Grazing Management, Resilience, and the Dynamics of a Fire-driven Rangeland System

We developed a stylized mathematical model to explore the effects of physical, ecological, and economic factors on the resilience of a managed fire-driven rangeland system. Depending on grazing pressure, the model exhibits one of three distinct configurations: a fire-dominated, grazing-dominated, or shrub-dominated rangeland system. Transaction costs and costs due to shrub invasion, via their effect on grazing decisions, strongly influence which stable configuration is occupied. This, in turn, determines the resilience of the rangeland system. These results are used to establish conditions under which management for profit is consistent with the maintenance of resilience. Received 2 January 2001; accepted 11 June 2001.     

The Use of Discontinuities and Functional Groups to Assess Relative Resilience in Complex Systems

It is evident when the resilience of a system has been exceeded and the system qualitatively changed. However, it is not clear how to measure resilience in a system prior to the demonstration that the capacity for resilient response has been exceeded. We argue that self-organizing human and natural systems are structured by a relatively small set of processes operating across scales in time and space. These structuring processes should generate a discontinuous distribution of structures and frequencies, where discontinuities mark the transition from one scale to another. Resilience is not driven by the identity of elements of a system, but rather by the functions those elements provide, and their distribution within and across scales. A self-organizing system that is resilient should maintain patterns of function within and across scales despite the turnover of specific elements (for example, species, cities). However, the loss of functions, or a decrease in functional representation at certain scales will decrease system resilience. It follows that some distributions of function should be more resilient than others. We propose that the determination of discontinuities, and the quantification of function both within and across scales, produce relative measures of resilience in ecological and other systems. We describe a set of methods to assess the relative resilience of a system based upon the determination of discontinuities and the quantification of the distribution of functions in relation to those discontinuities.     

Multiple Scales and the Maintenance of Biodiversity

The problem of multiple scales permeates the study of ecological process and pattern, uniting aspects of space, time, and organizational complexity. In particular, it supports the maintenance of biological diversity, allowing for the magnification of underlying patterns of variation in the physical environment to create many resources from few, through the evolutionary diversification of species' niches and life histories. Specialization to particular stages of a successional gradient facilitates coexistence of multiple types in the presence of uncorrelated local disturbances, such as gap formation, which reinitiate successional sequences. Superimposed upon such successional dynamics are the effects of multiple stable states and multiple successional pathways (Levin 1976), which increase diversity even more. Multiple stable states more generally raise the possibility of sudden flips of systems from one stable configuration to another, and with such changes may come huge changes in the biotic composition. On larger spatial scales and longer time scales, these flips may become correlated, resulting in the transformation of the landscape, or may result in sustained spatiotemporal mosaics of states. Instability at the local level may lead to the maintenance of biodiversity on broader scales. Finally, the ultimate scale mismatch involves that between the dynamics of natural systems and the cultural dynamics of human societies. Our ability to live sustainably in a global commons is dependent upon adjusting normative behavior, and tightening feedback loops more generally, so that individual actions serve the common good. Received 11 November 1999; accepted 18 February 2000     

Mobile Link Organisms and Ecosystem Functioning: Implications for Ecosystem Resilience and Management

Current natural resource management seldom takes the ecosystem functions performed by organisms that move between systems into consideration. Organisms that actively move in the landscape and connect habitats in space and time are here termed “mobile links.” They are essential components in the dynamics of ecosystem development and ecosystem resilience (that is, buffer capacity and opportunity for reorganization) that provide ecological memory (that is, sources for reorganization after disturbance). We investigated the effects of such mobile links on ecosystem functions in aquatic as well as terrestrial environments. We identify three main functional categories: resource, genetic, and process linkers and suggest that the diversity within functional groups of mobile links is a central component of ecosystem resilience. As the planet becomes increasingly dominated by humans, the magnitude, frequency, timing, spatial extent, rate, and quality of such organism-mediated linkages are being altered. We argue that global environmental change can lead to (a) the decline of essential links in functional groups providing pollination, seed dispersal, and pest control; (b) the linking of previously disconnected areas, for example, the spread of vector-borne diseases and invasive species; and (c) the potential for existing links to become carriers of toxic substances, such as persistent organic compounds. We conclude that knowledge of interspatial exchange via mobile links needs to be incorporated into management and policy-making decisions in order to maintain ecosystem resilience and hence secure the capacity of ecosystems to supply the goods and services essential to society. Received 23 April 2001; accepted 17 June 2002.     

Spatial Resilience of Coral Reefs

There have been several earlier studies that addressed the influence of natural disturbance regimes on coral reefs. Humans alter natural disturbance regimes, introduce new stressors, and modify background conditions of reefs. We focus on how coral reef ecosystems relate to disturbance in an increasingly human-dominated environment. The concept of ecosystem resilience—that is, the capacity of complex systems with multiple stable states to absorb disturbance, reorganize, and adapt to change—is central in this context. Instead of focusing on the recovery of certain species and populations within disturbed sites of individual reefs, we address spatial resilience—that is, the dynamic capacity of a reef matrix to reorganize and maintain ecosystem function following disturbance. The interplay between disturbance and ecosystem resilience is highlighted. We begin the identification of spatial sources of resilience in dynamic seascapes and exemplify and discuss the relation between “ecological memory” (biological legacies, mobile link species, and support areas) and functional diversity for seascape resilience. Managing for resilience in dynamic seascapes not only enhances the likelihood of conserving coral reefs, it also provides insurance to society by sustaining essential ecosystem services. Received 25 February 2000; accepted 31 January 2001.     

Understanding the Complexity of Economic, Ecological, and Social Systems

Hierarchies and adaptive cycles comprise the basis of ecosystems and social-ecological systems across scales. Together they form a panarchy. The panarchy describes how a healthy system can invent and experiment, benefiting from inventions that create opportunity while being kept safe from those that destabilize because of their nature or excessive exuberance. Each level is allowed to operate at its own pace, protected from above by slower, larger levels but invigorated from below by faster, smaller cycles of innovation. The whole panarchy is therefore both creative and conserving. The interactions between cycles in a panarchy combine learning with continuity. An analysis of this process helps to clarify the meaning of “sustainable development.” Sustainability is the capacity to create, test, and maintain adaptive capability. Development is the process of creating, testing, and maintaining opportunity. The phrase that combines the two, “sustainable development,” thus refers to the goal of fostering adaptive capabilities and creating opportunities. It is therefore not an oxymoron but a term that describes a logical partnership. Received 7 March 2001; accepted 16 March 2001.     

Original Articles: Plant Attribute Diversity, Resilience, and Ecosystem Function: The Nature and Significance of Dominant and Minor Species

This study tested an hypothesis concerning patterns in species abundance in ecological communities. Why do the majority of species occur in low abundance, with just a few making up the bulk of the biomass? We propose that many of the minor species are analogues of the dominants in terms of the ecosystem functions they perform, but differ in terms of their capabilities to respond to environmental stresses and disturbance. They thereby confer resilience on the community with respect to ecosystem function. Under changing conditions, ecosystem function is maintained when dominants decline or are lost because functionally equivalent minor species are able to substitute for them. We have tested this hypothesis with respect to ecosystem functions relating to global change. In particular, we identified five plant functional attributes—height, biomass, specific leaf area, longevity, and leaf litter quality—that determine carbon and water fluxes. We assigned values for these functional attributes to each of the graminoid species in a lightly grazed site and in a heavily grazed site in an Australian rangeland. Our resilience proposition was cast in the form of three specific hypotheses in relation to expected similarities and dissimilarities between dominant and minor species, within and between sites. Functional similarity—or ecological distance—was determined as the euclidean distance between species in functional attribute space. The analyses provide evidence in support of the resilience hypothesis. Specifically, within the lightly grazed community, dominant species were functionally more dissimilar to one another, and functionally similar species more widely separated in abundance rank, than would be expected on the basis of average ecological distances in the community. Between communities, depending on the test used, two of three, or three of four minor species in the lightly grazed community that were predicted to increase in the heavily grazed community did in fact do so. Although there has been emphasis on the importance of functional diversity in supporting the flow of ecosystem goods and services, the evidence from this study indicates that functional similarity (between dominant and minor species, and among minor species) may be equally important in ensuring persistence (resilience) of ecosystem function under changing environmental conditions.     

Contagious Disturbance, Ecological Memory, and the Emergence of Landscape Pattern

Landscapes are strongly shaped by the degree of interaction between pattern and process. This paper examines how ecological memory, the degree to which an ecological process is shaped by its past modifications of a landscape, influences landscape dynamics. I use a simulation model to examine how ecological memory shapes the landscape dynamics produced by the interaction of vegetative regrowth and fire. The model illustrated that increased ecological memory increased the strength and spatial extent of landscape pattern. The extent of these changes depended upon the relative rates of vegetative recovery and fire initiation. When ecological memory is strong, landscape pattern is persistent; pattern tends to be maintained rather than destroyed by fire. The generality of the simulation model suggests that these results may also apply to disturbance processes other than fire. The existence of ecological memory in ecosystems may allow processes to produce ecological pattern that can entrain other ecosystem variables. The methods presented in this paper to analyze pattern in model ecosystems could be used to detect such pattern in actual ecosystems. Received 14 November 2000; accepted 21 September 2001.     

Functional Richness and Relative Resilience of Bird Communities in Regions with Different Land Use Intensities

Abstract Empirical estimates of the function and resilience of communities under different management regimes can provide valuable information for sustainable natural resource management, but such estimates are scarce to date. We quantified the functional richness and relative resilience of bird communities inhabiting five regions in southeastern Australia that represented different management regimes. First, we show that functional richness and relative resilience were reduced at species-poor sites in all regions. Second, we show that bird communities in agricultural regions had fewer body mass groups and fewer functional groups than expected by chance. This suggests that both the function and the resilience of bird communities in agricultural regions were reduced. The likely mechanisms for the observed loss of function and relative resilience are: (1) the simplification of landscape texture resulting in selective extinction of certain body mass groups; and (2) the selective extinction of certain functional groups that are particularly sensitive to intensive land use. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ecological stability,evolutionary stability and the ESS maximum principle

Summary Since the fitness of each individual organism in a biological community may be affected by the strategies of all other individuals in the community, the essential element of a game exists. This game is an evolutionary game where the individual organisms (players) inherit their strategies from continuous play of the game through time. Here, the strategies are assumed to be constants associated with certain adaptive parameters (such as sunlight conversion efficiency for plants or body length in animals) in a set of differential equations which describe the population dynamics of the community. By means of natural selection, these parameters will evolve to a set of strategy values that natural selection, by itself, can no longer modify, i.e. an evolutionarily stable strategy (ESS). For a given class of models, it is possible to predict the outcome of this evolutionary process by determining ESSs using an ESS maximum principle. However, heretofore, the proof of this principle has been based on a limited set of conditions. Herein, we generalize the proof by removing certain restrictions and use instead the concept of an ecological stable equilibrium (ESE). Individuals in a biological community will be at an ESE if fixing the strategies used by the individuals results in stable population densities subject to perturbations in those densities. We present both necessary and sufficient conditions for an ESE to exist and then use the ESE concept to provide a very simple proof of the ESS maximum principle (which is a necessary condition for an ESS). A simple example is used to illustrate the difference between a strategy that maximizes fitness and one that satisfies the ESS maximum principle. In general they are different. We also look for ESEs in Lotka—Volterra competition and use the maximum principle to determine when an ESE will be an ESS. Finally, we examine the applicability of these ideas to matrix games.     

Resilience indicators: prospects and limitations for early warnings of regime shifts

In the vicinity of tipping points—or more precisely bifurcation points—ecosystems recover slowly from small perturbations. Such slowness may be interpreted as a sign of low resilience in the sense that the ecosystem could easily be tipped through a critical transition into a contrasting state. Indicators of this phenomenon of ‘critical slowing down (CSD)’ include a rise in temporal correlation and variance. Such indicators of CSD can provide an early warning signal of a nearby tipping point. Or, they may offer a possibility to rank reefs, lakes or other ecosystems according to their resilience. The fact that CSD may happen across a wide range of complex ecosystems close to tipping points implies a powerful generality. However, indicators of CSD are not manifested in all cases where regime shifts occur. This is because not all regime shifts are associated with tipping points. Here, we review the exploding literature about this issue to provide guidance on what to expect and what not to expect when it comes to the CSD-based early warning signals for critical transitions.     

植物功能多样性与功能群研究进展

综述了植物功能多样性与功能群研究的最新进展。介绍了植物功能群的定义及植物功能群的划分方法。在功能多样性与生态系统资源动态关系方面．抽样效应和生态位互补效应用来解释植物多样性在生态系统资源动态中的作用。功能多样性与生态系统的稳定性间的关系可以用生态冗余或生态保险概念来解释,这两个概念是一个问题的两个侧面,是多样性与生态系统功能争论的焦点。     

Functional Group Change within and across Scales following Invasions and Extinctions in the Everglades Ecosystem

Cross-scale resilience theory predicts that the combination of functional diversity within scales and functional redundancy across scales is an important attribute of ecosystems because it helps these systems resist minor ecological disruptions and regenerate after major disturbances such as hurricanes and fire. Using the vertebrate fauna of south Florida, we quantified how the loss of native species and invasion by nonnatives may alter functional group richness within and across scales. We found that despite large changes in species composition due to potential extinctions and successful invasions by nonnative species, functional group richness will not change significantly within scales, there will not be any significant loss of overall redundancy of ecology function across scales, and overall body mass pattern will not undergo substantial change. However, the types of functions performed will change, and this change may have profound effects on not only the Everglades ecosystem but on the entire landscape of south Florida. Received 14 November 2000; accepted 20 December 2001.     

Compounded Perturbations Yield Ecological Surprises

All species have evolved in the presence of disturbance, and thus are in a sense matched to the recurrence pattern of the perturbations. Consequently, disturbances within the typical range, even at the extreme of that range as defined by large, infrequent disturbances (LIDs), usually result in little long-term change to the system's fundamental character. We argue that more serious ecological consequences result from compounded perturbations within the normative recovery time of the community in question. We consider both physically based disturbance (for example, storm, volcanic eruption, and forest fire) and biologically based disturbance of populations, such as overharvesting, invasion, and disease, and their interactions. Dispersal capability and measures of generation time or age to first reproduction of the species of interest seem to be the important metrics for scaling the size and frequency of disturbances among different types of ecosystems. We develop six scenarios that describe communities that have been subjected to multiple perturbations, either simultaneously or at a rate faster than the rate of recovery, and appear to have entered new domains or “ecological surprises.” In some cases, three or more disturbances seem to have been required to initiate the changed state. We argue that in a world of ever-more-pervasive anthropogenic impacts on natural communities coupled with the increasing certainty of global change, compounded perturbations and ecological surprises will become more common. Understanding these ecological synergisms will be basic to environmental management decisions of the 21st century. Received 14 July 1998; accepted 18 September 1998.     

Ten Years of the Relative Risk Model and Regional Scale Ecological Risk Assessment

It has been 10 years since the publication of the relative risk model (RRM) for regional scale ecological risk assessment. The approach has since been used successfully for a variety of freshwater, marine, and terrestrial environments in North America, South America, and Australia. During this period the types of stressors have been expanded to include more than contaminants. Invasive species, habitat loss, stream alteration and blockage, temperature, change in land use, and climate have been incorporated into the assessments. Major developments in the RRM have included the extensive use of geographical information systems, uncertainty analysis using Monte Carlo techniques, and its application to retrospective assessments to determine causation. The future uses of the RRM include assessments for forestry and conservation management, an increasing use in invasive species evaluation, and in sustainability. Developments in risk communication, the use of Bayesian approaches, and in uncertainty analyses are on the horizon.     

Ecosystems and the Biosphere as Complex Adaptive Systems

Ecosystems are prototypical examples of complex adaptive systems, in which patterns at higher levels emerge from localized interactions and selection processes acting at lower levels. An essential aspect of such systems is nonlinearity, leading to historical dependency and multiple possible outcomes of dynamics. Given this, it is essential to determine the degree to which system features are determined by environmental conditions, and the degree to which they are the result of self-organization. Furthermore, given the multiple levels at which dynamics become apparent and at which selection can act, central issues relate to how evolution shapes ecosystems properties, and whether ecosystems become buffered to changes (more resilient) over their ecological and evolutionary development or proceed to critical states and the edge of chaos. Received 14 April 1998; accepted 26 May 1998.     

Structure and Dynamics in an Urban Landscape: Toward a Multiscale View

Ecosystems and city systems often form hierarchically structured landscapes whose spatial pattern is scale dependent. While trends in the upper tail of national city-size distributions leave the impression that fractal-scaling laws such as Zipf's law or the rank-size rule truly represent the essence of the system, the linearity depicted at aggregate scale actually obscures variation and discontinuity in the urban size-density function, including multimodalities evident in regional data sets. Tracing individual city trajectories through these hierarchical patterns reveals structural resilience at macroscopic scale, the punctuated growth of individual cities of differing sizes, the persistence and self-reinforcing character of spatial agglomeration, and a general need for further empirical investigation of the relationship between city size and growth. It also raises questions for future exploration, including the meaning of persistent departures from the power laws of traditional urban systems theory. Interpretation of such departures in the context of questions of jurisdictional scale in environmental management and “smart growth” policy adds a practical dimension to the research agenda. Received 14 November 2000; accepted 5 September 2001.     

A Procedure for Ecological Tiered Assessment of Risks (PETAR)

Several procedures for Ecological Risk Assessment (ERA) have been suggested. The use of these existing procedures often relies on availability of existing data and/or on large resources for acquisition of new ones. This paper presents a three-tiered procedure for retrospective evaluation of risks adapted to limited resources and scarce background information of relevance for risk assessments, such as in developing countries. The tiers require successively more detailed investigations. The approach assures that resources available for site-specific investigations are directed towards well-formulated questions raised during previous stages of the assessment. The first tier, the preliminary assessment, is a qualitative evaluation of existing information on anthropogenic stressors, sources of stressors and expected ecological effects. The second tier is a regional risk assessment; a semi-quantitative evaluation of ecological risks, over large geographical areas, which results in a ranking of sources and stressors having the greatest potential for ecological impact and ranking of subareas inside the study area more likely to be impacted. The final tier is a site-specific and quantitative risk assessment, at a smaller scale and requiring more resources, that incorporates methodologies for establishing causality between exposure to multiple stressors and effects on specific endpoints of ecological and societal relevance.