leaf traits and trait-environment relationships, but few studies have examined whether these predictions hold across different levels of organization, particularly within species. Here, we asked (1) whether the main assumptions of the LES (expected trait relationships and shifts in trait values across resource gradients) hold at the intraspecific level, and (2) how within-species trait correlations scale up to interspecific or among-community levels. We worked with leaf traits of saplings of woody species growing across light and soil N and P availability gradients in temperate rainforests of southern Chile. We found that ITV accounted for a large proportion of community-level variation in leaf traits (e.g., LMA and leaf P) and played an important role in driving community-level shifts in leaf traits across environmental gradients. Additionally, intraspecific leaf trait relationships were generally consistent with interspecific and community-level trait relationships and with LES predictions-e.g., a strong negative intraspecific LMA-leaf N correlation-although, most trait relationships varied significantly among species, suggesting idiosyncrasies in the LES at the intraspecific level.