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Santa Fe Institute. July 21, 2005 1:42 p.m.

Ch1-Pascual page 3

From Small to Large Ecological Networks in a
Dynamic World
Mercedes Pascual
Jennifer A. Dunne

Food webs are one of the most useful, and challenging, objects of study in ecology.
These networks of predator-prey interactions, conjured in Darwin’s image of a
“tangled bank,” provide a paradigmatic example of complex adaptive systems.
While it is deceptively easy to throw together simplified caricatures of feeding
relationships among a few taxa as can be seen in many basic ecology text books,
it is much harder to create detailed descriptions that portray a full range of
diversity of species in an ecosystem and the complexity of interactions among
them (fig. 1). Difficult to sample, difficult to describe, and difficult to model,
food webs are nevertheless of central practical and theoretical importance. The
interactions between species on different trophic (feeding) levels underlie the
flow of energy and biomass in ecosystems and mediate species’ responses to
natural and unnatural perturbations such as habitat loss. Understanding the
ecology and mathematics of food webs, and more broadly, ecological networks,
is central to understanding the fate of biodiversity and ecosystems in response
to perturbations.

Ecological Networks: Linking Structure to Dynamics in Food Webs,
edited by Mercedes Pascual and Jennifer A. Dunne, Oxford University Press. 3

Santa Fe Institute. July 21, 2005 1:42 p.m. Ch1-Pascual page 4

4 From Small to Large Ecological Networks in a Dynamic World

FIGURE 1 A detailed food web of Little Rock Lake, WI, with 997 feedings links among
92 taxa (Martinez 1991). Ecosystems are complex networks consisting of many species
that interact in many different ways with each other and their environment. Food webs
focus on the feeding relationships among co-occurring species in a particular habitat.
In this image, each node represents a “trophic species” that may be a biological species,
a group of species, a life-history stage of a species, or organic matter such as detritus
(Briand and Cohen 1984). Each trophic species in a food web is functionally distinct;
i.e., has a set of predators and prey that differ, even if only by one predator or prey
item, from those of other trophic species. This image was produced using FoodWeb3D
software written by R. J. Williams and provided by the Pacific Ecoinformatics and
Computational Ecology Lab www.foodwebs.org.

Research on ecological networks is also important for understanding the
consequences of biodiversity itself for ecosystem function. Much theoretical and
empirical food-web research, as well as other ecological research, has oriented
itself around various notions of stability (Box 1). Ultimately, stability properties
matter to the functioning of ecosystems and to the all-to-often unacknowledged
services they provide to humans (Box 2). While a large body of research addresses
the relationship between biodiversity and ecosystem functions such as primary
productivity, the ecological networks considered in those studies are generally
restricted to one or two trophic levels, and usually focus on interactions among
competitors for one or a few resources (Kinzig et al. 2002; but see Montoya et al.
2003). The importance of extending studies of biodiversity-ecosystem function
to include multiple trophic levels and complex predator-prey interactions has
been recently emphasized (Dobson et al. in press; Worm and Duffy 2003). A
better understanding of dynamics in large, complex networks is an important
intermediate step in addressing how the functioning of ecosystems is influenced
by structural properties of the underlying network.

• Resilience: Resilience is closely related to the concept of local asymptotic stability and measures how fast a stable system returns to equilibrium following a perturbation away from it. Chapter 7). Resilience is quantified as the absolute magnitude of the largest real part of any of the eigenvalues of the community matrix (Pimm and Lawton 1977. It specifies whether perturbations can initially grow and be amplified. Chen and Cohen 2001b). and in some cases. • Reactivity: Reactivity is one of the measures characterizing the short-term transient response of a locally stable system to a perturbation away from equi- librium. allowing for stability concepts related to more complex nonlinear dynamics.Santa Fe Institute. Close to equilibrium. such as cycles and chaos. For nonlinear systems. Written in this form. but this definition is better described by the terms resistance or robustness. 2005 1:42 p. July 21. The following measures move away from an emphasis on small perturbations in an arbitrarily small neighborhood of equilibria. it is well known that the possible coexistence of multiple equilibria (and other attractors) further limits the relevance of this form of stability. Chen and Cohen 2001b. the relationship between species diversity and community stability. dx/dt = Ax. whose dominant eigenvalue rules the exponential decay or growth of perturbations in the long term (May 1973. continued on next page In the rest of this introduction we provide a brief sketch of the historically central problem in trophic ecology. that govern the dynamics of perturbations (with the vector x specifying the deviations from equilibrium). Pimm 1982). nonlinear systems can be approximated by a linear set of equations. Reactivity is calculated as the maximum instantaneous rate at which perturbations away from equilib- rium can be amplified (Neubert and Caswell 1997. away from the focus on equilibrium behavior altogether. the so-called community matrix in the case of ecological networks. Other relevant quantities characterizing the “stability” of transients are the size and time of the maximum amplification of perturbations (Neubert and Caswell 1997). and sets the stage for outlining how the chapters in this book further develop issues of complexity and stability and . Ch1-Pascual page 5 Mercedes Pascual and Jennifer A. Ruiz-Moreno et al.m. Ecosystem resilience has also been defined as the magnitude of perturbation that can be tolerated before a change in system control and structure (Holling and Gunderson 2002). in spite of the eventual return of the system to equilibrium. the system is specified by a matrix A. This points the reader to a few of many key books and papers in food-web research (see also Box 3). Dunne 5 Box 1: Different definitions and concepts of “stability” used in ecology • Local Asymptotic Stability (LAS): An equilibrium is said to be locally sta- ble if arbitrarily small perturbations away from this steady state always decay. Pimm 1982). Local stability technically refers to asymptotic or long-term behavior. thus pro- viding no information on the short-term or transient response to perturbations (Neubert and Caswell 1997.

see Jansen (1987). and biomasses provide means of measuring biological diversity.g. Chen and Cohen 2001a). For the possible discrepancies between permanence and persistence. 2002b). 1990. 1) represent feeding relationships. permanence measures whether the system’s variables re- main bounded and positive. for example the like- lihood of cascading secondary extinctions resulting from primary biodiversity loss in ecological networks (e. Persistence is in turn determined via numerical simulation.g. expand into new areas of structure and dynamics. in which the species are deleted one at a time to examine if they can reinvade from arbitrarily small numbers. For more technical criteria (i.. Dunne et al. The qualitative theory of nonlinear differential equations has been applied to the QGAS of food webs (Cohen et al. Kokkoris et al. One way to determine permanence is by numerical invasibility analysis. Chapter 6). 1999).g. For a given species. • Robustness: Robustness focuses on the persistence of features of interest in a system’s response to perturbations. by examining the trajectories of the dynamical system for a large number of initial conditions and for a prescribed window of time (e. It has been measured in a variety of ways depending on the system being studied. see Law and Morton (1993) or Chen and Cohen (2001). whose identity. 2005 1:42 p.e. a system is said to be permanent if the boundary of the positive quadrant of state space is a repellor (Hofbauer and Sigmund 1998. It is global in the sense that this return is also independent of initial conditions. Law and Morton (1993). Thus.Santa Fe Institute. Species . QGAS can evaluate responses to large perturbations. Chen and Cohen 2001a).. particularly those the system does not nor- mally experience in its development or history (Jen 2003). • Variability: Variability measures the magnitude of fluctuations in species’ numbers. Martinez et al. • Invasibility: Invasibility measures the likelihood that new species are able to invade an established community of interacting species. abun- dances. Ch1-Pascual page 6 6 From Small to Large Ecological Networks in a Dynamic World Box 1 continued • Qualitative Global (Asymptotic) Stability (QGAS): Qualitative stability refers to the tendency of a system to return to equilibrium when interaction coefficients are specified only by their sign and not by their magnitude. Thus. sufficient conditions for permanence) applied to Lotka- Volterra systems. • Permanence and persistence: These measures do not rely on the existence of any single specific type of attractor but focus instead on whether species remain in the system.. especially within the context of broader theory related to networks and complex systems. It is generally evaluated in assembly models of ecological networks (e. that account for the connectivity of ecological networks.m.. it is computed as the coefficient of variation (standard deviation divided by the mean) of its abundance over time (Pimm 1984). The nodes of Darwin’s “tangled bank” are species. and Chen and Cohen (2001a). July 21. and potentially other interactions. The links between the nodes (fig. More technically.

g. Odum 1953.g. 1998. changing continuously in response to the state of the system itself. replicated. since it emerged out of May’s challenge to prior claims that complexity should enhance stability of ecosystems (e. Garlaschelli et al. Ways that ecologists come to understand the interplay between structure and dynamics of ecosystems can.. Holling 1973. even though complex webs of many interacting species are widely observed in nature. and more broadly.. Dunne 7 richness and connectance. and robustness (e. see also Box 1). 1973) on the relationship between complexity and ecosystem stability. Within ecology. which in turn emerged out of the very early recognition of the fundamental connection between ecological structure and function (Elton 1927). More recently. 1991). ethical fashion. Pimm 1984.m. 2002. This is largely due to methodological limitations.Santa Fe Institute. Connectance and species richness provide simple characterizations of eco- logical network structure by describing global properties of an entire network of interacting species. 2002a. persistence. Complex net- works are characterized not only by numerous components that interact. 1990a. and are adaptive. A much larger set of properties that describe other aspects of global structure as well as more local characteristics of species and links quickly accumulated to address the existence of regularities in the structure of food webs (Lawton 1989. have often been used in food-web research as basic measures of complex- ity. understanding their persistence remains puzzling.. Interestingly.g. Dunne et al. experimentation using microcosms is limited in the . It is basically impossible to manipulate diverse assemblages of species in natural systems in a controlled. Chen and Cohen 2001a. These simple measures are highlighted in the influential work of May (1972. but by interactions that are nonlinear. 2005). Species richness and connectance thus provided an early. McCann 2000. 2005 1:42 p.e. dynamical stability was equated with the mathematical concept of local stability.g. Dunne et al. permanence.. are distributed non-randomly. the proportion of possible interactions that actually occur. 1999. These characteristics obviously apply to food webs. Kokkoris et al. In this and other early theoretical food-web research. Ch1-Pascual page 7 Mercedes Pascual and Jennifer A. measured as the tendency of an arbi- trarily small perturbation to grow or contract in the proximity of an equilibrium point. and continuing. 2003. as well as the reverse. Cohen et al. McCann. influence research in other fields. Such integration has even deeper roots. July 21. to ecosystems. point of contact and integration between quantitative assessments of both dy- namics and structure. This core ecological notion continues to in- form much current research. including resilience. and other interpretations of ecosystem sta- bility have been used. MacArthur 1955). the explo- sion of research on network topology as a fundamental aspect of the study of complex systems (Strogatz 2001. Albert and Barabási 2002) has stimulated new efforts to unravel regularities of ecological network structure (e. Camacho et al. Pimm et al. Jordano et al. invasibility. i. one of the most challenging problems across many scientific fields today is the relationship between the structure of complex networks and their nonlinear dynamics (Strogatz 2001). Much has been said about the relevance of this concept in ecology (e. and should.. 2003).

July 21. Most structural studies focus on “snapshots” of ecological networks. the inclusion of plau- sible structural elements. 1995. 1978). nonlinearities. and there may be general patterns of how interaction strengths are distributed in ecological networks that relate to stability (Neutel et al. Since then. Consequences of such plasticity and other types of indirect effects. need to be explored . albeit somewhat arbitrary. May 1972. Understanding stability and persistence. Furthermore. de Ruiter et al. 1998) in simple models provide some glimpses at constraints that might lead to stability or persistence of diverse. ideally with governing processes. these early models often suggested that complexity gets in the way of stability (e. the notion of incorporating not only the presence or absence of links between species but the strength of those interactions is clearly important for dynamical modeling (starting with De Angelis 1975). 2002). the large and increasing body of ecological literature on higher-order interactions such as phenotypic plasticity (see review by Bolker et al. Dynamic aspects of structure may also prove crucial for our understanding of stability and persistence. Yodzis 1981. Earlier complexity-stability research has generally focused on simple dynamical models such as linear approximations close to equilibrium and Lotka-Volterra equations.g. and assembly processes that occur at both ecological and evolutionary time scales. and conducting nonlinear dynamical simu- lations of “virtual ecosystems” that are diverse and incorporate plausible biolog- ical processes is only now becoming computationally tractable. For example. in turn. temporal and spatial scale. changing in time and space as the result of the plasticity of ecological interactions. Lawton 1999). the central concern of conservation biology. A different issue concerns how ecological network structure is studied. which can play as important a role as direct effects (Menge 1995). 2005 1:42 p. may (or may not!) emerge (Brown 1995. However. can help us understand how and why ecosystems are robust or sensitive to human-related perturbations. simpler picture where coarse-grained patterns.. environmental variation. by compiling a master list of species and their observed in- teractions integrated over some sufficiently inclusive.Santa Fe Institute. complex sys- tems. McCann et al. Pimm 1982. structure is clearly dynamic at many scales. but presents its own challenges such as how to sensibly explore giant parameter spaces. 2003) demonstrates that the strength of interactions between species is not constant but varies in response to various indirect effects that extend beyond the densities of directly interacting species (Peacor and Werner 2001). recent efforts that focus on parasites and mutualistic relationships underscore the importance of extending the scope of ecological network studies to embrace other kinds of interactions. The interplay between ecological structure and dynamics has many nuances. While most observations and theory pertain to webs of trophic interactions.m. Pimm and Lawton 1977. McCann and Hastings 1997. Ch1-Pascual page 8 8 From Small to Large Ecological Networks in a Dynamic World diversity it can embrace. simple models may not be scalable to more complex systems or may oversimplify biology. and/or variable interaction strengths (De Angelis 1975. This strategy allows researchers to go beyond the con- tingent details of local dynamics of a few interacting species to look for a bigger.

m. The second section moves to the explicit coupling of structure and dynamics. STRUCTURE OF COMPLEX ECOLOGICAL NETWORKS Within the study of complex systems. Ch1-Pascual page 9 Mercedes Pascual and Jennifer A.Santa Fe Institute. July 21. From studies of the structure of the internet and the World Wide Web. The first section of this book focuses on the history of and recent advances in uncovering regularities and universal patterns in the structure of complex ecological networks. More challenges for the future are raised in the concluding chapter. which includes an outline of a series of open empirical and theoretical questions. and is followed by a set of chapters that address various aspects of adaptation at ecological and evolutionary time scales. one pervasive subject is the characteriza- tion of the structure of biotic and abiotic networks that include dozens to millions of nodes. Dunne 9 Box 2. A brief roadmap of these sections follows below. Linkages among structure. mutualism). 2005 1:42 p. with a focus initially on food webs that is followed by work on other types of interactions (parasitism. to those of the social links that underlie the transmission of innovation as well as . and ecosystem services systematically with regard to their impact on the dynamics and structure of high-dimensional systems. The importance of an ecological network approach to conservation and restoration is emphasized in the final section. dynamics.

For example. Barabási 2002. More importantly. Just as other kinds of network structure research have undergone a renaissance.g. Martinez et al. Car- tozo et al. provide some dramatic examples of the important role parasites can play in the dynamics of food webs (Dobson Chapter 4. While a few excellent studies have explicitly considered or focused on the role of parasites and parasitoids in ecological network structure (Huxham et al. Recent work has begun to expand this view to include other kinds of interactions. network research is becoming omnipresent across disciplines. Chapter 3). the links typically considered in the literature are con- ventional “predator-prey” interactions. their consequences include positive effects that go far beyond classic “negative” feeding effects. One important example is parasitism (Dobson et al. This inter- est has resulted in a number of novel studies that are building on a new generation of improved food-web data (Dunne Chapter 2. Recent analysis of these data demonstrates that parasitic links are not distributed randomly upon the under- lying trophic network (Warren et al. July 21.. in prep). and then benefits that plant by acting as a dispersal vector for its seeds. with important ecological and graph-theory contexts that extend to the present day (Dunne Chapter 2. there has been a recent renewed interest in potential generalities and simple models relating to the structure of food webs. While these are also a type of consumer-resource interaction.Santa Fe Institute. Strogatz 2003). Chapter 4). more are needed. 2000. their consideration is also likely to illuminate the role of size in determining network structure. Ch1-Pascual page 10 10 From Small to Large Ecological Networks in a Dynamic World infectious disease. Instead. These types of not-strictly-feeding interactions are revealing fascinating nonrandom network patterns that reflect coevolutionary . an an- imal feeds on the fruit of a plant. Mem- mott et al. ecologists were starting to characterize apparent gener- alities in the non-random structure of food-web networks (Cohen 1978) and to develop models inspired by graph theory to explain observed generalities (Co- hen and Newman 1985). non-regular structure of “real- world” networks (Watts and Strogatz 1998). spawning thousands of papers and a number of popular books (e. it demonstrates the critical role played by parasites in both the structure and the biomass flow of ecosystems. Well before the current re-popularization of research on the non-random. Leaper and Huxhman 2002). trophic and parasitic ones. 2005 1:42 p. 1996.m. Because parasites are typically smaller than their hosts. There is a rich and stormy history of empiricism and modeling related to complex food-web structure. 2003. in press) illustrates the enormous field efforts required to obtain the data on both trophic and parasitic interactions. For the most part. a clustering structure is apparent with measures that extend the concept of clustering coefficient (Dunne Chapter 2. The unintended introduction of parasites into ecosys- tems. Buchanan 2002. as informed by earlier graph theory (Erdös and Rényi 1960). Chapter 3) to networks with two types of links. Research on ecological network structure is also being expanded to include plant-animal mutualisms (Bascompte and Jordano Chapter 5). Watts 1999. Cartozo et al. 1999. Box C). Chapter 3). Cartozo et al. A recent study on coastal salt marshes (Lafferty et al. such as rinderspest in the Serengeti and lampreys into the Great Lakes.

but these still require a number of simplifying assumptions. Memmott et al. and represent an important future area of research (Bascompte and Jordano Chapter 5. Chen and Cohen 2001a). For example. Dunne 11 dynamics likely to have important consequences for conservation. Analytical results on these high-dimensional nonlinear systems are rare. Static (probabilistic) models to generate the non-random trophic structure of ecological communities (Dunne Chapter 2) provide a starting point to inves- tigate the dynamics of more realistic network models.. INTEGRATING ECOLOGICAL STRUCTURE AND DYNAMICS Two challenges are immediately obvious in addressing the dynamics of large networks. numerical simula- tions of the dynamical system itself must be performed. Interestingly.Santa Fe Institute.g. 2005 1:42 p. Chapter 14). Ch1-Pascual page 11 Mercedes Pascual and Jennifer A. as demonstrated with nu- merical explorations of parameter space and comparisons to the corresponding models for less realistic structures. An earlier example is found in the Lotka-Volterra cascade model (LVCM) which generates network structure via the cascade model. the niche model of Williams and Martinez (2000) successfully generates a network of links that shares many properties with empirical food webs. including random ones. The first one concerns the need to specify appropriate non-random structures consistent with empirical patterns. if almost non-existent. capable as such of generating a variety of non-equilibrium behaviors in- cluding cycles and chaos (Hastings and Powell 1991. This set of differential equations for the nonlinear population dynamics of the interacting species is based on a bioenergetic model (Yodzis and Innes 1992) describing realistic biology with relatively simple parameters. As an alternative. Memmott et al. a predecessor of the niche model (Cohen et al. such as similar values for specific parameters . July 21.m. 1990b). A dynamical model for trophic interactions can then be built upon the resulting structure (Martinez et al. Fussman and Heber 2002). Those con- sequences (e. 2004) are largely unexplored. Chapter 6). Such hybrid models provide a promising way to address feedbacks between structure and dynamics. allows for the persistence of a surprisingly large number of species. the qualitative global stability (Box 1) of this hybrid model was studied analytically in the limit of a large number of species and long time sequences. which couples structure and dynamics. and focus on ecological net- works in which the signs and not the numerical values of the interaction coef- ficients are considered. The resulting hybrid model..g. They provide stability conditions that can be evaluated numerically (e. The second one is that predator- prey and therefore food-web models are prototypical examples of nonlinear sys- tems. While mathematical analysis is difficult for low-dimensional nonlinear systems and usually impossible for large ones. the alternative of numerical exploration of parameter space through computer simulation is quickly limited by the size of the system being explored.

one can address the opposite question of which structures exhibit particular dynamical properties of interest. Genetic algorithms (GAs. Enquist and Niklas 2001. Chapter 6).. Raffaelli and Hall 2004). with regard to the use of the two different ecological currencies of numbers and energy.g. The few studies of the nonlinear behavior of plankton food-web models constructed with allometric scaling showed a striking propensity for unstable equilibria and pronounced fluctuations. but a substantial part of their work in this area concerned ecosystem models.g.g..g. 1997. Although this first application to food webs considers only lin- ear dynamics close to equilibrium. In the last decade. While earlier work had considered the effect of compartments on long-term stability. Network space is explored by an “evolutionary” process that selects for those networks with the desired dynamical properties. The final chapter on challenges for the future (Pascual et al. Instead of specifying a structure and ask- ing what are its dynamic consequences. where the links between species are not predefined. Yodzis 1982. (Chapter 7) provides an example of an application to the local stability of food webs. steady-state (often linear) considerations have been the norm. results in this chapter indicate that modularity may be critical in short-term transient responses. The use of the patterns them- selves in the formulation of dynamical models for ecological networks has not been pursued. This approach is not new to biological oceanographers and plankton ecologists.g.g.m. through a separate route not directly connected to trophic interactions. Chapter 15) returns to allometry and emphasizes the importance . terrestrial ecologists have become in- creasingly fascinated by allometric scalings in both physiological and community patterns (e. Platt and Denman 1978).. the nonlinear case can also be explored but is computationally more taxing (see Sporns and Tononi [2002] for an example in neurobiology). 2005 1:42 p. One potential avenue for realistic parame- terizations is given by allometric scalings describing how specific parameters vary as a power-law function of the size of organisms (e. They further allow the exploration of the large “network” space of possible structures. Ruiz-Moreno et al. Ch1-Pascual page 12 12 From Small to Large Ecological Networks in a Dynamic World across species. to reduce the enormous size of parameter space (Martinez et al. While much elegant theory has focused on explanations of these patterns. Emmerson and Raffaelli 2004). Mitchell 1996) provide one possi- ble numerical approach to explore the large space of parameters specifying the dynamics of networks (e.Santa Fe Institute. (Chapter 8) begin to explore the connection of al- lometry to food web theory. Moloney and Fields 1991). Sporns and Tononi 2002). and targeted steady-state dynamics and explanations for observed power-law distri- butions of biomass in pelagic ecosystems (e. Gillooly et al. July 21.. Williams 1997). The importance of considering short- term transient responses when evaluating the stability of food webs has also been recently emphasized by Chen and Cohen (2001b). West et al. and explores the relationship between network modularity and the response of the system to perturbations. Another approach to reduce the size of network and parameter space consists of constraining parameters or structure based on empirical findings (e.. Holland 1975.. raising questions about their applicability to nature (e.

and whether ecosystems become buffered to changes (more resilient) over their eco- logical and evolutionary development or proceed to critical states and the edge of chaos” (Levin 1998. Different chapters in this book (Ruiz-Moreno et al. and the related idea of “self-organized instabilities” (Solé et al. the view of ecosys- tems as “prototypical examples of complex adaptive systems” (Levin 1998) is very much based on properties of ecological networks. This is a powerful approach but one in which properties of the regional pool can exert important constraints on the local assembly process. Wilke and Chow Chapter 11) illustrate approaches from CAS. Chapter 10. Martinez Chapter 12. In fact. and levels of selection that are below that of the whole structure. emphasizing the need to better understand un- der what conditions ecological networks converge to these boundaries (Pascual et al. Several chapters (McKane and Drossel Chapter 9. Chapter 15). Pascual et al. Dunne 13 of considering deviations from both steady-state conditions and simple power laws in community patterns. the daunting size of network space can be tackled with theoretical approaches on the assembly of communities (Yodzis 1982. In addition. Chapter 15) outline a series of open questions at the interface of ecology and evolution. Peacor et al. Wilke and Chow Chapter 11). namely the nonrandom distribution and nonlinear nature of interactions between species. July 21. 2002) are discussed. . 2005 1:42 p. The central issue of “how evolution shapes ecosystem properties. Ch1-Pascual page 13 Mercedes Pascual and Jennifer A. Species are allowed to se- quentially invade a previously established local community from a predefined regional pool. and through this structure. to understand responses to perturbations. see also the section on “Community Assembly” in the review by Hall and Rafaelli 1993). Solé et al. computational ap- proaches centered on the concept of digital organisms begin to provide the tools to explore the assembly and dynamics of ecological networks in silico (Peacor et al. Chapter 7. Chapter 10. Law and Blackford 1992. Evo- lutionary processes can also influence dynamical properties by modifying inter- actions over ecological time scales if sufficiently fast. as has been shown with a predator-prey model formulated at the individual level (Hargvigsen and Levin 1997). Finally. see also Kauffman 1993) could have been explicitly written for the ecological networks that underlie ecosystems. 2002. The concept of instability boundaries.m. ECOLOGICAL NETWORKS AS EVOLVING. Post and Pimm 1983. ADAPTIVE SYSTEMS A number of theoretical studies are beginning to address how co-evolutionary processes shape the structure of the resulting food webs (McKane and Drossel Chapter 9). One avenue to define those constraints within the model itself is to incorporate explicitly the evolutionary time scale.Santa Fe Institute. their response to perturbations. parameter space is thus restricted by the trajectory of the assem- bly process itself for persistent local networks.

the dynamics of ecological networks are influenced by changes that are behavioral and modify the structure of interactions. in his still cited masterwork “The Ecology of Invasions by Animals and Plants.. predator switching was incorporated in plankton ecosystem models. Peacor and Werner 2001). in which predation on any one prey is also a function of the relative abundances of the other prey. Chapter 10.. 1990). there is significant scope for formulating models at the individual level and us- ing the models themselves to examine representations at the more aggregated population level (Bolker et al. It is increasingly recognized that interaction strength varies not just as a function of the two species directly involved in the interaction. continuously changing as a function of the state of the system. experiments and models have shown that phenotypic plasticity can have significant effects. Ives and Dobson 1987). and where necessary alter the pattern of food-chains in the world. Chapter 10). but with the abundance of other species in the network.. On the dynamic front. Changes in behavioral or physiological traits of a prey. 2005 1:42 p. where its stabilizing effect on equilibria has been well known for a long time (e.Santa Fe Institute. Another suite of examples is provided by phenotypic plasticity. Indirect interactions of this sort are not mediated by density but by traits. without upsetting the balance of their populations. although only low numbers of species have been considered so far (see review by Bolker et al. it is important to recognize that a main motivation behind the early thoughts on this subject was to address applied is- sues in the conservation and management of ecological systems. Phenotypic plasticity and other forms of higher-order interac- tions underscore the adaptive nature of food webs at shorter. Chapter 10). 2003.m. time scales (Peacor et al. Because these functional forms are not really known. and by predator (or pathogen) regulation of herbivores (Packer et al. For example.g. and contrast with the lethal effects of predators typically represented in graphs of food webs. These studies have also represented phe- notypic plasticity with a prescribed functional form at the population level (e. It is this last problem that has not by any means been solved. The structure itself must now be viewed as dynamic.g. and which is exacerbated every year by the spread of species . discussion in Peacor et al. 2003 ). Ch1-Pascual page 14 14 From Small to Large Ecological Networks in a Dynamic World Besides evolutionary and ecological time scales. driven by the presence of a preda- tor. 2003). Fasham et al. control. STABILITY AND ROBUSTNESS OF ECOLOGICAL NETWORKS Although the relationship between complexity and “stability” is a fascinating theoretical question that continues to prove mathematically and computation- ally challenging after many decades. Peacor et al. It was also considered in a recent theoretical study of large food webs to reexamine the question of complexity and stability (Kondoh 2003). behavioral. For example. affect in turn its own ability to forage and therefore its own predation rate (e. July 21.” Elton (1958) wrote that “the enormous problem still is to manage. Well-known examples of these so-called higher- order interactions are given by predator switching.g.

optimistic tone he added. if not overly.” The last section of this book treats the subject of ecological networks and their robustness to perturbations within the realm of conservation and restora- tion. and other factors vary over many or- ders of magnitude. The effects of habitat destruction and fragmentation are addressed. “Once the notion is grasped that complexity of populations is a property of the community.m. Solé et al. 2005 1:42 p.” where robustness is a type of stability that focuses on the persis- tence of features of interest in a system’s response to perturbations. (Chapter 13) emphasize spatial considerations from a network perspective. and of the structural features that underlie their dynamical responses to perturbations. The conservation of functioning ecosystems will only benefit from a deeper un- derstanding of the robustness and resilience of ecological networks. as well as the collapse of ecological networks following spatial perturbations and species’ extinctions. The case is convincingly made for the importance of a network. The problems have daunting scales of complexity. multi-trophic. interaction strengths. there is hardly any limit to the ways in which it could be introduced. rates of birth-death. they may also enhance our ability to preserve the invaluable diversity of nature. particularly those it does not normally experience in its development or history (Jen 2003). approach to conservation and restoration. Thus.Santa Fe Institute. Dunne 15 to new lands. Memmott et al. (Chapter 14) outline a series of open questions for future theory but also empirical work. One way of thinking about these issues is in the context of ecosystem “robustness. It is one of the defining scientific problems of the twenty-first century as only now do we have the computing power to examine problems which involve evolving nonlinear interactions among large numbers of different components whose sizes. to be studied and used in conservation. yet their solution will not only contribute to the development of the mathematics of com- plexity. July 21. a significant proportion of the quality and possibly even the per- sistence of human life is dependent upon the conservation of natural ecosystems. .” On a more. Ultimately. robustness is a useful way to think about ecosystem response to perturba- tions such as species loss. Ch1-Pascual page 15 Mercedes Pascual and Jennifer A.

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