Large-scale Patterns Of Species Richness Research Articles

Species richness patterns and the determinants of larch forests in China

Larch forests are important for species diversity, as well as soil and water conservation in mountain regions. In this study, we determined large-scale patterns of species richness in larch forests and identified the factors that drive these patterns. We found that larch forest species richness was high in southern China and low in northern China, and that patterns of species richness along an elevational gradient depend on larch forest type. In addition, we found that patterns of species richness in larch forests are best explained by contemporary climatic factors. Specifically, mean annual temperature and annual potential evapotranspiration were the most important factors for species richness of tree and shrub layers, while mean temperature of the coldest quarter and anomaly of annual precipitation from the Last Glacial Maximum to the present were the most important for that of herb layer and the whole community. Community structural factors, especially stand density, are also associated with the species richness of larch forests. Our findings that species richness in China's larch forests is mainly affected by energy availability and cold conditions support the ambient energy hypothesis and the freezing tolerance hypothesis.

The Importance of Including Soil Properties When Disentangling the Drivers of Species Richness: The Case of the Alpine Genus Saxifraga L. in China

Despite the numerous studies on the large-scale patterns of species richness, the spatial variation and determinants of species richness for alpine plant are still an outstanding question and critical to future biodiversity conservation. The genus Saxifraga is a typical alpine plant group with high species richness in the Himalaya-Hengduan mountain regions, China. We performed simple regression models and variance partitioning to assess the importance of different factors, especially soil-related ones, in driving Saxifraga richness patterns. The results showed that environmental energy, habitat heterogeneity, and soil heterogeneity together dominated the spatial variation of species richness. The coarse fragments volume of soil, elevation range, and soil heterogeneity, are positively related to Saxifraga richness. Soil slightly outperforms habitat heterogeneity in predicting the spatial variation of Saxifraga species richness with an explanatory power of 39.3% and 36.6%, respectively. Environmental energy, such as the maximum temperature of the warmest quarter, is negatively correlated with species richness and explains 44.8% of spatial variation of Saxifraga richness. Multiple regression models, including three variables, each representing energy, soil, and habitat heterogeneity, can only explain 53.1% variation of species richness. Variance partitioning outscored 26% of the shared effects of the three variables, while the independent effect of each variable is less than 10%. These results indicated the energy, soil, and habitat heterogeneity together are primary determinants of the spatial variation of Saxifraga species richness. However, there are probably other hidden factors predicting species richness variation due to the low explanatory power of the multiple regression models. Our study emphasizes the significance of soil properties in determining species richness patterns in China, especially for the alpine plant groups. The negative association of species richness with temperature suggests a potential threat of alpine biodiversity loss in HHM from future warming.

Scavenging in the Anthropocene: Human impact drives vertebrate scavenger species richness at a global scale

Understanding the distribution of biodiversity across the Earth is one of the most challenging questions in biology. Much research has been directed at explaining the species latitudinal pattern showing that communities are richer in tropical areas; however, despite decades of research, a general consensus has not yet emerged. In addition, global biodiversity patterns are being rapidly altered by human activities. Here, we aim to describe large-scale patterns of species richness and diversity in terrestrial vertebrate scavenger (carrion-consuming) assemblages, which provide key ecosystem functions and services. We used a worldwide dataset comprising 43 sites, where vertebrate scavenger assemblages were identified using 2,485 carcasses monitored between 1991 and 2018. First, we evaluated how scavenger richness (number of species) and diversity (Shannon diversity index) varied among seasons (cold vs. warm, wet vs. dry). Then, we studied the potential effects of human impact and a set of macroecological variables related to climatic conditions on the scavenger assemblages. Vertebrate scavenger richness ranged from species-poor to species rich assemblages (4-30 species). Both scavenger richness and diversity also showed some seasonal variation. However, in general, climatic variables did not drive latitudinal patterns, as scavenger richness and diversity were not affected by temperature or rainfall. Rainfall seasonality slightly increased the number of species in the community, but its effect was weak. Instead, the human impact index included in our study was the main predictor of scavenger richness. Scavenger assemblages in highly human-impacted areas sustained the smallest number of scavenger species, suggesting human activity may be overriding other macroecological processes in shaping scavenger communities. Our results highlight the effect of human impact at a global scale. As species-rich assemblages tend to be more functional, we warn about possible reductions in ecosystem functions and the services provided by scavengers in human-dominated landscapes in the Anthropocene.

Spatial Patterns and Determinants of the Diversity of Hemipteran Insects in the Qinghai-Tibetan Plateau

Large-scale patterns of species richness is an important issue in biogeography and ecology. The Qinghai-Tibetan Plateau (QTP) is a biodiversity hotspot in the world, which has an important status in the zoogeographical realms. Here, we analyzed the diversity patterns of Hemipteran insects in the QTP, and tested whether the patterns can be jointly explained by modern environmental as well as historical factors. A comprehensive geographic distribution dataset consisting of 1166 Hemipteran species, which belong to 510 genera and 53 families, was compiled and used in our analyses. Patterns of richness were mapped into a grid-based map with a spatial resolution of 0.5°x 0.5°. An unbalanced diversity pattern of the Hemipteran insects in the QTP was presented, with more species in the eastern and southern parts of the plateau, while few species in the northern and main surface of the plateau. The northwestern Sichuan, the southern Gansu, the southeastern Tibet, the northwestern Yunnan and the eastern Qinghai were identified as diversity hotspots of species richness. Further analyses based on General linear models and Random Forest indicated that the diversity patterns of Hemipteran insects were influenced by both contemporary environmental factors and historical factors (e.g. habitat heterogeneity, climate stability, energy availability). Specifically, the species richness patterns of all Hemipteran insects in the QTP have been mainly affected by elevation range, temperature annual range, min temperature of coldest month, mean temperature of coldest quarter and the temperature change since the Last Glacial Maximum. In contrast, the water-related variables have relatively small effects on species richness. In addition, although habitat heterogeneity was indicated the most important factor for different suborders of Hemiptera, the climate stability was another dominate factor for Heteroptera and Auchenorrhyncha, while Sternorrhyncha was more affected by historical climate change.

Integrating multiple species criteria and species hypotheses in subterranean biology

Decades of debates around the species problem have resulted in the emergence of a unified species concept with multiple criteria to delimit species taxa. Many biologists now agree to consider species as separately evolving segments of metapopulation lineages (i.e. the species concept), and to consider species taxa (i.e. the elementary units used in biodiversity science) as scientific hypotheses of separately evolving entities. In this framework, sets of species hypotheses are generated using different criteria (i.e. morphological distinguishability, genetic isolation) that mirror the properties expressed by species at different times and sequential orders during the extended and heterogeneous process of speciation. This conceptual and methodological advance in taxonomy has several implications for biodiversity science. First, species taxa represent a heterogeneous set of hypotheses whose properties are contingent on the heterogeneous, continuous and extended nature of speciation. Second, species databases need to integrate information on the diverse properties of species by attributing specimens to multiple species hypotheses generated using different delimitation criteria. Third, biodiversity science at large can provide novel insights into biodiversity processes by incorporating multiple species hypotheses into the analysis of biodiversity patterns. Here, we show how these implications have been taken into account by subterranean biologists. First, we briefly review the criteria and methods used to delimit species in subterranean biology and the diverse sets of species hypotheses they generated. Second, we present a new generation of species occurrence databases that integrate different species criteria and hypotheses while fully respecting the scientific rigor of taxonomy. Last, we show how incorporating multiple species hypotheses into macroecological analyses of European groundwater fauna bolsters our under­standing of the factors shaping large-scale patterns of species richness and geographic range size.

Trophic interactions among vertebrate guilds and plants shape global patterns in species diversity.

Trophic interactions play critical roles in structuring biotic communities. Understanding variation in trophic interactions among systems provides important insights into biodiversity maintenance and conservation. However, the relative importance of bottom-up versus top-down trophic processes for broad-scale patterns in biodiversity is poorly understood. Here, we used global datasets on species richness of vascular plants, mammals and breeding birds to evaluate the role of trophic interactions in shaping large-scale diversity patterns. Specifically, we used non-recursive structural equation models to test for top-down and bottom-up forcing of global species diversity patterns among plants and trophic guilds of mammals and birds (carnivores, invertivores and herbivores), while accounting for extrinsic environmental drivers. The results show that trophic linkages emerged as being more important to explaining species richness than extrinsic environmental drivers. In particular, there were strong, positive top-down interactions between mammal herbivores and plants, and moderate to strong bottom-up and/or top-down interactions between herbivores/invertivores and carnivores. Estimated trophic interactions for separate biogeographical regions were consistent with global patterns. Our findings demonstrate that, after accounting for environmental drivers, large-scale species richness patterns in plant and vertebrate taxa consistently support trophic interactions playing a major role in shaping global patterns in biodiversity. Furthermore, these results suggest that top-down forces often play strong complementary roles relative to bottom-up drivers in structuring biodiversity patterns across trophic levels. These findings underscore the importance of integrating trophic forcing mechanisms into studies of biodiversity patterns.

The influence of habitat heterogeneity and latitude on gamma diversity of the Nearctic Simuliidae, a ubiquitous group of stream-dwelling insects.

Among the most prominent, large-scale patterns of species richness are the increases in richness with decreasing latitude and with increasing habitat heterogeneity. Using the stream-dwelling larval and pupal stages of North American black flies (Diptera: Simuliidae), we address 3 broad questions about species richness: (i) Does a significant latitude-richness relationship exist? (ii) How does habitat heterogeneity influence gamma diversity? (iii) What is the sign (positive or negative) of the latitude-richness and the heterogeneity-richness relationships? We found no evidence that habitat heterogeneity influences gamma diversity. The estimated peak species richness for black flies in North America was at 50-53°N, which also corresponds with peak generic richness. All plesiomorphic, extant lineages of the Simuliidae in the Western Hemisphere are found in cool mountainous environments of North America, suggesting that peak richness at 50-53°N might be a signature of this phylogenetic pattern and a reflection of underlying historical processes.

Effects of Environment and Space on Species Turnover of Woody Plants across Multiple Forest Dynamic Plots in East Asia.

Species turnover is fundamental for understanding the mechanisms that influence large-scale species richness patterns. However, few studies have described and interpreted large-scale spatial variation in plant species turnover, and the causes of this variation remain elusive. In addition, the determinants of species turnover depend on the dispersal ability of growth forms. In this study, we explored the large-scale patterns of woody species turnover across the latitude gradient based on eight large stem-mapping plots (covering 184 ha forest) in East Asia. The patterns of woody species turnover increased significantly with increasing latitude differences in East Asia. For overall woody species, environment explained 36.30, 37.20, and 48.48% of the total variance in Jaccard’s (βj), Sorenson’s, (βs), and Simpson’s dissimilarity (βsim). Spatial factors explained 47.92, 48.39, and 41.38% of the total variance in βj, βs, and βsim, respectively. The effects of pure spatial and spatially structured environments were stronger than pure environmental effects for overall woody species. Our results support the hypothesis that the effect of neutral processes on woody species turnover is more important than the effect of the environment. Neutral processes explained more variation for turnover of tree species, and environmental factors explained more variation for the turnover of shrub species on a large scale. Therefore, trees and shrubs should be subjected to different protection strategies in future biodiversity conservation efforts.

Topographic heterogeneity and temperature amplitude explain species richness patterns of birds in the Qinghai-Tibetan Plateau.

Large-scale patterns of species richness have gained much attention in recent years; however, the factors that drive high species richness are still controversial in local regions, especially in highly diversified montane regions. The Qinghai–Tibetan Plateau (QTP) and the surrounding mountains are biodiversity hot spots due to a high number of endemic montane species. Here, we explored the factors underlying this high level of diversity by studying the relationship between species richness and environmental variables. The richness patterns of 758 resident bird species were summarized at the scale of 1°×1° grid cell at different taxonomic levels (order, family, genus, and species) and in different taxonomic groups (Passeriformes, Galliformes, Falconiformes, and Columbiformes). These richness patterns were subsequently analyzed against habitat heterogeneity (topographical heterogeneity and land cover), temperature amplitude (annual temperature, annual precipitation, precipitation seasonality, and temperature seasonality) and a vegetation index (net primary productivity). Our results showed that the highest richness was found in the southeastern part of the QTP, the eastern Himalayas. The lowest richness was observed in the central plateau of the QTP. Topographical heterogeneity and temperature amplitude are the primary factors that explain overall patterns of species richness in the QTP, although the specific effect of each environmental variable varies between the different taxonomic groups depending on their own evolutionary histories and ecological requirements. High species richness in the southeastern QTP is mostly due to highly diversified habitat types and temperature zones along elevation gradients, whereas the low species richness in the central plateau of the QTP may be due to environmental and energetic constraints, as the central plateau is harsh environment.

Unimodal latitudinal pattern of land-snail species richness across northern Eurasian lowlands.

Large-scale patterns of species richness and their causes are still poorly understood for most terrestrial invertebrates, although invertebrates can add important insights into the mechanisms that generate regional and global biodiversity patterns. Here we explore the general plausibility of the climate-based “water-energy dynamics” hypothesis using the latitudinal pattern of land-snail species richness across extensive topographically homogeneous lowlands of northern Eurasia. We established a 1480-km long latitudinal transect across the Western Siberian Plain (Russia) from the Russia-Kazakhstan border (54.5°N) to the Arctic Ocean (67.5°N), crossing eight latitudinal vegetation zones: steppe, forest-steppe, subtaiga, southern, middle and northern taiga, forest-tundra, and tundra. We sampled snails in forests and open habitats each half-degree of latitude and used generalized linear models to relate snail species richness to climatic variables and soil calcium content measured in situ. Contrary to the classical prediction of latitudinal biodiversity decrease, we found a striking unimodal pattern of snail species richness peaking in the subtaiga and southern-taiga zones between 57 and 59°N. The main south-to-north interchange of the two principal diversity constraints, i.e. drought stress vs. cold stress, explained most of the variance in the latitudinal diversity pattern. Water balance, calculated as annual precipitation minus potential evapotranspiration, was a single variable that could explain 81.7% of the variance in species richness. Our data suggest that the “water-energy dynamics” hypothesis can apply not only at the global scale but also at subcontinental scales of higher latitudes, as water availability was found to be the primary limiting factor also in this extratropical region with summer-warm and dry climate. A narrow zone with a sharp south-to-north switch in the two main diversity constraints seems to constitute the dominant and general pattern of terrestrial diversity across a large part of northern Eurasia, resulting in a subcontinental diversity hotspot of various taxa in this zone.

Patterns of species diversity and phylogenetic structure of vascular plants on the Qinghai‐Tibetan Plateau

Large-scale patterns of species richness and the underlying mechanisms regulating these patterns have long been the central issues in biogeography and macroecology. Phylogenetic community structure is a result of combined effects of contemporary ecological interactions, environmental filtering, and evolutionary history, and it links community ecology with biogeography and trait evolution. The Qinghai-Tibetan Plateau provides a good opportunity to test the influence of contemporary climate on shaping species richness because of its unique geological history, cold climate, and high biodiversity. In this study, based on high-resolution distributions of ˜9000 vascular plant species, we explored how species richness and phylogenetic structure of vascular plants correlate with climates on the highest (and species rich) plateau on the Earth. The results showed that most of the vascular plants were distributed on the eastern part of the plateau; there was a strong association between species richness and climate, even after the effects of habitat heterogeneity were controlled. However, the responses of richness to climate remarkably depended on life-forms. Richness of woody plants showed stronger climatic associations than that of herbaceous plants; energy and water availability together regulated richness pattern of woody plants; whereas water availability predominantly regulated richness pattern of herbaceous plants. The phylogenetic structure of vascular species clustered in most areas of the plateau, suggesting that rapid speciation and environment filtering dominated the assembly of communities on the plateau. We further propose that biodiversity conservation in this area should better take into account ecological features for different life-forms and phylogenetic lineages.

Geographical patterns in the beta diversity of China's woody plants: the influence of space, environment and range size

Beta diversity (i.e. species turnover rate across space) is fundamental for understanding mechanisms controlling large‐scale species richness patterns. However, the influences on beta diversity are still a matter of debate. In particular, the relative role of environmental and spatial processes (e.g. environmental niche versus dispersal limitation of species) remains elusive, and the influence of species range size has been poorly tested. Here, using distribution maps of 11 405 woody species in China (ca 9.6 × 106 km2), we investigated 1) the geographical and directional patterns of beta diversity for all woody species and species with different range sizes, and 2) compared the effects of environmental and spatial processes on these patterns. Beta diversity was calculated as the decay of similarity in species composition with increasing distance. Variables representing environmental energy, water availability, climatic seasonality, habitat heterogeneity and human activities were used to evaluate the effects of environmental processes, while spatial distance was used to assess the influence of spatial processes. The results indicated significant directional patterns of beta diversity: the similarity decay along the latitudinal gradient was 1.6–2.3 times faster than that along the longitudinal gradient. Beta diversity also increased with the decrease of species range size. As compared with spatial processes, environmental processes had stronger effects on longitudinal beta diversity and on the beta diversity of widely‐ranged species. This was opposite to the larger influence of spatial processes on latitudinal beta diversity and the beta diversity of narrowly‐ranged species. These results suggest that the distributions of narrowly‐ranged woody species in China may have not reached equilibrium with their environmental niches due to dispersal limitation induced by China's topography and/or their low dispersal ability. The projected rapid climatic changes will likely endanger such species. Species dispersal processes should be taken into account in future conservation strategies in China.

Disentangling the relative effects of ambient energy, water availability, and energy–water balance on pteridophyte species richness at a landscape scale in China

Understanding what factors generate geographic variation in species richness is a fundamental goal of ecology and biogeography. Water and energy are considered as the major environmental factors influencing large-scale patterns of species richness, but their roles vary among taxa and regions. Pteridophytes are an ideal group of organisms for examining the relationship between species richness and their environment because the distribution of pteridophytes is usually in equilibrium with contemporary climate to a greater degree than those of seed plants and most terrestrial vertebrates partly due to the lightness of their spores, which is highly capable of long-distance dispersal by wind, and partly due to their single-spore reproduction strategy. Using correlation and regression analyses and structural equation modeling technique, we examine the relationship of pteridophyte species richness in 151 localities from across China with environmental factors representing energy, water, and energy–water balance. We found that pteridophyte species richness is correlated to water availability more strongly than to ambient energy. Furthermore, we found that of all environmental variables considered, energy–water balance has played the most important role in regulating pteridophyte species richness gradients in China.

Patterns, determinants and models of woody plant diversity in China

What determines large-scale patterns of species richness remains one of the most controversial issues in ecology. Using the distribution maps of 11 405 woody species in China, we compared the effects of habitat heterogeneity, human activities and different aspects of climate, particularly environmental energy, water-energy dynamics and winter frost, and explored how biogeographic affinities (tropical versus temperate) influence richness-climate relationships. We found that the species richness of trees, shrubs, lianas and all woody plants strongly correlated with each other, and more strongly correlated with the species richness of tropical affinity than with that of temperate affinity. The mean temperature of the coldest quarter was the strongest predictor of species richness, and its explanatory power for species richness was significantly higher for tropical affinity than for temperate affinity. These results suggest that the patterns of woody species richness mainly result from the increasing intensity of frost filtering for tropical species from the equator/lowlands towards the poles/highlands, and hence support the freezing-tolerance hypothesis. A model based on these results was developed, which explained 76-85% of species richness variation in China, and reasonably predicted the species richness of woody plants in North America and the Northern Hemisphere.

Water limitation prevails over energy in European diversity gradients of sheetweb spiders (Araneae: Linyphiidae)

Across large spatial scales, species richness in many taxa is mainly determined by climatic variables. However, some of the mechanisms behind large-scale patterns of species richness and abundance are expected to act on the community level, and on a smaller scale than the resolution of the data commonly used for deriving these patterns. We studied the distribution of sheetweb spiders (Araneae: Linyphiidae) across Europe using both standardised samples from local habitats and large-scale inventories. In the first approach, we analysed species and individual numbers from standardised pitfall traps from 17 locations distributed from northern Sweden to Spain and Greece. We also calculated Simpson indices to correct for different activity densities. Additionally, we analysed whether diversity of sheetweb spiders is sensitive to habitat type. In the second approach, we investigated the known sheetweb spider species richness of European countries and large islands. In both datasets, species richness of sheetweb spiders reached a maximum at about 551N and declined towards the Mediterranean and the Arctic. In addition, species richness decreased from western (maritime) to eastern (continental) regions. Also Simpson diversity showed a hump-shaped relation to latitude, and was the only variable influenced by habitat type. In contrast to species richness and diversity, activity density increased monotonously with latitude. Towards the north, sheetweb spiders are likely limited by energy availability, towards the south, water limitation as well as bottom-up and top-down community interactions may reduce sheetweb spiders. Accordingly, their diversity does not follow the pattern of vertebrates, large insects and aquatic organisms, which depend more strongly on energy availability.

Global concordance in diversity patterns of vascular plants and terrestrial vertebrates

The factors that determine large-scale patterns of species richness are poorly understood. In particular, biologists have not determined the relative roles of taxon-specific characteristics that influence diversification and distribution, and region-specific features that promote and constrain diversity. We show that the numbers of species of vascular plants and of four terrestrial vertebrate taxa (mammals, birds, reptiles and amphibians) vary in parallel across 296 geographic areas covering most of the globe, even after accounting for sample area, climate, topographic heterogeneity and differences between continents. Thus, a common set of regional characteristics and processes appears to shape patterns of species richness in a diverse set of taxa, despite substantial differences in their biological traits.

Global Patterns of Diversification and Species Richness in Amphibians

Geographic patterns of species richness ultimately arise through the processes of speciation, extinction, and dispersal, but relatively few studies consider evolutionary and biogeographic processes in explaining these diversity patterns. One explanation for high tropical species richness is that many species-rich clades originated in tropical regions and spread to temperate regions infrequently and more recently, leaving little time for species richness to accumulate there (assuming similar rates of diversification in temperate and tropical regions). However, the major clades of anurans (frogs) and salamanders may offer a compelling counterexample. Most salamander families are predominately temperate in distribution, but the one primarily tropical clade (Bolitoglossinae) contains nearly half of all salamander species. Similarly, most basal clades of anurans are predominately temperate, but one largely tropical clade (Neobatrachia) contains approximately 96% of anurans. In this article, I examine patterns of diversification in frogs and salamanders and their relationship to large-scale patterns of species richness in amphibians. I find that diversification rates in both frogs and salamanders increase significantly with decreasing latitude. These results may shed light on both the evolutionary causes of the latitudinal diversity gradient and the dramatic but poorly explained disparities in the diversity of living amphibian clades.

EFFECTS OF REGIONAL VS. ECOLOGICAL FACTORS ON PLANT SPECIES RICHNESS: AN INTERCONTINENTAL ANALYSIS

Conclusions from past studies on the roles that historical and regional factors and contemporary and ecological factors have played in regulating large-scale patterns of species richness have been controversial. Conflicting past results were likely affected by differences in the range of environments analyzed and the scales of observation. Eastern North America and eastern Asia are ideal regions for examining the relative effects of historical and regional factors and contemporary and ecological factors on large-scale patterns of plant species richness because these two regions are closely matched in terms of climate and because their floras originated from the same paleoflora but have experienced different histories of development since the late Paleogene when climate cooling caused their separation. We report on a comprehensive data set of 471 floras ranging from 10 km2 to 4.7 x 10(6) km2 and spanning a wide range of climate and latitude (from 21 degrees to 55 degrees N) to examine whether the contribution of region relative to climate persists from small to large floras and increases from cooler to warmer climates. We found that eastern Asia is richer than eastern North America when sample area, maximum elevation, and climate are accounted for, that this difference diminishes toward higher latitudes, and that elevation plays a much stronger role in eastern Asia than in eastern North America. Our analysis reconciles contemporary/ecological and historical/regional explanations for species richness variation and helps explain why different conclusions have been reached by different authors working in the same geographical areas: the strength of the region effect itself varies with location and range of climatic conditions of the observations.

Does habitat use explain large scale species richness patterns of aquatic beetles in Europe?

Regularities in species richness are widely observed but controversy continues over its mechanistic explanation. Because richness patterns are usually a compound measure derived from taxonomically diverse species with different ecological requirements, these analyses may confound diverse causes of species numbers. Here we investigate species richness in the aquatic beetle fauna of Europe, separating major taxonomic groups and two major ecological types, species occurring in standing and running water bodies. We collated species distributions for 800+ species of water beetles in 15 regions across western Europe. Species number in any of these regions was related to three variables: total area size, geographic connectedness of the area, and latitude. Pooled species numbers were accurately predicted, but correlations were different for species associated with either running or standing water. The former were mostly correlated with latitude, while the latter were only correlated with the measure of connectedness or with area size. These differences were generally also observed in each of the four phylogenetically independent lineages of aquatic Coleoptera when analysed separately. We propose that effects of habitat, in this case possibly mediated by different long term persistence of running and standing water bodies, impose constraints at the population or local level which, if effective over larger temporal and spatial scales, determine global patterns of species richness.