Additive partitioning of aquatic invertebrate species diversity across multiple spatial scales

1. Additive partitioning of three measures of diversity (species richness, Shannon's diversity index H and Simpson's diversity D) was used to study the relationship between local and regional diversity of benthic macroinvertebrate communities of boreal lakes (littoral habitats) and streams (riffle habitats) across three spatial scales (sampling sites, ecoregions and biogeographic regions). 2. Alpha (α) and beta (β) diversity are defined as within‐habitat and between‐habitat diversity, respectively. According to the concept of additive partitioning, diversity can be partitioned across multiple spatial scales such that the total (γ) diversity on one spatial scale becomes within‐habitat (α) diversity at the next higher scale. Hence, the total diversity at one scale is determined by the α diversity and the between‐habitat diversity (β) at the next lower scale. Consequently, one of the advantages of additive partitioning is that it is possible to study simultaneously β diversity and the regional‐local species relationship and the scale dependence of α and β components. 3. For both lakes and streams α diversity was low for sites and ecoregions, whereas β diversity was high, indicating that among‐site factors are important in describing the variability among the lakes and streams studied here. 4. Weak, albeit significant, evidence was found for regional and local species saturation patterns. Multiple stepwise regression indicated that local processes might be more important in structuring lake‐littoral and stream‐riffle species assemblages than regional processes. From these results we conclude that environmental heterogeneity may act as an important factor contributing to species coexistence, resulting in the observed saturation patterns. 5. Our study supports the use of additive partitioning for identifying specific patterns of macroinvertebrate diversity on multiple spatial scales and the underlying processes generating these patterns. This information is needed to improve understanding of the relation between patterns and processes affecting (decreasing) trends in aquatic biodiversity.     

村落文化林与非文化林多尺度物种多样性加性分配

文化林是指村民按照文化传统、风俗习惯或宗教信仰自觉保护和管理的森林,具有一定社会文化功能。目前国内外对文化林物种多样性研究主要为定性描述,缺乏对文化林和非文化林生物多样性的定量比较及差异来源分析。利用物种多样性加性分配方法,将总的Gamma 多样性分成样格内的Alpha多样性以及样格间、样方间和林型间Beta多样性,对中国亚热带地区3个村落文化林的乔木层、灌木层、草本层和藤本层进行物种多样性的多尺度分析。调查发现:(1)文化林共有维管束植物296种,以苦槠,樟和米槠为优势种,非文化林共有维管束植物189种,以杉木、马尾松和毛竹为优势种。文化林乔木层和灌木层物种数显著高于非文化林,草本层和藤本层物种数差异不显著。(2)Beta多样性随尺度增大而增加,林型间Beta多样性最高,占区域总Gamma多样性的41.9%-62.8%,其次是样方间Beta多样性(18.6%-31.9%),对区域多样性贡献最小为样格内Alpha和样格间Beta多样性。(3)林型间的多样性对区域物种多样性的贡献中,文化林占主导作用,乔木层占54.4%-81.0%,灌木层占51.2%-60.2%,草本层占42.9%-64.1%,藤本层占49.9%-62.2%。(4) 物种累积-面积曲线表明,在各个尺度上,文化林物种多样性始终高于非文化林,从而在相同面积下保护了更多的物种。加性分配模型在多个空间尺度上阐明了Alpha和Beta多样性的变化,突出了文化林对区域物种多样性的贡献,对保护对象和保护范围的决策以及生物多样性的保护与恢复具有重要意义。     

Ant diversity partitioning across spatial scales: Ecological processes and implications for conserving Tropical Dry Forests

Several ecological and evolutionary processes can drive changes in diversity at different spatial scales. To determine the scale at which these processes are most influential, we hypothesized that (i) broad‐scale differences between ecoregions had greater influence on ant species richness and species turnover than local differences among fragments within ecoregions; and (ii) the degree of dissimilarity in ant species composition is larger between Tropical Dry Forest fragments and the surrounding vegetations than among Tropical Dry Forests located in different ecoregions, indicating that extant Tropical Dry Forests are relicts of a broader distribution of this vegetation. To examine ant diversity patterns, we built a nested hierarchical design on three spatial scales, ranging from fragments (local scale), Tropical Dry Forest + surroundings vegetation (landscape scale) and Brazilian ecoregions (regional scale). We used 450 sampling units (45 sampling units × two fragments × five ecoregions = 450). A null model based on the sample was used to identify variations in the random distribution across spatial scales. Spatial partitioning of ant diversity showed that observed β₁ diversity (between fragments) and β₂ diversity (among ecoregions) were higher than expected by chance. When the partitioning was analysed separately for each region, the observed β₁ diversity (Tropical Dry Forest and surrounding vegetation) was higher than expected by the null hypothesis in all ecoregions of Brazil. Based on species composition and diversity patterns, we stress the importance of creating more protected areas throughout the coverage area of Tropical Dry Forests, favouring a more efficient conservation process.     

Species richness change across spatial scales

Humans have elevated global extinction rates and thus lowered global scale species richness. However, there is no a priori reason to expect that losses of global species richness should always, or even often, trickle down to losses of species richness at regional and local scales, even though this relationship is often assumed. Here, we show that scale can modulate our estimates of species richness change through time in the face of anthropogenic pressures, but not in a unidirectional way. Instead, the magnitude of species richness change through time can increase, decrease, reverse, or be unimodal across spatial scales. Using several case studies, we show different forms of scale‐dependent richness change through time in the face of anthropogenic pressures. For example, Central American corals show a homogenization pattern, where small scale richness is largely unchanged through time, while larger scale richness change is highly negative. Alternatively, birds in North America showed a differentiation effect, where species richness was again largely unchanged through time at small scales, but was more positive at larger scales. Finally, we collated data from a heterogeneous set of studies of different taxa measured through time from sites ranging from small plots to entire continents, and found highly variable patterns that nevertheless imply complex scale‐dependence in several taxa. In summary, understanding how biodiversity is changing in the Anthropocene requires an explicit recognition of the influence of spatial scale, and we conclude with some recommendations for how to better incorporate scale into our estimates of change.     

Woody Encroachment Removal from Midwestern Oak Savannas Alters Understory Diversity across Space and Time

Recovering biodiversity is a common goal during restoration; however, for many ecosystems, it is not well understood how restoration influences species diversity across space and time. I examined understory species diversity and composition after woody encroachment removal in a large-scale savanna restoration experiment in central Iowa, United States. Over a 4-year time series, restoration had profound effects across space and time, increasing richness at local and site-level scales. Restoration sites had increased α (within sample) Simpson's diversity and α and γ (site level) species richness relative to control sites, although γ and β (among sample) Simpson's diversity, β richness, and α species evenness were not affected. Changes in richness were driven by graminoids at the α and γ scales and woody species (and some evidence for forbs) at the α scale. Interestingly, indicator species analysis revealed that at least some species from all functional groups were promoted by restoration, although no species were significant indicators of pre-treatment or control sites. Both savanna and nonsavanna species were indicators of restored sites. Restoration promoted exotic species at both scales, although species with spring phenologies were unaffected. Woody encroachment removal may be a means to promote species establishment in savannas; however, in this study, it resulted in establishment and proliferation of native and exotic and savanna and nonsavanna species. Future work might consider reintroduction of key savanna species to supplement those that have established. Work like this demonstrates the utility of restoration experiments for conducting research on large- and multiscale processes, such as species diversity.     

生物多样性分布格局的地史成因假说

生物多样性的大尺度分布格局是现代环境与地史过程共同作用的结果。本文从影响机制、参数选择及与现代气候假说的关系等方面回顾了地史成因假说的最新进展, 并得出以下认识: (1) 地史过程对生物多样性的分布格局有显著影响, 但地史过程与现代环境之间强烈的共线性使得两者的影响常叠加在一起; (2) 与广域物种的多样性相比, 地史过程更有利于解释狭域物种(或特有物种)的多样性; (3) 地史过程的参数选择是地史假说所面临的挑战之一, 目前所用的指标与现代环境具有显著的共线性, 难以直观地体现地史过程对生物多样性的影响, 对不同区域内物种系统发育过程的对比或者物种形成速率及灭绝速率分布格局的分析可能有助于评价地史成因假说的影响。     

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.     

Identifying potential drivers of distribution patterns of invasive Corbicula fluminea relative to native freshwater mussels (Unionidae) across spatial scales

This study aimed to identify the importance of ecological factors to distribution patterns of the invasive Clam (Corbicula fluminea) relative to native mussels (family: Unionidae) across seven rivers within the Mobile and Tennessee basins, Southeast United States. We quantitatively surveyed dense, diverse native mussel aggregations across 20 river reaches and estimated mussel density, biomass, and species richness along with density of invasive C. fluminea (hereafter Corbicula). We measured substrate particle size, velocity, and depth in quadrats where animals were collected. Additionally, we characterized reach scale environmental parameters including seston quantity and quality (% Carbon, % Nitrogen, % Phosphorous), water chemistry (ammonium [NH4+], soluble reactive phosphorous [SRP]), and watershed area and land cover. Using model selection, logistic regression, and multivariate analysis, we characterized habitat features and their association to invasive Corbicula within mussel beds. We found that Corbicula were more likely to occur and more abundant in quadrats with greater mussel biomass, larger substrate size, faster water velocity, and shallower water depth. At the reach scale, Corbicula densities increased where particle sizes were larger. Mussel richness, density, and biomass increased with watershed area. Water column NH4+ increased at reaches with more urban land cover. No land cover variables influenced Corbicula populations or mussel communities. The strong overlapping distribution of Corbicula and mussels support the hypothesis that Corbicula are not necessarily limited by habitat factors and may be passengers of change in rivers where mussels have declined due to habitat degradation. Whether Corbicula is facilitated by mussels or negatively interacts with mussels in these systems remains to be seen. Focused experiments that manipulate patch scale variables would improve our understanding of the role of species interactions (e.g., competition, predation, facilitation) or physical habitat factors in influencing spatial overlap between Corbicula and native mussels.     

Local–regional diversity relationship varies with spatial scale in lotic diatoms

Aim Ecologists have shown increasing interest in the relative roles of local and regional factors in structuring biotic communities. One approach to studying this is to examine the relationship between local species richness (LSR) and regional species richness (RSR). We examined the LSR–RSR relationship in stream diatoms, using two data sets that varied in spatial extent. At broad spatial extent ranging across drainage systems, we expected climatic and dispersal‐related factors to constrain LSR, thus resulting in a linear LSR–RSR relationship. However, at small spatial scales dispersal across sites should be unconstrained, resulting in strong local interactions and a weak or asymptotic LSR–RSR relationship. Location Boreal streams in Finland. Methods For data set 1, we sampled 15 stream riffles (localities) in each of eight drainage systems (regions), with the latitudinal gradient between the southernmost and northernmost sites being almost 1100 km. For data set 2, a locality for estimating LSR was a single stone, and each riffle represented a region for estimating RSR. We sampled 20 stones in each of eight riffles. We used linear regressions to examine the relationship between LSR and RSR across regions. We used both observed richness values, as well as values estimated with the Chao1 estimator. Results We found a relatively strong linear relationship between the Chao1‐estimated mean LSR and RSR (R² = 0.654, P = 0.015) across drainage systems. The slope of the regression was 0.643 and it did not differ from 1.0, thus indicating linearity. At the riffle scale, however, LSR and RSR were not linearly related, and the slope of the regression (0.039) differed significantly from 1.0, indicating curvilinearity. Main conclusions These results suggest that the relationship between mean LSR and RSR varies across spatial scales in diatoms – from significantly linear at large scales to curvilinear at small scales. These plots imply strong regional enrichment in stream diatoms across drainage systems. Their diversity is thus determined largely by the composition of the regional species pool, as also in many macroorganisms. In contrast, at small spatial scales the LSR–RSR relationship implied a hard limit to local diversity, reflecting the primacy of local processes.     

Elevational patterns and hierarchical determinants of biodiversity across microbial taxonomic scales

Microbial biogeography is gaining increasing attention due to recent molecular methodological advance. However, the diversity patterns and their environmental determinants across taxonomic scales are still poorly studied. By sampling along an extensive elevational gradient in subarctic ponds of Finland and Norway, we examined the diversity patterns of aquatic bacteria and fungi from whole community to individual taxa across taxonomic coverage and taxonomic resolutions. We further quantified cross‐phylum congruence in multiple biodiversity metrics and evaluated the relative importance of climate, catchment and local pond variables as the hierarchical drivers of biodiversity across taxonomic scales. Bacterial community showed significantly decreasing elevational patterns in species richness and evenness, and U‐shaped patterns in local contribution to beta diversity (LCBD). Conversely, no significant species richness and evenness patterns were found for fungal community. Elevational patterns in species richness and LCBD, but not in evenness, were congruent across bacterial phyla. When narrowing down the taxonomic scope towards higher resolutions, bacterial diversity showed weaker and more complex elevational patterns. Taxonomic downscaling also indicated a notable change in the relative importance of biodiversity determinants with stronger local environmental filtering, but decreased importance of climatic variables. This suggested that niche conservatism of temperature preference was phylogenetically deeper than that of water chemistry variables. Our results provide novel perspectives for microbial biogeography and highlight the importance of taxonomic scale dependency and hierarchical drivers when modelling biodiversity and species distribution responses to future climatic scenarios.     

Strong paleoclimatic legacies in current plant functional diversity patterns across Europe

Numerous studies indicate that environmental changes during the late Quaternary have elicited long‐term disequilibria between species diversity and environment. Despite its importance for ecosystem functioning, the importance of historical environmental conditions as determinants of FD (functional diversity) remains largely unstudied. We quantified the geographic distributions of plant FD (richness and dispersion) across Europe using distribution and functional trait information for 2702 plant species. We then compared the importance of historical and contemporary factors to determine the relevance of past conditions as predictors of current plant FD in Europe. For this, we compared the strength of the relationships between FD with temperature and precipitation stability since the LGM (Last Glacial Maximum), accessibility to LGM refugia, and contemporary environmental conditions (climate, productivity, soil, topography, and land use). Functional richness and dispersion exhibited geographic patterns with strong associations to the environmental history of the region. The effect size of accessibility to LGM refugia and climate stability since the LGM was comparable to that of the contemporary predictors. Both functional richness and dispersion increased with temperature stability since the LGM and accessibility to LGM refugia. Functional richness' geographic pattern was primarily associated with accessibility to LGM refugia growing degree‐days, land use heterogeneity, diversity of soil types, and absolute minimum winter temperature. Functional dispersion's geographic pattern was primarily associated with accessibility to LGM refugia growing degree‐days and absolute minimum winter temperature. The high explained variance and model support of historical predictors are consistent with the idea that long‐term variability in environmental conditions supplements contemporary factors in shaping FD patterns at continental scales. Given the importance of FD for ecosystem functioning, future climate change may elicit not just short‐term shifts in ecosystem functioning, but also long‐term functional disequilibria.     

Drivers of temporal changes in temperate forest plant diversity vary across spatial scales

Global biodiversity is affected by numerous environmental drivers. Yet, the extent to which global environmental changes contribute to changes in local diversity is poorly understood. We investigated biodiversity changes in a meta‐analysis of 39 resurvey studies in European temperate forests (3988 vegetation records in total, 17–75 years between the two surveys) by assessing the importance of (i) coarse‐resolution (i.e., among sites) vs. fine‐resolution (i.e., within sites) environmental differences and (ii) changing environmental conditions between surveys. Our results clarify the mechanisms underlying the direction and magnitude of local‐scale biodiversity changes. While not detecting any net local diversity loss, we observed considerable among‐site variation, partly explained by temporal changes in light availability (a local driver) and density of large herbivores (a regional driver). Furthermore, strong evidence was found that presurvey levels of nitrogen deposition determined subsequent diversity changes. We conclude that models forecasting future biodiversity changes should consider coarse‐resolution environmental changes, account for differences in baseline environmental conditions and for local changes in fine‐resolution environmental conditions.     

Assessing endemism at multiple spatial scales, with an example from the Australian vascular flora

Aim  To develop an approach for assessing the spatial scale of centres of endemism among species level data.
Location Australia.
Methods  Endemism is inherently scale dependent. Therefore, the Corrected Weighted Endemism (CWE) index used by Crisp et al. [ J. Biogeogr. (2001)28:183] is extended to account for species samples in local neighbourhoods as a Spatial CWE index. This then allows an analysis of how the degree of endemism of a location (cell) changes with spatial scale. The quality of the Spatial CWE index results are assessed using three spatial randomizations at the species level with and without preserving species richness and distributional patterns. We show that CWE is equivalent to beta diversity and predict that it should show high rates of change around centres of endemism.
Results  Similar patterns to those found by Crisp et al. using a data set of vascular flora from Australia are retrieved, but the extent to which they are scale dependent is more easily identified. For example, the Central Australian centre discounted by Crisp et al. is identified when a three-cell radius neighbourhood is used. However, the level of endemism in this centre is no greater than in the margins of many of the coastal centres of endemism. Most of the identified centres of endemism are better than random at all scales and are increasingly so as the spatial scale increases. As predicted, the highest rate of change in Spatial CWE (beta diversity) is most often between zero- and one-cell radius neighbours in most centres of endemism.
Main conclusions  The explicit incorporation of geographical space in analyses allows for a greater understanding of the scale-dependence of phenomena, in this case endemism and beta diversity.     

Grazer exclusion alters plant spatial organization at multiple scales,increasing diversity

Grazing is one of the most important factors influencing community structure and productivity in natural grasslands. Understanding why and how grazing pressure changes species diversity is essential for the preservation and restoration of biodiversity in grasslands. We use heavily grazed subalpine meadows in the Qinghai‐Tibetan Plateau to test the hypothesis that grazer exclusion alters plant diversity by changing inter‐ and intraspecific species distributions. Using recently developed spatial analyses combined with detailed ramet mapping of entire plant communities (91 species), we show striking differences between grazed and fenced areas that emerged at scales of just one meter. Species richness was similar at very small scales (0.0625 m²), but at larger scales diversity in grazed areas fell below 75% of corresponding fenced areas. These differences were explained by differences in spatial distributions; intra‐ and interspecific associations changed from aggregated at small scales to overdispersed in the fenced plots, but were consistently aggregated in the grazed ones. We conclude that grazing enhanced inter‐ and intraspecific aggregations and maintained high diversity at small scales, but caused decreased turnover in species at larger scales, resulting in lower species richness. Our study provides strong support to the theoretical prediction that inter‐ and intraspecific aggregation produces local spatial patterns that scale‐up to affect species diversity in a community. It also demonstrates that the impacts of grazing can manifest through this mechanism, lowering diversity by reducing spatial turnover in species. Finally, it highlights the ecological and physiological plant processes that are likely responding to grazing and thereby altering aggregation patterns, providing new insights for monitoring, and mediating the impacts of grazing.     

Life history determines biogeographical patterns of soil bacterial communities over multiple spatial scales

The extent to which the distribution of soil bacteria is controlled by local environment vs. spatial factors (e.g. dispersal, colonization limitation, evolutionary events) is poorly understood and widely debated. Our understanding of biogeographic controls in microbial communities is likely hampered by the enormous environmental variability encountered across spatial scales and the broad diversity of microbial life histories. Here, we constrained environmental factors (soil chemistry, climate, above‐ground plant community) to investigate the specific influence of space, by fitting all other variables first, on bacterial communities in soils over distances from m to 10² km. We found strong evidence for a spatial component to bacterial community structure that varies with scale and organism life history (dispersal and survival ability). Geographic distance had no influence over community structure for organisms known to have survival stages, but the converse was true for organisms thought to be less hardy. Community function (substrate utilization) was also shown to be highly correlated with community structure, but not to abiotic factors, suggesting nonstochastic determinants of community structure are important Our results support the view that bacterial soil communities are constrained by both edaphic factors and geographic distance and further show that the relative importance of such constraints depends critically on the taxonomic resolution used to evaluate spatio‐temporal patterns of microbial diversity, as well as life history of the groups being investigated, much as is the case for macro‐organisms.     

The partitioning of macroinvertebrate diversity across multiple spatial scales in the upper Modder River System,South Africa

A method for assessing the alpha and beta diversity components of a macroinvertebrate community across numerous spatial scales is presented. Findings were not empirically linked to ecological questions as the purpose of this study was primarily the demonstration of a diversity partitioning method. Sampling was carried out at three sites on the upper Modder River in the Free State Province, South Africa between April 2008 and January 2009. Communities were analysed by investigating the relative frequency of species in specific biotopes, a Similarity Profile (SIMPROF) and cluster analyses of the Bray‐Curtis similarities between samples, and the partitioning of species richness and Shannon diversity across multiple spatial scales. Findings revealed that sites showed significant clustering (SIMPROF P < 0.05; <20% Bray‐Curtis similarity), and the species frequencies indicated preference to selected microhabitats. Species richness and Shannon diversity of macroinvertebrates differed significantly (5% confidence levels) from randomly simulated values for sampling sites, biotopes and seasons indicating that diversity is clustered and not homogeneously distributed. The diversity partitioning could have potential in diversity assessment for conservation biology, land management and environmental impact assessments.     

Macroecological patterns of spider species richness across Europe

We analysed the pattern of covariation of European spider species richness with various environmental variables at different scales. Four layers of perception ranging from single investigation sites to the whole European continent were selected. Species richness was determined using published data from all four scales. Correlation analyses and stepwise multiple linear regression were used to relate richness to topographic, climatic and biotic variables. Up to nine environmental variables were included in the analyses (area, latitude, elevation range, mean annual temperature, local variation in mean annual temperature, mean annual precipitation, mean July temperature, local variation in mean July temperature, plant species richness). At the local and at the continental scale, no significant correlations with surface area were found, whereas at the landscape and regional scale, surface area had a significant positive effect on species richness. Factors that were positively correlated with species richness at both broader scales were plant species richness, elevation range, and specific temperature variables (regional scale: local variation in mean annual, and mean July temperature; continental scale: mean July temperature). Latitude was significantly negatively correlated with the species richness at the continental scale. Multiple models for spider species richness data accounted for up to 77% of the total variance in spider species richness data. Furthermore, multiple models explained variation in plant species richness up to 79% through the variables mean July temperature and elevation range. We conclude that these first continental wide analyses grasp the overall pattern in spider species richness of Europe quite well, although some of the observed patterns are not directly causal. Climatic variables are expected to be among the most important direct factors, although other variables (e.g. elevation range, plant species richness) are important (surrogate) correlates of spider species richness.