A continental measure of urbanness predicts avian response to local urbanization

Understanding species-specific relationships with their environment is essential for ecology, biogeography and conservation biology. Moreover, understanding how these relationships change with spatial scale is critical to mitigating potential threats to biodiversity. But methods which measure inter-specific variation in response to environmental parameters that are also generalizable across multiple spatial scales are scarce. We used broad-scale avian citizen science data, over continental Australia, integrated with remotely-sensed products, to produce a measure of urban-tolerance for a given species at a continental-scale. We then compared these urban-tolerances to modelled responses to urbanization at a local-scale, based on systematic sampling within four small cities. For 49 species which had sufficient data for modelling, we found a significant relationship (R² = 0.51) between continental-scale urbanness and local-scale urbanness. We also found that relatively few citizen science observations (~250) are necessary for reliable estimates of continental-scale species-specific urban scores to predict local-scale response to urbanization. Our approach demonstrates the applicability of broad-scale citizen science data, contrasting both the spatial grain and extent of standard point-count surveys generally only conducted at small spatial scales. Continental-scale responses in Australia are representative of small-scale responses to urbanization among four small cities in Australia, suggesting that our method of producing species-specific urban scores is robust and may be generalized to other locations lacking appropriate data.     

生态学中的尺度问题——尺度上推

尺度推绎是生态学理论和应用的核心。如何在一个异质景观中进行尺度推绎仍然是一个悬而未决的科学难题,是对当今生态学家在全球变化背景下研究环境问题的重大挑战。就目前的研究,一般可分为四大类尺度推绎途径:空间分析法(如分维分析法和小波分析法)、基于相似性的尺度上推方法、基于局域动态模型的尺度上推方法、随机(模型)法。基于相似性的尺度上推方法来源于生物学上的异量关联,可将其思想延伸至空间上,研究物种丰富度、自然河网、地形特征、生态学格局或过程变量和景观指数等。基于局域动态模型的尺度上推方法需要首先确定是否进行跨尺度推绎,以及是否考虑空间单元之间的水平相互作用和反馈,然后再应用具体的方法或途径,如简单聚合法、有效值外推法、直接外推法、期望值外推、显式积分法和空间相互作用模拟法等。随机(模型)法以其它尺度上推方法为基础,根据研究的是单个景观,还是多个景观,采用不同的途径。理解、定量和降低尺度推绎结果的不确定性已经变得越来越重要,但相关研究仍然极少。以上所有有关尺度推绎的方法、途径和结果分析共同构成了尺度推绎的概念框架。     

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.     

β‐diversity scaling patterns are consistent across metrics and taxa

β‐diversity (variation in community composition) is a fundamental component of biodiversity, with implications for macroecology, community ecology and conservation. However, its scaling properties are poorly understood. Here, we systematically assessed the spatial scaling of β‐diversity using 12 empirical large‐scale datasets including different taxonomic groups, by examining two conceptual types of β‐diversity and explicitly considering the turnover and nestedness components. We found highly consistent patterns across datasets. Multiple‐site β‐diversity (i.e. variation across multiple sites) scaling curves were remarkably consistent, with β‐diversity decreasing with sampled area according to a power law. For pairwise dissimilarities, the rates of increase of dissimilarity with geographic distance remained largely constant across scales, while grain size (or scale level) had a stronger effect on overall dissimilarity. In both analyses, turnover was the main contributor to β‐diversity, following total β‐diversity patterns closely, while the nestedness component was largely insensitive to scale changes. Our results highlight the importance of integrating both inter‐ and intraspecific aggregation patterns across spatial scales, which underpin substantial differences in community structure from local to regional scales.     

Spatial scales in the study of reef fishes: A theoretical perspective

Abstract Theoretical models imply that spatial scale derives its greatest importance through interactions between density-dependent processes and spatial variation in population densities and environmental variables. Such interactions cause population dynamics on large spatial scales to differ in important ways from predictions based on measurements of population dynamics at smaller scales, a phenomenon called the scale transition. These differences can account for large-scale population stability and species coexistence. The interactions between density dependence and spatial variation that lead to the scale transition can be understood by the process of non-linear averaging, which shows how variance originating on various spatial scales contributes to large-scale population dynamics. Variance originating below the scale of density dependence contributes less to the scale transition as the spatial scale of the variation declines, while variation originating on or above the scale of density dependence contributes independently of the spatial scale of the variation.     

Little evidence for climate effects on local-scale structure and dynamics of California kelp forest communities

Climatic variables such as temperature and precipitation play an important role in controlling local and regional scale differences in population dynamics and species distributions, and large-scale climatic events such as El Niño southern oscillation (ENSO) have been shown to affect population dynamics of key species in many ecosystems, particularly in kelp forests. Few studies have been able to evaluate the consequences of climate variables on the structure and dynamics of biological communities, in large part because the lack of data at appropriate spatial and temporal scales has made it difficult to adequately address local-scale responses of species and communities to such events over relevant time scales. Here, we combined an unprecedented dataset of kelp forest species' abundances from the Channel Islands, California with data for several local, regional, and global scale climatic variables to evaluate the temporal and spatial scale at which one can detect community-wide effects of climate variables, in particular ENSO events. We found large and significant local-scale differences in community structure, but these differences were not related to differences in climatic variables. Moreover, giant kelp abundance, which has been shown to be highly sensitive to water temperature and storm disturbance, was a poor predictor of community differences, and all communities tended to decline in abundance over the 20-year sampling period, suggesting a press perturbation to the system such as PDO cycles or sustained fishing pressure. Although ENSO events can have dramatic impacts on the abundance and distribution of giant kelp itself across the range of the species, such events appear to have little effect on local-scale kelp forest community structure or dynamics.     

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.     

尺度与密度:测定不同尺度下的种群密度

群落中的种群密度由于空间尺度的变化而存在着一定差异,那么,某一种群的密度随着空间尺度的变化会发生怎样的变化?抑或某一物种相对于另一物种而言,随着空间尺度的变化其密度会怎样变化?这是与尺度有关的种群密度问题,当属生态学的基本问题.该文提出这样的问题,并把不同尺度下的种群密度称之为尺度密度(scale density).O-Ring函数的实质是计算不同尺度下的种群密度.因此,在研究实例中,应用O-Ring函数计算了典型草原处于不同恢复阶段的羊草(Leymuschinensis)种群、米氏冰草(Agropyron michnoi)种群,以及米氏冰草种群相对于羊草种群在不同尺度下的种群密度,结果发现:羊草和米氏冰草2个种群的尺度密度,在小尺度范围内严重退化群落均高于恢复演替群落,这一结果验证了“胁迫梯度假说”,同时表明该结果是放牧胁迫下正相互作用所致;通过比较羊草种群与米氏冰草相对于羊草的尺度密度发现,在严重退化的群落中,羊草与米氏冰草的种间关联为负联结,这种负联结是由正相互作用引起的,而在恢复8年和恢复21年群落中,二者之间是正联结,当为竞争所致.该实例说明分析种群密度随尺度变化的规律对于深入认识生态学问题可能会有很大帮助.     

Scaling up from local competition to regional coexistence across two scales of spatial heterogeneity: insect larvae in the fruits of Apeiba membranacea

Species that live in patchy and ephemeral habitats can compete strongly for resources within patches at a small scale. The ramifications of these interactions for population dynamics and coexistence at regional scales will depend on the intraspecific and interspecific distributions of individuals among patches. Spatial heterogeneity due to independent aggregation of competitors among patchy habitats is an important mechanism maintaining species diversity. I describe regional patterns of aggregation for four species of insect larvae in the fruits of Apeiba membranacea, a Neotropical rainforest tree. This aggregation results from variation in densities at a small scale (among the fruits under a single tree), compounded by significant variation among trees in both mean densities and degrees of aggregation. Both the degrees of aggregation and mean densities are statistically independent within and across species at both spatial scales. I evaluate the regional consequences of these spatial patterns by using maximum likelihood methods to parameterize a model that includes both explicit measures of the strength of competition and spatial variation at both within- and among-tree spatial scales. Despite strong competitive interactions among these species, during 2 years the observed spatial variation at both scales combined was sufficient to explain the coexistence of these species, although other coexistence mechanisms may also operate simultaneously. The observed spatial variation at small spatial scales may not be sufficient for coexistence, indicating the importance of considering multiple sources of spatial heterogeneity when scaling up from experiments that investigate local interactions to regional patterns of coexistence.     

Scaling up from local competition to regional coexistence across two scales of spatial heterogeneity: insect larvae in the fruits of <Emphasis Type="Italic">Apeiba membranacea</Emphasis>

Species that live in patchy and ephemeral habitats can compete strongly for resources within patches at a small scale. The ramifications of these interactions for population dynamics and coexistence at regional scales will depend on the intraspecific and interspecific distributions of individuals among patches. Spatial heterogeneity due to independent aggregation of competitors among patchy habitats is an important mechanism maintaining species diversity. I describe regional patterns of aggregation for four species of insect larvae in the fruits of Apeiba membranacea, a Neotropical rainforest tree. This aggregation results from variation in densities at a small scale (among the fruits under a single tree), compounded by significant variation among trees in both mean densities and degrees of aggregation. Both the degrees of aggregation and mean densities are statistically independent within and across species at both spatial scales. I evaluate the regional consequences of these spatial patterns by using maximum likelihood methods to parameterize a model that includes both explicit measures of the strength of competition and spatial variation at both within- and among-tree spatial scales. Despite strong competitive interactions among these species, during 2 years the observed spatial variation at both scales combined was sufficient to explain the coexistence of these species, although other coexistence mechanisms may also operate simultaneously. The observed spatial variation at small spatial scales may not be sufficient for coexistence, indicating the importance of considering multiple sources of spatial heterogeneity when scaling up from experiments that investigate local interactions to regional patterns of coexistence.     

Scaling effects and spatio-temporal multilevel dynamics in epileptic seizures

Epileptic seizures are one of the most well-known dysfunctions of the nervous system. During a seizure, a highly synchronized behavior of neural activity is observed that can cause symptoms ranging from mild sensual malfunctions to the complete loss of body control. In this paper, we aim to contribute towards a better understanding of the dynamical systems phenomena that cause seizures. Based on data analysis and modelling, seizure dynamics can be identified to possess multiple spatial scales and on each spatial scale also multiple time scales. At each scale, we reach several novel insights. On the smallest spatial scale we consider single model neurons and investigate early-warning signs of spiking. This introduces the theory of critical transitions to excitable systems. For clusters of neurons (or neuronal regions) we use patient data and find oscillatory behavior and new scaling laws near the seizure onset. These scalings lead to substantiate the conjecture obtained from mean-field models that a Hopf bifurcation could be involved near seizure onset. On the largest spatial scale we introduce a measure based on phase-locking intervals and wavelets into seizure modelling. It is used to resolve synchronization between different regions in the brain and identifies time-shifted scaling laws at different wavelet scales. We also compare our wavelet-based multiscale approach with maximum linear cross-correlation and mean-phase coherence measures.     

Influences of catchment and corridor imperviousness on urban stream macroinvertebrate communities at multiple spatial scales

Resolving land cover hierarchy relationships in urban settings is important for defining the scale and type of management required to enhance stream health. We investigated associations between macroinvertebrate assemblages in urban streams of Hamilton, New Zealand, and environmental variables measured at multiple spatial scales comprising (i) local-scale physicochemical conditions, (ii) impervious area in multiple stream corridor widths (30, 50 and 100 m) along segments (sections of stream between tributary nodes) and for entire upstream networks, and (iii) total impervious area in stream segment sub-catchments and upstream catchments. Imperviousness was higher for stream segment sub-catchments than for entire catchments because of the agricultural headwaters of some urban streams. Imperviousness declined as corridor width declined at both segment and catchment scales reflecting the vegetated cover along most urban stream gullies. Upstream catchment imperviousness was strongly and inversely correlated with dissolved organic carbon concentration, whereas segment and upstream corridor scales were correlated with water temperature and pH. Corridor imperviousness appeared to be a stronger predictor than catchment imperviousness of Ephemeroptera, Plecoptera and Trichoptera taxa richness and the Quantitative Urban Community Index specifically developed to assess impacts of urbanisation. In contrast, imperviousness at all measured scales added only marginal improvement in assemblage-based models over that provided by the local-scale physicochemical variables of reach width, habitat quality, macrophyte cover, pH and dissolved oxygen concentration. These findings infer variable scales of influence affecting macroinvertebrate communities in urban streams and suggest that it may be important to consider local and corridor factors when determining mechanisms of urbanisation impacts and potential management options.     

Integration of local and regional species-area relationships from space-time species accumulation

A long-standing observation in community ecology is that the scaling of species richness, as exemplified by species-area curves, differs on local and regional scales. This decoupling of scales may be largely due to sampling processes (the increasing constraint imposed by sampling fewer individuals at fine scales), as distinct from ecological processes, such as environmental heterogeneity, that operate across scales. Removal of the sampling constraint from fine-scale richness estimates should yield species-area curves that behave like those of the regions in which they are embedded, but an effective method for this removal has not been available. We suggest an approach that incorporates the manner in which small areas accumulate species over time as a way to remove the signature of sampling processes from fine-scale species-area curves. We report for three species-rich grasslands from two continents how local plant species richness is distributed through time at multiple, nested spatial scales, and we ask whether sampling-corrected curves reflect the spatial scaling of richness of each larger floristic province. Our analysis suggests that fine-scale values of richness are highly constrained by sampling processes, but once these constraints are removed, the spatial scaling of species richness is consistent from the scale of individuals to that of an entire province.     

Assessing riverscape-scale variation in fish life history using banded sculpin (<Emphasis Type="Italic">Cottus carolinae</Emphasis>)

Advancing the field of fish ecology requires a shift in focus from describing patterns in species occurrences to understanding the mechanisms that regulate distributions and abundances across broad scales. For stream fish ecology, this includes understanding environmental mechanisms that regulate stream fish demographic properties at the scale of stream networks or riverscapes. Despite the fact that Banded Sculpin Cottus carolinae occupy a diversity of habitats and stream sizes across the southeastern United States, relatively little is known about the demography of this species. We assessed annual demographic properties (reproduction, growth, and survival) of C. carolinae collected monthly from four sites distributed longitudinally along the Roaring River riverscape in Tennessee to simultaneously describe life history attributes of the species and address riverscape-scale variation in population dynamics. Cottus carolinae lived for a maximum of four years, local populations were dominated by age-0 and age-1 individuals, reproduction began after one year, spawning occurred during December and January, and mean ova number was 398. A life history tradeoff between growth (robustness) and survival was evident at one site where water temperature and flow were least variable, otherwise life history attributes were consistent across the riverscape despite longitudinal changes in abiotic variables. Our work illustrates the potential for muted population responses to a strong hydrologic gradient in stream size and highlights the stability inherent with fish life history adaptations to natural environmental regimes across broad spatial scales.     

Spatial scaling of ecosystem C and N in a subtropical savanna landscape

Widely occurred woody encroachment in grass‐dominated ecosystems has the potential to influence soil organic carbon (SOC) and total nitrogen (TN) pools at local, regional, and global scales. Evaluation of this potential requires assessment of both pool sizes and their spatial patterns. We quantified SOC and TN, their relationships with soil and vegetation attributes, and their spatial scaling along a catena (hill‐slope) gradient in the southern Great Plains, USA where woody cover has increased substantially over the past 100 years. Quadrat variance analysis revealed spatial variation in SOC and TN at two scales. The larger scale variation (40–45 m) was approximately the distance between centers of woody plant communities and their adjoining herbaceous patches. The smaller scale variation (10 m) appeared to reflect the local influence of shrubs on SOC and TN. Litter, root biomass, shrub, and tree basal area (a proxy for plant age) exhibited not only similar spatial scales, but also strong correlations with SOC and TN, suggesting invasive woody plants alter both the storage and spatial scaling of SOC and TN through ecological processes related primarily to root turnover and, to a lesser extent litter production, as mediated by time of occupancy. Forb and grass biomass were not significantly correlated with SOC and TN suggesting that changes in herbaceous vegetation have not been the driving force for the observed changes in SOC and TN. Because SOC and TN varied at two scales, it would be inappropriate to estimate SOC and TN pools at broad scales by extrapolating from point sampling at fine scales. Sampling designs that capture variation at multiple scales are required to estimate SOC and TN pools at broader scales. Knowledge of spatial scaling and correlations will be necessary to design field sampling protocols to quantify the biogeochemical consequences of woody plant encroachment at broad scales.     

生态学研究中的尺度问题

尺度是生态学中的核心问题,时间和空间尺度包含于任何生态过程中,尺度在生态学研究中越来越显现出其重要性,原因是解决地球环境问题要求在大尺度上理解格局和过程,而以前生态学调查的数据主要是基于小尺度的,并且有许多研究表明,一个生态问题的结论在很大程度上取决于研究所采纳的尺度。但目前尺度研究还存在很多的问题;一些相关概念易与尺度混淆;缺乏成熟的尺度识别方法和系统完备的尺度转换方法等。针对这些问题,本文首先阐明了尺度的确切含义,并对尺度研究的发展历史,重要性作了问题的阐述：分析探讨了尺度与等级理论,格局的关系,及尺度识别,尺度转换的方法和发展趋势。     

空间幅度变化对景观格局分析的影响

景观格局指数是量化描述景观格局特征的主要方法之一,各种格局指数的尺度依赖性使比较分析景观格局特征和尺度推绎复杂化,分析不同指数随空间幅度变化的一般行为有助于景观格局分析结果的解释和降低空间尺度推绎的复杂性。研究以2种真实景观和27种模拟景观为分析对象,考查了16种常用的景观水平格局指数随空间幅度变化行为。根据这些指数因幅度变化行为可预测性把它们分为两类：第1类随幅度变化可预测性强,指数与幅度之间的关系可用简单的函数关系来表达;这类指数包括缀块数、边界总长、景观形状指数和缀块丰度密度;前两者随幅度增加呈幂函数形式增加,而缀块丰度密度随幅度增加呈幂函数下降,景观形状指数随幅度增加呈直线增加。第2类指数随幅度变化的可预测性较差,指数随幅度的变化存在多种可能(不同形式的增加、减小或保持不变),不可用一种或多种简单的函数关系来描述所有的情况。这类指数包括缀块丰度、缀块密度、边界密度、最大缀块指数、平均缀块面积、缀块面积标准差、缀块面积变异系数、平均缀块形状指数、面积加权平均缀块形状指数、双对数回归分维数、聚集度指数与Shannon多样性指数。第2类指数随幅度的变化行为受景观格局特征和指数本身算法的影响。总体上来说,第2类格局指数随幅度变化呈折线增加或减小;但当景观的缀块类型较多、各类型优势度比较均等、空间分布格局比较随机时,它们随幅度变化形为的可预测性增加,随幅度增加的函数关系主要有3种：幂函数减小、对数函数增加或直线增加,因指数和格局特征不同而异。研究的结果在揭示了常用景观指数随幅度变化的一般关系外,也启示我们在进行景观格局的比较分析,比较景观的幅度应相同或采用尺度效应图(scalograms)更有效。     

Environmental variables interact across spatial scales to structure trichopteran assemblages in Ouachita Mountain rivers

An important goal in aquatic ecology is to determine the interacting variables that regulate community structure; however, complex biotic and abiotic interactions coupled with the significance of scale have confounded the interpretation of community data. We evaluated stream and riparian habitat features in southeastern Oklahoma, USA at a range of spatial scales from local, in-stream variables to large-scale, regional characteristics to address the following questions: (1) How much variation in trichopteran community composition can be attributed to local, regional, and spatial variables? and (2) What environmental variables are most important in determining trichopteran community structure? We collected data on caddisfly community structure, local and regional environmental variables, and spatial location on the landscape from 25 sites in four rivers. We analyzed these data using canonical correspondence analysis (CCA) and variation partitioning. Our analysis explained approximately 60% of the variation in caddisfly community composition. We found that local and regional environmental variables were near equal in importance in governing caddisfly communities, with each accounting for approximately a quarter of the explained variation. Although pure spatial variables were less important, the amount of variation shared among spatial variables and local and regional variables was substantial, indicating that biogeographic history is also key to understanding caddisfly distributions. We also found a strong influence of human landuse (i.e., percent of land in agriculture, distance to roads) on caddisfly community composition. Our study indicated that communities are influenced by factors across scales, and that bioassessments should focus on not only local habitat conditions, but also incorporate larger-scale factors.     

Determinants of Spatial Distribution in a Bee Community: Nesting Resources,Flower Resources,and Body Size

Understanding biodiversity distribution is a primary goal of community ecology. At a landscape scale, bee communities are affected by habitat composition, anthropogenic land use, and fragmentation. However, little information is available on local-scale spatial distribution of bee communities within habitats that are uniform at the landscape scale. We studied a bee community along with floral and nesting resources over a 32 km² area of uninterrupted Mediterranean scrubland. Our objectives were (i) to analyze floral and nesting resource composition at the habitat scale. We ask whether these resources follow a geographical pattern across the scrubland at bee-foraging relevant distances; (ii) to analyze the distribution of bee composition across the scrubland. Bees being highly mobile organisms, we ask whether bee composition shows a homogeneous distribution or else varies spatially. If so, we ask whether this variation is irregular or follows a geographical pattern and whether bees respond primarily to flower or to nesting resources; and (iii) to establish whether body size influences the response to local resource availability and ultimately spatial distribution. We obtained 6580 specimens belonging to 98 species. Despite bee mobility and the absence of environmental barriers, our bee community shows a clear geographical pattern. This pattern is mostly attributable to heterogeneous distribution of small (<55 mg) species (with presumed smaller foraging ranges), and is mostly explained by flower resources rather than nesting substrates. Even then, a large proportion (54.8%) of spatial variability remains unexplained by flower or nesting resources. We conclude that bee communities are strongly conditioned by local effects and may exhibit spatial heterogeneity patterns at a scale as low as 500–1000 m in patches of homogeneous habitat. These results have important implications for local pollination dynamics and spatial variation of plant-pollinator networks.