Broad-scale Species Richness Patterns Research Articles

Genetic and species-level biodiversity patterns are linked by demography and ecological opportunity.

The processes that give rise to species richness gradients are not well understood, but may be linked to resource-based limits on the number of species a region can support. Ecological limits placed on regional species richness should also affect population demography, suggesting that these processes could also generate genetic diversity gradients. If true, we might better understand how broad-scale biodiversity patterns are formed by identifying the common causes of genetic diversity and species richness. We develop a hypothetical framework based on the consequences of regional variation in ecological limits set by resource availability and heterogeneity to simultaneously explain spatial patterns of species richness and neutral genetic diversity. Repurposing raw genotypic data spanning 38 mammal species sampled across 801 sites in North America, we show that estimates of genome-wide genetic diversity and species richness share spatial structure. Notably, species richness hotspots tend to harbor lower levels of within-species genetic variation. A structural equation model encompassing eco-evolutionary processes related to resource availability, habitat heterogeneity, and contemporary human disturbance supports the spatial patterns we detect. These results suggest broad-scale patterns of species richness and genetic diversity could both partly be caused by intraspecific demographic and evolutionary processes acting simultaneously across species.

A latitudinal phylogeographic diversity gradient in birds.

High tropical species diversity is often attributed to evolutionary dynamics over long timescales. It is possible, however, that latitudinal variation in diversification begins when divergence occurs within species. Phylogeographic data capture this initial stage of diversification in which populations become geographically isolated and begin to differentiate genetically. There is limited understanding of the broader implications of intraspecific diversification because comparative analyses have focused on species inhabiting and evolving in restricted regions and environments. Here, we scale comparative phylogeography up to the hemisphere level and examine whether the processes driving latitudinal differences in species diversity are also evident within species. We collected genetic data for 210 New World bird species distributed across a broad latitudinal gradient and estimated a suite of metrics characterizing phylogeographic history. We found that lower latitude species had, on average, greater phylogeographic diversity than higher latitude species and that intraspecific diversity showed evidence of greater persistence in the tropics. Factors associated with species ecologies, life histories, and habitats explained little of the variation in phylogeographic structure across the latitudinal gradient. Our results suggest that the latitudinal gradient in species richness originates, at least partly, from population-level processes within species and are consistent with hypotheses implicating age and environmental stability in the formation of diversity gradients. Comparative phylogeographic analyses scaled up to large geographic regions and hundreds of species can show connections between population-level processes and broad-scale species-richness patterns.

Using climate, energy, and spatial-based hypotheses to interpret macroecological patterns of North America chelonians

Our study investigates how factors, such as latitude, productivity, and several environmental variables, influence contemporary patterns of the species richness in North American turtles. In particular, we test several hypotheses explaining broad-scale species richness patterns on several species richness data sets: (i) total turtles, (ii) freshwater turtles only, (iii) aquatic turtles, (iv) terrestrial turtles only, (v) Emydidae, and (vi) Kinosternidae. In addition to spatial data, we used a combination of 25 abiotic variables in spatial regression models to predict species richness patterns. Our results provide support for multiple hypotheses related to broad-scale patterns of species richness, and in particular, hypotheses related to climate, productivity, water availability, topography, and latitude. In general, species richness patterns were positively associated with temperature, precipitation, diversity of streams, coefficient of variation of elevation, and net primary productivity. We also found that North America turtles follow the general latitudinal diversity gradient pattern (i.e., increasing species richness towards equator) by exhibiting a negative association with latitude. Because of the incongruent results among our six data sets, our study highlights the importance of considering phylogenetic constraints and guilds when interpreting species richness patterns, especially for taxonomic groups that occupy a myriad of habitats.

Linking diversity and distribution to understand biodiversity gradients and inform conservation assessments

Broad-scale patterns of species richness result from differential coexistence among species in distinct regions of the globe, determined by the species’ ranges and their properties such as size, shape and location. Thus, species richness and ranges are inherently linked. These two biodiversity features also yield primary information for conservation assessments. However, species richness and range size have been usually studied separately and no formal analytical link has been established. In my PhD thesis, I applied and extended a recently developed conceptual and methodological framework to study geographical association among species and similarity among sites. This range–diversity framework, along with stochastic simulation modelling, allowed me to jointly evaluate the relationship between diversity and distribution, to infer potential processes underlying composite patterns of phyllostomid bats, and to use this approach to inform conservation assessments for the Mexican avifauna. I highlight the need to explore composite patterns for understanding biodiversity patterns and show how combining diversity and distributional data can help describe complex biogeographical patterns, providing a transparent and explicit application for initial conservation assessments.

Global patterns of freshwater species diversity, threat and endemism.

AimGlobal-scale studies are required to identify broad-scale patterns in the distributions of species, to evaluate the processes that determine diversity and to determine how similar or different these patterns and processes are among different groups of freshwater species. Broad-scale patterns of spatial variation in species distribution are central to many fundamental questions in macroecology and conservation biology. We aimed to evaluate how congruent three commonly used metrics of diversity were among taxa for six groups of freshwater species.LocationGlobal.MethodsWe compiled geographical range data on 7083 freshwater species of mammals, amphibians, reptiles, fishes, crabs and crayfish to evaluate how species richness, richness of threatened species and endemism are distributed across freshwater ecosystems. We evaluated how congruent these measures of diversity were among taxa at a global level for a grid cell size of just under 1°.ResultsWe showed that although the risk of extinction faced by freshwater decapods is quite similar to that of freshwater vertebrates, there is a distinct lack of spatial congruence in geographical range between different taxonomic groups at this spatial scale, and a lack of congruence among three commonly used metrics of biodiversity. The risk of extinction for freshwater species was consistently higher than for their terrestrial counterparts.Main conclusionsWe demonstrate that broad-scale patterns of species richness, threatened-species richness and endemism lack congruence among the six freshwater taxonomic groups examined. Invertebrate species are seldom taken into account in conservation planning. Our study suggests that both the metric of biodiversity and the identity of the taxa on which conservation decisions are based require careful consideration. As geographical range information becomes available for further sets of species, further testing will be warranted into the extent to which geographical variation in the richness of these six freshwater groups reflects broader patterns of biodiversity in fresh water.

Environmental determinants of woody and herb plant species richness patterns in Great Britain

There is evidence that broad-scale species richness patterns of woody plants correlate with contemporary gradients of environmental energy inputs and water availability. However, climate—richness relationships for herbaceous species are less well known. Here we analyze the seed-bearing flora of Great Britain, quantifying the relationships between total, herb, and woody plant species richness and 14 environmental predictors measuring levels of energy, water, or combined energy and water. PCAs of the environmental variables identified the primary trends of environmental variation across Great Britain related to mean annual temperature and annual precipitation. OLS multiple regression and partial regression analyses identified mean annual temperature as the strongest driver of richness for all species groups. Similarly, reparameterized versions of published energy-water models for woody plant species richness and for angiosperm family richness gradients also identified energy variables as the primary predictors of the richness of all, woody, and herb species.

Testing the Water–Energy Theory on American Palms (Arecaceae) Using Geographically Weighted Regression

Water and energy have emerged as the best contemporary environmental correlates of broad-scale species richness patterns. A corollary hypothesis of water–energy dynamics theory is that the influence of water decreases and the influence of energy increases with absolute latitude. We report the first use of geographically weighted regression for testing this hypothesis on a continuous species richness gradient that is entirely located within the tropics and subtropics. The dataset was divided into northern and southern hemispheric portions to test whether predictor shifts are more pronounced in the less oceanic northern hemisphere. American palms (Arecaceae, n = 547 spp.), whose species richness and distributions are known to respond strongly to water and energy, were used as a model group. The ability of water and energy to explain palm species richness was quantified locally at different spatial scales and regressed on latitude. Clear latitudinal trends in agreement with water–energy dynamics theory were found, but the results did not differ qualitatively between hemispheres. Strong inherent spatial autocorrelation in local modeling results and collinearity of water and energy variables were identified as important methodological challenges. We overcame these problems by using simultaneous autoregressive models and variation partitioning. Our results show that the ability of water and energy to explain species richness changes not only across large climatic gradients spanning tropical to temperate or arctic zones but also within megathermal climates, at least for strictly tropical taxa such as palms. This finding suggests that the predictor shifts are related to gradual latitudinal changes in ambient energy (related to solar flux input) rather than to abrupt transitions at specific latitudes, such as the occurrence of frost.

Polychaete diversity and distribution patterns in Canadian marine waters

As changes to Canada’s marine biodiversity loom and the tools for assessing biodiversity advance, it becomes increasingly important to have baseline information about a region’s species so that future comparisons can be made. The polychaetes represent a conspicuous component of benthic communities in Canadian marine waters, yet information on species diversity and distribution is available only at the local scale. This study compiles polychaete species records from Canada over the last 150 years to assess broad-scale patterns of species richness, beta diversity (partitioned into turnover and nestedness), and species distributions in the surrounding Atlantic, Arctic, and Pacific Oceans. Records indicate the presence of 1200 species belonging to 68 families, while statistical extrapolation estimated a total of 1629 species. Pacific waters have the highest richness with 791 recorded species. The eastern and western regions of the Arctic Ocean have the highest faunal similarity, but the fewest endemic species, reflecting the recent ice-free status of this basin. Species turnover is highest between Pacific and Atlantic basins, decreases with geographic proximity, and is lowest in comparisons with Hudson Bay. Similar patterns of turnover between temperate and (sub)Arctic basins suggest that both Atlantic and Pacific faunas played an important role in Arctic recolonization following deglaciation. Despite the vast geographic area, nearly 13% of species are reported in all three oceans; however, many of these taxa likely represent species complexes. These results highlight the need for more detailed examination of many species with broad distributions and suggest that diversity in Canada is currently underestimated.

TESTS OF THE MID-DOMAIN HYPOTHESIS: A REVIEW OF THE EVIDENCE

Geographic variation of species richness is strongly correlated with environmental gradients. However, random arrangement of species distributions within a bounded domain can also theoretically produce richness gradients without underlying environmental gradients. This mid-domain effect (MDE) could serve as the null hypothesis against which to test effects of environmental variables, or as a component of a multivariate explanation of species-richness patterns. Recent reviews have concluded that there is a substantial MDE signature in observed geographical patterns of richness, based on correlations between observed patterns of richness and the predictions of mid-domain models. However, the mid-domain hypothesis makes additional powerful predictions about how richness should vary through space, and about the slope of the relationship between predicted and observed richness. Very few studies have tested these more powerful MDE predictions. Here, we reexamine the published mid-domain literature for agreement between observed patterns of richness and MDE predictions. We find that 50 of 53 published studies of MDEs showed significant deviations from the predictions of mid-domain models. When observed richness is correlated with MDE predictions, there are nearly always strongly collinear environmental gradients (e.g., in the Americas, climatic favorability and MDE-predicted richness are both maximal in the middle). Interpolation in sparsely sampled data can also give rise to spurious, apparently strong, mid-domain effects (e.g., the classic study of the Madagascan rain forest). We conclude that observed broad-scale patterns of species richness are not consistent with the mid-domain hypothesis.

Species Richness and Patterns of Invasion in Plants, Birds, and Fishes in the United States*

We quantified broad-scale patterns of species richness and species density (mean # species/km2) for native and non-indigenous plants, birds, and fishes in the continental USA and Hawaii. We hypothesized that the species density of native and non-indigenous taxa would generally decrease in northern latitudes and higher elevations following declines in potential evapotranspiration, mean temperature, and precipitation. County data on plants (n = 3004 counties) and birds (n=3074 counties), and drainage (6 HUC) data on fishes (n = 328 drainages) showed that the densities of native and non-indigenous species were strongly positively correlated for plant species (r = 0.86, P < 0.0001), bird species (r = 0.93, P<0.0001), and fish species (r = 0.41, P<0.0001). Multiple regression models showed that the densities of native plant and bird species could be strongly predicted (adj. R2 = 0.66 in both models) at county levels, but fish species densities were less predictable at drainage levels (adj. R2 = 0.31, P<0.0001). Similarly, non-indigenous plant and bird species densities were strongly predictable (adj. R2 = 0.84 and 0.91 respectively), but non-indigenous fish species density was less predictable (adj. R2 = 0.38). County level hotspots of native and non-indigenous plants, birds, and fishes were located in low elevation areas close to the coast with high precipitation and productivity (vegetation carbon). We show that (1) native species richness can be moderately well predicted with abiotic factors; (2) human populations have tended to settle in areas rich in native species; and (3) the richness and density of non-indigenous plant, bird, and fish species can be accurately predicted from biotic and abiotic factors largely because they are positively correlated to native species densities. We conclude that while humans facilitate the initial establishment, invasions of non-indigenous species, the spread and subsequent distributions of non-indigenous species may be controlled largely by environmental factors.

Concordance of species richness patterns among multiple freshwater taxa: a regional perspective

Geographical gradients in species richness and the degree to which different taxa show congru- ent patterns remain unknown for many taxonomic groups. Here, I examined broad-scale species richness patterns in five groups of freshwater organisms; macrophytes, dragonflies, stoneflies, aquatic beetles and fishes. The analyses were based on provincial distribution records in Denmark, Norway, Sweden and Fin- land. In general, variation in species richness across provinces was concordant among the groups, but stoneflies showed weaker negative relationships with the other taxonomic groups. Species richness in most groups decreased with increasing latitude and altitude, and a considerable part of the variation was ex- plained by mean July temperature. However, stoneflies showed a reversed pattern, with species richness correlating positively, albeit more weakly, with mean provincial altitude. Nevertheless, combined species richness of all five taxa showed a strong relationship with mean July temperature, accounting for 74% of variation in provincial species richness alone. Such temperature-controlled patterns suggest that regional freshwater biodiversity will strongly respond to climate change, with repercussions for local community organization in freshwater ecosystems in Fennoscandia.

The relative importance of evolutionary and environmental controls on broad-scale patterns of species richness in North America

Species richness varies greatly through space. It has been hypothesized that, over broad spatial scales, richness is determined by contemporary environmental characteristics, especially climatic energy. This hypothesis predicts that spatial variation of richness should be correlated with climatic variables. In contrast, it has also been hypothesized that differing net rates of speciation and subsequent dispersal may have given rise to contemporary patterns of species richness. The predictions of this hypothesis are less clear, but many proponents of the hypothesis derive the prediction that the average degree of evolutionary advancement of taxa should be related to patterns of richness. We have tested these predictions by examining spatial variation across North America of richness and of indices of evolutionary rate in the families Cicindelidae (tiger beetles), Cyprinidae (minnows), Catastomidae (suckers) and Percidae (perches and darters). We observed that richness in all three groups was related to climate in ways consistent with the richness-energy hypothesis. In contrast, richness in these four families was only weakly and inconsistently related to measures of evolutionary advancement. We conclude that, in the taxa for which we could find both phylogenies and distribution data, there is little reason to postulate evolutionary explanations for contemporary continental patterns of species richness in North America.