Using functional diversity patterns to explore metacommunity dynamics: a framework for understanding local and regional influences on community structure

The metacommunity concept, describing how local and regional scale processes interact to structure communities, has been successfully applied to patterns of taxonomic diversity. Functional diversity has proved useful for understanding local scale processes, but has less often been applied to understanding regional scale processes. Here, we explore functional diversity patterns within a metacommunity context to help elucidate how local and regional scale processes influence community assembly. We detail how each of the four metacommunity perspectives (species sorting, mass effects, patch dynamics, neutral) predict different patterns of functional beta‐ and alpha‐diversity and spatial structure along two key gradients: dispersal limitation and environmental conditions. We then apply this conceptual model to a case study from alpine tundra plant communities. We sampled species composition in 17 ‘sky islands’ of alpine tundra in the Colorado Rocky Mountains, USA that differed in geographic isolation and area (key factors related to dispersal limitation) and temperature and elevation (key environmental factors). We quantified functional diversity in each site based on specific leaf area, leaf area, stomatal conductance, plant height and chlorophyll content. We found that colder high elevation sites were functionally more similar to each other (decreased functional beta‐diversity) and had lower functional alpha‐diversity. Geographic isolation and area did not influence functional beta‐ or alpha‐diversity. These results suggest a strong role for environmental conditions structuring alpine plant communities, patterns consistent with the species sorting metacommunity perspective. Incorporating functional diversity into metacommunity theory can help elucidate how local and regional factors structure communities and provide a framework for observationally examining the role of metacommunity dynamics in systems where experimental approaches are less tractable.     

The relative importance of local conditions and regional processes in structuring aquatic plant communities

1. The structure of biological communities reflects the influence of both local environmental conditions and processes such as dispersal that create patterns in species’ distribution across a region. 2. We extend explicit tests of the relative importance of local environmental conditions and regional spatial processes to aquatic plants, a group traditionally thought to be little limited by dispersal. We used partial canonical correspondence analysis and partial Mantel tests to analyse data from 98 lakes and ponds across Connecticut (northeastern United States). 3. We found that aquatic plant community structure reflects the influence of local conditions (pH, conductivity, water clarity, lake area, maximum depth) as well as regional processes. 4. Only 27% of variation in a presence/absence matrix was explained by environmental conditions and spatial processes such as dispersal. Of the total explained, 45% was related to environmental conditions and 40% to spatial processes. 5. Jaccard similarity declined with Euclidean distance between lakes, even after accounting for the increasing difference in environmental conditions, suggesting that dispersal limitation may influence community composition in the region. 6. The distribution of distances among lakes where species occurred was associated with dispersal‐related functional traits, providing additional evidence that dispersal ability varies among species in ways that affect community composition. 7. Although environmental and spatial variables explained a significant amount of variation in community structure, a substantial amount of stochasticity also affects these communities, probably associated with unpredictable colonisation and persistence of the plants.     

The scale-dependent importance of habitat factors and dispersal limitation in structuring Great Lakes shoreline plant communities

Niche-based and neutral models of community structure posit distinct mechanisms underlying patterns in community structure; correlation between species’ distributions and habitat factors points to niche assembly while spatial pattern independent of habitat suggests neutral assembly via dispersal limitation. The challenge is to disentangle the relative contributions when both processes are operating, and to determine the scales at which each is important. We sampled shoreline plant communities on an island in Lake Michigan, varying the extent and the grain of sampling, and used both distance-based correlation methods and variance partitioning to quantify the proportion of the variation in plant species composition that was attributable to habitat factors and to spatial configuration independent of habitat. Our results were highly scale dependent. We found no distance decay of plant community similarity at the island scale (1−33 km). All of the explained variation (32%) in species composition among samples at this scale was attributed to habitat factors. However, at a site intensively sampled at a smaller scale (5−1,200 m), similarity of species composition did decay with distance. Using a coarse sampling grain (transects), habitat factors explained 40% of the variation, but the purely spatial component explained a comparable 22%. Analyzing plots within transects revealed variation in species composition that was still jointly determined by habitat and spatial factors (18 and 11% of the variance, respectively). For both grain sizes, most of the habitat component was spatially structured, reflecting an abrupt alongshore transition from sandy dunes to cobble beach. Space per se explained more variation in species composition at a second site where the habitat transition was more gradual; here, habitat acted as a less selective filter, allowing the signal of dispersal limitation to be detected more readily. We conclude that both adaptation to specific habitat factors and habitat-independent spatial position indicative of dispersal limitation determine plant species composition in this system. Our results support the prediction that dispersal limitation—a potentially, but not necessarily, neutral driver—is relatively more important at smaller scales.     

From species distributions to meta‐communities

The extent that biotic interactions and dispersal influence species ranges and diversity patterns across scales remains an open question. Answering this question requires framing an analysis on the frontier between species distribution modelling (SDM), which ignores biotic interactions and dispersal limitation, and community ecology, which provides specific predictions on community and meta‐community structure and resulting diversity patterns such as species richness and functional diversity. Using both empirical and simulated datasets, we tested whether predicted occurrences from fine‐resolution SDMs provide good estimates of community structure and diversity patterns at resolutions ranging from a resolution typical of studies within reserves (250 m) to that typical of a regional biodiversity study (5 km). For both datasets, we show that the imprint of biotic interactions and dispersal limitation quickly vanishes when spatial resolution is reduced, which demonstrates the value of SDMs for tracking the imprint of community assembly processes across scales.     

Geographical patterns in the beta diversity of China's woody plants: the influence of space,environment and range size

Beta diversity (i.e. species turnover rate across space) is fundamental for understanding mechanisms controlling large‐scale species richness patterns. However, the influences on beta diversity are still a matter of debate. In particular, the relative role of environmental and spatial processes (e.g. environmental niche versus dispersal limitation of species) remains elusive, and the influence of species range size has been poorly tested. Here, using distribution maps of 11 405 woody species in China (ca 9.6 × 10⁶ km²), we investigated 1) the geographical and directional patterns of beta diversity for all woody species and species with different range sizes, and 2) compared the effects of environmental and spatial processes on these patterns. Beta diversity was calculated as the decay of similarity in species composition with increasing distance. Variables representing environmental energy, water availability, climatic seasonality, habitat heterogeneity and human activities were used to evaluate the effects of environmental processes, while spatial distance was used to assess the influence of spatial processes. The results indicated significant directional patterns of beta diversity: the similarity decay along the latitudinal gradient was 1.6–2.3 times faster than that along the longitudinal gradient. Beta diversity also increased with the decrease of species range size. As compared with spatial processes, environmental processes had stronger effects on longitudinal beta diversity and on the beta diversity of widely‐ranged species. This was opposite to the larger influence of spatial processes on latitudinal beta diversity and the beta diversity of narrowly‐ranged species. These results suggest that the distributions of narrowly‐ranged woody species in China may have not reached equilibrium with their environmental niches due to dispersal limitation induced by China's topography and/or their low dispersal ability. The projected rapid climatic changes will likely endanger such species. Species dispersal processes should be taken into account in future conservation strategies in China.     

Macroecological drivers of zooplankton communities across the mountains of western North America

Disentangling the environmental and spatial drivers of biological communities across large scales increasingly challenges modern ecology in a rapidly changing world. Here, we investigate the hierarchical and trait‐based organization of regional and local factors of zooplankton communities at a macroscale of 1240 mountain lakes and ponds spanning western North America (California, USA, to Yukon Territory, Canada). Variation partitioning was used to test the hypothesized importance of climate, connectivity, catchment features, and exotic sportfish to zooplankton beta‐diversity in the context of key functional traits (body size and reproductive dispersal potential) given the pronounced environmental heterogeneity (e.g. thermal gradients), topographic barriers, and legacy of stocked fish in mountainous regions. Dispersal limitation was inferred from multispecies patch connectivity estimates based on nearest and average distances to occupied patches. Environmental heterogeneity best explained community composition as catchment/lake features (morphometry, land cover, and lithology) collectively captured greater variation than did climate (temperature, precipitation, and solar radiation), local stocking, or connectivity; however, single climatic variables captured the most variation individually. Macrospatial variation by larger obligate sexual species was better explained than that by smaller cyclically parthenogenetic asexual species. Our results provide several novel insights into the macroecology of zooplankton of the North American Cordillera, demonstrating their stronger associations to climatically driven aquatic‐terrestrial habitat coupling than dynamics arising from introduced salmonids, human land‐use, or species dispersal. These findings highlight the clear and important role of these communities as bioindicators of the limnological impacts of accelerating rates of climate change, as their responses appear relatively not confounded by local human perturbations or dispersal limitation.     

History,rather than contemporary processes,determines variation in macroinvertebrate diversity in artesian springs: the expansion hypothesis

1. The relative importance of contemporary and historical processes is a fundamental question in understanding patterns of biodiversity. We tested the hypothesis that species‐sorting into different habitats (limnocrenes, rheocrenes, helocrenes), rather than history, would account for the greatest variation in macroinvertebrate diversity in desert artesian springs of The Bonneville Basin, U.S.A. These springs were isolated at the valley scale c. 9000 years ago by aridity and high salinity. Thus, the valley scale will account for the greatest variation in community composition if history and dispersal limitation are important, whereas niche‐based sorting processes will be most important if habitat accounts for the greatest variation in community composition. 2. We identified 302 taxa from 280 sites and used a partial redundancy analysis, additive partitioning and classification strength (CS) to partition the variability in diversity among the springs. The valley scale accounted for more variation in community composition in limnocrene habitats (32.5%) than all other spatial and environmental variables combined. Valleys also accounted for 58% (additive partitioning) and 83% (CS) of the regional variation in diversity in analyses that included all three habitat types. That is, the average community similarity was 25% across the region, but increased to 41% within valleys. By contrast, habitat filtering did not account for significant variation in community composition in any of the analyses. Our study is one of the few suggesting the over‐riding importance of neutral processes in determining patterns of diversity (history and dispersal limitation). 3. The ‘expansion’ hypothesis suggests that the youthful age of a region, combined with slow dispersal by a fauna dominated by generalists, will maximise the imprint of history. These communities appear to exist in a pre‐equilibrial state, where the maximum carrying capacity has not been reached and niche space is plentiful. With time, we predict that local richness will increase while β‐diversity decreases as species expand their distribution across the region. Consequently, the importance of niche‐based processes may increase with time as the imprint of history fades.     

Niche versus neutrality in structuring the beta diversity of damselfly assemblages

1. Differences among communities in taxonomic composition – beta diversity – are frequently expected to result from taxon‐specific responses to spatial variation in ecological conditions, through niche partitioning. Such process‐derived patterns are in sharp contrast to arguments from neutral theory, where taxa are ecologically equivalent and beta diversity results primarily from dispersal limitation. 2. Here, we compared beta diversity among assemblages of damselflies (Odonata: Zygoptera), for which previous experiments have shown that niche differences maintain genera within a community, but patterns of relative abundance for species within each genus are shaped primarily by neutral dynamics. 3. Using null‐model and ordination‐based methods, we find that both genera and (in contrast to neutral theory) species assemblage composition vary across the landscape in a deterministic fashion, shaped by environmental and spatial factors. 4. While the observed patterns in species composition conflict with theory, we suggest that this a result of weak ecological filters acting to produce spatial variation in assemblages of ecologically similar species undergoing ecological drift within communities. Such patterns are especially likely in systems of relatively weak dispersers like damselflies.     

How beta diversity and the underlying causes vary with sampling scales in the Changbai mountain forests

This study aims to establish a relationship between the sampling scale and tree species beta diversity temperate forests and to identify the underlying causes of beta diversity at different sampling scales. The data were obtained from three large observational study areas in the Changbai mountain region in northeastern China. All trees with a dbh ≥1 cm were stem‐mapped and measured. The beta diversity was calculated for four different grain sizes, and the associated variances were partitioned into components explained by environmental and spatial variables to determine the contributions of environmental filtering and dispersal limitation to beta diversity. The results showed that both beta diversity and the causes of beta diversity were dependent on the sampling scale. Beta diversity decreased with increasing scales. The best‐explained beta diversity variation was up to about 60% which was discovered in the secondary conifer and broad‐leaved mixed forest (CBF) study area at the 40 × 40 m scale. The variation partitioning result indicated that environmental filtering showed greater effects at bigger grain sizes, while dispersal limitation was found to be more important at smaller grain sizes. What is more, the result showed an increasing explanatory ability of environmental effects with increasing sampling grains but no clearly trend of spatial effects. The study emphasized that the underlying causes of beta diversity variation may be quite different within the same region depending on varying sampling scales. Therefore, scale effects should be taken into account in future studies on beta diversity, which is critical in identifying different relative importance of spatial and environmental drivers on species composition variation.     

The role of environmental and spatial processes in structuring native and non‐native fish communities across thousands of lakes

Quantifying the role of spatial patterns is an important goal in ecology to further understand patterns of community composition. We quantified the relative role of environmental conditions and regional spatial patterns that could be produced by environmental filtering and dispersal limitation on fish community composition for thousands of lakes. A database was assembled on fish community composition, lake morphology, water quality, climatic conditions, and hydrological connectivity for 9885 lakes in Ontario, Canada. We utilized a variation partitioning approach in conjunction with Moran's Eigenvector Maps (MEM) and Asymmetric Eigenvector Maps (AEM) to model spatial patterns that could be produced by human‐mediated and natural modes of dispersal. Across 9885 lakes and 100 fish species, environmental factors and spatial structure explained approximately 19% of the variation in fish community composition. Examining the proportional role of spatial structure and environmental conditions revealed that as much as 90% of the explained variation in native species assemblage composition is governed by environmental conditions. Conversely on average, 67% of the explained variation in non‐native assemblage composition can be related to human‐mediated dispersal. This study highlights the importance of including spatial structure and environmental conditions when explaining patterns of community composition to better discriminate between the ecological processes that underlie biogeographical patterns of communities composed of native and non‐native fish species.     

Contrasting patterns and mechanisms of spatial turnover for native and exotic freshwater fish in Europe

Aim We compare the distribution patterns of native and exotic freshwater fish in Europe, and test whether the same mechanisms (environmental filtering and/or dispersal limitation) govern patterns of decrease in similarity of native and exotic species composition over geographical distance (spatial species turnover). Locations Major river basins of Europe. Methods Data related to geography, habitat diversity, regional climate and species composition of native and exotic freshwater fish were collated for 26 major European river basins. We explored the degree of nestedness in native and exotic species composition, and quantified compositional similarity between river basins according to the beta‐sim (independent of richness gradient) and Jaccard (dependent of richness gradient) indices of similarity. Multiple regression on distance matrices and variation‐partitioning approaches were used to quantify the relative roles of environmental filtering and dispersal limitation in shaping patterns of decreasing compositional similarity over geographical distance. Results Native and exotic species exhibited significant nested patterns of species composition, indicating that differences in fish species composition between river basins are primarily the result of species loss, rather than species replacement. Both native and exotic compositional similarity decreased significantly with increasing geographical distance between river basins. However, gradual changes in species composition with geographical distance were found only for exotic species. In addition, exotic species displayed a higher rate of similarity decay (higher species turnover rate) with geographical distance, compared with native species. Lastly, the majority of explained variation in exotic compositional similarity was uniquely related to geography, whereas native compositional similarity was either uniquely explained by geography or jointly explained by environment and geography. Main conclusions Our study suggests that large‐scale patterns of spatial turnover for exotic freshwater fish in Europe are generated by human‐mediated dispersal limitation, whereas patterns of spatial turnover for native fish result from both dispersal limitation relative to historical events (isolation by mountain ranges, glacial history) and environmental filtering.     

Dispersal ability links to cross‐scale species diversity patterns across the Eurasian Arctic tundra

Aim The role of dispersal in structuring biodiversity across spatial scales is controversial. If dispersal controls regional and local community assembly, it should also affect the degree of spatial species turnover as well as the extent to which regional communities are represented in local communities. Here we provide the first integrated assessment of relationships between dispersal ability and local‐to‐regional spatial aspects of species diversity across a large geographical area. Location Northern Eurasia. Methods Using a cross‐scale analysis covering local (0.64 m²) to continental (the Eurasian Arctic biome) scales, we compared slope parameters of the dissimilarity‐to‐distance relationship in species composition and the local‐to‐regional relationship in species richness among three plant‐like groups that differ in dispersal ability: lichens with the highest dispersal ability; mosses and moss allies with intermediate dispersal ability; and seed plants with the lowest dispersal ability. Results Diversity patterns generally differed between the three groups according to their dispersal ability, even after controlling for niche‐based processes. Increasing dispersal ability is linked to decreasing spatial species turnover and an increasing ratio of local to regional species richness. All comparisons supported our expectations, except for the slope of the local‐to‐regional relationship in species richness for mosses and moss allies which was not significantly steeper than that of seed plants. Main conclusions The negative link between dispersal ability and spatial species turnover and the corresponding positive link between dispersal ability and the ratio of local‐to‐regional species richness support the idea that dispersal affects community structure and diversity patterns across spatial scales.     

哀牢山亚热带中山湿性常绿阔叶林树种beta多样性格局形成的驱动力

Beta多样性通常指群落在时间和空间上物种组成的差异, 包括物种周转组分和物种丰富度差异组分。驱动beta多样性格局形成的生态过程决定了群落的时空动态, 然而关于beta多样性及其两个组分格局形成的驱动力还存在较多争议。以往研究表明, beta多样性的格局存在取样尺度的依赖性, 驱动其形成的生态过程在不同取样尺度下的相对重要性也随之改变。本研究以哀牢山亚热带中山湿性常绿阔叶林20 ha动态监测样地为研究对象, 在不同取样尺度上, 将样方间的Bray-Curtis指数分解为物种周转组分和物种丰富度差异组分, 通过典范冗余分析和方差分解的方法揭示环境过滤和扩散限制对于beta多样性及其两个组分格局形成的相对重要性及其尺度依赖性。结果表明: (1) beta多样性、物种周转组分和物种丰富度差异组分均随取样尺度的增大而减小。在不同取样尺度下, 物种周转组分对于beta多样性的贡献始终占主导地位。(2)随着取样尺度的增大, 环境过滤驱动beta多样性格局形成的相对重要性逐渐增加, 而扩散限制的相对重要性逐渐降低。本研究进一步证实了取样尺度在beta多样性格局形成及其驱动力定量评价中的重要性, 今后的研究需要进一步解析上述尺度效应的形成机制。     

Geographical and environmental controls of palm beta diversity in paleo-riverine terrace forests in Amazonian Peru

The palm (Arecaceae) community on low paleo-riverine terraces (terrace forest) in the north-western Amazon, is described, and we assessed the importance of environmental differences and geographic distance as drivers of its local (25² grain and 0–500 extent) and regional scale (500² grain and 0.3–143 km extent) beta diversity using ordination, multiple regressions on distance matrices and Indicator Species Analysis. A total of 15,869 individuals and 37 species of palm were sampled in 10 terrace forest transects, while 3758 individuals and 21 species were sampled in two adjacent floodplain forest transects for comparison. The terrace and floodplain forest were clearly different in their diversity and floristic composition. The relative importance of geographical distance and environmental difference as controls of terrace forest beta diversity was scale dependent, with environmental differences, notably in soil moisture, dominating at local scales and geographical distance dominating at regional scales. In fact, none of the environmental factors had a significant influence on regional-scale beta diversity. The geographical distance decay in floristic similarity was markedly steeper at local scale ( −0.25 km ⁻¹) than at regional scale ( −0.003 km⁻¹). Such a nonlinear decay is expected if simple dispersal limitation controls beta diversity. However, the absent flattening of the distance decay at the largest distances and the sub-Andean affinities of the westernmost palm communities suggest that large-scale biogeographical processes also contribute to the regional-scale beta diversity. Hereby our results indicate that not only local environment, but also dispersal limitation and biogeographical history can be important controls of the diversity and composition of local plant communities.     

Phylogenetic community structure and phylogenetic turnover across space and edaphic gradients in western Amazonian tree communities

Ecological and evolutionary processes influence community assembly at both local and regional scales. Adding a phylogenetic dimension to studies of species turnover allows tests of the extent to which environmental gradients, geographic distance and the historical biogeography of lineages have influenced speciation and dispersal of species throughout a region. We compare measures of beta diversity, phylogenetic community structure and phylobetadiversity (phylogenetic distance among communities) in 34 plots of Amazonian trees across white‐sand and clay terra firme forests in a 60 000 square kilometer area in Loreto, Peru. Dominant taxa in white‐sand forests were phylogenetically clustered, consistent with environmental filtering of conserved traits. Phylobetadiversity measures found significant phylogenetic clustering between terra firme communities separated by geographic distances of <200–300 km, consistent within recent local speciation at the watershed scale in the Miocene‐aged clay‐soil forests near the foothills of the Andes. Although both distance and habitat type yielded statistically significant effects on both species and phylogenetic turnover, the patterns we observed were more consistent with an effect of habitat specialization than dispersal limitation. Our results suggest a role for both broad‐scale biogeographic and evolutionary processes, as well as habitat specialization, influencing community structure in Amazonian forests.     

地理距离及环境差异对云南元江干热河谷植物群落beta多样性的影响

beta多样性反映了群落间物种组成的差异, 是生物多样性研究的热点之一。本研究通过对云南元江干热河谷41个植物群落样方进行调查, 用Jaccard相异系数表征物种beta多样性, 用样方之间的最近谱系距离(mean nearest taxon distance, MNTD)及平均谱系距离(mean pairwise distance, MPD)表征谱系beta多样性, 采用基于距离矩阵的多元回归和方差分解方法, 探讨了该区域干热河谷典型植物群落的物种beta多样性和谱系beta多样性与样方间环境差异(主要是气候)及地理距离之间的关系。结果表明: (1)群落间的地理距离和年平均温度差异对干热河谷植物群落的物种beta多样性和谱系beta多样性有显著影响; (2)地理距离对物种beta多样性和MNTD的影响最大; 地理距离和年平均温度差异对MPD的影响均较大; (3)样方间年平均温度与年平均降水量的差异和地理距离能够解释群落间beta多样性及谱系beta多样性11-13%的变异。以上结果表明, 生态位分化和扩散限制对该地区植物群落的beta多样性均有显著影响, 其中扩散限制的影响可能更大。此外, 人类活动等其他因素也很可能对元江干热河谷的群落组成具有非常重要的影响。     

Invertebrate Metacommunity Structure and Dynamics in an Andean Glacial Stream Network Facing Climate Change

Under the ongoing climate change, understanding the mechanisms structuring the spatial distribution of aquatic species in glacial stream networks is of critical importance to predict the response of aquatic biodiversity in the face of glacier melting. In this study, we propose to use metacommunity theory as a conceptual framework to better understand how river network structure influences the spatial organization of aquatic communities in glacierized catchments. At 51 stream sites in an Andean glacierized catchment (Ecuador), we sampled benthic macroinvertebrates, measured physico-chemical and food resource conditions, and calculated geographical, altitudinal and glaciality distances among all sites. Using partial redundancy analysis, we partitioned community variation to evaluate the relative strength of environmental conditions (e.g., glaciality, food resource) vs. spatial processes (e.g., overland, watercourse, and downstream directional dispersal) in organizing the aquatic metacommunity. Results revealed that both environmental and spatial variables significantly explained community variation among sites. Among all environmental variables, the glacial influence component best explained community variation. Overland spatial variables based on geographical and altitudinal distances significantly affected community variation. Watercourse spatial variables based on glaciality distances had a unique significant effect on community variation. Within alpine catchment, glacial meltwater affects macroinvertebrate metacommunity structure in many ways. Indeed, the harsh environmental conditions characterizing glacial influence not only constitute the primary environmental filter but also, limit water-borne macroinvertebrate dispersal. Therefore, glacier runoff acts as an aquatic dispersal barrier, isolating species in headwater streams, and preventing non-adapted species to colonize throughout the entire stream network. Under a scenario of glacier runoff decrease, we expect a reduction in both environmental filtering and dispersal limitation, inducing a taxonomic homogenization of the aquatic fauna in glacierized catchments as well as the extinction of specialized species in headwater groundwater and glacier-fed streams, and consequently an irreversible reduction in regional diversity.     

Congruence,but no cascade—Pelagic biodiversity across three trophic levels in Nordic lakes

Covariation in species richness and community structure across taxonomical groups (cross‐taxon congruence) has practical consequences for the identification of biodiversity surrogates and proxies, as well as theoretical ramifications for understanding the mechanisms maintaining and sustaining biodiversity. We found there to exist a high cross‐taxon congruence between phytoplankton, zooplankton, and fish in 73 large Scandinavian lakes across a 750 km longitudinal transect. The fraction of the total diversity variation explained by local environment alone was small for all trophic levels while a substantial fraction could be explained by spatial gradient variables. Almost half of the explained variation could not be resolved between local and spatial factors, possibly due to confounding issues between longitude and landscape productivity. There is strong consensus that the longitudinal gradient found in the regional fish community results from postglacial dispersal limitations, while there is much less evidence for the species richness and community structure gradients at lower trophic levels being directly affected by dispersal limitation over the same time scale. We found strong support for bidirectional interactions between fish and zooplankton species richness, while corresponding interactions between phytoplankton and zooplankton richness were much weaker. Both the weakening of the linkage at lower trophic levels and the bidirectional nature of the interaction indicates that the underlying mechanism must be qualitatively different from a trophic cascade.