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。     

Cattle select African savanna termite mound patches less when sharing habitat with wild herbivores

African savanna termite mounds function as nutrient‐rich foraging hotspots for different herbivore species, but little is known about their effects on the interaction between domestic and wild herbivores. Understanding such effects is important for better management of these herbivore guilds in landscapes where they share habitats. Working in a central Kenyan savanna ecosystem, we compared selection of termite mound patches by cattle between areas cattle accessed exclusively and areas they shared with wild herbivores. Termite mound selection index was significantly lower in the shared areas than in areas cattle accessed exclusively. Furthermore, cattle used termite mounds in proportion to their availability when they were the only herbivores present, but used them less than their availability when they shared foraging areas with wild herbivores. These patterns were associated with reduced herbage cover on termite mounds in the shared foraging areas, partly indicating that cattle and wild herbivores compete for termite mound forage. However, reduced selection of termite mound patches was also reinforced by higher leafiness of Brachiaria lachnantha (the principal cattle diet forage species) off termite mounds in shared than in unshared areas. Taken together, these findings suggest that during wet periods, cattle can overcome competition for termite mounds by taking advantage of wildlife‐mediated increased forage leafiness in the matrix surrounding termite mounds. However, this advantage is likely to dissipate during dry periods when forage conditions deteriorate across the landscape and the importance of termite mounds as nutrient hotspots increases for both cattle and wild herbivores. Therefore, we suggest that those managing for both livestock production and wildlife conservation in such savanna landscapes should adopt grazing strategies that could lessen competition for forage on termite mounds, such as strategically decreasing stock numbers during dry periods.     

Allometric Convergence in Savanna Trees and Implications for the Use of Plant Scaling Models in Variable Ecosystems

Theoretical models of allometric scaling provide frameworks for understanding and predicting how and why the morphology and function of organisms vary with scale. It remains unclear, however, if the predictions of ‘universal’ scaling models for vascular plants hold across diverse species in variable environments. Phenomena such as competition and disturbance may drive allometric scaling relationships away from theoretical predictions based on an optimized tree. Here, we use a hierarchical Bayesian approach to calculate tree-specific, species-specific, and ‘global’ (i.e. interspecific) scaling exponents for several allometric relationships using tree- and branch-level data harvested from three savanna sites across a rainfall gradient in Mali, West Africa. We use these exponents to provide a rigorous test of three plant scaling models (Metabolic Scaling Theory (MST), Geometric Similarity, and Stress Similarity) in savanna systems. For the allometric relationships we evaluated (diameter vs. length, aboveground mass, stem mass, and leaf mass) the empirically calculated exponents broadly overlapped among species from diverse environments, except for the scaling exponents for length, which increased with tree cover and density. When we compare empirical scaling exponents to the theoretical predictions from the three models we find MST predictions are most consistent with our observed allometries. In those situations where observations are inconsistent with MST we find that departure from theory corresponds with expected tradeoffs related to disturbance and competitive interactions. We hypothesize savanna trees have greater length-scaling exponents than predicted by MST due to an evolutionary tradeoff between fire escape and optimization of mechanical stability and internal resource transport. Future research on the drivers of systematic allometric variation could reconcile the differences between observed scaling relationships in variable ecosystems and those predicted by ideal models such as MST.     

Consistent spatial scaling of high-severity wildfire can inform expected future patterns of burn severity

Increasing wildfire activity in forests worldwide has driven urgency in understanding current and future fire regimes. Spatial patterns of area burned at high severity strongly shape forest resilience and constitute a key dimension of fire regimes, yet remain difficult to predict. To characterize the range of burn severity patterns expected within contemporary fire regimes, we quantified scaling relationships relating fire size to patterns of burn severity. Using 1615 fires occurring across the Northwest United States between 1985 and 2020, we evaluated scaling relationships within fire regimes and tested whether relationships vary across space and time. Patterns of high-severity fire demonstrate consistent scaling behaviour; as fire size increases, high-severity patches consistently increase in size and homogeneity. Scaling relationships did not differ substantially across space or time at the scales considered here, suggesting that as fire-size distributions potentially shift, stationarity in patch-size scaling can be used to infer future patterns of burn severity.     

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.     

Termite mounds differ in their importance for herbivores across savanna types,seasons and spatial scales

Herbivores do not forage uniformly across landscapes, but select for patches of higher nutrition and lower predation risk. Macrotermes mounds contain higher concentrations of soil nutrients and support grasses of higher nutritional value than the surrounding savanna matrix, attracting mammalian grazers that preferentially forage on termite mound vegetation. However, little is known about the spatial extent of such termite influence on grazing patterns and how it might differ in time and space. We measured grazing intensity in three African savanna types differing in rainfall and foliar nutrients and predicted that the functional importance of mounds for grazing herbivores would increase as the difference in foliar nutrient levels between mound and savanna matrix grasses increases and the mounds become more attractive. We expected this to occur in nutrient‐poor areas and during the dry season when savanna matrix grass nutrient levels are lower. Tuft use and grass N and P content were measured along transects away from termite mounds, enabling calculation of the spatial extent of termite influence on mammalian grazing. Using termite mound densities estimated from airborne light detection and ranging (LiDAR), we further upscaled field‐based results to determine the percentage of the landscape influenced by termite activity. Grasses in close proximity to termite mounds were preferentially grazed at all sites and in both seasons, but the strength of mound influence varied between savanna types and seasons. In the wet season, mounds had a relatively larger effect on grazers at the landscape scale in the nutrient‐poor, wetter savanna, whereas in the dry season the pattern was reversed with more of the landscape influenced at the nutrient‐rich, driest site. Our results reveal that termite mounds enhance the value of savanna landscapes for herbivores, but that their functional importance varies across savanna types and seasons.     

北京东灵山土壤动物多样性海拔格局的尺度推绎规律

识别群落内部各类群多样性格局的复杂性是生态学家面临的挑战,而尺度推绎规律是揭示复杂生态结构的有效途径之一。本研究利用多重分形的方法探索了不同海拔土壤动物多样性格局的尺度推绎规律,对比分析了凋落物层和土壤层之间多重分形谱的差异。结果表明: 与之前对植物群落的分析结果相似,土壤动物多样性尺度推绎规律同样具有幂律特征,如丰富度、Shannon多样性指数和Simpson多样性的倒数。凋落物层和土壤层中不同相对多度土壤动物的丰富度也具有幂律尺度推绎规律。凋落物层和土壤层中土壤动物多样性格局都具有多重分形特征,但凋落物层中多样性的分形结构比土壤层更均匀,且两层间优势类群与稀有类群的尺度推绎特征在多重分形谱上不同格局。幂律尺度推绎规律对于有着较高丰富度与多度的土壤动物同样存在,从而有助于揭示地下生物多样性的空间分布机制。     

Ecological application of wavelet analysis in the scaling of spatial distribution patterns of Ceratoides ewersmanniana

Ecological experiments are usually conducted on small scales, but the ecological and environmental issues are usually on large scales. Hence, there is a clear need of scaling. Namely, when we deal with patterns and processes on larger scales, a special connection needs to be established on the small scales that we are familiar with. Here we presented a wavelet analysis method that could build relationships between spatial distribution patterns on different scales. With this method, we also studied how spatial heterogeneity and distribution patterns changed with the scale. We investigated the distribution and the habitat of C. ewersmanniana in two plots (200 m × 200 m; the distance between these two plots is 15 km) at Mosuowan desert. The results demonstrated that spatial heterogeneity and distribution patterns were incorporated into larger scales when the wavelet scale varied from one (5 m) to four (20 m). However, if the wavelet scale was above five (25 m), the spatial distribution patterns varied placidly, the oscillation frequency of landforms stabilized at 110 m, and the dynamic quantity period of C. ewersmanniana stabilized at 115–125 m. We also identified signal mutation points with wavelet analysis and verified the heterogeneity degree of local space with position variance. We found that position variance decomposed the distribution patterns on large scales into small sampling plots, and the position with the largest variance also had the strongest heterogeneity. In a word, the wavelet analysis method could scale-up spatial distribution patterns and habitat heterogeneity. With this method and other methods derived from this one, such as wavelet scale, wavelet variance, position variance and extremely direct-viewing graphs, wavelet analysis could be widely applied in solving the scaling problem in ecological and environmental studies.     

生态学中的尺度及尺度转换方法

尺度作为生态学的重要范式,已经引起了广泛重视,但对尺度问题的研究还不够成熟.尺度具有多维性特点,即功能尺度、空间尺度、时间尺度等,但生态学研究的重点是空间和时间尺度.并且时空尺度还具有复杂性、变异性特征.尺度研究的根本目的在于通过适宜的空间和时间尺度来揭示和把握复杂的生态学规律.为此,科学有效的尺度选择和尺度转换方法不可或缺.常见的尺度转换方法有图示法、回归分析、变异函数、自相关分析、谱分析、分形和小波变换,同时遥感和地理信息系统技术在尺度研究中也发挥着重要作用.结合实例对上述方法进行了分析和论述,认为各种方法都有其内在的优势和不足,新方法的引入和应用对于尺度转换方法体系的充实和完善非常重要.有关尺度的研究将进一步加强,研究的重点是尺度变异性、不同尺度间的相互作用机制以及尺度转换方法等.     

Broad‐scale patterns in plant diversity vary between land uses in a variegated temperate Australian agricultural landscape

Plant diversity is threatened in many agricultural landscapes. Our understanding of patterns of plant diversity in these landscapes is mainly based on small‐scale (<1000 m²) observations of species richness. However, such observations are insufficient for detecting the spatial heterogeneity of vegetation composition. In a case‐study farm on the North‐West Slopes of New South Wales, Australia, we observed species richness at four scales (quadrat, patch, land use and landscape) across five land uses (grazed and ungrazed woodlands, native pastures, roadsides and crops). We applied two landscape ecological models to assess the contribution of these land uses to landscape species richness: (i) additive partitioning of diversity at multiple spatial scales, and (ii) a measure of habitat specificity – the effective number of species that a patch contributes to landscape species richness. Native pastures had less variation between patches than grazed and ungrazed woodlands, and hence were less species‐rich at the landscape scale, despite having similar richness to woodlands at the quadrat and patch scale. Habitat specificity was significantly higher for ungrazed woodland patches than all other land uses. Our results showed that in this landscape, ungrazed woodland patches had a higher contribution than the grazed land uses to landscape species richness. These results have implications for the conservation management of this landscape, and highlighted the need for greater consensus on the influence of different land uses on landscape patterns of plant diversity.     

Evidence for indigenous selection and distribution of the shea tree, Vitellaria paradoxa, and its potential significance to prevailing parkland savanna tree patterns in sub-Saharan Africa north of the equator

Aim Woody vegetation patterns in African savannas north of the equator are closely connected to human presence, but the distinctions between natural and anthropogenic landscapes have not been clear to many observers. Criteria for identifying savanna landscapes on a continuum of intensity of anthropic impact are explored. Methods A key savanna tree species, Vitellaria paradoxa (Sapotaceae), was used as model for evaluating anthropic impact. Fruits harvested from tree populations across the species range were analysed for variation in traits valued by indigenous peoples. A simple selection index was used to scale tree populations from a hypothetical wild state to a hypothetical domesticated state. Index values were compared with trait values along climate zone gradients and evaluated in the context of indigenous savanna management practices and historical species distribution reports. Results Trait values such as fruit size and shape, pulp sweetness, and kernel fat content show a significant influence of temperature and rainfall. At the same time, the mean values of groups of traits vary perpendicular to the general climatic zone gradient. Selection index values between Vitellaria populations vary up to sixfold, with highest values in central Burkina Faso. Comparison of present day Vitellaria distribution with historical range limits show range expansion by human migration. Main conclusions The prevalence of major economic tree species in the savannas of Africa north of the equator is a strong indicator of human involvement in tree dispersal. This conclusion is supported by paleobotanical evidence and by recent Vitellaria range expansion as a result of human migration. The presence of high mean values of several Vitellaria fruit traits in central Burkina Faso suggests that selection for desired characteristics has occurred. The impact of indigenous savanna peoples on woody species composition and spatial distribution is probably much greater than usually thought and is the result of a deliberate strategy of altering the landscape to provide needed human resources.     

Motivation and Benefits of Complex Systems Approaches in Ecology

Studies of complex systems in other disciplines provide models and analytical strategies for understanding ecosystems and landscapes. The emphasis is on invariant properties, particularly processes that create scaling relations over wide ranges of scale, both in time and space. Translations between levels of ecological organization may be accomplished by succinct characterizations of processes that operate at fine scales, followed by renormalization group analysis to reveal patterns at broad scales. The self-organized patterns found in simple ecosystem, landscape, and forest-fire models may be explained as feedback between the system state and control parameters. Critical phenomena and phase transitions are expected in open, dissipative systems where long-range correlations defy predictions based on average population densities, a concept that becomes irrelevant as nonstationary conditions prevail. Thus, complexity theory for open systems relates to the ecology of self-entailing ecosystems that function as their own environments and thereby create constraints through emergence. Received: 14 April 1998; accepted 26 May 1998     

Resource use by the dryad butterfly is scale-dependent

The factors shaping the ways in which animals use resources are a key element of conservation biology, but ecological studies on resource use typically neglect to consider how the study’s spatial scale may have affected the outcomes. We used the dryad butterfly, inhabiting xerothermic grassland and wet meadow, to test for differences in its resource use at two scales–habitat patch and landscape. Based on records of plant species composition from random points within four habitat patches and from points in 53 patches along surveyed transects, we compared the microhabitat preferences of the butterfly on the patch scale, and species occurrence and abundance patterns on the landscape scale. We distinguished four main groups of factors related to vegetation structure which affected the butterfly’s resource use—factors having similar effects on both spatial scales, factors operating primarily on one of the scales considered, factors relevant only on a single spatial scale, and factors operating on both scales but with effects differing between the two habitat types. We suggest that invertebrates may respond on two spatial levels or on only one, and conclude that larger-scale studies can meet the challenges of a sophisticated metapopulation approach and can give insight into the habitat characteristics affecting the persistence of species in landscapes. We stress the value of large-scale studies on species’ habitat preferences when planning conservation strategies, while pointing out that small-scale studies provide useful information about species ecology and behavior, especially if conducted in multiple habitats.     

Continuum or discrete patch landscape models for savanna birds? Towards a pluralistic approach

Conceptualising landscapes as a mosaic of discrete habitat patches is fundamental to landscape ecology, metapopulation theory and conservation biology. An emerging question in ecology is: when is the discrete patch model more appropriate than alternative and conceptually appealing models such as the continuum model? There is limited empirical testing of the utility of alternative landscape models compared to the discrete patch model for a range of species. In this paper, we constructed three alternative sets of models for testing the effect of landscape structure on diversity and abundance of a suite of woodland birds in a savanna landscape of northern Australia: the null model (only site‐scale habitat variables, landscape context not important), the continuum model, and the discrete patch model. We utilised high‐spatial resolution satellite images to quantify spatial gradients in tree cover density (the continuum model), and to then aggregate the fine‐scale heterogeneity in tree cover into discrete patches of trees, with grass cover forming the “matrix” (the discrete patch‐model). We then evaluated the relative importance of the alternative models using generalised linear models and an information theoretic approach. We found that the importance of the models varied among species, with no single model dominant. Species that move between open grassy areas and woody shelter responded well to the continuum model, reflecting the importance of gradients in density of forage (grasses) and cover (trees), while the discrete model performed best for species that forage in all vegetation strata, and nest predominantly in dense woody vegetation. This finding supports a pluralistic approach, highlighting the need for adopting and testing more than one landscape model in savanna landscapes, and in other landscapes that do not have a well defined patch structure.     

Evaluating empirical scaling relations of pattern metrics with simulated landscapes

Landscape patterns demonstrate scale-dependent properties that have been parsimoniously described by empirical scaling functions. These functions, derived from multiple-scale analysis of real landscapes, are evaluated here for their generality and robustness via a series of simulated landscapes with known landscape patterns. A factorial design was used to generate these landscapes, varying the number of classes, class abundance distribution, and patch dispersion. The results confirm that the three types of scaling relations were both general and robust. Type I metrics were predictable with simple scaling functions (e.g. power laws or linear functions); Type II metrics showed stair-case like response patterns and were essentially not predictable; Type III metrics exhibited erratic response patterns that were unpredictable in most cases. However, significant differences were found between real and simulated landscapes when landscape extent was increased. Systematic changes in grain size show that the predictability of scaling relations increases with the number of classes, the evenness of class abundance distribution, and the aggregation of patch dispersion. However, random patch dispersion seemed to enhance the predictability of scaling relations when changing spatial extent.     

An improved neutral landscape model for recreating real landscapes and generating landscape series for spatial ecological simulations

Many studies have assessed the effect of landscape patterns on spatial ecological processes by simulating these processes in computer‐generated landscapes with varying composition and configuration. To generate such landscapes, various neutral landscape models have been developed. However, the limited set of landscape‐level pattern variables included in these models is often inadequate to generate landscapes that reflect real landscapes. In order to achieve more flexibility and variability in the generated landscapes patterns, a more complete set of class‐ and patch‐level pattern variables should be implemented in these models. These enhancements have been implemented in Landscape Generator (LG), which is a software that uses optimization algorithms to generate landscapes that match user‐defined target values. Developed for participatory spatial planning at small scale, we enhanced the usability of LG and demonstrated how it can be used for larger scale ecological studies. First, we used LG to recreate landscape patterns from a real landscape (i.e., a mountainous region in Switzerland). Second, we generated landscape series with incrementally changing pattern variables, which could be used in ecological simulation studies. We found that LG was able to recreate landscape patterns that approximate those of real landscapes. Furthermore, we successfully generated landscape series that would not have been possible with traditional neutral landscape models. LG is a promising novel approach for generating neutral landscapes and enables testing of new hypotheses regarding the influence of landscape patterns on ecological processes. LG is freely available online.     

Local contagion and regional compression: habitat selection drives spatially explicit,multiscale dynamics of colonisation in experimental metacommunities

Habitat selection, including oviposition site choice, is an important driver of community assembly in freshwater systems. Factors determining patch quality are assessed by many colonising organisms and affect colonisation rates, spatial distribution and community structure. For many species, the presence/absence of predators is the most important factor affecting female oviposition decisions. However, individual habitat patches exist in complex landscapes linked by processes of dispersal and colonisation, and spatial distribution of factors such as predators has potential effects beyond individual patches. Perceived patch quality and resulting colonisation rates depend both on risk conditions within a given patch and on spatial context. Here we experimentally confirm the role of one context‐dependent processes, spatial contagion, functioning at the local scale, and provide the first example of another context‐dependent process, habitat compression, functioning at the regional scale. Both processes affect colonisation rates and patterns of spatial distribution in naturally colonised experimental metacommunities.     

Points, patches, and regions: scaling soil biogeochemical patterns in an urbanized arid ecosystem

Cities are rapidly growing throughout the world and are altering biologic processes in many regions, with global consequences. Urbanization in the Phoenix, USA metropolitan region has dramatically altered regional ecosystem patterns, but little is known about how these changes have influenced soil organic matter, total nitrogen, and the distribution of nitrogen stable isotopes. Because urban development is a phenomenon occurring at multiple scales, ecological consequences of urbanization will likely differ between individual patches and the entire metropolitan region. To investigate such changes we conducted spatially explicit surveys including three dominant land‐use types in this region: native desert, agriculture, and mesic residential. These data were combined for analysis with previously collected samples from a synoptic regional survey. A landscape scaling approach was implemented to compare the dependence of soil variability on the sampled extent and the uncertainty associated with scaling from points to patches, land‐use types, and the Phoenix metropolitan region. The multiple‐scale analysis of soil properties showed that variation in total soil nitrogen, soil organic matter, and δ⁵N content of soils differed between patch and regional scales. The majority of variation in the urbanized patch types was exhibited between patches while for the native desert the majority of variation was observed within individual patches. These differences show the impact of urbanization on the scaling relations of ecosystem components. Overall, urbanization in this region appears to have increased soil organic matter by 44%, total nitrogen by 48%, and has elevated δ¹⁵N by 21%.     

State‐space modeling reveals habitat perception of a small terrestrial mammal in a fragmented landscape

相似文献   

Bistability,Spatial Interaction,and the Distribution of Tropical Forests and Savannas

Recent work has indicated that tropical forest and savanna can be alternative stable states under a range of climatic conditions. However, dynamical systems theory suggests that in case of strong spatial interactions between patches of forest and savanna, a boundary between both states is only possible at conditions in which forest and savanna are equally stable, called the ‘Maxwell point.’ Frequency distributions of MODIS tree-cover data at 250 m resolution were used to estimate such Maxwell points with respect to the amount and seasonality of rainfall in both South America and Africa. We tested on a 0.5° scale whether there is a larger probability of local coexistence of forests and savannas near the estimated Maxwell points. Maxwell points for South America and Africa were estimated at 1760 and 1580 mm mean annual precipitation and at Markham’s Seasonality Index values of 50 and 24 %. Although the probability of local coexistence was indeed highest around these Maxwell points, local coexistence was not limited to the Maxwell points. We conclude that critical transitions between forest and savanna may occur when climatic changes exceed a critical value. However, we also conclude that spatial interactions between patches of forest and savanna may reduce the hysteresis that can be observed in isolated patches, causing more predictable forest-savanna boundaries than continental-scale analyses of tree cover indicate. This effect could be less pronounced in Africa than in South America, where the forest-savanna boundary is substantially affected by rainfall seasonality.