Spatio-temporal dynamics of species richness in coastal fish communities

Determining patterns of change in species richness and the processes underlying the dynamics of biodiversity are of key interest within the field of ecology, but few studies have investigated the dynamics of vertebrate communities at a decadal temporal scale. Here, we report findings on the spatio-temporal variability in the richness and composition of fish communities along the Norwegian Skagerrak coast having been surveyed for more than half a century. Using statistical models incorporating non-detection and associated sampling variance, we estimate local species richness and changes in species composition allowing us to compute temporal variability in species richness. We tested whether temporal variation could be related to distance to the open sea and to local levels of pollution. Clear differences in mean species richness and temporal variability are observed between fjords that were and were not exposed to the effects of pollution. Altogether this indicates that the fjord is an appropriate scale for studying changes in coastal fish communities in space and time. The year-to-year rates of local extinction and turnover were found to be smaller than spatial differences in community composition. At the regional level, exposure to the open sea plays a homogenizing role, possibly due to coastal currents and advection.     

Global patterns and predictors of tropical reef fish species richness

In the marine realm, the tropics host an extraordinary diversity of taxa but the drivers underlying the global distribution of marine organisms are still under scrutiny and we still lack an accurate global predictive model. Using a spatial database for 6336 tropical reef fishes, we attempted to predict species richness according to geometric, biogeographical and environmental explanatory variables. In particular, we aimed to evaluate and disentangle the predictive performances of temperature, habitat area, connectivity, mid‐domain effect and biogeographical region on reef fish species richness. We used boosted regression trees, a flexible machine‐learning technique, to build our predictive model and structural equation modeling to test for potential ‘mediation effects’ among predictors. Our model proved to be accurate, explaining 80% of the total deviance in fish richness using a cross‐validated procedure. Coral reef area and biogeographical region were the primary predictors of reef fish species richness, followed by coast length, connectivity, mid‐domain effect and sea surface temperature, with interactions between the region and other predictors. Important indirect effects of water temperature on reef fish richness, mediated by coral reef area, were also identified. The relationship between environmental predictors and species richness varied markedly among biogeographical regions. Our analysis revealed that a few easily accessible variables can accurately predict reef fish species richness. They also highlight concerns regarding ongoing environmental declines, with region‐specific responses to variation in environmental conditions predicting a variable response to anthropogenic impacts.     

Richness, structure and functioning in metazoan parasite communities

Ecosystem functioning, characterized by components such as productivity and stability, has been extensively linked with diversity in recent years, mainly in plant ecology. The aim of our study was thus to quantify general relationships between diversity, community structure and ecosystem functions in metazoan parasite communities. We used data on parasite communities from 15 species of marine fish hosts from coastal Chile. The volumetric abundance (volume of all parasite species per individual host, in mm³) was used as a surrogate for productivity. Species diversity was measured using both species richness and evenness, while community structure was estimated using the co‐occurrence indices V‐ratio, C‐score and a new C‐score_s index standardized for the number of host replicates. After correcting for fish size, 47% of host species show no relationship, 13% show a hump shaped curve and 40% show positive monotonic relationships between productivity and parasite richness across all host individuals in a sample. We obtained a logarithmically decreasing relationship between evenness and productivity for all fish species, and propose a ‘dominance‐resistance’ hypothesis based on immunity to explain this pattern. The stability of the parasite community, measured as the coefficient of variation in productivity among individual hosts, was strongly and positively related to mean species richness across the 15 host species. The C‐score_s index, based on the number of checkerboard units in the host‐parasite presence/absence matrix, increases linearly with mean productivity across the 15 host species, suggesting that parasite communities tend to be more structured when they are more productive. This is the likely reason why linear relationships between richness and productivity were not observed consistently in all fish species. Parasite communities provide some clear patterns for the diversity–ecosystem functioning debate in ecology, although other factors, such as the history of community assembly, may also influence these patterns.     

Protist Diversity and Seasonal Dynamics in Skagerrak Plankton Communities as Revealed by Metabarcoding and Microscopy

Protist community composition and seasonal dynamics are of major importance for the production of higher trophic levels, such as zooplankton and fish. Our aim was to reveal how the protist community in the Skagerrak changes through the seasons by combining high‐throughput sequencing and microscopy of plankton collected monthly over two years. The V4 region of the 18S rRNA gene was amplified by eukaryote universal primers from the total RNA/cDNA. We found a strong seasonal variation in protist composition and proportional abundances, and a difference between two depths within the euphotic zone. Highest protist richness was found in late summer‐early autumn, and lowest in winter. Temperature was the abiotic factor explaining most of the variation in diversity. Dinoflagellates was the most abundant and diverse group followed by ciliates and diatoms. We found about 70 new taxa recorded for the first time in the Skagerrak. The seasonal pattern in relative read abundance of major phytoplankton groups was well in accordance with microscopical biovolumes. This is the first metabarcoding study of the protist plankton community of all taxonomic groups and through seasons in the Skagerrak, which may serve as a baseline for future surveys to reveal effects of climate and environmental changes.     

The roles of community biomass and species pools in the regulation of plant diversity

Considerable debate has developed over the importance of community biomass and species pools in the regulation of community diversity. Attempts to explain patterns of plant diversity as a function of community biomass or productivity have been only partially successful and, in general, have explained only a fraction of the observed variation in diversity. At the same time, studies that have focused on the importance of species pools have led some to conclude that diversity is primarily regulated in the short term by the size of the species pool rather than by biotic interactions. In this paper, I explore how community biomass and species pools may work in combination to regulate diversity in herbaceous plant communities. To address this problem, I employ a simple model in which the dynamics of species richness are a function of aboveground community biomass and environmentally controlled gradients in species pools. Model results lead to two main predictions about the role of biomass regulation: (1) Seasonal dynamics of richness will tend to follow a regular oscillation, with richness rising to peak values during the early to middle portion of the growing season and then declining during the latter part of the season. (2) Seasonal dieback of aboveground tissues facilitates the long‐term maintenance of high levels of richness in the community. The persistence of aboveground tissues and accumulation of litter are especially important in limiting the number of species through the suppression of recruitment. Model results also lead to two main predictions about the role of species pools: (1) The height and position of peak richness relative to community biomass will be influenced by the rate at which the species pool increases as available soil resources increase. (2) Variations in nonresource environmental factors (e.g. soil pH or soil salinity) have the potential to regulate species pools in a way that is uncorrelated with aboveground biomass. Under extreme conditions, such nonresource effects can create a unimodal envelope of biomass–richness values. Available evidence from the literature provides partial support for these predictions, though additional data are needed to provide more convincing tests.     

Dissecting spatiotemporal patterns of functional diversity through the lens of Darwin's naturalization conundrum

Darwin's naturalization conundrum describes the paradigm that community assembly is regulated by two opposing processes, environmental filtering and competitive interactions, which predict both similarity and distinctiveness of species to be important for establishment. Our goal is to use long‐term, large‐scale, and high‐resolution temporal data to examine diversity patterns over time and assess whether environmental filtering or competition plays a larger role in regulating community assembly processes. We evaluated Darwin's naturalization conundrum and how functional diversity has changed in the Laurentian Great Lakes fish community from 1870 to 2010, which has experienced frequent introductions of non‐native species and extirpations of native species. We analyzed how functional diversity has changed over time by decade from 1870 to 2010 at three spatial scales (regional, lake, and habitat) to account for potential noninteractions between species at the regional and lake level. We also determined which process, environmental filtering or competitive interactions, is more important in regulating community assembly and maintenance by comparing observed patterns to what we would expect in the absence of an ecological mechanism. With the exception of one community, all analyses show that functional diversity and species richness has increased over time and that environmental filtering regulates community assembly at the regional level. When examining functional diversity at the lake and habitat level, the regulating processes become more context dependent. This study is the first to examine diversity patterns and Darwin's conundrum by integrating long‐term, large‐scale, and high‐resolution temporal data at multiple spatial scales. Our results confirm that Darwin's conundrum is highly context dependent.     

The relationship between species richness and ecosystem variability is shaped by the mechanism of coexistence

Theory relating species richness to ecosystem variability typically ignores the potential for environmental variability to promote species coexistence. Failure to account for fluctuation‐dependent coexistence may explain deviations from the expected negative diversity–ecosystem variability relationship, and limits our ability to predict the consequences of increases in environmental variability. We use a consumer‐resource model to explore how coexistence via the temporal storage effect and relative nonlinearity affects ecosystem variability. We show that a positive, rather than negative, diversity–ecosystem variability relationship is possible when ecosystem function is sampled across a natural gradient in environmental variability and diversity. We also show how fluctuation‐dependent coexistence can buffer ecosystem functioning against increasing environmental variability by promoting species richness and portfolio effects. Our work provides a general explanation for variation in observed diversity–ecosystem variability relationships and highlights the importance of conserving regional species pools to help buffer ecosystems against predicted increases in environmental variability.     

Temporal and spatial changes in benthic invertebrate trophic networks along a taxonomic richness gradient

Species interactions underlie most ecosystem functions and are important for understanding ecosystem changes. Representing one type of species interaction, trophic networks were constructed from biodiversity monitoring data and known trophic links to assess how ecosystems have changed over time. The Baltic Sea is subject to many anthropogenic pressures, and low species diversity makes it an ideal candidate for determining how pressures change food webs. In this study, we used benthic monitoring data for 20 years (1980–1989 and 2010–2019) from the Swedish coast of the Baltic Sea and Skagerrak to investigate changes in benthic invertebrate trophic interactions. We constructed food webs and calculated fundamental food web metrics evaluating network horizontal and vertical diversity, as well as stability that were compared over space and time. Our results show that the west coast of Sweden (Skagerrak) suffered a reduction in benthic invertebrate biodiversity by 32% between the 1980s and 2010s, and that the number of links, generality of predators, and vulnerability of prey have been significantly reduced. The other basins (Bothnian Sea, Baltic Proper, and Bornholm Basin) do not show any significant changes in species richness or consistent significant trends in any food web metrics investigated, demonstrating resilience at a lower species diversity. The decreased complexity of the Skagerrak food webs indicates vulnerability to further perturbations and pressures should be limited as much as possible to ensure continued ecosystem functions.     

Effects of foundation species genotypic diversity on subordinate species richness in an assembling community

Foundation (dominant or matrix) species play a key role in structuring plant communities, influencing processes from population to ecosystem scales. However, the effects of genotypic diversity of foundation species on these processes have not been thoroughly assessed in the context of assembling plant communities. We modified the classical filter model of community assembly to include genotypic diversity as part of the biotic filter. We hypothesized that the proportion of fit genotypes (i.e. competitively superior and dominant) affects niche space availability for subordinate species to establish with consequence for species diversity. To test this hypothesis, we used an individual‐based simulation model where a foundation species of varying genotypic diversity (number of genotypes and variability among genotypes) competes for space with subordinate species on a spatially heterogeneous lattice. Our model addresses a real and practical problem in restoration ecology: choosing the level of genetic diversity of re‐introduced foundation and subordinate species. Genotypic diversity of foundation species significantly affected equilibrium community diversity, measured as species richness, either positively or negatively, depending upon environmental heterogeneity. Increases in genotypic diversity gave the foundation species a wider niche breadth. Under conditions of high environmental heterogeneity, this wider niche breadth decreased niche space for other species, lowering species richness with increased genotypic diversity until the genotypes of the foundation species saturated the landscape. With a low level of environmental heterogeneity, increasing genotypic diversity caused the foundation species niche breadth to be overdispersed, resulting in a weak positive relationship with species richness. Under these conditions, some genotypes are maladapted to the environment lowering fitness of the foundation species. These effects of genotypic diversity were secondary to the larger effects of overall foundation species fitness and environmental heterogeneity. The novel aspect of incorporating genotype diversity in combination with environmental heterogeneity in community assembly models include predictions of either positive or negative relationships between species diversity and genotypic diversity depending on environmental heterogeneity, and the conditions under which these factors are potentially relevant. Mechanistically, differential niche availability is imposed by the foundation species.     

Stream flow, salmon and beaver dams: roles in the structuring of stream fish communities within an anadromous salmon dominated stream

The current paradigm of fish community distribution is one of a downstream increase in species richness by addition, but this concept is based on a small number of streams from the mid-west and southern United States, which are dominated by cyprinids. Further, the measure of species richness traditionally used, without including evenness, may not be providing an accurate reflection of the fish community. We hypothesize that in streams dominated by anadromous salmonids, fish community diversity will be affected by the presence of the anadromous species, and therefore be influenced by those factors affecting the salmonid population. Catamaran Brook, New Brunswick, Canada, provides a long-term data set to evaluate fish community diversity upstream and downstream of an obstruction (North American beaver Castor canadensis dam complex), which affects distribution of Atlantic salmon Salmo salar. The Shannon Weiner diversity index and community evenness were calculated for sample sites distributed throughout the brook and over 15 years. Fish community diversity was greatest upstream of the beaver dams and in the absence of Atlantic salmon. The salmon appear to depress the evenness of the community but do not affect species richness. The community upstream of the beaver dams changes due to replacement of slimy sculpin Cottus cognatus by salmon, rather than addition, when access is provided. Within Catamaran Brook, location of beaver dams and autumn streamflow interact to govern adult Atlantic salmon spawner distribution, which then dictates juvenile production and effects on fish community. These communities in an anadromous Atlantic salmon dominated stream do not follow the species richness gradient pattern shown in cyprinid-dominated streams and an alternative model for stream fish community distribution in streams dominated by anadromous salmonids is presented. This alternative model suggests that community distribution may be a function of semipermeable obstructions, streamflow and the distribution of the anadromous species affecting resident stream fish species richness, evenness, biomass and production.     

Demersal fish diversity of the isolated Rockall plateau compared with the adjacent west coast shelf of Scotland

This study compares the diversity of the demersal fish assemblage of an isolated shelf sea, the Rockall plateau, with that of the nearest adjacent continental shelf sea ecosystem, the west coast of Scotland. Bottom trawl surveys were carried out between 1986 and 2008 on the Rockall plateau and the adjacent shelf sea to the west of Scotland. All demersal fish were identified and counted. Analyses of species richness and abundance were undertaken. The fish assemblage of the Rockall plateau was less diverse and the proportional representation of species was markedly different. A number of species common at Rockall were rare on the west coast shelf and, in general, there were fewer common and temporally stable species and more rare and ephemeral species at Rockall. Some species absent from Rockall have life‐history stages associated with inshore habitat. The Rockall plateau fish assemblage can be described as an impoverished subset of the north‐west European shelf sea fish assemblage. It is likely that there are constraints on diversity imposed by the relatively small area of the Rockall plateau and its degree of isolation by depth, distance and ocean currents. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 138–147.     

Variation in freshwater fish assemblages along a regional elevation gradient in the northern Andes,Colombia

Studies on elevation diversity gradients have covered a large number of taxa and regions throughout the world; however, studies of freshwater fish are scarce and restricted to examining their changes along a specific gradient. These studies have reported a monotonic decrease in species richness with increasing elevation, but ignore the high taxonomic differentiation of each headwater assemblage that may generate high β‐diversity among them. Here, we analyzed how fish assemblages vary with elevation among regional elevation bands, and how these changes are related to four environmental clines and to changes in the distribution, habitat use, and the morphology of fish species. Using a standardized field sampling technique, we assessed three different diversity and two structural assemblage measures across six regional elevation bands located in the northern Andes (Colombia). Each species was assigned to a functional group based on its body shape, habitat use, morphological, and/or behavioral adaptations. Additionally, at each sampling site, we measured four environmental variables. Our analyses showed: (1) After a monotonic decrease in species richness, we detected an increase in richness in the upper part of the gradient; (2) diversity patterns vary depending on the diversity measure used; (3) diversity patterns can be attributed to changes in species distribution and in the richness and proportions of functional groups along the regional elevation gradient; and (4) diversity patterns and changes in functional groups are highly correlated with variations in environmental variables, which also vary with elevation. These results suggest a novel pattern of variation in species richness with elevation: Species richness increases at the headwaters of the northern Andes owing to the cumulative number of endemic species there. This highlights the need for large‐scale studies and has important implications for the aquatic conservation of the region.     

Deep‐branching Novel Lineages and High Diversity of Haptophytes in the Skagerrak (Norway) Uncovered by 454 Pyrosequencing

Microalgae in the division Haptophyta may be difficult to identify to species by microscopy because they are small and fragile. Here, we used high‐throughput sequencing to explore the diversity of haptophytes in outer Oslofjorden, Skagerrak, and supplemented this with electron microscopy. Nano‐ and picoplanktonic subsurface samples were collected monthly for 2 yr, and the haptophytes were targeted by amplification of RNA/cDNA with Haptophyta‐specific 18S ribosomal DNA V4 primers. Pyrosequencing revealed higher species richness of haptophytes than previously observed in the Skagerrak by microscopy. From ca. 400,000 reads we obtained 156 haptophyte operational taxonomic units (OTUs) after rigorous filtering and 99.5% clustering. The majority (84%) of the OTUs matched environmental sequences not linked to a morphological species, most of which were affiliated with the order Prymnesiales. Phylogenetic analyses including Oslofjorden OTUs and available cultured and environmental haptophyte sequences showed that several of the OTUs matched sequences forming deep‐branching lineages, potentially representing novel haptophyte classes. Pyrosequencing also retrieved cultured species not previously reported by microscopy in the Skagerrak. Electron microscopy revealed species not yet genetically characterised and some potentially novel taxa. This study contributes to linking genotype to phenotype within this ubiquitous and ecologically important protist group, and reveals great, unknown diversity.     

Homogenization of fish assemblages in different lake depth strata at local and regional scales

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Imprint of monsoonal patterns on the fish assemblage in coastal waters of south‐east India: a case study

The composition, species richness and diversity of a coastal fish assemblage from the Kalpakkam coast of south‐east India are described along with temporal distribution patterns related to seasonal fluctuations in dissolved oxygen, salinity, pH, chlorophyll‐a, phytoplankton and zooplankton species richness and density. A total of 244 fish species belonging to 21 orders, 87 families and 163 genera were recorded. The fish assemblage was dominated by reef‐associated species, followed by demersal species. The majority of the species (63%) are widely distributed in the western Indo‐Pacific as well as in the central Indo‐Pacific. Jaccard's coefficient analysis showed three distinct seasonal patterns of fish occurrence: pre‐monsoon (PrM), monsoon (M) and post‐monsoon (PoM). The maximum number of species was during the PrM period, followed by the PoM and M periods. Species occurrence analysis showed Sardinella longiceps to be dominant during PrM and M periods, Leiognathus dussumieri during the M period and Secutor insidiator and Secutor ruconius during the M and PoM periods. Canonical correspondence analysis indicated that salinity and rainfall were the two most influential environmental factors strongly correlated with temporal variation in the fish assemblage. The physico‐chemical conditions, in combination with factors such as greater food availability and shelter, might control the seasonal local distribution of the ichthyofauna in these Indian coastal waters.     

Genetic and community similarities are correlated in endemic-rich springs of the northern Chihuahuan Desert

Aim Species diversity and genetic diversity within a taxon are intrinsic parts of global biodiversity. These two levels of biodiversity can show strong correlation due to a variety of reasons (i.e. parallel processes affecting both communities and populations, genotypes of a numerically or functionally dominant species affecting community composition, a species assemblage selecting for a particular genotype by affecting its selection regime). We examined correlations between species and genetic biodiversity in four isolated endemic‐rich spring systems in a hot desert and their potential link to environmental variables and physical isolation. Location Chihuahuan Desert spring systems in the Pecos River basin of New Mexico and Texas, USA. Methods We compared species richness of fish and benthic macroinvertebrate communities to within‐population allelic richness of amphipods (monophyletic Gammarus spp.) and Pecos gambusia (Gambusia nobilis) using Mantel tests. We also compared pairwise community similarities with pairwise genetic identities of populations among the same groups. We tested correlations among diversity, similarity and environmental variables after controlling for the effects of spatial distances using partial Mantel tests. We partitioned genetic and species diversity into three spatial scales (i.e. individual springs, individual spring systems, the entire region) using AMOVA and partition . Results We found strong correlations between invertebrate species richness and mosquitofish allelic richness. We found even stronger correlations of amphipod and gambusia genetic identities with fish and invertebrate community similarities; these were best explained by geographic distance rather than abiotic environmental factors. Most of the taxa and communities exhibited the largest proportion of diversity at the regional level. Main conclusions Our results suggest that drift and migration are the mechanisms that best explain our observations, and although α‐diversity among genes and species may not be strongly correlated, the pattern of species and allelic complementarity among these groups seems to be concordant at the regional level.     

The carrying capacity for species richness

The idea that the number of species within an area is limited by a specific capacity of that area to host species is old yet controversial. Here, we show that the concept of carrying capacity for species richness can be as useful as the analogous concept in population biology. Many lines of empirical evidence indicate the existence of limits of species richness, at least at large spatial and phylogenetic scales. However, available evidence does not support the idea of diversity limits based on limited niche space; instead, carrying capacity should be understood as a stable equilibrium of biodiversity dynamics driven by diversity‐dependent processes of extinction, speciation and/or colonization. We argue that such stable equilibria exist even if not all resources are used and if increasing species richness increases the ability of a community to use resources. Evaluating the various theoretical approaches to modelling diversity dynamics, we conclude that a fruitful approach for macroecology and biodiversity science is to develop theory that assumes that the key mechanism leading to stable diversity equilibria is the negative diversity dependence of per‐species extinction rates, driven by the fact that population sizes of species must decrease with an increasing number of species owing to limited energy availability. The recently proposed equilibrium theory of biodiversity dynamics is an example of such a theory, which predicts that equilibrium species richness (i.e., carrying capacity) is determined by the interplay of the total amount of available resources, the ability of communities to use those resources, environmental stability that affects extinction rates, and the factors that affect speciation and colonization rates. We argue that the diversity equilibria resulting from these biodiversity dynamics are first‐order drivers of large‐scale biodiversity patterns, such as the latitudinal diversity gradient.     

Elevational Gradients in Fish Diversity in the Himalaya: Water Discharge Is the Key Driver of Distribution Patterns

Background

Studying diversity and distribution patterns of species along elevational gradients and understanding drivers behind these patterns is central to macroecology and conservation biology. A number of studies on biogeographic gradients are available for terrestrial ecosystems, but freshwater ecosystems remain largely neglected. In particular, we know very little about the species richness gradients and their drivers in the Himalaya, a global biodiversity hotspot.

Methodology/Principal Findings

We collated taxonomic and distribution data of fish species from 16 freshwater Himalayan rivers and carried out empirical studies on environmental drivers and fish diversity and distribution in the Teesta river (Eastern Himalaya). We examined patterns of fish species richness along the Himalayan elevational gradients (50–3800 m) and sought to understand the drivers behind the emerging patterns. We used generalized linear models (GLM) and generalized additive models (GAM) to examine the richness patterns; GLM was used to investigate relationship between fish species richness and various environmental variables. Regression modelling involved stepwise procedures, including elimination of collinear variables, best model selection, based on the least Akaike’s information criterion (AIC) and the highest percentage of deviance explained (D²). This maiden study on the Himalayan fishes revealed that total and non-endemic fish species richness monotonously decrease with increasing elevation, while endemics peaked around mid elevations (700–1500 m). The best explanatory model (synthetic model) indicated that water discharge is the best predictor of fish species richness patterns in the Himalayan rivers.

Conclusions/Significance

This study, carried out along one of the longest bioclimatic elevation gradients of the world, lends support to Rapoport’s elevational rule as opposed to mid domain effect hypothesis. We propose a species-discharge model and contradict species-area model in predicting fish species richness. We suggest that drivers of richness gradients in terrestrial and aquatic ecosystems are likely to be different. These studies are crucial in context of the impacts of unprecedented on-going river regulation on fish diversity and distribution in the Himalaya.     

Context‐dependent functional dispersion across similar ranges of trait space covered by intertidal rocky shore communities

Functional diversity is intimately linked with community assembly processes, but its large‐scale patterns of variation are often not well understood. Here, we investigated the spatiotemporal changes in multiple trait dimensions (“trait space”) along vertical intertidal environmental stress gradients and across a landscape scale. We predicted that the range of the trait space covered by local assemblages (i.e., functional richness) and the dispersion in trait abundances (i.e., functional dispersion) should increase from high‐ to low‐intertidal elevations, due to the decreasing influence of environmental filtering. The abundance of macrobenthic algae and invertebrates was estimated at four rocky shores spanning ca. 200 km of the coast over a 36‐month period. Functional richness and dispersion were contrasted against matrix‐swap models to remove any confounding effect of species richness on functional diversity. Random‐slope models showed that functional richness and dispersion significantly increased from high‐ to low‐intertidal heights, demonstrating that under harsh environmental conditions, the assemblages comprised similar abundances of functionally similar species (i.e., trait convergence), while that under milder conditions, the assemblages encompassed differing abundances of functionally dissimilar species (i.e., trait divergence). According to the Akaike information criteria, the relationship between local environmental stress and functional richness was persistent across sites and sampling times, while functional dispersion varied significantly. Environmental filtering therefore has persistent effects on the range of trait space covered by these assemblages, but context‐dependent effects on the abundances of trait combinations within such range. Our results further suggest that natural and/or anthropogenic factors might have significant effects on the relative abundance of functional traits, despite that no trait addition or extinction is detected.