Scaling biodiversity responses to hydrological regimes

Of all ecosystems, freshwaters support the most dynamic and highly concentrated biodiversity on Earth. These attributes of freshwater biodiversity along with increasing demand for water mean that these systems serve as significant models to understand drivers of global biodiversity change. Freshwater biodiversity changes are often attributed to hydrological alteration by water‐resource development and climate change owing to the role of the hydrological regime of rivers, wetlands and floodplains affecting patterns of biodiversity. However, a major gap remains in conceptualising how the hydrological regime determines patterns in biodiversity's multiple spatial components and facets (taxonomic, functional and phylogenetic). We synthesised primary evidence of freshwater biodiversity responses to natural hydrological regimes to determine how distinct ecohydrological mechanisms affect freshwater biodiversity at local, landscape and regional spatial scales. Hydrological connectivity influences local and landscape biodiversity, yet responses vary depending on spatial scale. Biodiversity at local scales is generally positively associated with increasing connectivity whereas landscape‐scale biodiversity is greater with increasing fragmentation among locations. The effects of hydrological disturbance on freshwater biodiversity are variable at separate spatial scales and depend on disturbance frequency and history and organism characteristics. The role of hydrology in determining habitat for freshwater biodiversity also depends on spatial scaling. At local scales, persistence, stability and size of habitat each contribute to patterns of freshwater biodiversity yet the responses are variable across the organism groups that constitute overall freshwater biodiversity. We present a conceptual model to unite the effects of different ecohydrological mechanisms on freshwater biodiversity across spatial scales, and develop four principles for applying a multi‐scaled understanding of freshwater biodiversity responses to hydrological regimes. The protection and restoration of freshwater biodiversity is both a fundamental justification and a central goal of environmental water allocation worldwide. Clearer integration of concepts of spatial scaling in the context of understanding impacts of hydrological regimes on biodiversity will increase uptake of evidence into environmental flow implementation, identify suitable biodiversity targets responsive to hydrological change or restoration, and identify and manage risks of environmental flows contributing to biodiversity decline.     

Do generalized scaling laws exist for species abundance distribution in mountains?

Knowing the global pattern of species diversity is a central goal of the science of ecology, and scaling laws can be useful for analysis of cross-scale biodiversity patterns. An elevational gradient in a warm temperate zone of the Donglingshan mountains (China) is used to test the scaling laws of species abundance distribution using multifractal analysis. We show that the power law scaling relationship holds for not just the classical SAR (species–area relationship for richness), but also for Shannon and Simpson diversity. In fact, we find power-laws in the generalized species abundance distribution at all stratal levels of the forest (trees, shrubs and herbs). The fact that these laws exist across a heterogeneous landscape representing a strong bioclimatic gradient suggests that biodiversity scaling laws may be more robust than previously thought.     

The macroecology of infectious diseases: a new perspective on global‐scale drivers of pathogen distributions and impacts

Identifying drivers of infectious disease patterns and impacts at the broadest scales of organisation is one of the most crucial challenges for modern science, yet answers to many fundamental questions remain elusive. These include what factors commonly facilitate transmission of pathogens to novel host species, what drives variation in immune investment among host species, and more generally what drives global patterns of parasite diversity and distribution? Here we consider how the perspectives and tools of macroecology, a field that investigates patterns and processes at broad spatial, temporal and taxonomic scales, are expanding scientific understanding of global infectious disease ecology. In particular, emerging approaches are providing new insights about scaling properties across all living taxa, and new strategies for mapping pathogen biodiversity and infection risk. Ultimately, macroecology is establishing a framework to more accurately predict global patterns of infectious disease distribution and emergence.     

Spatial scaling of extinction rates: Theory and data reveal nonlinearity and a major upscaling and downscaling challenge

Aim

Biodiversity loss is a key component of biodiversity change and can impact ecosystem services. However, estimation of the loss has focused mostly on per‐species extinction rates measured over a limited number of spatial scales, with little theory linking small‐scale extirpations to global extinctions. Here, we provide such a link by introducing the relationship between area and the number of extinctions (number of extinctions–area relationship; NxAR) and between area and the proportion of extinct species (proportion of extinctions–area relationship; PxAR). Unlike static patterns, such as the species–area relationship, NxAR and PxAR represent spatial scaling of a dynamic process. We show theoretical and empirical forms of these relationships and we discuss their role in perception and estimation of the current extinction crisis.

Location

U.S.A., Europe, Czech Republic and Barro Colorado Island (Panama).

Time period

1500–2009.

Major taxa studied

Vascular plants, birds, butterflies and trees.

Methods

We derived the expected forms of NxAR and PxAR from several theoretical frameworks, including the theory of island biogeography, neutral models and species–area relationships. We constructed NxAR and PxAR from five empirical datasets collected over a range of spatial and temporal scales.

Results

Although increasing PxAR is theoretically possible, empirical data generally support a decreasing PxAR; the proportion of extinct species decreases with area. In contrast, both theory and data revealed complex relationships between numbers of extinctions and area (NxAR), including nonlinear, unimodal and U‐shaped relationships, depending on region, taxon and temporal scale.

Main conclusions

The wealth of forms of NxAR and PxAR explains why biodiversity change appears scale dependent. Furthermore, the complex scale dependence of NxAR and PxAR means that global extinctions indicate little about local extirpations, and vice versa. Hence, effort should be made to understand and report extinction rates as a scale‐dependent problem. In this effort, estimation of scaling relationships such as NxAR and PxAR should be central.     

The role of evolutionary processes in producing biodiversity patterns, and the interrelationships between taxonomic, functional and phylogenetic biodiversity

Patterns of biodiversity are ultimately the product of speciation and extinction. Speciation serves as the biodiversity pump while extinction serves as the agent that culls global to local levels of biodiversity. Linking these central processes to global and local patterns of biodiversity is a key challenge in both ecology and evolution. This challenge necessarily requires a simultaneous consideration of the species, phylogenetic, and functional diversity across space and the tree of life. In this review, I outline a research framework for biodiversity science that considers the evolutionary and ecological processes that generate and cull levels of biodiversity and that influence the inter-relationships between species, phylogenetic, and functional diversity. I argue that a biodiversity synthesis must begin with a consideration of the inherently ecological process of speciation and end with how global biodiversity is filtered into local-scale plant communities. The review ends with a brief outlook on future challenges for those studying biodiversity, including outstanding hypotheses that need testing and key data limitations.     

A statistical theory for sampling species abundances

The pattern of species abundances is central to ecology. But direct measurements of species abundances at ecologically relevant scales are typically unfeasible. This limitation has motivated a long-standing interest in the relationship between the abundance distribution in a large, regional community and the distribution observed in a small sample from the community. Here, we develop a statistical sampling theory to describe how observed patterns of species abundances are influenced by the spatial distributions of populations. For a wide range of regional-scale abundance distributions we derive exact expressions for the sampled abundance distributions, as a function of sample size and the degree of conspecific spatial aggregation. We show that if populations are randomly distributed in space then the sampled and regional-scale species-abundance distribution typically have the same functional form: sampling can be expressed by a simple scaling relationship. In the case of aggregated spatial distributions, however, the shape of a sampled species-abundance distribution diverges from the regional-scale distribution. Conspecific aggregation results in sampled distributions that are skewed towards both rare and common species. We discuss our findings in light of recent results from neutral community theory, and in the context of estimating biodiversity.     

Network isolation and local diversity in neutral metacommunities

Biologists seek an understanding of the biological and environmental factors determining local community diversity. Recent advances in metacommunity ecology, and neutral theory in particular, highlight the importance of dispersal processes interacting with the spatial structure of a landscape for generating spatial patterns and maintaining biodiversity. The relative spatial isolation of a community is traditionally thought to have a large influence on local diversity. However, isolation remains an elusive concept to quantify, particularly in metacommunities with complex spatial structure. We represent the metacommunity as a network of local communities, and use network centrality measures to quantify the isolation of a local community. Using spatially explicit neutral theory, we examine how node position predicts variation in alpha diversity across a metacommunity. We find that diversity increases with node centrality in the network, but only when centrality is measured on a given scale in the network that widens with increasing dispersal rates and narrows with increasing evolutionary rates. More generally, complex biodiversity patterns form only when the underlying geography has structure on this critical scale. This provides a framework for understanding the influence of spatial geographic structure on global biodiversity patterns.     

Field theory for biogeography: a spatially explicit model for predicting patterns of biodiversity

Predicting the variation of biodiversity across the surface of the Earth is a fundamental issue in ecology, and in this article we focus on one of the most widely studied spatial biodiversity patterns: the species–area relationship (SAR). The SAR is a central tool in conservation, being used to predict species loss following global climate change, and is striking in its universality throughout different geographical regions and across the tree of life. In this article we draw upon the methods of quantum field theory and the foundation of neutral community ecology to derive the first spatially explicit neutral prediction for the SAR. We find that the SAR has three phases, with a power law increase at intermediate scales, consistent with decades of documented empirical patterns. Our model also provides a building block for incorporating non-neutral biological variation, with the potential to bridge the gap between neutral and niche-based approaches to community assembly.
Ecology Letters (2010) 13: 87–95     

土壤动物群落空间格局和构建机制研究进展

群落空间格局和构建机制一直是生态学研究的核心内容。在生物多样性严重丧失的背景下, 揭示群落空间格局及其构建机制, 有助于深刻理解生物多样性丧失的原因, 更有助于应对生物多样性保护等重大生态环境问题。然而, 陆地生态系统的研究多集中于地上生物群落, 对地下生态系统, 尤其是土壤动物空间格局和构建机制的研究尚不充分。事实上, 土壤动物多样性是全球生物多样性的关键组成之一, 是地下生态系统结构和功能维持的重要部分。对土壤动物空间格局和构建机制的研究, 能明确不同空间尺度条件下土壤动物多样性的维持机制。土壤动物群落常在多种空间尺度形成复杂的空间分布格局, 因此, 本文首先介绍了不同空间尺度主要土壤动物群落的空间自相关性特征, 阐述了土壤动物群落斑块和孔隙镶嵌分布的复杂空间格局。继而阐明这种空间格局主要受生物间作用、环境过滤和随机扩散的调控, 并说明这三个过程对土壤动物群落的调控能力和作用方式。作者提出, 这三个过程仍是今后土壤动物群落空间格局和构建机制研究的重点内容, 需要进一步加强以土壤动物为研究对象的群落构建理论的验证和发展。我国土壤动物群落空间格局和构建机制起步较晚, 希望本文能够促进我国土壤动物生态学相关领域的研究。     

Scaling‐up biodiversity‐ecosystem functioning research

A rich body of knowledge links biodiversity to ecosystem functioning (BEF), but it is primarily focused on small scales. We review the current theory and identify six expectations for scale dependence in the BEF relationship: (1) a nonlinear change in the slope of the BEF relationship with spatial scale; (2) a scale‐dependent relationship between ecosystem stability and spatial extent; (3) coexistence within and among sites will result in a positive BEF relationship at larger scales; (4) temporal autocorrelation in environmental variability affects species turnover and thus the change in BEF slope with scale; (5) connectivity in metacommunities generates nonlinear BEF and stability relationships by affecting population  synchrony at local and regional scales; (6) spatial scaling in food web structure and diversity will generate scale dependence in ecosystem functioning. We suggest directions for synthesis that combine approaches in metaecosystem and metacommunity ecology and integrate cross‐scale feedbacks. Tests of this theory may combine remote sensing with a generation of networked experiments that assess effects at multiple scales. We also show how anthropogenic land cover change may alter the scaling of the BEF relationship. New research on the role of scale in BEF will guide policy linking the goals of managing biodiversity and ecosystems.     

青藏高原高寒草地地下生物多样性: 进展、问题与展望

栖息于土壤中的微生物和微型动物种类繁多、数量巨大, 在对地上生物多样性的调控和在生态系统功能与服务的维系中, 具有举足轻重的作用。虽然对土壤微生物以及土壤动物已经开展了广泛的调查, 但是整体上对于地下生物多样性的分布格局、驱动机制及其对全球变化的响应与适应过程, 仍缺乏深刻的认识。青藏高原是全球变化的敏感区域, 其中高寒草地是高原最主要的植被类型, 占高原面积的60%左右, 在高寒生态系统生物多样性维持中具有重要意义。近年来, 已有大量研究关注于高寒草地地下生物多样性, 但是缺乏系统的总结与论述。基于此, 本文从细菌、真菌、古菌、线虫、节肢动物五大土壤生物类群出发, 阐述了青藏高原高寒草地的地下物种丰富度、分布格局及其影响因素, 重点探讨了它们对气候变化和人类活动的响应, 并就未来高寒草地地下生物多样性亟需关注的关键问题进行了展望, 包括: (1)地下各个生物类群的分布格局、各类群之间的联系及驱动机制; (2)地上与地下生物多样性耦联的机制; (3)地下生物多样性对生态系统功能和健康的影响; (4)地下生物多样性的调控实验研究。     

Scale dependency of diversity components estimated from primary biodiversity data and distribution maps

Different sources of information about biodiversity may lead to unrealistic or biased estimation of its components, with different patterns according to the scale of analysis. In this study, we analyse patterns of species richness at the local (average alpha) and regional (gamma) scales, and the relationship between them (Whittaker's beta), in central Mexico, using as a source of data for the species' distributions: (1) museum specimen occurrence data for birds, and (2) distribution maps based on ecological niche models developed and refined by experts. We performed analyses at five spatial resolutions (1/32°−1/2°). Scale changes (grain and extent) affected significantly the estimates of average alpha, gamma, and beta. Use of raw occurrence data vs. distribution maps yielded contrasting results, with raw data underestimating alpha and overestimating beta, as functions of area. As regards species–area relationships, our results suggest a natural decomposition of factors into an area-invariant component (related to alpha), and an area dependent factor (related to beta). Most of our results are maintained in a null model that randomizes occurrences without changing observed range-size distributions. From this result we argue that average alpha and Whittaker's beta capture little information about the spatial covariation of species distribution patterns.     

The ‘rotiferologist’ effect and other global correlates of species richness in monogonont rotifers

Global biodiversity patterns are often driven by different environmental variables at different scales. However, it is still controversial whether there are general trends, whether similar processes are responsible for similar patterns, and/or whether confounding effects such as sampling bias can produce misleading results. Our aim is twofold: 1) assessing the global correlates of diversity in a group of microscopic animals little analysed so far, and 2) inferring the influence of sampling intensity on biodiversity analyses. As a case study, we choose rotifers, because of their high potential for dispersal across the globe. We assembled and analysed a new worldwide dataset of records of monogonont rotifers, a group of microscopic aquatic animals, from 1960 to 1992. Using spatially explicit models, we assessed whether the diversity patterns conformed to those commonly obtained for larger organisms, and whether they still held true after controlling for sampling intensity, variations in area, and spatial structure in the data. Our results are in part analogous to those commonly obtained for macroorganisms (habitat heterogeneity and precipitation emerge as the main global correlates), but show some divergence (potential absence of a latitudinal gradient and of a large‐scale correlation with human population). Moreover, the effect of sampling effort is remarkable, accounting for >50% of the variability; this strong effect may mask other patterns such as latitudinal gradients. Our study points out that sampling bias should be carefully considered when drawing conclusions from large‐scale analyses, and calls for further faunistic work on microorganisms in all regions of the world to better understand the generality of the processes driving global patterns in biodiversity.     

Selecting for extinction: nonrandom disease-associated extinction homogenizes amphibian biotas

Studying the patterns in which local extinctions occur is critical to understanding how extinctions affect biodiversity at local, regional and global spatial scales. To understand the importance of patterns of extinction at a regional spatial scale, we use data from extirpations associated with a widespread pathogenic agent of amphibian decline, Batrachochytrium dendrobatidis ( Bd ) as a model system. We apply novel null model analyses to these data to determine whether recent extirpations associated with Bd have resulted in selective extinction and homogenization of diverse tropical American amphibian biotas. We find that Bd -associated extinctions in this region were nonrandom and disproportionately, but not exclusively, affected low-occupancy and endemic species, resulting in homogenization of the remnant amphibian fauna. The pattern of extirpations also resulted in phylogenetic homogenization at the family level and ecological homogenization of reproductive mode and habitat association. Additionally, many more species were extirpated from the region than would be expected if extirpations occurred randomly. Our results indicate that amphibian declines in this region are an extinction filter, reducing regional amphibian biodiversity to highly similar relict assemblages and ultimately causing amplified biodiversity loss at regional and global scales.     

微生物多样性及其分布的研究进展:模式与过程

生物多样性产生和维持机制长期以来是生态学研究的核心问题。人们通过实验观察和理论推导揭示了很多群落生物多样性的共有模式及其背后的生态学过程,而相关的工作主要集中在动植物等大型生物中。微生物是地球上数量最多、分布最广,以及在生态系统过程中发挥极其重要作用的生物类群,但人们对其群落多样性的认识还非常有限。将介绍目前人们对能够独立自由生活的微生物多样性分布模式的探索,总结对其背后的生物学和生态学过程的研究现状,并探讨当前面临的挑战。     

Seasonal and spatial hydrological variability drives aquatic biodiversity in a flood‐pulsed,sub‐tropical wetland

1. Flood‐pulsed wetlands make vital contributions to local and global biodiversity. However, the patterns and controls of spatial and temporal variation in aquatic biodiversity in flood‐pulsed wetlands are not well understood. We analysed the relationship between variation in hydrological regime and the patterns of aquatic biodiversity in a large pristine flood‐pulsed wetland, the Okavango Delta, Botswana. 2. Surveys of water chemistry, diatoms and macroinvertebrates were conducted over the seasonal phases of the flood pulse. Hydrological variables of flood frequency and hydroperiod class were collated from 16 years of satellite images. Multivariate regression trees and generalised least squares regression were used to determine the chief controls of community composition and taxon richness. 3. Hydroperiod class, phase of the flood and conductivity explained 32% and 43% of the variation in diatom and invertebrate taxon richness, respectively. There was a negative relationship between hydroperiod class and invertebrate taxon richness on the rising, peak and receding flood, whereas at low flood there was no significant relationship. Multivariate regression tree analysis revealed hydroperiod class, phase of the flood and conductivity as the dominant forces shaping invertebrate and diatom community composition. 4. Seasonal and spatial variation in hydrological conditions are the principal drivers of variation in aquatic biodiversity in flood‐pulsed wetlands. In pristine flood‐pulsed wetlands, increased productivity caused by the arrival of the flood waters appears to override disturbance and connectivity in shaping taxon richness and community composition. Thus, these data suggest that the maintenance of a rich mosaic of habitats covering a broad range of hydroperiod is the key to preserving aquatic biodiversity and natural ecosystem function in flood‐pulsed wetlands.     

Human impacts on the species-area relationship in reef fish assemblages

The relationship between species richness and area is one of the oldest, most recognized patterns in ecology. Here we provide empirical evidence for strong impacts of fisheries exploitation on the slope of the species–area relationship (SAR). Using comparative field surveys of fish on protected and exploited reefs in three oceans and the Mediterranean Sea, we show that exploitation consistently depresses the slope of the SAR for both power-law and exponential models. The magnitude of change appears to be proportional to fishing intensity. Results are independent of taxonomic resolution and robust across coral and rocky reefs, sampling protocols and statistical methods. Changes in species richness, relative abundance and patch occupancy all appear to contribute to this pattern. We conclude that exploitation pressure impacts the fundamental scaling of biodiversity as well as the species richness and spatial distribution patterns of reef fish. We propose that species–area curves can be sensitive indicators of community-level changes in biodiversity, and may be useful in quantifying the human imprint on reef biodiversity, and potentially elsewhere.     

High diversity,abundance and distinct fish assemblages on submerged coral reef pinnacles compared to shallow emergent reefs

Coral Reefs - Coral reefs exhibit consistent patterns in biodiversity across multiple spatial scales, from local to global clines in species richness, abundance and community structure. Knowledge...     

Macroecological patterns in soil communities

Aim To review published evidence regarding the factors that influence the geographic variation in diversity of soil organisms at different spatial scales. Location Global. Methods A search of the relevant literature was conducted using the Web of Science and the author's personal scientific database as the major sources. Special attention was paid to include seminal studies, highly cited papers and/or studies highlighting novel results. Results Despite their significant contribution to global biodiversity, our taxonomic knowledge of soil biota is still poor compared with that of most above‐ground organisms. This is particularly evident for small‐bodied taxa. Global patterns of soil biodiversity distribution have been poorly documented and are thought to differ significantly from what is reported above‐ground. Based on existing data, it appears that microorganisms do not respond to large‐scale environmental gradients in the same way as metazoans. Whereas soil microflora seem to be mainly represented by cosmopolitan species, soil animals respond to altitudinal, latitudinal or area gradients in the same way as described for above‐ground organisms. At local scales, there is less evidence that local factors regulate above‐ and below‐ground communities in the same way. Except for a few taxa, the humpbacked response to stress and disturbance gradients doesn't seem to apply underground. Soil communities thus appear weakly structured by competition, although competitive constraints may account for assembly rules within specific taxa. The main factor constraining local soil biodiversity is the compact and heterogeneous nature of soils, which provides unrivalled potential for niche partitioning, thus allowing high levels of local biodiversity. This heterogeneity is increased by the impact of ecosystem engineers that generate resource patchiness at a range of spatio‐temporal scales.     

Inferring macro‐ecological patterns from local presence/absence data

Biodiversity provides support for life, vital provisions, regulating services and has positive cultural impacts. It is therefore important to have accurate methods to measure biodiversity, in order to safeguard it when we discover it to be threatened. For practical reasons, biodiversity is usually measured at fine scales whereas diversity issues (e.g. conservation) interest regional or global scales. Moreover, biodiversity may change across spatial scales. It is therefore a key challenge to be able to translate local information on biodiversity into global patterns. Many databases give no information about the abundances of a species within an area, but only its occurrence in each of the surveyed plots. In this paper, we introduce an analytical framework (implemented in a ready‐to‐use R code) to infer species richness and abundances at large spatial scales in biodiversity‐rich ecosystems when species presence/absence information is available on various scattered samples (i.e. upscaling). This framework is based on the scale‐invariance property of the negative binomial. Our approach allows to infer and link within a unique framework important and well‐known biodiversity patterns of ecological theory, such as the species accumulation curve (SAC) and the relative species abundance (RSA) as well as a new emergent pattern, which is the relative species occupancy (RSO). Our estimates are robust and accurate, as confirmed by tests performed on both in silico‐generated and real forests. We demonstrate the accuracy of our predictions using data from two well‐studied forest stands. Moreover, we compared our results with other popular methods proposed in the literature to infer species richness from presence to absence data and we showed that our framework gives better estimates. It has thus important applications to biodiversity research and conservation practice.