Advances in Ecological Understanding: By Kuhnian Revolution or Conceptual Evolution? Author(s): Robert T. Paine Source: Ecology, Vol. 83, No.

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Ecology, 83(6), 2002, pp. 1553-1559

C 2002 by the Ecological Society of America

ADVANCES IN ECOLOGICAL UNDERSTANDING: BY KUHNIAN REVOLUTION OR CONCEPTUAL EVOLUTION?

ROBERT T. PAINE

Department of Zoology, University of Washington, Seattle, Washington 98195-1800 USA

Abstract. Ecology, although a rapidly evolving science, has experienced few if any paradigm shifts or revolutions in the sense of Thomas Kuhn because it is a pluralistic, multiple-causation discipline. Instead, what appears to characterize conceptual development are fads or "bandwagons," favored themes, or transient foci of interest, that wax, wane, and even recover in popularity. These are often centered on characteristics of the real world which cannot be denied. I identify and discuss two: postlarval recruitment to an established population and physical disturbance to an intertidal assemblage. The richness of ecological thought has been broadened immeasurably by acceptance of the following themes: the necessity to distinguish open from closed systems, that equilibrium may exist only in an abstract sense, and that controlled manipulation can reveal hitherto unanticipated dynamics. These findings have expanded our understanding of nature's complexity (a conceptual evolution) rather than relegated most prior generalizations to the dustbin of history (a paradigm shift).
Key words: disturbance; ecological history; intertidal; Kuhnian revolution; mussel; Mytilus californianus; paradigm; recruitment; rocky shore; Semibalanus cariosus; settlement.

INTRODUCTION

"Paradigm" dramatically entered the active vocabulary of science with the publication of Kuhn's The Structure of Scientific Revolutions (1962). The concept of normal science punctuated by revolts led by young, disenchanted enthusiasts (Wade 1977, Wilkins 1996) provides definitive hallmarks for identifying paradigm shifts or revolutions. The interpretational difficulty of what constitutes a legitimate revolution is enormous and depends critically on whether Kuhn's examples, drawn from the physical sciences, can apply to biology. Subtle definitional challenges abound: How does one characterize a revolution or paradigm shift? What is its genesis? How sweeping must it be? However, I believe there is a more fundamental dichotomy. Concepts in the physical sciences, Kuhn's subject matter, are predisposed to contradiction by single well-conceived observations or experiments. The linear or binary nature of such constructs permits challenge (Strohman 1997). Thus, revolutions are not difficult to recognize in other disciplines: Copernicus' heliocentric theory, the rediscovery of Mendel and particulate inheritance, the theory of continental drift, and Michelson-Morley's disproving of the existence of an ether permeating the entire universe. At times the disciplinary restructuring or redirecting was abrupt and relatively rapid (Michelson-Morley), at other times, the "revolution" simply catalyzed a gradual replacement of the older views (Copernicus). On the other hand, the biological sciences at levels
Manuscriptreceived 3 November 2000; revised 16 July 2001; accepted 26 July 2001; Special Featureaccepted 7 October 2001. For reprintsof this Special Feature, see footnote 1, p. 1479.

of the organism and above are plagued by multiple causation, generating what McIntosh (1987) describes as ecology's pluralism. Thus, I will argue, paradigm shifts are apt to be less obvious in our field, if they truly exist at all. For instance, did Elton (1927) change ecology with his concept of food cycle (a.k.a. food web)? I believe the answer is no. Brilliant tour de force that his book was, it is probably best described as an important advance in a lengthy quest to understand natural complexity. Darwin (1859) had previously expressed a deep interest in such issues and the multiplicity of associated interactions. Slightly later, Forbes (1877) openly described the importance of interspecific relationships. Thinking about trophic structure has continued to evolve (e.g., Lindeman 1942, Cohen 1978, Pimm 1982). This 100-plus year quest for understanding is best described as a progression but one that is uneven through time as interest waxed and waned. While there have been interpretative disagreements, revolutions are nowhere to be found. I suggest that what ecology does have in profusion are fads or bandwagons. "Bandwagon" is defined (Gove 1971) as "a party, faction, or other element that attracts adherents by its timeliness, vigor, showmanship, or novelty." I do not employ the term in a pejorative sense: these are transient foci of interest, usually only temporarily in vogue. Table 1 provides my partial list of bandwagons in ecology, assembled by comparing early texts (Chapman 1931, Allee et al. 1949) with their modern counterparts (Begon et al. 1990, Krebs 1994). I searched these texts for commonalities, with special sensitivity to novel additions and shifts in emphasis in an effort to calibrate my own perspective. My list includes subjects identified as ma-

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TABLE 1.

ROBERT T. PAINE Some bandwagons in ecology. Sub-themes Null models Optkmality Population regulation Trophic dynamics Web complexity Interaction strengths Indirect effects Disturbance Species-abundance patterns Equilibrium/nonequilibrium Metapopulations Ecological efficiencies Functional redundancy Services

Ecology, Vol. 83, No. 6

Major themes Niches

Food webs

Succession Stability and diversity

Ecosystems

jor themes, each of which incorporate relatively minor, more specialized components. Collectively, they reflect substantial refinement and progress unpunctuated by revolts. Thus, a bandwagon is a temporarily popular focus or research agenda, sort of an ecological supernova which bursts into prominence and then fades with time. No one doubts their reality or importance. Interest usually wanes because technological limits have been reached (energetics and bomb calorimetry), theory development has outpaced data acquisition (the stabilitydiversity relationship), or fatal conceptual flaws are uncovered (communities as superorganisms). On the other hand, paradigm shifts require substantive precursors (i.e., a belief system to be overthrown). Argument exists whether evolutionary change driven by natural selection qualifies as a paradigm shift (Wilkins 1996). The Watson-Crick model for the structure of DNA does not: it was a de novo discovery in a continuum of change, and was not opposed by an entrenched group of elder scientists (Wilkins 1996, Strohman 1997). All bandwagons identified in Table 1 are deeply rooted in ecological reality, i.e., they are not hypothesized, invisible constructs. Although my list is certainly incomplete, the omitted topics largely reflect difficulty in defining what constitutes a bandwagon as opposed to continuing background science. "Designer assemblages" (Ruesink and Srivastava 2001) are experimentally popular now. Will they persist or expand in use, or will their artificial construction lead to questions about the robustness of the results? Austin (1999) identified a historical dichotomy between botanical interests in gradient analyses and ordination on one hand, and more mechanistic experimental pursuits by animal ecologists on the other, as a difference between paradigms. Are these differences legitimate or are they simply constraints or advantages imposed by the research material? I would have described these as tactical differences because the methodology, not the fundamental question, differs. My list reflects personal biases and

ways of organizing and subdividing some of the myriad subjects that compose ecology and generate its pluralistic nature. Thus, species do appear to have individualistic lifestyles (niches), to replace one another through time (succession), to eat and be eaten (food webs), and to inhabit environments influenced by both physical and biological stressors (ecosystems). Our vocabulary is of necessity general and not precise; the term bandwagon is intended to catch the essence or flavor of a topic of interest, whether it passes quietly or persists. For instance, ideas on change (succession) began with Warming (1896) and Cowles (1899), were codified by Clements (1916), overextended and debated (Phillips 1934, Tansley 1935), mechanistically expanded (Drury and Nisbet 1973, Connell and Slatyer 1977), and remain the subject of intense investigation (Turner and Dale 1998). I cannot conceive of a revolution that could or will replace our increasingly sophisticated and mechanistically detailed understanding of how ecosystems respond to disruption. The same trajectories of refinement and increased appreciation can be identified for ecological niches, food web construction, foraging of organisms, variation in species richness, factors influencing primary and secondary production, and even global warming and sea-level change. All share a common biological heritage: the enormous complexity of natural systems. They also share the following trait: multiple causation or explanation is commonplace and unavoidable, and thus numerous alternative hypotheses are the norm for dissecting patterns. This fact, by itself, tends to insulate generalizable (and popular) ecological views from radical changes or scientific revolutions. Because ecology is not a linear discipline, breaking or challenging one logical strand is unlikely to generate a major overhaul. In this sense MacArthur's (1955) views on stability should also apply to paradigm shifts. In the following paragraphs I provide two examples of bandwagon development based on personal involvement and interest, and I argue that they are so fundamentally and naturally important that they will not, and cannot, be replaced. Detailed refinement, not revolutionary displacement, is the sole alternative.
RECRUITMENT LIMITATION

Individual species populations grow or shrink, reach nuisance proportion or go extinct, based on their reproductive potential and their survival. Limitation of effective reproductive output was a key component of Darwin's views on natural selection. The topic of spawner:recruit ratios became central to fishery biology more than a century ago (Sinclair 1988). The primary issue then, as now, was whether the reproductive output of a stock could be related quantitatively to the number of individuals recruiting to that stock after their prolonged exposure to a host of unknown mortality factors. The relative contributions of pre- and postrecruitment mortality have important implications for management

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and harvest schedules and for conservation. I consider the issue to be a bandwagon because of its "timeliness" and continuing importance, and one unlikely to be subject to a scientific revolution. The opportunity to test this view occurred in the early 1980s when recruitment was rediscovered (Underwood et al. 1983, Connell 1985, Gaines and Roughgarden 1985), named "supply-side ecology" (Lewin 1986), and advertised as a fatal and missing flaw in understanding rocky intertidal shore dynamics (Underwood et al. 1983). One reality is certain: populations will inexorably go extinct, locally and globally, without replacement. Effective reproduction is thus essential, and studies which ignore the dynamics of replacement should be suspect. The issue has been especially dramatic in marine ecology, where many species release gametes into a water mass of unknown dimension, with unknown (and perhaps unlikely) fertilization success, spend variable dispersal intervals in currents only vaguely described, and then settle onto the shore. Settlement often involves a dramatic change in morphology, lifestyle, and sources of mortality. Recruitment occurs when specific identity can be determined; this technically demanding determination often is made some time after settlement, during which interval the settled population has been subject to numerous sources of mortality when it is maximally vulnerable (Caley et al. 1996). Employing this operationally effective definition, however, impairs the ability to determine the

relative significance of pre- vs. postsettlement processes. The settlement/recruitment phase involves much basic biology. The details vary from species to species depending on life history traits, with mobile species being potentially more vexing subjects than sessile invertebrates. The fundamental question, whether it is conditions during the transport phase in the ocean or postsettlement dynamics which determine the character of the postsettlement population, is crucial. Because annual or subannual species are apt to spend a greater fraction of their lifetime being subject to the vagaries of transport, I would anticipate an enhanced effect of presettlement factors. For perennials that conclusion is much less certain and depends on their potential longevity since, for instance, perenniality (Cole 1954) and storage effects (Warnerand Chesson 1985) can insulate the species from years of near-zero reproductive output. As an example (Fig. 1), I have measured since 1978 the recruitment of the barnacle Semibalanus cariosus in newly formed patches in mussel beds at Tatoosh Island, Washington, USA. The pattern is one of enormous temporal variation; some years are excessive, while others fail to attain a replacement level. If this species were an annual, it would require -90 recruits per 100 cm2 to saturate that surface after one year, given perfect survival (i.e., no benthic interactions leading to individual death) and known growth rates (i.e., Dayton 1971, Paine and Levin 1981). That level of re-

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Ecology,Vol. 83, No. 6

cruitment is achieved in only eight of the 22 yr, presumably eventually dooming an annual species. S. cariosus, however, is not an annual and possibly lives -20 yr (Connell 1972, Sebens and Lewis 1985). If these barnacles reproduce in only five of these years, a very conservative estimate, the recruitment must be 18 barnacles per year on average, again without mortality, to saturate the space. This level is exceeded in 15 of the 22 years S. cariosus, like redwood trees, is not ecologically threatened by occasional settlement/recruitment failures. Two attributes of the recruitment limitation issue remain unresolved. First, massive, benthic-based mortality must often occur before recruitment and after settlement (e.g., Wethey 1984). This mortality will have nothing to do with the planktonic interval, its existence is experimentally testable, and for perennial species, it challenges the notion of primary determination of adult population traits by events during the dispersal phase. As a related aside, the excessive mortality characteristic of postrecruitment crowding due to settlement binges (Barnes and Powell 1950), capable of eliminating entire recruit classes, should further cloud the recruit-spawner relationship. Second, because variation in recruitment has been recognized for decades (Hjort 1914, Thorson 1946), and is totally natural, there can be no paradigm shifts. There can be refinements, increased mechanistic understanding, better modeling of the transition from planktonic to benthic existence, or increased appreciation of the relative contributions to population dynamics of mortality during the transport and postsettlement phases for our theories of variation in recruitment, but radical restructuring of our views is not possible. Thus, important refinements (i.e., conceptual evolution) should continue to characterize our interpretation of life history dynamics spanning two very different environments.
DISTURBANCE

Field ecologists have probably always been aware of change, and certainly our discipline's first major theory, the notions of primary and secondary succession (Warming 1896, Cowles 1899, Clements 1905, 1916) were developed in the context of change. Classic publications such as Watt's (1947) pattern and process paper identified cyclical dynamics and therefore no single general termination to a successional sequence. Because the initial focus was on terrestrial landscapes, it should come as no surprise that fire was broadly acknowledged as a generating pattern. Heinselman's (1973) graphic maps suggested how fire influenced forests in Minnesota, and studies in Yellowstone National Park (e.g., Taylor 1973) helped to change our national fire suppression mania. Even earlier, Turner (1935) had attributed localized age-structure difference in pines to a region's tornado history. Perhaps the culmination of this trend was Pickett and White's (1985) book, The Ecology of Natural Disturbance and Patch Dynamics.

Physical disturbances to landscapes continue to attract the attention of ecologists, and it seems certain that this bandwagon has not run its due course. The primary reasons are obvious: disturbance, perhaps especially when ultimately expressed as global change, is an inexorable fact of life, and ecologists are still on the steep portion of their learning curve (Turner and Dale 1998). The following paragraphs explore one dimension of disturbance ecology in an intensely studied rocky intertidal system. The Cape Flattery region of Washington State in general, and Tatoosh Island in particular, has been the venue of a variety of such studies (Dayton 1971, Levin and Paine 1974, Paine and Levin 1981, Dethier 1984). Beds of the mussel Mytilus californianus owe their internal spatial patterns to waves which remove mussels and thus produce gaps. The gaps or patches are "born" (disturbance) and "die" (recovery) at rates which are patch-size dependent. Levin and Paine (1974) and Paine and Levin (1981) modeled this system, assuming that settlement into patches was independent of events occurring to mussel larvae during their planktonic interval, that the individual mussel beds had no defining history, and therefore that disturbance intensity during a given winter interval was independent of that site's local history. The latter assumption seemed realistic, and permitted averaging disturbance traits (gap size at formation) across many sites. Long-term observations suggest that these assumptions were premature. Disruption to 26 independent (spatially distinct) mussel beds has either been measured directly or estimated each spring since 1974. Earlier data in this time series are relatively sparse. Fig. 2 presents the mean annual disturbance for 25 sites scattered around the south, west, and north periphery of Tatoosh Island. A site from which the starfish Pisaster ochraceus was removed was deleted from the tally. The original data, expressed as the percentage of mussel bed removed each winter, were arcsine squareroot transformed, averaged for the island, and then back transformed. Visual inspection suggests a pattern of peaks and troughs. A cold water winter in 1975-1976, extreme El Nifio conditions during the winter of 19821983, and a brief but severe February cold snap which killed almost 15% of the island's mussels in 1988-1989 stand out in the data. The 1982-1983 event was particularly destructive to mussels close to their upper limit, mussels which are always small (<3 cm shell length) and usually unaffected by wave forces. Environmental correlates of the 1989-1990 mortality are unknown, as are those for 1998-1999. The 1996-1997 mortality can be attributed to an exceptionally severe November storm, one which even destroyed breakwaters 50 km to the east. If I include data recorded in photographs from my first scientific visit to Tatoosh, substantial M. californianus mortality also occurred in the winter of 1967-1968. Thus the pattern from June 1968 to June 2001 suggests peak disruption intervals

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of 8, 7, 6, 1, 7, and 2 yr. Fourier transformation and analysis reveal a slow peak at 7.7-yr intervals and a faster one at 2.3-yr intervals. What do these data suggest about the disturbance regime to mussels on Tatoosh? First, they share no common environmental cause. Some El Niio intervals are characterized by major disturbance (>13%) whereas others are not (1997-1998, early 1990s). Severe storms and deep freezes also generate major mortality events. If one dismisses the 1989 freeze because it primarily impacted small mussels usually invulnerable to wave stresses, and accepts the photographic evidence of 1968, the five peaks average 6.2 yr apart. This mean is reflected in the time series identification of a 7.7-yr periodicity and an estimate of a minimal rotation period of 7-10 yr in Paine and Levin (1981). This periodicity implies a temporal pattern to disturbance and that years are not truly independent of one another. The underlying biological reason is that as mussels grow, their susceptibility to wave force increases substantially, and beyond some size the probability of destruction, on intertidal wave-swept shores approaches unity (Denny et al. 1985). The 2.3-yr periodicity remains unexplained. Can such revelations alter the reality that winter waves carve patches in outer coast mussel beds? Certainly not. At best the realization will help expand and calibrate more sophisticated spatial models dealing with the endless cycles of disruption and recovery which generate the mosaic of pattern so essential to and characteristic of these environments. In a broader context, disturbance processes, whether marine or terrestrial, interrupt successional trajectories. New and interesting biological detail and model refinement will continue to characterize this fertile subdivision of ecol-

ogy. These will fine-tune this bandwagon (reducing squeaks in the wheels?); they will not overturn the cart which seems robust in its diversity and central to much of ecology.
AN ALTERNATIVE VIEW

I have reasoned that because ecology in the broadest sense is a pluralistic, multiple-causation discipline, revolutions are unlikely. That does not imply that conceptual changes have not occurred. The clearest example of a possible legitimate paradigm shift known to me is MacArthur and Wilson's (1963, 1967) theory of island biogeography. Their hypotheses meet most of Kuhn's criteria for revolution: the authors' relatively youthful ages and the redirection of an important focus of ecology (biogeography) which broke from the prior standard analysis of an island's area and assessments of its biota to suggest dynamical alternatives. There have been other shifts, but they are more tactical and general rather than specific. Two have been in progress for decades. Connell (1961), Paine (1966), and Dayton (1971) introduced the notion (though hardly for the first time; see Paine 1994 for details) that field experiments could reveal subtle yet important dynamics influencing species distribution and abundance. Experimental, manipulation-based hypothesis testing is now commonplace. This approach represents an attitude adjustment; its application has certainly changed our views on how the natural world is organized and the relationship between process and pattern. A second refocusing directed the attention of theoreticians from the relatively straightforward realm of equilibrial situations toward those in which transitional stages are important and outcomes are context dependent. Caswell (1978) caught the essence of this shift

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from closed to open systems, and equilibrial to nonequilibrial dynamics. It remains surprising to me that this change was initially lead by theoreticians, for instance, Skellam (1951), Horn and MacArthur (1972), and Levin (1974). If there is a balance of nature, it is difficult to recognize (Pimm 1991). We inhabit and observe a dynamic and changing world. One response has been a new area of ecology, metapopulations, a term coined by Levins (1969) and, since, broadly applied because it permits local extinction, emphasizes the fundamental importance of dispersal, and is an increasingly important cornerstone of conservation biology. This is a conceptual advance and unlikely to generate a true revolution. Its genesis, like most other ecological changes, was produced by recognizing the reality of complexity in the natural world and reacting to it. I conclude with two predictions of future shifts in emphasis. Both will influence how we think about the organic universe but will probably alter few fundamental concepts. First, Gee (1999) has suggested that cladistics, the identification of statistically unbiased evolutionary relationships between species, is a developing area. I concur. Cladistics bears substantive implications for niche interrelationships and biogeography. Second, ecologists have long been aware of the disarming yet unrealistic assumptions of Lotka-Volterra dynamics, those mass-action equations in which species behave like randomly interacting molecules. Spatially explicit individualistic models concerning very local domains can provide an alternative (Levin 1999). The consequences will be sweeping, more in the way field empirical ecologists derive inspiration from theory and design experiments to strengthen or challenge the relationship, than in the normal off-hand dismissal of theoretical predictions which are untestable and seem unrealistic. Ecology as a discipline has evolved rapidly in concert with increasingly altered ecosystems (Lawton 2000). Ecological variation provided a cornerstone for Darwin and remains an immutable and unchallengeable reality. Ecology is awash in a sea of facts and factoids: their explanation requires a conceptual foundation that must, eventually, encompass much that remains currently elusive. These numerous details inspire theoretical construction and at the same time frustrate empiricists. They also lend stability to central ideas and diminish the possibility of radical new conceptual advances. Thus temporary research frontiers or bandwagons should continue to wax or wane as ecologists expand their understanding of nature's complexity. That fact alone insulates us from the intellectual trauma of sweeping paradigm shifts, and should enhance a capacity to describe as quantitatively as possible long recognized features and dynamics common to populations, communities, and ecosystems.
ACKNOWLEDGMENTS

Makah Tribal Council and the U.S. Coast Guard. Words cannot express my gratitude for the privilege of working at this unique site. Financial support has come from the National Science and Mellon Foundations. My personal wrestling match with a small corner of the philosophy of science has not been easy. My colleagues Jennifer Ruesink and Shahid Naeem have been provocative foils; Tom Daniel graciously provided the Fourier time-series analysis. Last, I thank the Western Society of Naturalists for the opportunity to express my heterodox views and the Hall Family Educational Trust for funding my participation.
LITERATURE CITED

The centerpiece of my contribution, the very long-term measurements at Tatoosh, is based on permissions from the

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