Community Completeness: Linking Local and Dark Diversity within the Species Pool Concept

Biodiversity of ecological communities has been examined widely. However, comparisons of observed species richness are limited because they fail to reveal what part of the differences are caused by natural variation in species pool size and what part is due to dark diversity – the absence of suitable species from a species pool. In other words, conventional biodiversity inventories do not convey information about how complete local plant communities are. We therefore propose the community completeness concept – a new perspective on the species pool framework. In order to ascertain community completeness, we need to estimate the extent of dark diversity, for which several methods are under development. We recommend the Community Completeness Index based on a log-ratio (or logistic) expression: ln(observed richness/dark diversity). This metric offers statistical advantages over other methods (e.g. the proportion of observed richness from the species pool). We discuss how community completeness can be related to long-term and successional community stability, landscape properties and disturbance patterns as well as to a variety of biotic interactions within and among trophic levels. The community completeness concept is related to but distinctive from the alpha-beta-gamma diversity approach and the community saturation phenomenon. The Community Completeness Index is a valuable metric for comparing biodiversity of different ecosystems for nature conservation. It can be used to measure the success of ecological restoration and vulnerability to invasion by alien species. In summary, community completeness is an interface between observed local observed species richness and dark diversity, which can be useful both in theoretical and applied biodiversity research.     

Dark diversity reveals importance of biotic resources and competition for plant diversity across habitats

Species richness is the most commonly used metric to quantify biodiversity. However, examining dark diversity, the group of missing species which can potentially inhabit a site, can provide a more thorough understanding of the processes influencing observed biodiversity and help evaluate the restoration potential of local habitats. So far, dark diversity has mainly been studied for specific habitats or large‐scale landscapes, while less attention has been given to variation across broad environmental gradients or as a result of local conditions and biotic interactions. In this study, we investigate the importance of local environmental conditions in determining dark diversity and observed richness in plant communities across broad environmental gradients. Using the ecospace concept, we investigate how these biodiversity measures relate to abiotic gradients (defined as position), availability of biotic resources (defined as expansion), spatiotemporal extent of habitats (defined as continuity), and species interactions through competition. Position variables were important for both observed diversity and dark diversity, some with quadratic relationships, for example, plant richness showing a unimodal response to soil fertility corresponding to the intermediate productivity hypothesis. Interspecific competition represented by community mean Grime C had a negative effect on plant species richness. Besides position‐related variables, organic carbon was the most important variable for dark diversity, indicating that in late‐succession habitats such as forests and shrubs, dark diversity is generally low. The importance of highly competitive species indicates that intermediate disturbance, such as grazing, may facilitate higher species richness and lower dark diversity.     

茂县土地岭植被恢复过程中物种多样性动态特征

植被恢复是退化生态系统重建的重要途径,植被恢复过程物种多样性的变化反映了植被的恢复程度.通过群落调查和多样性分析,研究了岷江上游土地岭植被恢复过程中群落物种多样性特征.结果表明： 恢复过程中6类不同类型群落分别表现其对于不同环境特征、干扰及更新方式等的响应;森林是较灌丛更适合当地环境状况的植被类型;人工恢复无干扰和轻度干扰群落的多样性相对较高,是较好的恢复模式.重度干扰使得1年生植物与地下芽植物比例增加,其它口食性较好的多年生草本减少.较强的干扰是群落无法更新、长期处于灌丛阶段且多样性较低的重要原因.本地区人工恢复群落在更新进程和多样性维持上优于自然更新群落,种植华山松加速了本地区植被演替进程.建议以适合恢复区域的多种恢复配置方式进行造林,并避免较强干扰,可以加速群落演替进程并保持恢复群落较高的物种丰富度与多样性.     

Hierarchical effects of environmental filters on the functional structure of plant communities: a case study in the French Alps

Understanding the influence of the environment on the functional structure of ecological communities is essential to predict the response of biodiversity to global change drivers. Ecological theory suggests that multiple environmental factors shape local species assemblages by progressively filtering species from the regional species pool to local communities. These successive filters should influence the various components of community functional structure in different ways. In this paper, we tested the relative influence of multiple environmental filters on various metrics of plant functional trait structure (i.e. ‘community weighted mean trait’ and components of functional trait diversity, i.e. functional richness, evenness and divergence) in 82 vegetation plots in the Guisane Valley, French Alps. For the 211 sampled species we measured traits known to capture key aspects of ecological strategies amongst vascular plant species, i.e. leaf traits, plant height and seed mass (LHS). A comprehensive information theory framework, together with null model based resampling techniques, was used to test the various environmental effects. Particular community components of functional structure responded differently to various environmental gradients, especially concerning the spatial scale at which the environmental factors seem to operate. Environmental factors acting at a large spatial scale (e.g. temperature) were found to predominantly shape community weighted mean trait values, while fine‐scale factors (topography and soil characteristics) mostly influenced functional diversity and the distribution of trait values among the dominant species. Our results emphasize the hierarchical nature of ecological forces shaping local species assemblage: large‐scale environmental filters having a primary effect, i.e. selecting the pool of species adapted to a site, and then filters at finer scales determining species abundances and local species coexistence. This suggests that different components of functional community structure will respond differently to environmental change, so that predicting plant community responses will require a hierarchical multi‐facet approach.     

Seed size- and density-related hidden treatments in common biodiversity experiments

Aims With a few exceptions, most well-known field biodiversity experiments on ecosystem functioning have been conducted in plant communities (especially grasslands) in which different numbers of species are planted as treatments. In these experiments, investigators have either kept the total seed weight or seed number constant across treatment plots. However, although in some cases attempts have been made to randomly choose species for planting from a designated species pool, the issue of possible 'hidden treatments' remains unsolved. Particularly, the total and relative abundance among species and across treatments could still affect the results. This study aims to determine whether treatments related to planted seed abundance and seed size may contribute to observed productivity.Methods We re-analyzed data from four biodiversity experiments based on a common seeding design (i.e. diversity treatments).Important findings We show that diversity (richness) treatments usually involve a hidden treatment related to the planted seeds (i.e. weight, number and seed size) that ultimately affect plant density. Thus, the un-intended hidden treatment of seeding more seeds on more diverse plots contributes to the productivity to some degree. Such derivative but often neglected hidden treatments are important for further improvement of experimental design and have significant implications in ecological restoration.     

怒江河谷入侵植物与乡土植物丰富度的分布格局与影响因子

外来物种入侵严重威胁着乡土植物多样性并削弱了生态系统服务功能。本文基于滇西北怒江河谷植被调查的样方数据, 从群落水平研究了乡土和入侵植物多样性的空间分布格局, 以及地形、气候、人类干扰等因子对两种格局的影响。本研究共记录到外来入侵植物26种, 隶属于13科21属; 乡土植物1,145种, 分属于158科628属。沿着怒江河谷, 入侵植物物种丰富度随纬度与海拔的增加而减少; 乡土物种丰富度则随纬度增加而增加, 并在海拔梯度上呈单峰格局。运用广义线性模型分析公路边缘效应(反映生境干扰)、气候、地形和土壤等环境因素对物种丰富度分布格局的影响。等级方差分离的结果显示, 公路两侧的生境干扰对入侵种和乡土种的丰富度格局均具有首要影响。在自然环境因子中, 降水量是入侵植物丰富度的主要限制因子, 而乡土物种丰富度则主要受到地形因子尤其是坡向的影响。结构方程模型的分析结果也表明, 乡土植物和入侵植物丰富度之间的负相关关系反映了二者对环境响应的差异。本文结果支持物种入侵的资源可利用性限制假说, 并强调了人类活动对生物多样性的负面影响; 乡土植物或已较好地适应了干旱河谷气候, 但并没有显示出对外来物种入侵的抵抗作用。     

水淹干扰下水库水滨带植物群落谱系结构与物种多样性格局

群落谱系结构将物种系统发育信息整合到群落中，给生物多样性研究提供了全新视角。为探讨水淹干扰下水库水滨带植物群落物种多样性维持机制，以丹江口水库水滨带为研究区域，以南水北调中线一期工程运行前的原有水滨带植被为研究对象，分析不同水淹干扰强度下群落谱系结构和物种多样性及其与环境因素的关系。结果表明：(1)随着水淹干扰强度增加，群落物种丰富度、物种多样性和谱系多样性显著降低。谱系结构指数虽无显著差异，但随着水淹干扰强度增加呈现由发散到聚集的变化趋势。表明水滨带植物群落物种多样性维持机制是生境过滤和生态位分化共同驱动的结果，并且随着水淹干扰强度增加呈现生境过滤作用增加、物种间竞争排斥作用减弱的生态过程影响规律。(2)坡度、土壤质地、速效氮和全氮含量对整个水滨带植物群落物种丰富度、物种多样性和谱系多样性影响显著。此外，土层厚度和速效磷对物种丰富度和土层厚度对谱系多样性也影响显著。而仅有速效氮含量对谱系结构指数影响显著；轻度水淹干扰下坡向、速效磷含量与多样性指数显著相关；中度水淹干扰下坡度、土层厚度、土壤质地、土壤总氮含量与多样性指数显著相关，土壤pH、全氮和全磷含量与谱系结构指数显著相关；重度水淹...     

More species,but all do the same: contrasting effects of flood disturbance on ground beetle functional and species diversity

The role of habitat disturbance on biodiversity is central as it promotes changes in ecological systems. That said, still little is known about the functional consequences of such changes. Functional diversity can be used to revealing more mechanistically the disturbance effects on communities by considering the richness and the distribution of traits among the species. Here we analyzed the response of functional and species diversity of ground beetles to flood disturbance to better understand the functioning of alluvial invertebrate communities. Ground beetles were sampled in periodically flooded grasslands along the Elbe River in Germany. We used generalized linear mixed effects models to unveil the relationships between flood disturbance, species and functional diversity, respectively. We measured different components of functional diversity (functional richness, evenness, dispersion, and divergence) and analyzed species diversity by means of rarefied species richness, abundances, evenness and Simpson's diversity. We found contrasting relationships in that most species diversity measures peaked at highest disturbance levels, while most functional diversity measures decreased with increasing disturbance intensities. Inversed relationships between species and functional diversity are rarely observed, as most studies report on positive correlations. We explain increasing species diversity with a higher amount of resources available in highly disturbed sites. Decreasing functional diversity is best explained through the convergence of species traits by flood disturbance and uneven resource exploitation in highly disturbed plots (low functional evenness), suggesting strong impacts from functionally different generalist species in floodchannels. We show that the amount of resources available, and how these resources are exploited, play major roles in the functioning of floodplain ground beetle communities.     

Small-scale plant species richness in calcareous grasslands determined by the species pool, community age and shoot density

Variation in small-scale plant species richness was examined between twenty discrete sites of calcareous alvar grasslands in the western Estonian coastal area. This community type is very species rich at the small-scale. The pattern of diversity has often been explained by variation in ecological conditions which makes it possible to avoid competitive exclusion. Here we test an alternative species pool hypothesis. The species pool was defined as the set of species present in a community, but excluding all those species whose presence may be connected with exceptional environmental conditions for this community. Monte Carlo modelling was used to determine the strengths of the relationships between the non-independent variables: richness vs species pool and average shoot density vs richness. Site age was characterized by its elevation above sea level, since there is a continuous land uplift in the study area. Species richness was positively correlated to the size of the species pool, community age and vegetation density. We conclude that historical processes on both the regional and local levels, which determine the arrival of any particular species to a target community, and not environmental heterogeneity, are responsible for the variations in species richness between communities of this type.     

Dispersal and neutral sampling mediate contingent effects of disturbance on plant beta‐diversity: a meta‐analysis

A major challenge in ecology, conservation and global‐change biology is to understand why biodiversity responds differently to similar environmental changes. Contingent biodiversity responses may depend on how disturbance and dispersal interact to alter variation in community composition (β‐diversity) and assembly mechanisms. However, quantitative syntheses of these patterns and processes across studies are lacking. Using null‐models and meta‐analyses of 22 factorial experiments in herbaceous plant communities across Europe and North America, we show that disturbance diversifies communities when dispersal is limited, but homogenises communities when combined with increased immigration from the species pool. In contrast to the hypothesis that disturbance and dispersal mediate the strength of niche assembly, both processes altered β‐diversity through neutral‐sampling effects on numbers of individuals and species in communities. Our synthesis suggests that stochastic effects of disturbance and dispersal on community assembly play an important, but underappreciated, role in mediating biotic homogenisation and biodiversity responses to environmental change.     

The relationship between local and regional diversity of indigenous forest fauna in KwaZulu-Natal Province, South Africa

The relationship between local and regional diversity was tested by regressing local community richness against regional species diversity for three taxa, birds, butterflies and mammals, in subtropical forest. The quadratic model best fits the relationship between local and regional diversity for birds. Local bird species richness is theoretically independent of the size of the regional pool of species and may represent saturated communities. A linear model best describes the relationship for mammals and butterflies. For mammals, the slope is shallow (0.264) and regional richness overestimates local species richness, suggesting communities are undersaturated. Extinction filtering may explain this pattern. Past climatic changes have filtered out many mammalian species, these changes have been too recent for autochthanous speciation, and the relatively low vagility of mammals has prevented extensive recolonisation. Differences in the nature of the diversity relationship between taxa are as much due to independent evolutionary histories as to differences in vagility and colonising potential. A pervasive role is suggested for regional biogeographic processes in the development of faunal assemblage structure. Large-scale processes are not considered in current conservation plans. We encourage the shift of conservation emphasis from local ecological processes and species interactions, to whole communities and consideration of regional processes.     

Constraints on colonization and species richness along a grassland productivity gradient: the role of propagule availability

An important goal in ecology is to discern under what habitat conditions community structure is primarily regulated by local ecological interactions and under what conditions community structure is more regulated by the pool of available colonists. I conducted a seed addition experiment in successional grassland to evaluate the relative significance of neighbourhood biotic interactions and propagule availability in regulating plant colonization and species richness along a natural gradient of grassland productivity. In undisturbed field plots, seed additions of 34 species led to an increase in species richness in locations of low productivity, an effect that declined in magnitude as productivity increased. In disturbed plots, seed additions led to a relatively constant increase in species richness at all levels of productivity. The results support the hypothesis that the role of propagule availability in regulating colonization dynamics and species richness declines in significance relative to local-scale competitive interactions as habitat productivity increases.     

Species pools and the "hump-back" model of plant species diversity: an empirical analysis at a relevant spatial scale

We investigated the relative roles of productivity, the species pool, and spatial habitat structure in determining local species richness (alpha diversity) of plant communities within a single, well-defined landscape unit, at spatial and ecological scales where the relationship between community productivity and species diversity often assumes a unimodal or "hump-back" form. At high levels of productivity, the decrease-phase of the unimodal model of the diversity-productivity relationship is typically explained as the dynamic outcome of increased competitive exclusion, but it may also be the passive consequence of a small pool of species possessing attributes necessary to competitively survive in high-fertility environments. We conducted statistical analyses of previously collected data to determine whether variations in local richness in the herbaceous vegetation of a Slovakian mountain valley were best explained by habitat productivity itself (which presumably leads to more intense competition) or by the sizes of the relevant community species pools. We also used measures of spatial habitat structure to investigate the extent to which habitat patchiness influenced patterns of species diversity. In the study system, both community biomass and size of the species pools contributed significantly to local species richness, but the positive effect of the species pools was about twice as important as the negative effect of biomass. The combined area of related associations (alliance area), association perimeter, and habitat patch geometry were all closely related to species pool size.     

Parallel responses of species and genetic diversity to El Niño Southern Oscillation-induced environmental destruction

Species diversity within communities and genetic diversity within species are two fundamental levels of biodiversity. Positive relationships between species richness and within-species genetic diversity have recently been documented across natural and semi-natural habitat islands, leading Vellend to suggest a novel macro-ecological pattern termed the species-genetic diversity correlation. We tested whether this prediction holds for areas affected by recent habitat disturbance using butterfly communities in east Kalimantan, Indonesia. Here, we show that both strong spatial and temporal correlations exist between species and allelic richness across rainforest habitats affected by El Niño Southern Oscillation-induced disturbance. Coupled with evidence that changes in species richness are a direct result of local extirpation and lower recruitment, these data suggest that forces governing variation at the two levels operate over parallel and short timescales, with implications for biodiversity recovery following disturbance. Remnant communities may be doubly affected, with reductions in species richness being associated with reductions in genetic diversity within remnant species.     

The Relationship between Local Species Richness and Species Pool: A Case Study from the High Mountains of the Greater Caucasus

The causes of linear relationships between local species richness and the size of the actual species pool in closed subalpine meadow communities and open plant communities of the alpine stony substrate (the Greater Caucasus Mountains) were analyzed using a computer simulation model. The results demonstrated that this relationship is insufficient evidence for the variation of local species richness among communities is wholly or partly determined by regional processes (the species-pool hypothesis). A relatively proportional ratio between these variables can also arise where local species richness and the size of the species pool both depend on local processes, or where local species richness is determined by local factors alone while the size of the species pool is determined by both local and regional factors.     

The Importance of Primary Tropical Rain Forest For Species Diversity: An Investigation Using Arboreal Ants as an example

The arboreal ant communities of a primary lowland rain forest and three differently disturbed forests that lie close together forming an anthropogenic disturbance gradient were collected with insecticide fogging. Combined samples from all trees (87 foggings) comprised 153,504 ants sorted to 331 morphospecies. The primary forest ant fauna was characterized by high species richness and 53 foggings were necessary to collect communities representatively. Another 63 species of ants were found in the disturbed forests indicating a large regional species pool that might exceed 420 species of arboreal ants. Anthropogenic disturbance caused a change in the taxonomic composition, diversity and structure of ant communities. Community size was a predictor of species richness in the severely disturbed forest types but not in the old secondary or primary forest. Ant abundance had declined significantly in the disturbed forests and only 10% of the primary forest’s species were collected in the most disturbed forest type. In each of the secondary forests a change in the frequency distribution of species was observed and a small number of species had gained numerical dominance. Analysis of species associations indicates that the strength of species interactions changed with the degree of forest disturbance. These changes were still clearly recognizable after 40 years of forest regeneration despite optimal conditions for colonization from the adjacent primary forest, demonstrating that the time scale needed for forest recovery after anthropogenic disturbance is very long.     

Reducing dispersal limitation via seed addition increases species richness but not above‐ground biomass

Seed dispersal limitation, which can be exacerbated by a number of anthropogenic causes, can result in local communities having fewer species than they might potentially support, representing a potential diversity deficit. The link between processes that shape natural variation in diversity, such as dispersal limitation, and the consequent effects on productivity is less well known. Here, we synthesised data from 12 seed addition experiments in grassland communities to examine the influence of reducing seed dispersal limitation (from 1 to 60 species added across experiments) on species richness and productivity. For every 10 species of seed added, we found that species richness increased by about two species. However, the increase in species richness by overcoming seed limitation did not lead to a concomitant increase in above‐ground biomass production. This highlights the need to consider the relationship between biodiversity and ecosystem functioning in a pluralistic way that considers both the processes that shape diversity and productivity simultaneously in naturally assembled communities.     

Quantitative partitioning of regional and local processes shaping regional diversity patterns

Much recent research explaining plant community diversity has focused on comparing the relative impacts of regional and local processes. We employed a novel analysis to quantify the effects of multiple regional and local processes on species richness, and to make quantitative comparisons of those effects across two sites that differ in plot-level species richness, productivity and environmental conditions. While abiotic stress and competition limited richness within the communities at both sites, only differences in the overall pool of species at the site, likely resulting from long-distance dispersal and climate fluctuations, explained the differences in plot-level richness between sites. Patterns in local richness may be driven by a temporal storage effect, with greater richness in the site with greater climatic variability. By identifying both the factors that impact diversity within communities and those that vary systematically across communities, our integrated approach provides a better understanding of regional diversity patterns.     

Indicators for plant species richness in pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) forests of Germany

Forestry is obliged to record as well as maintain and/or enhance biological diversity in forests due to national and international agreements. Accordingly, it is necessary to work out methodological approaches for the assessment of biodiversity in forests. In the study presented here, we focus on the total plant species pool (563 vascular plant and bryophyte species) of pine (Pinus sylvestris L.) forests in NE Germany to identify indicators for plant species richness. We distinguished several groups like “herb”, “grass-like”, “woody”, “endangered”, and “exotic species”, for which we detected indicators for low (class #1), intermediate (class #2), and high (class #3) species numbers. From a total of 84 species, which were identified by a three-step procedure, most indicators were found for class #3. Only few indicators have been revealed for intermediate species numbers, i.e. class #2. With help of Ellenberg’s ecological indicator values and information on the main occurrence in Central European vegetation types and plant communities, respectively, we characterized the indicator species ecologically. The ecological site preferences of the indicator species in general reflect the fact that species richness is highest in base-rich, light, and anthropogenically disturbed pine forests. On the contrary, species-poor forests were revealed by indicators, which mainly occur on acidic sites. It is concluded that a considerable set of indicators for species richness can help facilitate biodiversity assessments in forestry and ecosystem restoration practice. Electronic Supplementary Material  The online version of this article (doi: ) contains supplementary material, which is available to authorized users.     

Regional enrichment of local assemblages is robust to variation in local productivity, abiotic gradients, and heterogeneity

Theory predicts that the effects of regional richness on the richness of local communities may depend on the productivity, resource availability, and/or heterogeneity of local sites. Using the wetland plant communities of 50 independent streams as 'regions', we tested whether: (1) local richness in 1-m² quadrats and 50-m stream segments was positively related to regional richness, even after environmental influences were considered; and (2) the effect of regional richness would interact with the effects of biomass, soil moisture, and/or heterogeneity on local richness. In models that explained up to 88% of variation in local richness, we found that richness at both local scales was positively related to regional richness, and that regional richness did not interact with any of the environmental gradients that also shaped local richness. We conclude that species availability from the regional pool may consistently enrich local communities, even while other constraints on local richness operate.