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Advances in Ecological Understanding: By Kuhnian Revolution or Conceptual Evolution?

Author(s): Robert T. PaineSource: Ecology, Vol. 83, No. 6 (Jun., 2002), pp. 1553-1559Published by: Ecological Society of AmericaStable URL: http://www.jstor.org/stable/3071973.

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Ecology, 83(6), 2002, pp. 1553-1559C 2002 by the Ecological Society of America

ADVANCES IN ECOLOGICAL UNDERSTANDING: BY KUHNIANREVOLUTION OR CONCEPTUAL EVOLUTION?ROBERT T. PAINE

Department of Zoology, University of Washington, Seattle, Washington 98195-1800 USAAbstract. Ecology, although a rapidly evolving science, has experienced few if anyparadigm shifts or revolutions in the sense of Thomas Kuhn because it is a pluralistic,multiple-causation discipline. Instead, what appears to characterize conceptual developmentare 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 worldwhich cannot be denied. I identify and discuss two: postlarval recruitment to an establishedpopulation and physical disturbance to an intertidal assemblage. The richness of ecologicalthought has been broadened immeasurably by acceptance of the following themes: thenecessity to distinguish open from closed systems, that equilibrium may exist only in anabstract sense, and that controlled manipulation can reveal hitherto unanticipated dynamics.These findings have expanded our understanding of nature's complexity (a conceptualevolution) rather than relegated most prior generalizations to the dustbin of history (a

paradigm shift).Key words: disturbance; ecological history; intertidal; Kuhnian revolution; mussel; Mytiluscalifornianus; paradigm; recruitment; rocky shore; Semibalanus cariosus; settlement.

INTRODUCTIONParadigm dramatically entered the active vocab-

ulary of science with the publication of Kuhn's TheStructure of Scientific Revolutions (1962). The conceptof normal science punctuated by revolts led by young,disenchanted enthusiasts (Wade 1977, Wilkins 1996)provides definitive hallmarks for identifying paradigmshifts or revolutions. The interpretational difficulty ofwhat constitutes a legitimate revolution is enormousand depends critically on whether Kuhn's examples,drawn from the physical sciences, can apply to biology.Subtle definitional challenges abound: How does onecharacterize a revolution or paradigm shift? What is itsgenesis? How sweeping must it be? However, I believethere is a more fundamental dichotomy. Concepts inthe physical sciences, Kuhn's subject matter, are pre-disposed to contradiction by single well-conceived ob-servations or experiments. The linear or binary natureof such constructs permits challenge (Strohman 1997).Thus, revolutions are not difficult to recognize in otherdisciplines: Copernicus' heliocentric theory, the redis-covery of Mendel and particulate inheritance, the the-ory of continental drift, and Michelson-Morley's dis-proving of the existence of an ether permeating theentire universe. At times the disciplinary restructuringor redirecting was abrupt and relatively rapid (Mi-chelson-Morley), at other times, the revolution sim-ply catalyzed a gradual replacement of the older views(Copernicus).

On the other hand, the biological sciences at levelsManuscriptreceived 3 November 2000; revised 16 July 2001;accepted26 July 2001; Special Featureaccepted7 October 2001.For reprintsof this Special Feature, see footnote 1, p. 1479.

of the organism and above are plagued by multiplecausation, generating what McIntosh (1987) describesas ecology's pluralism. Thus, I will argue, paradigmshifts are apt to be less obvious in our field, if theytruly exist at all. For instance, did Elton (1927) changeecology with his concept of food cycle (a.k.a. foodweb)? I believe the answer is no. Brilliant tour de forcethat his book was, it is probably best described as animportant advance in a lengthy quest to understandnatural complexity. Darwin (1859) had previously ex-pressed a deep interest in such issues and the multi-plicity of associated interactions. Slightly later, Forbes(1877) openly described the importance of interspecificrelationships. Thinking about trophic structure has con-tinued to evolve (e.g., Lindeman 1942, Cohen 1978,Pimm 1982). This 100-plus year quest for understand-ing is best described as a progression but one that isuneven 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 profusionare fads or bandwagons. Bandwagon is defined(Gove 1971) as a party, faction, or other element thatattracts adherents by its timeliness, vigor, showman-ship, or novelty. I do not employ the term in a pe-jorative sense: these are transient foci of interest, usu-ally only temporarily in vogue. Table 1 provides mypartial list of bandwagons in ecology, assembled bycomparing early texts (Chapman 1931, Allee et al.1949) with their modern counterparts (Begon et al.1990, Krebs 1994). I searched these texts for com-monalities, with special sensitivity to novel additionsand shifts in emphasis in an effort to calibrate my ownperspective. My list includes subjects identified as ma-

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1554 ROBERT T. PAINE Ecology, Vol. 83, No. 6TABLE 1. Some bandwagons in ecology.

Major themes Sub-themesNiches Null models

OptkmalityPopulation regulationFood webs Trophic dynamics

Web complexityInteraction strengthsIndirect effectsSuccession DisturbanceStability and diversity Species-abundance patternsEquilibrium/nonequilibriumMetapopulationsEcosystems Ecological efficienciesFunctional redundancyServices

jor themes, each of which incorporate relatively minor,more specialized components. Collectively, they reflectsubstantial refinement and progress unpunctuated byrevolts.

Thus, a bandwagon is a temporarily popular focusor research agenda, sort of an ecological supernovawhich bursts into prominence and then fades with time.No one doubts their reality or importance. Interest usu-ally wanes because technological limits have beenreached (energetics and bomb calorimetry), theory de-velopment has outpaced data acquisition (the stability-diversity relationship), or fatal conceptual flaws areuncovered (communities as superorganisms). On theother hand, paradigm shifts require substantive pre-cursors (i.e., a belief system to be overthrown). Ar-gument exists whether evolutionary change driven bynatural selection qualifies as a paradigm shift (Wilkins1996). The Watson-Crick model for the structure ofDNA does not: it was a de novo discovery in a con-tinuum of change, and was not opposed by an en-trenched group of elder scientists (Wilkins 1996, Stroh-man 1997).All bandwagons identified in Table 1 are deeply root-ed in ecological reality, i.e., they are not hypothesized,invisible constructs. Although my list is certainly in-complete, the omitted topics largely reflect difficultyin defining what constitutes a bandwagon as opposedto continuing background science. Designer assem-blages (Ruesink and Srivastava 2001) are experimen-tally popular now. Will they persist or expand in use,or will their artificial construction lead to questionsabout the robustness of the results? Austin (1999) iden-tified a historical dichotomy between botanical inter-ests in gradient analyses and ordination on one hand,and more mechanistic experimental pursuits by animalecologists on the other, as a difference between para-digms. Are these differences legitimate or are they sim-ply constraints or advantages imposed by the researchmaterial? I would have described these as tactical dif-ferences because the methodology, not the fundamentalquestion, differs. My list reflects personal biases and

ways of organizing and subdividing some of the myriadsubjects that compose ecology and generate its plural-istic nature. Thus, species do appear to have individ-ualistic lifestyles (niches), to replace one anotherthrough time (succession), to eat and be eaten (foodwebs), and to inhabit environments influenced by bothphysical and biological stressors (ecosystems). Our vo-cabulary is of necessity general and not precise; theterm bandwagon is intended to catch the essence orflavor of a topic of interest, whether it passes quietlyor persists. For instance, ideas on change (succession)began with Warming (1896) and Cowles (1899), werecodified by Clements (1916), overextended and debated(Phillips 1934, Tansley 1935), mechanistically ex-panded (Drury and Nisbet 1973, Connell and Slatyer1977), and remain the subject of intense investigation(Turner and Dale 1998). I cannot conceive of a revo-lution that could or will replace our increasingly so-phisticated and mechanistically detailed understandingof how ecosystems respond to disruption. The sametrajectories of refinement and increased appreciationcan be identified for ecological niches, food web con-struction, foraging of organisms, variation in speciesrichness, factors influencing primary and secondaryproduction, and even global warming and sea-levelchange. All share a common biological heritage: theenormous complexity of natural systems. They alsoshare the following trait: multiple causation or expla-nation is commonplace and unavoidable, and thus nu-merous alternative hypotheses are the norm for dis-secting patterns. This fact, by itself, tends to insulategeneralizable (and popular) ecological views from rad-ical changes or scientific revolutions. Because ecologyis not a linear discipline, breaking or challenging onelogical strand is unlikely to generate a major overhaul.In this sense MacArthur's (1955) views on stabilityshould also apply to paradigm shifts.In the following paragraphs I provide two examplesof bandwagon development based on personal involve-ment and interest, and I argue that they are so funda-mentally and naturally important that they will not, andcannot, be replaced. Detailed refinement, not revolu-tionary displacement, is the sole alternative.

RECRUITMENT LIMITATIONIndividual species populations grow or shrink, reach

nuisance proportion or go extinct, based on their re-productive potential and their survival. Limitation ofeffective reproductive output was a key component ofDarwin's views on natural selection. The topic ofspawner:recruitratios became central to fishery biologymore than a century ago (Sinclair 1988). The primaryissue then, as now, was whether the reproductive outputof a stock could be related quantitatively to the numberof individuals recruiting to that stock after their pro-longed exposure to a host of unknown mortality factors.The relative contributions of pre- and postrecruitmentmortality have important implications for management



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FIG. 1. Interannual ariation n recruitment f the barnacleSemibalanus ariosus at Tatoosh sland,Washington,USA.Densitywas estimatedonce postsettlementndividualswerelarge enoughto be identified.Dashed ines representhe 18 and90 individualsper100cm2 evels of replacement assumingzeropostrecruitment ortality) equiredo saturate hesubstratumif therewere, respectively, ive yearsand one yearof recruitment.

and harvest schedules and for conservation. I considerthe issue to be a bandwagon because of its timelinessand continuing importance, and one unlikely to be sub-ject to a scientific revolution.The opportunity to test this view occurred in the early1980s when recruitment was rediscovered (Underwoodet al. 1983, Connell 1985, Gaines and Roughgarden1985), named supply-side ecology (Lewin 1986),and advertised as a fatal and missing flaw in under-standing rocky intertidal shore dynamics (Underwoodet al. 1983). One reality is certain: populations willinexorably go extinct, locally and globally, without re-placement. Effective reproduction is thus essential, andstudies which ignore the dynamics of replacementshould be suspect. The issue has been especially dra-matic in marine ecology, where many species releasegametes into a water mass of unknown dimension, withunknown (and perhaps unlikely) fertilization success,spend variable dispersal intervals in currents onlyvaguely described, and then settle onto the shore. Set-tlement often involves a dramatic change in morphol-ogy, lifestyle, and sources of mortality. Recruitmentoccurs when specific identity can be determined; thistechnically demanding determination often is madesome time after settlement, during which interval thesettled population has been subject to numerous sourc-es of mortality when it is maximally vulnerable (Caleyet al. 1996). Employing this operationally effective def-inition, however, impairs the ability to determine the

relative significance of pre- vs. postsettlement pro-cesses.The settlement/recruitment phase involves much ba-sic biology. The details vary from species to speciesdepending on life history traits, with mobile species

being potentially more vexing subjects than sessile in-vertebrates. The fundamental question, whether it isconditions during the transport phase in the ocean orpostsettlement dynamics which determine the characterof the postsettlement population, is crucial. Becauseannual or subannual species are apt to spend a greaterfraction of their lifetime being subject to the vagariesof transport, I would anticipate an enhanced effect ofpresettlement factors. For perennials that conclusion ismuch less certain and depends on their potential lon-gevity since, for instance, perenniality (Cole 1954) andstorage effects (Warnerand Chesson 1985) can insulatethe species from years of near-zero reproductive output.As an example (Fig. 1), I have measured since 1978the recruitment of the barnacle Semibalanus cariosusin newly formed patches in mussel beds at TatooshIsland, Washington, USA. The pattern is one of enor-mous temporal variation; some years are excessive,while others fail to attain a replacement level. If thisspecies were an annual, it would require -90 recruitsper 100 cm2 to saturate that surface after one year, givenperfect survival (i.e., no benthic interactions leadingto individual death) and known growth rates (i.e., Day-ton 1971, Paine and Levin 1981). That level of re-

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Ecology,Vol. 83, No. 6cruitment is achieved in only eight of the 22 yr, pre-sumably eventually dooming an annual species. S. car-iosus, however, is not an annual and possibly lives -20yr (Connell 1972, Sebens and Lewis 1985). If thesebarnacles reproduce in only five of these years, a veryconservative estimate, the recruitment must be 18 bar-nacles per year on average, again without mortality, tosaturate the space. This level is exceeded in 15 of the22 years S. cariosus, like redwood trees, is not eco-logically threatened by occasional settlement/recruit-ment failures.

Two attributes of the recruitment limitation issue re-main unresolved. First, massive, benthic-based mor-tality must often occur before recruitment and aftersettlement (e.g., Wethey 1984). This mortality willhave nothing to do with the planktonic interval, itsexistence is experimentally testable, and for perennialspecies, it challenges the notion of primary determi-nation of adult population traits by events during thedispersal phase. As a related aside, the excessive mor-tality characteristic of postrecruitment crowding due tosettlement binges (Barnes and Powell 1950), capableof eliminating entire recruit classes, should furthercloud the recruit-spawner relationship. Second, be-cause variation in recruitment has been recognized fordecades (Hjort 1914, Thorson 1946), and is totally nat-ural, there can be no paradigm shifts. There can berefinements, increased mechanistic understanding, bet-ter modeling of the transition from planktonic to ben-thic existence, or increased appreciation of the relativecontributions to population dynamics of mortality dur-ing the transportand postsettlement phases for our the-ories of variation in recruitment, but radical restruc-turing of our views is not possible. Thus, importantrefinements (i.e., conceptual evolution) should contin-ue to characterize our interpretation of life history dy-namics spanning two very different environments.

DISTURBANCEField ecologists have probably always been awareof change, and certainly our discipline's first major

theory, the notions of primary and secondary succes-sion (Warming 1896, Cowles 1899, Clements 1905,1916) were developed in the context of change. Classicpublications such as Watt's (1947) pattern and processpaper identified cyclical dynamics and therefore no sin-gle general termination to a successional sequence. Be-cause the initial focus was on terrestrial landscapes, itshould come as no surprise that fire was broadly ac-knowledged as a generating pattern. Heinselman's(1973) graphic maps suggested how fire influenced for-ests in Minnesota, and studies in Yellowstone NationalPark (e.g., Taylor 1973) helped to change our nationalfire suppression mania. Even earlier, Turner(1935) hadattributed localized age-structure difference in pines toa region's tornado history. Perhaps the culmination ofthis trend was Pickett and White's (1985) book, TheEcology of Natural Disturbance and Patch Dynamics.

Physical disturbances to landscapes continue to attractthe attention of ecologists, and it seems certain that thisbandwagon has not run its due course. The primaryreasons are obvious: disturbance, perhaps especiallywhen ultimately expressed as global change, is an in-exorable fact of life, and ecologists are still on the steepportion of their learning curve (Turnerand Dale 1998).The following paragraphsexplore one dimension ofdisturbance ecology in an intensely studied rocky in-tertidal system. The Cape Flattery region of Washing-ton State in general, and Tatoosh Island in particular,has been the venue of a variety of such studies (Dayton1971, Levin and Paine 1974, Paine and Levin 1981,Dethier 1984). Beds of the mussel Mytilus californi-anus owe their internal spatial patterns to waves whichremove mussels and thus produce gaps. The gaps orpatches are born (disturbance) and die (recovery)at rates which are patch-size dependent. Levin andPaine (1974) and Paine and Levin (1981) modeled thissystem, assuming that settlement into patches was in-dependent of events occurring to mussel larvae duringtheir planktonic interval, that the individual musselbeds had no defining history, and therefore that dis-turbance intensity during a given winter interval wasindependent of that site's local history. The latter as-sumption seemed realistic, and permitted averagingdisturbance traits (gap size at formation) across manysites.

Long-term observations suggest that these assump-tions were premature. Disruption to 26 independent(spatially distinct) mussel beds has either been mea-sured directly or estimated each spring since 1974. Ear-lier data in this time series are relatively sparse. Fig.2 presents the mean annual disturbance for 25 sitesscattered around the south, west, and north peripheryof Tatoosh Island. A site from which the starfish Pi-saster ochraceus was removed was deleted from thetally. The original data, expressed as the percentage ofmussel bed removed each winter, were arcsine square-root transformed, averaged for the island, and then backtransformed. Visual inspection suggests a pattern ofpeaks and troughs. A cold water winter in 1975-1976,extreme El Nifio conditions during the winter of 1982-1983, and a brief but severe February cold snap whichkilled almost 15% of the island's mussels in 1988-1989stand out in the data. The 1982-1983 event was par-ticularly destructive to mussels close to their upperlimit, mussels which are always small (


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FIG. 2. Interannual variation in the proportion of the mussel Mytilus californianus removed during winter disturbancesfrom 25 spatially isolated sites at Tatoosh Island. Errorbars represent + 1 SE and are based on arcsine square-root transformeddata.

of 8, 7, 6, 1, 7, and 2 yr. Fourier transformation andanalysis reveal a slow peak at 7.7-yr intervals and afaster one at 2.3-yr intervals.

What do these data suggest about the disturbanceregime to mussels on Tatoosh? First, they share nocommon environmental cause. Some El Niio intervalsare characterized by major disturbance (>13%) where-as others are not (1997-1998, early 1990s). Severestorms and deep freezes also generate major mortalityevents. If one dismisses the 1989 freeze because itprimarily impacted small mussels usually invulnerableto wave stresses, and accepts the photographic evidenceof 1968, the five peaks average 6.2 yr apart. This meanis reflected in the time series identification of a 7.7-yrperiodicity and an estimate of a minimal rotation periodof 7-10 yr in Paine and Levin (1981). This periodicityimplies a temporal pattern to disturbance and that yearsare not truly independent of one another. The under-lying biological reason is that as mussels grow, theirsusceptibility to wave force increases substantially, andbeyond some size the probability of destruction, onintertidal wave-swept shores approaches unity (Dennyet al. 1985). The 2.3-yr periodicity remains unex-plained.

Can such revelations alter the reality that winterwaves carve patches in outer coast mussel beds? Cer-tainly not. At best the realization will help expand andcalibrate more sophisticated spatial models dealingwith the endless cycles of disruption and recoverywhich generate the mosaic of pattern so essential toand characteristic of these environments. In a broadercontext, disturbance processes, whether marine or ter-restrial, interruptsuccessional trajectories. New and in-teresting biological detail and model refinement willcontinue to characterize this fertile subdivision of ecol-

ogy. These will fine-tune this bandwagon (reducingsqueaks in the wheels?); they will not overturn the cartwhich seems robust in its diversity and central to muchof ecology.

AN ALTERNATIVE VIEWI have reasoned that because ecology in the broadest

sense is a pluralistic, multiple-causation discipline, rev-olutions are unlikely. That does not imply that con-ceptual changes have not occurred. The clearest ex-ample of a possible legitimate paradigm shift knownto me is MacArthur and Wilson's (1963, 1967) theoryof island biogeography. Their hypotheses meet most ofKuhn's criteria for revolution: the authors' relativelyyouthful ages and the redirection of an important focusof ecology (biogeography) which broke from the priorstandard analysis of an island's area and assessmentsof its biota to suggest dynamical alternatives.

There have been other shifts, but they are more tac-tical and general rather than specific. Two have beenin progress for decades. Connell (1961), Paine (1966),and Dayton (1971) introduced the notion (though hard-ly for the first time; see Paine 1994 for details) thatfield experiments could reveal subtle yet important dy-namics influencing species distribution and abundance.Experimental, manipulation-based hypothesis testing isnow commonplace. This approach represents an atti-tude adjustment; its application has certainly changedour views on how the natural world is organized andthe relationship between process and pattern.

A second refocusing directed the attention of theo-reticians from the relatively straightforward realm ofequilibrial situations toward those in which transitionalstages are important and outcomes are context depen-dent. Caswell (1978) caught the essence of this shift



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1558 ROBERT T. PAINE Ecology, Vol. 83, No. 6

from closed to open systems, and equilibrial to non-equilibrial dynamics. It remains surprising to me thatthis change was initially lead by theoreticians, for in-stance, Skellam (1951), Horn and MacArthur (1972),and Levin (1974). If there is a balance of nature, it isdifficult to recognize (Pimm 1991). We inhabit andobserve a dynamic and changing world. One responsehas been a new area of ecology, metapopulations, aterm coined by Levins (1969) and, since, broadly ap-plied because it permits local extinction, emphasizesthe fundamental importance of dispersal, and is an in-creasingly important cornerstone of conservation bi-ology. This is a conceptual advance and unlikely togenerate a true revolution. Its genesis, like most otherecological changes, was produced by recognizing thereality of complexity in the natural world and reactingto it.

I conclude with two predictions of future shifts inemphasis. Both will influence how we think about theorganic universe but will probably alter few funda-mental concepts. First, Gee (1999) has suggested thatcladistics, the identification of statistically unbiasedevolutionary relationships between species, is a de-veloping area. I concur. Cladistics bears substantiveimplications for niche interrelationships and biogeog-raphy. Second, ecologists have long been aware of thedisarming yet unrealistic assumptions of Lotka-Vol-terra dynamics, those mass-action equations in whichspecies behave like randomly interacting molecules.Spatially explicit individualistic models concerningvery local domains can provide an alternative (Levin1999). The consequences will be sweeping, more in theway field empirical ecologists derive inspiration fromtheory and design experiments to strengthen or chal-lenge the relationship, than in the normal off-hand dis-missal of theoretical predictions which are untestableand seem unrealistic.

Ecology as a discipline has evolved rapidly in con-cert with increasingly altered ecosystems (Lawton2000). Ecological variation provided a cornerstone forDarwin and remains an immutable and unchallengeablereality. Ecology is awash in a sea of facts and factoids:their explanation requires a conceptual foundation thatmust, eventually, encompass much that remains cur-rently elusive. These numerous details inspire theoret-ical construction and at the same time frustrate empir-icists. They also lend stability to central ideas and di-minish the possibility of radical new conceptual ad-vances. Thus temporary research frontiers orbandwagons should continue to wax or wane as ecol-ogists expand their understanding of nature's com-plexity. That fact alone insulates us from the intellec-tual trauma of sweeping paradigm shifts, and shouldenhance a capacity to describe as quantitatively as pos-sible long recognized features and dynamics commonto populations, communities, and ecosystems.

ACKNOWLEDGMENTSThe centerpiece of my contribution, the very long-termmeasurements at Tatoosh, is based on permissions from the

Makah Tribal Council and the U.S. Coast Guard. Words can-not express my gratitude for the privilege of working at thisunique site. Financial support has come from the NationalScience and Mellon Foundations. My personal wrestlingmatch with a small corner of the philosophy of science hasnot been easy. My colleagues Jennifer Ruesink and ShahidNaeem have been provocative foils; Tom Daniel graciouslyprovided the Fourier time-series analysis. Last, I thank theWestern Society of Naturalists for the opportunity to expressmy heterodox views and the Hall Family Educational Trustfor funding my participation.LITERATURE CITED

Allee, W. C., A. E. Emerson, 0. Park, T. Park, and K. P.Schmidt. 1949. Principles of animal ecology. W. B. Saun-ders, Philadelphia, Pennsylvania, USA.Austin, M. P. 1999. A silent clash of paradigms: some in-consistencies in community ecology. Oikos 86:170-178.Barnes, H., and H. T. Powell. 1950. General morphology andsubsequent elimination of barnacle populations, Balanuscrenatus and B. balanoides, after a heavy initial settlement.Journal of Animal Ecology 19:175-179.Begon, M., J. L. Harper, and C. R. Townsend. 1990. Ecology.Blackwell Scientific, Oxford, UK.Caley, M. J., M. H. Carr, M. A. Hixon, T. P. Hughes, G. P.Jones, and B. A. Menge. 1996. Recruitment and the localdynamics of open marine populations. Annual Review ofEcology and Systematics 27:477-500.Caswell, H. 1978. Predator-mediated coexistence: a non-equilibrium model. American Naturalist 112:127-154.Chapman, R. N. 1931. Animal ecology. McGraw-Hill, NewYork, New York, USA.Clements, F E. 1905. Research methods in ecology. Uni-versity Publishing, Lincoln, Nebraska, USA.Clements, F E. 1916. Plant succession: an analysis of thedevelopment of vegetation. Carnegie Institution of Wash-ington 242:1-512.Cohen, J. E. 1978. Food webs and niche space. PrincetonUniversity Press, Princeton, New Jersey, USA.Cole, L. C. 1954. The population consequences of life history

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