Introduction: Between Ecology and Evolutionary BiologyAuthor(s): James P. Collins, John Beatty and Jane MaienscheinSource: Journal of the History of Biology, Vol. 19, No. 2, Reflections on Ecology andEvolution (Summer, 1986), pp. 169-180Published by: SpringerStable URL: http://www.jstor.org/stable/4330971Accessed: 29-05-2018 17:55 UTCJSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a widerange of content in a trusted digital archive. We use information technology and tools to increase productivity andfacilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available athttp://about.jstor.org/termsSpringer is collaborating with JSTOR to digitize, preserve and extend access to Journal ofthe History of BiologyThis content downloaded from 129.219.247.33 on Tue, 29 May 2018 17:55:39 UTCAll use subject to http://about.jstor.org/terms
Introduction: Between Ecology andEvolutionary BiologyJAMES P. COLLINSDepartment of ZoologyArizona State UniversityTempe, Arizona 85287JOHN BEATTYDepartment of Ecology and Behavioral BiologyUniversity of MinnesotaMinneapolis, Minnesota 55455JANE MAIENSCHEINDepartment of PhilosophyArizona State UniversityTempe, Arizona 85287Ecology emerged as a self-conscious discipline during the lastdecade of the nineteenth century, growing out of a heterogeneousmix of fi'elds. Indeed, its roots are as different as field naturalhistory and expefimental physiology.' Given that ecology was (andremains) such a heterogeneous enterprise, it is unlikely that anysingle perspective will suffice to describe its history. Several verydifferent approaches have already proved fruitful. Ronald Tobeyhas discussed the changing importance of applied versus nonmission-oriented research in the development of ecology. SharonKingsland has emphasized the relative merits of theoretical versusempirical research at various points in the history of the discipline.Robert McIntosh has taken the very heterogeneity of the disciplineas his perspective on its history.2 The papers that follow approachthe history of ecology from yet another standpoint, namely, thechanging role of evolutionary theory in the solution of ecologicalproblems.In On the Origin of Species Darwin frequently brought evolu1. Robert McIntosh, 7The Background of' L coloy (Cambridge: UniversityPress. 1 985), p. 26.2. Ronald C. Tobey, Saving the P'rairies: 7he Life Cycle of the FoundingSchool of American Plant Ecology, 1895-1955 (Berkeley: University of Cali-fornia Press, 1981); Sharon E. Kingsland, Modeling Nature: E;pisodes in theHistory of lopulation Ecology (Chicago: University of Chicago Press, 1985);McIntosh, Baic-kgroundtd. Sec also R. Colwell, "'The Evolution of Ecology," Amer.Zool., 25 (1985), 771-777.Journal of the History of Biology, Vol. 19, No. 2 (Summer 1 986), pp. 169-180.C) 1986 by D. Reidel Publishing Company.This content downloaded from 129.219.247.33 on Tue, 29 May 2018 17:55:39 UTCAll use subject to http://about.jstor.org/terms
COLLINS, BEATIrY, AND MAIENSCHEINtionary considerations to bear on the distribution and abundanceof organisms, and on the relations of organisms to their environments. Inasmuch as these areas eventually constituted the domainof ecology, Darwin is often cited as an early advocate of evolu-tionary approaches to ecological problems. Not all subsequentecologists have followed Darwin in this regard. We asked theauthors of the following papers to consider why ecologists sometimes have found evolutionary concepts significant for theirresearch and sometimes have not. In addition, some of ourcontributors have explored the degree to which evolutionarybiologists have sometimes and sometimes not found ecologicalconcepts significant.The ensuing articles illustrate how complicated the answers tosuch questions can be. The complications undoubtedly reflect thediffuse and changing identity of ecology in the late nineteenth andtwentieth centuries. Historians and philosophers are hardly inagreement on what constitutes a discipline, much less on howdisciplines emerge and establish their indentities. The difficultiesare compounded when it comes to studying the history of a youngdiscipline like ecology.During the late nineteenth century biologists most often soughtexplanations for the distribution and abundance of organismsthrough studies of natural history - especially biogeography.Those explanations were historical and evolutionary, and fre-quently incorporated the concept of adaptation via natural selec-tion. By the very end of the century, however, botanical biogeographers in particular had begun to offer mechanistic, ratherthan historical, explanations for the distributions of species. Theirobjective was to account for distribution and abundance of plantsnot in terms of the evolutionary histories of those plants, but moredirectly, in terms of the physiological abilities of those plants toadjust to some environments but not to others.Eugenius Warming was an early proponent of this new approach to the study of plant distribution - an approach that hecalled ecological plant geography. In his paper here WilliamColeman summarizes Warming's vision of this new geography as adeparture from the more common methods of the time.- An3. William Coleman, "Evolution into Ecology? The Strategy of Warming'sEcological Plant geography," J. Hist. Biol., this issue.170This content downloaded from 129.219.247.33 on Tue, 29 May 2018 17:55:39 UTCAll use subject to http://about.jstor.org/terms
Introduction: Between Ecology and Evolutionary Biologyecological approach to plant distribution supposedly provided abetter explanation of community structure. According to thephysiological approach, communities of plants in different parts ofa country, or even the world, might look quite similar (have asimilar physiognomy) if they grew in comparable environments.Although having different evolutionary histories, species of plantsin a community would look alike because environments withsimilar biotic and abiotic attributes would favor plants withcomparable physiological capacities. Evolutionary history was not,therefore, of as much consequence for the study of communitystructure as was physiology. The concept of adaptation to theenvironment was very useful for Warming, although he was notmuch concerned with the historical, evolutionary process leadingto it. For Warming ecology was the science of understanding howphysiological plant-environment and plant-plant interactions re-sulted in groupings of organisms into communities that wereunevenly distributed geographically.The physiological relationship between plant and environmentprovided an important theoretical framework within which thenew science of ecology could begin to develop and distinguishitself from the closely allied discipline of biogeography. Bothrelied on the background perspective provided by the concept ofadaptation. But the biogeographers were more interested in theevolution of adaptation. In particular, biogeographic explanationsfor distribution and abundance depended on adaptive divergenceof taxa within a region, and on subsequent migration. Fundamentally, such biogeographic explanations were historical and evolutionary. In contrast, ecologists argued that present environmentalconditions could be invoked to account for plant distribution andabundance, that is, in a manner more analogous to a physiologicalexplanation.Joel Hagen also discusses the physiological roots of ecology,though not so much to point out the difference between physiological and historical approaches as to emphasize the lure ofphysiology's experimental approach.4 Early ecologists such asWarming, Andreas Schimper, and Fredenrc Clements urged ecologists to use the rigorous, experimental methods of physiology4. Joel B. Hagen, Ecologists and Taxonomists: Divergent Traditions inTwentieth-Century Plant Geography," J. Hlist. Biol., this issue. See also EugeneCittadino, "Ecology and the Professionalization of Botany in America, lX8901905," Stud. Hist. Biol. 4(1980), 1 71-198.171This content downloaded from 129.219.247.33 on Tue, 29 May 2018 17:55:39 UTCAll use subject to http://about.jstor.org/terms
COLLINS, BEATTY, AND MAIENSCHEINeffectively making ecology field physiology. Their new science wasto be quantitative and experimental following the model providedby physiology, a model highly regarded by biologists of the time.As Garland Allen has argued, there was at the turn of the centurya dramatic movement toward the use of experiments as exemplified by physiology. While the revolutionary nature of this shiftremains debatable, it seems clear that more biologists, includingecologists, began to use experiments in their research.5 Hagendemonstrates the importance of this experimentation, but showsalso that this new approach did not completely replace traditionalplant geographies, which continued to rely on historically orientedexplanations.The physiological interests of many early ecologists are wellreflected, as Coleman and Hagen both note, in the "superorganis-mic" analogies that these researchers used when characterizingcommunities. Just as traditional physiology attended to the development and functioning of individual organisms, these early eco-logists treated the development and functioning of individual,superorganismic communities.We have thus far discussed only community-level investigations.One might expect to find more interest in Darwinian evolutionarytheory among those investigating the ecology of individual populations and species than among those studying communities.Populations and species are, after all, the things that evolve bynatural selection. If evolutionary biology is going to be relevant toecology, one would expect to find it at least in population ecology.For the first two decades of this century, however, Darwinianevolutionary theory was under considerable attack.6 In particular,it was seriously doubted whether evolutionary change could occuras a result of natural selection of small mutations. A number ofalternative evolutionary agents, the most prominent includinglarge-scale and directed mutations, were accorded great influence.So it is not surprising that evolution by natural selection wasnot the favored mode of explanation of population growth andcomposition.5. Garland E. Allen, Life Science in the Twentieth Century (Cambridge:Cambridge University Press, 1975); Jane Maienschein, Ronald Rainger, andKeith Benson, "Introduction: Were American Morphologists in Revolt?" J. Hist.Biol., 14 (1981); 83-87.6. Peter Bowler, Eclipse of Darwinism (Baltimore: Johns Hopkins UniversityPress, 1983).172This content downloaded from 129.219.247.33 on Tue, 29 May 2018 17:55:39 UTCAll use subject to http://about.jstor.org/terms
Introduction: Between Ecology and Evolutionary BiologyThere were notable exceptions. Among the several topicstreated in his paper, William Kimler discusses the career ofEdward Poulton, an entomologist and a staunch defender ofDarwinism during the first decades of this century, when this wasnot such a popular position.7 Poulton saw the science of ecology asa means of promoting studies of the role of natural selection inaccounting for population composition, specifically intraspecificvariation. He was especially interested in intraspecific differencesin mimicry, which he denied were due simply to mutation pres-sure. He attributed the maintenance of such differences to selection instead. For Poulton understanding the selection pressuresinvolved was an ecological problem.Kimler's point is not that Poulton was representative of thegeneral influence that Darwinism had on ecology in the firstdecades of the century. Poulton was exceptional in this respect.But he serves as a contrasting case, to show what sort of influenceDarwinism might have had (and what influence it eventually had)in distinction to what prevailed at the time. After Poulton there developed in Britain a strongly pro-Darwinist school of ecologicalgenetics. Among its members were R. A. Fisher, E. B. Ford, A. J.Cain, and P. M. Shepard. The Darwinian character of the Britishschool was also exceptional in its time, a point to which we shallreturn shortly.So-called theoretical population ecology developed considerably in the twenties and thirties, still prior to the general acceptance of selection of small mutations as a viable mode ofevolutionary change. That population ecology should not haveassumed a Darwinian character in light of the period of its initialdevelopment is understandable. The reasons for that fact are com-plicated, however, as Sharon Kingsland explains in her article.8The attitude toward evolutionary theory differed among earlypopulation ecologists. W. R. Thompson, for one, positively eschewed evolutionary approaches to population dynamics and evengrew suspicious of mathematical approaches to the subject because of the success of Fisher's mathematical population genetics,which in turn supported the importance of natural selection, which7. William C. Kimler, "Advantage, Adaptiveness, and Evolutionary Ecology,"J. Hist. Biol., this issue.8. Sharon E. Kingsland, "Mathematical Figments, Biological Facts: Population Ecology in the Thirties," J. Hist. Biol., this issue.173This content downloaded from 129.219.247.33 on Tue, 29 May 2018 17:55:39 UTCAll use subject to http://about.jstor.org/terms
COLLINS, BEAYFY, AND MAIENSCHEINThompson did not accept. Although population ecologists likeA. J. Lotka and Vito Volterra explicitly proposed to formulategeneral theories of "evolution," the equations of population growththat they (and Raymond Pearl, whom Kingsland discusses elsewhere) formulated - the equations that formed the theoreticalcore of population ecology - were hardly "evolutionary" in anystandard sense of that term.9 Truly evolutionary considerations ofpopulation dynamics (for example, notions of r-selection andK-selection) were much longer in coming.Kingsland's article helps us to understand why a Darwinianperspective was not a point of convergence among Lotka, Volterra, Pearl, Thompson, and other population ecologists like R. N.Chapman, G. F. Gause, and A. J. Nicholson. Perhaps mostimportant in this regard, these individuals had very differentbackgrounds in the sciences and very different goals. Certainlythey did not all identify themselves as ecologists. Consider, forinstance, that Thompson was an entomologist; Nicholson, also anentomologist, collaborated with V. Bailey, a physicist; Lotka was ademographer and mathematician, and Volterra a mathematician.Pearl's career is more difficult to categorize because of hisinterests in demography, biostatistics, genetics, and evolutionarytheory.The reasons that these investigators had for pursuing themathematics of population dynamics were similar only in theirshared quest to achieve some generally applicable theories. Lotkasought a law of evolution that would be as general as the laws ofthermodynamics, especially the second law. Pearl pursued ageneral law of population growth. Volterra sought to develop amathematical, general theory of evolution. Thompson wanted tounderstand in general terms the relationship between parasites andtheir hosts. In developing their theories, these researchers usedanalogies borrowed from different disciplines to motivate theirmathematical reasoning. Lotka, as is clear from the point above,drew on physical chemistry, whereas Volterra and Pearl usedanalogies from physics, especially theory concerning movement ofparticles in a gas.A significant improvement in the understanding of microevolution was accomplished in the twenties and thirties, thanks inlarge part to the pathbreaking studies of Fisher, Sewall Wright,9. Kingsland, Modeling Nature.174This content downloaded from 129.219.247.33 on Tue, 29 May 2018 17:55:39 UTCAll use subject to http://about.jstor.org/terms
Introduction: Between Ecology and Evolutionary BiologyJ. B. S. Haldane, and Sergei Chetverikov. Applications of thetheory by Theodosius Dobzhansky, Ernst Mayr, G. G. Simpson,Bernhard Rensch, and G. Ledyard Stebbins during the thirties andforties, represent the major empirical advances. This chain of developments has come to be referred to as the "evolutionarysynthesis,"'" The synthesis made it clear that natural selection ofsmall variations could be effective in directing evolutionarychange. It did not, however, establish natural selection as the chiefmeans of evolutionary change - at least, not at first. As Steven JayGould has pointed out in a number of articles, proponents of thesynthesis early on - in the thirties and early forties - attributedconsiderable significance to evolution by random drift. Only in thelater forties and fifties did the synthesis "harden," to use Gould'sterm, in favor of the all-importance of evolution by naturalselection.''William Provine, in a talk at this conference (taken in part froma forthcoming book and a previous essay), used the fact of thisfurther delay in the general acceptance of natural selection toaccount for the late impact of evolutionary thinking in ecology.'2Provine did not neglect the pro-Darwinian character of the Britishschool of ecological genetics, but pointed out that the position of1(0. William B. Provine, The Origins of Theoretical Population Genetics(Chicago: University of Chicago Press, 1971); Ernst Mayr and William B.Provine, eds., The Evolutionary Synthesis (Cambridge, Massachusetts: HarvardUniversity Press, 1980); Ernst Mayr, The Growth of Biological Thought (Cambridge, Massachusetts: Harvard University Press, 1982); Mark Adams, "Towardsa Synthesis: Population Concepts in Russian Evolutionary Thought," J. Hist.Biol., 3(1970), 107-129.11. Steven Jay Gould, "G. G. Simpson, Paleontology, and the ModernSynthesis," in Mayr and Provine, The Evolutionary Synthesis; idem, Introductionto the Columbia Classics in Evolution Series reprint of the first edition ofTheodosius Dobzhansky's Genetics and the Origin of Species (New York:Columbia University Press, 1982); idem, "The Hardening of the Synthesis," inDimensions of Darwinism, ed. Marjorie Grene (Cambridge: Cambridge University Press, 1983), pp. 71-93.12. William B. Provine, "Ecological Genetics: D)rosophila pseludoobscura,(epaea nemoralis and hortensis, and 'anaxia (lominula," unpublished paperpresented at the conference "Reflections on Ecology and Evolutionary Biology,"Arizona State University, Tempe, Arizona, March 1-2, 1985. See also idem,Sewall Wright: Geneticist and Evolutionist (Chicago: University of Chicago Press,1986); and idem, "The Development of Wright's Theory of Evolution: Systematics, Adaptation, and Drift," in Grene, Dimensions of D)arwinism, pp. 437.175This content downloaded from 129.219.247.33 on Tue, 29 May 2018 17:55:39 UTCAll use subject to http://about.jstor.org/terms
COLLINS, BEATTY, AND MAIENSCHEINthe British school was, in the thirties and forties, not the positionof general consent.In a related paper at the conference (part of a much longermanuscript in progress), Malcolm Kottler discussed the career ofthe British ecologist and evolutionist David Lack.'3 Lack's analysisof "character displacement" among the various species of Gala-pagos finches is by now a legendary example of evolutionaryecology. Lack explained the differences in beak morphology of thefinches in terms of selection for ability to utilize different foodresources - that is, in terms of the advantages of escapingcompetition for the same resources. This combination of ecological thinking (such as his use of the notion of interspecificcompetition) and evolutionary thinking (such as his use of thenotion of natural selection) has made Lack one of the mostfrequently cited evolutionary ecologists. But Lack originally considered the differences among the Galapagos finches to be amatter of random drift. Thus, he neatly
natural selection. If evolutionary biology is going to be relevant to ecology, one would expect to find it at least in population ecology. For the first two decades of this century, however, Darwinian evolutionary theory was under considerable attack.6 In particular, it was seriously doubted whether evolutionary change could occur
Department of Ecology and Evolutionary Biology Ecology and Evolutionary Biology Graduate Programs The department comprises a large number of biologists with a variety of research interests. 3 broad overlapping themes capture the interests and activities in EEB — biodiversity and macroevolution, ecology and global change biology, and .
population ecology) and then subsequently covering interactions between species in a community (i.e., community ecology). However, to facilitate completion of the final paper, I have recently switched to covering community ecology and ecosystem ecology before population ecology. As both ecology and evolution have to be covered in the same .
evolutionary biology. From this point of view, some authors have tried to extend the Darwinian theory universally beyond the domain of evolutionary biology (Cf. Dawkins, 1983), using the three principles of evolutionary theory (inheritance or retention, variation, and adaptation) as a heuristic for evolutionary economic theorizing (Campbell, 1965).
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human behaviour Natural selection 3 Box 1.1 Speciation and the evolutionary processes 4 Asking the right questions 5 Box 1.2 Reductionism vs holism 7 Approaches to the study of human behaviour 8 Human behavioural ecology 8 Evolutionary psychology 10 Box 1.3 The problem of external validity 11 Environment of evolutionary adaptedness12 Box 1.4 .
2nd Grade – Launching with . Voices in the Park by Anthony Browne (lead from the Third Voice) My First Tooth is Gone by student (student authored work from Common Core Student Work Samples) A Chair for my Mother by Vera B. William Moonlight on the River by Robert McCloskey One Morning in Maine by Robert McCloskey, Roach by Kathy (student authored work from www.readingandwritingproject.com .