Effects of dispersal mode on the environmental and spatial correlates of nestedness and species turnover in pond communities

Advances in metacommunity theory have made a significant contribution to understanding the drivers of variation in biological communities. However, there has been limited empirical research exploring the expression of metacommunity theory for two fundamental components of beta diversity: nestedness and species turnover. In this paper, we examine the influence of local environmental and a range of spatial variables (hydrological connectivity, proximity and overall spatial structure) on total beta diversity and the nestedness and turnover components of beta diversity for the entire macroinvertebrate community and active and passively dispersing taxa within pond habitats. High beta diversity almost entirely reflects patterns of species turnover (replacement) rather than nestedness (differences in species richness) in our dataset. Local environmental variables were the main drivers of total beta diversity, nestedness and turnover when the entire community was considered and for both active and passively dispersing taxa. The influence of spatial processes on passively dispersing taxa, total beta diversity and nestedness was significantly greater than for actively dispersing taxa. Our results suggest that species sorting (local environmental variables) operating through niche processes was the primary mechanism driving total beta diversity, nestedness and turnover for the entire community and active and passively dispersing taxa. In contrast, spatial factors (hydrological connectivity, proximity and spatial eigenvectors) only exerted a secondary influence on the nestedness and turnover components of beta diversity.     

Partitioning global patterns of freshwater fish beta diversity reveals contrasting signatures of past climate changes

Here, we employ an additive partitioning framework to disentangle the contribution of spatial turnover and nestedness to beta diversity patterns in the global freshwater fish fauna. We find that spatial turnover and nestedness differ geographically in their contribution to freshwater fish beta diversity, a pattern that results from contrasting influences of Quaternary climate changes. Differences in fish faunas characterized by nestedness are greater in drainage basins that experienced larger amplitudes of Quaternary climate oscillations. Conversely, higher levels of spatial turnover are found in historically unglaciated drainage basins with high topographic relief, these having experienced greater Quaternary climate stability. Such an historical climate signature is not clearly detected when considering the overall level of beta diversity. Quantifying the relative roles of historical and ecological factors in explaining present-day patterns of beta diversity hence requires considering the different processes generating these patterns and not solely the overall level of beta diversity.     

Projected impacts of climate change on spatio‐temporal patterns of freshwater fish beta diversity: a deconstructing approach

Aim To assess the potential impacts of future climate change on spatio‐temporal patterns of freshwater fish beta diversity. Location Adour–Garonne River Basin (France). Methods We first applied an ensemble modelling approach to project annually the future distribution of 18 fish species for the 2010–2100 period on 50 sites. We then explored the spatial and temporal patterns of beta diversity by distinguishing between its two additive components, namely species turnover and nestedness. Results Taxonomic homogenization of fish assemblages was projected to increase linearly over the 21st century, especially in the downstream parts of the river gradient. This homogenization process was almost entirely caused by a decrease in spatial species turnover. When considering the temporal dimension of beta diversity, our results reveal an overall pattern of decreasing beta diversity along the upstream–downstream river gradient. In contrast, when considering the turnover and nestedness components of temporal beta diversity we found significant U‐shaped and hump‐shaped relationships, respectively. Main conclusions Future climate change is projected to modify the taxonomic composition of freshwater fish assemblages by increasing their overall similarity over the Adour–Garonne River Basin. Our findings suggest that the distinction between the nestedness and turnover components of beta diversity is not only crucial for understanding the processes shaping spatial beta‐diversity patterns but also for identifying localities where the rates of species replacement are projected to be greatest. Specifically we recommend that future conservation studies should not only consider the spatial component of beta diversity but also its dynamic caused by climate warming.     

Partitioning the turnover and nestedness components of beta diversity

Aim  Beta diversity (variation of the species composition of assemblages) may reflect two different phenomena, spatial species turnover and nestedness of assemblages, which result from two antithetic processes, namely species replacement and species loss, respectively. The aim of this paper is to provide a unified framework for the assessment of beta diversity, disentangling the contribution of spatial turnover and nestedness to beta-diversity patterns.
Innovation  I derive an additive partitioning of beta diversity that provides the two separate components of spatial turnover and nestedness underlying the total amount of beta diversity. I propose two families of measures of beta diversity for pairwise and multiple-site situations. Each family comprises one measure accounting for all aspects of beta diversity, which is additively decomposed into two measures accounting for the pure spatial turnover and nestedness components, respectively. Finally, I provide a case study using European longhorn beetles to exemplify the relevance of disentangling spatial turnover and nestedness patterns.
Main conclusion  Assigning the different beta-diversity patterns to their respective biological phenomena is essential for analysing the causality of the processes underlying biodiversity. Thus, the differentiation of the spatial turnover and nestedness components of beta diversity is crucial for our understanding of central biogeographic, ecological and conservation issues.     

Assembly mechanisms determining high species turnover in aquatic communities over regional and continental scales

Niche and neutral processes drive community assembly and metacommunity dynamics, but their relative importance might vary with the spatial scale. The contribution of niche processes is generally expected to increase with increasing spatial extent at a higher rate than that of neutral processes. However, the extent to what community composition is limited by dispersal (usually considered a neutral process) over increasing spatial scales might depend on the dispersal capacity of composing species. To investigate the mechanisms underlying the distribution and diversity of species known to have great powers of dispersal (hundreds of kilometres), we analysed the relative importance of niche processes and dispersal limitation in determining beta‐diversity patterns of aquatic plants and cladocerans over regional (up to 300 km) and continental (up to 3300 km) scales. Both taxonomic groups were surveyed in five different European regions and presented extremely high levels of beta‐diversity, both within and among regions. High beta‐diversity was primarily explained by species replacement (turnover) rather than differences in species richness (i.e. nestedness). Abiotic and biotic variables were the main drivers of community composition. Within some regions, small‐scale connectivity and the spatial configuration of sampled communities explained a significant, though smaller, fraction of compositional variation, particularly for aquatic plants. At continental scale (among regions), a significant fraction of compositional variation was explained by a combination of spatial effects (exclusive contribution of regions) and regionally‐structured environmental variables. Our results suggest that, although dispersal limitation might affect species composition in some regions, aquatic plant and cladoceran communities are not generally limited by dispersal at the regional scale (up to 300 km). Species sorting mediated by environmental variation might explain the high species turnover of aquatic plants and cladocerans at regional scale, while biogeographic processes enhanced by dispersal limitation among regions might determine the composition of regional biotas.     

Environmental drivers of ant species richness and composition across the Argentine Pampas grassland

Understanding the underlying mechanisms causing diversity patterns is a fundamental objective in ecology and science‐based conservation biology. Energy and environmental‐heterogeneity hypotheses have been suggested to explain spatial changes in ant diversity. However, the relative roles of each one in determining alpha and beta diversity patterns remain elusive. We investigated the main factors driving spatial changes in ant (Hymenoptera, Formicidae) species richness and composition (including turnover and nestedness components) along a 500 km longitudinal gradient in the Pampean region of Argentina. Ants were sampled using pitfall traps in 12 sample sites during the summer. We performed a model selection approach to analyse responses of ant richness and composition dissimilarity to environmental factors. Then, we computed a dissimilarity partitioning of the contributions of spatial turnover and nestedness to total composition dissimilarity. Temporal habitat heterogeneity and temperature were the primary factors explaining spatial patterns of epigean ant species richness across the Pampas. The distance decay in species composition similarity was best accounted by temperature dissimilarity, and turnover had the greatest contribution to the observed beta diversity pattern. Our findings suggest that both energy and environmental‐heterogeneity‐related variables are key factors shaping richness patterns of ants and niche‐based processes instead of neutral processes appear to be regulating species composition of ant assemblages. The major contribution of turnover to the beta diversity pattern indicated that lands for potential reconversion to grassland should represent the complete environmental gradient of the Pampean region, instead of prioritizing a single site with high species richness.     

A meta‐analysis of nestedness and turnover components of beta diversity across organisms and ecosystems

Aim

The number of studies investigating the nestedness and turnover components of beta diversity has increased substantially, but our general understanding of the drivers of turnover and nestedness remains elusive. Here, we examined the effects of species traits, spatial extent, latitude and ecosystem type on the nestedness and turnover components of beta diversity.

Location

Global.

Time period

1968–2017.

Major taxa studied

From bacteria to mammals.

Methods

From the 99 studies that partition total beta diversity into its turnover and nestedness components, we assembled 269 and 259 data points for the pairwise and multiple site beta‐diversity metrics, respectively. Our data covered a broad variation in species dispersal type, body size and trophic position. The data were from freshwater, marine and terrestrial realms, and encompassed geographical areas from the tropics to near polar regions. We used linear modelling as a meta‐regression tool to analyse the data.

Results

Pairwise turnover, multiple site turnover and total beta diversity all decreased significantly with latitude. In contrast, multiple site nestedness showed a positive relationship with latitude. Beta‐diversity components did not generally differ among the realms. The turnover component and total beta diversity increased with spatial extent, whereas nestedness was scale invariant for pairwise metrics. Multiple site beta‐diversity components did not vary with spatial extent. Surprisingly, passively dispersed organisms had lower turnover and total beta diversity than flying organisms. Body size showed a relatively weak relationship with beta diversity but had important interactions with trophic position, thus also affecting beta diversity via interactive effects. Producers had significantly higher average pairwise turnover and total beta diversity than carnivores.

Main conclusions

The present results provide evidence that species turnover, being consistently the larger component of total beta diversity, and nestedness are related to the latitude of the study area and intrinsic organismal features. We showed that two beta‐diversity components had generally opposing patterns with regard to latitude. We highlight that beta‐diversity partition may give additional insights into the underlying causes of spatial variability in biotic communities compared with total beta diversity alone.     

西藏横断山区溪流细菌beta多样性组分对气候和水体环境的响应

理解沿环境或空间梯度的群落组成变化(即beta多样性)一直是生态学和保护生物学的中心问题, 且beta多样性的形成机制及其对环境的响应已成为当前生物多样性研究的热点问题。本文以西藏横断山区怒江和澜沧江两个流域入江溪流中的细菌为研究对象, 使用Baselga的beta多样性分解方法, 基于Sørensen相异性指数将细菌的beta多样性分解为周转(turnover)和嵌套(nestedness)两个组分, 探究了细菌beta多样性及其分解组分随海拔距离的分布模式, 并且衡量了环境、气候和空间因子的相对重要性。结果表明, 两个流域中细菌的群落结构显著不同。两个流域的细菌总beta多样性和周转组分随海拔距离的增加而增加, 周转组分占总beta多样性的比例较大。气候和环境因子是两个流域中细菌总beta多样性及周转过程的重要预测因子, 并且所有的显著因子均为正相关, 其中环境因子中相关性最高的为海拔距离(R ²= 0.408, P < 0.001), 而气候因子中相关性最高的为年均温差(R ²= 0.417, P < 0.001)。方差分解结果暗示嵌套组分主要受空间扩散的影响; 总beta多样性和周转组分在环境较恶劣的澜沧江主要受环境过滤的影响, 而在环境较温和的怒江主要受空间扩散和环境过滤的共同影响。此外, 较为恶劣的环境条件会增加细菌的总beta多样性和周转率, 并且会形成更强的环境筛选作用去影响细菌群落的物种组成。我们的研究表明对西藏横断山区水体细菌多样性的保护需要从整个流域入手, 而非少量的生物多样性热点地区。     

Contrasting patterns and mechanisms of spatial turnover for native and exotic freshwater fish in Europe

Aim We compare the distribution patterns of native and exotic freshwater fish in Europe, and test whether the same mechanisms (environmental filtering and/or dispersal limitation) govern patterns of decrease in similarity of native and exotic species composition over geographical distance (spatial species turnover). Locations Major river basins of Europe. Methods Data related to geography, habitat diversity, regional climate and species composition of native and exotic freshwater fish were collated for 26 major European river basins. We explored the degree of nestedness in native and exotic species composition, and quantified compositional similarity between river basins according to the beta‐sim (independent of richness gradient) and Jaccard (dependent of richness gradient) indices of similarity. Multiple regression on distance matrices and variation‐partitioning approaches were used to quantify the relative roles of environmental filtering and dispersal limitation in shaping patterns of decreasing compositional similarity over geographical distance. Results Native and exotic species exhibited significant nested patterns of species composition, indicating that differences in fish species composition between river basins are primarily the result of species loss, rather than species replacement. Both native and exotic compositional similarity decreased significantly with increasing geographical distance between river basins. However, gradual changes in species composition with geographical distance were found only for exotic species. In addition, exotic species displayed a higher rate of similarity decay (higher species turnover rate) with geographical distance, compared with native species. Lastly, the majority of explained variation in exotic compositional similarity was uniquely related to geography, whereas native compositional similarity was either uniquely explained by geography or jointly explained by environment and geography. Main conclusions Our study suggests that large‐scale patterns of spatial turnover for exotic freshwater fish in Europe are generated by human‐mediated dispersal limitation, whereas patterns of spatial turnover for native fish result from both dispersal limitation relative to historical events (isolation by mountain ranges, glacial history) and environmental filtering.     

Rethinking the relationship between nestedness and beta diversity: a comment on Baselga (2010)

Baselga [Partitioning the turnover and nestedness components of beta diversity. Global Ecology and Biogeography, 19 , 134–143, 2010] proposed pairwise (β_nes) and multiple‐site (β_NES) beta‐diversity measures to account for the nestedness component of beta diversity. We used empirical, randomly created and idealized matrices to show that both measures are only partially related to nestedness and do not fit certain fundamental requirements for consideration as true nestedness‐resultant dissimilarity measures. Both β_nes and β_NES are influenced by matrix size and fill, and increase or decrease even when nestedness remains constant. Additionally, we demonstrate that β_NES can yield high values even for matrices with no nestedness. We conclude that β_nes and β_NES are not true measures of the nestedness‐resultant dissimilarity between sites. Actually, they quantify how differences in species richness that are not due to species replacement contribute to patterns of beta diversity. Finally, because nestedness is a special case of dissimilarity in species composition due to ordered species loss (or gain), the extent to which differences in species composition is due to nestedness can be measured through an index of nestedness.     

Seasonal patterns of taxonomic and functional beta diversity in submerged macrophytes at a fine scale

Spatiotemporal variation in community composition is of considerable interest in ecology. However, few studies have focused on seasonal variation patterns in taxonomic and functional community composition at the fine scale. As such, we conducted seasonal high‐density sampling of the submerged macrophyte community in Hongshan Bay of Erhai Lake in China and used the generalized dissimilarity model (GDM) to evaluate the effects of environmental factors and geographic distance on taxonomic and functional beta diversity as well as corresponding turnover and nestedness components. At the fine scale, taxonomic turnover and nestedness as well as functional turnover and nestedness showed comparable contributions to corresponding taxonomic and functional beta diversity, with different importance across seasons. All taxonomic and functional dissimilarity metrics showed seasonal variation. Of note, taxonomic beta diversity was highest in summer and lowest in winter, while functional beta diversity showed the opposite pattern. Taxonomic and functional turnover showed similar change patterns as taxonomic and functional beta diversity. Taxonomic nestedness was low in summer and high in winter. Functional nestedness was also lower in summer. These results suggest that under extreme environmental conditions, both turnover and nestedness can exist at the fine scale and seasonal community composition patterns in submerged macrophytes should be considered. Future investigations on community assembly mechanisms should pay greater attention to long‐term dynamic characteristics and functional information.     

高原鼠兔干扰对高寒草甸植物物种和功能性状beta多样性的影响

采用野外空间多点同步取样,分析了高原鼠兔干扰对高寒草甸植物物种beta多样性和植物功能性状beta多样性的影响,确定了高原鼠兔干扰下高寒草甸植物物种和功能性状beta多样性的变化途径,分别提出了高原鼠兔干扰区域内,基于植物物种多样性和功能性状多样性的高寒草甸植物多样性维持策略。结果表明,高原鼠兔干扰使高寒草甸植物物种相似性显著降低了28.1%,植物功能相似性降低了28.7%。尽管高原鼠兔干扰没有改变高寒草甸植物物种和功能性状beta多样性的变化途径,且对植物物种和功能性状的嵌套组分不存在显著影响,但高原鼠兔干扰显著降低了植物物种和功能性状周转组分所占的比例,降幅分别为36.6%和34.3%。高原鼠兔干扰区域内,高寒草甸植物物种beta多样性的变化以周转为主导(周转占比81.4%;嵌套占比:18.6%),植物功能性状beta多样性的变化以嵌套为主导(嵌套占比64.9%;周转占比35.1%)。因此,针对划定的高原鼠兔干扰区,需要同时保护区域内所有高原鼠兔栖息地(多位点保护),以达到维持植物物种多样性的目的,而可以仅通过保护该区域内植物功能性状丰富的位点,即可维持较高的植物功能多样性。     

Historical climate‐change influences modularity and nestedness of pollination networks

The structure of species interaction networks is important for species coexistence, community stability and exposure of species to extinctions. Two widespread structures in ecological networks are modularity, i.e. weakly connected subgroups of species that are internally highly interlinked, and nestedness, i.e. specialist species that interact with a subset of those species with which generalist species also interact. Modularity and nestedness are often interpreted as evolutionary ecological structures that may have relevance for community persistence and resilience against perturbations, such as climate‐change. Therefore, historical climatic fluctuations could influence modularity and nestedness, but this possibility remains untested. This lack of research is in sharp contrast to the considerable efforts to disentangle the role of historical climate‐change and contemporary climate on species distributions, richness and community composition patterns. Here, we use a global database of pollination networks to show that historical climate‐change is at least as important as contemporary climate in shaping modularity and nestedness of pollination networks. Specifically, on the mainland we found a relatively strong negative association between Quaternary climate‐change and modularity, whereas nestedness was most prominent in areas having experienced high Quaternary climate‐change. On islands, Quaternary climate‐change had weak effects on modularity and no effects on nestedness. Hence, for both modularity and nestedness, historical climate‐change has left imprints on the network structure of mainland communities, but had comparably little effect on island communities. Our findings highlight a need to integrate historical climate fluctuations into eco‐evolutionary hypotheses of network structures, such as modularity and nestedness, and then test these against empirical data. We propose that historical climate‐change may have left imprints in the structural organisation of species interactions in an array of systems important for maintaining biological diversity.     

小兴安岭阔叶红松林地表甲虫Beta多样性

Beta多样性用来衡量集群内物种组成的变异性,可以被分解为空间物种转换和物种集群镶嵌两个组分,是揭示群落构建机制的重要基础。目前开展了较多的地上生态系统beta多样性研究,然而地下生态系统beta多样性进展缓慢。以小兴安岭凉水和丰林自然保护区为研究地区,于2015年8、10月采用陷阱法对阔叶红松林进行调查,揭示地表甲虫(步甲科、隐翅虫科、葬甲科)的beta多样性。结果表明:(1)凉水共发现39种、856只地表甲虫,丰林共发现43种、1182只地表甲虫。8月凉水明显具有较高的全部甲虫(三个科的总和)物种多样性和丰富度,10月正好相反。(2)凉水和丰林之间地表甲虫beta多样性的差异仅发现于8月的步甲科和葬甲科之间。(3)凉水和丰林地表甲虫的beta多样性主要由空间物种转换组成,物种集群镶嵌对beta多样性的贡献很小,说明地表甲虫物种组成变异主要由本地物种之间较高的转换引起。研究表明小兴安岭阔叶红松林地表甲虫的beta多样性主要由空间物种转换组成,在揭示群落构建机制过程中,其内部物种交换和环境调控不容忽视。     

The role of habitat and daily activity patterns in explaining the diversity of mountain Neotropical dung beetle assemblages

The interaction between land use and climate change is expected to strongly affect species distributions along high elevation landscapes. We aimed to test the effect of climatic variables on community metrics among five types of land use in a high elevation landscape. We described dung beetle spatial and temporal taxonomic and functional diversity patterns, and partitioned β‐diversity into turnover and nestedness components. The interaction between land use and daily period of activity mostly drives abundance, functional richness and functional diversity, but not dung beetle species richness. Unlike Neotropical lowlands, species richness and abundance in open environments are similar to those existing in forests. Temperature is an important predictor of abundance and functional divergence. There is a higher spatial component of the taxonomic β‐diversity, which is highly driven by species turnover. The temporal component of the taxonomic β‐diversity was strongly driven by nestedness, where night assemblages are sub‐sets, although not entirely, of diurnal assemblages. For functional diversity, the temporal β‐diversity was much higher than the spatial β‐diversity, but both were similarly represented by functional group turnover and nestedness. The composition of nocturnal and diurnal assemblages is clearly different, even more than the differences observed between habitats. However, taxonomic turnover is the dominant force between sampling sites while nestedness dominates the daily pattern. This means that forest habitats are unlikely to act as shelters for grassland species under a scenario of rising temperature.     

Differential response to abiotic stress controls species distributions at biogeographic transition zones

Understanding range limits is critical to predicting species responses to climate change. Subtropical environments, where many species overlap at their range margins, are cooler, more light‐limited and variable than tropical environments. It is thus likely that species respond variably to these multi‐stressor regimes and that factors other than mean climatic conditions drive biodiversity patterns. Here, we tested these hypotheses for scleractinian corals at their high‐latitude range limits in eastern Australia and investigated the role of mean climatic conditions and of parameters linked to abiotic stress in explaining the distribution and abundance of different groups of species. We found that environmental drivers varied among taxa and were predominantly linked to abiotic stress. The distribution and abundance of tropical species and gradients in species richness (alpha diversity) and turnover (beta diversity) were best explained by light limitation, whereas minimum temperatures and temperature fluctuations best explained gradients in subtropical species, species nestedness and functional diversity. Variation in community structure (considering species composition and abundance) was most closely linked to the combined thermal and light regime. Our study demonstrates the role of abiotic stress in controlling the distribution of species towards their high‐latitude range limits and suggests that, at biogeographic transition zones, robust predictions of the impacts of climate change require approaches that account for various aspects of physiological stress and for species abundances and characteristics. These findings support the hypothesis that abiotic stress controls high‐latitude range limits and caution that projections solely based on mean temperature could underestimate species’ vulnerabilities to climate change.     

Species and individual replacements contribute more than nestedness to shape vertebrate scavenger metacommunities

Understanding the mechanisms that organize biodiversity is central in ecology and conservation. Beta diversity links local (alfa) and regional (gamma) diversity, giving insight into how communities organize spatially. Metacommunity ecology provides the framework to interpret regional and local processes interacting to shape communities. However, the lack of metacommunity studies for large vertebrates may limit the understanding and compromise the preservation of ecosystem functions and services. We aim to understand the mechanisms underlying differences in species composition among vertebrate scavenger communities ? which provide key ecosystem functions, e.g. carrion consumption ? within a metacommunity context. We obtained species richness and abundances at scavenger communities consuming ungulate carcasses monitored through motion‐triggered remote cameras in seven terrestrial ecosystems in Spain. We partitioned beta diversity to decompose incidence‐based (species presence/absence) and abundance‐based dissimilarities into their components (turnover/balanced variation and nestedness/abundance gradient, respectively). We identified the environmental factors explaining the observed patterns. The vertebrate scavenger metacommunity consisted of 3101 individuals from 30 species. Changes in composition among ecosystems were mostly (> 84%) due to species or individual replacement (i.e. turnover or balanced variation). Species or individual loss/gain (i.e. nestedness or abundance gradient) accounted for 13–16% of these changes. Mean carcass weight, elevation and habitat diversity were the main factors explaining species/individual replacement. Our findings suggest that local processes such as species‐sorting through habitat heterogeneity would dominate scavenger metacommunity dynamics together with stochastic forces (i.e. related to carrion unpredictability and scavenging being a widespread strategy among vertebrates). The presence of structured patterns (i.e. nestedness) in beta diversity could reflect a role of deterministic processes: mass‐effects through dispersal and defaunation. Vultures are long‐distance foragers and functionally dominant species, which would connect local assemblages within the metacommunity, supporting scavenger diversity and functions across space. These results highlight the importance of managing vertebrate scavenger assemblages within a metacommunity context.     

Patterns and drivers of Nothobranchius killifish diversity in lowland Tanzania

Temporary pools are seasonal wetland habitats with specifically adapted biota, including annual Nothobranchius killifishes that survive habitat desiccation as diapausing eggs encased in dry sediment. To understand the patterns in the structure of Nothobranchius assemblages and their potential in wetland conservation, we compared biodiversity components (alpha, beta, and gamma) between regions and estimated the role and sources of nestedness and turnover on their diversity. We sampled Nothobranchius assemblages from 127 pools across seven local regions in lowland Eastern Tanzania over 2 years, using dip net and seine nets. We estimated species composition and richness for each pool, and beta and gamma diversity for each region. We decomposed beta diversity into nestedness and turnover components. We tested nestedness in three main regions (Ruvu, Rufiji, and Mbezi) using the number of decreasing fills metric and compared the roles of pool area, isolation, and altitude on nestedness. A total of 15 species formed assemblages containing 1–6 species. Most Nothobranchius species were endemic to one or two adjacent regions. Regional diversity was highest in the Ruvu, Rufiji, and Mbezi regions. Nestedness was significant in Ruvu and Rufiji, with shared core (N. melanospilus, N. eggersi, and N. janpapi) and common (N. ocellatus and N. annectens) species, and distinctive rare species. Nestedness apparently resulted from selective colonization rather than selective extinction, and local species richness was negatively associated with altitude. The Nothobranchius assemblages in the Mbezi region were not nested, and had many endemic species and the highest beta diversity driven by species turnover. Overall, we found unexpected local variation in the sources of beta diversity (nestedness and turnover) within the study area. The Mbezi region contained the highest diversity and many endemic species, apparently due to repeated colonizations of the region rather than local diversification. We suggest that annual killifish can serve as a flagship taxon for small wetland conservation.     

Historical legacies in world amphibian diversity revealed by the turnover and nestedness components of Beta diversity

Historic processes are expected to influence present diversity patterns in combination with contemporary environmental factors. We hypothesise that the joint use of beta diversity partitioning methods and a threshold-based approach may help reveal the effect of large-scale historic processes on present biodiversity. We partitioned intra-regional beta diversity into its turnover (differences in composition caused by species replacements) and nestedness-resultant (differences in species composition caused by species losses) components. We used piecewise regressions to show that, for amphibian beta diversity, two different world regions can be distinguished. Below parallel 37, beta diversity is dominated by turnover, while above parallel 37, beta diversity is dominated by nestedness. Notably, these regions are revealed when the piecewise regression method is applied to the relationship between latitude and the difference between the Last Glacial Maximum (LGM) and the present temperature but not when present energy-water factors are analysed. When this threshold effect of historic climatic change is partialled out, current energy-water variables become more relevant to the nestedness-resultant dissimilarity patterns, while mountainous areas are associated with higher spatial turnover. This result suggests that nested patterns are caused by species losses that are determined by physiological constraints, whereas turnover is associated with speciation and/or Pleistocene refugia. Thus, the new threshold-based view may help reveal the role of historic factors in shaping present amphibian beta diversity patterns.     

Amphibian Beta Diversity in the Brazilian Atlantic Forest: Contrasting the Roles of Historical Events and Contemporary Conditions at Different Spatial Scales

Current patterns of biodiversity distribution result from a combination of historical and contemporary processes. Here, we compiled checklists of amphibian species to assess the roles of long-term climate stability (Quaternary oscillations), contemporary environmental gradients and geographical distance as determinants of change in amphibian taxonomic and phylogenetic composition in the Brazilian Atlantic Forest. We calculated beta diversity as both variation in species composition (CBD) and phylogenetic differentiation (PBD) among the assemblages. In both cases, overall beta diversity was partitioned into two basic components: species replacement and difference in species richness. Our results suggest that the CBD and PBD of amphibians are determined by spatial turnover. Geographical distance, current environmental gradients and long-term climatic conditions were complementary predictors of the variation in CBD and PBD of amphibian species. Furthermore, the turnover components between sites from different regions and between sites within the stable region were greater than between sites within the unstable region. On the other hand, the proportion of beta-diversity due to species richness difference for both CBD and PBD was higher between sites in the unstable region than between sites in the stable region. The high turnover components from CBD and PBD between sites in unstable vs stable regions suggest that these distinct regions have different biogeographic histories. Sites in the stable region shared distinct clades that might have led to greater diversity, whereas sites in the unstable region shared close relatives. Taken together, these results indicate that speciation, environmental filtering and limited dispersal are complementary drivers of beta-diversity of amphibian assemblages in the Brazilian Atlantic Forest.