Strong influence of regional species pools on continent-wide structuring of local communities

There is a long tradition in ecology of evaluating the relative contribution of the regional species pool and local interactions on the structure of local communities. Similarly, a growing number of studies assess the phylogenetic structure of communities, relative to that in the regional species pool, to examine the interplay between broad-scale evolutionary and fine-scale ecological processes. Finally, a renewed interest in the influence of species source pools on communities has shown that the definition of the source pool influences interpretations of patterns of community structure. We use a continent-wide dataset of local ant communities and implement ecologically explicit source pool definitions to examine the relative importance of regional species pools and local interactions for shaping community structure. Then we assess which factors underlie systematic variation in the structure of communities along climatic gradients. We find that the average phylogenetic relatedness of species in ant communities decreases from tropical to temperate regions, but the strength of this relationship depends on the level of ecological realism in the definition of source pools. We conclude that the evolution of climatic niches influences the phylogenetic structure of regional source pools and that the influence of regional source pools on local community structure is strong.     

Testing for local species saturation with nonindependent regional species pools

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

Disentangling the importance of ecological niches from stochastic processes across scales

Deterministic theories in community ecology suggest that local, niche-based processes, such as environmental filtering, biotic interactions and interspecific trade-offs largely determine patterns of species diversity and composition. In contrast, more stochastic theories emphasize the importance of chance colonization, random extinction and ecological drift. The schisms between deterministic and stochastic perspectives, which date back to the earliest days of ecology, continue to fuel contemporary debates (e.g. niches versus neutrality). As illustrated by the pioneering studies of Robert H. MacArthur and co-workers, resolution to these debates requires consideration of how the importance of local processes changes across scales. Here, we develop a framework for disentangling the relative importance of deterministic and stochastic processes in generating site-to-site variation in species composition (β-diversity) along ecological gradients (disturbance, productivity and biotic interactions) and among biogeographic regions that differ in the size of the regional species pool. We illustrate how to discern the importance of deterministic processes using null-model approaches that explicitly account for local and regional factors that inherently create stochastic turnover. By embracing processes across scales, we can build a more synthetic framework for understanding how niches structure patterns of biodiversity in the face of stochastic processes that emerge from local and biogeographic factors.     

The drivers of woody species richness and density in a Neotropical savannah

Environmental filtering prevents species without certain attributes from occurring in local communities. Traits respond differently to different abiotic factors, assembling communities with varying composition along environmental gradients. Here, we measured proxies of soil fertility, disturbance by fire, response and physiological traits to assess how these variables interact to determine woody species richness and density in a Neotropical savannah. We explicitly incorporated our assumptions about how different abiotic filters influence different subsets of traits into a statistical model using structural equation modelling, yielding a more accurate representation of the assembly process. Fire had an effect on resistance traits, whereas soil fertility influenced physiological traits. Resistance traits explained both the richness and density of plots, whereas physiological traits explained only the density. Fewer fire events led to richer and denser plots. Similarly, areas with lower cation exchange capacity assembled less dense communities. Furthermore, we showed that structural equation modelling yielded a realistic representation of the bivariate interactions of distinct environmental filters with different subsets of traits.     

The formation of species pools: historical habitat abundance affects current local diversity

Aim Explanations of biogeographic diversity patterns have emphasized the role of large‐scale processes that determine species pools, whereas explanations of local patterns have not. We address the hypothesis that local diversity patterns are also primarily dependent on the size of the available species pools, which are expected to be large when the particular habitat type has been evolutionary more abundant, or in unproductive habitats due to shorter generation time and hence higher diversification rates. Location The Canary Islands. Methods We determined the geographic distribution and habitat requirements of all native vascular plant species in the Canary Islands. Species pools for each habitat type on particular islands were further split into two categories according to origin: either originating due to local diversification or due to natural immigration. The dependence of historical diversification and diversification rate on habitat type, area, age, altitude and distance to the mainland was tested with general linear mixed models weighed according to the Akaike information criterion. Results The largest portion of the local variation in plant species diversity was attributed to the historic (pre‐human) habitat area, although island age was also important. The diversification rate was higher in unproductive habitats of coastal scrub and summit vegetation. Main conclusion Our study supports the species pool hypothesis, demonstrating that natural local patterns of species diversity in different habitats mirror the abundance of those particular habitats in evolutionary history. It also supports the community‐level birth rate hypothesis, claiming that stressful conditions result in higher diversification rates. We conclude that much of the observed local variation in plant diversity can be attributed to the differing sizes of species pools evolved under particular habitat conditions, and that historic parameters are far more important determinants of local diversity than suggested by ecological theory.     

Estimating species richness using environmental DNA

The foundation for any ecological study and for the effective management of biodiversity in natural systems requires knowing what species are present in an ecosystem. We assessed fish communities in a stream using two methods, depletion‐based electrofishing and environmental DNA metabarcoding (eDNA) from water samples, to test the hypothesis that eDNA provides an alternative means of determining species richness and species identities for a natural ecosystem. In a northern Indiana stream, electrofishing yielded a direct estimate of 12 species and a mean estimated richness (Chao II estimator) of 16.6 species with a 95% confidence interval from 12.8 to 42.2. eDNA sampling detected an additional four species, congruent with the mean Chao II estimate from electrofishing. This increased detection rate for fish species between methods suggests that eDNA sampling can enhance estimation of fish fauna in flowing waters while having minimal sampling impacts on fish and their habitat. Modern genetic approaches therefore have the potential to transform our ability to build a more complete list of species for ecological investigations and inform management of aquatic ecosystems.     

伊洛河流域草本植物群落物种多样性

生物多样性沿环境梯度的变化是生物多样性研究的重要内容,环境梯度包含了多种环境因子(海拔高度、水热条件、人类扰动等)的综合。以伊洛河流域草本植物群落为对象,沿河从入黄河口到河源地选取典型样地调查研究伊洛河流域草本植物群落物种多样性及其分布格局。结果表明:物种丰富度和Shannon-Wiener多样性指数沿河均稍呈"S"型曲线变化,不同群落类型中分布格局差别不大,各群落类型中的物种丰富度和多样性均呈现出中游丘陵山地交界区最高,上游河源区次之,下游平原地区最低的趋势;β多样性指数的变化趋势与α多样性较一致,总体上呈现出中游丘陵山地区物种更替速率较快,平原区更替较慢;在流域内上游河源地属于自然植被区,人为干扰较轻,具有较高的物种多样性,物种替代主要受物种的竞争扩散能力和生境条件的制约;在下游平原农业区,人类活动强烈,区域内以人工生态系统为主,物种组成简单,物种替代具有跳跃性的特征,主要受人类活动的制约;在中游从自然生态系统向农业生态系统的过渡区域,人类活动的扰动有一定的强度,导致该区域内自然分布种和伴人种混合生长,具有较高的物种多样性和较快的物种替代速率。总体上伊洛河流域草本植物群落物种多样性分布格局强烈的受到人类活动的影响,物种替代速率较高。     

局域和区域过程共同控制着群落的物种多样性:种库假说

解释群落的物种多样性大小是生态学研究的一个重要的理论和实践问题。人们提出了群落物种多样性的多种假说, Zobel等人提出的种库假说(species pool hypothesis)是生物多样性理论研究的重要发展。该假说认为, 一个群落的物种多样性不仅与环境条件和生态过程(ecological process)(如竞争、捕食)有关, 也受区域种库(regional species pool)的限制。区域种库是指一个地区可进入某一群落的潜在物种数量, 它由地史过程(如冰期、地质年代)和区域过程(物种形成、迁移扩散以及消亡)所决定。按照种库假说, 某一生境类型的面积越大, 地质年代越古老, 物种形成的机会也就越多, 因而能适应和分布于该生境的物种也就越多, 实际群落中的物种丰富度也就越高。种库在空间上主要有两个层次: 区域种库和实际种库, 前者指某一生境所拥有的潜在物种数量, 主要由生物地理过程(biogeographic processes)所决定; 后者则为调查的群落中实际出现的物种数量, 主要由竞争等生态过程和区域种库共同决定。本文对种库假说的基本概念、主要内容、种库确定方法等作了介绍, 并阐述了作者对这些问题的理解和认识。     

Patterns of species richness and turnover in endemic amphibians of the Guineo-Congolian rain forest

Aim

The African Guineo-Congolian (GC) region is a global biodiversity hotspot with high species endemism, bioclimatic heterogeneity, complex landscape features, and multiple biogeographic barriers. Bioclimatic and geographic variables influence global patterns of species richness and endemism, but their relative importance varies across taxa and regions and is poorly understood for many faunas. Here, we test the hypothesis that turnover in endemic amphibians of the GC biodiversity hotspot is influenced mainly by the geographic distance between grid cells and secondarily by rainfall- and temperature-related variables.

Location

West and Central Africa.

Major Taxa Studied

Amphibians.

Methods

We compiled species-occurrence records via field sampling, online databases, and taxonomic literature. Our study used 1205 unique georeferenced records of 222 amphibian species endemic to the GC region. Patterns of species richness were mapped onto a grid with a spatial resolution of 0.5° × 0.5°. We estimated weighted endemism and tested whether endemism was higher than the expected species richness (randomization test). We quantified species turnover using generalized dissimilarity modelling to evaluate the processes underlying observed patterns of species richness in GC endemic amphibians. We explored bioregionalization using agglomerative hierarchical clustering based on the unweighted pair group method with arithmetic averages.

Results

We identified seven areas within the lower GC region – forests in Cameroon, Gabon, Southern Nigeria, Equatorial Guinea, Republic of Congo, Democratic Republic of Congo, and Cote d'Ivoire – as having high species richness of endemic amphibians. The randomization test returned four major areas of significant weighted endemism: Nigeria-Cameroon mountains, forest regions of the Democratic Republic of Congo, Cote d'Ivoire, and Ghana. Our analysis revealed five bioregions for amphibian endemism, four of which were located within the lower Guineo-Congolian forest. Species turnover was strongly related to the geographic distance between grid cells; contributing bioclimatic variables included precipitation of the warmest quarter, mean temperature of the wettest quarter, and mean diurnal temperature range.

Main Conclusions

Our results indicate that geographic distance between grid cells is the primary determinant of turnover in GC endemic amphibians, with secondary but significant effects of rainfall- and temperature-related variables. Our study identifies key areas of endemic amphibian richness that could be prioritized for conservation actions.     

Disentangling drivers of spatial autocorrelation in species distribution models

Species distribution models (SDMs) are frequently used to understand the influence of site properties on species occurrence. For robust model inference, SDMs need to account for the spatial autocorrelation of virtually all species occurrence data. Current methods do not routinely distinguish between extrinsic and intrinsic drivers of spatial autocorrelation, although these may have different implications for conservation. Here, we present and test a method that disentangles extrinsic and intrinsic drivers of spatial autocorrelation using repeated observations of a species. We focus on unknown habitat characteristics and conspecific interactions as extrinsic and intrinsic drivers, respectively. We model the former with spatially correlated random effects and the latter with an autocovariate, such that the spatially correlated random effects are constant across the repeated observations whereas the autocovariate may change. We tested the performance of our model on virtual species data and applied it to observations of the corncrake Crex crex in the Netherlands. Applying our model to virtual species data revealed that it was well able to distinguish between the two different drivers of spatial autocorrelation, outperforming models with no or a single component for spatial autocorrelation. This finding was independent of the direction of the conspecific interactions (i.e. conspecific attraction versus competitive exclusion). The simulations confirmed that the ability of our model to disentangle both drivers of autocorrelation depends on repeated observations. In the case study, we discovered that the corncrake has a stronger response to habitat characteristics compared to a model that did not include spatially correlated random effects, whereas conspecific interactions appeared to be less important. This implies that future conservation efforts should primarily focus on maximizing habitat availability. Our study shows how to systematically disentangle extrinsic and intrinsic drivers of spatial autocorrelation. The method we propose can help to correctly identify the main drivers of species distributions.     

Multiple source pools for Galápagos plant species richness: a critical analysis of the line of sight connectivity index

An index (C_i*E) combining the number of line‐of‐sight islands (C_i) within a radius i and target island elevation (E) has been proposed as an improved predictive model of plant species richness (S_t) in the Galápagos Archipelago. We examined this index critically and found that several major flaws preclude it from being a useful predictive tool for the archipelago. Although the number of collecting trips to an island was reported over 20 years ago to have substantial predictive value for reported plant species richness in the Galápagos Islands, this relationship was ignored in multiple regression analyses of the index. When we included the number of collecting trips in different multiple regression analyses of the index, C_i*E had less predictive power than collecting trips or ceased to be significant at all. Additionally, the strong significant relationship between elevation and area in the Galápagos Archipelago results in area having a major confounding influence on the C_i*E index. When elevation is removed from the C_i*E index, the predictive power of C_i is far less than area alone. Finally, the data used to construct and correlate the C_i*E index with (S_t) were based only on a subset of the islands and species lists that were incomplete or out of date. Species richness on islands can be related to the interaction of different factors, so development and testing of indices like C_i*E is not inappropriate. However, it is important to examine the interrelationships among the components of these indices thoroughly, and not ignore the effect of factors already known to have high predictive power. We propose several ways in which more meaningful indices of source pool(s) capacity can be constructed.