Effects of diversity on diversity: consequences of competition and facilitation

Diversity in one group of species or genotypes is often correlated with diversity in a second group – prominent examples including native vs exotic species, and genetic diversity in a focal species vs species diversity in the rest of the community. I used simulation models to investigate the roles of competition and facilitation among species or genotypes in creating diversity–diversity relationships, with a focus on facilitation, which has received little theoretical attention. When competitive interactions dominate, increasing diversity in one group reduces diversity in the second group via filling of available niche space. Facilitation can create positive diversity–diversity relationships via a sampling effect, whereby a strong facilitator of the second group is more likely to be present as diversity increases in the first group, and also via one group acting as a source of biotic heterogeneity (i.e. diversifying selection) on the second group. However, the biotic heterogeneity effect is expected only under restricted conditions – with asymmetric facilitation, only during a transient period, or only over a small range of species diversity levels – and therefore seems unlikely to operate within trophic levels in natural communities. More generally, the simultaneous operation of competition and facilitation results in several different diversity–diversity relationships and underlying mechanisms. The results clarify the potential roles of positive and negative interactions in creating diversity–diversity relationships, and in determining the outcome of community dynamics in general. This study also highlights some important difficulties in incorporating facilitation into ecological theory for communities with many species.     

Genetic diversity and species diversity of stream fishes covary across a land-use gradient

Genetic diversity and species diversity are expected to covary according to area and isolation, but may not always covary with environmental heterogeneity. In this study, we examined how patterns of genetic and species diversity in stream fishes correspond to local and regional environmental conditions. To do so, we compared population size, genetic diversity and divergence in central stonerollers (Campostoma anomalum) to measures of species diversity and turnover in stream fish assemblages among similarly sized watersheds across an agriculture–forest land-use gradient in the Little Miami River basin (Ohio, USA). Significant correlations were found in many, but not all, pair-wise comparisons. Allelic richness and species richness were strongly correlated, for example, but diversity measures based on allele frequencies and assemblage structure were not. In-stream conditions related to agricultural land use were identified as significant predictors of genetic diversity and species diversity. Comparisons to population size indicate, however, that genetic diversity and species diversity are not necessarily independent and that variation also corresponds to watershed location and glaciation history in the drainage basin. Our findings demonstrate that genetic diversity and species diversity can covary in stream fish assemblages, and illustrate the potential importance of scaling observations to capture responses to hierarchical environmental variation. More comparisons according to life history variation could further improve understanding of conditions that give rise to parallel variation in genetic diversity and species diversity, which in turn could improve diagnosis of anthropogenic influences on aquatic ecosystems.     

On the relationship between plant species diversity and genetic diversity of Plantago lanceolata (Plantaginaceae) within and between grassland communities

Aims and Methods The relationship between genetic diversity and species diversity and the underlying mechanisms are of both fundamental and applied interest. We used amplified fragment length polymorphism (AFLP) and vegetation records to investigate the association between genetic diversity of Plantago lanceolata and plant species diversity using 15 grassland communities in central Germany. We used correlation and partial correlation analyses to examine whether relationships between genetic and species diversity were direct or mediated by environmental differences between habitats.Important findings Both within- and between-population genetic diversity of P. lanceolata were significantly positively correlated with plant species diversity within and between sites. Simple and partial correlations revealed that the positive correlations indirectly resulted from the effects of abiotic habitat characteristics on plant species diversity and, via abundance, on genetic diversity of P. lanceolata. Thus, they did not reflect a direct causal relationship between plant species diversity and genetic diversity of P. lanceolata, as would have been expected based on the hypothesis of a positive relationship between plant species diversity and niche diversity.     

Lessons from the fish: a multi‐species analysis reveals common processes underlying similar species‐genetic diversity correlations

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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.     

Connections between species diversity and genetic diversity

Species diversity and genetic diversity remain the nearly exclusive domains of community ecology and population genetics, respectively, despite repeated recognition in the literature over the past 30 years of close parallels between these two levels of diversity. Species diversity within communities and genetic diversity within populations are hypothesized to co‐vary in space or time because of locality characteristics that influence the two levels of diversity via parallel processes, or because of direct effects of one level of diversity on the other via several different mechanisms. Here, we draw on a wide range of studies in ecology and evolution to examine the theoretical underpinnings of these hypotheses, review relevant empirical literature, and outline an agenda for future research. The plausibility of species diversity–genetic diversity relationships is supported by a variety of theoretical and empirical studies, and several recent studies provide direct, though preliminary support. Focusing on potential connections between species diversity and genetic diversity complements other approaches to synthesis at the ecology–evolution interface, and should contribute to conceptual unification of biodiversity research at the levels of genes and species.     

Environmental heterogeneity,species diversity and co‐existence at different spatial scales

The positive relationship between spatial environmental heterogeneity and species diversity is a widely accepted concept, generally associated with niche limitation. However, niche limitation cannot account for negative heterogeneity–diversity relationships (HDR) revealed in several case studies. Here we explore how HDR varies at different spatial scales and provide novel theories for small‐scale species co‐existence that explain both positive and negative HDR. At large spatial scales of heterogeneity (e.g. landscape level), different communities co‐exist, promoting large regional species pool size and resulting in positive HDR. At smaller scales within communities, species co‐existence can be enhanced by increasing the number of different patches, as predicted by the niche limitation theory, or alternatively, restrained by heterogeneity. We conducted meta‐regressions for experimental and observational HDR studies, and found that negative HDRs are significantly more common at smaller spatial scales. We propose three theories to account for niche limitation at small spatial scales. (1) Microfragmentation theory: with increasing spatial heterogeneity, large homogeneous patches lose area and become isolated, which in turn restrains the establishment of new plant individuals and populations, thus reducing species richness. (2) Heterogeneity confounded by mean: when heterogeneity occurs at spatial scales smaller than the size of individual plants, which forage through the patches, species diversity can be either positively or negatively affected by a change in the mean of an environmental factor. (3) Heterogeneity as a separate niche axis: the ability of species to tolerate heterogeneity at spatial scales smaller than plant size varies, affecting HDR. We conclude that processes other than niche limitation can affect the relationship between heterogeneity and diversity.     

Plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses

Questions: What is the observed relationship between plant species diversity and spatial environmental heterogeneity? Does the relationship scale predictably with sample plot size? What are the relative contributions to diversity patterns of variables linked to productivity or available energy compared to those corresponding to spatial heterogeneity? Methods: Observational and experimental studies that quantified relationships between plant species richness and within‐sample spatial environmental heterogeneity were reviewed. Effect size in experimental studies was quantified as the standardized mean difference between control (homogeneous) and heterogeneous treatments. For observational studies, effect sizes in individual studies were examined graphically across a gradient of plot size (focal scale). Relative contributions of variables representing spatial heterogeneity were compared to those representing available energy using a response ratio. Results: Forty‐one observational and 11 experimental studies quantified plant species diversity and spatial environmental heterogeneity. Observational studies reported positive species diversity‐spatial heterogeneity correlations at all points across a plot size gradient from ～1.0 × 10^?1 to ～1.0 × 10¹¹ m², although many studies reported spatial heterogeneity variables with no significant relationships to species diversity. The cross‐study effect size in experimental studies was not significantly different from zero. Available energy variables explained consistently more of the variance in species richness than spatial heterogeneity variables, especially at the smallest and largest plot sizes. Main conclusions: Species diversity was not related to spatial heterogeneity in a way predictable by plot size. Positive heterogeneity‐diversity relationships were common, confirming the importance of niche differentiation in species diversity patterns, but future studies examining a range of spatial scales in the same system are required to determine the role of dispersal and available energy in these patterns.     

Genetic variation of loci potentially under selection confounds species–genetic diversity correlations in a fragmented habitat

Positive species–genetic diversity correlations (SGDCs) are often thought to result from the parallel influence of neutral processes on genetic and species diversity. Yet, confounding effects of non‐neutral mechanisms have not been explored. Here, we investigate the impact of non‐neutral genetic diversity on SGDCs in high Andean wetlands. We compare correlations between plant species diversity and genetic diversity (GD) calculated with and without loci potentially under selection (outlier loci). The study system includes 2188 specimens from five species (three common aquatic macroinvertebrate and two dominant plant species) that were genotyped for 396 amplified fragment length polymorphism loci. We also appraise the importance of neutral processes on SGDCs by investigating the influence of habitat fragmentation features. Significant positive SGDCs were detected for all five species (mean SGDC = 0.52 ± 0.05). While only a few outlier loci were detected in each species, they resulted in significant decreases in GD and in SGDCs. This supports the hypothesis that neutral processes drive species–genetic diversity relationships in high Andean wetlands. Unexpectedly, the effects on genetic diversity GD of the habitat fragmentation characteristics in this study increased with the presence of outlier loci in two species. Overall, our results reveal pitfalls in using habitat features to infer processes driving SGDCs and show that a few loci potentially under selection are enough to cause a significant downward bias in SGDC. Investigating confounding effects of outlier loci thus represents a useful approach to evidence the contribution of neutral processes on species–genetic diversity relationships.     

Relationships between spatial environmental heterogeneity and plant species diversity on a limestone pavement

No empirical studies have examined the relationship between diversity and spatial heterogeneity across unimodal species richness gradients. We determined the relationships between diversity and environmental factors for 144 0.18 m² plots in a limestone pavement alvar in southern Ontario, Canada, including within-plot spatial heterogeneity in soil depth, microtopography and microsite composition. Species richness was unimodally related to mean soil depth and relative elevation. Microsite heterogeneity and soil depth heterogeneity were positively correlated with species richness, and the richness peaks of the unimodal gradients correspond to the maximally spatially heterogeneous plots. The best predictive models of species richness and evenness, however, showed that other factors, such as ramet density and flooding, are the major determinants of diversity in this system. The findings that soil depth heterogeneity had effects on diversity when the effects of mean soil depth were factored out, and that unimodal richness peaks were associated with high spatial heterogeneity in environmental factors represent significant contributions to our understanding of how spatial heterogeneity might contribute to diversity maintenance in plant communities.     

Riverine fish diversity varies according to geographical isolation and land use modification

Understanding the environmental factors driving species‐genetic diversity correlations (SGDCs) is critical for designing appropriate conservation and management strategies to protect biodiversity. Yet, few studies have explored the impact of changing land use patterns on SGDCs specifically in aquatic communities. This study examined patterns of genetic diversity in roach (Rutilus rutilus L.) together with fish species composition across 19 locations in a large river catchment, spanning a gradient in land use. Our findings show significant correlations between some, but not all, species and genetic diversity end points. For example, genetic and species differentiation showed a weak but significant linear relationship across the Thames catchment, but additional diversity measures such as allelic richness and fish population abundance did not. Further examination of patterns in species and genetic diversity indicated that land use intensification has a modest effect on fish diversity compared to the combined influence of geographical isolation and land use intensification. These results indicate that environmental changes in riparian habitats have the potential to amplify shifts in the composition of stream fish communities in poorly connected river stretches. Conservation and management strategies for fish populations should, therefore, focus on enhancing connectivity between river stretches and limit conversion of nearby land to arable or urban use to maintain current levels of biodiversity.     

Dispersal and neutral sampling mediate contingent effects of disturbance on plant beta‐diversity: a meta‐analysis

A major challenge in ecology, conservation and global‐change biology is to understand why biodiversity responds differently to similar environmental changes. Contingent biodiversity responses may depend on how disturbance and dispersal interact to alter variation in community composition (β‐diversity) and assembly mechanisms. However, quantitative syntheses of these patterns and processes across studies are lacking. Using null‐models and meta‐analyses of 22 factorial experiments in herbaceous plant communities across Europe and North America, we show that disturbance diversifies communities when dispersal is limited, but homogenises communities when combined with increased immigration from the species pool. In contrast to the hypothesis that disturbance and dispersal mediate the strength of niche assembly, both processes altered β‐diversity through neutral‐sampling effects on numbers of individuals and species in communities. Our synthesis suggests that stochastic effects of disturbance and dispersal on community assembly play an important, but underappreciated, role in mediating biotic homogenisation and biodiversity responses to environmental change.     

A truce with neutral theory: local deterministic factors, species traits and dispersal limitation together determine patterns of diversity in stream invertebrates

1. Studies seeking to explain local patterns of diversity have typically relied on niche explanations, reflected in correlations with local environmental conditions, or neutral theory, invoking dispersal processes and speciation. 2. We used macroinvertebrate community data from 10 streams that varied independently in local ecological conditions and spatial proximity. Neutral theory predicts that similarity in communities will be negatively associated with distance between sites, while niche theory suggests that community similarity will be positively associated with similarity in local ecological conditions. 3. Similarity in total invertebrate, grazer and predator assemblages showed negative relationships with distance and, for grazers and predators, positive relationships with local ecological conditions. However, the best model predicting community similarity in all three cases included aspects of both local ecological conditions and distance between sites. 4. When assemblages were analysed according to dispersal ability, high-dispersal species were shown to be freely accessing all sites and community similarity was not well predicted by either local ecology or spatial separation. Assemblages of species with low and moderate dispersal ability were best predicted by combined models, including distance between sites and local ecological factors. 5. The results suggest that the perceived dichotomy between neutral and local environmental processes in determining local patterns of diversity may not be useful. Neutral and niche processes structured these communities differentially depending on trophic level and species traits. 6. We emphasize the potential for both dispersal processes and local environmental conditions to explain local patterns of diversity.     

Variation in habitat connectivity generates positive correlations between species and genetic diversity in a metacommunity

An increasing number of studies are simultaneously investigating species diversity (SD) and genetic diversity (GD) in the same systems, looking for ‘species– genetic diversity correlations’ (SGDCs). From negative to positive SGDCs have been reported, but studies have generally not quantified the processes underlying these correlations. They were also mostly conducted at large biogeographical scales or in recently degraded habitats. Such correlations have not been looked for in natural networks of connected habitat fragments (metacommunities), and the underlying processes remain elusive in most systems. We investigated these issues by studying freshwater snails in a pond network in Guadeloupe (Lesser Antilles). We recorded SD and habitat characteristics in 232 ponds and assessed GD in 75 populations of two species. Strongly significant and positive SGDCs were detected in both species. Based on a decomposition of SGDC as a function of variance–covariance of habitat characteristics, we showed that connectivity (opportunity of water flow between a site and the nearest watershed during the rainy season) has the strongest contribution on SGDCs. More connective sites received both more alleles and more species through immigration resulting in both higher GD and higher SD. Other habitat characteristics did not contribute, or contributed negatively, to SGDCs. This is true of the desiccation frequency of ponds during the dry season, presumably because species markedly differ in their ability to tolerate desiccation. Our study shows that variation in environmental characteristics of habitat patches can promote SGDCs at metacommunity scale when the studied species respond homogeneously to these environmental characteristics.     

Phylogenetic structure illuminates the mechanistic role of environmental heterogeneity in community organization

1. Diversity begets diversity. Numerous published positive correlations between environmental heterogeneity and species diversity indicate ubiquity of this phenomenon. Nonetheless, most assessments of this relationship are phenomenological and provide little insight into the mechanism whereby such positive association results. 2. Two unresolved issues could better illuminate the mechanistic basis to diversity begets diversity. First, as environmental heterogeneity increases, both productivity and the species richness that contributes to that productivity often increase in a correlated fashion thus obscuring the primary driver. Second, it is unclear how species are added to communities as diversity increases and whether additions are trait based. 3. We examined these issues based on 31 rodent communities in the central Mojave Desert. At each site, we estimated rodent species richness and characterized environmental heterogeneity from the perspectives of standing primary productivity and number of seed resources. We further examined the phylogenetic structure of communities by estimating the mean phylogenetic distance (MPD) among species and by comparing empirical phylogenetic distances to those based on random assembly from a regional species pool. 4. The relationship between rodent species diversity and environmental heterogeneity was positive and significant. Moreover, diversity of resources accounted for more unique variation than did total productivity, suggesting that variety and not total amount of resource was the driver of increased rodent diversity. Relationships between environmental heterogeneity and phylogenetic distance were negative and significant; species were significantly phylogenetically over-dispersed in communities of low environmental heterogeneity and became more clumped as environmental heterogeneity increased. 5. Results suggest that species diversity increases with environmental heterogeneity because a wider variety of resources allow greater species packing within communities.     

Limiting similarity and functional diversity along environmental gradients

Recent developments in community models emphasize the importance of incorporating stochastic processes (e.g. ecological drift) in models of niche‐structured community assembly. We constructed a finite, spatially explicit, lottery model to simulate the distribution of species in a one‐dimensional landscape with an underlying gradient in environmental conditions. Our framework combines the potential for ecological drift with environmentally‐mediated competition for space in a heterogeneous environment. We examined the influence of niche breadth, dispersal distances, community size (total number of individuals) and the breadth of the environmental gradient on levels of species and functional trait diversity (i.e. differences in niche optima). Three novel results emerge from this model: (1) niche differences between adjacent species (e.g. limiting similarity) increase in smaller communities, because of the interaction of competitive effects and finite population sizes; (2) immigration from a regional species pool, stochasticity and niche‐assembly generate a bimodal distribution of species residence times (‘transient’ and ‘resident’) under a heterogeneous environment; and (3) the magnitude of environmental heterogeneity has a U‐shaped effect on diversity, because of shifts in species richness of resident vs. transient species. These predictions illustrate the potential importance of stochastic (although not necessarily neutral) processes in community assembly.     

Species diversity in local neutral communities

We extend the neutral theory of macroecology by deriving biodiversity models (relative species abundance and species-area relationships) in a local community-metacommunity system in which the local community is embedded within the metacommunity. We first demonstrate that the local species diversity patterns converge to that of the metacommunity as the size (scale) of the embedded local community increases. This result shows that in continuous landscapes no sharp boundaries dividing the communities at the two scales exist; they are an artificial distinction made by the current spatially implicit neutral theory. Second, we remove the artificial restriction that speciation cannot occur in a local community, even if the effects of local speciation are small. Third, we introduce stochasticity into the immigration rate, previously treated as constant, and demonstrate that local species diversity is a function not only of the mean but also of the variance in immigration rate. High variance in immigration rates reduces species diversity in local communities. Finally, we show that a simple relationship exists between the fundamental diversity parameter of neutral theory and Simpson's index for local communities. Derivation of this relationship extends recent work on diversity indices and provides a means of evaluating the effect of immigration on estimates of the fundamental diversity parameter derived from relative species abundance data on local communities.     

How diversification rates and diversity limits combine to create large-scale species-area relationships

Species-area relationships (SARs) have mostly been treated from an ecological perspective, focusing on immigration, local extinction and resource-based limits to species coexistence. However, a full understanding across large regions is impossible without also considering speciation and global extinction. Rates of both speciation and extinction are known to be strongly affected by area and thus should contribute to spatial patterns of diversity. Here, we explore how variation in diversification rates and ecologically mediated diversity limits among regions of different sizes can result in the formation of SARs. We explain how this area-related variation in diversification can be caused by either the direct effects of area or the effects of factors that are highly correlated with area, such as habitat diversity and population size. We also review environmental, clade-specific and historical factors that affect diversification and diversity limits but are not highly correlated with region area, and thus are likely to cause scatter in observed SARs. We present new analyses using data on the distributions, ages and traits of mammalian species to illustrate these mechanisms; in doing so we provide an integrated perspective on the evolutionary processes shaping SARs.     

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.     

Environmental heterogeneity and the spatial structure of fern species diversity in one hectare of old-growth forest

The precise relationship between species diversity and spatial heterogeneity has not often been investigated using quantitative and repeatable measures of environmental variation. In this study, we map the metre-level distribution of fern species in one hectare of old-growth forest and test for a relationship between diversity and heterogeneity of physical features and soil conditions. The nineteen species recorded in the hectare were non-randomly distributed and varied greatly in abundance and spatial aggregation. Different. Species distributions were not independent of one another: three groups were formed with species which occurred together significantly more often than random expectation. Physical and soil conditions were highly variable and spatially auto correlated from the 5 m scale up to the extent of the whole hectare. Based on the sites where they grew, species differed in their references for soil moisture, fertility and pH. Fern diversity was highest at sites with high soil moisture and low soil fertility: however, there was no relationship between diversity and the environmental variance within quadrats. Unpredictable spatial distribution patterns produced by processes of dispersal and immigration may obscure any relationship between diversity and spatial heterogeneity at this fine scale.