Dispersal limitation and environmental heterogeneity shape scale-dependent diversity patterns in plant communities

Understanding the large-scale distribution of species diversity requires distinguishing two of the primary factors that cause compositional differences: dispersal limitation and environmental variation. In a community with a naturally discontinuous spatial structure, we asked (1) at what scale(s) nonrandom variation in species composition occurs and (2) at what scale(s) such variation is associated with spatial separation, indicative of dispersal limitation, and at what scale(s) variation is associated with environmental heterogeneity? We sampled 50 seeps (small wetlands) on five serpentine outcrops. Using a randomization model, we showed that additive beta diversity (a measure of community dissimilarity) was lower than random within seeps and higher than random among both seeps and outcrops. Using Mantel tests, we showed that plant community dissimilarity, in both the full seep assemblage as well as in a subset of seep endemics, at the two larger scales was associated with different forms of environmental heterogeneity and, at the largest scale, was also associated with geographic distance. We conclude that diversity in this system is shaped by multiple scales of heterogeneity and by dispersal limitation at the largest scale.     

Genotypic richness and phenotypic dissimilarity enhance population performance

Increases in biodiversity can result from an increase in species richness, as well as from a higher genetic diversity within species. Intraspecific genetic diversity, measured as the number of genotypes, can enhance plant primary productivity and have cascading effects at higher trophic levels, such as an increase in herbivore and predator richness. The positive effects of genotypic mixtures are not only determined by additive effects, but also by interactions among genotypes, such as facilitation or inhibition. However, so far there has been no effort to predict the extent of such effects. In this study, we address the question of whether the magnitude of the effect of genotype number on population performance can be explained by the extent of dissimilarity in key traits among genotypes in a mixture. We examine the relative contribution of genotype number and phenotypic dissimilarity among genotypes to population performance of the soil arthropod, Orchesella cincta. Nearly homogeneous genotypes were created from inbred isofemale lines. Phenotypic dissimilarity among genotypes was assessed in terms of three life-history traits that are associated with population growth rate, i.e., egg size, egg development time, and juvenile growth rate. A microcosm experiment with genotype mixtures consisting of one, two, four, and eight genotypes, showed that genotypic richness strongly increased population size and biomass production and was associated with greater net diversity effects. Most importantly, there was a positive log-linear relationship between phenotypic dissimilarity in a mixture and the net diversity effects for juvenile population size and total biomass. In other words, the degree of phenotypic dissimilarity among genotypes determined the magnitude of the genotypic richness effect, although this relationship leveled off at higher values of phenotypic dissimilarity. Although the exact mechanisms responsible for these effects are currently unknown, similar advantages of trait dissimilarity have been found among species. Hence, to better understand population performance, genotype number and phenotypic dissimilarity should be considered collectively.     

Measuring biological diversity

The diversity of a set of species refers to the joint dissimilarity of the species in the set. This paper discusses the measurement of diversity from the set of pairwise distances between the species in the set. A measure called the effective number of species is developed from a non-parametric probability inequality and is shown to have a simple interpretation in terms of comparing linear experiments.     

Conserving alpha and beta diversity in wood-production landscapes

International demand for wood and other forest products continues to grow rapidly, and uncertainties remain about how animal communities will respond to intensifying resource extraction associated with woody bioenergy production. We examined changes in alpha and beta diversity of bats, bees, birds, and reptiles across wood production landscapes in the southeastern United States, a biodiversity hotspot that is one of the principal sources of woody biomass globally. We sampled across a spatial gradient of paired forest land-uses (representing pre and postharvest) that allowed us to evaluate biological community changes resulting from several types of biomass harvest. Short-rotation practices and residue removal following clearcuts were associated with reduced alpha diversity (−14.1 and −13.9 species, respectively) and lower beta diversity (i.e., Jaccard dissimilarity) between land-use pairs (0.46 and 0.50, respectively), whereas midrotation thinning increased alpha (+3.5 species) and beta diversity (0.59). Over the course of a stand rotation in a single location, biomass harvesting generally led to less biodiversity. Cross-taxa responses to resource extraction were poorly predicted by alpha diversity: correlations in responses between taxonomic groups were highly variable (−0.2 to 0.4) with large uncertainties. In contrast, beta diversity patterns were highly consistent and predictable across taxa, where correlations in responses between taxonomic groups were all positive (0.05–0.4) with more narrow uncertainties. Beta diversity may, therefore, be a more reliable and information-rich indicator than alpha diversity in understanding animal community response to landscape change. Patterns in beta diversity were primarily driven by turnover instead of species loss or gain, indicating that wood extraction generates habitats that support different biological communities.     

Spatial clustering of habitat structure effects patterns of community composition and diversity

Natural ecosystems often show highly productive habitats that are clustered in space. Environmental disturbances are also often nonrandomly distributed in space and are either intrinsically linked to habitat quality or independent in occurrence. Theoretical studies predict that configuration and aggregation of habitat patch quality and disturbances can affect metacommunity composition and diversity, but experimental evidence is largely lacking. In a metacommunity experiment, we tested the effects of spatially autocorrelated disturbance and spatial aggregation of patch quality on regional and local richness, among-community dissimilarity, and community composition. We found that spatial aggregation of patch quality generally increased among-community dissimilarity (based on two measures of beta diversity) of communities containing protozoa and rotifers in microcosms. There were significant interacting effects of landscape structure and location of disturbances on beta diversity, which depended in part on the specific beta diversity measures used. Effects of disturbance on composition and richness in aggregated landscapes were generally dependent on distance and connectivity among habitat patches of different types. Our results also show that effects of disturbances in single patches cannot directly be extrapolated to the landscape scale: the predictions may be correct when only species richness is considered, but important changes in beta diversity may be overlooked. There is a need for biodiversity and conservation studies to consider the spatial aggregation of habitat quality and disturbance, as well as connectivity among spatial aggregations.     

Hydrologic connection between ponds positively affects macrophyte alpha and gamma diversity but negatively affects beta diversity

Connections between habitat patches can positively influence the number of species in respective patches, providing a basis for preferentially conserving interconnected patches. However, from a regional perspective, it is not known whether conserving multiple sets of interconnected habitat patches would include more species (i.e., show higher gamma diversity) than conserving multiple, unconnected, solitary patches. We studied aquatic macrophytes in 15 sets of unidirectionally interconnected ponds and 19 unconnected ponds and also tested whether alpha and beta diversity, expressed as the number of species and dissimilarity in species composition, respectively, differed between connected and unconnected ponds. We found that gamma diversity was higher in connected ponds than in unconnected ponds, even after controlling for surface area. This resulted from a higher alpha diversity in connected ponds, despite lower beta diversity. These results suggest that connections between habitat patches positively influence diversity at both local and regional scales. When the total surface area available for conservation is limited, interconnected habitat patches should be preferentially conserved.     

The apportionment of quadratic entropy: a useful alternative for partitioning diversity in ecological data

Many methods that study the diversity within hierarchically structured populations have been developed in genetics. Among them, the analysis of molecular variance (AMOVA) (Excoffier et al., 1992) has the advantage of including evolutionary distances between individuals. AMOVA is a special case of a far more general statistical scheme produced by Rao (1982a; 1986) and called the apportionment of quadratic entropy (APQE). It links diversity and dissimilarity and allows the decomposition of diversity according to a given hierarchy. We apply this framework to ecological data showing that APQE may be very useful for studying diversity at various spatial scales. Moreover, the quadratic entropy has a critical advantage over usual diversity indices because it takes into account differences between species. Finally, the differences that can be incorporated in APQE may be either taxonomic or functional (biological traits), which may be of critical interest for ecologists.     

Tree species diversity and its relationship to stand parameters and geomorphology features in the eastern Black Sea region forests of Turkey

We studied the effects of stand parameters (crown closure, basal area, stand volume, age, mean stand diameter number of trees, and heterogeneity index) and geomorphology features (elevation, aspect and slope) on tree species diversity in an example of untreated natural mixed forest stands in the eastern Black Sea region of Turkey. Tree species diversity and basal area heterogeneity in forest ecosystems are quantified using the Shannon-Weaver and Simpson indices. The relationship between tree species diversity basal area heterogeneity stand parameters and geomorphology features are examined using regression analysis. Our work revealed that the relationship between tree species diversity and stand parameters is loose with a correlation coefficient between 0.02 and 0.70. The correlation of basal area heterogeneity with stand parameters fluctuated between 0.004 and 0.77 (R2). According to our results, stands with higher tree species diversity are characterised by higher mean stand diameter number of diameter classes, basal area and lower homogeneity index value. Considering the effect of geomorphology features on tree species or basal area heterogeneity we found that all investigated relationships are loose with R < or = 0.24. A significant correlation was detected only between tree species diversity and aspect. Future work is required to verify the detected trends in behaviour of tree species diversity if it is to estimate from the usual forest stand parameters and topography characteristics.     

Interactions among bacterial-feeding nematode species at different levels of food availability

Accurate prediction of the biodiversity–ecosystem functioning relationship requires adequate understanding of the interactions among species in a community. Effects of species diversity on ecosystem functioning are usually considered more pronounced with increasing functional dissimilarity, although species within functional groups may also perform non-identical functions and interact with each other. Here we present results of a laboratory experimental study aimed at elucidating whether interspecific interactions among species within a single nematode trophic group, bacterivores, (1) affect population development and community structure, and (2) depend on food availability. We studied the population growth of Rhabditis (Pellioditis) marina, a rhabditid nematode known to favour very high food densities when in monoculture, and of Diplolaimelloides meyli and D. oschei, congeneric Monhysteridae known to perform better in monocultures at intermediate food availability. Both Diplolaimelloides species showed significantly different patterns of food-density dependence in combination culture compared to monoculture. At very high food availability, the rhabditid nematode facilitated growth of both monhysterid species, probably as a result of down-regulation of bacterial density. At the lowest food availabilities, the presence of even low numbers of monhysterid nematodes lead to exclusion of the rhabditid, which at such low food availability has a very inefficient food uptake. At intermediate food availabilities, abundances of both Diplolaimelloides species were strongly depressed in the combination culture, as a result of food depletion by the rhabditid, indirect inhibitory interactions between the two congeneric species, or both. The complexity of the species interactions render predictions on the outcome and functional consequences of changes in within-trophic-group diversity highly problematic.     

扁刺栲-华木荷群系次生林林下物种多样性分析

对四川中亚热带扁刺栲(Castanopsis platyacantha)—华木荷(Schima sinensis)群系人工杉木林、水杉林、日本落叶松林和天然次生林林下植物进行了群落调查和物种多样性分析。结果表明：(1)杉木林、水杉林、日本落叶松林和次生扁刺栲—华木荷林(20a)的物种丰富度分别为62，59，53和32；Simpson多样性指数分别为7．95，7．08，9．24和5．38；Shannon—Wiener多样性指数分别为3．67，3．3l，3．64和2．46。(2)人工林林下植物群落之间的群落系数和相似度系数分别为0．45～0．60和0．62～0．82；人工林林下植物群落与次生阔叶林林下植物群落之间的群落系数和相似度系数分别为0．19～0．53和0．55～0．77。(3)人工林林下物种多样性比次生林高的主要原因可能是风倒木和站干死木导致人工林乔木层郁闭度减小，增加了林下的光照，从而改蛮了林下的微生境。表6参19。     

Inequality in paleorecords

Paleorecords provide information on past environmental variability, and help define ecological reference conditions by means of changes in their characteristics (accumulation rate, geochemical composition, density, etc.). A measure of temporal dissimilarity, which has traditionally been used in dendrochronology and is called "mean sensitivity," only focuses on first-order time-series lags. In this paper mean sensitivity was extended to all possible lags to derive a mean sensitivity function (MSF). The MSF is equivalent to a one-dimensional form of the paired relative madogram, a tool used in geostatistics to quantify spatial dependence. We then showed that the sum of madograms for all possible time-series lags is encapsulated by a single parameter, the Gini coefficient. This parameter has long been used by econometricians, social scientists, and ecologists as a synthetic, quantitative measure of inequality and diversity. Considering the connection between the MSF and the madogram, and the convenience of summarizing data heterogeneity with a single number, the Gini coefficient is therefore particularly appropriate for succinctly evaluating the diversity of paleorecords. An example of this application is provided by focusing on public domain dendrochronological data for the western conterminous United States.     

Are ecological groups of species optimal for forest dynamics modelling?

The huge diversity of tree species in tropical rain-forests makes the modelling of its dynamics a difficult task. One-way to deal with it is to define species groups. A classical approach for building species groups consists in grouping species with nearby characteristics, using cluster analysis. A group of species is then characterized by the same list of attributes as a single species, and it is incorporated in the model of forest dynamics in the same way as a single species. In this paper, a new approach for building species group is proposed. It relies on the discrepancy between model predictions when all species are considered separately, and model predictions when species groups are used. An aggregation error that quantifies the bias in model predictions that results from species grouping is thus defined. We then define the optimal species grouping as the one that minimizes the aggregation error. Using data from a tropical rain-forest in French Guiana and a toy model of forest dynamics, this new method for species grouping is confronted to the classical method based on cluster analysis of the species characteristics, and to a combined method based on a cluster analysis that uses the aggregation error as a dissimilarity between species. The optimal species grouping is quite different from the classical species grouping. The ecological interpretation of the optimal groups is difficult, as there is no direct linkage between the species characteristics and the way that they are grouped. The combined approach yields species groups that are closed to the optimal ones, with much less computations. The optimal species groups are thus specific to the model of forest dynamics and lack the generality of those of the classical method, that in turn are not optimal.     

Threatened Status, Rarity, and Diversity as Alternative Selection Measures for Protected Areas: A Test Using Afrotropical Antelopes

A major aim of conservation today is the maintenance of biodiversity. Practically, this pursuit might involve protecting a representative sample of the current biotic diversity (where diversity can have a variety of different meanings as in Vane-Wright et al. 1991), safeguarding species with traits that may be correlated with susceptibility to extinction (see International Council for Bird Preservation 1992), or protecting those species that are currently categorized as under short-term threat of extinction. Priority areas for conservation may vary, however, depending on which of these three approaches is taken. We investigated the designation of priority areas using these different approaches for Afrotropical antelope. Sites were selected on the basis of (1) biotic diversity—simple species richness and taxonomic diversity; (2) uniqueness of the fauna relative to other sites—how geographically restricted the component species were; and (3) degree of endangerment of the fauna. When insufficient sites to represent all the species could be selected, there was little agreement between the priority sites selected using the different methods. Sites selected by each approach were also generally poor at representing the diversity components ranked highly by other approaches. Also, many of the species were represented in only one site in the selected network, which on its own probably does not represent a viable population for the species. Therefore, it is important that the precise aims and consequences of any selection procedure be understood. A combination of different approaches, emphasizing different aspects of biodiversity and implemented sequentially, may be the best compromise for preserving a full range of biotic diversity.     

A method for assigning species into groups based on generalized Mahalanobis distance between habitat model coefficients

Habitat association models are commonly developed for individual animal species using generalized linear modeling methods such as logistic regression. We considered the issue of grouping species based on their habitat use so that management decisions can be based on sets of species rather than individual species. This research was motivated by a study of western landbirds in northern Idaho forests. The method we examined was to separately fit models to each species and to use a generalized Mahalanobis distance between coefficient vectors to create a distance matrix among species. Clustering methods were used to group species from the distance matrix, and multidimensional scaling methods were used to visualize the relations among species groups. Methods were also discussed for evaluating the sensitivity of the conclusions because of outliers or influential data points. We illustrate these methods with data from the landbird study conducted in northern Idaho. Simulation results are presented to compare the success of this method to alternative methods using Euclidean distance between coefficient vectors and to methods that do not use habitat association models. These simulations demonstrate that our Mahalanobis-distance-based method was nearly always better than Euclidean-distance-based methods or methods not based on habitat association models. The methods used to develop candidate species groups are easily explained to other scientists and resource managers since they mainly rely on classical multivariate statistical methods.     

Does plant species co-occurrence influence soil mite diversity?

Few studies have considered whether plant taxa can be used as predictors of belowground faunal diversity in natural ecosystems. We examined soil mite (Acari) diversity beneath six grass species at the Konza Prairie Biological Station, Kansas, USA. We tested the hypotheses that soil mite species richness, abundance, and taxonomic diversity are greater (1) beneath grasses in dicultures (different species) compared to monocultures (same species), (2) beneath grasses of higher resource quality (lower C:N) compared to lower resource quality, and (3) beneath heterogeneous mixes of grasses (C3 and C4 grasses growing together) compared to homogeneous mixes (C3 or C4 grasses) using natural occurrences of plant species as treatments. This study is the first to examine the interaction between above- and belowground diversity in a natural setting with species-level resolution of a hyper-diverse taxon. Our results indicate that grasses in diculture supported a more species and phylogenetically rich soil mite fauna than was observed for monocultures and that this relationship was significant at depth but not in the upper soil horizon. We noted that mite species richness was not linearly related to grass species richness, which suggests that simple extrapolations of soil faunal diversity based on plant species inventories may underestimate the richness of associated soil mite communities. The distribution of mite size classes in dicultures was considerably different than those for monocultures. There was no difference in soil mite richness between grass combinations of differing resource quality, or resource heterogeneity.     

入侵植物加拿大一枝黄花和本地一枝黄花的遗传多样性比较

为了阐明加拿大一枝黄花成功入侵的机制,利用简单序列重复区间标记(ISSR)方法对加拿大一枝黄花和本地一枝黄花的遗传多样性进行比较研究。从100条引物中筛选出12条引物用于PCR扩增,利用POPGEN32软件对2种一枝黄花进行遗传多样性分析。结果表明,加拿大一枝黄花在物种水平上的多态位点百分率为95.19%,Nei’s基因多样性指数为0.308 5,Shannon’s信息指数为0.415 8;本地一枝黄花在物种水平上的多态位点百分率(89.80%)、Nei’s基因多样性指数(0.249 1)和Shannon’s信息指数(0.383 4)都比加拿大一枝黄花小。加拿大一枝黄花和本地一枝黄花居群间遗传分化系数分别为0.118 2和0.131 3,居群内变异分别为0.881 8和0.868 7,表明2个物种居群间的遗传分化不明显,遗传一致度高,且主要的遗传变异存在于居群内。入侵植物加拿大一枝黄花具有较高遗传多样性,且高于本地一枝黄花,这可能是加拿大一枝黄花成功入侵的原因之一。     

Animals on marine flowers: does the presence of flowering shoots affect mobile epifaunal assemblage in an eelgrass meadow?

Eelgrass, Zostera marina, produces two types of shoots: morphologically simple vegetative shoots and highly branched flowering (reproductive) shoots, the latter found only in summer months. We examined whether the abundance and diversity of mobile epifaunal assemblage are affected by the presence of flowering shoots in an eelgrass meadow of Otsuchi Bay, northeastern Japan. Comparisons of epifauna in natural vegetation revealed that density and species richness did not differ significantly between sites consisting of both flowering and vegetative shoots, and those only of vegetative shoots. A transplant experiment, conducted to examine the colonization rates of epifauna to defaunated eelgrass planted with different combination of vegetative and flowering shoots, showed no obvious variation in abundance and species richness. At species level, the density of some species such as a tanaid Zeuxo sp. and a polychaete Platynereis sp. was higher at sites and/or treatments with flowering shoots, whereas that of some gastropods, such as Lirularia iridescens and Siphonacmea oblongata was higher at sites without flowering shoots. The species-specific response led to dissimilarity of epifaunal assemblage between sites and among treatments with different densities of vegetative and flowering shoots. Similar patterns observed for natural vegetation and the transplant experiment suggest that the variation in assemblage structure is caused by habitat selection of each species, for example, the utilization of flowering shoots as feeding ground and nursery by Zeuxo sp.     

南岳森林群落生物多样性研究(Ⅱ)广济寺森林群落植物物种多样性测定

本文根据实地调查资料及其统计分析结果,选用了24个物种多样性指数公式,并采用重要值、株数或鲜重以及优势度或盖度等作为测度指标,对南岳广济寺森林群落植物物种多样性进行了研究。结果表明,南岳广济寺森林属落多样性较高,接近山地季雨林的水平;此外,不同测度指标对物种多样性指数计算值并无明显的规律性。     

Beta diversity at different spatial scales: plant communities in organic and conventional agriculture.

Biodiversity studies that guide agricultural subsidy policy have generally compared farming systems at a single spatial scale: the field. However, diversity patterns vary across spatial scales. Here, we examined the effects of farming system (organic vs. conventional) and position in the field (edge vs. center) on plant species richness in wheat fields at three spatial scales. We quantified alpha-, beta-, and gamma-diversity at the microscale in 800 plots, at the mesoscale in 40 fields, and at the macroscale in three regions using the additive partitioning approach, and evaluated the relative contribution of beta-diversity at each spatial scale to total observed species richness. We found that alpha-, beta-, and gamma-diversity were higher in organic than conventional fields and higher at the field edge than in the field center at all spatial scales. In both farming systems, beta-diversity at the meso- and macroscale explained most of the overall species richness (up to 37% and 25%, respectively), indicating considerable differences in community composition among fields and regions due to environmental heterogeneity. The spatial scale at which beta-diversity contributed the most to overall species richness differed between rare and common species. Total richness of rare species (present in < or = 5% of total samples) was mainly explained by differences in community composition at the meso- and macroscale (up to 27% and 48%, respectively), but only in organic fields. Total richness of common species (present in > or = 25% of total samples) was explained by differences in community composition at the micro- and mesoscale (up to 29% and 47%, respectively), i.e., among plots and fields, independent of farming system. Our results show that organic farming made the greatest contribution to total species richness at the meso (among fields) and macro (among regions) scale due to environmental heterogeneity. Hence, agri-environment schemes should exploit this large-scale contribution of beta-diversity by tailoring schemes at regional scales to maximize dissimilarity between conservation areas using geographic information systems rather than focusing entirely at the classical local-field scale, which is the current practice.     

川西山区苹果属（Malus）植物野生种表型相似性和地理亲缘

用表型数量分析方法，结合地理分布对川西苹果属11个野生种进行研究，探讨其种间关系与地理分布的联系．得结论如下：1．不同系数的WPGMA和UPGMA聚类结果基本相同．表型数量骤类分析能大体反映近缘种间的亲缘关系；2．滇池海棠与沧江海棠在形态上高度相似，聚类图上极靠近，分布上相重叠，在塑定上有时感到困难；3．花叶海棠与变叶海棠和稻城海棠与小金海棠，从表型及分布上看显然是地理替代种；4．本文所列各种，大都分属两大植物区系范围内，例如花叶海棠属中国-日本区东范围内，变叶海棠属中国-喜马拉雅区系范围内；5．发现沿康定-理塘-巴塘（约（N）30°）存在一条南北物种分布的分界地带线，称康-巴线；沿岷江上游和大渡河流域也存在一条明显的地带线，称岷江-大渡河带，这一带就是中国-喜马拉雅区系和中国-日本区系的交界地带，通过这一地带有些向南分布的种可以越过康-巴线以东地区到达金沙江边，向北分布的种有时也可越过康-巴线以东达到岷江上游。在这南北和东西交界带上，不同物种会集，无疑是物种分化以及种质渗入十分活跃的地带；6．与其它地区相比较，四川西部山区是我国野生苹果植物分化的多样中心。