Importance of temporal variability at different spatial scales for diversity of floodplain aquatic communities

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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。     

Environmental heterogeneity,species diversity and co‐existence at different spatial scales

The positive relationship between spatial environmental heterogeneity and species diversity is a widely accepted concept, generally associated with niche limitation. However, niche limitation cannot account for negative heterogeneity–diversity relationships (HDR) revealed in several case studies. Here we explore how HDR varies at different spatial scales and provide novel theories for small‐scale species co‐existence that explain both positive and negative HDR. At large spatial scales of heterogeneity (e.g. landscape level), different communities co‐exist, promoting large regional species pool size and resulting in positive HDR. At smaller scales within communities, species co‐existence can be enhanced by increasing the number of different patches, as predicted by the niche limitation theory, or alternatively, restrained by heterogeneity. We conducted meta‐regressions for experimental and observational HDR studies, and found that negative HDRs are significantly more common at smaller spatial scales. We propose three theories to account for niche limitation at small spatial scales. (1) Microfragmentation theory: with increasing spatial heterogeneity, large homogeneous patches lose area and become isolated, which in turn restrains the establishment of new plant individuals and populations, thus reducing species richness. (2) Heterogeneity confounded by mean: when heterogeneity occurs at spatial scales smaller than the size of individual plants, which forage through the patches, species diversity can be either positively or negatively affected by a change in the mean of an environmental factor. (3) Heterogeneity as a separate niche axis: the ability of species to tolerate heterogeneity at spatial scales smaller than plant size varies, affecting HDR. We conclude that processes other than niche limitation can affect the relationship between heterogeneity and diversity.     

Urbanization drives cross‐taxon declines in abundance and diversity at multiple spatial scales

The increasing urbanization process is hypothesized to drastically alter (semi‐)natural environments with a concomitant major decline in species abundance and diversity. Yet, studies on this effect of urbanization, and the spatial scale at which it acts, are at present inconclusive due to the large heterogeneity in taxonomic groups and spatial scales at which this relationship has been investigated among studies. Comprehensive studies analysing this relationship across multiple animal groups and at multiple spatial scales are rare, hampering the assessment of how biodiversity generally responds to urbanization. We studied aquatic (cladocerans), limno‐terrestrial (bdelloid rotifers) and terrestrial (butterflies, ground beetles, ground‐ and web spiders, macro‐moths, orthopterans and snails) invertebrate groups using a hierarchical spatial design, wherein three local‐scale (200 m × 200 m) urbanization levels were repeatedly sampled across three landscape‐scale (3 km × 3 km) urbanization levels. We tested for local and landscape urbanization effects on abundance and species richness of each group, whereby total richness was partitioned into the average richness of local communities and the richness due to variation among local communities. Abundances of the terrestrial active dispersers declined in response to local urbanization, with reductions up to 85% for butterflies, while passive dispersers did not show any clear trend. Species richness also declined with increasing levels of urbanization, but responses were highly heterogeneous among the different groups with respect to the richness component and the spatial scale at which urbanization impacts richness. Depending on the group, species richness declined due to biotic homogenization and/or local species loss. This resulted in an overall decrease in total richness across groups in urban areas. These results provide strong support to the general negative impact of urbanization on abundance and species richness within habitat patches and highlight the importance of considering multiple spatial scales and taxa to assess the impacts of urbanization on biodiversity.     

Monitoring large herbivore diversity at different scales: comparing direct and indirect methods

Monitoring of large herbivores is central to research and management activities in many protected areas. Monitoring programs were originally developed to estimate (trends in) population sizes of individual species. However, emphasis is shifting increasingly towards conservation of diversity and communities instead of individual species, as a growing literature shows the importance of herbivore diversity for ecosystem functioning. We argue that the design of monitoring programs has not yet been adapted well to this new conservation paradigm. Using large herbivore census data from Hluhluwe-iMfolozi Park, South Africa, we studied how monitoring methodology (observational counts vs. dung counts) and spatial scale interact in influencing estimates of large herbivore species richness and diversity. Dung counts resulted in higher herbivore species richness and diversity estimates than direct observational counts, especially at finer monitoring resolutions (grid cells smaller than 25 km²). At monitoring resolutions coarser than 25 km² both methods gave comparable diversity estimates. The methods also yielded different spatial diversity estimates, especially at finer resolutions. Grid cells with high diversity according to the dung count data did not necessarily have high diversity according to the observational counts, as shown by low correlation of grid cell values of both methods. We discuss these results in the light of estimates of the sampling effort of each method and, hence, suggest new monitoring designs that are more suitable for tracking temporal and spatial trends in large herbivore diversity and community composition.     

景观格局与植物多样性的关系及其空间尺度效应——以浑善达克沙地为例

景观格局是植物多样性的重要影响因素,为了探明沙地景观格局与植物多样性的关系及其空间尺度效应,以浑善达克沙地腹地为研究区,采用样方法进行植物群落学调查,基于GIS和遥感分析技术,分析景观格局指数,计算了10个尺度(100,200,300,400,500,600,700,800,900,1000m)下33个景观格局指数,利用双变量相关性分析法筛选出不同尺度下与植物多样性显著相关的指数;运用逐步回归的方法识别不同尺度下对植物多样性贡献显著的景观格局指数,并进行拟合检验。研究结果表明:景观格局指数对物种多样性影响具有显著的尺度依赖性。在所有空间尺度,景观格局指数主要对Shannon多样性指数和β指数影响较大;在100m尺度下,Shannon指数逐步回归R~2值达到最大(0.138);β指数在900m尺度下R2值达到最大值(0.2)。拟合检验发现,在500m尺度下,均匀度指数、Shannon多样性指数和β多样性指数实测值和预测值拟合均达到显著水平(P0.05),故认为500m尺度下,景观格局对植物多样性影响较大。因此,在不同尺度下管理和保护沙地植物多样性,应综合考虑景观格局指数的尺度效应,根据需要建立和优化有利于植物多样性的景观格局。     

Habitat amount drives the functional diversity and nestedness of anuran communities in an Atlantic Forest fragmented landscape

Natural environments disturbed by human activities can suffer from species extinctions, but some can still harbor high taxonomic diversity. However, disturbances may have impacts beyond the species level, if the species lost represent unique functions in the ecosystem. In this study, we evaluated to what extent the amount of habitat can determine the functional diversity and nestedness of amphibian communities in an Atlantic Forest fragmented landscape in Brazil, and if there is a threshold of habitat amount beyond which there is severe loss of functional diversity. As species responses may depend on their habitat type, we performed the analyses for three different sets of species: all species, forest‐dependent species, and generalist species. We also evaluated the relative importance of turnover and nestedness components to total functional dissimilarity among sites. Habitat amount affected functional diversity of frogs, especially for forest‐dependent species where a linear reduction was detected. The functional dissimilarity among sites was mostly explained by the nestedness component. The reduction of functional diversity was mediated by an ordered loss of traits, leading to a functionally nested metacommunity. These sensitive traits were closely related to habits and reproductive modes that depend on rivers and streams. The maintenance of functional diversity of frogs in fragmented landscapes must rely on the conservation of both terrestrial and aquatic environments, as some species and their traits can disappear from remnants of native vegetation lacking some specific habitats (e.g. streams). Abstract in Portuguese is available with online material.     

晋中盆地人类聚居地植物多样性分异规律

在城市化的发展过程中,生物多样性受到的影响日趋严重,因此开展这一方面的研究显得非常重要。以晋中盆地为研究对象,用随机抽样法选取86个村庄驻地、48个乡镇驻地及9个县市驻地,然后在各个驻地建成区内分别设置调查样方,运用Gleason丰富度指数、Whittaker Beta多样性指数、Jaccard相似性指数和t-检验对建成区内野生植物、栽培植物及总植物物种进行分析,同时探索总结植物多样性分异规律。结果表明:沿村庄-乡镇-县市梯度,野生植物平均丰富度降低,栽培植物平均丰富度增加,总植物物种平均丰富度先降低后增加;野生植物、栽培植物和总植物物种的β多样性指数均降低;植物相似度方面,村庄与乡镇的相似性最高,乡镇与县市次之,村庄与县市的相似性最低。同时,调查过程中发现,沿"村庄-乡镇-县市"梯度,城市化水平的差异与人为干扰强度的不同是引起盆地内植物多样性变化的重要原因。     

On the scaling patterns of species spatial distribution and association

Understanding how species distribution (occupancy and spatial autocorrelation) and association (that is, multi-species co-distribution) change across scales is fundamental to unlocking the pattern formation in population ecology and macroecology. Based on the Bayesian rule and join-count statistics, I present here a mathematical model that can demonstrate the effect of spatial scale on the observation of species distribution and association. Results showed that the intensity of spatial autocorrelation and species association declines when the grain in the spatial analysis increases, although the category of species distribution (aggregated or segregated) and association (positive or negative) remains the same. Random distribution and species independence were proved to be scale-free. Regardless of the possible patterns of species distribution and association, species tend to be randomly distributed and independent from each other when scaling-up (an increasing grain), reflecting a percolation process. This model, thus, grasps the statistical essence of species scaling pattern and presents a step forward for unveiling mechanisms behind species distributional and macroecological patterns.     

Environmental and spatial drivers of spider diversity at contrasting microhabitats

The relative importance of environmental and spatial drivers of animal diversity varies across scales, but identifying these scales can be difficult if a sampling design does not match the scale of the target organisms' interaction with their habitat. In this study, we quantify and compare the effects of environmental variation and spatial proximity on ground‐dwelling spider assemblages sampled from three distinct microhabitat types (open grassland, logs, trees) that recur across structurally heterogeneous grassy woodlands. We used model selection and multivariate procedures to compare the effects of different environmental attributes and spatial proximity on spider assemblages at each microhabitat type. We found that species richness and assemblage composition differed among microhabitat types. Bare ground cover had a negative effect on spider richness under trees, but a positive effect on spider richness in open grassland. Turnover in spider assemblages from open grassland was correlated with environmental distance, but not geographic distance. By contrast, turnover in spiders at logs and trees was correlated with geographic distance, but not environmental distance. Our study suggests that spider assemblages from widespread and connected open grassland habitat were more affected by environmental than spatial gradients, whereas spiders at log and tree habitats were more affected by spatial distance among these discrete but recurring microhabitats. Deliberate selection and sampling of small‐scale habitat features can provide robust information about the drivers of arthropod diversity and turnover in landscapes.     

毛乌素沙地植被不同恢复阶段植物群落物种多样性、功能多样性和系统发育多样性

进入21世纪以来, 中国荒漠化恢复取得显著成效, 荒漠化、沙化土地面积持续减少, 植被覆盖度大幅提升, 但关于植被恢复过程中生物多样性如何变化的研究不足, 这制约着对荒漠化恢复成效的全面评估。本文基于群落调查和叶功能性状(叶片厚度、叶片干物质含量、比叶面积和叶片密度)的测定, 分析了毛乌素沙地不同恢复阶段(半固定沙地、固定沙地、结皮覆盖沙地和草本植物覆盖沙地)的植物群落物种多样性、功能多样性和系统发育多样性特征。结果表明: (1)多数叶功能性状的系统发育信号不显著, 表明环境因子对研究区植物功能性状的塑造作用很强。(2)对于α多样性, 结皮覆盖沙地的物种多样性(Shannon-Wiener多样性, H)、物种丰富度(S)、功能丰富度(FRic)及系统发育多样性(PD)指数均显著低于其他恢复阶段, 而其他3个阶段间无显著差异; 这些指数间均显著正相关, 表明物种多样性、功能多样性和系统发育多样性在植被恢复过程中协同变化。(3) β多样性指数随恢复阶段间隔增加而逐渐增大, 表明物种组成、功能属性及系统发育关系随植被恢复持续变化, 且半固定沙地到固定沙地的群落物种组成、功能属性及系统发育关系更替最快, 导致群落间差异最大。(4)固定沙地、结皮覆盖沙地和草本植物覆盖沙地群落系统发育结构均趋向于发散, 表明竞争排斥是群落构建的主要驱动力; 而半固定沙地群落系统发育结构无一致规律, 表明群落构建可能受到生境过滤和竞争排斥的综合作用。研究结果可为植被建设与管理提供参考, 为毛乌素沙地生态保育和生物多样性的保护提供科学依据。     

Variation of plant diversity components in different scales in relation to grazing and climatic conditions

Background: Understanding the role of livestock grazing on plant diversity can be improved by an accurate measurement of diversity at all hierarchical scales due to the changeability of diversity components in space.

Aims: We evaluated the effects of grazing on plant species diversity at different scales of all common and rare species in two regions that have different climatic conditions (arid vs. semi-arid).

Methods: In each region, we collected abundant data of plant species from a nested sampling design that consisted of local (80 plots) and regional (16 sites) scales. We partitioned total species diversity (γ) into within plots (α_l), among plots (β_l) and among sites (β₂) using the additive partitioning.

Results: Diversity among sites contributed the most to total diversity for all and rare plant species in both regions. In addition, α₁ and β₁ diversities in ungrazed areas were greater than those in grazed areas for all and common species in both climates.

Conclusion: Abandonment of grazing after 10 years resulted in significant regeneration of common species at the local scale, with no change in rare species. We conclude that low grazing intensity is likely to be an important tool for conservation of plant diversity in which all scales should be considered.     

Amount and spatial distribution of habitats influence occupancy and dispersal of frogs at multiple scales in agricultural landscape

Agriculture changes the aquatic and terrestrial habitats used by animals, affecting their responses to matrix permeability. Here, we evaluated the impacts that resulted from the replacement of native vegetation with pastures on habitat occupancy, colonisation and local extinction of two Neotropical frogs with contrasting ecological strategies, Leptodactylus bufonius and L. chaquensis. We conducted fieldwork during two reproductive seasons in 50 temporary ponds in the Brazilian Chaco. We used site occupancy models in a multi-scale approach to identify landscape changes affecting population parameters and to determine the scale of interaction between species and the landscape. At local (10 m) and scales ≤400 m, increased pasture proportion limited the availability of bare soil required by males of L. bufonius to build mud chambers for reproduction and decreased proportion of shrublands affecting L. chaquensis occupancy. At larger spatial scales (>400 m), landscape modification limited dispersal of the smaller species L. bufonius. We found that the amount of habitat available is important in maintaining population parameters such as occupancy. However, our results highlight that the spatial distribution of habitats may also play an important role in the persistence and mobility of frogs in agricultural landscapes and that it is possible to identify a scale of effect in such anthropic landscapes. We recommend the consideration of reproductive and dispersal requirements of amphibian species, along with body size, as predictors of the spatial scale for management of populations in farmlands.