Abstract. It is often claimed that we do not understand the forces driving the global diversity gradient. However, an extensive literature suggests that contemporary climate constrains terrestrial taxonomic richness over broad geographic extents. Here, we review the empirical literature to examine the nature and form of the relationship between climate and richness. Our goals were to document the support for the climatically based energy hypothesis, and within the constraints imposed by correlative analyses, to evaluate two versions of the hypothesis: the productivity and ambient energy hypotheses. Focusing on studies extending over 800 km, we found that measures of energy, water, or water-energy balance explain spatial variation in richness better than other climatic and non-climatic variables in 82 of 85 cases. Even when considered individually and in isolation, water/ energy variables explain on average over 60% of the variation in the richness of a wide range of plant and animal groups. Further, water variables usually represent the strongest predictors in the tropics, subtropics, and warm temperate zones, whereas energy variables (for animals) or water-energy variables (for plants) dominate in high latitudes. We conclude that the interaction between water and energy, either directly or indirectly (via plant productivity), provides a strong explanation for globally extensive plant and animal diversity gradients, but for animals there also is a latitudinal shift in the relative importance of ambient energy vs. water moving from the poles to the equator. Although contemporary climate is not the only factor influencing species richness and may not explain the diversity pattern for all taxonomic groups, it is clear that understanding water-energy dynamics is critical to future biodiversity research. Analyses that do not include water-energy variables are missing a key component for explaining broad-scale patterns of diversity.

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Broad-scale variation in taxonomic richness is strongly correlated with climate. Many mechanisms have been hypothesized to explain these patterns; however, testable predictions that would distinguish among them have rarely been derived. Here, we examine several prominent hypotheses for climate-richness relationships, deriving and testing predictions based on their hypothesized mechanisms. The Ôenergy-richness hypothesisÕ (also called the Ômore individuals hypothesisÕ ) postulates that more productive areas have more individuals and therefore more species. More productive areas do often have more species, but extant data are not consistent with the expected causal relationship from energy to numbers of individuals to numbers of species. We reject the energy-richness hypothesis in its standard form and consider some proposed modifications. The Ôphysiological tolerance hypothesisÕ postulates that richness varies according to the tolerances of individual species for different sets of climatic conditions. This hypothesis predicts that more combinations of physiological parameters can survive under warm and wet than cold or dry conditions. Data are qualitatively consistent with this prediction, but are inconsistent with the prediction that species should fill climatically suitable areas. Finally, the Ôspeciation rate hypothesisÕ postulates that speciation rates should vary with climate, due either to faster evolutionary rates or stronger biotic interactions increasing the opportunity for evolutionary diversification in some regions. The biotic interactions mechanism also has the potential to amplify shallower, underlying gradients in richness. Tests of speciation rate hypotheses are few (to date), and their results are mixed.

Aim We surveyed the empirical literature to determine how well six diversity hypotheses account for spatial patterns in species richness across varying scales of grain and extent.Location Worldwide.Methods We identified 393 analyses ('cases') in 297 publications meeting our criteria. These criteria included the requirement that more than one diversity hypothesis was tested for its relationship with species richness. We grouped variables representing the hypotheses into the following 'correlate types': climate/ productivity, environmental heterogeneity, edaphics/nutrients, area, biotic interactions and dispersal/history (colonization limitation or other historical or evolutionary effect). For each case we determined the 'primary' variable: the one most strongly correlated with taxon richness. We defined 'primacy' as the proportion of cases in which each correlate type was represented by the primary variable, relative to the number of times it was studied. We tested for differences in both primacy and mean coefficient of determination of the primary variable between the hypotheses and between categories of five grouping variables: grain, extent, taxon (animal vs. plant), habitat medium (land vs. water) and insularity (insular vs. connected).Results Climate/productivity had the highest overall primacy, and environmental heterogeneity and dispersal/history had the lowest. Primacy of climate/ productivity was much higher in large-grain and large-extent studies than at smaller scales. It was also higher on land than in water, and much higher in connected systems than in insular ones. For other hypotheses, differences were less pronounced. Throughout, studies on plants and animals showed similar patterns. Coefficients of determination of the primary variables differed little between hypotheses and across the grouping variables, the strongest effects being low means in the smallest grain class and for edaphics/nutrients variables, and a higher mean for water than for land in connected systems but vice versa in insular systems. We highlight areas of data deficiency. Main conclusionsOur results support the notion that climate and productivity play an important role in determining species richness at large scales, particularly for non-insular, terrestrial habitats. At smaller extents and grain sizes, the primacy of the different types of correlates appears to differ little from null expectation. In our analysis, dispersal/history is rarely the best correlate of species richness, but this may reflect the difficulty of incorporating historical factors into regression models, and the collinearity between past and current climates. Our findings are consistent with the view that climate determines the capacity for species richness. However, its influence is less evident at smaller spatial scales, probably because (1) studies small in extent tend to sample little climatic range, and (2) at large

Species richness, the simplest index of biodiversity, varies greatly over broad spatial scales. Richness-climate relationships often account for >80% of the spatial variance in richness. However, it has been suggested that richness-climate relationships differ significantly among geographic regions and that there is no globally consistent relationship. This study investigated the global patterns of species and family richness of angiosperms in relation to climate. We found that models relating angiosperm richness to mean annual temperature, annual water deficit, and their interaction or models relating richness to annual potential evapotranspiration and water deficit are both globally consistent and very strong and are independent of the diverse evolutionary histories and functional assemblages of plants in different parts of the world. Thus, effects of other factors such as evolutionary history, postglacial dispersal, soil nutrients, topography, or other climatic variables either must be quite minor over broad scales (because there is little residual variation left to explain) or they must be strongly collinear with global patterns of climate. The correlations shown here must be predicted by any successful hypothesis of mechanisms controlling richness patterns.

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In population ecology, there has been a fundamental controversy about the relative importance of competition-driven (densitydependent) population regulation vs. abiotic influences such as temperature and precipitation. The same issue arises at the community level; are population sizes driven primarily by changes in the abundances of cooccurring competitors (i.e., compensatory dynamics), or do most species have a common response to environmental factors? Competitive interactions have had a central place in ecological theory, dating back to Gleason, Volterra, Hutchison and MacArthur, and, more recently, Hubbell's influential unified neutral theory of biodiversity and biogeography. If competitive interactions are important in driving year-to-year fluctuations in abundance, then changes in the abundance of one species should generally be accompanied by compensatory changes in the abundances of others. Thus, one necessary consequence of strong compensatory forces is that, on average, species within communities will covary negatively. Here we use measures of community covariance to assess the prevalence of negative covariance in 41 natural communities comprising different taxa at a range of spatial scales. We found that species in natural communities tended to covary positively rather than negatively, the opposite of what would be expected if compensatory dynamics were important. These findings suggest that abiotic factors such as temperature and precipitation are more important than competitive interactions in driving year-to-year fluctuations in species abundance within communities.biological interactions ͉ community dynamics ͉ negative covariance ͉ neutral models ͉ zero-sum

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology. Abstract.The contemporary literature accepts that disturbance strongly influences patterns of species diversity, and that the relationship is peaked, with a maximum at intermediate levels of disturbance. We tested this hypothesis using a compilation of published species diversity-disturbance relationships that were gleaned from a literature search of papers published from 1985 through 1996 and from references therein. We identified 116 species richness-, 53 diversity-, and 28 evenness-disturbance relationships in the literature, which we grouped according to shape of relationship (nonsignificant, peaked, negative monotonic, positive monotonic, or U-shaped). We tested the relationships between the strength and shapes of these relationships and attributes of the community, disturbance, and sampling and study design. Nonsignificant relationships were the most common, comprising 35% of richness, 28% of diversity, and 50% of evenness studies. Peaked responses were reported in only 16% of richness, 19% of diversity, and 11% of evenness cases. Explained variation in the three measures of diversity was variable among studies but averaged -50%. It was higher when few samples and few disturbance levels were examined and when organisms within the samples were not exhaustively censused, suggesting that procedural artifact contributes to these relationships. Explained variation was also higher in studies in which disturbance was measured as a gradient of time passed since the last disturbance (mean r2 = 61%), vs. studies of spatial variation in richness (mean r2 = 42%). Peaked richness relationships had the greatest odds of being observed when sampled area and actual evapotranspiration were small, when disturbances were natural rather than anthropogenic in origin, and when few disturbance levels were examined. Thus, on average, diversity-disturbance relationships do not have consistently high r2 and are not as consistently peaked as the contemporary consensus would suggest. It has been hypothesized that in communities where post-disturbance succession is not driven by competitive hierarchies, disturbance intensity and frequency may have little effect on species richness (Reice 1985, Chesson and Huntly 1997). Huston (1979, 1994) suggests productivity may influence the shape of the diversity-disturbance relationship. Unpredictable, severe, episodic disturbances may have greater effects on richness than do predictable, moderate events to which communities may adapt (Reice et al. 1990). Wootton's (1998) models predict that, in multitrophic systems, IDH should hold for species competing ...