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.     

Spatial modelling of succession-disturbance dynamics in forest ecosystems: Concepts and examples

Over the last few decades it has become increasingly obvious that disturbance, whether natural or anthropogenic in origin, is ubiquitous in ecosystems. Disturbance-related processes are now considered to be important determinants of the composition, structure and function of ecological systems. However, because disturbance and succession processes occur across a wide range of spatio-temporal scales their empirical investigation is difficult. To counter these difficulties much use has been made of spatial modelling to explore the response of ecological systems to disturbance(s) occurring at spatial scales from the individual to the landscape and above, and temporal scales from minutes to centuries. Here we consider such models by contrasting two alternative motivations for their development and use: prediction and exploration, with a focus on forested ecosystems. We consider the two approaches to be complementary rather than competing. Predictive modelling aims to combine knowledge (understanding and data) with the goal of predicting system dynamics; conversely, exploratory models focus on developing understanding in systems where uncertainty is high. Examples of exploratory modelling include model-based explorations of generic issues of criticality in ecological systems, whereas predictive models tend to be more heavily data-driven (e.g. species distribution models). By considering predictive and exploratory modelling alongside each other, we aim to illustrate the range of methods used to model succession and disturbance dynamics and the challenges involved in the model-building and evaluation processes in this arena.     

Interactions Across Spatial Scales among Forest Dieback,Fire, and Erosion in Northern New Mexico Landscapes

Abstract Ecosystem patterns and disturbance processes at one spatial scale often interact with processes at another scale, and the result of such cross-scale interactions can be nonlinear dynamics with thresholds. Examples of cross-scale pattern-process relationships and interactions among forest dieback, fire, and erosion are illustrated from northern New Mexico (USA) landscapes, where long-term studies have recently documented all of these disturbance processes. For example, environmental stress, operating on individual trees, can cause tree death that is amplified by insect mortality agents to propagate to patch and then landscape or even regional-scale forest dieback. Severe drought and unusual warmth in the southwestern USA since the late 1990s apparently exceeded species-specific physiological thresholds for multiple tree species, resulting in substantial vegetation mortality across millions of hectares of woodlands and forests in recent years. Predictions of forest dieback across spatial scales are constrained by uncertainties associated with: limited knowledge of species-specific physiological thresholds; individual and site-specific variation in these mortality thresholds; and positive feedback loops between rapidly-responding insect herbivore populations and their stressed plant hosts, sometimes resulting in nonlinear “pest” outbreak dynamics. Fire behavior also exhibits nonlinearities across spatial scales, illustrated by changes in historic fire regimes where patch-scale grazing disturbance led to regional-scale collapse of surface fire activity and subsequent recent increases in the scale of extreme fire events in New Mexico. Vegetation dieback interacts with fire activity by modifying fuel amounts and configurations at multiple spatial scales. Runoff and erosion processes are also subject to scale-dependent threshold behaviors, exemplified by ecohydrological work in semiarid New Mexico watersheds showing how declines in ground surface cover lead to non-linear increases in bare patch connectivity and thereby accelerated runoff and erosion at hillslope and watershed scales. Vegetation dieback, grazing, and fire can change land surface properties and cross-scale hydrologic connectivities, directly altering ecohydrological patterns of runoff and erosion. The interactions among disturbance processes across spatial scales can be key drivers in ecosystem dynamics, as illustrated by these studies of recent landscape changes in northern New Mexico. To better anticipate and mitigate accelerating human impacts to the planetary ecosystem at all spatial scales, improvements are needed in our conceptual and quantitative understanding of cross-scale interactions among disturbance processes.     

缀块性和缀块动态：Ⅲ.生态与进化效应

自然界各种等级系统都普遍存在时间和空间的缀块性。它反映了系统内部或系统间的时空异质性,影响着生态学过程。不同缀块的大小、形状、边际性质以及缀块间的距离等空间分布特征构成缀块性的差异,并控制生态过程的速率。某一空间特有的缀块性也是生态过程的结果。把这种因果关系在时间轴上     

Spatial analyses of intertidal assemblages on sheltered rocky shores

Abstract Understanding processes in complex assemblages depends on good understanding of spatial and temporal patterns of structure at various spatial scales. There has been little quantitative information about spatial patterns and natural temporal changes in intertidal assemblages on sheltered rocky shores in temperate Australia. Natural changes and responses to anthropogenic disturbances in these habitats cannot be accurately measured and assessed without quantitative data on patterns of natural variability in space and through time. This paper describes some suitable quantitative methods for examining spatial and temporal patterns of diversity and abundances of highshore, midshore and lowshore intertidal assemblages and the important component species for a number of shores in a bay that has not been severely altered by human disturbance. Despite a diverse flora and fauna on these shores, the midshore and lowshore assemblages on sheltered shores were characterized by a few species which were also the most important in discriminating among assemblages on a shore and, for each assemblage, among different shores. The same set of species was also important for measuring small-scale patchiness within each assemblage (i.e. between replicate sites on a shore). Therefore, these data provide a rationale for selecting species that are useful for measuring differences and changes in abundance among places and times at different scales and, hence, can be used in the more complex sampling designs necessary to detect environmental impacts. There was considerable spatial variability in all assemblages and all species (or taxa) examined at scales of metres, tens of metres and kilometres. There were no clear seasonal trends for most measures, with as much or more variability at intervals of 3 months as from year to year. Most interactions between spatial and temporal measures were at the smallest scale, with different sites on the same shore generally showing different changes from time to time. The cause(s) of this apparently idiosyncratic variability1 were not examined, but some potential causes are discussed. These data are appropriate for testing hypotheses about the applicability of these findings to other relatively undisturbed sheltered shores, about effects of different anthropogenic disturbances on sheltered intertidal assemblages and to test hypotheses about differences in intertidal assemblages on sheltered versus wave-exposed shores.     

Effects of wildfire, rainfall and region on desert lizard assemblages: the importance of multi-scale processes

Vertebrate populations are influenced by environmental processes that operate at a range of spatial and temporal scales. Wildfire is a disturbance that can affect vertebrate populations across large spatial scales, although vertebrate responses are frequently influenced by processes operating at smaller spatial scales such as topography, interspecific interactions and regional history. Here, we investigate the effects of a broad-scale wildfire on lizard assemblages in a desert region. We predicted that a rainfall gradient within the region affected by the wildfire would influence lizard responses to the fire by encouraging post-fire succession to proceed more rapidly in high-rainfall areas, and would be enabled in turn by more rapid vegetation recovery. To test our prediction, we censused lizards, measured rainfall, undertook vegetation surveys and sampled invertebrate abundance across burnt and unburnt habitat ecotones within three regional areas situated along a gradient of long-term annual rainfall. Lizard diversity was not affected by fire or region and lizard abundance was influenced only by region. Lizard assemblage composition was also only influenced by region, but this did not relate to differences in rainfall or habitat as we had predicted. Regional differences in lizard assemblages related instead to food availability. The observed differences also likely reflected regional differences in the strength of biotic interactions with predators and changes in land use. Our study shows that assemblage responses to a disturbance were not uniform within a large desert region and instead were influenced by other environmental processes operating simultaneously at multiple temporal and spatial scales.     

The future in and of criticality assessments

In recent literature, the concept of criticality aspires to provide a multifaceted risk assessment of resource supply shortage. However, most existing methodologies for the criticality assessment of raw materials are restricted to a fixed temporal and spatial reference system. They provide a snapshot in time of the equilibrium between supply and demand/economic importance and do not account for temporal changes of their indicators. The static character of criticality assessments limits the use of criticality methodologies to short‐term policy making of raw materials. In the current paper, we argue for an enhancement of the criticality framework to account for three key dynamic characteristics, namely changes of social, technical, and economic features; consideration of the spatial dimension in site‐specific assessments; and impact of changing governance frameworks. We illustrate how these issues were addressed in studies outside of the field of criticality and identify the dynamic parameters that influence resource supply and demand based on a review of studies that belong to the general field of resource supply and demand. The parameters are grouped in seven categories: extraction, social, economic, technical, policy, market dynamics, and environmental. We explore how these parameters were considered in the reviewed studies and propose ways and specific examples of addressing the dynamic effects in the criticality indicators. Furthermore, we discuss the current work on future scenarios to provide reference points for indicator benchmarks. The insights and guidelines derived from the review and our recommendations for future research set the foundations for an enhanced dynamic and site‐specific criticality assessment framework.     

Multiscale responses of soil stability and invasive plants to removal of non-native grazers from an arid conservation reserve

Disturbances and ecosystem recovery from disturbance both involve numerous processes that operate on multiple spatial and temporal scales. Few studies have investigated how gradients of disturbance intensity and ecosystem responses are distributed across multiple spatial resolutions and also how this relationship changes through time during recovery. We investigated how cover of non-native species and soil-aggregate stability (a measure of vulnerability to erosion by water) in surface and subsurface soils varied spatially during grazing by burros and cattle and whether patterns in these variables changed after grazer removal from Mojave National Preserve, California, USA. We compared distance from water and number of ungulate defecations — metrics of longer-term and recent grazing intensity, respectively, — as predictors of our response variables. We used information-theoretic analyses to compare hierarchical linear models that accounted for important covariates and allowed for interannual variation in the disturbance–response relationship at local and landscape scales. Soil stability was greater under perennial vegetation than in bare interspaces, and surface soil stability decreased with increasing numbers of ungulate defecations. Stability of surface samples was more affected by time since removal of grazers than was stability of subsurface samples, and subsurface soil stability in bare spaces was not related to grazing intensity, time since removal, or any of our other predictors. In the high rainfall year (2003) after cattle had been removed for 1–2 years, cover of all non-native plants averaged nine times higher than in the low-rainfall year (2002). Given the heterogeneity in distribution of large-herbivore impacts that we observed at several resolutions, hierarchical analyses provided a more complete understanding of the spatial and temporal complexities of disturbance and recovery processes in arid ecosystems.     

Pattern formation in a spatial plant-wrack model with tide effect on the wrack

Spatial patterns are a subfield of spatial ecology, and these patterns modify the temporal dynamics and stability properties of population densities at a range of spatial scales. Localized ecological interactions can generate striking large-scale spatial patterns in ecosystems through spatial self-organization. Possible mechanisms include oscillating consumer–resource interactions, localized disturbance–recovery processes, and scale-dependent feedback. However, in this paper, our main aim is to study the effect of tide on the pattern formation of a spatial plant-wrack model. We discuss the changes of the wavelength, wave speed, and the conditions of the spatial pattern formation, according to the dispersion relation formula. Both the mathematical analysis and numerical simulations reveal that the tide has great influence on the spatial pattern. More specifically, typical traveling spatial patterns can be obtained. Our obtained results are consistent with the previous observation that wracks exhibit traveling patterns, which is useful to help us better understand the dynamics of the real ecosystems.     

Closing the larval loop: linking larval ecology to the population dynamics of marine benthic invertebrates

The majority of marine benthic invertebrates exhibit a complex life cycle that includes separate planktonic larval, and bottom-dwelling juvenile and adult phases. To understand and predict changes in the spatial and temporal distributions, abundances, population growth rate, and population structure of a species with such a complex life cycle, it is necessary to understand the relative importance of the physical, chemical and biological properties and processes that affect individuals within both the planktonic and benthic phases. To accomplish this goal, it is necessary to study both phases within a common, quantitative framework defined in terms of some common currency. This can be done efficiently through construction and evaluation of a population dynamics model that describes the complete life cycle.

Two forms that such a model might assume are reviewed: a stage-based, population matrix model, and a model that specifies discrete stages of the population, on the bottom and in the water column, in terms of simultaneous differential equations that may be solved in both space and time. Terms to be incorporated in each type of model can be formulated to describe the critical properties and processes that can affect populations within each stage of the life cycle. For both types of model it is shown how this might be accomplished using an idealized balanomorph barnacle as an example species. The critical properties and processes that affect the planktonic and benthic phases are reviewed. For larvae, these include benthic adult fecundity and fertilization success, growth and larval stage duration, mortality, larval behavior, dispersal by currents and turbulence, and larval settlement. It is possible to predict or estimate empirically all of the key terms that should be built into the larval and benthic components of the model. Thus, the challenge of formulating and evaluating a full life cycle model is achievable. Development and evaluation of such a model will be challenging because of the diverse processes which must be considered, and because of the disparities in the spatial and temporal scales appropriate to the benthic and planktonic larval phases. In evaluating model predictions it is critical that sampling schemes be matched to the spatial and temporal scales of model resolution.     

Multifractal growth in periphyton communities

Periphyton is an aquatic community composed by algae, bacteria, fungi, and other microorganisms that can develop a complex architecture comparable to tropical forests. We analyzed the spatial pattern of a periphyton community along a succession developed in experimental tanks. Our aim was to identify regularities that may help us to explain the patchiness of this community. Therefore, we estimated the spatial pattern of periphyton biomass using a non‐destructive image analysis technique to obtain a temporal series of the spatial distribution. These were analyzed using multifractal techniques. Multifractals are analogous to fractals but they look at the geometry of quantities instead of the geometry of pattern. To use these techniques the object of study must show scale invariance and then can be characterized by a spectra of fractal dimensions. Self‐organization describes the evolution of complex structures that emerge spontaneously driven internally by variations of the system itself. The spatial distribution of biomass showed scale invariance at all stages of succession and as the periphyton developed in a homogeneous landscape, in a demonstration of self‐organized behavior. Self‐organization to a critical state (SOC) is presented in the complex systems literature as a general explanation for scale invariance in nature. SOC requires a mechanism where the history of past events in a place influence the actual dynamics, this was termed ecological memory. The scale invariance was found from the very beginning of the succession thus self‐organized criticality is a very improbable explanation for the pattern because there would be not enough time for the build‐up of ecological memory. Positive interactions between algae and bacteria, and the existence of different spatial scales of colonization and growth are the likely causes of this pattern. Our work is a demonstration of how large scale patterns emerge from local biotic interactions.     

Wildfire and forest harvest disturbances in the boreal forest leave different long-lasting spatial signatures

Aims

Natural disturbances leave long-term legacies that vary among landscapes and ecosystem types, and which become integral parts of successional processes at a given location. As humans change land use, not only are immediate post-disturbance patterns altered, but the processes of recovery themselves are likely altered by the disturbance. We assessed whether short-term effects on soil and vegetation that distinguish wildfire from forest harvest persist over 60 years after disturbance in boreal black spruce forests, or post-disturbance processes of recovery promote convergence of the two disturbance types.

Methods

Using semi-variograms and Principal Coordinates of Neighbour Matrices, we formulated precise, a priori spatial hypotheses to discriminate spatial signatures following wildfire and forest harvest both over the short- (16–18 years) and long-term (62–98 years).

Results

Both over the short- and the long-term, wildfire generated a wide spectrum of responses in soil and vegetation properties at different spatial scales, while logging produced simpler patterns corresponding to the regular linear pattern of harvest trails and to pre-disturbance ericaceous shrub patches that persist between trails.

Conclusions

Disturbance by harvest simplified spatial patterns associated with soil and vegetation properties compared to patterns associated with natural disturbance by fire. The observed differences in these patterns between disturbance types persist for over 60 years. Ecological management strategies inspired by natural disturbances should aim to increase the complexity of patterns associated with harvest interventions.     

Implications of recurrent disturbance for genetic diversity

Exploring interactions between ecological disturbance, species’ abundances and community composition provides critical insights for ecological dynamics. While disturbance is also potentially an important driver of landscape genetic patterns, the mechanisms by which these patterns may arise by selective and neutral processes are not well‐understood. We used simulation to evaluate the relative importance of disturbance regime components, and their interaction with demographic and dispersal processes, on the distribution of genetic diversity across landscapes. We investigated genetic impacts of variation in key components of disturbance regimes and spatial patterns that are likely to respond to climate change and land management, including disturbance size, frequency, and severity. The influence of disturbance was mediated by dispersal distance and, to a limited extent, by birth rate. Nevertheless, all three disturbance regime components strongly influenced spatial and temporal patterns of genetic diversity within subpopulations, and were associated with changes in genetic structure. Furthermore, disturbance‐induced changes in temporal population dynamics and the spatial distribution of populations across the landscape resulted in disrupted isolation by distance patterns among populations. Our results show that forecast changes in disturbance regimes have the potential to cause major changes to the distribution of genetic diversity within and among populations. We highlight likely scenarios under which future changes to disturbance size, severity, or frequency will have the strongest impacts on population genetic patterns. In addition, our results have implications for the inference of biological processes from genetic data, because the effects of dispersal on genetic patterns were strongly mediated by disturbance regimes.     

Ecological Thresholds: The Key to Successful Environmental Management or an Important Concept with No Practical Application?

An ecological threshold is the point at which there is an abrupt change in an ecosystem quality, property or phenomenon, or where small changes in an environmental driver produce large responses in the ecosystem. Analysis of thresholds is complicated by nonlinear dynamics and by multiple factor controls that operate at diverse spatial and temporal scales. These complexities have challenged the use and utility of threshold concepts in environmental management despite great concern about preventing dramatic state changes in valued ecosystems, the need for determining critical pollutant loads and the ubiquity of other threshold-based environmental problems. In this paper we define the scope of the thresholds concept in ecological science and discuss methods for identifying and investigating thresholds using a variety of examples from terrestrial and aquatic environments, at ecosystem, landscape and regional scales. We end with a discussion of key research needs in this area.     

Local climate mediates spatial and temporal variation in carabid beetle communities in three forests in Mount Odaesan,Korea

1. Global environmental change can dramatically alter the composition of floral and faunal communities, and elucidating the mechanisms underlying this process is important for predicting its outcomes. Studies on global climate change have mostly focused on statistical summaries within wide spatial and temporal scales; less attention has been paid to variability in microclimates at narrower spatial and temporal scales. 2. The microclimate is the suite of climatic conditions measured in a local area. Environmental variables at the microclimatic scale can be critical for the ecology of organisms inhabiting each area. The effect of spatial and temporal changes in the microclimate on the ecology of carabid beetle communities in three sites on Mount Odaesan, Korea was examined. 3. Carabid beetle communities and quantified site‐specific environmental factors from measurements of air temperature, air humidity, light intensity and soil temperature over 5 years (2010–2015) were surveyed. 4. It was found that microclimatic variables and the patterns of temporal changes in carabid beetle communities differed between the three sites within the single mountain system. Microclimatic variables influencing temporal changes in beetle communities also differed between the sites. Therefore, it is suggested that variation in local microclimates affects spatial and temporal variation in carabid beetle communities at a local scale. 5. The present results demonstrate the importance of regular surveys of communities at local scales. Such surveys are expected to reveal an additional fraction of variation in communities and underlying processes that have been overlooked in studies of global community patterns and change.     

土壤呼吸作用时空动态变化及其影响机制研究与展望

测定不同陆地生态系统土壤呼吸速率及其时空波动, 阐明其影响因子, 对于全球碳素平衡预算和全球变化潜在效应估计是最为基本的数据。然而, 有关土壤呼吸作用变异性及其影响因素的知识仍存在局限性, 一些关键的过程和机制还有待阐明。该文综述了近年来土壤呼吸作用时空动态规律、影响机制和模拟方面的研究进展, 指出环境因子和生物因子共同驱动着土壤呼吸作用的时间动态变化; 土壤呼吸作用在不同时间尺度上还具有明显的空间异质性, 这主要是植被覆盖、根系分布、主要的环境因素和土壤特性空间分布的异质性造成的。生物因子是影响土壤呼吸作用时空动态变化的主要因素之一。然而, 目前所使用的土壤呼吸作用经验模型通常利用土壤温度、土壤湿度或者两者的交互作用模拟土壤呼吸作用动态变化, 但没有考虑生物因子的影响, 这可能会导致明显的偏差和错误。因此, 为了精确估算土壤呼吸作用, 必须解决土壤呼吸作用小尺度上的空间变异性; 加强不同时间尺度上生物要素对土壤呼吸作用动态变化的影响研究; 除了气候因子外, 土壤呼吸作用经验模型应该纳入生物因子等其它影响因素作为变量, 用以提高模型模拟的正确性和准确性。     

Simple temporal models for ecological systems with complex spatial patterns

Spatial patterns are ubiquitous in nature. Because these patterns modify the temporal dynamics and stability properties of population densities at a range of spatial scales, their effects must be incorporated in temporal ecological models that do not represent space explicitly. We demonstrate a connection between a simple parameterization of spatial effects and the geometry of clusters in an individual‐based predator–prey model that is both nonlinear and stochastic. Specifically we show that clusters exhibit a power‐law scaling of perimeter to area with an exponent close to unity. In systems with a high degree of patchiness, similar power‐law scalings can provide a basis for applying simple temporal models that assume well‐mixed conditions.     

Natural,potential and actual vegetation in North America

The potential natural vegetation (PNV) concept has parallel applications in Europe and North America. Paleoecological studies in parts of North America provide records of vegetation patterns and dynamics under little or no human disturbance. Something resembling PNV emerges at millennial temporal scales and at regional to subcontinental spatial scales. However, at finer spatial and temporal scales, actual vegetation often displays properties of inertia, contingency and hysteresis, most frequently because of climatic variability across multiple timescales and the episodic nature of disturbance and establishment. Thus, in the absence of human disturbance, the actual vegetation that develops at a site may not resemble a particular PNV ideal, but could instead represent one of any number of potential outcomes constrained by historically contingent processes. PNV may best be viewed as an artificial construct, with utility in some settings. Its utility may diminish and even be detrimental in a rapidly changing environment.     

Collective predator evasion: Putting the criticality hypothesis to the test

According to the criticality hypothesis, collective biological systems should operate in a special parameter region, close to so-called critical points, where the collective behavior undergoes a qualitative change between different dynamical regimes. Critical systems exhibit unique properties, which may benefit collective information processing such as maximal responsiveness to external stimuli. Besides neuronal and gene-regulatory networks, recent empirical data suggests that also animal collectives may be examples of self-organized critical systems. However, open questions about self-organization mechanisms in animal groups remain: Evolutionary adaptation towards a group-level optimum (group-level selection), implicitly assumed in the “criticality hypothesis”, appears in general not reasonable for fission-fusion groups composed of non-related individuals. Furthermore, previous theoretical work relies on non-spatial models, which ignore potentially important self-organization and spatial sorting effects. Using a generic, spatially-explicit model of schooling prey being attacked by a predator, we show first that schools operating at criticality perform best. However, this is not due to optimal response of the prey to the predator, as suggested by the “criticality hypothesis”, but rather due to the spatial structure of the prey school at criticality. Secondly, by investigating individual-level evolution, we show that strong spatial self-sorting effects at the critical point lead to strong selection gradients, and make it an evolutionary unstable state. Our results demonstrate the decisive role of spatio-temporal phenomena in collective behavior, and that individual-level selection is in general not a viable mechanism for self-tuning of unrelated animal groups towards criticality.     

缀块性和缀块动态:Ⅱ.描述与分析

在生态学中,“缀块”(patch)的概念早已在植被群落和海洋生态系统中浮游动植物空间分布研究中广泛应用。A.S.Watte早在1947年就强调缀块在植物群落结构分析中的重要性。实际上,缀块存在于地球的任何地方。森林可