Scaling processes and problems

A personal view is presented of current approaches to scaling processes and variables in the context of better understanding ecosystem function and predicting the consequences of global environmental change. Issues considered include spatial and temporal scales of interest, the scaling process, scaling strategy, scaling problems, heterogeneity, patchiness and non-linearity, aggregation methodology and feedbacks. Knowledge of processes in plants and vegetation is largely at small scales. The transfer of this knowledge up to larger spatial and longer temporal scales is an open-ended process with potential errors arising from heterogeneity and patchiness in the distribution of processes and non-linearities in the functional relationships between processes and environmental variables. Scaling now covers several orders of magnitude with respect to spatial and temporal scales with attendant risks of propagating errors. At larger scales the wide diversity of vegetation classes poses a problem, and it is suggested that this can be countered by classifying classes of vegetation (not species) into a small number of ‘functional types’ of vegetation. Scaling through summation of component processes and through derivation of appropriately averaged parameters is considered. However, the increasing role of feedbacks at larger spatial and longer temporal scales is an essential feature of the scaling process. Thus, understanding the feedbacks and including them in upscaling schemes is a major priority. A scaling strategy is outlined to minimize the propagation of errors. Because the scaling process is open-ended it is essential that good models are used and tested at each increase in scale.     

Scale of habitat connectivity and colonization in fragmented nuthatch populations

Studies of effects of landscape pattern on population dynamics should consider the spatial scale at which habitat connectivity varies relative to the spatial scale of the species' behavioral response. In this paper, I investigate the relationship between the degree of connectivity of wooded patches measured at different spatial scales and the colonization of these patches by nuthatch Sitta europaea populations. I used different threshold distances to calculate the degree of habitat connectivity. A threshold distance is the distance beyond which the probability of successfully bridging the distance by dispersing individuals is assumed to decrease rapidly. The sum of the degree of connectivity over all patches in a region provides a measure for the overall degree of connectivity. Based on this measure, I could first give an indication under what conditions species may experience effects of constrained dispersal in that region. Second, the degree of connectivity of the individual patches was related with the observed colonizations of these patches. The degree of habitat connectivity measured for threshold distances of ea 2.4-3 km best explains the colonization probability of unoccupied patches. These threshold distances give an indication of the distances covered by dispersing nuthatches that led to successful colonizations.     

Effects of matrix heterogeneity on animal dispersal: from individual behavior to metapopulation-level parameters

Mounting theoretical and empirical evidence shows that matrix heterogeneity may have contrasting effects on metapopulation dynamics by contributing to patch isolation in nontrivial ways. We analyze the movement properties during interpatch dispersal in a metapopulation of Iberian lynx (Lynx pardinus). On a daily temporal scale, lynx habitat selection defines two types of matrix habitats where individuals may move: open and dispersal habitats (avoided and used as available, respectively). There was a strong and complex impact of matrix heterogeneity on movement properties at several temporal scales (hourly and daily radiolocations and the entire dispersal event). We use the movement properties on the hourly temporal scale to build a simulation model to reconstruct individual dispersal events. The two most important parameters affecting model predictions at both the individual (daily) and metapopulation scales were related to the movement capacity (number of movement steps per day and autocorrelation in dispersal habitat) followed by the parameters representing the habitat selection in the matrix. The model adequately reproduced field estimates of population-level parameters (e.g., interpatch connectivity, maximum and final dispersal distances), and its performance was clearly improved when including the effect of matrix heterogeneity on movement properties. To assume there is a homogeneous matrix results in large errors in the estimate of interpatch connectivity, especially for close patches separated by open habitat or corridors of dispersal habitat, showing how important it is to consider matrix heterogeneity when it is present. Movement properties affect the interaction of dispersing individuals with the landscape and can be used as a mechanistic representation of dispersal at the metapopulation level. This is so when the effect of matrix heterogeneity on movement properties is evaluated under biologically meaningful spatial and temporal scales.     

Landscape heterogeneity at multiple spatial scales enhances spider species richness in an agricultural landscape

Landscape supplementation, which enhances densities of organisms by combination of different landscape elements, is likely common in heterogeneous landscapes, but its prevalence and effects on species richness have been little explored. Using grassland-dwelling spiders in an agricultural landscape, we postulated that richness and abundances of major constituent species are both highest in intermediate mixtures of forests and paddy fields, and that this effect derives from multi-scale landscape heterogeneity. We collected spiders in 35 grasslands in an agricultural landscape in Japan and determined how species richness and abundances of major species related to local and landscape factors across different spatial scales. We used a generalized linear model to fit data, created all possible combinations of variables at 15 spatial scales, and then explored the best models using Akaike's information criterion. Species richness showed a hump-shaped pattern in relation to surrounding forest cover, and the spatial scale determining this relationship was a 300–500-m radius around the study sites. Local variables were of minor importance for species richness. Abundances of major species also exhibited a hump-shaped pattern when plotted against forest cover. Thus, a combination of paddy fields and forests is important for enhancement of grassland spider species richness and abundance, suggesting habitat supplementation. The effective spatial scales determining abundances varied, ranging from 200 to >1000 m, probably representing different dispersal abilities. Landscape compositional heterogeneity at multiple spatial scales may be thus crucial for the maintenance of species diversity.     

A wavelet‐based approach to evaluate the roles of structural and functional landscape heterogeneity in animal space use at multiple scales

The functional relationship between habitat utilization and landscape spatial heterogeneity is fundamental to understanding the spatial nature of animal distribution across scales. Although structural and functional properties of landscape spatial heterogeneity can have different consequences for animal species, few studies have explicitly considered both forms of heterogeneity, partly due to the lack of general methods for direct assessment of scale‐specific associations between variables. We present a wavelet‐based approach to evaluate the roles of structural and functional landscape spatial heterogeneity in animal space use at multiple spatial scales. As a case study, we examined scale‐specific space use patterns of American black bears Ursus americanus in response to structural and functional spatial heterogeneity as well as spatial patterns of vegetation age‐classes in a Canadian boreal forest. We found strong differences in the effects of structural and functional spatial heterogeneity and the scales at which they are associated with the patterns of habitat use by black bears. Functional heterogeneity alone affected space use at 800 and 1600‐m scales, but had significant effects when interacting with structural heterogeneity at 400, 800, and 1600‐m scales. Compared with male bears, female black bears were most sensitive to patterns of forage abundance at intermediate scales, or more specifically, in young and regenerating forests that provide abundant soft mast in boreal forests. Our study highlights the importance of accounting for scale‐dependent properties of (structural and functional) spatial heterogeneity in assessing the ecological effects of landscape components and the effectiveness of the wavelet transform technique in identifying such scale‐specific relationships.     

Anthropogenic impact on habitat connectivity: A multidimensional human footprint index evaluated in a highly biodiverse landscape of Mexico

Evaluating the cumulative effects of the human footprint on landscape connectivity is crucial for implementing policies for the appropriate management and conservation of landscapes. We present an adjusted multidimensional spatial human footprint index (SHFI) to analyze the effects of landscape transformation on the remnant habitat connectivity for 40 terrestrial mammal species representative of the Trans-Mexican Volcanic System in Michoacán (TMVS_Mich), in western central Mexico. We adjusted the SHFI by adding fragmentation and habitat loss to its original three components: land use intensity, time of human landscape intervention, and biophysical vulnerability. The adjusted SHFI was applied to four scenarios: one grouping all species and three grouping several species by habitat spatial requirements. Using the SHFI as a dispersal resistance surface and applying a circuit theory based approach, we analyzed the effects of cumulative human impact on habitat connectivity in the different scenarios. For evaluating the relationship between habitat loss and connectivity, we applied graph theory-based equivalent connected area (ECA) index. Results show over 60% of the TMVS_Mich has high SHFI values, considerably lowering current flow for all species. Nevertheless, the effect on connectivity of human impact is higher for species with limited dispersal capacity (100–500 m). Our approach provides a new form of evaluating human impact on habitat connectivity that can be applied to different scales and landscapes. Furthermore, the approach is useful for guiding discussions and implementing future biodiversity conservation initiatives that promote landscape connectivity as an adaptive strategy for climate change.     

At what spatial scale(s) do mammals respond to urbanization?

Spatial scale is fundamental in understanding species–landscape relationships because species’ responses to landscape characteristics typically vary across scales. Nonetheless, such scales are often unidentified or unreliably predicted by theory. Many landscapes worldwide are urbanizing, yet the spatial scaling of species’ responses to urbanization is poorly understood. We investigated the spatial scaling of urbanization effects on a community of 15 mammal species using ~60 000 wildlife detections collected from a constellation of 207 camera traps across an extensive urban park system. We embedded a bivariate Gaussian kernel in hierarchical multi-species models to determine two scales of effect (a scale of maximal effect and a broader scale of cumulative landscape effect) for two biological responses (occupancy and site visit frequency) across two seasons (winter and summer) for each species. We then assessed whether scales of effect varied according to theoretical predictions associated with biological responses and species traits (body size and mobility). Scales of effect ranged from < 50 m to > 9000 m and varied among species, but not as predicted by theory. Species’ occupancy generally showed a weak response to urbanization and the scale of this effect was both highly uncertain and consistent across species. We did not detect any relationship between scales of effect and species’ body size or mobility, nor was there any evident pattern of scaling across biological response or seasons. These results imply that 1) urbanization effects on mammals manifest across a very broad spectrum of spatial scales, and 2) current theories that a priori predict the scale at which urbanization affects mammals may be of limited use within a given system. Overall, this study suggests that developing general theory regarding the scaling of species–landscape relationships requires additional empirical work conducted across multiple species, systems and timescales.     

Spatiotemporally explicit demographic modelling supports a joint effect of historical barriers to dispersal and contemporary landscape composition on structuring genomic variation in a red‐listed grasshopper

Inferring the processes underlying spatial patterns of genomic variation is fundamental to understand how organisms interact with landscape heterogeneity and to identify the factors determining species distributional shifts. Here, we use genomic data (restriction site‐associated DNA sequencing) to test biologically informed models representing historical and contemporary demographic scenarios of population connectivity for the Iberian cross‐backed grasshopper Dociostaurus hispanicus, a species with a narrow distribution that currently forms highly fragmented populations. All models incorporated biological aspects of the focal taxon that could hypothetically impact its geographical patterns of genomic variation, including (a) spatial configuration of impassable barriers to dispersal defined by topographic landscapes not occupied by the species; (b) distributional shifts resulting from the interaction between the species bioclimatic envelope and Pleistocene glacial cycles; and (c) contemporary distribution of suitable habitats after extensive land clearing for agriculture. Spatiotemporally explicit simulations under different scenarios considering these aspects and statistical evaluation of competing models within an Approximate Bayesian Computation framework supported spatial configuration of topographic barriers to dispersal and human‐driven habitat fragmentation as the main factors explaining the geographical distribution of genomic variation in the species, with no apparent impact of hypothetical distributional shifts linked to Pleistocene climatic oscillations. Collectively, this study supports that both historical (i.e., topographic barriers) and contemporary (i.e., anthropogenic habitat fragmentation) aspects of landscape composition have shaped major axes of genomic variation in the studied species and emphasizes the potential of model‐based approaches to gain insights into the temporal scale at which different processes impact the demography of natural populations.     

Landscape‐ and small‐scale determinants of grassland species diversity: direct and indirect influences

Species richness is influenced both by mechanisms occurring at landscape scales, such as habitat availability, and local‐scale processes, that are related to abiotic conditions and plant–plant interactions. However, it is rarely tested to what extent local species richness can be explained by the combined effect of factors measured at multiple spatial scales. In this study, we quantified the simultaneous influence of historical landscape‐scale factors (past human population density, and past habitat availability – an index combining area and connectivity) and small‐scale environmental conditions (shrub cover, and heterogeneity of light, soil depth, and other soil environmental variables) on plant species richness in dry calcareous grasslands (alvars). By applying structural equation modelling (SEM) we found that both landscape conditions and local environmental factors had significant direct and indirect (i.e. through the modification of another factor), effects on species richness. At the landscape scale, we found a direct positive influence of historical habitat availability on species richness, and indirect positive influence of past human population (via its effects on historical habitat availability). At small scales, we found a positive direct influence of light heterogeneity and shrub cover on species richness. Conversely, we found that small‐scale soil environmental heterogeneity, which was mainly determined by soil depth heterogeneity, had a negative effect on species richness. Our study indicates that patterns of species richness in alvar grasslands are positively influenced by the anthropogenic management regime that maintained the landscape habitat conditions in the past. However, the abandonment of management, leading to shrub invasion and increased competition for light resources also influenced species richness. In contrast to the positive heterogeneity–diversity relationship we found that soil heterogeneity reduced species richness. Environmental heterogeneity, occurring at the plant neighbourhood scale (i.e. centimetres), can increase the isolation among suitable soil patches and thus hinder the normal functioning of populations. The combination of previous knowledge of the system with new ecological theories facilitates disentangling how species richness responds to complex relationships among factors operating at multiple scales.     

Contrasting demographic and genetic estimates of dispersal in the endangered Coahuilan box turtle: a contemporary approach to conservation

The evolutionary viability of an endangered species depends upon gene flow among subpopulations and the degree of habitat patch connectivity. Contrasting population connectivity over ecological and evolutionary timescales may provide novel insight into what maintains genetic diversity within threatened species. We employed this integrative approach to evaluating dispersal in the critically endangered Coahuilan box turtle (Terrapene coahuila) that inhabits isolated wetlands in the desert‐spring ecosystem of Cuatro Ciénegas, Mexico. Recent wetland habitat loss has altered the spatial distribution and connectivity of habitat patches; and we therefore predicted that T. coahuila would exhibit limited movement relative to estimates of historic gene flow. To evaluate contemporary dispersal patterns, we employed mark–recapture techniques at both local (wetland complex) and regional (intercomplex) spatial scales. Gene flow estimates were obtained by surveying genetic variation at nine microsatellite loci in seven subpopulations located across the species’ geographical range. The mark–recapture results at the local spatial scale reveal frequent movement among wetlands that was unaffected by interwetland distance. At the regional spatial scale, dispersal events were relatively less frequent between wetland complexes. The complementary analysis of population genetic substructure indicates strong historic gene flow (global F_ST = 0.01). However, a relationship of genetic isolation by distance across the geographical range suggests that dispersal limitation exists at the regional scale. Our approach of contrasting direct and indirect estimates of dispersal at multiple spatial scales in T. coahuila conveys a sustainable evolutionary trajectory of the species pending preservation of threatened wetland habitats and a range‐wide network of corridors.     

Importance of estimating matrix quality for modeling species distribution in complex tropical landscapes: a test with Atlantic forest small mammals

Information to guide decision making is especially urgent in human dominated landscapes in the tropics, where urban and agricultural frontiers are still expanding in an unplanned manner. Nevertheless, most studies that have investigated the influence of landscape structure on species distribution have not considered the heterogeneity of altered habitats of the matrix, which is usually high in human dominated landscapes. Using the distribution of small mammals in forest remnants and in the four main altered habitats in an Atlantic forest landscape, we investigated 1) how explanatory power of models describing species distribution in forest remnants varies between landscape structure variables that do or do not incorporate matrix quality and 2) the importance of spatial scale for analyzing the influence of landscape structure. We used standardized sampling in remnants and altered habitats to generate two indices of habitat quality, corresponding to the abundance and to the occurrence of small mammals. For each remnant, we calculated habitat quantity and connectivity in different spatial scales, considering or not the quality of surrounding habitats. The incorporation of matrix quality increased model explanatory power across all spatial scales for half the species that occurred in the matrix, but only when taking into account the distance between habitat patches (connectivity). These connectivity models were also less affected by spatial scale than habitat quantity models. The few consistent responses to the variation in spatial scales indicate that despite their small size, small mammals perceive landscape features at large spatial scales. Matrix quality index corresponding to species occurrence presented a better or similar performance compared to that of species abundance. Results indicate the importance of the matrix for the dynamics of fragmented landscapes and suggest that relatively simple indices can improve our understanding of species distribution, and could be applied in modeling, monitoring and managing complex tropical landscapes.     

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。     

Scale-specific correlations between habitat heterogeneity and soil fauna diversity along a landscape structure gradient

Habitat heterogeneity contributes to the maintenance of diversity, but the extent that landscape-scale rather than local-scale heterogeneity influences the diversity of soil invertebrates—species with small range sizes—is less clear. Using a Scottish habitat heterogeneity gradient we correlated Collembola and lumbricid worm species richness and abundance with different elements (forest cover, habitat richness and patchiness) and qualities (plant species richness, soil variables) of habitat heterogeneity, at landscape (1 km²) and local (up to 200 m²) scales. Soil fauna assemblages showed considerable turnover in species composition along this habitat heterogeneity gradient. Soil fauna species richness and turnover was greatest in landscapes that were a mosaic of habitats. Soil fauna diversity was hump-shaped along a gradient of forest cover, peaking where there was a mixture of forest and open habitats in the landscape. Landscape-scale habitat richness was positively correlated with lumbricid diversity, while Collembola and lumbricid abundances were negatively and positively related to landscape spatial patchiness. Furthermore, soil fauna diversity was positively correlated with plant diversity, which in turn peaked in the sites that were a mosaic of forest and open habitat patches. There was less evidence that local-scale habitat variables (habitat richness, tree cover, plant species richness, litter cover, soil pH, depth of organic horizon) affected soil fauna diversity: Collembola diversity was independent of all these measures, while lumbricid diversity positively and negatively correlated with vascular plant species richness and tree canopy density. Landscape-scale habitat heterogeneity affects soil diversity regardless of taxon, while the influence of habitat heterogeneity at local scales is dependent on taxon identity, and hence ecological traits, e.g. body size. Landscape-scale habitat heterogeneity by providing different niches and refuges, together with passive dispersal and population patch dynamics, positively contributes to soil faunal diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Dispersal of fleshy-fruited species: a matter of spatial scale?

The processes associated with the dispersal of fleshy-fruited species have been an important focus of ecological research during the last two decades. These processes include fruit removal, seed rain, seed predation, seed bank dynamics, germination and establishment. Some of them interfere with the mutualistic interaction of frugivorous birds and fleshy-fruited plants. We might expect such interference to be most pronounced where the intensity of the different processes has a spatial distribution similar to that of the original seed shadow. The central theme of this review is that the main processes associated with dispersal and recruitment act at different spatial scales. To investigate this idea, about 140 publications on dispersal of fleshy-fruited species from 1980 to 2000 were screened for the spatial scaling of these processes. Microhabitat, habitat, landscape, region and biome were the five spatial scales most commonly used. However, the representation of the different scales was not fully balanced; large-scale studies were scarce and most publications considered only one scale.The review reveals some trends in scaling of the main processes of plant dispersal and recruitment. Seed dispersal by birds and seed predation by rodents are strongly determined at the habitat level, and several studies report negative results for contrasts between microhabitats. Germination and seedling establishment, on the other hand, appear to be mainly influenced by differences between microhabitats, though information on larger scales is scarce. Genetic differentiation and phenology of fruiting have mostly been investigated at the habitat, landscape and regional scale, whereas information on the abundance of frugivorous birds and patterns in plant distribution results are available across the full range of scales from the level of the microhabitat to the region and biome. Future research should be directed to the major gaps in our knowledge, i.e. regional and zonal comparisons of the processes associated with dispersal. They should also be more sensitive to scale issues and ideally should have a multi-scaled design.     

Spatial graphs highlight how multi-generational dispersal shapes landscape genetic patterns

Current approaches that compare spatial genetic structure of a given species and the dispersal of its mobile phase can detect a mismatch between both patterns mainly due to processes acting at different temporal scales. Genetic structure result from gene flow and other evolutionary and demographic processes over many generations, while dispersal predicted from the mobile phase often represents solely one generation on a single time-step. In this study, we present a spatial graph approach to landscape genetics that extends connectivity networks with a stepping-stone model to represent dispersal between suitable habitat patches over multiple generations. We illustrate the approach with the case of the striped red mullet Mullus surmuletus in the Mediterranean Sea. The genetic connectivity of M. surmuletus was not correlate with the estimated dispersal probability over one generation, but with the stepping-stone estimate of larval dispersal, revealing the temporal scale of connectivity across the Mediterranean Sea. Our results highlight the importance of considering multiple generations and different time scales when relating demographic and genetic connectivity. The spatial graph of genetic distances further untangles intra-population genetic structure revealing the Siculo-Tunisian Strait as an important corridor rather than a barrier for gene flow between the Western- and Eastern Mediterranean basins, and identifying Mediterranean islands as important stepping-stones for gene flow between continental populations. Our approach can be easily extended to other systems and environments.     

Beetle diversity in a matrix of old-growth boreal forest: influence of habitat heterogeneity at multiple scales

The relative contribution of compositional and structural heterogeneity on biodiversity is currently ambiguous because field studies generally integrate these two sources of habitat heterogeneity into a single index. We established the relationship between species richness of ground-dwelling and flying beetles and compositional and structural attributes of forest heterogeneity. The relationship was evaluated at two spatial scales: the scale of forest stand, corresponding to an 11.3  m radius, and the scale of landscape, corresponding to either a 400 or 800  m radius. Seventy stands were sampled in the matrix of old-growth boreal forest of the North Shore region of Québec, Canada, during the summers of 2004 and 2005. A total of 133 ground-dwelling beetle species (range: 4–42 species per site) were captured in the pitfall traps and 251 flying species (range 16–58 species per site) in flight-interception traps. We found that the most relevant type of heterogeneity to explain variations in species richness and the significance of landscape scale information varied between groups of beetles. Compositional heterogeneity (i.e. the number of species of forest trees and shrubs) at the stand scale best predicted species richness in ground-dwelling beetles. On the other hand, it was the combined influence of structural and compositional habitat heterogeneity at stand and landscape scales that best explained richness patterns in flying beetles. Our study outlines the significance of considering multiple types and spatial scales of habitat heterogeneity when describing patterns of species richness.     

Effect of the landscape matrix on gene flow in a coastal amphibian metapopulation

Functional connectivity is crucial for the persistence of a metapopulation, because migration among subpopulations enables recolonization and counteracts genetic drift, which is especially important in small subpopulations. We studied the degree and drivers of connectivity among occupied patches of a coastal dune metapopulation of the Natterjack Toad (Epidalea calamita Laurenti), on the basis of microsatellite variation. As spatial landscape heterogeneity is expected to influence dispersal and genetic structure, we analyzed which landscape features affect functional connectivity and to what extent. Sixty different landscape resistance scenarios as well as the isolation-by-distance model were compared using two landscape genetics approaches. We identified three subpopulations with unidirectional levels of gene flow among the two most geographically separated subpopulations, while inferred gene flow into the geographically intermediate subpopulation was limited. Urbanization and vegetation height negatively affected connectivity. Low estimates of genetic diversity and effective population size indicate that conservation measures in the smallest and most isolated subpopulation are required.     

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。     

Functional spatial scale of community composition change in response to windthrow disturbance in a deciduous temperate forest

Community dynamics in local habitats are affected by landscape characteristics such as the area and connectivity of surrounding habitats at a functional spatial scale where the community responds to landscape structure. However, the functional spatial scale at which community composition is affected by landscape structure has never been explored. We assessed the functional spatial scales of composition change in birds and in three types of arthropod communities (canopy, forest-floor and flying ones) with regard to landscape heterogeneity resulting from a large typhoon in a temperate forest of Japan. We examined the effects of tree-fall disturbance on the communities at various spatial scales, with special attention to compositional evenness. The spatial scale of the best-fitting model, which was selected from models fitted to the disturbance area at stepwise spatial scales, was interpreted as the community-specific functional spatial scale. The composition of all communities studied was all significantly dependent on gap area. The functional spatial scale was highest in birds (370 m in radius), intermediate in flying arthropods (90 m) and lowest in canopy and forest-floor arthropods (10 m). This result may reflect typical dispersal ability and the spatial range of resource use in the community. Compositional changes in each community were consistent with theory regarding traits and responses of component taxa, although the enhancement of evenness was observed only in the arthropod communities. These results imply that management and reserve selection based on functional spatial scales can be effective in the conservation of biodiversity and ecosystem services at the community level.