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Jonathan D.

Phillips

Divergence, Semitivity, and Noneguilibrium in Ecosystems

Contemporay theoretical debate in ecology and biogeography is often focused on equilibrium vs. nonequilibrium behavior in ecosystems and on the nature and source of ecosystemdynamics. I t is suggested that these debates be recast in terms of the way ecosystems develop and respond to disturbances, rather than in terms of concepts often importedfrom mathematics, physics, and otherfields. Using nonlinear dynamical systems theoy, it is shown that key theoretical implications can be cast in terms of geoecologtcallysignijcant phenomenologiessuch as divergent evolution, sensitivity to initial conditions and small disturbances, historical contin ency, and path dqendence. Examples show these phenomena are widely observe in ecosystems. Ecological and biogeographical the0y can be problematized from within eography and ecology rather than fizzy, abstract concepts such as equilibrium, se f-organization, “balanceof nature, or chaos. Complexity, sensitivity, vartabilit , nonsteady states, and other concepts often associated with nonequilibrium or comp exity-the0y frameworks have manvestations that are evident in observable ecological phenomena, in addition to the0y and models.

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1. INTRODUCTION

A fundamental debate in biogeography and ecology is whether, or the extent to which, communities and ecosystems follow a developmental athway leading toward a stable, steady-state equilibrium condition. The absence of, eviation from, or variation in such monotonic developmental pathways is likewise a focus of debate, particularly on the roles and relative importance of external disturbances, intrinsic complex dynamics, and historical or path dependencies. The purpose of this aper is not to provide a comprehensive review or critique of these debates. Rather, &e goal is to attempt to redirect the focus to observable manifestations of (non)equilbrium,(in)stability, and other henomena in ecosystems. Rather than an abstract debate over whether grasslan s, for example, are equilibrium or nonequilibrium systems, research should address specific testable phenomena such as divergent vs. convergent

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Critiques and challenges, tempered by encouragements, from several anonymous referees helped improve this paper.

Jonathun D.Phillips i s a professor in the Department o f Geography, Universityo f Kentucky
(j p@uky.edu).

Geographical Analysis, Vol. 36, No. 4 (October2004) The Ohio State University
Submitted: March 1,2003. Revised version accepted: November 17,2003.

370 / Geographical Analysis evolution and sensitivityto disturbances. The intent is to show that such an approach does not involve abandoning metatheory in favor of pure empiricism, because there are direct links between the behaviors of nonlinear dynamical systems and measurable, observable, geoecological phenomena. Recent decades have seen shifts, or at least broadening, of viewpoints on equilibrium in bio eography and ecology, as documented by Per (ZOOZ), amon others. Even thoug a nonequilibrium view, emphasizing the role o disturbance an chance events, has become common (or even dominant; Perry ZOOZ),the debate is still significant. First, classicial equilibrium concepts are still quite prominent in applied ecology, geography, and resource management. The whole exercise of delineating ecoregions, for example, is implicitly or explicitly based on the concept of equilibrium, climax communities, at least at a hi h level of generalization.The concept and practice of ecosystem restoration, for anot er example, is often linked to the idea of a “natural,”equilibrium ecosystem which can be maintained in a steady state. Second, classical equilibrium concepts are not necessarily at odds with modem ecological thought. Disturbance, for instance, can be treated as a riodic factor necessary to maintain equilibrium, rather than as a source of non- or g-equilibrium. Third, basic equilibrium vs. non-equilibrium debates are still prominent in some fields, such as the study of semi-arid grazing systems (e.g. Illius and OConnor 1999; Sullivan and Rohde 2002). Fourth, acceptance of nonequilibrium viewpoints is uite variable, ranging from those who view nonequilibrium as the rule and norm, to ose who recognize that equilibrium or nonequilibrium may both be possible, and common. Finally, independently of the points above, there is substantial controversy over the extent to which nonequilibrium behavior is linked to external factors versus intrinsic complex dynamics of ecosystems. Equilibrium is defined in various, and often imprecise, ways in geo raphy and ecology. Here the term is used to refer to a steady-state, whereb small uctuations around a constant mean condition. For example, stea y-state equilibrium matter implies that the rate of litter additions is roughly balanced by the so that soil organic matter remains a roximately constant. A steady-state grassland community, for another example, wouf)d)be characterized by an overall species composition that does not vary much, even as individual plants and patches come and go. The link between observable ecosystem properties and histories and theory is accomplished here via nonlinear dynamical systems (NDS) theory, This should not be construed as an implicit claim of primacy for NDS theory. However, NDS theory and methods are critical or relevant to many of the concepts of nonequilibrium ecosystems, and at the very least this work should serve as an illustration of how complex systems concepts can be translated into or interpreted in the context of real-world ecosystem patterns, processes, and histories. The purpose of this paper is not to provide a comprehensive overview of nonlinear dynamics in geography or ecology. Rather, the goal is to identify complex nonlinear phenomenologiesof ecosystems and

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and other sciences that are to a laboratory or simulaand even concepts modification.

ver ent; originally similar ve etation plots, for example, become progressively more

dif erent over time. The con ’tions under which the development of ecosystems and communities is divergent or convergent are of fundamental ecological importance

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Jonathan D. Phillips / 371 and interest. In studies to assess or predict ecolo 'cal responses to climate or other environmental change, for example, Mitchell an Csillag (2001) and Hobbs (1994) point out that understandin the stability of communities may be more important than predicting levels of pro uctivity or other ecological particulars. Having acknowledged that there are no claims for comprehensiveness, note that the examples are selective and biased toward subjects reflecting the interests and experience of the author, and they should not be interpreted as suggesting chaos is more common or significant in these ecosystems or subfields as opposed to other areas of biogeography and ecology.

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2. BACKGROUND: CHAOS AND INSTABILITY

The starting oint is an ecological system with n components that at least potentially affect, an are affected by, each other, Thus the analysis of ecological systems can be approached based on dynamical equation systems and interaction matrices. As these methods are described in some detail in geographical and ecological contexts elsewhere (Logofet 1993; Pahl-Wostl 1995; Phillips 1992; 1999a), this section will focus on a few key points linking nonlinear dynamical systems theory and methods with observed geographicaland ecological patterns and behaviors. The stability of a system to small perturbations can be determined based on the interaction (Jacobian) matrix even if only qualitative information (positive,negative, or zero entries) is available. Dynamical instability in this sense is identical to deterministic chaos. Chaotic systems are often said to have sensitive dependence on initial conditions because even minuscule differences in startingvalues lead to divergent results later. This terminology has been unfortunately misleading for paleoecology, as it superficially implies the ability to estimate or infer initial conditions from the present state of a system. Ecologically, sensitivityto initial conditions is best understood as fndependence of initial conditions; even minor, initially ecologically insignificant variations in starting conditions could lead to a very different system state. Chaotic systems are also sensitive to small disturbances. Sensitivityto initial conditions is usually stressed in the chaos theory literature because of the predominance of numerical modeling. In field studies, as opposed to numerical models and laboratory e eriments, initial conditions are unknowable. And because ecologists often deal with ong time periods, disturbances are likely. Sensitivity to perturbations is therefore at least as important as sensitivity to initial conditions when the goal is to understand or redict system res onses to disturbance, or to reconstruct system states or behavior rom clues in the p eorecord. In a stable, nonchaotic system, development is convergent-that is, variations in initial conditions are reduced, on average, over time. Unstable, chaotic systems are divergent, as variations in initial conditions (on average) are exa erated and increase over time. Observations and measurements of convergence or %vergence, independent of external controls and forcings, can therefore be linked directly to dynamical stability. There is also a direct link between Kolmogorov (K-) entropy and chaos, whereby finite positive K-entropy is associated with instability and chaos. K-entropy, in essence, measures the change in statistical (Shannon)entropy-a well-known and widely used tool in spatial analysis-as a system evolves (Oono 1978; Culling 1988).

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3. ECOLOGICAL MANIFESTATIONS

Based on the discussion above, ecological signatures or manifestations of instabili can be identified. Because they are based on ecologicalphenomena, they can be s t u x ied independently of chaos theory and NDS formalisms, though the latter are often vital to understanding, interpreting, and modeling these phenomena (Pahl-Wostl

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1995; Philli s 1999a; Stone and Ezrati 1996).This section should not be understood to suggest t at ecological s stems are alwa s divergent, subject to abrupt change and historical contingency, an characterized y multiple equilibria, as there are examples of convergence, gradual change, time-independence, and single stable equilibria. Even where the phenomena listed below do exist, they may be unrelated to nonlinear dynamics.

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3.1. Divergent Evolution Progressive differentiation of communities,ecosystems, and landscapes is diagnostic of instability and chaos if the divergence occurs in the absence of, or is disproportionately large relative to, variability in environmental controls and forcings. The latter is an important consideration, as nonchaotic divergence may also occur, associated with obvious, measurable variations in environmental controls or external forcings. Divergence is most readily visualized in a spatial context, whereby initially similar locations become increasingly different, on average, over time. An example would be increasin patchiness of species composition or soil nutrients. Systems characterized by sta le equilibrium, or progression toward such a state, are convergent. Initial variations or minor perturbations are gradually obscured as the steady state is approached or restored. Convergence and divergence are two fundamentally different forms of evolution that are directly linked to equilibrium and nonequilibrium. In the case of convergence, whatever the starting points or the variations in initial conditions, the ecosystem inexorably roceeds toward some articular end state (Clementsian succession is a classic exam e). The latter may be efined specifically (an oak-hicko climax forest, for examp e), or generally (steady state or a critical threshold state, or instance). Disturbances may interru t or delay but do not stop this progression. Divergence indicates g a t the landscape does not move toward a predetermined destination, and in fact becomes more diverse over time. Initial differences become, on average, magnified. The effects of perturbations or disturbances, internal or external, tend to persist and grow over time. Where this occurs in the absence of, or independently of, external controls or forcings, it is indicative of chaos. Further, divergence that is disproportionatelylarge compared to the initial trigger is evidence of instability and chaos. There are several examples from the literature, shown in Table 1. Many of the examples are not grounded in NDS theory. I argue that divergence that is demonstrably large and long-lived relative to the variations or perturbations that initiated it must reflect dynamical instability and chaos. Acceptance of this argument, however, is not necessary for recognition that this key implication of NDS theory can be directly linked to observable, geoecologically significant phenomena. A few studies address divergence as sensitivity to initial conditions within theoretical and methodological frameworks explicitly linked to nonlinear dynamical systems theory and landscape sensitivity. These are summarized in Table 2.

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3.2. Abrupt Changes and Sensitivity

Chaotic systems are characterized by sensitivity to small perturbations, implying that small disturbances have disproportionately lar e and long-lived effects. When es of responses occur near a threshold, t ere may be an abrupt chan e these (often c aracterized as a flip or switch) in the state of an ecological system. Thres olds can, of course, be transgressed due to gradual ongoing changes or large forcing events. Abrupt chan es triggered by dis ro ortionately small (even undetectable) disturbances are attri f Jutable to chaos. T is 'stinction is critical for equilibrium vs. nonequilibrium in ecosystems. Any ecosystem, stable or unstable, may change abruptly in response to large disturbances (for example, volcanic eruptions, tsunamis,

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TABLE 1 Empirical Examples of Divergence in Ecological Systems.

D ylanrl land d e g r d t i o n .

Increasing variability over time in the form of a progressively more complex attern of vegetation and soil nutrient resources. Initially more uniform distribution of vegetation, soil nutrients and carbon, and surface erosion diverges into an increasin ly atchy distrihution (Abraham, Parsons, and Wainwright 199%; ghlesinger et al. 1990; 1996). Plant-soil-microclimateinteractions. Microscale heterogeneil increases due to pro ressive modifications by ve etation; patc y vegetationbare soifmosaics develo from initiay more uniform patterns (Beatty 1987; Hardy and 11bert 1995; Holtmeier and Brolll992; Wilson and A p e w 1992). Increasin ly complex mosaic of marsh plant communities, salt Coastal marshes. pans, andgopen water due to complex interactions between ve eta tion, substrate, erosioddeposition, and h droperiod (Boston 1%3Hackney et al. 1996; Nyrnan et al. 1993; T994; Pethick 1974).Vertical accretion on tidal flats and subsequent salt marsh development is sensitive to the distributions and grazin predation of benthic microalgae in the initial stages of mudfiat5evelopment (Cola 197Q). Mutual adjustments of vegetation and environmental factors such Positive vegetation feedbacks. as microclimate, hydrology, fire regimes, and soils produce divergence in the form of sh ened ecological boundaries and local community “switches”(%son and Agnew 1992). In forest communities, individual trees create nutrient-rich miSelf-reinforcingtree effects. crosites that encourage future tree establishment; trees preferentially reoccupy the same microsites over multi le forest y t i o n s , with pedologic effects increasingTocal soil variability Phillips and Marion 2004; Van Lear, Kapeluck, and Carroll 2000). Initial recruitment in floodplain forests controlled by variations in geomorphic processes persists and is enhanced in the composition of older forests (Robertson and Augspurger 1999). Alternative stable vegetation states within a single landsca e and Grazing effects. vegetation discontinuities, arise after grazing due to compfei interactions between animal im acts, ve etation, microclimate, soil moisture, and soil erosion &allin f988; 1989; Hobbs 1994; Laycock 1991; Friedel 1991; Tausch,%i and, and Burkhardt 1993; Walker 1993; Westoby, Walker, and ioy-bleir 1989). Climate-vegetation feedbacks. Over Quaterna timescales feedback between precipitation and development ofya root mne is characterized by instabilities that lead to differential res onse and divergence into tro ical forest, savanna, semiarid, and Lsert environments from simifar initial conditions (Lapenis and Shabalova 1994). Evolution of biodiversity. Isoto e trends across the Permo-Triassicbounda do not su port mod& of biodiversity collapse and recovery baseyon a singufar event. Rather, the transitions seem to reflect multiple ecosystem stable states and abrupt responses to small changes durin forcings of a complex nonlinear biotic system (de Wit et af%%!)?

TABLE 2 Examples of Studies Showing Sensitivity to Initial Conditions in Ecological Systems.

Coastal wetland$.

Unstable, chaotic interactions between deposition, marsh surface elevation, hydroperiod, and vegetation produce increasingly complex marsh shorelines and s atial atterns of marsh and open water over time (Phillips 1989; 1592; 1b9b; Rankey 2003). Grasslandlwoodland transitions. Historical contingency and sensitivity to intitial conditions, linked to thresholds of grazin ressure, control subsequent successional pathways (Miles et al. &?). Red Spruce decline. Statistical models that best fit tree-rin width data indicate deterministic chaos and sensitivity to initial condftions that lead to rogressive differentiation in forest stand condition (Van Deusen 199Of:

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urbanization), or to gradual changes that go on long enough or whose cumulative effects are sufficient. Very little of the extensive literature on disturbance ecology has been interpreted in light of the magnitude and longevity of responses relative to the size and duration of the disturbance, but it is likely such an interpretation would yield numerous additional examples of sensitivityto small perturbations. Some selected exam les are presented in Table 3. Again, many of the studies cited were not conducte using NDS theory and methods, and some might argue that the results therefore do not necessarily constitute proof or evidence of chaos and instability. Here I sidestep that argument to simply int out that an important im lication of NDS theory-that many systems may be ighly sensitive to small pertur ations and to initial conditions-can be linked to well-documented cases where ecosystems indeed display such sensitivity,

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3.3. Historical Contingency and Path Dependence There is an ever-increasing mass of evidence that historical contin ency and path dependence are critical in ecology and biogeography (Currie and Na elhoffer 2002; Hendry and McGlade 1995; Foster 2000; Mailly, Kimmins, and Busing 2000; Nystrom and Folke 2001; Parker and Pickett 1998; Peterson 2002; Polakow and Dunne 2001; Sinton et al. 2000; Tausch, Wi and, and Burkhardt 1993;Whittaker,Willis, and Field 2001). Instability and chaos cfctate a degree of historical contingency in that such systems have a "memory" of disturbances or initial variations. Historical continency and path dependence can also be due to inheritance of persistent or relict forms or features, or to simple conditionality (e.g., whether or not a site is farmed,

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TABLE 3 Examples of Studies Showing Sensitivityto Small Perturbations and Disturbances in Ecological Systems.
Lake twphic status. Coastal wetlands.

Seminrtd ecosystems.

Ternfirme lowlandforest. Vegetation-wtnd interactions.

Vegetation change. Forest composition.

Multiple equlNbrtum states.

Many lake eco tems sensitive to small erturbations often manifested as abrupt switcxs between clear and tugid or oligotrophideutrophic conditions (Lau and Lane 2002). Effects of local disturbances persist and row dispro ortionately large and long-lived relative to magnitude andgduration otthe i n i t i a l perturbation (Orson and Howes 1992; Orson, Simpson, and Good 1992; Grace and Guntspergen 1999). Minor vegetation change leads to disproportionately large eomorphic response, with semipermanent soil and vegetation results fibrahams, Parsons, and Wainwright 1995; Nicolau et al. 1996; Puigdefabregas et al. 1996; Parsons, Abrahams, and Wainwright 1996). Steady-state biogeochemical cycles unstable and sensitive to minor changes (Brinkmann 1989). Small vegetation disturbances in dunes lead to unstable self-acceleration and formation of unvegetated blowouts (Barth 1982; Gares and Nordstrom 1995); wind-sha d forests linked to perturbations in wind velocity fields (Robertson 1 9 x Vegetation change in data set of 15,000 experimental lots is unpredictable, due manly to growth and ersistence of loci disturbances or influence of s m d local variations (&& 1998). Chance events (gap disturbances) contribute more, and niche partitioning less, than expected to the maintenance of tree species richness in forest gaps (Brokaw and Busing 2000). Complex forest structures arise from contingent interactions amon climate, landforms, stand condition, and disturbance (Sinton et al. 2 d ) . Persistence and exa eration of local disturbances important in creating multiple successionygthways and e uilibrium states (Cattelino et al. 1979; DeAn elis and aterhouse 198q; Hobbs, 1994; Tausch, Wigand, and Burkhar%t 1993).

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burned, grazed, etc.). Where historical contingencycan be directly linked to progressive exaggeration of initial variations (independent of external factors), or to the disproportionately large and persistent effects of perturbations, then chaotic dynamics are indicated.
3.4. Multiple Equilibrla

Many ecological systems have multiple possible equilibria, as opposed to a single preordained equilibrium condition, such as a climatic climax community. In some cases this is due to the instability of equilibria, which makes it unlikely that any given equilibrium state can persist for long in light of the small disturbances that inevitably occur. Multiple stable states have been much discussed in ecology, and these discussions do not necessarily relate to NDS principles. However, d amical instability and chaos can be responsible for the presence of multiple stab e states. For example, Thornes's (1985) NDS model found that the interaction of vegetation cover and soil erosion was dynamically unstable. Any disturbance to the system would tip it to one of two stable states, associated with erodedho vegetation or full vegetation coverho erosion conditions. Subsequent work has confirmed the applicability of Thornes's model (Abrahams, Parsons, and Wainwri ht 1995; Parsons, Abrahams, and Wainwright 1996; Nicolau et al. 1996; Puigde ibregas and Sanchez 1996). Similar phenomena have been identified in lake ecosystems (Lau and Lane 2002).

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4. TESTING AND VALIDATION

Analtytical techniques for testing and validating NDS-based hypotheses in ecology and geography are treated at length elsewhere (Cushing et al. 2003; Logofet 1994; Pahl-Wostl 1995; Phillips 1999% 1999b; 2000). Where time series are of suitable length and quality, a variety of methods exist for detectin complex nonlinear dynamics, and for distinguishingdeterministic from stochastic orcings. These are reviewed in the context of h drology by Sivakumar (2004);the same approaches can be applied to biogeographicJand ecological time series. Experimental investigations of chaos in ecology are outlined and illustrated by Cushing et al. (2003).This section will focus on other means for interpreting potential field evidence of complex nonlinear dynamics, and for distinguishing chaos and instability from other potential causes of spatial differentiation, abrupt switches, path dependence, and multiple equilibria.

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4.1. Chronosequences
A chronosequence represents an ergodic space-for-time substitution. The ideal chronosequence consists of a series of sites or environments where all the factors controlling ecosystems except time or age are invariant, so that the sequence can be assumed to represent a temporal progression. The chronosequence a proach has been most often applied in edology and soil geography (Huggett 1998 but also in plant ecology (Foster and T man 2000). It is fiendishly difficult to identi a situation where factors other than age or time are negligibly variable, and even en there is the problem of the longer, and thus different, histories of the older members of the chronosequence. Nonetheless, given the long time scales and historical investigations that often characterize physical geography and ecolo chronosequences may offer insight into temporal trends that is not otherwise avai able. A good chronosequence, by definition, means that there are no si nificant variations in geology, climate, disturbance regime, or other factors that in uence ecosystem development. Thus the hypothesis of increasing variability over time associated with divergence can be directly tested, as any increase in variabili must occur due to the unstable, chaotic magnification of the effects of minor initi variations or small perturbations. Any relevant metric of the spatial variability or richness of the phe-

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nomenon of interest can be computed for each stage of the chronosequence to determine whether variability is increasing. Generally,

where DI is a divergence index, and V is some measure of variability measured at two time increments t and t - x . DI > 0 indicates divergent and DI < 0 convergent behavior. While not appealing directly to NDS theory, Gracheva, Targulian, and Zamotaev (2001)explicitly developed their model for interpreting tropical and subtropical soil chronosequences in terms of divergent or convergent pathways. Barrett’s (2001) study of a strand plain soil chronosequence in Michigan used the statistical scatter in chronofunctions (soil property vs. time regressions)as a measure of variability, which showed increasing scatter over time. She attributed this to chaotic pedogenesis, as did Phillips (1993),who used simple measures of soil taxonomic richness to demonstrate increases in variability in a North Carolina chronosequence. Chronosequence-based tests of NDS hypotheses (or use of NDS theory to interpret chronosequences) has not been directly applied in biogeography or to ecological sequences, but such application would be reasonably straightforward,using a variety of available metrics for biodiversity and landscape heterogeneity (Gustafson 1998; Haines-Young and Chopping 1996; Ibanez et al. 1995; Pike 2000).
4.2. Historical Data

The same basic logic represented by equation (1)can be ap lied, with caveats, to any historical representation of ecosystem change or ecolo *ca!)dynamics (for exameoecological reconstrucple, sequential aerial photogra hs or satellite images, p aY tions, long-term monitoring). T e caveats are that one must be able to show, or at least reasonably assume, that the time periods or episodes are not characterized by significant changes in inputs or environmental controls, or by large disturbancesthat is, the temporal slices must be analagous to a chronosequence in that each stage or sample differs only in the time factor.

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4.3. Variability within Elemnta y Areas If we define an elementary area A, as a contiguous finite area with constant or negli ‘bly variable environmental controls, then a similar logic to the above can be app ed in the spatial domain. If variables controlling habitat, for example, are constant within Adto some acce table level of measurement precision, then (again, by definition) ecological variabi ‘ty within A, may be attributable to the dynamically unstable exaggeration of the effects of variations in initial conditions or persistence of the effects of small disturbances. In some cases this may re uire accounting or adjusting for the effects of chance (for example, in seed dispersal ,depending on the phenomenon of interest. There are many spatial statistical techniques for apportioning within-unit vs. between-unit variability, any of which mi ht be employed to test for chaos and dynamical instability if the units represent e ementary areas as defined above. Other techniques, such as specieshrea relationshi s, may also be employed. Define an area of interest A, subdivided into n uniform e ementary areas A,, such that C A, = A. The most typical relationship relating species richness (number of different species) to area is of the form

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S=cAb

(2)

Jonathan D. Phillips / 377 where S is richness and c and b are empirical parameters. Speciedarea curves can also be derived for each elementary area,
Sl = c1A,hi

(3)

Then

where the overbars indicate mean values and m = C S,/S to adjust for species which are counted in more than one elementary area. Because
-

FIA,himn=cAb

(4)

the ratio gI/bindicates the relative importance of intrinsic variability attributable to complex nonlinear dynamics to that associated with environmental heterogeneity (that is, the habitat difference between the A J . This method is described more completely by Phillips (2001). For example, Phillips and Marion (2004) hypothesized that short-range soil variability in forest soils in Arkansas was due to instabilities associated with the effects of individual trees on soil morphology. One test of thz hypothesis was a richnesdarea analysis of 16 elementary area plots, where the b,/b ratio of 1.15 indicated that within-plot variation of soil taxa, consistent with the hypothesis, was greater than between-plot.
4.4. Landscape and Ecosystem Entropy

As noted earlier, there is a direct relationship between Kolmogorov (K-) entropy, representing the change in Shannon entropy as a dynamical system evolves, and chaos. K-entropy is given by

where p , represents the proportion of pixels, cells, sample sites, area, etc., in category i. The categories could represent vegetation communities, classifications of spectral data, soil types, etc. Hk > 0 indicates chaos, as finite positive K-entropy is related to the Lyapunov exponents by

Recall that a positive Lyapunov exponent is required for deterministicchaos. The application of entrop in a chronosequence or historical context as described above is straightforward.Ac rditionally, in some cases ecosystems and landscapes may be interpreted to determine the nature and direction of entropy changes, and thus whether there is positive K-entropy. This ap lication of K-entropy in an interpretive context is illustrated for tidal creek networks y Rankey (2003),for soils and weathering profiles by Phillips (ZOOO),and for soil landscapes by Ibanez et al. (1994) and Phillips, Gares, and Slattery (1999).For exam le, if it is clear that that vegetation type A is encroaching into types B and C, then i e effects of these changes in ( p , p b , p,) can be examined to see if entropy is increasing or decreasing.

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4.5. Magnitude o f Perturbations

Chaotic systems are characterized by sensitivity to small perturbations, whereby small disturbances have disproportionatelylarge and long-lived effects. Thus in some cases a test of whether abrupt changes, historical contingency, or multiple states might be attributable to dynamical instability and chaos can be based on whether a perturbation (or variation in initial conditions) is “small“relative to the magnitude of the resulting change. This can be based on the magnitude of the perturbation in terms of longevity, area of impact, energy (or ower, force, shear stress),or mass. For example, a perturbation magnitude index cou d be based on

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PMZ, = A$,/&,

(7)

where Atp and At, represent the duration of the perturbation and the response, respectively. Similarly,

where ”p and A, are the footprint or area of the erturbation and response. Analagous PMZ’s could be developed with respect to mass, orce, or other parameters. A small perturbation would be associated with PMZ << 1. For example, a pine tree in Louisiana would have a life span on the order of 100 years or less, while fossil tree casts in soils of south Louisiana may be Pleistocene in age (Mossa and Schumacher 1993). Thus the pedologic influences of the trees have a PMZ, < 0.01, indicating a small perturbation and unstable exaggeration of its effects. Likewise, an invasive weed inoculating a 1 ha field and subsequently establishing dominance over a 1,000 ha area would produce PMZ, = 0.001. An invader that is seeded over 1,OOO ha, but whose local range is subsequently reduced to 100 ha has PMZa = 10, which represents a stable system where the effects of perturbations are damped.

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4.6. A Note on Scale

It has often been said, only partly in jest, that the answer to every question in hysical geography is “it depends on the scale.’’ Stability, instability, and other m A e s t a tions of NDS are emergent-that is, they appear or disappear as time frames, spatial resolutions, and levels of detail are changed. This has obvious relevance for this paper, in the context of large and small perturbations and the magnitude of changes or responses relative to that of disturbances. There are also issues regarding the resolution of temporal and spatial data vis-a-vis the relevant scales of nonlinear dynamics, and the practical problems of determining (for example, in a chronosequence or an elementary area) just what constitutes homogeneity or constancy. The identification and significance of nonlinear dynamics will, indeed, depend on the spatial and temporal scale. Thus it is inap ropriate to state that a grassland community or a forest ecosystem is unstable (sta le) or divergent (convergent)without specifying the spatiotemporal frame of reference.

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5. DISCUSSION

5.1. Nonlinear Dynamics in Landscape Ecology and Biogeography The review above, though incomplete and selective, suffices to make the point that nonlinear dynamics as manifested in ecosystems are fully consistent with many existing and emerging threads of inquiry in biogeography and landscape ecology, includ-

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ing considerations of nonequilibrium systems, multiple stable states, historical contingency, path dependence, and divergent development. The presence and possibility of complex nonlinear dynamics is also important in that it may complicate (or inform) efforts to interpret historical evidence such as paleoecological data. Due to dynamical instabilities and chaos, small perturbations or short-lived disturbances rather than larger environmental change may be responsible for changes or switches. Due to multiple equilibria, there may not be a one-to-one correspondence between system states and environmental controls. In some cases complex nonlinear dynamics can provide explanations of phenomena not otherwise explained. In climatology there are numerous examples where nonlinear d amical instabilities provide the only plausible, or the most widely accepted, exp anations for phenomena such as abrupt shifts observed in paleoclimatic proxy data. Stone and Ezrati (1996) ve a number of ecological exam les. Nonlinear formalisms and metho s may also provide tools for un erstanding, explaining, and describing complex ecological systems. Chaos is sometimes interpreted to mean that prediction is impossible, and that for practical purposes chaotic systems are indistinguishablefrom genuinely random systems. If an irregular pattern is shown to be chaotic, this actually poses several advantages. Chaos indeed inhibits some forms of redictability and limits the range of forecasts. However, chaos by definition implies tfe presence of underlying deterministic dynamics. Finding the underlying generator (particularly in terms of processes or environmental controls) of pseudorandomness may be an important step to redictability. While chaotic systems produce complicated patterns that are unpre 'ctable in detail over long timeddistances or many interations, these atterns are often perfectly predictable a few iterations into the future. Further, wEl'le the details are irregular and unpredictable, chaotic patterns have well-behaved statistical moments. By definition, chaos occurs within deterministically defined boundaries: unlike true randomness, all outcomes are not equally possible, and the entire hase space is not potentially filled. These properties have yet to e well exploited in bio eography and ecosystem science, but examples are available from other field-base sciences. For example, nonlinear models performed better than linear models in forecasting surf zone sediment suspension ( affe and Rubin 1996). The recurrence patterns of lake-volume fluctuations were s own to be better modeled and forecast by nonlinear models than by classical linear methods of time series analysis (Sangoyami, Lall, and Abarbanel 1996).The methods used in these studies capitalized on two important characteristics of chaotic sequences: the ability to predict deterministicall a few iterations ahead, and the role of histo in conditioning subsequent values o a time series. Neither is possible in a system at is treated as linear and random.

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5.2. Concretization

If ecosystems are in or progressing inexorably towards stable steady-state equilibrium, they will be characterized by convergent evolution, insensitivity to minor variations in initial conditions, and the ability to recover from and negate the effects of small disturbances. If ecosystems are not in or progressing toward a stable steady state, they are likely to be characterized by divergent evolution, sensitivity to initial conditions, and the unstable exaggeration of the effects of small perturbations. Historical contingency and ath dependence may occur in equilibrium or nonequilibrium systems, but in the atter case should be linked to sensitivityto initial conditions and small perturbations. Thus at least some implications of nonequilibrium and equilibrium in ecosystems can be directly linked both to theoretical formalisms (in this case nonlinear dynamical systems theory) and to observable, geographically and ecologically meaningful henomenologies. These phenomenologiesrelate directly to issues of the role and efp ects

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of disturbances, spatial heterogeneity, and evolution and historical development of ecosystems. It seems, then, that b shifting the terms of debate from equilibrium, chaos, selforganization,and so f o A to ecological1 meaningful phenomenologies such as divergence and disturbance responses, the ebates can be concretized. The controversies can be firmly rounded in geographical and historical observables without losing connections to de ates grounded in metatheories such as NDS theory.

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6. SUMMARY AND RESEARCH AGENDA

Sullivan and Rohde (2002,p. 1,595) write that “a debate in ecology rages over the sources and types of dynamic behavior driving ecological systems.” This study suggests that in many cases intrinsic properties of nonlinear ecosystems drive the dynamic behavior, and that these are manifested in measurable, observable, ecologically meaningful phenomena in real-world ecosystems. Perry (2002,p. 354) writes, “It has been realized that s ace can fundamentallyalter the dynamics and outcomes of ecological processes. &s has been largely achieved through theoretical, model-based studies; the challenge remains to empiricallytest and explore the large body of spatial ecological theory.” In the case of NDS, the implications of theory can be directly translated into ecological phenomenologies that are empirically testable. It is likely that this is, or will, also be the case with other threads of spatial ecological theory, This suggests that there should be more links between models and observational data. This can be accomplished not only by theorists linking their work to field studies and experiments, but also b the examination of historical contingency, multiple equilibria, etc., in the context o NDS and other theories. This encompasses the need to develop hypotheses regarding ecosystem dynamics and spatial ecology that are testable on the basis of observations in nature. Chaos, instability, and other forms of nonequilibrium do not preclude predictability but do provide a new context for predictabili We need to know the range, accuracy, precision, and uncertainty of predictions, w ich are all profoundly influenced by phenomena such as chaos. Where nonequilibrium exists, this further implies a critical need to assess the extent to which unpredictability and uncertainty are inherent in system structures and dynamics, as opposed to being associated with stochastic forcings, large degrees of freedom, and information inadequacies. The new context for predictability may well be one where we recognize that we can perhaps never accurately forecast the time and spatial coordinates of pest invasions or ecotone shifts, but can move toward accurate and recise identification of synoptic situations, where such events are more or less like y, and better identification of trigger mechanisms. This research agenda should be problematized from within ecology and geography rather than NDS or com lexity theory. The fundamental manifestations of chaos, for example, in ecology can e linked to observable ecological phenemona. Ecosystems themselves, as opposed to models, should be the primary source of ideas on ecosystem dynamics.

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