What Is the Role of Ecology in Understanding Ecosystem Resilience ?

I the article that begins on page 489 of this issue of BioScience, Andrew J. Kerkhoff and Brian J. Enquist make an important synthesis from two hitherto largely separate fields of biological inquiry. One is the conceptually well-established, although not uncontentious, area of allometric scaling laws. The other is the less-established area known as resilience. The concept of resilience, as applied to an ecosystem, is loosely defined as the ability of the system to maintain its function when faced with novel disturbance. The concept is related to stability, but with its focus on maintenance of function and novel disturbance, resilience uniquely encompasses aspects of society’s reliance on ecosystem services and increasing anthropogenic change. Thus, scientists from many different backgrounds recognize the societal importance of resilience and are intellectually intrigued by resilience concepts. While ecologists generally seem to share this perspective, many are also frustrated by the diversity of definitions for resilience and the complex role of ecology in this area. These issues arise partially from the history of resilience research, which has been performed by very different groups of scientists, and partially from the inherent difficulties of integrating interdisciplinary research. Indeed, resilience is still an evolving concept. Since resilience research is not driven solely by ecology, the challenge for ecologists is to understand resilience perspectives from multiple fields in order to better integrate traditional ecology with modern perspectives and research on resilience. A brief history of resilience Holling (1973) and Pimm (1984) used a mathematical approach to illustrate resilience concepts in ecology. Parallel development of resilience concepts by these two researchers and others resulted in two dominant paradigms. Engineering resilience (Pimm 1984) is the length of time that a system takes to return to equilibrium following perturbation (i.e., disturbance). Ecological resilience (Holling 1973) is the amount of perturbation a system can withstand before it moves into a different basin of attraction or stability domain; there are several variations on this general theme, including the idea of system “flips” (Scheffer et al. 2001), which may make it hard to return a system to a stability domain that it has left. These paradigms of resilience are now seen primarily as analogies, at least partially because of the difficulties in interpreting these mathematical models clearly in an empirical, ecological context. Much of the current research on resilience relevant to ecosystems has broadened to include multiple fields and moved to a complex adaptive systems (CAS) approach. Complex adaptive systems are composed of diverse components with localized interactions and a selection process (Levin 1998). Scientists from many fields, interested in the response of CAS to novel disturbance, define resilience as maintenance of system function when confronted with some perturbation that exceeds the historical range of variation. Two main types of research outside of traditional ecology comprise much current resilience research. First, researchers taking a CAS approach in engineering, computer science, physics, and other fields look for commonalities among their systems to elucidate which characteristics of CAS enhance resilience (often termed “robustness” in the CAS context). These general principles can be applied to ecosystems. The robustness projects carried out at the Santa Fe Institute (www.santafe.edu) are a clear example of this type of research. Second, researchers interested in sustainable development and adaptive management use a CAS approach in studying coupled social–ecological systems (SES). The SES approach differs from that of traditional ecology by placing equal weight on the human and ecological dimensions of ecosystem function and maintenance. The Resilience Alliance (www.resalliance.org) has spearheaded much of this SES research. Differences in approaches among traditional ecology, SES, and CAS in nonecological fields, though rarely articulated, may dissuade ecologists from working with the resilience concept. In part, these differences have developed because researchers from different fields highlight the facets of resilience that are most important for their primary field, thus contributing to the vagueness of the resilience concept as perceived by ecologists. In addition, researchers comparing CAS across fields or working within SES frequently are problem driven and use a case study approach. In contrast, traditional ecology is dominated by hypothetico–deductive reasoning and experimentation. These differences make it difficult for ecoloWhat Is the Role of Ecology in Understanding Ecosystem Resilience?