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.     

Elevational patterns and hierarchical determinants of biodiversity across microbial taxonomic scales

Microbial biogeography is gaining increasing attention due to recent molecular methodological advance. However, the diversity patterns and their environmental determinants across taxonomic scales are still poorly studied. By sampling along an extensive elevational gradient in subarctic ponds of Finland and Norway, we examined the diversity patterns of aquatic bacteria and fungi from whole community to individual taxa across taxonomic coverage and taxonomic resolutions. We further quantified cross‐phylum congruence in multiple biodiversity metrics and evaluated the relative importance of climate, catchment and local pond variables as the hierarchical drivers of biodiversity across taxonomic scales. Bacterial community showed significantly decreasing elevational patterns in species richness and evenness, and U‐shaped patterns in local contribution to beta diversity (LCBD). Conversely, no significant species richness and evenness patterns were found for fungal community. Elevational patterns in species richness and LCBD, but not in evenness, were congruent across bacterial phyla. When narrowing down the taxonomic scope towards higher resolutions, bacterial diversity showed weaker and more complex elevational patterns. Taxonomic downscaling also indicated a notable change in the relative importance of biodiversity determinants with stronger local environmental filtering, but decreased importance of climatic variables. This suggested that niche conservatism of temperature preference was phylogenetically deeper than that of water chemistry variables. Our results provide novel perspectives for microbial biogeography and highlight the importance of taxonomic scale dependency and hierarchical drivers when modelling biodiversity and species distribution responses to future climatic scenarios.     

Evidence of stochasticity driving anuran metacommunity structure in the Pantanal wetlands

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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.     

Bird diversity patterns in Neotropical temperate farmlands: The role of environmental factors and trophic groups in the spring and autumn

Productivity, habitat heterogeneity and environmental similarity are of the most widely accepted hypotheses to explain spatial patterns of species richness and species composition similarity. Environmental factors may exhibit seasonal changes affecting species distributions. We explored possible changes in spatial patterns of bird species richness and species composition similarity. Feeding habits are likely to have a major influence in bird–environment associations and, given that food availability shows seasonal changes in temperate climates, we expect those associations to differ by trophic group (insectivores or granivores). We surveyed birds and estimated environmental variables along line‐transects covering an E‐W gradient of annual precipitation in the Pampas of Argentina during the autumn and the spring. We examined responses of bird species richness to spatial changes in habitat productivity and heterogeneity using regression analyses, and explored potential differences between seasons of those responses. Furthermore, we used Mantel tests to examine the relationship between species composition similarity and both the environmental similarity between sites and the geographic distance between sites, also assessing differences between seasons in those relationships. Richness of insectivorous birds was directly related to primary productivity in both seasons, whereas richness of seed‐eaters showed a positive association with habitat heterogeneity during the spring. Species composition similarity between assemblages was correlated with both productivity similarity and geographic proximity during the autumn and the spring, except for insectivore assemblages. Diversity within main trophic groups seemed to reflect differences in their spatial patterns as a response to changes between seasons in the spatial patterns of food resources. Our findings suggest that considering different seasons and functional groups in the analyses of diversity spatial pattern could contribute to better understand the determinants of biological diversity in temperate climates.     

Unravelling the correlates of species richness and ecological uniqueness in a metacommunity of urban pond insects

City ponds have the potential to harbour a rich biodiversity of aquatic insects despite being located in an urban landscape. However, our current knowledge on the correlates of pond biodiversity is limited and even less is known about the factors that influence the ecological uniqueness of urban ponds. The multiple environmental gradients, at different spatial scales, that may affect biodiversity and ecological uniqueness of urban ponds can thus be seen both as an opportunity and as a challenge for a study. In this study, we aimed to fill this gap by focusing on aquatic insect assemblages in 51 ponds in the Swedish city of Stockholm, using a metacommunity perspective. We found that species richness was primarily determined by the density of aquatic insects, water depth and proportion of buildings around the pond. The uniqueness of ponds was estimated as local contributions to beta diversity (LCBD), and it was primarily related to the proportion of arable land and industry around the ponds. With regard to the metacommunity we found two interesting patterns. First, there was a negative relationship between richness and LCBD. Second, biodiversity was spatially independent, suggesting that spatially-patterned dispersal did not structure species richness or LCBD. These last two patterns are important when considering conservation efforts of biodiversity in city ponds. We hence suggest that the conservation of insect biodiversity in urban pond should consider the surroundings of the ponds, and that high-richness ponds are not necessarily those that require most attention because they are not ecologically the most unique.     

Plant diversity,biogeography and environment in Iberia: Patterns and possible causal factors

Abstract. We associated patterns of plant diversity with possible causal factors by considering 93 local regions in the Iberian Peninsula and Balearic Islands with respect to biogeography, environmental favourability, and environmental heterogeneity, and their relationship with measured species diversity at four different scales: mean local species richness standardized at a grain of 100 m², total species richness in a community type within a region (regional community richness), mean compositional similarity, and mosaic diversity. Local regions in biogeographic transition zones to the North African and Atlantic floras had higher regional community richness and greater mosaic diversity than did non‐transitional regions, whereas no differences existed in mean local species richness or mean compositional similarity. Mean local species richness was positively related to environmental favourability as measured by actual evapotranspiration, but negatively related to total precipitation and temporal heterogeneity in precipitation. Mean local species richness was greatest in annual grassland and dwarf shrubland communities, and on calcareous bedrock types. Regional community richness was similarly related to actual evapotranspiration and total precipitation, but in addition was positively related to spatial heterogeneity in topography and soil water holding capacity. Mean compositional similarity decreased with increasing spatial heterogeneity and temperature seasonality. Mosaic diversity, a measure of complexity, increased with increasing local and regional richness. We hypothesize that these relationships can be explained by four ecological and evolutionary classes of causal factors: numbers of individuals, intermediate environments, limits to adaptation, and niche variation. These factors operate at various scales and manifest themselves in various ways. For example, at the site level, apparently processes that increase the number of individuals increase mean local species richness, but at the level of the entire region no such effects were found.     

A resource-based conceptual model of plant diversity that reassesses causality in the productivity–diversity relationship

Biogeographical studies frequently reveal positive correlations between species richness and estimates of environmental water and/or energy. A popular interpretation of this relationship relates the supply of water and energy to productivity, and then, in turn, to richness. Productivity–diversity theories are now legion, yet none has proved sufficiently intuitive to gain broad acceptance. Like productivity, heterogeneity is known to influence diversity at fine spatial scales, yet the possibility that richness might relate to water–energy dynamics at coarse spatial scales via a heterogeneity‐generating mechanism has received little attention. In this paper we outline such a conceptual model for plants that is internally consistent and testable. We believe it may help to explain the capacity of environments receiving different inputs of water and energy to support variable numbers of species at a range of spatial scales, the pervasive correlation between productivity and richness, some exceptions to the productivity–diversity relationship, the form of productivity–diversity curves and the link between richness and environmental ‘harshness’. The model may also provide an answer to one of the most venerable puzzles in the field of diversity studies: why high inputs of water and energy correspond to more species rather than simply more individuals.     

Evaluating the role of regional and local processes in structuring a larval trematode metacommunity of Helisoma trivolvis

Metacommunity theory has advanced our understanding of how local and regional processes affect the structure of ecological communities. While parasites have largely been omitted from metacommunity research, parasite communities can provide the large sample sizes and discrete boundaries often required for evaluating metacommunity patterns. Here, we used assemblages of flatworm parasites that infect freshwater snails (Helisoma trivolvis) to evaluate three questions: 1) what factors affect individual host infections within ponds? 2) Is the parasite metacommunity structured among ponds? And 3) what is the relative role of local versus regional processes in determining metacommunity structure and species richness among ponds? We examined 10 821 snails from 96 sites in five park complexes in the San Francisco Bay area, California, and found 953 infections from six parasite groups. At the within‐pond level, infection status of host snails correlated positively with individual snail size and pond infection prevalence for all six parasite groups. Using an ordination method to test for metacommunity structure, we found that the parasite metacommunity was organized in a non‐random pattern with species responding individually along an environmental gradient. Based on a model selection approach involving local and regional predictors, parasite species richness and metacommunity structure correlated with both local abiotic (pH and total dissolved nitrogen) and biotic (non‐host mollusk density, and H. trivolvis biomass) factors, with little support for regional predictors. Overall, this trematode metacommunity most closely followed the predictions from the species sorting or mass effects metacommunity paradigm, in which community diversity is filtered by local site characteristics.     

Tree Diversity,Environmental Heterogeneity,and Productivity in a Mexican Tropical Dry Forest1

Species diversity–environmental heterogeneity (D–EH) and species diversity–productivity (D–P) relationships have seldom been analyzed simultaneously even though such analyses could help to understand the processes underlying contrasts in species diversity among sites. Here we analyzed both relationships at a local scale for a highly diverse tropical dry forest of Mexico. We posed the following questions: (1) are environmental heterogeneity and productivity related?; (2) what are the shapes of D–EH and D–P relationships?; (3) what are individual, and interactive, contributions of these two variables to the observed variance in species diversity?; and (4) are patterns affected by sample size, or by partitioning into average local diversity and spatial species turnover? All trees (diameter at breast height ≥5 cm) within twenty‐six 0.2‐ha transects were censused; four environmental variables associated with water availability were combined into an environmental heterogeneity index; aboveground standing biomass was used as a productivity estimator. Simple and multiple linear and nonlinear regression models were run. Environmental heterogeneity and productivity were not correlated. We found consistently positive log‐linear D–EH and D–P relationships. Productivity explained a larger fraction of among‐transect variance in species diversity than did environmental heterogeneity. No effects of sample size were found. Different components of diversity varied in sensitivity to environmental heterogeneity and productivity. Our results suggest that species' differentiation along water availability gradients and species exclusion at the lowest productivity (driest) sites occur simultaneously, independently, and in a scale‐dependent fashion on the tree community of this forest.     

Urban ponds as an aquatic biodiversity resource in modified landscapes

Urbanization is a global process contributing to the loss and fragmentation of natural habitats. Many studies have focused on the biological response of terrestrial taxa and habitats to urbanization. However, little is known regarding the consequences of urbanization on freshwater habitats, especially small lentic systems. In this study, we examined aquatic macro‐invertebrate diversity (family and species level) and variation in community composition between 240 urban and 782 nonurban ponds distributed across the United Kingdom. Contrary to predictions, urban ponds supported similar numbers of invertebrate species and families compared to nonurban ponds. Similar gamma diversity was found between the two groups at both family and species taxonomic levels. The biological communities of urban ponds were markedly different to those of nonurban ponds, and the variability in urban pond community composition was greater than that in nonurban ponds, contrary to previous work showing homogenization of communities in urban areas. Positive spatial autocorrelation was recorded for urban and nonurban ponds at 0–50 km (distance between pond study sites) and negative spatial autocorrelation was observed at 100–150 km and was stronger in urban ponds in both cases. Ponds do not follow the same ecological patterns as terrestrial and lotic habitats (reduced taxonomic richness) in urban environments; in contrast, they support high taxonomic richness and contribute significantly to regional faunal diversity. Individual cities are complex structural mosaics which evolve over long periods of time and are managed in diverse ways. This facilitates the development of a wide range of environmental conditions and habitat niches in urban ponds which can promote greater heterogeneity between pond communities at larger scales. Ponds provide an opportunity for managers and environmental regulators to conserve and enhance freshwater biodiversity in urbanized landscapes whilst also facilitating key ecosystem services including storm water storage and water treatment.     

Factors influencing geographic patterns in diversity of forest bird communities of eastern Connecticut,USA

At regional scales, the most important variables associated with diversity are latitudinally‐based temperature and net primary productivity, although diversity is also influenced by habitat. We examined bird species richness, community density and community evenness in forests of eastern Connecticut to determine whether: 1) spatial and seasonal patterns exist in diversity, 2) energy explains the greatest proportion of variation in diversity parameters, 3) variation in habitat explains remaining diversity variance, and 4) seasonal shifts in diversity provide clues about how environmental variables shape communities. We sought to discover if our data supported predictions of the species–energy hypothesis. We used the variable circular plot technique to estimate bird populations and quantified the location, elevation, forest type, vegetation type, canopy cover, moisture regime, understory density and primary production for the study sites. We found that 1) summer richness and population densities are roughly equal in northeastern and southeastern Connecticut, whereas in winter both concentrate toward the coast, 2) variables linked with temperature explained much of the patterns in winter diversity, but energy‐related variables showed little relationship to summer diversity, 3) the effect of habitat variables on diversity parameters predominated in summer, although their effect was weak, 4) contrary to theory, evenness increased from summer to winter, and 5) support for predictions of species–energy theory was primarily restricted to winter data. Although energy and habitat played a role in explaining community patterns, they left much of the variance in regional diversity unexplained, suggesting that a large stochastic component to diversity also may exist.     

The richness–heterogeneity relationship differs between heterogeneity measures within and among habitats

The positive monotonic relationship between habitat heterogeneity and species richness is a cornerstone of ecology. Recently, it was suggested that this relationship should be unimodal rather than monotonic due to a tradeoff between environmental heterogeneity and population sizes, which increases local species extinctions at high heterogeneity levels. Here, we studied the richness–heterogeneity relationship for an avian community using two different environmental variables, foliage‐height diversity and cover type diversity. We analyzed the richness–heterogeneity within different habitat types (grasslands, savannas, or woodlands) and at the landscape scale. We found strong evidence that both positive and unimodal relationships exist at the landscape scale. Within habitats we found positive relationships between richness and heterogeneity in grasslands and woodlands, and unimodal relationships in savannas. We suggest that the length of the environmental heterogeneity gradient (which is affected by both spatial scale and the environmental variable being analyzed) affects the type of the richness–heterogeneity relationship. We conclude that the type of the relationship between species richness and environmental heterogeneity is non‐ubiquitous, and varies both within and among habitats and environmental variables.     

Temporal patterns of diversity, abundance and evenness for invertebrate communities from coastal freshwater and brackish water rock pools

Aquatic invertebrate data were collected from 49 erosional, Jamaican,rock pools between 1989 and 1998 and used to describe temporal patterns ofspecies diversity. This unique series of pools on the north coast of Jamaica,classified as either brackish (31) or freshwater (18), was used to determinehowdiversity changes over time, whether there was a difference between poolclassifications, and the impacts of environmental variables. Mean communitymetrics (richness, diversity, evenness, abundance) were not significantlydifferent between freshwater and brackish pools. However, there weresignificantdifferences among the eight sampling dates and differences over time dependedonpool classification. Measures of diversity for freshwater pools were relativelyconstant over time, implying little change at the community level. Brackishpools showed significant differences over time in species richness, totalabundance, and evenness implying that community composition and structure werenot static but changed in response to either environmental or biotic changes(possibly initiated by environmental change).Some temporal changes in community metrics could be linked to temporal changesin environmental variables. In brackish water pools, a significant increase inpool salinity between January 1991 and January 1992 corresponded to an increasein species richness, likely due to an increase in marine fauna. Similarly,changes in abundance and evenness corresponded to changes in temperature,dissolved oxygen, and pH. In addition, physicochemical variables used in thisstudy were shown to affect community metrics and those relationships dependedonpool classification. Most relationships between community metrics andenvironmental variables were negative with the exception of Simpson's diversityindex for which positive relationships were found. This may indicate that, aspool conditions become less favorable, a few species flourish and dominate thecommunity.     

Broad-scale spatial patterns of canopy cover and pond morphology affect the structure of a Neotropical amphibian metacommunity

Spatial and environmental processes influence species composition at distinct scales. Previous studies suggested that the distribution of larval anurans at the landscape-scale is influenced by environmental gradients related to adult breeding site selection, such as pond canopy cover, but not by water chemistry. However, the combined effects of spatial, pond morphology, and water chemistry variables on metacommunity structure of larval anurans have not been analyzed yet. We used a partial redundancy analysis with variation partitioning to analyze the relative influence of pond morphology (e.g., depth, area, and aquatic vegetation), water chemistry, and spatial variables on a tadpole metacommunity from southeastern Brazil. We predict that pond morphology and canopy cover will influence the metacommunity at broad spatial scales, while water chemistry would play a larger role at finer scales. We found that broad-scale spatial patterns of pond canopy cover and pond morphology strongly influenced metacommunity structure, with water chemistry being not significant. Additionally, species composition was spatially autocorrelated at short distances. We suggest that the reproductive behavior of adult anurans is driving tadpole metacommunity dynamics, since pond morphology, but not water chemistry affects breeding site selection by adults. Our results contribute to the understanding of amphibian species diversity in tropical wetlands.     

Temporal variability in the spatial and environmental determinants of functional metacommunity organization – stream fish in a human‐modified landscape

1. Quantifying the relative importance of environmental filtering versus regional spatial structuring has become an intensively studied area in the context of metacommunity ecology. However, most studies have evaluated the role of environmental and spatial processes using taxonomic data sets of single snapshot surveys. 2. Here, we examined temporal changes in patterns and possible processes behind the functional metacommunity organization of stream fishes in a human‐modified landscape. Specifically, we (i) studied general changes in the functional composition of fish assemblages among 40 wadeable stream sites during a 3‐year study period in the catchment area of Lake Balaton, Hungary, (ii) quantified the relative importance of spatial and environmental factors as determinants of metacommunity structure and (iii) examined temporal variability in the relative role of spatial and environmental processes for this metacommunity. 3. Partial triadic analysis showed that assemblages could be effectively ordered along a functional gradient from invertebrate consuming species dominated by the opportunistic life‐history strategy, to assemblages with a diverse array of functional attributes. The analysis also revealed that functional fish assemblage structure was moderately stable among the sites between the sampling periods. 4. Despite moderate stability, variance partitioning using redundancy analyses (RDA) showed considerable temporal variability in the contribution of environmental and spatial factors to this pattern. The analyses also showed that environmental variables were, in general, more important than spatial ones in determining metacommunity structure. Of these, natural environmental variables (e.g. altitude, velocity) proved to be more influential than human‐related effects (e.g. pond area, % inhabited area above the site, nutrient enrichment), even in this landscape with relatively low variation in altitude and stream size. 5. Pond area was, however, the most important human stressor variable that was positively associated with the abundance of non‐native species with diverse functional attributes. The temporal variability in the relative importance of environmental and spatial factors was probably shaped by the release of non‐native fish from fish ponds to the stream system during flood events. 6. To conclude, both spatial processes and environmental control shape the functional metacommunity organization of stream fish assemblages in human‐modified landscapes, but their importance can vary in time. We argue, therefore, that metacommunity studies should better consider temporal variability in the ecological mechanisms (e.g. dispersal limitation, species sorting) that determine the dynamics of landscape‐level community organization.     

Altitude effects on spatial components of vascular plant diversity in a subarctic mountain tundra

Environmental gradients are caused by gradual changes in abiotic factors, which affect species abundances and distributions, and are important for the spatial distribution of biodiversity. One prominent environmental gradient is the altitude gradient. Understanding ecological processes associated with altitude gradients may help us to understand the possible effects climate change could have on species communities. We quantified vegetation cover, species richness, species evenness, beta diversity, and spatial patterns of community structure of vascular plants along altitude gradients in a subarctic mountain tundra in northern Sweden. Vascular plant cover and plant species richness showed unimodal relationships with altitude. However, species evenness did not change with altitude, suggesting that no individual species became dominant when species richness declined. Beta diversity also showed a unimodal relationship with altitude, but only for an intermediate spatial scale of 1 km. A lack of relationships with altitude for either patch or landscape scales suggests that any altitude effects on plant spatial heterogeneity occurred on scales larger than individual patches but were not effective across the whole landscape. We observed both nested and modular patterns of community structures, but only the modular patterns corresponded with altitude. Our observations point to biotic regulations of plant communities at high altitudes, but we found both scale dependencies and inconsistent magnitude of the effects of altitude on different diversity components. We urge for further studies evaluating how different factors influence plant communities in high altitude and high latitude environments, as well as studies identifying scale and context dependencies in any such influences.     

Spatio-temporal drivers of different oomycete beta diversity components in Brazilian rivers

Disentangling the role of mechanisms driving metacommunity structure is fundamental for conservation strategies. Several studies have been done in aquatic communities; however, little is known about the factors driving oomycete communities. This research aimed to investigate beta diversity patterns and assess the role of environmental (chemical, physical, and hydrologic), spatial, and temporal (sampling months) factors in driving oomycete beta diversity in a spatial extent of 33 km from two Brazilian rivers. We took water samples in 10 sites quarterly, from August 2017 to May 2018. The partition of beta diversity into its components – species replacement and richness difference – was performed using the Jaccard dissimilarity index. Distance-based redundancy analysis and variation partitioning were used to assess the relationship between explanatory variables and beta diversity. We found that beta diversity was spatially and temporally high, and the replacement component was the main driver of the oomycete metacommunity’s beta diversity. Replacement and total beta diversity were explained mainly by spatial location and the month of sampling, while the richness difference was more associated with the environmental variables chlorophyll a and ammonia. Our findings suggest that dispersal limitation (spatial) and temporal factors are the main drivers of the total beta diversity and replacement in the oomycete metacommunity, while species sorting (environmental factor) influences the richness difference. Accordingly, that taking temporal factors into account in metacommunity studies is important to explain beta diversity patterns, especially in rivers with remarkable variability in hydrological regime and under eutrophic conditions.