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.     

Regional enrichment of local assemblages is robust to variation in local productivity, abiotic gradients, and heterogeneity

Theory predicts that the effects of regional richness on the richness of local communities may depend on the productivity, resource availability, and/or heterogeneity of local sites. Using the wetland plant communities of 50 independent streams as 'regions', we tested whether: (1) local richness in 1-m² quadrats and 50-m stream segments was positively related to regional richness, even after environmental influences were considered; and (2) the effect of regional richness would interact with the effects of biomass, soil moisture, and/or heterogeneity on local richness. In models that explained up to 88% of variation in local richness, we found that richness at both local scales was positively related to regional richness, and that regional richness did not interact with any of the environmental gradients that also shaped local richness. We conclude that species availability from the regional pool may consistently enrich local communities, even while other constraints on local richness operate.     

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.     

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.     

局域与区域生态过程对太白山牛皮桦林林隙内草本植物物种丰富度的影响

以太白山牛皮桦林林隙内草本植物总数为区域物种丰富度,分别采用0.25m2和1m2样方重复抽样的丰富度平均值为局域物种丰富度,来探讨区域物种多样性变异对局域多样性的影响。结果显示:(1)0.25m2和1m2局域物种丰富度与区域物种丰富度显著相关(r=0.791和r=0.861),且随区域物种丰富度的增加而增加;同时,林隙面积也能显著增加局域和区域物种丰富度。(2)控制林隙面积变量的多元回归分析显示,0.25m2和1m2局域物种丰富度与区域物种丰富度存在显著线性回归关系(R2=0.642和R2=0.743);方差分离分析显示,林隙面积仅能解释0.25m2和1m2局域物种丰富度变异的4.0%和4.4%,而区域物种丰富度能解释25.8%和35.3%。研究表明,区域物种丰富度变异在一定程度上决定着局域物种丰富度的组成。     

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.     

Are local patterns of anthropoid primate diversity related to patterns of diversity at a larger scale?

Aims (1) To determine the relationship between local and regional anthropoid primate species richness. (2) To establish the spatial and temporal scale at which the ultimate processes influencing patterns of primate species coexistence operate. Location Continental landmasses of Africa, South America and Asia (India to China, and all islands as far south as New Guinea). Methods The local–regional species richness relationship for anthropoid primates is estimated by regressing local richness against regional richness (independent variable). Local richness is estimated in small, replicate local assemblages sampled in regions that vary in total species richness. A strong linear relationship is taken as evidence that local assemblages are unsaturated and local richness results from proportional sampling of the regional pool. An asymptotic curvilinear relationship is interpreted to reflect saturated communities, where strong biotic interactions limit local richness and local processes structure the species assemblage. As a further test of the assumption of local assemblage saturation, we looked for density compensation in high‐density local primate assemblages. Results The local–regional species richness relationship was linear for Africa and South America, and the slope of the relationship did not differ between the two continents. For Asia, curvilinearity best described the relationship between local and regional richness. Asian primate assemblages appear to be saturated and this is confirmed by density compensation among Asian primates. However, density compensation was also observed among African primates. The apparent assemblage saturation in Asia is not a species–area phenomenon related to the small size of the isolated islands and their forest blocks, since similar low local species richness occurs in large forests on mainland and/or peninsular Asia. Main conclusions In Africa and South America local primate assemblage composition appears to reflect the influence of biogeographic processes operating on regional spatial scales and historical time scales. In Asia the composition of primate assemblages are by‐and‐large subject to ecological constraint operating over a relatively small spatial and temporal scale. The possible local influence of the El Niño Southern Oscillations on the evolution and selection of life‐history characteristics among Asian primates, and in determining local patterns of primate species coexistence, warrants closer inspection.     

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’.     

Local–regional species richness relationships are linear at very small to large scales in west-central Pacific corals

If local communities are saturated with species, the relationship between local and regional species richness [the local species richness (LSR)–regional species richness (RSR) relationship] is predicted to become increasingly curvilinear at more local spatial scales. This study tested whether the LSR–RSR relationship for coral species was linear or curvilinear at three local scales across the west-central Pacific Ocean, along a regional biodiversity gradient that includes the world’s most diverse coral assemblages. The local scales comprised transects 10^0–2 m apart, sites 10^3–4 m apart and islands 10^4–6 m apart. The LSR–RSR relationship was never significantly different from linear at any scale. When the Chao1 estimator was used to predict true RSR and LSR, all relationships were also strongly linear. We conclude that local assemblages are open to regional influences even when the local scale is very small relative to the regional scale, and even in extraordinarily rich regions.     

The Relationship between Local Species Richness and Species Pool: A Case Study from the High Mountains of the Greater Caucasus

The causes of linear relationships between local species richness and the size of the actual species pool in closed subalpine meadow communities and open plant communities of the alpine stony substrate (the Greater Caucasus Mountains) were analyzed using a computer simulation model. The results demonstrated that this relationship is insufficient evidence for the variation of local species richness among communities is wholly or partly determined by regional processes (the species-pool hypothesis). A relatively proportional ratio between these variables can also arise where local species richness and the size of the species pool both depend on local processes, or where local species richness is determined by local factors alone while the size of the species pool is determined by both local and regional factors.     

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.     

Diversity patterns of stygobiotic crustaceans across multiple spatial scales in Europe

1. Using species distribution data from 111 aquifers distributed in nine European regions, we examined the pairwise relationships between local species richness (LSR), dissimilarity in species composition among localities, and regional species richness (RSR). In addition, we quantified the relative contribution of three nested spatial units – aquifers, catchments and regions – to the overall richness of groundwater crustaceans.
2. The average number of species in karst and porous aquifers (LSR) varied significantly among regions and was dependent upon the richness of the regional species pool (RSR). LSR–RSR relationships differed between habitats: species richness in karstic local communities increased linearly with richness of the surrounding region, whereas that of porous local communities levelled off beyond a certain value of RSR.
3. Dissimilarity in species composition among aquifers of a region increased significantly with increasing regional richness because of stronger habitat specialisation and a decrease in the geographic range of species among karst aquifers. Species turnover among karst aquifers was positively related to RSR, whereas this relationship was not significant for porous aquifers.
4. The contribution of a given spatial unit to total richness increased as size of the spatial unit increased, although 72% of the overall richness was attributed to among-region diversity. Differences in community composition between similar habitats in different regions were typically more pronounced than between nearby communities from different habitats.
5. We conclude by calling for biodiversity assessment methods and conservation strategies that explicitly integrate the importance of turnover in community composition and habitat dissimilarity at multiple spatial scales.     

Patterns of species richness for blackflies (Diptera: Simuliidae) in the Nearctic and Neotropical regions

Abstract.  1. Patterns of simuliid species richness were examined over a variety of scales at 532 stream sites in the Nearctic (394) and Neotropical (138) regions. In Nearctic streams, species richness of immature blackflies both within and across ecoregions and over two seasons was examined. Stream variables at each site included seston, width, depth, velocity, discharge, conductivity, pH, dissolved oxygen, water temperature, dominant streambed-particle size, canopy cover, and riparian vegetation. These variables were subjected to a principal component analysis and derived principal components were related back to richness, using regression analysis. At the level of the stream reach, richness was not highly correlated with single-point measurements of stream conditions.
2. Using data from both Nearctic and Neotropical sites, the effect of regional richness on local richness was examined. As regional richness increased, local diversity reached an asymptote in which further increases in regional richness were not matched by increases in local richness. Hence, simuliid communities are best described as saturated (type II) communities, consistent with the current view of lotic communities as non-equilibrium systems.
3. The well-established pattern of greater species richness in tropical regions was not observed in this study. To the contrary, blackfly richness is higher in temperate streams than in tropical streams at both local and regional scales.     

Non-interactive fish communities in the coastal streams of North-western France

1. Spatial patterns of freshwater fish species at regional and local scales were investigated to explore the possible role of interspecific interactions in influencing distribution and abundance within communities occupying coastal streams of North-Western France.
2. Nine sites from nine streams situated in the same biogeographical region were sampled annually over the 6-year period from 1990 to 1995.
3. Similar habitats (sites) with richer regional colonization pools exhibited proportionally richer local communities in terms of number of species, total density and total biomass of individuals. Furthermore, no negative relationships were found between density and biomass of each of the most common species and local species richness.
4. Results of dynamic regression models (applied to the above-mentioned species) suggest an absence of strong competition between all pairs of species.
5. The evidence on lack of density compensation for species-poor communities and absence of perceptible interspecific competition between species suggest that the communities studied are non-interactive.
6. Two main explanations can be advanced. First, the local abundance of species in the communities studied could be determined through differential responses to unpredictable environmental changes, rather than through biological interactions. Second, as a result of historical events, the communities studied are reduced in congeneric species which can limit, in turn, the influence of interspecific competition in structuring these communities.
7. These results underline the strong influence of regional processes in shaping local riverine fish communities and minimize the possible influence of species interactions in governing these communities.     

Random sampling, abundance–extinction dynamics and niche-filtering immigration constraints explain the generation of species richness gradients

Aim  The paradigm that species' patterns of distribution, abundance and coexistence are the result of adaptations of the species to their niches has recently been challenged by evidence that similar patterns may be generated by simple random processes. We argue here that a better understanding of macroecological patterns requires an integration of both ecological and neutral stochastic approaches. We demonstrate the utility of such an integrative approach by testing the sampling hypothesis in a species–energy relationship of forest bird species.
Location  A Mediterranean biome in Catalonia, Spain.
Methods  To test the sampling hypothesis we designed a metacommunity model that reproduces the stochastic sampling from a regional pool to predict local species richness variation. Four conceptually different sampling procedures were evaluated.
Results  We showed that stochastic sampling processes predicted a substantial part (over 40%) of the observed variation in species richness, but left considerable variation unexplained. This remaining variation in species richness may be better understood as the result of alternative ecological processes. First, the sampling model explained more variation in species richness when the probability that a species colonises a new locality was assumed to increase with its niche width, suggesting that ecological differences between species matter when it comes to explaining macroecological patterns. Second, extinction risk was significantly lower for species inhabiting high-energy regions, suggesting that abundance–extinction processes play a significant role in shaping species richness patterns.
Main conclusions  We conclude that species–energy relationships may not simply be understood as a result of either ecological or random sampling processes, but more likely as a combination of both.     

Dispersal ability links to cross‐scale species diversity patterns across the Eurasian Arctic tundra

Aim The role of dispersal in structuring biodiversity across spatial scales is controversial. If dispersal controls regional and local community assembly, it should also affect the degree of spatial species turnover as well as the extent to which regional communities are represented in local communities. Here we provide the first integrated assessment of relationships between dispersal ability and local‐to‐regional spatial aspects of species diversity across a large geographical area. Location Northern Eurasia. Methods Using a cross‐scale analysis covering local (0.64 m²) to continental (the Eurasian Arctic biome) scales, we compared slope parameters of the dissimilarity‐to‐distance relationship in species composition and the local‐to‐regional relationship in species richness among three plant‐like groups that differ in dispersal ability: lichens with the highest dispersal ability; mosses and moss allies with intermediate dispersal ability; and seed plants with the lowest dispersal ability. Results Diversity patterns generally differed between the three groups according to their dispersal ability, even after controlling for niche‐based processes. Increasing dispersal ability is linked to decreasing spatial species turnover and an increasing ratio of local to regional species richness. All comparisons supported our expectations, except for the slope of the local‐to‐regional relationship in species richness for mosses and moss allies which was not significantly steeper than that of seed plants. Main conclusions The negative link between dispersal ability and spatial species turnover and the corresponding positive link between dispersal ability and the ratio of local‐to‐regional species richness support the idea that dispersal affects community structure and diversity patterns across spatial scales.     

Community assembly time and the relationship between local and regional species richness

Many previous studies have assumed that a linear relationship between local and regional species richness indicates that communities are limited by regional processes, while a saturating relationship suggests that species interactions restrict local richness. We show theoretically that the relationship between local and regional richness changes in a consistent fashion with assembly time in interacting communities. Communities show saturation in their early assembly stages because only a subset of the regional pool may colonize a locality. At intermediate assembly times, communities will appear unsaturated until significant competitive exclusion occurs. Finally, when communities reach equilibrium, we found saturation as a result of resource competition resulting in the dominance of a limited number of species. We show that habitat size and species fecundity are important in determining the time needed for the community to reach equilibrium and thus affect the relationship between local and regional species richness. Our results suggest the number of coexisting species is a function of local and regional processes whose relative influences might vary over time and that research using the relationship between local and regional species richness to infer mechanisms limiting species richness must have knowledge of the assembly time of the community.     

Evolution of termite functional diversity: analysis and synthesis of local ecological and regional influences on local species richness

Aim To (1) describe termite functional diversity patterns across five tropical regions using local species richness sampling of standardized areas of habitat; (2) assess the relative importance of environmental factors operating at different spatial and temporal scales in influencing variation in species representation within feeding groups and functional taxonomic groups across the tropics; (3) achieve a synthesis to explain the observed patterns of convergence and divergence in termite functional diversity that draws on termite ecological and biogeographical evidence to‐date, as well as the latest evidence for the evolutionary and distributional history of tropical rain forests. Location Pantropical. Methods A pantropical termite species richness data set was obtained through sampling of eighty‐seven standardized local termite diversity transects from twenty‐nine locations across five tropical regions. Local‐scale, intermediate‐scale and large‐scale environmental data were collected for each transect. Standardized termite assemblage and environmental data were analysed at the levels of whole assemblages and feeding groups (using components of variance analysis) and at the level of functional taxonomic groups (using correspondence analysis and canonical correspondence analysis). Results Overall species richness of local assemblages showed a greater component of variation attributable to local habitat disturbance level than to region. However, an analysis accounting for species richness across termite feeding groups indicated a much larger component of variation attributable to region. Mean local assemblage body size also showed the greater overall significance of region compared with habitat type in influencing variation. Ordination of functional taxonomic group data revealed a primary gradient of variation corresponding to rank order of species richness within sites and to mean local species richness within regions. The latter was in the order: Africa > south America > south‐east Asia > Madagascar > Australia. This primary gradient of species richness decrease can be explained by a decrease in species richness of less dispersive functional taxonomic groups feeding on more humified food substrates such as soil. Hence, the transects from more depauperate sites/regions were dominated by more dispersive functional taxonomic groups feeding on less humified food substrates such as dead wood. Direct gradient analysis indicated that ‘region’ and other large‐scale factors were the most important in explaining patterns of local termite functional diversity followed by intermediate‐scale geographical and site variables and, finally, local‐scale ecological variables. Synthesis and main conclusions Within regions, centres of termite functional diversity lie in lowland equatorial closed canopy tropical forests. Soil feeding termite evolution further down food substrate humification gradients is therefore more likely to have depended on the long‐term presence of this habitat. Known ecological and energetic constraints upon contemporary soil feeders lend support for this hypothesis. We propose further that the anomalous distribution of termite soil feeder species richness is partly explained by their generally very poor dispersal abilities across oceans. Evolution, radiation and dispersal of soil feeder diversity appears to have been largely restricted to what are now the African and south American regions. The inter‐regional differences in contemporary local patterns of termite species richness revealed by the global data set point to the possibility of large differences in consequent ecosystem processes in apparently similar habitats on different continents.     

The macroecology of reef fish agonistic behaviour

Understanding the interplay between processes operating at large and small spatiotemporal scales in shaping biotic interactions remains challenging. Recent studies illustrate how phenotypic specialization, species life-history traits and/or resource partitioning recurrently underlie the structure of mutualistic interactions in terrestrial ecosystems along large latitudinal gradients of biodiversity. However, we know considerably less about how local processes interact with large-scale patterns of biodiversity in modulating biotic interactions in the marine realm. Considering agonistic behaviour as a proxy for contest competition, we empirically investigate whether the structure of reef fish agonistic interactions is conserved across a 34 000-km longitudinal gradient of biodiversity. By sampling coral reefs using standardized remote underwater video, we found recurrent patterns of fish agonistic behaviour in disparate communities distributed across five biogeographic provinces of the Pacific and Atlantic oceans. While the sheer number of species increases with regional richness, the number of aggressive disputes at the habitat scale is similar across communities. We then combined generalized linear models and network theory to reveal that, the emergent structure of local agonistic networks is not modular but instead recurrently display a nested structure, with a core of highly interactive site-attached herbivores of the Pomacentridae family. Therefore, despite the increase in the number of species involved in agonistic interactions toward speciose communities, the network structure is conserved along the longitudinal richness gradient because local disputes are mostly driven by closely-related, functionally-similar species. These findings suggest that evolutionary and local processes interact in modulating reef fish agonistic behaviour and that fine-scale niche-partitioning can structure the ecological networks in marine ecosystems.     

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.