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.     

From richer to poorer: successful invasion by freshwater fishes depends on species richness of donor and recipient basins

Evidence for the theory of biotic resistance is equivocal, with experiments often finding a negative relationship between invasion success and native species richness, and large‐scale comparative studies finding a positive relationship. Biotic resistance derives from local species interactions, yet global and regional studies often analyze data at coarse spatial grains. In addition, differences in competitive environments across regions may confound tests of biotic resistance based solely on native species richness of the invaded community. Using global and regional data sets for fishes in river and stream reaches, we ask two questions: (1) does a negative relationship exist between native and non‐native species richness and (2) do non‐native species originate from higher diversity systems. A negative relationship between native and non‐native species richness in local assemblages was found at the global scale, while regional patterns revealed the opposite trend. At both spatial scales, however, nearly all non‐native species originated from river basins with higher native species richness than the basin of the invaded community. Together, these findings imply that coevolved ecological interactions in species‐rich systems inhibit establishment of generalist non‐native species from less diverse communities. Consideration of both the ecological and evolutionary aspects of community assembly is critical to understanding invasion patterns. Distinct evolutionary histories in different regions strongly influence invasion of intact communities that are relatively unimpacted by human actions, and may explain the conflicting relationship between native and non‐native species richness found at different spatial scales.     

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.     

Evidence on ecotone concepts from switch,environmental and anthropogenic ecotones

Abstract. Four contrasting ecotones were sampled to address three questions: (1) Are there ‘ecotonal’ species, (2) Do ecotones possess higher (or lower) species richness than the adjacent communities? and (3) Are exotic species more likely to occur in ecotones? One ecotone was edaphic, one was apparently caused by a positive‐feedback switch, one was environmental/anthropogenic and one was entirely anthropogenic. The exact position of each ecotone was established from the spatial change in ordination scores. Ecotonal species, in the sense of species mainly restricted to the ecotone at the site, were present in all four ecotones. All but one of the ecotonal species were native. The switch ecotone and the purely anthropogenic ecotone also contained native species that were significantly more frequent in the ecotone than in either adjacent community. Species richness was intermediate between that of the two adjacent communities in three of the ecotones. In the environmental/anthropogenic ecotone, species richness was higher than in adjacent communities, but not significantly so. There were appreciable numbers of exotic species in the two ecotones with anthropogenic influence, one of which had a proportion of exotic species intermediate between the two adjacent communities. Contrary to theory, the proportion of exotic species in the second ecotone was significantly lower than in either adjacent community. We conclude that all three features we examined depend on the particular ecological conditions and the ecology of the species present; they are not intrinsic properties of ecotones.     

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.     

生态位分化与森林群落物种多样性维持研究展望

一直以来,生态学家和进化生物学家对森林群落物种多样格局及其形成机制持有不同的观点。虽然Robert Ricklefs将进化和生态过程整合的观点已经被群落生态学家广泛接受,但是区域物种进化历史以及局域群落微进化过程是否能够影响群落生态学过程以及这些过程如何影响群落结构和动态还有待商榷。经典的生态位理论同时强调了种间和种内生态位分化对群落多样性维持的影响。但是生态学家普遍认为种间差异足以代表群落内个体间的相互作用关系,并且由于进化过程导致的种内分化往往涉及较长的时间尺度,因此,虽然种内差异是自然选择的重要材料,物种对环境的适应性进化过程所导致的种内分化对群落构建的影响往往被生态学家所忽视。为此,通过回顾种间和个体生态位分化的研究历史,对两类研究分别进行简要阐述,强调在今后的群落生态学研究中需要考虑个体分化对局域群落构建的影响。     

A critical analysis of the ubiquity of linear local–regional richness relationships

Synthesis Despite theoretical criticisms, the ubiquity of linear relationships between local and regional species richness has long been used to justify it as a valid framework to conclude that local communities are not saturated with species. However, we reanalyzed published studies with a new unbiased method and found no prevalence of linear relationships and more than 40% of misclassifications, including textbook examples. We thus conclude that the prevailing argument in favor of associating a valid ecological interpretation to local–regional species richness plots, its ubiquity, is not sustained, and that ecologists could use for instance metacommunity theory to make inference on the strength of local and regional processes. Identifying the relative importance of regional and local processes to local species diversity is a central issue to many questions in basic and applied ecology. One widely‐used method is to plot local species richness against its regional richness to infer whether regional or local processes determine local diversity. However, this method increases the tendency to find regional prevalence as suggested by a recent simulation. We reanalyzed studies in the literature with an unbiased method and found no prevalence of either regional or local processes. In addition, almost 40% of the studies and 50% of the ecology textbook examples using the traditional method were misclassified. Our findings reinforce the need of alternative, novel tools identified by for instance metacommunity theory to go beyond the studies of local–regional relationships in the ecological literature that focus on the interdependence of regional and local processes.     

An intercontinental comparison of dung beetle diversity between two mediterranean-climatic regions: local versus regional and historical influences

The species richness of biological communities is influenced by both local ecological, regional ecological, and historical factors. The relative importance of these factors may be deduced by comparison between communities in climatically and ecologically equivalent, but geographically and historically separate regions of the world. This claim is based on the hypothesis that community processes driven by similar local ecological factors lead to convergence in species richness whereas those driven by differing regional or historical factors lead to divergence. An intercontinental comparison between the winter rainfall regions of South Africa and the Iberian Peninsula showed that overall species richness of dung beetles was dissimilar at local, subregional and regional scales in Scarabaeidae s. str. but similar at all scales in Aphodiinae. Removal of species widespread in the summer rainfall region of Africa or the temperate region of Europe (regional component) resulted in dissimilarity in species richness of mediterranean endemics at all scales in both dung beetle taxa. However, the lines joining each set of species richness values were parallel which may indicate similarities in processes between different mediterranean climatic regions despite slight differences in latitudinal range. The dominant pattern of dissimilarity or non-convergence may be related primarily to intercontinental differences in regional biogeographical and evolutionary history (faunal dispersal, glaciation effects in relation to geographical barriers to dispersal, speciation history, long-term disturbance history). The limited pattern of similarity or convergence in overall species richness of Aphodiinae may be a chance result or primarily related to intercontinental similarities in local ecological factors.     

Testing for local species saturation with nonindependent regional species pools

Identifying the factors controlling local community structure is a central problem in ecology. Ecologists frequently use regression to test for a nonlinear saturating relationship between local community richness and regional species pool richness, suggesting that species interactions limit the number of locally coexisting species. However, communities in different regions are not independent if regions share species. We present a Monte Carlo test for whether an observed local-regional richness relationship is significantly different from that expected when regions are nonindependent and species interactions do not limit community membership. We illustrate this test with data from experimental microcosm communities. A conventional F -test suggests a significant saturating relationship between realized community richness and species pool richness. However, the Monte Carlo test fails to reject the null hypothesis that species interactions do not affect community richness. Strong species interactions do not necessarily set an absolute upper limit to the number of locally coexisting species.     

Do plant communities exist? Evidence from scaling‐up local species‐area relations to the regional level

Abstract. One long tradition in ecology is that discrete communities exist, at least in the sense that there are areas of relatively uniform vegetation, with more rapid change in species composition between them. The alternative extreme view is the Self‐similarity concept – that similar community variation occurs at all spatial scales. We test between these two by calculating species‐area curves within areas of vegetation that are as uniform as can be found, and then extrapolating the within‐community variation to much larger areas, that will contain many ‘communities’. Using the Arrhenius species‐area model, the extrapolations are remarkably close to the observed number of species at the regional/country level. We conclude that the type of heterogeneity that occurs within ‘homogeneous’ communities is sufficient to explain species richness at much larger scales. Therefore, whilst we can speak of ‘communities’ for convenience, the variation that certainly exists at the ‘community’ level can be seen as only a larger‐scale manifestation of micro‐habitat variation.     

The relative of species pools in determining plant species richness: an alternative explanation of species coexistence?

Explanations of the pattern of species have traditionally relied on small-scale, local processes occurring in ecological time. Differences in species richness have associated with different mechanisms avoiding competition, such as spatiotemporal heterogeneity (weaker competitors may find a more favourable place or time) or environmental stress (competition is assumed to be less intensive under difficult conditions). More recently, large-scale process have been taken into account, raising such questions as: which plant species may potentially grow in a certain community? Are evolutionary processes and species dispersal responsible for the differences between communities? The species-pool theory attempts to answer these general questions, and information about species pools is needed for the design of experiments where the number of species in a community is manipulated.     

A unified model explains commonness and rarity on coral reefs

Abundance patterns in ecological communities have important implications for biodiversity maintenance and ecosystem functioning. However, ecological theory has been largely unsuccessful at capturing multiple macroecological abundance patterns simultaneously. Here, we propose a parsimonious model that unifies widespread ecological relationships involving local aggregation, species‐abundance distributions, and species associations, and we test this model against the metacommunity structure of reef‐building corals and coral reef fishes across the western and central Pacific. For both corals and fishes, the unified model simultaneously captures extremely well local species‐abundance distributions, interspecific variation in the strength of spatial aggregation, patterns of community similarity, species accumulation, and regional species richness, performing far better than alternative models also examined here and in previous work on coral reefs. Our approach contributes to the development of synthetic theory for large‐scale patterns of community structure in nature, and to addressing ongoing challenges in biodiversity conservation at macroecological scales.     

The alpha–beta–regional relationship: providing new insights into local–regional patterns of species richness and scale dependence of diversity components

Ecologists frequently regress local species richness on regional species richness to draw inferences about the processes that structure local communities. A more promising approach is to quantify the contributions of alpha and beta diversity to regional diversity (the ABR approach) using additive partitioning. We applied this approach to four local–regional relationships based on data from 583 arboreal beetle species collected in a hierarchically nested sampling design. All four local–regional relationships exhibited proportional sampling, yet the ABR approach indicated that each was produced by a different combination of alpha and beta richness. Using the results of the ABR analysis, we also analysed the scale dependence of alpha and beta using a hierarchical linear model. Alpha diversity contributed less than expected to regional diversity at the finest spatial scale and more than expected at the broadest spatial scale. A switch in relative dominance from beta to alpha diversity with increasing spatial scale suggested scale transitions in ecological processes. Analysing the scale dependence of diversity components using the ABR approach furthers our understanding about the additivity of species diversity in biological communities.     

Local versus landscape-scale effects of anthropogenic land-use on forest species richness

The study investigated the effects of human-induced landscape patterns on species richness in forests. For 80 plots of fixed size, we measured human disturbance (categorized as urban/industrial and agricultural land areas), at ‘local’ and ‘landscape’ scale (500 m and 2500 m radius from each plot, respectively), the distance from the forest edge, and the size and shape of the woody patch. By using GLM, we analyzed the effects of disturbance and patch-based measures on both total species richness and the richness of a group of specialist species (i.e. the ‘ancient forest species’), representing more specific forest features. Patterns of local species richness were sensitive to the structure and composition of the surrounding landscape. Among the landscape components taken into account, urban/industrial land areas turned out as the most threatening factor for both total species richness and the richness of the ancient forest species. However, the best models evidenced a different intensity of the response to the same disturbance category as well as a different pool of significant variables for the two groups of species. The use of groups of species, such as the ancient forest species pool, that are functionally related and have similar ecological requirements, may represent an effective solution for monitoring forest dynamics under the effects of external factors. The approach of relating local assessment of species richness, and in particular of the ancient forest species pool, to land-use patterns may play an important role for the science-policy interface by supporting and strengthening conservation and regional planning decision making.     

Palaeontology., adaptation and community ecology: A response to Walter and Pater son (1994)

Abstract This paper challenges Walter and Paterson's (1994) assertion that the community concept ought to be abandoned because of recent palaeontological evidence pointing to the ‘individualistic’ nature of biological communities. The ‘individualistic’ versus ‘superorganismic’ community concepts might provide good grist for the philosophical mill, but have little practical relevance to contemporary community ecology. Ecologists define communities in terms of current species distributions and interactions, and seek to integrate the roles of both biotic and abiotic factors influencing species distributions. There is no assumption of tight co-evolution among component species; Walter and Paterson confuse ‘organization’ with ‘co-adaptation’. Nor, contrary to the authors’ claims, is there an implicit assumption that all community patterns are caused by competition. For most ecologists, the ‘competition debate’ ended a decade ago. Walter and Paterson's view that competition is rarely, if ever, important in structuring communities is not even held by the main protaganists of the ‘competition is not so important’ school of the 1980s, and is in direct contradiction of the extensive, more recent literature on the subject. It entirely ignores plant ecology. Many of Walter and Paterson's misunderstandings appear to arise from the false premise that explanation of adaptation should be the ultimate goal of any ecological discipline. The authors are hostile to community ecology because, if communities are individualistic, then little light can be shed on species adaptations. Fortunately, most ecologists are not so preoccupied with adaptation.     

Hierarchical effects of environmental filters on the functional structure of plant communities: a case study in the French Alps

Understanding the influence of the environment on the functional structure of ecological communities is essential to predict the response of biodiversity to global change drivers. Ecological theory suggests that multiple environmental factors shape local species assemblages by progressively filtering species from the regional species pool to local communities. These successive filters should influence the various components of community functional structure in different ways. In this paper, we tested the relative influence of multiple environmental filters on various metrics of plant functional trait structure (i.e. ‘community weighted mean trait’ and components of functional trait diversity, i.e. functional richness, evenness and divergence) in 82 vegetation plots in the Guisane Valley, French Alps. For the 211 sampled species we measured traits known to capture key aspects of ecological strategies amongst vascular plant species, i.e. leaf traits, plant height and seed mass (LHS). A comprehensive information theory framework, together with null model based resampling techniques, was used to test the various environmental effects. Particular community components of functional structure responded differently to various environmental gradients, especially concerning the spatial scale at which the environmental factors seem to operate. Environmental factors acting at a large spatial scale (e.g. temperature) were found to predominantly shape community weighted mean trait values, while fine‐scale factors (topography and soil characteristics) mostly influenced functional diversity and the distribution of trait values among the dominant species. Our results emphasize the hierarchical nature of ecological forces shaping local species assemblage: large‐scale environmental filters having a primary effect, i.e. selecting the pool of species adapted to a site, and then filters at finer scales determining species abundances and local species coexistence. This suggests that different components of functional community structure will respond differently to environmental change, so that predicting plant community responses will require a hierarchical multi‐facet approach.     

The species pool hypothesis: the necessity to shift emphasis

The species pool of a biological community is determined as a group of species that inhabit some area and potentially can be included in a given community. The species pool hypothesis, i.e. the assumption that the size of species pool strongly influences species richness of local community can be confirmed if there is positive linear relationship between these two variables. The results of hypothesis testing however are not obvious. For example, correlation between local richness and species pool size can be caused by their dependence on the third variable--capacity of environment. It seems that in case of decreasing area occupied by local community the environmental conditions become more important than species pool size. If that is true, the influence of species pool on local species richness is not significant. However one can estimate the degree of unsaturation of species pool on the basis of relationships between the number of species in small locations occupied by similar local communities and their species pool. We think, that study of local and regional species richness should shift the emphasis--from the analysis of species pool influence on local community richness to the estimation of historical, ecological and anthropogenic factors in variation of species pool size. The local species richness should be considered rather as a tool (allowing to compare the species capacity of biological communities), than as an object of such study.     

Quantitative partitioning of regional and local processes shaping regional diversity patterns

Much recent research explaining plant community diversity has focused on comparing the relative impacts of regional and local processes. We employed a novel analysis to quantify the effects of multiple regional and local processes on species richness, and to make quantitative comparisons of those effects across two sites that differ in plot-level species richness, productivity and environmental conditions. While abiotic stress and competition limited richness within the communities at both sites, only differences in the overall pool of species at the site, likely resulting from long-distance dispersal and climate fluctuations, explained the differences in plot-level richness between sites. Patterns in local richness may be driven by a temporal storage effect, with greater richness in the site with greater climatic variability. By identifying both the factors that impact diversity within communities and those that vary systematically across communities, our integrated approach provides a better understanding of regional diversity patterns.     

Assessing the impacts of imperfect detection on estimates of diversity and community structure through multispecies occupancy modeling

Detecting all species in a given survey is challenging, regardless of sampling effort. This issue, more commonly known as imperfect detection, can have negative impacts on data quality and interpretation, most notably leading to false absences for rare or difficult‐to‐detect species. It is important that this issue be addressed, as estimates of species richness are critical to many areas of ecological research and management. In this study, we set out to determine the impacts of imperfect detection, and decisions about thresholds for inclusion in occupancy, on estimates of species richness and community structure. We collected data from a stream fish assemblage in Algonquin Provincial Park to be used as a representation of ecological communities. We then used multispecies occupancy modeling to estimate species‐specific occurrence probabilities while accounting for imperfect detection, thus creating a more informed dataset. This dataset was then compared to the original to see where differences occurred. In our analyses, we demonstrated that imperfect detection can lead to large changes in estimates of species richness at the site level and summarized differences in the community structure and sampling locations, represented through correspondence analyses.     

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.