Conceptions of adaptation have varied in the history of genetic Darwinism depending on whether what is taken to be focal is the process of adaptation, adapted states of populations, or discrete adaptations in individual organisms. I argue that Theodosius Dobzhansky’s view of adaptation as a dynamical process contrasts with so-called “adaptationist” views of natural selection figured as “design-without-a-designer” of relatively discrete, enumerable adaptations. Correlated with these respectively process and product oriented approaches to adaptive natural selection are divergent pictures of organisms (...) themselves as developmental wholes or as “bundles” of adaptations. While even process versions of genetical Darwinism are insufficiently sensitive to the fact much of the variation on which adaptive selection works consists of changes in the timing, rate, or location of ontogenetic events, I argue that articulations of the Modern Synthesis influenced by Dobzhansky are more easily reconciled with the recent shift to evolutionary developmentalism than are versions that make discrete adaptations central. (shrink)

Intellectual and professional reforms in evolutionary studies between 1935 and 1950 included substantial expansion, diversification, and realignment of community infrastructure. Theodosius Dobzhansky, Julian Huxley and Alfred Emerson organized the Society for the Study of Speciation at the 1939 AAAS Columbus meeting as one response to concerns about ‘isolation’ and ‘lack of contact’ among speciation workers worried about ‘dispersed’ and ‘scattered’ resources in this newly robust ‘borderline’ domain. Simply constructed, the SSS sought neither the radical reorganization of specialities nor the creation (...) of some new discipline. Instead, it was designed to facilitate: to simplify exchange of information and to provide a minimally invasive avenue for connecting disparate researchers. Emerson served as SSS secretary and was its principal agent. After publishing one block of publications, however, the SSS became ‘quiescent’. Anxious to promote his own agenda, Ernst Mayr tried to manoeuvre around Emerson in an effort to revitalize the project. After meeting impediments, he moved his efforts elsewhere. The SSS was too short-lived to merit a claim for major impact within the community; however, it reveals important features of community activity during the synthesis period and stands in contrast to later efforts by George Simpson, Dobzhansky, and Mayr. (shrink)

I propose we abandon the unit concept of "the evolutionary synthesis". There was much more to evolutionary studies in the 1920s and 1930s than is suggested in our commonplace narratives of this object in history. Instead, four organising threads capture much of evolutionary studies at this time. First, the nature of species and the process of speciation were dominating, unifying subjects. Second, research into these subjects developed along four main lines, or problem complexes: variation, divergence, isolation, and selection. Some calls (...) for 'synthesis' focused on these problem complexes (sometimes on one of these; other times, all). In these calls, comprehensive and pluralist compendia of plausibly relevant elements were preferred over reaching consensus about the value of particular formulae. Third, increasing confidence in the study of common problems coincided with methodological and epistemic changes associated with experimental taxonomy. Finally, the surge of interest in species problems and speciation in the 1930s is intimately tied to larger trends, especially a shifting balance in the life sciences towards process-based biologies and away from object-based naturalist disciplines. Advocates of synthesis in evolution supported, and were adapting to, these larger trends. (shrink)

The Committee on Common Problems of Genetics, Paleontology, and Systematics (United States National Research Council) marks part of a critical transition in American evolutionary studies. Launched in 1942 to facilitate cross-training between genetics and paleontology, the Committee was also designed to amplify paleontologist voices in modern studies of evolutionary processes. During coincidental absences of founders George Gaylord Simpson and Theodosius Dobzhansky, an opportunistic Ernst Mayr moved into the project's leadership. Mayr used the opportunity for programmatic reforms he had been pursuing (...) elsewhere for more than a decade. These are evident in the Bulletins he distributed under Committee auspices. In his brief tenure as Committee leader, Mayr gained his first substantial foothold within the coalescing community infrastructure of evolutionary studies. Carrying this momentum forward led Mayr directly into the project to launch the journal Evolution. The sociology of interdisciplinary activity provides useful tools for understanding the Committee's value in the broad sweep of change in evolutionary studies during the synthesis period. (shrink)

David Lack of Oxford University and V. C. Wynne- Edwards of Aberdeen University were renowned ornithologists with contrasting views of the modern synthesis which deeply influenced their interpretation and explanation of bird behavior. In the 1950's and 60's Lack became the chief advocate of neo-Darwinism with respect to avian ecology, while Wynne- Edwards developed his theory of group selection. Lack 's position was consistent with the developing focus on individual level adaptation, which was a core concept of the modern synthesis. (...) Alternatively, Wynne- Edwards viewed the emphasis on populations as the most important development provided by the modern synthesis. In this paper, I present the development of these two positions and trace their roots in the literature of the synthesis. Through an analysis of Lack 's 1966 critique of Wynne- Edwards I conclude that Wynne- Edwards was, in many ways, justified in his pursuit of group level explanations. (shrink)

Ernst Mayr''s distinction between ultimate and proximate causes is justly considered a major contribution to philosophy of biology. But how did Mayr come to this philosophical distinction, and what role did it play in his earlier scientific work? I address these issues by dividing Mayr''s work into three careers or phases: 1) Mayr the naturalist/researcher, 2) Mayr the representative of and spokesman for evolutionary biology and systematics, and more recently 3) Mayr the historian and philosopher of biology. If we want (...) to understand the role of the proximate/ultimate distinction in Mayr''s more recent career as a philosopher and historian, then it helps to consider hisearlier use of the distinction, in the course of his research, and in his promotion of the professions of evolutionary biology and systematics. I believe that this approach would also shed light on some other important philosophical positions that Mayr has defended, including the distinction between essentialism: and population thinking. (shrink)

Of all the scientists discussed by Mitman, Keller, and Taylor, Odum stands out most as the technocrat, the social engineer. But less obvious candidates, like Allee, also fancied themselves in this capacity: “Our task as biologists and as citizens of a civilized country, is a practical engineering job.” Allee had in mind the establishment of an international cooperative order based on his biological principles. He apparently did not recognize the extent to which his principles were themselves an engineering feat: he (...) had already constructed a world in which eternal peace and order were possible.To an engineer in the traditional sense, the world is changeable, but not in all respects; there are constraints, and these constraints are taken very seriously. Scientists acting as engineers, in the traditional sense, must also pay attention to constraints. But scientists sometimes also take the option of engineering the very constraints, intellectually reconstructing the world so that it can (supposedly) be physically manipulated in the desired direction. There seems to be a lot of engineering, in the extended sense, going on in the very interesting stories that Mitman, Keller, and Taylor tell. (shrink)

This work on behavioural ecology covers: natural selection and life histories; exploitation of resources; sexual selection and reproductive strategies; co-operation and conflict.

Arthur G. Tansley never accepted Frederic E. Clements’ view that succession is a developmental process whose final stage, the climax formation, is determined primarily by regional climate and that all other types of vegetation are some kind of successional stage or arrested successional stage. Tansley was convinced that in a given region a variety of environmental factors could produce different kinds of climax formations. At the heart of their dispute was Clements’ organicist view of succession, i.e., the formation was a (...) complex organism with an ontogeny and phylogeny. As early as 1905, Tansley offered an alternative to Clements’ complex organism, the quasi-organism, but Clements in private and public rejected this compromise. Tansley and other plant ecologists continued to criticize Clements’ theories for the next 20 years, but with no impact on Clements. John Phillips, a South African plant ecologist who was a follower of Clements, published a series of papers in 1934 and 1935 defending Clementsian ecology. These papers were triggered by the publication of a letter by another ecologist working in Africa who claimed that there was a strong correlation between soils and various kinds of climax vegetation, which was contrary to what was predicted by Phillips and Clements. In 1935, Tansley published an attack on Phillips and Clements and their developmental theory of succession. In it, he proposed the concept of the ecosystem as a way to get around Clements’ monoclimax theory by making the physical environment (e.g., soil chemistry, soil texture, soil moisture) as important a factor as climate, plants and other organisms in determining the composition and characteristics of ecological entities, i.e., ecosystems. Tansley’s ecosystem concept quickly replaced Clements’ monoclimax theory as a dominant paradigm in ecology. (shrink)

Arthur G. Tansley never accepted Frederic E. Clements’ view that succession is a developmental process whose final stage, the climax formation, is determined primarily by regional climate and that all other types of vegetation are some kind of successional stage or arrested successional stage. Tansley was convinced that in a given region a variety of environmental factors could produce different kinds of climax formations. At the heart of their dispute was Clements’ organicist view of succession, i.e., the formation was a (...) complex organism with an ontogeny and phylogeny. As early as 1905, Tansley offered an alternative to Clements’ complex organism, the quasi-organism, but Clements in private and public rejected this compromise. Tansley and other plant ecologists continued to criticize Clements’ theories for the next 20 years, but with no impact on Clements. John Phillips, a South African plant ecologist who was a follower of Clements, published a series of papers in 1934 and 1935 defending Clementsian ecology. These papers were triggered by the publication of a letter by another ecologist working in Africa who claimed that there was a strong correlation between soils and various kinds of climax vegetation, which was contrary to what was predicted by Phillips and Clements. In 1935, Tansley published an attack on Phillips and Clements and their developmental theory of succession. In it, he proposed the concept of the ecosystem as a way to get around Clements’ monoclimax theory by making the physical environment (e.g., soil chemistry, soil texture, soil moisture) as important a factor as climate, plants and other organisms in determining the composition and characteristics of ecological entities, i.e., ecosystems. Tansley’s ecosystem concept quickly replaced Clements’ monoclimax theory as a dominant paradigm in ecology. (shrink)

This paper compares and contrasts three groups that conducted biological research at Yale University during overlapping periods between 1910 and 1970. Yale University proved important as a site for this research. The leaders of these groups were Ross Granville Harrison, Grace E. Pickford, and G. Evelyn Hutchinson, and their members included both graduate students and more experienced scientists. All produced innovative research, including the opening of new subfields in embryology, endocrinology and ecology respectively, over a long period of time. Harrison's (...) is shown to have been a classic research school; Pickford's and Hutchinson's were not. Pickford's group was successful in spite of her lack of departmental or institutional position or power. Hutchinson and his graduate and post-graduate students were extremely productive but in diverse areas of ecology. His group did not have one focused area of research or use one set of research tools. The paper concludes that new models for research groups are needed, especially for those, like Hutchinson's, that included much field research. (shrink)

Ever since the early decades of this century, there have emerged a number of competing schools of ecology that have attempted to weave the concepts underlying natural resource management and natural-historical traditions into a formal theoretical framework. It was widely believed that the discovery of the fundamental mechanisms underlying ecological phenomena would allow ecologists to articulate mathematically rigorous statements whose validity was not predicated on contingent factors. The formulation of such statements would elevate ecology to the standing of a rigorous (...) scientific discipline on a par with physics. However, there was no agreement as to the fundamental units of ecology. Systems ecologists sought to identify the fundamental organization that tied the physical and biological components of ecosystems into an irreducible unit: the ecosystem was their fundamental unit. Population ecologists sought, instead, to identify the biological mechanisms regulating the abundance and distribution of plant and animal species: to these ecologists, the individual organism was the fundamental unit of ecology, and the physical environment was nothing more than a stage upon which the play of individuals in perennial competition took place. As Joel Hagen has pointed out, the two schools were thus dividied by fundamentally different and irreconcilable assumptions about the nature of ecosystems.Notwithstanding these divisive efforts to elevate the image of ecology, the discipline remained in the shadows of American academia until the mid-1960s, when systems ecologists succeeded in projecting ecology onto the national scene. They did so by seeking closer involvement with practical problems: they argued before Congress that their approach to the theoretical problems of ecology was uniquely suited to the solution of the impending “environmental crisis.” With the establishment of the International Biological Program, they succeeded in attracting unprecedented levels of funding for systems ecology research. Theoretical population ecologists, on the other hand, found themselves consigned to the outer regions of this new institutional landscape. The systems ecologists' successful capture of the limelight and the purse brought the divisions between them and population ecologists into sharper relief — hence the hardening of the division of ecology observed by Hagen.45I have argued that the population biologist Richard Levins, prompted by these institutional developments, sought to challenge the social position of systems ecology, and to assert the intellectual priority of theoretical population ecology. He attempted to do so by articulating a nontrivial and rather carefully thought out classification of ecological models that led to the disqualification of systems analysis as a legitimate approach to the study of ecological phenomena. I have suggested that — ultimately —Levins's case against systems analysis in ecology rested on the view that an aspiration to realism and prediction was incompatible with an interest in theoretical issues, a concern that he equated with the search for generality. He sought to reinforce this argument by exploiting the fact that systems ecologists had staked their future on the provision of technical solutions to the problems of the “environmental crisis”: he associated systems ecologists' aspiration to realism and precision with a concern for practical issues, trading on the widely accepted view that practical imperatives are incompatible with the aims of scientific inquiry.46 These are plausible, but nonetheless questionable, claims which have now become an integral part of ecological knowledge. And finally, I hope to have shown how even the most abstract levels of scientific argument are shaped by political considerations, and how discussions of the conceptual development of modern ecology might benefit from a greater consideration of its historical and social dimensions.47. (shrink)

John Greene has dismissed the evolutionary ethics of Simpson as a case in which science was “only a tool, a weapon, in defense of positions that were essentially religious and philosophical.”57 This position adopts an amorphous view of science, in which a scientific theory can be construed to support practically any rhetorical position. The relationship between theory and rhetoric, however, is more complex; it is interactive, with the theory and the rhetoric influencing and supporting one another. It is no coincidence (...) that Allee, Emerson, and Simpson all arrived at a biological basis of democracy and a naturalistic ethics during a period when the future of world politics and man's own morality were in question. Allee's commitment to world peace certainly antedated his theory of sociality, just as Simpson's commitment to democracy undoubtedly preceded his evolutionary views.By adopting a particular theory, however, the biologist necessarily imposes constraints on the corresponding rhetoric. Allee and Emerson could not, for example, have stressed the importance of the individual in democracy, given the orientation and framework of their biological research. There were differences among the social philosophies of Allee, Emerson, and Simpson; these differences depended, in part, on the specific evolutionary metaphors to which they subscribed. Allee's ideas with respect to cooperation were distinct from Emerson's, and both men differed strongly from Simpson on the role of the individual in evolution. The Chicago school was united by a conceptual framework that emphasized the population as the unit of selection, and the importance of cooperation in nature. But cooperation could play many roles: as a unifying principle for a theory of sociality, as an integrating mechanism in physiological functionalism, and as a biological source of hope for a society in the grips of a world war. (shrink)

With the rejection of group selectionist derivations of ecological phenomena so incisively given by George Williams in 1966,43 Nicholson's long-ignored messages met with acceptance. Species benefit became, explicitly, incidental. But the reorientation was not just about a point of ecological theory. It was more fundamentally about theoretical style, the element shared by Wynne-Edwards' work and the newer, evolutionary ecology. That current approach is well expressed in an already classic paper by the British plant ecologist John Harper: Ultimately all the discoveries (...) of descriptive, production or population ecology must find their meaning in evolutionary phenomena... Evolutionary thinking concentrates attention on the behaviour of the individual and his descendants. If nothing in biology has meaning except in the light of evolution and if evolution is about individuals and their descendants — i.e. fitness — we should not expect to reach any depth of understanding from studies that are based at the level of the superindividual... What we see as the organised behaviour of systems is the result of the fate of individuals.44The emphasis in this passage is on style of thinking more than on study matter. What is so different from earlier ecologists is the rejection of the superorganism and its associated baggage of selfimposed constraints in an end-oriented process, such as balance of nature or species-level adaptation.The hallmark of post-1959 evolutionary ecology is the recognition, the making explicit of the question that the selective, genetical process of fitness (fates of genes) has to be separated from the observations of different levels of adaptiveness. An easy conflation of cause and result is no longer satisfying; now attempts to disentangle the complex strands tying population phenomena to the different levels of selection are providing the excitement of a new research program. In this program evolutionary theory is used to define the appropriate questions, as exemplified by the organization of the textbook Evolutionary Ecology.45 Our discussion of the invention of a new approach, with a characteristic style and set of interests, must also be about how the practitioners of ecology have perceived themselves. The early attempts of Poulton and Nicholson were not taken up by the vigorously growing science of ecology, which developed first - and intentionally — along a narrower, hierarchically restrained set of questions about population. The depth of Poulton's and Nicholson's submergence is indicated by the fact that even Wynne-Edwards did not argue in the manner of one rescuing a minor tradition. Rather, he thought he had a new approach, that of placing evolution in the foundations of ecology. Although this approach was not exactly original, we should be cautious in grouping all evolutionary concerns within ecology in one intellectual tradition. The particular style of selectionist deduction invigorated by Wynne-Edwards was successful only in the volatile situation of a rapidly growing discipline, full of self-critical examination and new rigor. Whereas in 1930 evolutionary concerns did not provide that rigor, by 1960 they could - and did. (shrink)

Robert MacArthur's mathematical ecology is often regarded as ahistorical and has been criticized by historically oriented ecologists and philosophers for ignoring the importance of history. I clarify and defend his approach, especially his use of simple mathematical models to explain patterns in data and to generate predictions that stimulate empirical research. First I argue that it is misleading to call his approach ahistorical because it is not against historical explanation. Next I distinguish three kinds of criticism of his approach and (...) argue that his approach is compatible with the first two of them. Finally, I argue that the third kind of criticism, advanced by Kim Sterelny and Paul Griffiths, is largely irrelevant to MacArthur's approach. ‡I am especially grateful to Thomas Nickles for encouragement and helpful comments on earlier versions of this paper. Thanks also to Guy Hoelzer, Stephen Jenkins, and Jay Odenbaugh for comments on an earlier draft, Kim Sterelny for clarifications of the Tasmania example, Gregory Mikkelson for references, and the audience at PSA 2006 for discussions. †To contact the author, please write to: Department of History and Philosophy of Science, University of Pittsburgh, 1017 Cathedral of Learning, Pittsburgh, PA 15260; e-mail: [email protected]. (shrink)

Ecology has often been characterized as an immature scientific discipline. This paper explores some of the sources of this alleged immaturity. I argue that the perception of immaturity results primarily from the fact that historically ecologists have based their work upon two very different approaches to research.

This book is a sustained examination of issues in the philosophy of ecology that have been a source of controversy since the emergence of ecology as an explicit scientific discipline. The controversies revolve around the idea of a balance of nature, the possibility of general ecological knowledge and the role of model-building in ecology. The Science of the Struggle for Existence is also a detailed treatment of these issues that incorporates both a comprehensive investigation of the relevant ecological literature and (...) the development of an explicit theoretical framework in the philosophy of science. It addresses issues in the philosophy of ecology that are of particular importance for the deployment of ecology in the solution of environmental problems. It will have a cross-disciplinary appeal and will interest students and professionals in science, the philosophy of science, and environmental studies as well as policy-makers. (shrink)