Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity–ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m⁻² and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m⁻². One year after seed addition, local plant species richness had increased on average by six species m⁻² (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m⁻² (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump‐shaped model appears to be the limiting outline of the natural diversity–productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.     

Community patterns in source-sink metacommunities

We present a model of a source-sink competitive metacommunity, defined as a regional set of communities in which local diversity is maintained by dispersal. Although the conditions of local and regional coexistence have been well defined in such systems, no study has attempted to provide clear predictions of classical community-wide patterns. Here we provide predictions for species richness, species relative abundances, and community-level functional properties (productivity and space occupation) at the local and regional scales as functions of the proportion of dispersal between communities. Local (alpha) diversity is maximal at an intermediate level of dispersal, whereas between-community (beta) and regional (gamma) diversity decline as dispersal increases because of increased homogenization of the metacommunity. The relationships between local and regional species richness and the species rank abundance distributions are strongly affected by the level of dispersal. Local productivity and space occupation tend to decline as dispersal increases, resulting in either a hump-shaped or a positive relationship between species richness and productivity, depending on the scale considered (local or regional). These effects of dispersal are buffered by decreasing species dispersal success. Our results provide a niche-based alternative to the recent neutral-metacommunity model and have important implications for conservation biology and landscape management.     

Ecosystem productivity is associated with bacterial phylogenetic distance in surface marine waters

Understanding the link between community diversity and ecosystem function is a fundamental aspect of ecology. Systematic losses in biodiversity are widely acknowledged but the impact this may exert on ecosystem functioning remains ambiguous. There is growing evidence of a positive relationship between species richness and ecosystem productivity for terrestrial macro‐organisms, but similar links for marine micro‐organisms, which help drive global climate, are unclear. Community manipulation experiments show both positive and negative relationships for microbes. These previous studies rely, however, on artificial communities and any links between the full diversity of active bacterial communities in the environment, their phylogenetic relatedness and ecosystem function remain hitherto unexplored. Here, we test the hypothesis that productivity is associated with diversity in the metabolically active fraction of microbial communities. We show in natural assemblages of active bacteria that communities containing more distantly related members were associated with higher bacterial production. The positive phylogenetic diversity–productivity relationship was independent of community diversity calculated as the Shannon index. From our long‐term (7‐year) survey of surface marine bacterial communities, we also found that similarly, productive communities had greater phylogenetic similarity to each other, further suggesting that the traits of active bacteria are an important predictor of ecosystem productivity. Our findings demonstrate that the evolutionary history of the active fraction of a microbial community is critical for understanding their role in ecosystem functioning.     

Connectivity, scale-dependence, and the productivity–diversity relationship

We surveyed freshwater ponds (localities) nested within watersheds (regions) to evaluate the relationship between productivity and animal species richness at different spatial scales. In watersheds where the ponds were relatively distant from one another (likely reducing the level of interpond dispersal of many organisms), we found a scale‐dependent productivity–diversity relationship; at local scales (among ponds), diversity was a hump‐shaped function of productivity, whereas at regional scales (among watersheds), diversity monotonically increased with productivity. Furthermore, this relationship emerged because there was a strong relationship between productivity and pond‐to‐pond species compositional differences. Alternatively, in watersheds where ponds were relatively close together (likely leading to higher rates of dispersal of many organisms), we found no scale‐dependence; diversity was a hump‐shaped function of productivity at both local and regional scales. Here, the relationship between species compositional dissimilarity and productivity was much weaker. We conclude that whether or not scale‐dependence is observed in productivity–diversity relationships will depend, at least in part, on the degree of connectivity among localities within regions.     

New insights into relationships between active and dormant organisms,phylogenetic diversity and ecosystem productivity

Marine microbes make up a key part of ocean food webs and drive ocean chemistry through a range of metabolic processes. A fundamental question in ecology is whether the diversity of organisms in a community shapes the ecological functions of that community. While there is substantial evidence to support a positive link between diversity and ecological productivity for macro‐organisms in terrestrial environments, this relationship has not previously been verified for marine microbial communities. One factor complicating the understanding of this relationship is that many marine microbes are dormant and are easily dispersed by ocean currents, making it difficult to ensure that the organisms found in a given environmental sample accurately reflect processes occurring in that environment. Another complication is that, due to microbes great range of genotypic and phenotypic variability, communities with distantly related species may have greater range of metabolic functions than communities have the same richness and evenness, but in which the species present are more closely related to each other. In this issue of Molecular Ecology, Galand et al. (2015) provide compelling evidence that the most metabolically active communities are those in which the nondormant portion of the microbial community has the highest phylogenetic diversity. They also illustrate that focusing on the active portion of the community allows for detection of temporal patterns in community structure that would not be otherwise evident. The authors’ point out that the presence of many dormant organisms that do not contribute to ecosystem functioning is a feature that makes microbial ecosystems fundamentally different from macro‐ecosystems and that this difference needs to be accounted for in microbial ecology theory.     

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.     

Dispersal and species diversity: a meta-analysis

Species diversity in communities of interacting organisms is thought to be enhanced by dispersal, yet mechanisms predicting this have little to say about what effects differing rates of dispersal have on diversity and how dispersal affects diversity at larger spatial scales. I performed meta‐analyses on 23 studies comprising 50 experiments that manipulated species migration and measured community richness or diversity to test three hypotheses: that dispersal increases local diversity; that this effect depends on the rate of dispersal, specifically, that local diversity should be maximized at intermediate dispersal rates or else linearly related to dispersal rate; and that regional diversity may be either unaffected or negatively impacted by dispersal because dispersal tends to homogenize local communities. I found that immigration increased local diversity. Further, in animal studies, diversity appears maximized at intermediate dispersal rates but not with plant studies; however, more standardized studies are needed. Finally, results are ambiguous as to what happens at larger scales, with studies finding either declines or no change in regional diversity with dispersal. Taken together, these results reveal that dispersal has a complex, spatially contingent relationship with patterns of species diversity.     

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.     

Dispersal enhances beta diversity in nectar microbes

Dispersal is considered a key driver of beta diversity, the variation in species composition among local communities, but empirical tests remain limited. We manipulated dispersal of nectar‐inhabiting bacteria and yeasts via flower‐visiting animals to examine how dispersal influenced microbial beta diversity among flowers. Contrary to the prevailing view that dispersal lowers beta diversity, we found beta diversity was highest when dispersal was least limited. Our analysis suggested that this unexpected pattern might have resulted from stronger priority effects under increased dispersal. Dispersal is highly stochastic, generating variability in species arrival history and consequently the potential for community divergence via priority effects, in these and likely many other microbial, plant, and animal communities. Yet most previous experiments eliminated this possibility. We suggest that the positive effects of dispersal on beta diversity, like the one we report here, may have been greatly underappreciated.     

The effects of species pool,dispersal and competition on the diversity–productivity relationship

Aim The diversity–productivity relationship is a controversial issue in ecology. Diversity is sometimes seen to increase with productivity but a unimodal relationship has often been reported. Competitive exclusion was cited initially to account for the decrease of diversity at high productivity. Subsequently, the roles of evolutionary history (species pool size) and dispersal rate have been acknowledged. We explore how the effects of species pool, dispersal and competition combine to produce different diversity–productivity relationships. Methods We use a series of simulations with a spatially explicit, individual‐based model. Following empirical expectations, we used four scenarios to characterize species pool size along the productivity gradient (uniformly low and high, linear increase and unimodal). Similarly, the dispersal rate varied along the productivity gradient (uniformly low and high, and unimodal). We considered both neutral communities and communities with competitive exclusion. Results and main conclusions Our model predicts that competitive interactions will result in unimodal diversity–productivity relationships. The model often predicts unimodal patterns in neutral communities as well, although the decline in richness at high productivity is less than in competing communities. A positive diversity–productivity relationship is simulated for neutral communities when the species pool size increases with productivity and the dispersal rate is high. This scenario is probably more widespread in nature than the others since positive diversity–productivity relationships have been observed more frequently than previously expected, especially in the tropics and for woody species. Our simulated effects of species pool, dispersal and competition on diversity patterns can be linked to empirical observations to uncover mechanisms behind the diversity–productivity relationship.     

Assembly mechanisms determining high species turnover in aquatic communities over regional and continental scales

Niche and neutral processes drive community assembly and metacommunity dynamics, but their relative importance might vary with the spatial scale. The contribution of niche processes is generally expected to increase with increasing spatial extent at a higher rate than that of neutral processes. However, the extent to what community composition is limited by dispersal (usually considered a neutral process) over increasing spatial scales might depend on the dispersal capacity of composing species. To investigate the mechanisms underlying the distribution and diversity of species known to have great powers of dispersal (hundreds of kilometres), we analysed the relative importance of niche processes and dispersal limitation in determining beta‐diversity patterns of aquatic plants and cladocerans over regional (up to 300 km) and continental (up to 3300 km) scales. Both taxonomic groups were surveyed in five different European regions and presented extremely high levels of beta‐diversity, both within and among regions. High beta‐diversity was primarily explained by species replacement (turnover) rather than differences in species richness (i.e. nestedness). Abiotic and biotic variables were the main drivers of community composition. Within some regions, small‐scale connectivity and the spatial configuration of sampled communities explained a significant, though smaller, fraction of compositional variation, particularly for aquatic plants. At continental scale (among regions), a significant fraction of compositional variation was explained by a combination of spatial effects (exclusive contribution of regions) and regionally‐structured environmental variables. Our results suggest that, although dispersal limitation might affect species composition in some regions, aquatic plant and cladoceran communities are not generally limited by dispersal at the regional scale (up to 300 km). Species sorting mediated by environmental variation might explain the high species turnover of aquatic plants and cladocerans at regional scale, while biogeographic processes enhanced by dispersal limitation among regions might determine the composition of regional biotas.     

Regional species pools control community saturation in lake phytoplankton

Recent research has highlighted that positive biodiversity–ecosystem functioning relationships hold for all groups of organisms, including microbes. Yet, we still lack understanding regarding the drivers of microbial diversity, in particular, whether diversity of microbial communities is a matter of local factors, or whether metacommunities are of similar importance to what is known from higher organisms. Here, we explore the driving forces behind spatial variability in lake phytoplankton diversity in Fennoscandia. While phytoplankton biovolume is best predicted by local phosphorus concentrations, phytoplankton diversity (measured as genus richness, G) only showed weak correlations with local concentrations of total phosphorus. By estimating spatial averages of total phosphorus concentrations on various scales from an independent, spatially representative lake survey, we found that close to 70 per cent of the variability in local phytoplankton diversity can be explained by regionally averaged phosphorus concentrations on a scale between 100 and 400 km. Thus, the data strongly indicate the existence of metacommunities on this scale. Furthermore, we show a strong dependency between lake productivity and spatial community turnover. Thus, regional productivity affects beta-diversity by controlling spatial community turnover, resulting in scale-dependent productivity-diversity relationships. As an illustration of the interaction between local and regional processes in shaping microbial diversity, our results offer both empirical support and a plausible mechanism for the existence of common scaling rules in both the macrobial and the microbial worlds. We argue that awareness of regional species pools in phytoplankton and other unicellular organisms may critically improve our understanding of ecosystems and their susceptibility to anthropogenic stressors.     

Dispersal rates affect species composition in metacommunities of Sarracenia purpurea inquilines

Dispersal among local communities can have a variety of effects on species composition and diversity at local and regional scales. Local conditions (e.g., resource and predator densities) can have independent effects, as well as interact with dispersal, to alter these patterns. Based on metacommunity models, we predicted that local diversity would show a unimodal relationship with dispersal frequency. We manipulated dispersal frequencies, resource levels, and the presence of predators (mosquito larvae) among communities found in the water-filled leaves of the pitcher plant Sarracenia purpurea. Diversity and abundance of species of the middle trophic level, protozoa and rotifers, were measured. Increased dispersal frequencies significantly increased regional species richness and protozoan abundance while decreasing the variance among local communities. Dispersal frequency interacted with predation at the local community scale to produce patterns of diversity consistent with the model. When predators were absent, we found a unimodal relationship between dispersal frequency and diversity, and when predators were present, there was a flat relationship. Intermediate dispersal frequencies maintained some species in the inquiline communities by offsetting extinction rates. Local community composition and the degree of connectivity between communities are both important for understanding species diversity patterns at local and regional scales.     

Salmon‐derived nutrients drive diatom beta‐diversity patterns

1. Pacific salmon are a textbook example of migratory animals that transfer nutrients between ecosystems, but little is known about how salmon‐derived nutrients (SDN) affect the biodiversity of recipient freshwater ecosystems. We examined paleolimnological records from six Alaskan lakes to define how changes in SDN from sockeye salmon (Oncorhynchus nerka) influenced sedimentary diatom community structure and beta‐diversity among lakes and through time. 2. Using an isotopic mixing model, we showed that SDN loading could account for >80% of the lake total nitrogen budgets and strongly regulated diatom community composition. Spatial dissimilarity in diatom communities was positively related to differences in SDN among lakes (r² = 0.69, P < 0.01, n = 10). Likewise, temporal dissimilarity in diatom communities was positively related to differences in SDN in a sediment core with substantial variation in salmon spawner dynamics between 1700 and 1950 AD (r² = 0.34, P < 0.01, n = 19). Finally, beta‐diversity metrics quantifying temporal turnover within each lake’s sediment record were also positively related to the variance in SDN loading among lakes (r² = 0.88, P < 0.05, n = 5). Mean SDN was only negatively correlated to temporal diatom beta‐diversity. 3. Spatially subsidised systems often receive temporally variable resource inputs, and thus, it is not surprising that, unlike previous studies, we found that resource variability was the key driver of community composition and beta‐diversity. In habitats that receive strongly fluctuating external nutrient loads, environment heterogeneity may overweigh stochastic community processes. In addition, freshwater diatoms are characterised by great dispersal capabilities and short life cycles and therefore may be a more sensitive indicator for evaluating the role of resource variability than previously used model organisms. These results suggest that productivity–diversity relationship vary with the nature of nutrient loading and the life history of the community studied. 4. Overall, our study highlights that the transport of nutrients by sockeye salmon across ecosystem boundaries is a significant driver of algal community and biodiversity in nursery lakes, mainly through changing the magnitude of nutrient variation. As such, freshwater species diversity in regions like the U.S. Pacific Northwest may become impoverished where there have been long‐term declines in salmon populations and decreases in nutrient variability among lakes.     

Regional effects as important determinants of local diversity in both marine and terrestrial systems

Community ecologists have struggled to create unified theories across diverse ecosystems, but it has been difficult to acertain whether marine and terrestrial communities differ in the mechanisms responsible for structure and dynamics. One apparent difference between marine and terrestrial ecology is that the influence of regional processes on local populations and communities is better established in the marine literature. We examine three potential explanations: 1) influential early studies emphasized local interactions in terrestrial communities and regional dispersal in marine communities. 2) regional‐scale processes are actually more important in marine than in terrestrial communities. 3) recruitment from a regional species pool is easier to study in marine than terrestrial communities. We conclude that these are interrelated, but that the second and especially the third explanations are more important than the first. We also conclude that in both marine and terrestrial systems, there are ways to improve our understanding of regional influences on local community diversity. In particular, we advocate examining local vs regional diversity relationships at localities within environmentally similar regions that differ in their diversity either because of their sizes or their varying degrees of isolation from a species source.     

Combined effects of zooplankton grazing and dispersal on the diversity and assembly mechanisms of bacterial metacommunities

Effects of dispersal and the presence of predators on diversity, assembly and functioning of bacterial communities are well studied in isolation. In reality, however, dispersal and trophic interactions act simultaneously and can therefore have combined effects, which are poorly investigated. We performed an experiment with aquatic metacommunities consisting of three environmentally different patches and manipulated dispersal rates among them as well as the presence or absence of the keystone species Daphnia magna. Daphnia magna reduced both local and regional diversity, whereas dispersal increased local diversity but decreased beta‐diversity having no net effect on regional diversity. Dispersal modified the assembly mechanisms of bacterial communities by increasing the degree of determinism. Additionally, the combination of the D. magna and dispersal increased the importance of deterministic processes, presumably because predator‐tolerant taxa were spread in the metacommunity via dispersal. Moreover, the presence of D. magna affected community composition, increased community respiration rates but did not affect bacterial production or abundance, whereas dispersal slightly increased bacterial production. In conclusion, our study suggests that predation by a keystone species such as D. magna and dispersal additively influence bacterial diversity, assembly processes and ecosystem functioning.     

A heatwave increases turnover and regional dominance in microbenthic metacommunities

While the effect of the global biodiversity crisis on local species loss is still debated, there is empirical evidence for major changes in local biodiversity attributed to increased species turnover. In communities exposed to a climate stressor, species turnover can lead to increased dominance of well-adapted species and consequently to an overall decline in species diversity. Despite the known importance of species turnover for community dynamics and functioning, experimental results on the connection between biodiversity loss and species turnover are scarce. We still do not fully understand which specific factors increase the rate of change in species composition, especially when considering natural compared to artificially lab assembled communities. In the present study, we experimentally tested whether a heatwave and dispersal increased species turnover and decreased species diversity in natural benthic diatom communities with different initial species compositions. We found that on the local scale, dispersal had overall positive effects on species richness while the relationship between exposure to the heatwave, species turnover, and diversity depended on initial community composition. However, on the regional (i.e. metacommunity) scale, exposure to the heatwave and dispersal both increased turnover and decreased Shannon diversity by almost 50%. Turnover in these metacommunities was not caused by a loss of species, but rather by a change in dominance patterns leading to homogenization, and consequently decreased diversity. Our study shows that climate change can destabilize community composition and degrade species diversity, but still after ca. 15 generations does not decrease the number of species in the community, demonstrating that the response of species diversity and richness to changing conditions can be fundamentally decoupled on ecological time scales.     

Reducing dispersal limitation via seed addition increases species richness but not above‐ground biomass

Seed dispersal limitation, which can be exacerbated by a number of anthropogenic causes, can result in local communities having fewer species than they might potentially support, representing a potential diversity deficit. The link between processes that shape natural variation in diversity, such as dispersal limitation, and the consequent effects on productivity is less well known. Here, we synthesised data from 12 seed addition experiments in grassland communities to examine the influence of reducing seed dispersal limitation (from 1 to 60 species added across experiments) on species richness and productivity. For every 10 species of seed added, we found that species richness increased by about two species. However, the increase in species richness by overcoming seed limitation did not lead to a concomitant increase in above‐ground biomass production. This highlights the need to consider the relationship between biodiversity and ecosystem functioning in a pluralistic way that considers both the processes that shape diversity and productivity simultaneously in naturally assembled communities.     

Modularity along organism dispersal gradients challenges a prevailing view of abrupt transitions in animal landscape perception

A common property of landscapes and metacommunities is the occurrence of abrupt shifts in connectivity along gradients of individual dispersal abilities. Animals with short‐range dispersal capability perceive fragmented landscapes, but organisms moving across critical thresholds perceive continuous landscapes. This qualitative shift in landscape perception may determine several attributes of local communities and the dynamics of whole metacommunities. Modularity describes the existence in some communities of relatively high numbers of mutual connections favoring the movement of neighboring individuals (even when each individual is able to reach any patch in the landscape). Local patch linkages and metacommunity connectivity along gradients of dispersal ability have been reported frequently. However, the intermediate level of structure captured by modularity has not been considered. We evaluated landscape connectivity and modularity along gradients of individual dispersal abilities. Random landscapes with different degrees of cell aggregation and occupancy were simulated; we also analyzed ten real ecosystems. An expected, a shift in landscape connectivity was always detected; modularity consistently decreased gradually along dispersal gradients in both simulated networks and empirical landscapes. Neutral metacommunities within simulated landscapes demonstrated that modularity and connectivity may reflect landscape traits in the shaping of metacommunity diversity. Average beta‐diversity was strongly associated with modularity, particularly with low migration rates, while connectivity trends tracked changes in beta‐diversity at intermediate to high migrations rates. Consequently, while some species are able to perceive abrupt transitions in the landscape, many others probably experience a gradual continuum in landscape perception, contrary to predictions from previous analyses. Furthermore, the gradual behavior of modularity indicates that it may represent an exceptional early‐warning tool that measures system distance to tipping points. Our study highlights the multiple perceptions that different species may have of a single landscape and shows, for the first time, a theoretical and empirical relationship between landscape modularity, and metacommunity diversity.