Strong influence of regional species pools on continent-wide structuring of local communities

There is a long tradition in ecology of evaluating the relative contribution of the regional species pool and local interactions on the structure of local communities. Similarly, a growing number of studies assess the phylogenetic structure of communities, relative to that in the regional species pool, to examine the interplay between broad-scale evolutionary and fine-scale ecological processes. Finally, a renewed interest in the influence of species source pools on communities has shown that the definition of the source pool influences interpretations of patterns of community structure. We use a continent-wide dataset of local ant communities and implement ecologically explicit source pool definitions to examine the relative importance of regional species pools and local interactions for shaping community structure. Then we assess which factors underlie systematic variation in the structure of communities along climatic gradients. We find that the average phylogenetic relatedness of species in ant communities decreases from tropical to temperate regions, but the strength of this relationship depends on the level of ecological realism in the definition of source pools. We conclude that the evolution of climatic niches influences the phylogenetic structure of regional source pools and that the influence of regional source pools on local community structure is strong.     

Native and introduced fish species richness in French lakes: local and regional influences

Aim To analyse the importance of local and regional influences on the patterns of species richness in natural and man‐made lakes and to infer the impacts of human‐mediated introductions on these patterns. Location France. Methods Species occurrence data were gathered for 25 natural and 51 man‐made lakes. Analysis is based on regression models of local richness against their related regional richness and lake environmental variables. Results Local native richness was mostly controlled by the regional richness. Conversely, local total richness was mainly explained by local variables. These statements apply to both natural and man‐made lakes. Lacustrine systems displayed weak resistance to invaders. Main conclusions Species introductions have apparently contributed to saturate fish communities in these systems even if no clear negative effect on the survival of native species (i.e. species extinction) is detectable so far.     

Are communities saturated? On the relationship between α, β and γ diversity

A popular way to suggest a regional influence on local species diversity has been to plot local versus regional diversity. The form of these curves has been interpreted as evidence for or against "community saturation" due to species interactions. This interpretation, however, is unwarranted. Using the concepts of α, β and γ diversity, I show that local–regional richness curves are determined by the way total diversity is partitioned between its α and β components, which itself is a matter of scale. Changing the scale of the local community amounts to changing the scale at which the heterogeneity of the interactions between organisms and their environment manifests itself, and hence the balance between α and β diversity. Community saturation may occur because of physical limitations, but there are no theoretical grounds for the belief that species interactions set an absolute upper limit to diversity at any scale. A distinction between different meanings of the concept of "saturation" is proposed to clarify this issue. I argue that the challenge now is to understand the relationship between α and β diversity at multiple scales, and the processes that determine it.     

The relationship between local and regional diatom richness is mediated by the local and regional environment

Aim In this continental study, species richness at local (LSR) and regional (RSR) scales was correlated and examined as a function of stream (local) and watershed (regional) environment in an effort to elucidate what factors control diatom biodiversity across scales. Location Conterminous United States. Methods Data on diatom richness, stream conditions and watershed properties were generated by the US Geological Survey. In the present investigation, RSR was estimated as the total diatom richness in a hydrologic study unit and, together with stream and watershed characteristics, was included in stepwise multiple regressions of LSR. The unique and shared contributions of RSR, stream and watershed environment to the explained variance in LSR were determined by variance partitioning. RSR was regressed against stream and basin features averaged per study unit. Results LSR responded most strongly to variability in stream manganese concentration and RSR. Other predictors included stream discharge and iron concentration, soil organic matter content and fertilization, and proportions of open water, barren land and forest in the watershed. Variance partitioning revealed that RSR had the lowest independent contribution to explained variance in LSR. Multiple regressions identified average stream iron concentration as the most important predictor of RSR. Main conclusions Local micronutrient concentration was the major predictor of LSR, followed by RSR. Since average micronutrient supply in the region was the chief determinant of RSR, it is proposed that micronutrients had both a direct effect on LSR and an indirect effect through RSR. The same argument is extended to watershed features with an impact on stream trophic status, because of their substantial contributions to the explained variance in both LSR and RSR. Considering that the major proportion of LSR variance explained by RSR originated from the covariance of RSR with stream and watershed properties, it is concluded that the LSR–RSR relationship was mediated by the local and regional environment.     

Toward a better understanding of the regional causes of local community richness

Despite widespread acknowledgement that local ecological communities are profoundly shaped by regional-scale influences, including evolutionary and biogeographic processes, this perspective has yet to be widely incorporated into ecological research. Drawing on recent research, we propose four steps towards making regional influences a stronger part of research on the richness of local communities: (1) identifying the regional-scale causes of variation in species richness in the systems ecologists study; (2) testing for effects of regional richness on local richness, using improved observational and experimental analyses to overcome earlier problems; (3) simultaneously analysing environmental influences on regional and local species richness as well as the influence of regional richness on local richness and (4) considering the potential reciprocal effects of local processes on regional richness. In conclusion, we suggest some ways that similar approaches could be applied to other aspects of community structure beyond species richness.     

Processes involved in species saturation of ground‐dwelling ant communities (Hymenoptera,Formicidae)

The species saturation hypothesis in ground‐dwelling ant communities was tested, the relationship between local and regional species richness was studied and the possible processes involved in this relationship were evaluated in the present paper. To describe the relationship between local and regional species richness, the ground‐dwelling ant fauna of 10 forest remnants was sampled, using 10 1 m² quadrats in each remnant. The ants were extracted from the litter by using Winkler sacs. Using regression analyses, an asymptotic pattern between local and regional species richness was detected. This saturated pattern may be related to three processes: (i) high interspecific competition; (ii) habitat species specialization; or (iii) stochastic equilibrium. It is concluded that non‐interactive processes, such as stochastic equilibrium and habitat specialization may act as factors regulating species richness in this community. The predominance of locally restricted species, in all sampled remnants, seems to indicate the occurrence of a high degree of habitat specialization by the ant species. This result is evidence for the hypothesis that community saturation has been generated by non‐interactive processes. Although ants are frequently described as highly interactive, it is possible that interspecific competition is not important in the structuring of ground‐dwelling ant communities.     

Effects of regional pool size on local diversity in small-scale annual plant communities

The shape of the relationship between local and regional richness has been used to infer the relative strength of local vs. regional processes in structuring communities. While most empirical work has demonstrated linear relationships, recent reviews suggest that prior work has contained biases by selecting communities from different biological provinces and by defining large-scale local communities. We evaluate the utility of inferring interaction strength from local–regional relationships by examining 11 years of data from small-scale, annual plant communities at one site with strong local interactions. Both winter and summer annual plant assemblages exhibited linear rather than saturating relationships over a wide range of regional pool sizes. Our empirical work supports recent theory suggesting that local–regional relationships cannot be used to infer the strength of interactions in communities.     

The relationship between local and regional diversity of indigenous forest fauna in KwaZulu-Natal Province, South Africa

The relationship between local and regional diversity was tested by regressing local community richness against regional species diversity for three taxa, birds, butterflies and mammals, in subtropical forest. The quadratic model best fits the relationship between local and regional diversity for birds. Local bird species richness is theoretically independent of the size of the regional pool of species and may represent saturated communities. A linear model best describes the relationship for mammals and butterflies. For mammals, the slope is shallow (0.264) and regional richness overestimates local species richness, suggesting communities are undersaturated. Extinction filtering may explain this pattern. Past climatic changes have filtered out many mammalian species, these changes have been too recent for autochthanous speciation, and the relatively low vagility of mammals has prevented extensive recolonisation. Differences in the nature of the diversity relationship between taxa are as much due to independent evolutionary histories as to differences in vagility and colonising potential. A pervasive role is suggested for regional biogeographic processes in the development of faunal assemblage structure. Large-scale processes are not considered in current conservation plans. We encourage the shift of conservation emphasis from local ecological processes and species interactions, to whole communities and consideration of regional processes.     

The implications of palaeontological evidence for theories of ecological communities and species richness

Abstract Palaeontological evidence raises several questions that relate to current explanations of ecological communities, to the classification of communities and to interpretations of species richness. The first question relates to the stability of species detected in the fossil record. Coupled with that is the issue of incidental association of species on the same trophic level through differential effects of climatic change on the different species. Such observations are seen to support the ‘individualistic’ concept of communities. Recent statements about this concept leave unresolved questions about the acquisition of adaptation, and about the place of adaptation theory in theories of ecological communities and interpretations of ‘regional species richness’. At issue is whether there is justification for continuing to classify communities as a basis for understanding them. There is good reason to reject this approach for one in which questions about communities and ‘local’ and ‘regional’ species richness are replaced by more specific and basic questions about the relationship between adaptation, distribution and abundance, and ecological interactions. Some recent efforts to incorporate species theory into community theory fail because their basis remains the flawed concept of ‘local community’.     

The relationships between local and regional species richness and spatial turnover

Aim To determine the empirical relationships between species richness and spatial turnover in species composition across spatial scales. These have remained little explored despite the fact that such relationships are fundamental to understanding spatial diversity patterns. Location South‐east Scotland. Methods Defining local species richness simply as the total number of species at a finer resolution than regional species richness and spatial turnover as turnover in species identity between any two or more areas, we determined the empirical relationships between all three, and the influence of spatial scale upon them, using data on breeding bird distributions. We estimated spatial turnover using a measure independent of species richness gradients, a fundamental feature which has been neglected in theoretical studies. Results Local species richness and spatial turnover exhibited a negative relationship, which became stronger as larger neighbourhood sizes were considered in estimating the latter. Spatial turnover and regional species richness did not show any significant relationship, suggesting that spatial species replacement occurs independently of the size of the regional species pool. Local and regional species richness only showed the expected positive relationship when the size of the local scale was relatively large in relation to the regional scale. Conclusions Explanations for the relationships between spatial turnover and local and regional species richness can be found in the spatial patterns of species commonality, gain and loss between areas.     

Local–regional diversity relationship varies with spatial scale in lotic diatoms

Aim Ecologists have shown increasing interest in the relative roles of local and regional factors in structuring biotic communities. One approach to studying this is to examine the relationship between local species richness (LSR) and regional species richness (RSR). We examined the LSR–RSR relationship in stream diatoms, using two data sets that varied in spatial extent. At broad spatial extent ranging across drainage systems, we expected climatic and dispersal‐related factors to constrain LSR, thus resulting in a linear LSR–RSR relationship. However, at small spatial scales dispersal across sites should be unconstrained, resulting in strong local interactions and a weak or asymptotic LSR–RSR relationship. Location Boreal streams in Finland. Methods For data set 1, we sampled 15 stream riffles (localities) in each of eight drainage systems (regions), with the latitudinal gradient between the southernmost and northernmost sites being almost 1100 km. For data set 2, a locality for estimating LSR was a single stone, and each riffle represented a region for estimating RSR. We sampled 20 stones in each of eight riffles. We used linear regressions to examine the relationship between LSR and RSR across regions. We used both observed richness values, as well as values estimated with the Chao1 estimator. Results We found a relatively strong linear relationship between the Chao1‐estimated mean LSR and RSR (R² = 0.654, P = 0.015) across drainage systems. The slope of the regression was 0.643 and it did not differ from 1.0, thus indicating linearity. At the riffle scale, however, LSR and RSR were not linearly related, and the slope of the regression (0.039) differed significantly from 1.0, indicating curvilinearity. Main conclusions These results suggest that the relationship between mean LSR and RSR varies across spatial scales in diatoms – from significantly linear at large scales to curvilinear at small scales. These plots imply strong regional enrichment in stream diatoms across drainage systems. Their diversity is thus determined largely by the composition of the regional species pool, as also in many macroorganisms. In contrast, at small spatial scales the LSR–RSR relationship implied a hard limit to local diversity, reflecting the primacy of local processes.     

Using historical biogeography to test for community saturation

Saturation is the idea that a community is effectively filled with species, such that no more can be added without extinctions. This concept has important implications for many areas of ecology, such as species richness, community assembly, invasive species and climate change. Here, we illustrate how biogeography can be used to test for community saturation, when combined with data on local species richness, phylogeny and climate. We focus on a clade of frogs (Terrarana) and the impact of the Great American Biotic Interchange on patterns of local richness in Lower Middle America and adjacent regions. We analyse data on species richness at 83 sites and a time‐calibrated phylogeny for 363 species. We find no evidence for saturation, and show instead that biotic interchange dramatically increased local richness in the region. We suggest that historical biogeography offers thousands of similar long‐term natural experiments that can be used to test for saturation.     

The species pool concept applied to forests in a fragmented landscape: dispersal limitation versus habitat limitation

Abstract. The species pool concept has been used as a theoretical framework for understanding local community richness. A significant problem in putting the concept into practice is the lack of methods for determining the size of the species pool. We tested the hypothesis that species composition of recent forests is primarily determined by the species composition of neighbouring older forests against the null‐hypothesis that species are a random sample of the species occurring in the study area. Forest plant species composition of recently established fragments was significantly correlated with species composition in neighbouring older forests (i.e. the local species pool). When older forest within a neighbourhood of 1000m radius is considered, seed dispersal sources can be found for 91% of the flora in the recent forests. For an individual fragment, dispersal is a much more important determinant of species presence than the environment, with an average of 46% of the total pool excluded from local pools by dispersal limitation and only 8% excluded by environmental limitations. The species richness of recent forests is on average 23% of the local species pool. Several hypotheses are proposed for this low percentage, such as asymmetric competition due to the early successional state or the limited colonization period.     

Habitat selection behaviour links local and regional scales in aquatic systems

The role of habitat selection behaviour in the assembly of natural communities is an increasingly important theme in ecology. At the same time, ecologists and conservation biologists are keenly interested in scale and how processes at scales from local to regional interact to determine species distributions and patterns of biodiversity. How important is habitat selection in generating observed patterns of distribution and diversity at multiple spatial scales? In theory, habitat selection in response to interacting species can generate both positive and negative covariances among species distributions and create the potential to link processes of community assembly across multiple scales. Here I demonstrate that habitat selection by treefrogs in response to the distribution of fish predators functions at both the regional scale among localities and the local scale among patches within localities, implicating habitat selection as a critical link between local communities and the regional dynamics of metacommunities in complex landscapes.