Moving beyond abundance distributions: neutral theory and spatial patterns in a tropical forest

Assessing the relative importance of different processes that determine the spatial distribution of species and the dynamics in highly diverse plant communities remains a challenging question in ecology. Previous modelling approaches often focused on single aggregated forest diversity patterns that convey limited information on the underlying dynamic processes. Here, we use recent advances in inference for stochastic simulation models to evaluate the ability of a spatially explicit and spatially continuous neutral model to quantitatively predict six spatial and non-spatial patterns observed at the 50 ha tropical forest plot on Barro Colorado Island, Panama. The patterns capture different aspects of forest dynamics and biodiversity structure, such as annual mortality rate, species richness, species abundance distribution, beta-diversity and the species–area relationship (SAR). The model correctly predicted each pattern independently and up to five patterns simultaneously. However, the model was unable to match the SAR and beta-diversity simultaneously. Our study moves previous theory towards a dynamic spatial theory of biodiversity and demonstrates the value of spatial data to identify ecological processes. This opens up new avenues to evaluate the consequences of additional process for community assembly and dynamics.     

Spatio-temporal dynamics of species richness in coastal fish communities

Determining patterns of change in species richness and the processes underlying the dynamics of biodiversity are of key interest within the field of ecology, but few studies have investigated the dynamics of vertebrate communities at a decadal temporal scale. Here, we report findings on the spatio-temporal variability in the richness and composition of fish communities along the Norwegian Skagerrak coast having been surveyed for more than half a century. Using statistical models incorporating non-detection and associated sampling variance, we estimate local species richness and changes in species composition allowing us to compute temporal variability in species richness. We tested whether temporal variation could be related to distance to the open sea and to local levels of pollution. Clear differences in mean species richness and temporal variability are observed between fjords that were and were not exposed to the effects of pollution. Altogether this indicates that the fjord is an appropriate scale for studying changes in coastal fish communities in space and time. The year-to-year rates of local extinction and turnover were found to be smaller than spatial differences in community composition. At the regional level, exposure to the open sea plays a homogenizing role, possibly due to coastal currents and advection.     

The role of spatial scale and the perception of large-scale species-richness patterns

Despite two centuries of exploration, our understanding of factors determining the distribution of life on Earth is in many ways still in its infancy. Much of the disagreement about governing processes of variation in species richness may be the result of differences in our perception of species‐richness patterns. Until recently, most studies of large‐scale species‐richness patterns assumed implicitly that patterns and mechanisms were scale invariant. Illustrated with examples and a quantitative analysis of published data on altitudinal gradients of species richness (n = 204), this review discusses how scale effects (extent and grain size) can influence our perception of patterns and processes. For example, a hump‐shaped altitudinal species‐richness pattern is the most typical (c. 50%), with a monotonic decreasing pattern (c. 25%) also frequently reported, but the relative distribution of patterns changes readily with spatial grain and extent. If we are to attribute relative impact to various factors influencing species richness and distribution and to decide at which point along a spatial and temporal continuum they act, we should not ask only how results vary as a function of scale but also search for consistent patterns in these scale effects. The review concludes with suggestions of potential routes for future analytical exploration of species‐richness patterns.     

空间尺度对地形异质性-物种多样性关系的影响:粒度和幅度

空间尺度是影响我们理解生态学格局和过程的关键因素.目前已有多种关于物种多样性分布格局形成机制的假说且研究者未达成共识,原因之一是空间尺度对物种多样性分布格局的环境影响因子的解释力和相对重要性有重要影响.地形异质性是物种多样性分布格局的重要影响因素.本文综述了在地形异质性-物种多样性关系的研究中,不同空间粒度和幅度对研究...     

哀牢山亚热带中山湿性常绿阔叶林树种beta多样性格局形成的驱动力

Beta多样性通常指群落在时间和空间上物种组成的差异, 包括物种周转组分和物种丰富度差异组分。驱动beta多样性格局形成的生态过程决定了群落的时空动态, 然而关于beta多样性及其两个组分格局形成的驱动力还存在较多争议。以往研究表明, beta多样性的格局存在取样尺度的依赖性, 驱动其形成的生态过程在不同取样尺度下的相对重要性也随之改变。本研究以哀牢山亚热带中山湿性常绿阔叶林20 ha动态监测样地为研究对象, 在不同取样尺度上, 将样方间的Bray-Curtis指数分解为物种周转组分和物种丰富度差异组分, 通过典范冗余分析和方差分解的方法揭示环境过滤和扩散限制对于beta多样性及其两个组分格局形成的相对重要性及其尺度依赖性。结果表明: (1) beta多样性、物种周转组分和物种丰富度差异组分均随取样尺度的增大而减小。在不同取样尺度下, 物种周转组分对于beta多样性的贡献始终占主导地位。(2)随着取样尺度的增大, 环境过滤驱动beta多样性格局形成的相对重要性逐渐增加, 而扩散限制的相对重要性逐渐降低。本研究进一步证实了取样尺度在beta多样性格局形成及其驱动力定量评价中的重要性, 今后的研究需要进一步解析上述尺度效应的形成机制。     

Common spatial patterns of trees in various tropical forests: Small trees are associated with increased diversity at small spatial scales

Tropical forests are notable for their high species diversity, even on small spatial scales, and right‐skewed species and size abundance distributions. The role of individual species as drivers of the spatial organization of diversity in these forests has been explained by several hypotheses and processes, for example, stochastic dilution, negative density dependence, or gap dynamics. These processes leave a signature in spatial distribution of small trees, particularly in the vicinity of large trees, likely having stronger effects on their neighbors. We are exploring species diversity patterns within the framework of various diversity‐generating hypotheses using individual species–area relationships. We used the data from three tropical forest plots (Wanang—Papua New Guinea, Barro Colorado Island—Panama, and Sinharaja—Sri Lanka) and included also the saplings (DBH ≥ 1 cm). Resulting cross‐size patterns of species richness and evenness reflect the dynamics of saplings affected by the distribution of large trees. When all individuals with DBH ≥1 cm are included, ~50% of all tree species from the 25‐ or 50‐ha plot can be found within 35 m radius of an individual tree. For all trees, 72%–78% of species were identified as species richness accumulators, having more species present in their surroundings than expected by null models. This pattern was driven by small trees as the analysis of DBH >10 cm trees showed much lower proportion of accumulators, 14%–65% of species identified as richness repellers and had low richness of surrounding small trees. Only 11%–26% of species had lower species evenness than was expected by null models. High proportions of species richness accumulators were probably due to gap dynamics and support Janzen–Connell hypothesis driven by competition or top‐down control by pathogens and herbivores. Observed species diversity patterns show the importance of including small tree size classes in analyses of the spatial organization of diversity.     

Biome transitions as centres of diversity: habitat heterogeneity and diversity patterns of West African bat assemblages across spatial scales

It is widely accepted that species diversity is contingent upon the spatial scale used to analyze patterns and processes. Recent studies using coarse sampling grains over large extents have contributed much to our understanding of factors driving global diversity patterns. This advance is largely unmatched on the level of local to landscape scales despite being critical for our understanding of functional relationships across spatial scales. In our study on West African bat assemblages we employed a spatially explicit and nested design covering local to regional scales. Specifically, we analyzed diversity patterns in two contrasting, largely undisturbed landscapes, comprising a rainforest area and a forest‐savanna mosaic in Ivory Coast, West Africa. We employed additive partitioning, rarefaction, and species richness estimation to show that bat diversity increased significantly with habitat heterogeneity on the landscape scale through the effects of beta diversity. Within the extent of our study areas, habitat type rather than geographic distance explained assemblage composition across spatial scales. Null models showed structure of functional groups to be partly filtered on local scales through the effects of vegetation density while on the landscape scale both assemblages represented random draws from regional species pools. We present a mixture model that combines the effects of habitat heterogeneity and complexity on species richness along a biome transect, predicting a unimodal rather than a monotonic relationship with environmental variables related to water. The bat assemblages of our study by far exceed previous figures of species richness in Africa, and refute the notion of low species richness of Afrotropical bat assemblages, which appears to be based largely on sampling biases. Biome transitions should receive increased attention in conservation strategies aiming at the maintenance of ecological and evolutionary processes.     

Intra-annual patterns of saproxylic beetle assemblages inhabiting Mediterranean oak forests

The temporal factor has been widely neglected in studies related to species distribution. However, knowing the species phenology throughout the year could be a key element in ecological studies, more so considering the current climate change framework. The intra-annual temporal patterns of the saproxylic beetle assemblage inhabiting the oak forests in the western Iberian Peninsula were evaluated in terms of species richness and composition. Moreover, we analysed the temporal patterns of species richness and body size, with respect to the abiotic factors of temperature and relative humidity. The marked seasonality of the Mediterranean weather governed, to a great extent, the temporal patterns of the assemblage, as both species richness and body size showed a positive correlation with temperature and a negative correlation with relative humidity. The high temperatures reached in August limited the activity of certain species, causing a marked decrease in species richness. The results revealed that species body size decreased in the colder months of the year and vice versa, which could fit with the inverse of Bergmann’s rule; nevertheless, studies are needed to address the contribution of species physiology and autoecology to the observed temporal patterns. Regarding species composition, the partition of beta diversity in its two components showed a constant process of species turnover throughout the year. We conclude that, due to global warming, the study of temporal patterns of assemblages is becoming all the more important, since it could significantly affect the spatio-temporal distribution of species and the interactions between them.     

Metacommunity‐scale biodiversity regulation and the self‐organised emergence of macroecological patterns

There exist a number of key macroecological patterns whose ubiquity suggests that the spatio‐temporal structure of ecological communities is governed by some universal mechanisms. The nature of these mechanisms, however, remains poorly understood. Here, we probe spatio‐temporal patterns in species richness and community composition using a simple metacommunity assembly model. Despite making no a priori assumptions regarding biotic spatial structure or the distribution of biomass across species, model metacommunities self‐organise to reproduce well‐documented patterns including characteristic species abundance distributions, range size distributions and species area relations. Also in agreement with observations, species richness in our model attains an equilibrium despite continuous species turnover. Crucially, it is in the neighbourhood of the equilibrium that we observe the emergence of these key macroecological patterns. Biodiversity equilibria in models occur due to the onset of ecological structural instability, a population‐dynamical mechanism. This strongly suggests a causal link between local community processes and macroecological phenomena.     

Understanding landscape patterns of temporal variability in avian populations to improve environmental impact assessments

It is recognized that wildlife populations exhibit spatial and temporal variability in patterns of species richness across heterogeneous landscapes. This phenomenon can prove problematic for environmental practitioners when attempting to complete comprehensive environmental assessments (EAs) with limited field surveys. A better understanding of regional spatio-temporal patterns in population dynamics should enhance site-level decision-making. In this study, the variability of seasonal data across the Credit River Watershed, southern Ontario, is assessed for a hierarchy of conservation measures including species richness, and two conservation wildlife response guilds based on primary habitat and area sensitivity. Bird populations were monitored at 24 forest monitoring plots across the watershed by the authors twice a season from 2003 to 2010 following the protocol of Environment Canada's Forest Bird Monitoring Program. The monitoring plots are located within four land management zones identified as 1) urban, 2) transitional, 3) escarpment and 4) rural. Data from the monitoring program are used to compare species richness among plots across the watershed and among land management zones. In addition, the variability of records from each plot over the 8 year period was determined by means of the Coefficient of Variation (CV) statistic. The mean variability of these records at each site within each land management zone was determined in order to assess whether the temporal variability of bird records might affect the integrity of short term assessments. Finally, a one-way ANOVA was applied to learn whether the result of short-term assessments may be further compounded by differences in the response of selected bird guilds to landscape heterogeneity. The results show that there is a significant difference in mean richness of forest birds among the four management zones. The ANOVAs indicate that significant difference is due to the temporal variability of a) breeding forest interior birds rather than edge birds or generalist species and b) breeding area sensitive species rather than area non-sensitive species. Recommendations are made that environmental assessments targeting forest interior bird populations need to plan sampling strategies that recognize this variability, especially for sites within the transitional and urban zones. Planning in the transition or urbanizing landscape should incorporate landscape ecology principles to sustain current levels of richness in forest species.     

The impact of population processes on patterns of species richness: Lessons from elevational gradients

In the last few years, considerable headway has been made towards understanding patterns of species richness along latitudinal and elevational gradients, mostly by focussing on the influences of surface area, climatic factors, evolutionary history, and stochastic processes. However, the potential impact of population-level processes in determining or modifying patterns of species richness has largely been neglected, partly due to the difficulty of gathering such data for numerous species along geographical or ecological gradients. Based on two empirical examples, I here show that dispersal and the resulting source-sink effects modify patterns of plant species richness along elevation gradients, and that the inclusion or exclusion of such sink populations alters the perception of the diversity patterns and hence our interpretation of them. I argue that population processes should be taken into account when studying patterns of species richness, especially at scales at which dispersal is common in the taxon under consideration.     

Are local patterns of anthropoid primate diversity related to patterns of diversity at a larger scale?

Aims (1) To determine the relationship between local and regional anthropoid primate species richness. (2) To establish the spatial and temporal scale at which the ultimate processes influencing patterns of primate species coexistence operate. Location Continental landmasses of Africa, South America and Asia (India to China, and all islands as far south as New Guinea). Methods The local–regional species richness relationship for anthropoid primates is estimated by regressing local richness against regional richness (independent variable). Local richness is estimated in small, replicate local assemblages sampled in regions that vary in total species richness. A strong linear relationship is taken as evidence that local assemblages are unsaturated and local richness results from proportional sampling of the regional pool. An asymptotic curvilinear relationship is interpreted to reflect saturated communities, where strong biotic interactions limit local richness and local processes structure the species assemblage. As a further test of the assumption of local assemblage saturation, we looked for density compensation in high‐density local primate assemblages. Results The local–regional species richness relationship was linear for Africa and South America, and the slope of the relationship did not differ between the two continents. For Asia, curvilinearity best described the relationship between local and regional richness. Asian primate assemblages appear to be saturated and this is confirmed by density compensation among Asian primates. However, density compensation was also observed among African primates. The apparent assemblage saturation in Asia is not a species–area phenomenon related to the small size of the isolated islands and their forest blocks, since similar low local species richness occurs in large forests on mainland and/or peninsular Asia. Main conclusions In Africa and South America local primate assemblage composition appears to reflect the influence of biogeographic processes operating on regional spatial scales and historical time scales. In Asia the composition of primate assemblages are by‐and‐large subject to ecological constraint operating over a relatively small spatial and temporal scale. The possible local influence of the El Niño Southern Oscillations on the evolution and selection of life‐history characteristics among Asian primates, and in determining local patterns of primate species coexistence, warrants closer inspection.     

Contribution of rarity and commonness to patterns of species richness

There is little understanding in ecology as to how biodiversity patterns emerge from the distribution patterns of individual species. Here we consider the question of the contributions of rare (restricted range) and common (widespread) species to richness patterns. Considering a species richness pattern, is most of the spatial structure, in terms of where the peaks and troughs of diversity lie, caused by the common species or the rare species (or neither)? Using southern African and British bird richness patterns, we show here that commoner species are most responsible for richness patterns. While rare and common species show markedly different species richness patterns, most spatial patterning in richness is caused by relatively few, more common, species. The level of redundancy we found suggests that a broad understanding of what determines the majority of spatial variation in biodiversity may be had by considering only a minority of species.     

Contrasting spatial and temporal global change impacts on butterfly species richness during the 20th century

Regional patterns of species richness are often explained by models using temperature or measures habitat suitability. Generally, species richness is positively associated with temperature, and negatively associated with habitat degradation. While these models have been well tested across spatial scales, they have rarely been tested on a temporal scale – in part due to the difficulty in ascertaining accurate historical data at an appropriate resolution. In this study, we compared the results of temporal and spatial models, each incorporating two predictors of species richness: temperature, and human population density (as a surrogate of human-related habitat impacts). We found that the change in species richness from the early to late part of the 20th century was positively correlated with temperature change, and negatively correlated with human population density change. When we compared these results to two spatial models using contemporary and historic data, the spatial effects of temperature on butterfly richness were similar to its temporal effects, while the effect of human population density through time is the opposite of its spatial effect. More generally, the assumption that spatial patterns are equivalent to temporal ones when applying macroecological data to global change is clearly unreliable.     

The small‐island effect: fact or artefact?

Positive relationships between species richness and sampling area are perhaps the most pervasive patterns in nature. However, the shape of species–area relationships is often highly variable, for reasons that are poorly understood. One such source of variability is the "small-island effect", which refers to a decrease in the capacity of sampling area to predict species richness on small islands. Small-island effects have been attributed to a variety of processes, including spatial subsidies, habitat characteristics and ocean-born disturbances. Here, we show that small-island effects can be generated by logarithmic data transformations, which are commonly applied to both axes of species–area relationships. To overcome this problem, we derive several null models to test for non-random variability in the capacity of island area to predict species richness and apply them to data sets on island plant communities in Canada and New Zealand. Both archipelagos showed evidence for small-island effects using traditional breakpoint regression techniques on log-log axes. However, null model analyses revealed different results. The capacity of sampling area to predict species richness in the Canadian archipelago was actually lowest at intermediate island size classes. In the New Zealand archipelago, island area was similarly capable of predicting species richness across the full range of island sizes, indicating the small-island effect detected by breakpoint regression is an artifact of logarithm data transformation. Overall results show that commonly used regression techniques can generate spurious small-island effects and that alternative analytic procedures are needed to detect non-random patterns in species richness on small islands.     

The relationships between local and regional species richness and spatial turnover

Aim To determine the empirical relationships between species richness and spatial turnover in species composition across spatial scales. These have remained little explored despite the fact that such relationships are fundamental to understanding spatial diversity patterns. Location South‐east Scotland. Methods Defining local species richness simply as the total number of species at a finer resolution than regional species richness and spatial turnover as turnover in species identity between any two or more areas, we determined the empirical relationships between all three, and the influence of spatial scale upon them, using data on breeding bird distributions. We estimated spatial turnover using a measure independent of species richness gradients, a fundamental feature which has been neglected in theoretical studies. Results Local species richness and spatial turnover exhibited a negative relationship, which became stronger as larger neighbourhood sizes were considered in estimating the latter. Spatial turnover and regional species richness did not show any significant relationship, suggesting that spatial species replacement occurs independently of the size of the regional species pool. Local and regional species richness only showed the expected positive relationship when the size of the local scale was relatively large in relation to the regional scale. Conclusions Explanations for the relationships between spatial turnover and local and regional species richness can be found in the spatial patterns of species commonality, gain and loss between areas.     

Species-energy relationships in deep-sea molluscs

Consensus is growing among ecologists that energy and the factors influencing its utilization can play overarching roles in regulating large-scale patterns of biodiversity. The deep sea--the world's largest ecosystem--has simplified energetic inputs and thus provides an excellent opportunity to study how these processes structure spatial diversity patterns. Two factors influencing energy availability and use are chemical (productive) and thermal energy, here represented as seafloor particulate organic carbon (POC) flux and temperature. We related regional patterns of benthic molluscan diversity in the North Atlantic to these factors, to conduct an explicit test of species-energy relationships in the modern day fauna of the deep ocean. Spatial regression analyses in a model-averaging framework indicated that POC flux had a substantially higher relative importance than temperature for both gastropods and protobranch bivalves, although high correlations between variables prevented definitive interpretation. This contrasts with recent research on temporal variation in fossil diversity from deep-sea cores, where temperature is generally a more significant predictor. These differences may reflect the scales of time and space at which productivity and temperature operate, or differences in body size; but both lines of evidence implicate processes influencing energy utilization as major determinants of deep-sea species diversity.     

Spatial predictions at the community level: from current approaches to future frameworks

A fundamental goal of ecological research is to understand and model how processes generate patterns so that if conditions change, changes in the patterns can be predicted. Different approaches have been proposed for modelling species assemblage, but their use to predict spatial patterns of species richness and other community attributes over a range of spatial and temporal scales remains challenging. Different methods emphasize different processes of structuring communities and different goals. In this review, we focus on models that were developed for generating spatially explicit predictions of communities, with a particular focus on species richness, composition, relative abundance and related attributes. We first briefly describe the concepts and theories that span the different drivers of species assembly. A combination of abiotic processes and biotic mechanisms are thought to influence the community assembly process. In this review, we describe four categories of drivers: (i) historical and evolutionary, (ii) environmental, (iii) biotic, and (iv) stochastic. We discuss the different modelling approaches proposed or applied at the community level and examine them from different standpoints, i.e. the theoretical bases, the drivers included, the source data, and the expected outputs, with special emphasis on conservation needs under climate change. We also highlight the most promising novelties, possible shortcomings, and potential extensions of existing methods. Finally, we present new approaches to model and predict species assemblages by reviewing promising ‘integrative frameworks’ and views that seek to incorporate all drivers of community assembly into a unique modelling workflow. We discuss the strengths and weaknesses of these new solutions and how they may hasten progress in community‐level modelling.     

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.     

黄河流域被子植物和陆栖脊椎动物丰富度格局及其影响因子

生物多样性的大尺度空间分布格局及其形成机制一直是生态学和生物地理学的核心内容。黄河流域是我国重要的生态屏障, 明确该区域动植物多样性分布格局及其影响因素, 对我国黄河流域生态保护和高质量发展具有重要意义。本研究通过收集黄河流域被子植物和陆栖脊椎动物分布数据, 结合气候、环境异质性和人类活动等信息, 探讨了黄河流域被子植物和陆栖脊椎动物物种丰富度格局及其主要影响因素。结果表明, 黄河流域被子植物和陆栖脊椎动物物种丰富度在区域尺度具有相似的分布格局: 南部山地动植物物种丰富度最高, 而东部高寒区和北部干旱区物种丰富度最低。回归树模型表明, 冠层高度范围和净初级生产力范围分别是黄河流域被子植物和陆栖脊椎动物物种丰富度最重要的预测因子; 当移除空间自相关影响后, 环境异质性和气候因子依然对区域尺度的动植物物种丰富度具有较高且相似的解释度。表明环境异质性和气候共同决定了黄河流域被子植物和陆栖脊椎动物物种丰富度格局, 而人类使用土地面积并不是影响黄河流域动植物物种丰富度格局的主要因子。因此, 在未来的研究中若针对不同区域筛选出更精准的环境驱动因子或选用更多不同类别的环境异质性因子进行分析, 将有助于更深入理解物种多样性格局的成因。