Disentangling niche and neutral influences on community assembly: assessing the performance of community phylogenetic structure tests

Patterns of phylogenetic relatedness within communities have been widely used to infer the importance of different ecological and evolutionary processes during community assembly, but little is known about the relative ability of community phylogenetics methods and null models to detect the signature of processes such as dispersal, competition and filtering under different models of trait evolution. Using a metacommunity simulation incorporating quantitative models of trait evolution and community assembly, I assessed the performance of different tests that have been used to measure community phylogenetic structure. All tests were sensitive to the relative phylogenetic signal in species metacommunity abundances and traits; methods that were most sensitive to the effects of niche-based processes on community structure were also more likely to find non-random patterns of community phylogenetic structure under dispersal assembly. When used with a null model that maintained species occurrence frequency in random communities, several metrics could detect niche-based assembly when there was strong phylogenetic signal in species traits, when multiple traits were involved in community assembly, and in the presence of environmental heterogeneity. Interpretations of the causes of community phylogenetic structure should be modified to account for the influence of dispersal.     

Phylogenetic community ecology and the role of social dominance in sponge-dwelling shrimp

When functional traits are evolutionarily conserved, phylogenetic relatedness can serve as a proxy for ecological similarity to examine whether functional differences among species mediate community assembly. Using phylogenetic- and trait-based analyses, we demonstrate that sponge-dwelling shrimp (Synalpheus) assemblages are structured by size-based habitat filtering, interacting with competitive exclusion mediated by social system. Most shrimp communities were more closely related and/or more similar in size than randomized communities, consistent with habitat filtering facilitated by phylogenetically conserved body size. Those sponges with greater space heterogeneity hosted shrimp communities with greater size diversity, corroborating the importance of size in niche use. However, communities containing eusocial shrimp - which cooperatively defend territories - were less phylogenetically related and less similar in size, suggesting that eusociality enhances competitive ability and drives competitive exclusion. Our analyses demonstrate that community assembly in this diverse system occurs via traits mediating niche use and differential competitive ability.     

Facilitation can increase the phylogenetic diversity of plant communities

With the advent of molecular phylogenies the assessment of community assembly processes has become a central topic in community ecology. These processes have focused almost exclusively on habitat filtering and competitive exclusion. Recent evidence, however, indicates that facilitation has been important in preserving biodiversity over evolutionary time, with recent lineages conserving the regeneration niches of older, distant lineages. Here we test whether, if facilitation among distant-related species has preserved the regeneration niche of plant lineages, this has increased the phylogenetic diversity of communities. By analyzing a large worldwide database of species, we showed that the regeneration niches were strongly conserved across evolutionary history. Likewise, a phylogenetic supertree of all species of three communities driven by facilitation showed that nurse species facilitated distantly related species and increased phylogenetic diversity.     

Community disassembly under global change: Evidence in favor of the stress‐dominance hypothesis

Ecological theory suggests that communities are not random combinations of species but rather the results of community assembly processes filtering and sorting species that are able to coexist together. To date, such processes (i.e., assembly rules) have been inferred from observed spatial patterns of biodiversity combined with null model approaches, but relatively few attempts have been made to assess how these processes may be changing through time. Specifically, in the context of the ongoing biodiversity crisis and global change, understanding how processes shaping communities may be changing and identifying the potential drivers underlying these changes become increasingly critical. Here, we used time series of 460 French freshwater fish communities and assessed both functional and phylogenetic diversity patterns to determine the relative importance of two key assembly rules (i.e., habitat filtering and limiting similarity) in shaping these communities over the last two decades. We aimed to (a) describe the temporal changes in both functional and phylogenetic diversity patterns, (b) determine to what extent temporal changes in processes inferred through the use of standardized diversity indices were congruent, and (c) test the relationships between the dynamics of assembly rules and both climatic and biotic drivers. Our results revealed that habitat filtering, although already largely predominant over limiting similarity, became more widespread over time. We also highlighted that phylogenetic and trait‐based approaches offered complementary information about temporal changes in assembly rules. Finally, we found that increased environmental harshness over the study period (especially higher seasonality of temperature) led to an increase in habitat filtering and that biological invasions increased functional redundancy within communities. Overall, these findings underlie the need to develop temporal perspectives in community assembly studies, as understanding ongoing temporal changes could provide a better vision about the way communities could respond to future global changes.     

Fire modifies the phylogenetic structure of soil bacterial co‐occurrence networks

Fire alters ecosystems by changing the composition and community structure of soil microbes. The phylogenetic structure of a community provides clues about its main assembling mechanisms. While environmental filtering tends to reduce the community phylogenetic diversity by selecting for functionally (and hence phylogenetically) similar species, processes like competitive exclusion by limiting similarity tend to increase it by preventing the coexistence of functionally (and phylogenetically) similar species. We used co‐occurrence networks to detect co‐presence (bacteria that co‐occur) or exclusion (bacteria that do not co‐occur) links indicative of the ecological interactions structuring the community. We propose that inspecting the phylogenetic structure of co‐presence or exclusion links allows to detect the main processes simultaneously assembling the community. We monitored a soil bacterial community after an experimental fire and found that fire altered its composition, richness and phylogenetic diversity. Both co‐presence and exclusion links were more phylogenetically related than expected by chance. We interpret such a phylogenetic clustering in co‐presence links as a result of environmental filtering, while that in exclusion links reflects competitive exclusion by limiting similarity. This suggests that environmental filtering and limiting similarity operate simultaneously to assemble soil bacterial communities, widening the traditional view that only environmental filtering structures bacterial communities.     

The tangled link between β‐ and γ‐diversity: a Narcissus effect weakens statistical inferences in null model analyses of diversity patterns

Understanding the structure of and spatial variability in the species composition of ecological communities is at the heart of biogeography. In particular, there has been recent controversy about possible latitudinal trends in compositional heterogeneity across localities (β‐diversity). A gradient in the size of the regional species pool alone can be expected to impose a parallel gradient on β‐diversity, but whether β‐diversity also varies independently of the size of the species pool remains unclear. A recently suggested methodological approach to correct latitudinal β‐diversity gradients for the species pool effect is based on randomization null models that remove the effects of gradients in α‐ and γ‐diversity on β‐diversity. However, the randomization process imposes constraints on the variability of α‐diversity, which in turn force γ‐ and β‐diversity to become interdependent, such that any change in one is mirrored in the other. We argue that simple null model approaches are inadequate to discern whether correlations between α‐, β‐ and γ‐diversity reflect processes of ecological interest or merely differences in the size of the species pool among localities. We demonstrate that this kind of Narcissus effect may also apply to other metrics of spatial or phylogenetic species distribution. We highlight that Narcissus effects may lead to artificially high rejection rates for the focal pattern (Type II errors) and caution that these errors have not received sufficient attention in the ecological literature.     

Species matter: the role of competition in the assembly of congeneric bacteria

Interspecific competition is an important driver of community assembly in plants and animals, but phylogenetic evidence for interspecific competition in bacterial communities has been elusive. This could indicate that other processes such as habitat filtering or neutral processes are more important in bacterial community assembly. Alternatively, this could be a consequence of the lack of a consistent and meaningful species definition in bacteria. We hypothesize that competition in bacterial community assembly has gone undetected at least partly because overly broad measures of bacterial diversity units were used in previous studies. First, we tested our hypothesis in a simulation where we showed that how species are defined can dramatically affect whether phylogenetic overdispersion (a signal consistent with competitive exclusion) will be detected. Second, we demonstrated that using finer-scale Operational Taxonomic Units (OTUs) (with more stringent 16S rRNA sequence identity cutoffs or based on fast-evolving protein coding genes) in natural populations revealed previously undetected overdispersion. Finally, we argue that bacterial ecotypes, diversity units incorporating ecological and evolutionary theory, are superior to OTUs for the purpose of studying community assembly.     

Identifying models of trait‐mediated community assembly using random forests and approximate Bayesian computation

Ecologists often use dispersion metrics and statistical hypothesis testing to infer processes of community formation such as environmental filtering, competitive exclusion, and neutral species assembly. These metrics have limited power in inferring assembly models because they rely on often‐violated assumptions. Here, we adapt a model of phenotypic similarity and repulsion to simulate the process of community assembly via environmental filtering and competitive exclusion, all while parameterizing the strength of the respective ecological processes. We then use random forests and approximate Bayesian computation to distinguish between these models given the simulated data. We find that our approach is more accurate than using dispersion metrics and accounts for uncertainty in model selection. We also demonstrate that the parameter determining the strength of the assembly processes can be accurately estimated. This approach is available in the R package CAMI; Community Assembly Model Inference. We demonstrate the effectiveness of CAMI using an example of plant communities living on lava flow islands.     

Phylogenetic similarity and structure of Agaricomycotina communities across a forested landscape

The Agaricomycotina are a phylogenetically diverse group of fungi that includes both saprotrophic and mycorrhizal species, and that form species – rich communities in forest ecosystems. Most species are infrequently observed, and this hampers assessment of the role that environmental heterogeneity plays in determining local community composition and in driving β‐diversity. We used a combination of phenetic (TRFLP) and phylogenetic approaches [Unifrac and Net Relatedness Index (NRI)] to examine the compositional and phylogenetic similarity of Agaricomycotina communities in forest floor and surface soil of three widely distributed temperate upland forest ecosystems (one, xeric oak – dominated and two, mesic sugar maple dominated). Generally, forest floor and soil communities had similar phylogenetic diversity, but there was little overlap of species or evolutionary lineages between these two horizons. Forest floor communities were dominated by saprotrophic species, and were compositionally and phylogenetically similar in all three ecosystems. Mycorrhizal species represented 30% to 90% of soil community diversity, and these communities differed compositionally and phylogenetically between ecosystems. Estimates of NRI revealed significant phylogenetic clustering in both the forest floor and soil communities of only the xeric oak‐dominated forest ecosystem, and may indicate that this ecosystem acts as a habitat filter. Our results suggest that environmental heterogeneity strongly influences the phylogenetic β‐diversity of soil inhabiting Agaricomycotina communities, but has only a small influence on forest floor β‐diversity. Moreover, our results suggest that the strength of community assembly processes, such as habitat filtering, may differ between temperate forest ecosystems.     

Phylogenetic diversity and community assembly in a naturally fragmented system

We sought to assess effects of fragmentation and quantify the contribution of ecological processes to community assembly by measuring species richness, phylogenetic, and phenotypic diversity of species found in local and regional plant communities. Specifically, our fragmented system is Craters of the Moon National Monument and Preserve, Idaho, USA. CRMO is characterized by vegetated islands, kipukas, that are isolated in a matrix of lava. We used floristic surveys of vascular plants in 19 kipukas to create a local species list to compare traditional dispersion metrics, mean pairwise distance, and mean nearest taxon distance (MPD and MNTD), to a regional species list with phenotypic and phylogenetic data. We combined phylogenetic and functional trait data in a novel machine‐learning model selection approach, Community Assembly Model Inference (CAMI), to infer probability associated with different models of community assembly given the data. Finally, we used linear regression to explore whether the geography of kipukas explained estimated support for community assembly models. Using traditional metrics of MPD and MNTD neutral processes received the most support when comparing kipuka species to regional species. Individually no kipukas showed significant support for overdispersion. Rather, five kipukas showed significant support for phylogenetic clustering using MPD and two kipukas using MNTD. Using CAMI, we inferred neutral and filtering models structured the kipuka plant community for our trait of interest. Finally, we found as species richness in kipukas increases, model support for competition decreases and lower elevation kipukas show more support for habitat filtering models. While traditional phylogenetic community approaches suggest neutral assembly dynamics, recently developed approaches utilizing machine learning and model choice revealed joint influences of assembly processes to form the kipuka plant communities. Understanding ecological processes at play in naturally fragmented systems will aid in guiding our understanding of how fragmentation impacts future changes in landscapes.     

放牧干扰梯度下川西亚高山植物群落的组合机理

为了阐明放牧干扰对川西亚高山区域植物群落的组合过程以及群落结构的影响, 研究了放牧干扰梯度下的功能群均匀度和群落谱系结构的变化趋势。结果显示: 在干扰较轻的阔叶林与针叶林样地, 部分样方的功能群均匀度显著高于无效模型, 随着干扰梯度的增强, 功能群均匀度呈线性下降, 样方平均值从0.930降至0.840, 其高于无效模型的次数也逐渐降低, 干扰程度较大的草甸中出现部分样方的功能群均匀度显著低于无效模型。随着干扰程度的增强, 群落的谱系结构指数也呈逐渐上升趋势, 净关联指数平均值由-0.634逐渐增加至2.360, 邻近类群指数由-0.158上升至2.179。草甸与低矮灌丛受干扰较为严重, 其大部分样方的谱系结构指数显著高于随机群落, 表明干扰群落的谱系结构呈聚集分布。功能群均匀度与谱系结构的变化趋势一致, 表明生境筛滤效应与种间竞争作用的平衡决定着群落的组合过程。干扰降低了竞争作用, 促进了少数耐干扰功能群的优势地位, 造成功能群均匀度下降, 同时通过生境筛滤作用, 使群落的谱系结构呈现出聚集分布; 而未干扰的群落中由于竞争作用的效应, 功能群均匀度较高, 谱系结构也更加分散。研究区域植物群落的功能群均匀度与物种丰富度呈负相关, 表明物种间特别是相似物种间的竞争限制了群落的物种多样性。研究结果说明, 生态位分化和物种间的相互竞争在物种共存与群落组合中具有重要作用。     

Climate and Species Richness Predict the Phylogenetic Structure of African Mammal Communities

We have little knowledge of how climatic variation (and by proxy, habitat variation) influences the phylogenetic structure of tropical communities. Here, we quantified the phylogenetic structure of mammal communities in Africa to investigate how community structure varies with respect to climate and species richness variation across the continent. In addition, we investigated how phylogenetic patterns vary across carnivores, primates, and ungulates. We predicted that climate would differentially affect the structure of communities from different clades due to between-clade biological variation. We examined 203 communities using two metrics, the net relatedness (NRI) and nearest taxon (NTI) indices. We used simultaneous autoregressive models to predict community phylogenetic structure from climate variables and species richness. We found that most individual communities exhibited a phylogenetic structure consistent with a null model, but both climate and species richness significantly predicted variation in community phylogenetic metrics. Using NTI, species rich communities were composed of more distantly related taxa for all mammal communities, as well as for communities of carnivorans or ungulates. Temperature seasonality predicted the phylogenetic structure of mammal, carnivoran, and ungulate communities, and annual rainfall predicted primate community structure. Additional climate variables related to temperature and rainfall also predicted the phylogenetic structure of ungulate communities. We suggest that both past interspecific competition and habitat filtering have shaped variation in tropical mammal communities. The significant effect of climatic factors on community structure has important implications for the diversity of mammal communities given current models of future climate change.     

Beta‐diversity of central European forests decreases along an elevational gradient due to the variation in local community assembly processes

Beta‐diversity has been repeatedly shown to decline with increasing elevation, but the causes of this pattern remain unclear, partly because they are confounded by coincident variation in alpha‐ and gamma‐diversity. We used 8795 forest vegetation‐plot records from the Czech National Phytosociological Database to compare the observed patterns of beta diversity to null‐model expectations (beta‐deviation) controlling for the effects of alpha‐ and gamma‐diversity. We tested whether β‐diversity patterns along a 1200 m elevation gradient exclusively depend on the effect of varying species pool size, or also on the variation of the magnitude of community assembly mechanisms determining the distribution of species across communities (e.g. environmental filtering, dispersal limitation). The null model we used is a novel extension of an existing null‐model designed for presence/absence data and was specifically designed to disrupt the effect of community assembly mechanisms, while retaining some key features of observed communities such as average species richness and species abundance distribution. Analyses were replicated in ten subregions with comparable elevation ranges. Beta‐diversity declined along the elevation gradient due to a decrease in gamma‐diversity, which was steeper than the decrease in alpha‐diversity. This pattern persisted after controlling for alpha‐ and gamma‐diversity variation, and the results were robust when different resampling schemes and diversity metrics were used. We conclude that in temperate forests the pattern of decreasing beta‐diversity with elevation does not exclusively depend on variation in species pool size, as has been hypothesized, but also on variation in community assembly mechanisms. The results were consistent across resampling schemes and diversity measures, thus supporting the use of vegetation‐plot databases for understanding patterns of beta‐diversity at the regional scale.     

喀斯特退化天坑不同坡向植物群落系统发育与功能性状结构

喀斯特退化天坑地下森林主要分布在不同坡向的倒石坡上,探究环境过滤、竞争排斥和随机过程在天坑地下森林群落构建中的作用,有助于揭示天坑物种避难所价值,为天坑植物多样性的保护及坑外地表生态恢复提供科学的参考。以云南沾益深陷塘退化天坑为例,综合运用功能性状生态学和系统发育方法,研究不同坡向植物群落的土壤环境特征、功能性状变异、系统发育信号及系统发育多样性。研究表明：（1）不同坡向土壤环境差异较大,相对于南坡和北坡而言,东坡的土壤环境变异程度最大。（2）各坡向具有不同的优势种群,其中壳斗科植物种群多度较大,在北坡和东坡的黄背栎（Quercus pannosa）种群数量较大,在南坡则是青冈（Cyclobalanopsis glauca）多度最大。各优势物种功能性状在不同坡向存在差异,整体上变异程度中等,但南坡优势物种变异程度最大。（3）叶宽、长宽比、叶厚度、比叶面积具有较强且显著的系统发育信号（K>1,P<0.05）,性状进化过程中受遗传因素影响较大;而叶长、叶面积、叶干重、叶干物质含量、最大树高系统发育信号弱（K<1）,性状进化过程中受生境因素影响较大。（4）各坡向植物群落构建过程由生态位理论的环境过滤和竞争排斥作用主导,坡向从北坡-东坡-南坡的转变过程中,植物群落构建中环境过滤作用逐渐减小,竞争排斥作用逐渐增大。北坡可能较长时间内都是以旱生型植物为主要建群种,而南坡则可能最终演替成亚热带湿润气候型的顶极群落。     

Taxonomic,functional, and phylogenetic β‐diversity patterns of stream fish assemblages in tropical agroecosystems

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Integrating trait and phylogenetic distances to assess scale‐dependent community assembly processes

Biodiversity is structured by multiple mechanisms that are dependent, at least in part, on ecological similarities and differences among species. Integrating traits and phylogenies in diversity metrics may provide deeper insight into community assembly processes across spatial scales. However, different traits are influenced by processes at different spatial scales, and it is not clear how trait‐spatial scale mismatches skew our ability to detect assembly patterns. An additional complexity is how phylogenetic distances, which might capture unmeasured traits, reflect spatially dependent processes. Here we analyze a freshwater zooplankton dataset from 91 ponds and show that different traits are associated with processes at different spatial scales. We first assessed the response of individual traits to processes at both α‐ and β‐scales, and then quantified the power of different combinations of traits and phylogenetic distances to reveal environmental and spatial drivers of α‐ and β‐diversity. We found that explanatory power was maximised when we accounted for environmental and spatial drivers with single, but different traits for α‐ and β‐diversity. Using the most appropriate trait for each spatial scale outperformed phylogenetic information, but phylogenetic information outperformed the same traits when these were used at the wrong spatial scale, and all outperformed taxonomic analyses that ignore trait and phylogenetic information. We demonstrate that accounting for species’ similarities and differences provides important information about dominant assembly mechanisms at different spatial scales, and that phylogeny is especially useful when measured traits are uninformative at a given spatial scale or when there is lack of trait data. Our study also indicates, however, that trait‐scale mismatches among phylogenetically conserved traits may affect the performance of phylogenetic indices compared to indices that account only for the best single trait at each spatial scale.     

Evidence for community assembly constraints during succession in dune slack plant communities

Community assembly during succession can be constrained by both local and regional factors. Despite an increasing regional species pool size during succession, we found a limit on the number of species in 1 × 1 m plots in dune slacks. Three alternative hypotheses (habitat heterogeneity, dispersal limitation and niche limitation) explaining this community saturation were tested. A null model analysis showed that species richness in the plots had an unusually low variance suggesting that beta habitat diversity was not likely to explain the limitation on species richness. Because we did not find a correlation between the distribution of species over the slack and their dispersal capacity, we also excluded the dispersal limitation hypothesis. Finally, a guild proportionality analysis revealed that the abundances of forb, graminoid and ruderal species showed low an unusually low variance over all age classes involved. This provides evidence for nonrandom community assembly during succession, likely to be determined by competitive exclusion among species of the same guild.     

Neutral processes and species sorting in benthic microalgal community assembly: effects of tidal resuspension

Benthic microalgae (BMA) provide vital food resources for heterotrophs and stabilize sediments with their extracellular secretions. A central goal in ecology is to understand how processes such as species interactions and dispersal, contribute to observed patterns of species abundance and distribution. Our objectives were to assess the effects of sediment resuspension on microalgal community structure. We tested whether taxa‐abundance distributions could be predicted using neutral community models (NCMs) and also specific hypotheses about passive migration: (i) As migration decreases in sediment patches, BMA α‐diversity will decrease, and (ii) As migration decreases, BMA community dissimilarity (β‐diversity) will increase. Co‐occurrence indices (checkerboard score and variance ratio) were also computed to test for deterministic factors, such as competition and niche differentiation, in shaping communities. Two intertidal sites (mudflat and sand bar) differing in resuspension regime were sampled throughout the tidal cycle. Fluorometry and denaturing gradient gel electrophoresis were utilized to investigate diatom community structure. Observed taxa‐abundances fit those predicted from NCMs reasonably well (R² of 0.68–0.93), although comparisons of observed local communities to artificial randomly assembled communities rejected the null hypothesis that diatom communities were assembled solely by stochastic processes. No co‐occurrence tests indicated a significant role for competitive exclusion or niche partitioning in microalgal community assembly. In general, predictions about relationships between migration and species diversity were supported for local community dynamics. BMA at low tide (lowest migration) exhibited reduced α‐diversity as compared to periods of immersion at both mudflat and sand bar sites. β‐diversity was higher during low tide emersion on the mudflat, but did not differ temporally at the sand bar site. In between‐site metacommunity comparisons, low‐ and high‐resuspension sites exhibited distinct community compositions while the low‐energy mudflats contained higher microalgal biomass and greater α‐diversity. To our knowledge this is the first study to test the relevance of neutral processes in structuring marine microalgal communities. Our results demonstrate a prominent role for stochastic factors in structuring local BMA community assembly, although unidentified nonrandom processes also appear to play some role. High passive migration, in particular, appears to help maintain species diversity and structure communities in both sand and muddy habitats.     

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.     

Underdispersion and overdispersion of traits in terrestrial snail communities on islands

Understanding and disentangling different processes underlying the assembly and diversity of communities remains a key challenge in ecology. Species can assemble into communities either randomly or due to deterministic processes. Deterministic assembly leads to species being more similar (underdispersed) or more different (overdispersed) in certain traits than would be expected by chance. However, the relative importance of those processes is not well understood for many organisms, including terrestrial invertebrates. Based on knowledge of a broad range of species traits, we tested for the presence of trait underdispersion (indicating dispersal or environmental filtering) and trait overdispersion (indicating niche partitioning) and their relative importance in explaining land snail community composition on lake islands. The analysis of community assembly was performed using a functional diversity index (Rao's quadratic entropy) in combination with a null model approach. Regression analysis with the effect sizes of the assembly tests and environmental variables gave information on the strength of under‐ and overdispersion along environmental gradients. Additionally, we examined the link between community weighted mean trait values and environmental variables using a CWM‐RDA. We found both trait underdispersion and trait overdispersion, but underdispersion (eight traits) was more frequently detected than overdispersion (two traits). Underdispersion was related to four environmental variables (tree cover, habitat diversity, productivity of ground vegetation, and location on an esker ridge). Our results show clear evidence for underdispersion in traits driven by environmental filtering, but no clear evidence for dispersal filtering. We did not find evidence for overdispersion of traits due to diet or body size, but overdispersion in shell shape may indicate niche differentiation between snail species driven by small‐scale habitat heterogeneity. The use of species traits enabled us to identify key traits involved in snail community assembly and to detect the simultaneous occurrence of trait underdispersion and overdispersion.