The hypothesis of sympatric speciation as the dominant generator of endemism in a global hotspot of biodiversity

Allopatric or sympatric speciation influence the degree to which closely related species coexist in different manners, altering the patterns of phylogenetic structure and turnover among and between communities. The objective of this study was to examine whether phylogenetic community structure and turnover in the Brazilian Atlantic Forest permit conclusions about the dominant process for the formation of extant angiosperm richness of tree species. Therefore, we analyzed phylogenetic community structure (MPD, MNTD) as well as taxonomic (Jaccard similarity) and phylogenetic turnover (betaMPD, betaMNTD) among and between 49 tree communities distributed among three different habitat types. Mean annual precipitation and mean annual temperature in each survey area were estimated. Phylogenetic community structure does not differ between habitat types, although MPD reduces with mean annual temperature. Jaccard similarity decreases and betaMNTD increases with spatial distance and environmental differences between study sites. Spatial distance explains the largest portions of variance in the data, indicating dispersal limitation and the spatial aggregation of recently formed taxa, as betaMNTD is related to more recent evolutionary events. betaMPD, that is related to deep evolutionary splits, shows no spatial or environmental pattern, indicating that older clades are equally distributed across the Brazilian Atlantic Forest. While similarity pattern indicates dispersal limitations, the spatial turnover of betaMNTD is consistent with a high degree of sympatric speciation generating extant diversity and endemism in the Brazilian Atlantic Forest. More comprehensive approaches are necessary to reduce spatial sampling bias, uncertainties regarding angiosperm diversification patterns and confirm sympatric speciation as the dominant generator for the formation of extant species diversity in the Brazilian Atlantic Forest.     

黄土高原腹地人工林下草本层群落构建机制

群落构建机制是生态学研究的核心论题。生态位理论和中性理论是阐明群落构建的两种主要理论, 但这两种理论分别强调的环境过滤和扩散限制对群落构建的影响尚存争议。该研究以黄土高原人工林下草本层群落为研究对象, 将群落物种组成数据与地理空间、气候、地形以及生物因子相结合, 运用随机森林模型(RF)和基于距离矩阵的多元回归方法(MRM), 探究了各个因子的重要性, 并通过邻体主坐标矩阵(PCNM)和基于距离的冗余分析(db-RDA)分别筛选显著的空间结构和环境因子, 最后结合筛选出的PCNM特征值和环境变量进行变差分解。研究结果显示: 1)林下草本层群落组成相似性随地理距离和生境差异的增加而减小。2)地理距离与生境差异共同解释了群落组成相似性变化的47.8%。其中, 空间因子和环境因子的独立解释率分别为14.1%和9.8%, 两者的联合解释率为23.9%。3)地理距离和年降水量是引起研究区内物种组成变化的关键因子, 且地理距离的重要性大于年降水量。在黄土高原腹地, 扩散限制与环境过滤共同主导了人工林下草本层群落的构建过程。     

Nitrogen deposition mediates the effects and importance of chance in changing biodiversity

Nitrogen deposition is changing biodiversity on Earth. We need to understand the underlying mechanisms to conserve biodiversity better. Both selection and chance are potential mechanisms, and they may operate concurrently. Then, what are the respective effects of selection and chance, what is their relative importance and how do they change with increasing nitrogen deposition rate? Here, we performed a 6‐year nitrogen addition experiment (0–28 g N/m²/year) in a typical steppe ecosystem of Inner Mongolia to investigate the community structure of plants, bacteria and ammonia‐oxidizing Archaea (AOA). We developed an experimentally based calculation method to first separate the structural variations between plots into the effects of selection (S) and chance (C), and then calculate their relative importance. Our results showed that as nitrogen addition rate increased, S for both plants and bacteria increased, but C for plants first increased and then decreased, and C for bacteria also increased; meanwhile, both S and C for AOA changed nonlinearly. As nitrogen addition rate increased, the importance of chance decreased on the whole for all these communities, but it decreased nonlinearly for plants and bacteria, with a local increase at certain intermediate rates. At all treatments, the importance of chance was <0.5 for plants, but >0.5 for AOA. These results demonstrated that nitrogen deposition changed biodiversity by mediating the effects and importance of chance, implicating different strategies should be adopted in conserving biodiversity according to nitrogen deposition rate and community properties.     

Inferring the parameters of the neutral theory of biodiversity using phylogenetic information and implications for tropical forests

We develop a statistical method to infer the parameters of Hubbell's neutral model of biodiversity using data on local species abundances and their phylogenetic relatedness. This method uses the approximate Bayesian computation (ABC) approach, where the data are summarized into a small number of informative summary statistics. We used three statistics: the number of species in the sample, Shannon H index of evenness and Shao and Sokal's B ₁ index of phylogenetic tree imbalance. Our approach was found to outperform previous methods, illustrating the potential of ABC methods in ecology. Applying it to four large tropical forest tree data sets, the best-fit immigration rates m were found to be two orders of magnitude smaller and regional diversities θ larger than previously reported for the same data. This implies that neutral-compatible regional pools of tropical trees should extend over continental scales, and that m measures, in this context, mostly the frequency of long-distance dispersal events.     

Accounting for dispersal and biotic interactions to disentangle the drivers of species distributions and their abundances

Although abiotic factors, together with dispersal and biotic interactions, are often suggested to explain the distribution of species and their abundances, species distribution models usually focus on abiotic factors only. We propose an integrative framework linking ecological theory, empirical data and statistical models to understand the distribution of species and their abundances together with the underlying community assembly dynamics. We illustrate our approach with 21 plant species in the French Alps. We show that a spatially nested modelling framework significantly improves the model's performance and that the spatial variations of species presence-absence and abundances are predominantly explained by different factors. We also show that incorporating abiotic, dispersal and biotic factors into the same model bring new insights to our understanding of community assembly. This approach, at the crossroads between community ecology and biogeography, is a promising avenue for a better understanding of species co-existence and biodiversity distribution.     

Disentangling niche and neutral influences on community assembly: assessing the performance of community phylogenetic structure tests

Patterns of phylogenetic relatedness within communities have been widely used to infer the importance of different ecological and evolutionary processes during community assembly, but little is known about the relative ability of community phylogenetics methods and null models to detect the signature of processes such as dispersal, competition and filtering under different models of trait evolution. Using a metacommunity simulation incorporating quantitative models of trait evolution and community assembly, I assessed the performance of different tests that have been used to measure community phylogenetic structure. All tests were sensitive to the relative phylogenetic signal in species metacommunity abundances and traits; methods that were most sensitive to the effects of niche-based processes on community structure were also more likely to find non-random patterns of community phylogenetic structure under dispersal assembly. When used with a null model that maintained species occurrence frequency in random communities, several metrics could detect niche-based assembly when there was strong phylogenetic signal in species traits, when multiple traits were involved in community assembly, and in the presence of environmental heterogeneity. Interpretations of the causes of community phylogenetic structure should be modified to account for the influence of dispersal.     

Responses of species abundance distribution patterns to spatial scaling in subtropical secondary forests

To quantify and assess the processes underlying community assembly and driving tree species abundance distributions(SADs) with spatial scale variation in two typical subtropical secondary forests in Dashanchong state‐owned forest farm, two 1‐ha permanent study plots (100‐m × 100‐m) were established. We selected four diversity indices including species richness, Shannon–Wiener, Simpson and Pielou, and relative importance values to quantify community assembly and biodiversity. Empirical cumulative distribution and species accumulation curves were utilized to describe the SADs of two forests communities trees. Three types of models, including statistic model (lognormal and logseries model), niche model (broken‐stick, niche preemption, and Zipf‐Mandelbrodt model), and neutral theory model, were estimated by the fitted SADs. Simulation effects were tested by Akaike's information criterion (AIC) and Kolmogorov–Smirnov test. Results found that the Fagaceae and Anacardiaceae families were their respective dominance family in the evergreen broad‐leaved and deciduous mixed communities. According to original data and random sampling predictions, the SADs were hump‐shaped for intermediate abundance classes, peaking between 8 and 32 in the evergreen broad‐leaved community, but this maximum increased with size of total sampled area size in the deciduous mixed community. All niche models could only explain SADs patterns at smaller spatial scales. However, both the neutral theory and purely statistical models were suitable for explaining the SADs for secondary forest communities when the sampling plot exceeded 40 m. The results showed the SADs indicated a clear directional trend toward convergence and similar predominating ecological processes in two typical subtropical secondary forests. The neutral process gradually replaced the niche process in importance and become the main mechanism for determining SADs of forest trees as the sampling scale expanded. Thus, we can preliminarily conclude that neutral processes had a major effect on biodiversity patterns in these two subtropical secondary forests but exclude possible contributions of other processes.     

地理距离及环境差异对云南元江干热河谷植物群落beta多样性的影响

beta多样性反映了群落间物种组成的差异, 是生物多样性研究的热点之一。本研究通过对云南元江干热河谷41个植物群落样方进行调查, 用Jaccard相异系数表征物种beta多样性, 用样方之间的最近谱系距离(mean nearest taxon distance, MNTD)及平均谱系距离(mean pairwise distance, MPD)表征谱系beta多样性, 采用基于距离矩阵的多元回归和方差分解方法, 探讨了该区域干热河谷典型植物群落的物种beta多样性和谱系beta多样性与样方间环境差异(主要是气候)及地理距离之间的关系。结果表明: (1)群落间的地理距离和年平均温度差异对干热河谷植物群落的物种beta多样性和谱系beta多样性有显著影响; (2)地理距离对物种beta多样性和MNTD的影响最大; 地理距离和年平均温度差异对MPD的影响均较大; (3)样方间年平均温度与年平均降水量的差异和地理距离能够解释群落间beta多样性及谱系beta多样性11-13%的变异。以上结果表明, 生态位分化和扩散限制对该地区植物群落的beta多样性均有显著影响, 其中扩散限制的影响可能更大。此外, 人类活动等其他因素也很可能对元江干热河谷的群落组成具有非常重要的影响。     

Constraints on trait combinations explain climatic drivers of biodiversity: the importance of trait covariance in community assembly

Trade‐offs maintain diversity and structure communities along environmental gradients. Theory indicates that if covariance among functional traits sets a limit on the number of viable trait combinations in a given environment, then communities with strong multidimensional trait constraints should exhibit low species diversity. We tested this prediction in winter annual plant assemblages along an aridity gradient using multilevel structural equation modelling. Univariate and multivariate functional diversity measures were poorly explained by aridity, and were surprisingly poor predictors of community richness. By contrast, the covariance between maximum height and seed mass strengthened along the aridity gradient, and was strongly associated with richness declines. Community richness had a positive effect on local neighbourhood richness, indicating that climate effects on trait covariance indirectly influence diversity at local scales. We present clear empirical evidence that declines in species richness along gradients of environmental stress can be due to increasing constraints on multidimensional phenotypes.     

Insights into the historical construction of species-rich biomes from dated plant phylogenies, neutral ecological theory and phylogenetic community structure

Analytical methods are now available that can date all nodes in a molecular phylogenetic tree with one calibration, and which correct for variable rates of DNA substitution in different lineages. Although these techniques are approximate, they offer a new tool to investigate the historical construction of species-rich biomes. Dated phylogenies of globally distributed plant families often indicate that dispersal, even across oceans, rather than plate tectonics, has generated their wide distributions. By contrast, there are indications that animal lineages have undergone less long distance dispersal. Dating the origin of biome-specific plant groups offers a means of estimating the age of the biomes they characterize. However, rather than a simple emphasis on biome age, we stress the importance of studies that seek to unravel the processes that have led to the accumulation of large numbers of species in some biomes. The synthesis of biological inventory, systematics and evolutionary biology offered by the frameworks of neutral ecological theory and phylogenetic community structure offers a promising route for future work.     

Testing the ability of species distribution models to infer variable importance

Models of species’ distributions and niches are frequently used to infer the importance of range- and niche-defining variables. However, the degree to which these models can reliably identify important variables and quantify their influence remains unknown. Here we use a series of simulations to explore how well models can 1) discriminate between variables with different influence and 2) calibrate the magnitude of influence relative to an ‘omniscient’ model. To quantify variable importance, we trained generalized additive models (GAMs), Maxent and boosted regression trees (BRTs) on simulated data and tested their sensitivity to permutations in each predictor. Importance was inferred by calculating the correlation between permuted and unpermuted predictions, and by comparing predictive accuracy of permuted and unpermuted predictions using AUC and the continuous Boyce index. In scenarios with one influential and one uninfluential variable, models failed to discriminate reliably between variables when training occurrences were < 8–64, prevalence was > 0.5, spatial extent was small, environmental data had coarse resolution and spatial autocorrelation was low, or when pairwise correlation between environmental variables was |r| > 0.7. When two variables influenced the distribution equally, importance was underestimated when species had narrow or intermediate niche breadth. Interactions between variables in how they shaped the niche did not affect inferences about their importance. When variables acted unequally, the effect of the stronger variable was overestimated. GAMs and Maxent discriminated between variables more reliably than BRTs, but no algorithm was consistently well-calibrated vis-à-vis the omniscient model. Algorithm-specific measures of importance like Maxent's change-in-gain metric were less robust than the permutation test. Overall, high predictive accuracy did not connote robust inferential capacity. As a result, requirements for reliably measuring variable importance are likely more stringent than for creating models with high predictive accuracy.     

Testing the independent species' arrangement assertion made by theories of stochastic geometry of biodiversity

The assertion that the spatial location of different species is independent of each other is fundamental in major ecological theories such as neutral theory that describes a stochastic geometry of biodiversity. However, this assertion has rarely been tested. Here we use techniques of spatial point pattern analysis to conduct a comprehensive test of the independence assertion by analysing data from three large forest plots with different species richness: a species-rich tropical forest at Barro Colorado Island (Panama), a tropical forest in Sinharaja (Sri Lanka), and a temperate forest in Changbaishan (China). We hypothesize that stochastic dilution effects owing to increasing species richness overpower signals of species associations, thereby yielding approximate species independence. Indeed, the proportion of species pairs showing: (i) no significant interspecific association increased with species richness, (ii) segregation decreased with species richness, and (iii) small-scale interspecific interaction decreased with species richness. This suggests that independence may indeed be a good approximation in the limit of very species-rich communities. Our findings are a step towards a better understanding of factors governing species-rich communities and we propose a hypothesis to explain why species placement in species-rich communities approximates independence.     

The role of climate,water and biotic interactions in shaping biodiversity patterns in arid environments across spatial scales

Aim

Desert ecosystems, with their harsh environmental conditions, hold the key to understanding the responses of biodiversity to climate change. As desert community structure is influenced by processes acting at different spatial scales, studies combining multiple scales are essential for understanding the conservation requirements of desert biota. We investigated the role of environmental variables and biotic interactions in shaping broad and fine‐scale patterns of diversity and distribution of bats in arid environments to understand how the expansion of nondesert species can affect the long‐term conservation of desert biodiversity.

Location

Levant, Eastern Mediterranean.

Methods

We combine species distribution modelling and niche overlap statistics with a statistical model selection approach to integrate interspecific interactions into broadscale distribution models and fine‐scale analysis of ecological requirements. We focus on competition between desert bats and mesic species that recently expanded their distribution into arid environment following anthropogenic land‐use changes.

Results

We show that both climate and water availability limit bat distributions and diversity across spatial scales. The broadscale distribution of bats was determined by proximity to water and high temperatures, although the latter did not affect the distribution of mesic species. At the fine‐scale, high levels of bat activity and diversity were associated with increased water availability and warmer periods. Desert species were strongly associated with warmer and drier desert types. Range and niche overlap were high among potential competitors, but coexistence was facilitated through fine‐scale spatial partitioning of water resources.

Main conclusions

Adaptations to drier and warmer conditions allow desert‐obligate species to prevail in more arid environments. However, this competitive advantage may disappear as anthropogenic activities encroach further into desert habitats. We conclude that reduced water availability in arid environments under future climate change projections pose a major threat to desert wildlife because it can affect survival and reproductive success and may increase competition over remaining water resources.     

Global patterns in the shape of species geographical ranges reveal range determinants

Aim Do species range shapes follow general patterns? If so, what mechanisms underlie those patterns? We show for 11,582 species from a variety of taxa across the world that most species have similar latitudinal and longitudinal ranges. We then seek to disentangle the roles of climate, extrinsic dispersal limitation (e.g. barriers) and intrinsic dispersal limitation (reflecting a species’ ability to disperse) as constraints of species range shape. We also assess the relationship between range size and shape. Location Global. Methods Range shape patterns were measured as the slope of the regression of latitudinal species ranges against longitudinal ranges for each taxon and continent, and as the coefficient of determination measuring the degree of scattering of species ranges from the 1:1 line (i.e. latitudinal range = longitudinal range). Two major competing hypotheses explaining species distributions (i.e. dispersal or climatic determinism) were explored. To this end, we compared the observed slopes and coefficients of determination with those predicted by a climatic null model that estimates the potential range shapes in the absence of dispersal limitation. The predictions compared were that species distribution shapes are determined purely by (1) intrinsic dispersal limitation, (2) extrinsic dispersal limitations such as topographic barriers, and (3) climate. Results  Using this methodology, we show for a wide variety of taxa across the globe that species generally have very similar latitudinal and longitudinal ranges. However, neither neutral models assuming random but spatially constrained dispersal, nor models assuming climatic control of species distributions describe range shapes adequately. The empirical relationship between the latitudinal and longitudinal ranges of species falls between the predictions of these competing models. Main conclusions We propose that this pattern arises from the combined effect of macroclimate and intrinsic dispersal limitation, the latter being the major determinant among restricted‐range species. Hence, accurately projecting the impact of climate change onto species ranges will require a solid understanding of how climate and dispersal jointly control species ranges.     

太行山北段地区荆条灌丛和三裂绣线菊灌丛群落谱系结构

灌丛是太行山地区最重要的生态系统类型之一, 灌丛群落生物多样性的维持及其生态系统服务功能对京津冀地区的生态安全具有重要作用。本研究选择太行山最具代表性的两种灌木群落——荆条(Vitex negundo var. heterophylla)灌丛和三裂绣线菊(Spiraea trilobata)灌丛为研究对象, 利用净亲缘关系指数(net relatedness index, NRI)和多元回归等方法探究了两种灌丛群落构建机制的异同及主要的环境影响因子, 同时还利用谱系结构主坐标分析(principal coordinates of phylogenetic structure, PCPS)对决定群落谱系结构的关键系统发育节点进行了探究。结果表明: 两种灌丛群落内灌木植物的物种多样性没有显著差异, 但谱系结构差异显著。三裂绣线菊群落表现出显著的谱系发散趋势, 而荆条群落谱系聚集程度高于三裂绣线菊群落, 但未表现出显著的谱系结构。三裂绣线菊灌丛群落构建的驱动机制是生态位分化, 荆条灌丛中生境过滤作用有所增加, 与生态位分化共同驱动其群落构建过程。与干旱胁迫相关的生境过滤因素增加是荆条灌丛群落谱系聚集程度增加的重要原因。PCPS二维排序结果表明: 荆条灌丛群落谱系趋向聚集与其群落内缺乏蔷薇目、壳斗目等亲缘关系较远的类群有关, 而三裂绣线菊灌丛群落内物种则包含较多的演化分支。总体而言, 环境过滤不是决定太行山地区灌丛群落构建的主要驱动因素, 但水分条件仍然是影响区域群落谱系结构的重要因素。