The spatial scaling of beta diversity

Beta diversity is an important concept used to describe turnover in species composition across a wide range of spatial and temporal scales, and it underpins much of conservation theory and practice. Although substantial progress has been made in the mathematical and terminological treatment of different measures of beta diversity, there has been little conceptual synthesis of potential scale dependence of beta diversity with increasing spatial grain and geographic extent of sampling. Here, we evaluate different conceptual approaches to the spatial scaling of beta diversity, interpreted from ‘fixed’ and ‘varying’ perspectives of spatial grain and extent. We argue that a ‘sliding window’ perspective, in which spatial grain and extent covary, is an informative way to conceptualize community differentiation across scales. This concept more realistically reflects the varying empirical approaches that researchers adopt in field sampling and the varying scales of landscape perception by different organisms. Scale dependence in beta diversity has broad implications for emerging fields in ecology and biogeography, such as the integration of fine‐resolution ecogenomic data with large‐scale macroecological studies, as well as for guiding appropriate management responses to threats to biodiversity operating at different spatial scales.     

Fifteen important questions in the spatial ecology of diatoms

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Metacommunity Structure of Stream Insects across Three Hierarchical Spatial scales

A major challenge in community ecology is to understand the underlying factors driving metacommunity (i.e., a set of local communities connected through species dispersal) dynamics. However, little is known about the effects of varying spatial scale on the relative importance of environmental and spatial (i.e., dispersal related) factors in shaping metacommunities and on the relevance of different dispersal pathways. Using a hierarchy of insect metacommunities at three spatial scales (a small, within‐stream scale, intermediate, among‐stream scale, and large, among‐sub‐basin scale), we assessed whether the relative importance of environmental and spatial factors shaping metacommunity structure varies predictably across spatial scales, and tested how the importance of different dispersal routes vary across spatial scales. We also studied if different dispersal ability groups differ in the balance between environmental and spatial control. Variation partitioning showed that environmental factors relative to spatial factors were more important for community composition at the within‐stream scale. In contrast, spatial factors (i.e., eigenvectors from Moran's eigenvector maps) relative to environmental factors were more important at the among‐sub‐basin scale. These results indicate that environmental filtering is likely to be more important at the smallest scale with highest connectivity, while dispersal limitation seems to be more important at the largest scale with lowest connectivity. Community variation at the among‐stream and among‐sub‐basin scales were strongly explained by geographical and topographical distances, indicating that overland pathways might be the main dispersal route at the larger scales among more isolated sites. The relative effect of environmental and spatial factors on insect communities varied between low and high dispersal ability groups; this variation was inconsistent among three hierarchical scales. In sum, our study indicates that spatial scale, connectivity, and dispersal ability jointly shape stream metacommunities.     

Biodiversity and Conservation of Diptera in Heterogeneous Land Mosaics: A Fly's Eye View

Classical landscape ecology views spatial heterogeneity of habitats at relatively large 'human scales', and it is at such scales that most decisions of land management and nature conservation are made. The present paper makes use of a wider range of spatial scales to examine land mosaics from the 'fly point of view'. Taking examples from the Diptera faunas of mountainous land mosaics, it is demonstrated that: (i) large scale, 'patch content' landscape management has a direct bearing on Diptera community structure, (ii) borders between large scale patches are not necessarily perceived by flies (or other insects) in the same way as we perceive them, (iii) border complexity between patches at any scale may be as important as patch content as an axis of habitat definition. In this sense, 'border' is not to be confused with 'edge effects'. It is concluded that attention to both patch content and patch border complexity of land mosaics, viewed at the relevant spatial scales, is necessary for future successful conservation of Diptera biodiversity and for the efficient use of these insects in environmental assessment studies.     

Assembly rules of ground-foraging ant assemblages are contingent on disturbance, habitat and spatial scale

Aim A major endeavour of community ecology is documenting non‐random patterns in the composition and body size of coexisting species, and inferring the processes, or assembly rules, that may have given rise to the observed patterns. Such assembly rules include species sorting resulting from interspecific competition, aggregation at patchily distributed resources, and co‐evolutionary dynamics. However, for any given taxon, relatively little is known about how these patterns and processes change through time and vary with habitat type, disturbance history, and spatial scale. Here, we tested for non‐random patterns of species co‐occurrence and body size in assemblages of ground‐foraging ants and asked whether those patterns varied with habitat type, disturbance history, and spatial scale. Location Burned and unburned forests and fens in the Siskiyou Mountains of southern Oregon and northern California, USA. Methods We describe ground‐foraging ant assemblages sampled over two years in two discrete habitat types, namely Darlingtonia fens and upland forests. Half of these sites had been subject to a large‐scale, discrete disturbance – a major fire – in the year prior to our first sample. We used null model analyses to compare observed species co‐occurrence patterns and body‐size distributions in these assemblages with randomly generated assemblages unstructured by competition both within (i.e. at a local spatial scale) and among (i.e. at a regional scale) sites. Results At local spatial scales, species co‐occurrence patterns and body‐size ratios did not differ from randomness. At regional scales, co‐occurrence patterns were random or aggregated, and there was evidence for constant body‐size ratios of forest ants. Although these patterns varied between habitats and years, they did not differ between burned and unburned sites. Main conclusions Our results suggest that the operation of assembly rules depends on spatial scale and habitat type, but that it was not affected by disturbance history from fire.     

Scale dependence of temperature as an abiotic driver of species' distributions

Aim Scale dependence of patterns and processes remains one of the major unresolved problems in ecology. The responses of ecosystems to environmental stressors are reported to be strongly scale dependent, but projections of the effects of climate change on species' distributions are still restricted to particular scales and knowledge about scale dependence is lacking. Here we propose that the scale dependence of those species' niche dimensions related to climate change is strongly related to the strength of climatic cross‐scale links. More specifically, we hypothesize that the strong cross‐scale links between micro‐ and macroclimatic conditions are related to high cross‐scale similarity (low scale dependence) of species' realized temperature niches and, thus, species' spatial distributions. Location This study covers seven orders of magnitude of spatial scale, ranging from local‐scale (below a metre) and regional‐scale (kilometre) investigations in central European wetland ecosystems to continental‐scale (thousands of kilometres) studies of species' distributions. Methods We combined data on the spatial occurrence of species (vegetation records at local and regional scales, digitized distribution maps at the continental scale) with information about the corresponding temperature regime of vascular plant species occurring in environmentally stable wetland ecosystems characterized by strong cross‐scale links between micro‐ and macroclimatic conditions. Results We observed high cross‐scale similarity of the characteristics of species temperature niches across seven orders of magnitude of spatial scale. However, the importance of temperature as an abiotic driver decreased nonlinearly with decreasing scale, suggesting greater importance of additional (biotic) drivers of species' occurrence at small spatial scales. Main conclusions We report high cross‐scale similarity of realized temperature niches for species inhabiting ecosystems where small‐scale environmental noise is low and cross‐scale links between micro‐ and macroclimatic conditions are strong. By highlighting a strong relationship between abiotic and biotic cross‐scale similarity, our results will help to improve niche‐based species distribution modelling, one of the major assessment tools for determining the ecological effects of climate change.     

Community assembly is a race between immigration and adaptation: eco‐evolutionary interactions across spatial scales

Both ecological and evolutionary mechanisms have been proposed to describe how natural communities become assembled at both regional and biogeographical scales. Yet, these theories have largely been developed in isolation. Here, we unite these separate views and develop an integrated eco‐evolutionary framework of community assembly. We use a simulation approach to explore the factors determining the interplay between ecological and evolutionary mechanisms systematically across spatial scales. Our results suggest that the same set of ecological and evolutionary processes can determine community assembly at both regional and biogeographical scales. We find that the importance of evolution and community monopolization effects, defined as the eco‐evolutionary dynamics that occur when local adaptation of early established immigrants is fast enough to prevent the later immigration of better pre‐adapted species, are not restricted to adaptive radiations on remote islands. They occur at dispersal rates of up to ten individuals per generation, typical for many species at the scale of regional metacommunities. Dispersal capacity largely determines whether ecological species sorting or evolutionary monopolization structure metacommunity diversity and distribution patterns. However, other factors related to the spatial scale at which community assembly processes are acting, such as metacommunity size and the proportion of empty patches, also affect the relative importance of ecology versus evolution. We show that evolution often determines community assembly, and this conclusion is robust to a wide range of assumptions about spatial scale, mode of reproduction, and environmental structure. Moreover, we found that community monopolization effects occur even though species fully pre‐adapted to each habitat are abundant in the metacommunity, a scenario expected a priori to prevent any meaningful effect of evolution. Our results strongly support the idea that the same eco‐evolutionary processes underlie community assembly at regional and biogeographical scales.     

Island ecology and contingent theory: the role of spatial scale and taxonomic bias

Scale, the scale dependency of patterns and processes, and the ways that organisms scale their responses to these patterns and processes are central to island and landscape ecology. Here, we take a database of studies in island ecology and investigate how studies have changed over a 40-year period with respect to spatial scale and organisms studied. We demonstrate that there have been changes in the spatial scale of islands studied and that there is taxonomic bias in favour of vertebrates in island ecological studies when compared to scientific publications as a whole. We discuss how such taxonomic bias may have arisen and discuss the implications for ecology and biogeography.     

Understanding ecological change across large spatial,temporal and taxonomic scales: integrating data and methods in light of theory

The difficulty of integrating multiple theories, data and methods has slowed progress towards making unified inferences of ecological change generalizable across large spatial, temporal and taxonomic scales. However, recent progress towards a theoretical synthesis now provides a guiding framework for organizing and integrating all primary data and methods for spatiotemporal assemblage‐level inference in ecology. In this paper, we describe how recent theoretical developments can provide an organizing paradigm for linking advances in data collection and methodological frameworks across disparate ecological sub‐disciplines and across large spatial and temporal scales. First, we summarize the set of fundamental processes that determine change in multispecies assemblages across spatial and temporal scales by reviewing recent theoretical syntheses of community ecology. Second, we review recent advances in data and methods across the main sub‐disciplines concerned with ecological inference across large spatial, temporal and taxonomic scales, and organize them based on the primary fundamental processes they include, rather than the spatiotemporal scale of their inferences. Finally, we highlight how iteratively focusing on only one fundamental process at a time, but combining all relevant spatiotemporal data and methods, may reduce the conceptual challenges to integration among ecological sub‐disciplines. Moreover, we discuss a number of avenues for decreasing the practical barriers to integration among data and methods. We aim to reconcile the recent convergence of decades of thinking in community ecology and macroecology theory with the rapid progress in spatiotemporal approaches for assemblage‐level inference, at a time where a robust understanding of spatiotemporal change in ecological assemblages is more crucial than ever to conserve biodiversity.     

Spatial scale types and measurement of edge effects in ecology

Classification of spatial scales and measurement of edge effects in ecology were reviewed. The spatial scales can be classified into large scale (biome ecotone), meso-scale (ecological ecotone) and small scale (community ecotone) through the formation and maintenance of edge effects in ecology based on the synthetic analysis of published literatures. The biome ecotone is influenced by climate, regional dominant vegetation and terrain environment. The ecological ecotone is usually distributed in the transitional region with remarkable habitat heterogeneity. It connects adjacent ecosystems and affects the flow of energy and nutrient. Nowadays, study on edge effects in ecology mainly focuses on boundary sensitivity which associates with urban-rural ecotone, forest-grassland ecotone, agro-pastoral ecotone, forest-farmland ecotone, water-land ecotone and forest-swamp ecotone. As to the community ecotone which links with different patches to the interior of the community, previous studies focused on community edge, gap edge and treelines. The borderlines of different biome ecotones and the gradients of species distribution in the biome ecotones have been investigated through the method of quantitative ecology. The dynamic change, location and width of the ecological ecotone have been studied using the Geographic Information System (GIS), Remote Sensing (RS) and Global Positioning System (GPS) technologies and the landscape ecology theory. As important indicators, distance from edge, population, structure and diversity determined for establishing models can be applied to measure the intensity of edge effects and decide the positive or negative impact on communities. Although study on the edge effects in ecology was mostly reported at the meso-scale and small scale, study at large scale should be paid more attention as it is the potential value in ecology and global change fields.     

边缘效应的空间尺度与测度

综述了边缘效应的空间尺度类型以及在不同尺度上的测度方法.基于大量的研究整合,认为边缘效应空间尺度的划分,可以根据空间尺度的不同以及边缘效应形成和维持因素,分为大中小3个尺度类型,即大尺度的生物群区交错带、中尺度的景观类型之间的生态交错带和小尺度的斑块(生态系统)之间的群落交错区.大尺度主要是以植被气候带为标志的生物群区间的边缘效应,这种地带性的交错区主要受大气环境条件的影响.中尺度类型主要包括城乡交错带、林草交错带、农牧交错带等类型,是不同生态系统要素的空间交接地带,在物质能量等相互流动的作用下变得更为复杂.小尺度水平上是指斑块之间的交错所形成的边缘效应,受小地形等微环境条件及生物非生物等因子的制约,研究主要集中在群落边缘、林窗边缘和林线交错带等方面.对边缘效应测度的定量化研究有助于更加深入理解边缘效应.在大尺度水平上,边缘效应测度的研究主要是应用数量生态学等方法,研究不同气候带之间界线的划分及其物种分布的梯度规律性.中尺度水平上应用景观生态学的3S技术等方法,侧重于研究交错带的动态变化趋势及位置宽度的判定.小尺度水平上通过对距离边缘的长度,各群落中种群的数量、结构、多样性等定量指标的测定来构建测度公式,从而对边缘效应的强度进行量化,并反映边缘对群落的正负效应.总体上看,主要集中于中小尺度上,未来应该强化大尺度边缘效应测度的研究.     

Spatial hierarchical approach in community ecology: a way beyond high context-dependency and low predictability in local phenomena

Patterns and functioning of communities, which are determined by a set of processes operating at a large variety of spatial and temporal scales, exhibit quite high context-dependency and low predictability at the fine spatial scales at which recent studies have concentrated. More attention to broader scale and across-scale phenomena may be useful to search for general patterns and rules in communities. In this context, it is effective to incorporate hierarchical spatial scale explicitly into the experimental and sampling design of field studies, an approach referred to here as the spatial hierarchical approach, focusing on a particular assemblage in which biological interaction and species life history are well known. The spatial hierarchical approach can provide insight into the effects of scale in operating processes and answers to a number of important questions in community ecology such as: (1) detection of patterns and processes in spatiotemporal variability in communities, including how to explain the partitioning of pattern information of species diversity at a broad scale into finer scales, and the pattern of spatial variability of community properties at the finest spatial scale; (2) evaluation of changes in patterns observed in macroecology at finer scales; (3) testing of models explaining the coexistence of competing species; and (4) detection of latitudinal patterns in spatiotemporal variability in communities and their causal processes.     

Earlier breeding,lower success: does the spatial scale of climatic conditions matter in a migratory passerine bird?

Following over 20 years of research on the climatic effects on biodiversity we now have strong evidence that climate change affects phenology, fitness, and distribution ranges of different taxa, including birds. Bird phenology likely responds to changes in local weather. It is also affected by climatic year‐to‐year variations on larger scales. Although such scale‐related effects are common in ecology, most studies analyzing the effects of climate change were accomplished using climatic information on a single spatial scale. In this study, we aimed at determining the scale‐dependent sensitivity of breeding phenology and success to climate change in a migratory passerine bird, the barn swallow (Hirundo rustica). For both annual broods, we investigated effects of local weather (local scale) and the North Atlantic Oscillation (NAO, large scale) on the timing of breeding and breeding success. Consistent with previous studies in migratory birds we found that barn swallows in Eastern Germany bred progressively earlier. At the same time, they showed reduced breeding success over time in response to recent climatic changes. Responses to climatic variation were observed on both local and large climatic scales, but they differed with respect to the ecological process considered. Specifically, we found that the timing of breeding was primarily influenced by large‐scale NAO variations and to a lesser extent by local weather on the breeding grounds. Conversely, climatic conditions on the local scale affected breeding success, exclusively. The observed decrease in breeding success over years is likely a consequence of scale‐related mismatches between climatic conditions during different breeding phases. This provides further evidence that a species' response of earlier breeding may not be enough to cope with climate change. Our results emphasize the importance of considering the response of ecological processes along different climatic scales in order to better understand the complexity of climate change effects on biodiversity.     

A sense of scale: Foraging cetaceans' use of scale‐dependent multimodal sensory systems

Research on cetacean foraging ecology is central to our understanding of their spatial and behavioral ecology. Yet, functional mechanisms by which cetaceans detect prey across different scales remain unclear. Here, I postulate that cetaceans utilize a scale‐dependent, multimodal sensory system to assess and increase prey encounters. I review the literature on cetacean sensory systems related to foraging ecology, and hypothesize the effective scales of each sensory modality to inform foraging opportunities. Next, I build two “scale‐of‐senses” schematics for the general groups of dolphins and baleen whales. These schematics illustrate the hypothetical interchange of sensory modalities used to locate and discriminate prey at spatial scales ranging from 0 m to 1,000 km: (1) vision, (2) audition (sound production and sound reception), (3) chemoreception, (4) magnetoreception, and somatosensory perception of (5) prey, or (6) oceanographic stimuli. The schematics illustrate how a cetacean may integrate sensory modalities to form an adaptive foraging landscape as a function of distance to prey. The scale‐of‐senses schematic is flexible, allowing for case‐specific application and enhancement with improved cetacean sensory data. The framework serves to improve our understanding of functional cetacean foraging ecology, and to develop new hypotheses, methods, and results regarding how cetaceans forage at multiple scales.     

Universal scaling of species‐abundance distributions across multiple scales

The scale‐dependent species abundance distribution (SAD) is fundamental in ecology, but few spatially explicit models of this pattern have thus far been studied. Here we show spatially explicit neutral model predictions for SADs over a wide range of spatial scales, which appear to match empirical patterns qualitatively. We find that the assumption of a log‐series SAD in the metacommunity made by spatially implicit neutral models can be justified with a spatially explicit model in the large area limit. Furthermore, our model predicts that SADs on multiple scales are characterized by a single, compound parameter that represents the ratio of the survey area to the species’ average biogeographic range (which is in turn set by the speciation rate and the dispersal distance). This intriguing prediction is in line with recent empirical evidence for a universal scaling of the species‐area curve. Hence we hypothesize that empirical SAD patterns will show a similar universal scaling for many different taxa and across multiple spatial scales.     

A conceptual framework for the spatial analysis of functional trait diversity

Functional trait diversity is a popular tool in modern ecology, mainly used to infer assembly processes and ecosystem functioning. Patterns of functional trait diversity are shaped by ecological processes such as environmental filtering, species interactions and dispersal that are inherently spatial, and different processes may operate at different spatial scales. Adding a spatial dimension to the analysis of functional trait diversity may thus increase our ability to infer community assembly processes and to predict change in assembly processes following disturbance or land‐use change. Richness, evenness and divergence of functional traits are commonly used indices of functional trait diversity that are known to respond differently to large‐scale filters related to environmental heterogeneity and dispersal and fine‐scale filters related to species interactions (competition). Recent developments in spatial statistics make it possible to separately quantify large‐scale patterns (variation in local means) and fine‐scale patterns (variation around local means) by decomposing overall spatial autocorrelation quantified by Moran's coefficient into its positive and negative components using Moran eigenvector maps (MEM). We thus propose to identify the spatial signature of multiple ecological processes that are potentially acting at different spatial scales by contrasting positive and negative components of spatial autocorrelation for each of the three indices of functional trait diversity. We illustrate this approach with a case study from riparian plant communities, where we test the effects of disturbance on spatial patterns of functional trait diversity. The fine‐scale pattern of all three indices was increased in the disturbed versus control habitat, suggesting an increase in local scale competition and an overall increase in unexplained variance in the post‐disturbance versus control community. Further research using simulation modeling should focus on establishing the proposed link between community assembly rules and spatial patterns of functional trait diversity to maximize our ability to infer multiple processes from spatial community structure.     

Fish species diversity among spatial scales of altered temperate rivers

Aim The alteration of flowing systems over the past century has led to significant changes to the processes that drive these complex environments as well as to the scales at which these processes act. Recently, efforts have begun in earnest to restore some semblance of ecosystem diversity, but there is little understanding of exactly on what spatial scale or scales biotic diversity is responding. We investigated the manner in which fish diversity is partitioned at multiple spatial scales in two rivers in the central United States. Location The Missouri and Illinois rivers of the central United States. Methods We analysed how fish diversity was partitioned within the Illinois River and Missouri River systems by sampling each river under hierarchical frameworks that allowed analysis at section (large), reach (intermediate), and site (small) scales. We tested the hypothesis that there are scale‐dependent responses of fish diversity using an additive partitioning approach. Results Site alpha diversity was significantly higher than expected in both the Illinois and Missouri rivers. The relative contribution of alpha diversity to total diversity at a given spatial scale increased for the Illinois River, but not for the Missouri River, in that the highest alpha diversity contribution peaked at the reach scale. Diversity patterns from both rivers suggest that diversity at the site scale plays a significant role in determining the overall diversity in these systems. However, there is a substantial contribution at larger scales that warrants consideration when attempts are being made to protect or restore diversity and other ecosystem parameters. Main conclusions Understanding the variation of diversity in riverine systems is crucial for providing insight not only into how biotic communities respond to scale‐dependent factors, but also into the underlying abiotic and biotic factors that generate patterns of diversity across scales. These insights, in turn, are important for ensuring that restoration and management activities are targeting the appropriate scales for remediation. A lack of understanding of this issue could have negative outcomes for the recovery of a community in a restoration scenario, as well as resulting in a low economic return on restoration investments, which could hinder future efforts.     

Spatial variation of species diversity across scales in an old-growth temperate forest of China

Species richness and abundance are the two most important diversity variables. Species abundance is additive when aggregated across spatial scale, whereas species richness is non-additive. This study analyzes the effect of spatial scale and site on species abundance and richness in a 25-ha temperate forest plot in the Changbai Mountains, northeastern China. The result shows that species abundance and richness are not only dependent on spatial scales, but also dependent on site. Species abundance responds linearly to changes of spatial scale with no intersection in different sites of the study area. However, although species richness also increases with the increase of spatial scale, there are some intersections for the different sites, suggesting that a species-rich site does not always have a high value if the spatial scale is changed. In all, with respect to additive variables, it is relatively easy to extrapolate them from one spatial scale to another spatial scale, as they and the spatial scale usually form a linear relationship. In contrast, non-additive variables are difficult to extrapolate across spatial scales, because they often respond nonlinearly to spatial scale changes. In order to extrapolate these non-additive variables across spatial scales, it is necessary to estimate the relationships between them and spatial scales. As a result, extrapolation of information among spatial scales may be possible, but very difficult, especially for non-additive variables. Because the 25-ha Changbai plot is very small compared to the extent of the world temperate forests, and the vegetation is a relatively uniform type, more such studies in other ecosystems are needed before theories and generalization about scaling effects can be formulated.