Environmentalists consider invasions by exotic species of plants and animals to be one of the most serious environmental problems we face today, as well as one of the leading causes of biodiversity loss. We argue that in order to develop and enact sensible policies, it is crucial to consider two philosophical questions: What exactly makes a species native or exotic, and What values are at stake? We focus on the first of these two questions, and offer some preliminary suggestions with (...) regard to the second. Through a series of case studies, we show that it is not always clear whether a species is native or exotic. We identify five possible criteria that could be used for distinguishing natives from exotics. Rather than identifying one of these criteria as the 'correct' one, we suggest that the concepts of 'native' and ' exotic ' function more like what some philosophers have called cluster concepts. That is, there are several characteristics that are typical of native species, and a corresponding set of characteristics that are typical of exotic species. None of these characteristics is either necessary or sufficient for identifying a species as either native or exotic. We then identify several of the values that are at stake in dealing with exotic species, and we suggest that policies need to avoid being overly simplistic. (shrink)

Scientists use models to understand the natural world, and it is important not to conflate model and nature. As an illustration, we distinguish three different kinds of populations in studies of ecology and evolution: theoretical, laboratory, and natural populations, exemplified by the work of R.A. Fisher, Thomas Park, and David Lack, respectively. Biologists are rightly concerned with all three types of populations. We examine the interplay between these different kinds of populations, and their pertinent models, in three examples: the notion (...) of “effective” population size, the work of Thomas Park on /Tribolium/ populations, and model-based clustering algorithms such as /Structure/. Finally, we discuss ways to move safely between three distinct population types while avoiding confusing models and reality. (shrink)

Many biological investigations are organized around a small group of species, often referred to as ‘model organisms’, such as the fruit fly Drosophila melanogaster. The terms ‘model’ and ‘modelling’ also occur in biology in association with mathematical and mechanistic theorizing, as in the Lotka–Volterra model of predator-prey dynamics. What is the relation between theoretical models and model organisms? Are these models in the same sense? We offer an account on which the two practices are shown to have different epistemic characters. (...) Theoretical modelling is grounded in explicit and known analogies between model and target. By contrast, inferences from model organisms are empirical extrapolations. Often such extrapolation is based on shared ancestry, sometimes in conjunction with other empirical information. One implication is that such inferences are unique to biology, whereas theoretical models are common across many disciplines. We close by discussing the diversity of uses to which model organisms are put, suggesting how these relate to our overall account. 1 Introduction2 Volterra and Theoretical Modelling3 Drosophila as a Model Organism4 Generalizing from Work on Model Organisms5 Phylogenetic Inference and Model Organisms6 Further Roles of Model Organisms6.1 Preparative experimentation6.2 Model organisms as paradigms6.3 Model organisms as theoretical models6.4 Inspiration for engineers6.5 Anchoring a research community7 Conclusion. (shrink)

The status of population genetics has become hotly debated among biologists and philosophers of biology. Many seem to view population genetics as relatively unchanged since the Modern Synthesis and have argued that subjects such as development were left out of the Synthesis. Some have called for an extended evolutionary synthesis or for recognizing the insignificance of population genetics. Yet others such as Michael Lynch have defended population genetics, declaring "nothing in evolution makes sense except in the light of population genetics" (...) (a twist on Dobzhansky's famous slogan that "nothing in biology makes sense except in the light of evolution"). Missing from this discussion is the use of population genetics to shed light on ecology and vice versa, beginning in the 1940s and continuing until the present day. I highlight some of that history through an overview of traditions such as ecological genetics and population biology, followed by a slightly more in-depth look at a contemporary study of the endangered California Tiger Salamander. I argue that population genetics is a powerful and useful tool that continues to be used and modified, even if it isn't required for all evolutionary explanations or doesn't incorporate all the causal factors of evolution. (shrink)

A community, for ecologists, is a unit for discussing collections of organisms. It refers to collections of populations, which consist (by definition) of individuals of a single species. This is straightforward. But communities are unusual kinds of objects, if they are objects at all. They are collections consisting of other diverse, scattered, partly-autonomous, dynamic entities (that is, animals, plants, and other organisms). They often lack obvious boundaries or stable memberships, as their constituent populations not only change but also move in (...) and out of areas, and in and out of relationships with other populations. Familiar objects have identifiable boundaries, for example, and if communities do not, maybe they are not objects. Maybe they do not exist at all. The question this possibility suggests, of what criteria there might be for identifying communities, and for determining whether such communities exist at all, has long been discussed by ecologists. This essay addresses this question as it has recently been taken up by philosophers of science, by examining answers to it which appeared a century ago and which have framed the continuing discussion. (shrink)

The complexity and heterogeneity of causes influencing ecology’s domain challenge its capacity to generate a general theory without exceptions, raising the question of whether ecology is capable, even in principle, of achieving the sort of theoretical success enjoyed by physics. Weber has argued that competition theory built around the Competitive Exclusion Principle (especially Tilman’s resource-competition model) offers an example of ecology identifying a law-like causal regularity. However, I suggest that as Weber presents it, the CEP is not yet a causal (...) regularity. Instead, I argue that the scientific understanding in Tilman’s theory takes a different form. The theory explains through a structure I call “channeling explanation” which does not depend on deduction from general laws, but rather builds on constraints and trade-offs represented in state-space. Recognizing this structure supports the more general point that ecology and other so-called special sciences can reveal novel theoretical approaches to philosophy of science when approached with openness to their uniqueness. (shrink)

This paper surveys recent philosophy of biology. It aims to introduce outsiders to the field to the recent literature (which is reviewed in the footnotes) and the main recent debates. I concentrate on three of these: recent critiques of the replicator/vehicle distinction and its application to the idea of the gene as the unit of section; the recent defences of group selection and the idea that standard alternatives to group selection are in fact no more than a disguised form of (...) group selection; and recent ideas on the role of selection in evolution, especially the role of selection in structuring the large-scale history of life. The paper connects philosophy of biology to some more general problems in the philosophy of science, and concludes with a few suggestions about unfinished business. (shrink)

The niche is allegedly the conceptual bedrock underpinning the most prominent, and some would say most important, theorizing in ecology. We argue this point of view is more aspirational than veridical. Rather than critically dissect existing definitions of the concept, the supposedly significant work it is thought to have done in ecology is our evaluative target. There is no denying the impressive mathematical sophistication and theoretical ingenuity of the ecological modeling that invokes ‘niche’ terminology. But despite the pervasive labeling, we (...) demonstrate that niche talk is nothing more than a gloss on theory developed without it, that doesn’t need it, and that doesn’t benefit from it. (shrink)

In this article, I consider the term “environment” in various claims and models by evolutionists and ecologists. I ask whether “environment” is amenable to a philosophical explication, in the same way some key terms of evolutionary theorizing such as “fitness,” “species,” or more recently “population” have been. I will claim that it cannot. In the first section, I propose a typology of theoretical terms, according to whether they are univocal or equivocal, and whether they have been the object of formal (...) or conceptual attempts of clarification. “Environment” will appear to be in a position similar to “population,” yet almost no extant attempt has been made to make sense of its meaning and reference. In the second section, I will present several theoretical claims or issues that refer to “environment” in apparently very diverse ways, but always supposing a contrastive term, which is not always the same. The third section directly considers models in evolution and in ecology, and asks “where” in them is the environment term. The fourth section proposes that “environment” refers to a distinction between a varying and an invariant term, but shows that this does not exhaust its meaning, which requires making more conceptual differences. Then, I take on the suggestion that there might be two “environment” terms, one in ecology and one in evolution, but show that is not the case. Finally I center on the specific notion of “complex environment,” which is the object of several research programs, and propose a typology of “complex environments” across theories. (shrink)

Ecology in principle is tied to evolution, since communities and ecosystems result from evolution and ecological conditions determine fitness values. Yet the two disciplines of evolution and ecology were not unified in the twentieth-century. The architects of the Modern Synthesis, and especially Julian Huxley, constantly pushed for such integration, but the major ideas of the Synthesis—namely, the privileged role of selection and the key role of gene frequencies in evolution—did not directly or immediately translate into ecological science. In this paper (...) I consider five stages through which the Synthesis was integrated into ecology and distinguish between various ways in which a possible integration was gained. I start with Elton’s animal ecology, then consider successively Ford’s ecological genetics in the 1940s, the major textbook Principles of animal ecology edited by Allee et al., and the debates over the role of competition in population regulation in the 1950s, ending with Hutchinson’s niche concept and McArthur and Wilson’s Principles of Island Biogeography viewed as a formal transposition of Modern Synthesis explanatory schemes. I will emphasize the key role of founders of the Synthesis at each stage of this very nonlinear history. (shrink)

Evolutionary, ecological and ethical studies are, at the same time, specific scientific disciplines and, from an historical point of view, structurally linked domains of research. In a context of environmental crisis, the need is increasingly emerging for a connecting epistemological framework able to express a common or convergent tendency of thought and practice aimed at building, among other things, an environmental policy management respectful of the planet’s biodiversity and its evolutionary potential.

During the 1970s, a "revolution" in American paleobiology took place. It came about in part because a group of mostly young, ambitious paleontologists adapted many of the quantitative methodologies and techniques developed in fields including biology and ecology over the previous several decades to their own discipline. Stephen Jay Gould, who was then just beginning his career, joined others in articulating a singular vision for transforming paleontology from an isolated and often ignored science to a "nomothetic discipline" that could sit (...) at evolution's "high table." Over the course of a single decade, between 1970 and 1980, this transformation had in large part been accomplished. Among those most centrally involved in this process were Gould, Thomas Schopf, David Raup, and Gould's graduate student Jack Sepkoski, all of whom made major contributions in theoretical and quantitative analysis of the fossil record and evolutionary history. Recognizing that an ideological agenda was not enough, Gould and others developed and promoted new outlets, technologies, and pedagogical strategies to nurture their new discipline. This paper describes this process of transformation, and presents Sepkoski's education and participation as exemplary of the "new model paleontologist", which Gould hoped to produce. (shrink)

The Lotka–Volterra predator-prey-model is a widely known example of model-based science. Here we reexamine Vito Volterra’s and Umberto D’Ancona’s original publications on the model, and in particular their methodological reflections. On this basis we develop several ideas pertaining to the philosophical debate on the scientific practice of modeling. First, we show that Volterra and D’Ancona chose modeling because the problem in hand could not be approached by more direct methods such as causal inference. This suggests a philosophically insightful motivation for (...) choosing the strategy of modeling. Second, we show that the development of the model follows a trajectory from a “how possibly” to a “how actually” model. We discuss how and to what extent Volterra and D’Ancona were able to advance their model along that trajectory. It turns out they were unable to establish that their model was fully applicable to any system. Third, we consider another instance of model-based science: Darwin’s model of the origin and distribution of coral atolls in the Pacific Ocean. Darwin argued more successfully that his model faithfully represents the causal structure of the target system, and hence that it is a “how actually” model. (shrink)

This paper explores the work of Nicolas Rashevsky, a Russian émigré theoretical physicist who developed a program in "mathematical biophysics" at the University of Chicago during the 1930s. Stressing the complexity of many biological phenomena, Rashevsky argued that the methods of theoretical physics -- namely mathematics -- were needed to "simplify" complex biological processes such as cell division and nerve conduction. A maverick of sorts, Rashevsky was a conspicuous figure in the biological community during the 1930s and early 1940s: he (...) participated in several Cold Spring Harbor symposia and received several years of funding from the Rockefeller Foundation. However, in contrast to many other physicists who moved into biology, Rashevsky's work was almost entirely theoretical, and he eventually faced resistance to his mathematical methods. Through an examination of the conceptual, institutional, and scientific context of Rashevsky's work, this paper seeks to understand some of the reasons behind this resistance. (shrink)

Organizational ecology is commonly seen as a Darwinian research program that seeks to explain the diversity of organizational structures, properties and behaviors as the product of selection in past social environments in a similar manner as evolutionary biology seeks to explain the forms, properties and behaviors of organisms as consequences of selection in past natural environments. We argue that this explanatory strategy does not succeed because organizational ecology theory lacks an evolutionary mechanism that could be identified as the principal cause (...) of organizational diversity. The “evolution” of organizational populations by means of selection, which organizational ecologists put forward as the mechanism responsible for the extant diversity of organizational forms, is not evolution in any proper sense, because organizational populations do not have what it takes to participate in evolutionary processes. This implies that organizational ecology is not a Darwinian research program and that it cannot explain organizational diversity. (shrink)

Hull's recent work in evolutionary epistemology is marred by a deep tension. While he maintains that conceptual and biological evolution are both driven by selection processes, he also claims that only the former is globally progressive. In this paper I formulate this tension and present four possible responses (including Hull's). I argue that Hull's position rests on the assumption that there is a goal which is sufficiently general to describe most scientific activity yet precise enough to guide research. Working from (...) within Hull's framework, I argue that a non-progressionist stance is both preferable and more consistent with Hull's basic commitments. (shrink)

This paper revisits the debate over whether the study of facilitation requires ecologists to revise their understanding of the relationship between realized and fundamental niches as conceptualized by Hutchinson. Following Rodriguez-Cabal et al., I argue against Bruno et al.’s claim that facilitation can make a species’ realized niche larger than its fundamental niche. However, I also maintain that the abstract Hutchinsonian conceptualization of the niche makes a whole range of facilitative interactions—which I propose to call ameliorative facilitation—invisible to niche-based approaches (...) to the study of ecological communities. I propose a way to incorporate ameliorative facilitation into such approaches. My proposal involves supplementing the Hutchinsonian realized/fundamental dyad with a third concept: the potential niche. This concept was introduced by ecologists studying the effects of environmental change on species distributions, but I show how it could also be fruitfully used in facilitation studies. I argue that this proposed solution is more appealing than Stachowicz’s suggestion that Hutchinson’s realized/fundamental contrast be applied to a spatial-geographical, as opposed to an abstract-conceptual, notion of the niche. (shrink)

We invite systematic consideration of the metaphors of cycles and circulation as a long-term theme in the history of the life and environmental sciences and medicine. Ubiquitous in ancient religious and philosophical traditions, especially in representing the seasons and the motions of celestial bodies, circles once symbolized perfection. Over the centuries cyclic images in western medicine, natural philosophy, natural history and eventually biology gained independence from cosmology and theology and came to depend less on strictly circular forms. As potent ‘canonical (...) icons’, cycles also interacted with representations of linear and irreversible change, including arrows, arcs, scales, series and trees, as in theories of the Earth and of evolution. In modern times life cycles and reproductive cycles have often been held to characterize life, in some cases especially female life, while human efforts selectively to foster and disrupt these cycles have harnessed their productivity in medicine and agriculture. But strong cyclic metaphors have continued to link physiology and climatology, medicine and economics, and biology and manufacturing, notably through the relations between land, food and population. From the grand nineteenth-century transformations of matter to systems ecology, the circulation of molecules through organic and inorganic compartments has posed the problem of maintaining identity in the face of flux and highlights the seductive ability of cyclic schemes to imply closure where no original state was in fact restored. More concerted attention to cycles and circulation will enrich analyses of the power of metaphors to naturalize understandings of life and their shaping by practical interests and political imaginations. (shrink)

Ruth Myrtle Patrick was a pioneering ecologist and taxonomist whose extraordinary career at the Academy of Natural Sciences of Philadelphia spanned over six decades. In 1947, an opportunity arose for Patrick to lead a new kind of river survey for the Pennsylvania Sanitary Water Board to study the effects of pollution on aquatic organisms. Patrick leveraged her already extensive scientific network, which included ecologist G. Evelyn Hutchinson, to overcome resistance within the Academy, establish a new Department of Limnology, and carry (...) out the survey, which was a resounding success and brought much needed money to the Academy. As demand for her expertise grew among industrial companies, such as the chemical company DuPont, Patrick became more active in the world of applied science. She repurposed data and instruments from her river surveys to run new experiments, test ecological theories, and conduct long-term ecological studies. Through these studies, she advanced an argument that biologist Thomas Lovejoy dubbed the “Patrick principle,” the idea that the ecological health of a body of water could be measured by the relative abundance and diversity of species living there. Patrick was elected to the National Academy of Sciences in 1970, became a board member of DuPont in 1975, and received two of the most prestigious awards in ecology: the Eminent Ecologist Award from the Ecological Society of America in 1972 and the Tyler Ecology Award in 1975. This article analyzes Patrick’s unusual success in bridging the worlds of science and industry and her unusual ability to cross, and redefine, the perceived boundary between basic and applied fields in biology. It argues that Patrick’s position at the Academy, an institution of natural history that was both willing and able to accept money from industrial corporations, is key to understanding her success in, and influence on, the field of river ecology. (shrink)

This Ph.D. thesis provides a pilosophical account of the structure of the evolutionary synthesis of the 1930s and 40s. The first, more historical part analyses how classical genetics came to be integrated into evolutionary thinking, highlighting in particular the importance of chromosomal mapping of Drosophila strains collected in the wild by Dobzansky, but also the work of Goldschmidt, Sumners, Timofeeff-Ressovsky and others. The second, more philosophical part attempts to answer the question wherein the unity of the synthesis consisted. I argue (...) that it exemplifies a weak of form of explanatory unification. (shrink)

Many standard philosophical accounts of scientific practice fail to distinguish between modeling and other types of theory construction. This failure is unfortunate because there are important contrasts among the goals, procedures, and representations employed by modelers and other kinds of theorists. We can see some of these differences intuitively when we reflect on the methods of theorists such as Vito Volterra and Linus Pauling on the one hand, and Charles Darwin and Dimitri Mendeleev on the other. Much of Volterra's and (...) Pauling's work involved modeling; much of Darwin's and Mendeleev's did not. In order to capture this distinction, I consider two examples of theory construction in detail: Volterra's treatment of post-WWI fishery dynamics and Mendeleev's construction of the periodic system. I argue that modeling can be distinguished from other forms of theorizing by the procedures modelers use to represent and to study real-world phenomena: indirect representation and analysis. This differentiation between modelers and non-modelers is one component of the larger project of understanding the practice of modeling, its distinctive features, and the strategies of abstraction and idealization it employs. (shrink)

The contents of the British Ornithologists' Union's journal, "The Ibis," during the first half of the 20th century illustrates some of the transformations that have taken place in the naturalist tradition. Although later generations of ornithologists described these changes as logical and progressive, their historical narratives had more to do with legitimizing the infiltration of the priorities of evolutionary theory, ecology, and ethology than analyzing the legacy of the naturalist tradition on its own terms. Despite ornithologists' claim that the journal's (...) increasing focus on "biology" represented a natural development after the preliminary phase of systematics and geographical ornithology, in fact a small group campaigned to bring the priorities of population ecology, behavior, and selection theory into the journal and British ornithology more generally. The problems involved in this transition highlight the importance of methodological and institutional context in determining and reinforcing appropriate research programs for ornithologists. Comparing the discipline-building rhetoric of moderns with the contents of the past illustrates how modern evaluations of 19th century research programs have been enmeshed in ornithologists' endeavors to forge new identities for traditional disciplines. (shrink)

Wildlife populations in the northern reaches of the globe have long been observed to fluctuate or cycle periodically, with dramatic increases followed by catastrophic crashes. Focusing on the early work of Charles S. Elton, this article analyzes how investigations into population cycles shaped the development of Anglo-American animal ecology during the 1920s–1930s. Population cycling revealed patterns that challenged ideas about the “balance” of nature; stimulated efforts to quantify population data; and brought animal ecology into conversation with intellectual debates about natural (...) selection. Elton used the problem of understanding wildlife population cycles to explore a central tension in ecological thought: the relative influences of local conditions (food supply, predation) and universal forces (such as climate change and natural selection) in regulating wild animal populations. He also sought patronage and built research practices and the influential Bureau of Animal Population around questions of population regulation during the 1930s. Focusing on disease as a local population regulator that could interact with global climatic influences, Elton facilitated an interdisciplinary and population-based approach in early animal ecology. Elton created a network of epidemiologists, conservationists, pathologists and mathematicians, who contributed to population cycle research. I argue that, although these people often remained peripheral to ecology, their ideas shaped the young discipline. Particularly important were the concepts of abundance, density, and disease; and the interactions between these factors and natural selection. However, Elton’s reliance on density dependence unwittingly helped set up conditions conducive to the development of controversies in animal ecology in later years. While ecologists did not come to consensus on the ultimate causes of population cycles, this phenomenon was an important early catalyst for the development of theory and practice in animal ecology. (shrink)

I claim that the balance of nature metaphoris shorthand for a paradigmatic view of natureas a beneficent force. I trace the historicalorigins of this concept and demonstrate that itoperates today in the discipline of populationecology. Although it might be suspected thatthis metaphor is a pre-theoretic description ofthe more precisely defined notion ofequilibrium, I demonstrate that balance ofnature has constricted the meaning ofmathematical equilibrium in population ecology.As well as influencing the meaning ofequilibrium, the metaphor has also loaded themathematical term with values.Environmentalists (...) and critics use thisconflation of meaning and value to theiradvantage. This interplay between the balanceof nature and equilibrium fits aninteractionist interpretation of the role ofmetaphor in science. However, it seems theinteraction is asymmetric, and the balance ofnature metaphor has had a larger influence onmathematical equilibrium than vice versa. Thisdisproportionate influence suggests that themetaphor was and continues to be a constitutivepart of ecological theories. (shrink)

I present an attempt at an explication of the ecological theory of interspecific competition, including its explanatory role in community ecology and evolutionary biology. The account given is based on the idea that law-like statements play an important role in scientific theories of this kind. I suggest that the principle of competitive exclusion is such a law, and that it is evolutionarily invariant. The principle's empirical status is defended and implications for the ongoing debates on the existence of biological laws (...) are discussed. (shrink)

Philosophers of science have examined The Theory of Island Biogeography by Robert MacArthur and E. O. Wilson (1967) mainly due to its important contribution to modeling in ecology, but they have not examined it as a representative case of ecological explanation. In this paper, I scrutinize the type of explanation used in this paradigmatic work of ecology. I describe the philosophy of science of MacArthur and Wilson and show that it is mechanistic. Based on this account and in light of (...) contributions to the mechanistic conception of explanation due to Craver (2007), and Bechtel and Richardson (1993), I argue that MacArthur and Wilson use a mechanistic approach to explain the species-area relationship. In light of this examination, I formulate a normative account of mechanistic explanation in ecology. Furthermore, I argue that it offers a basis for methodological unification of ecology and solves a dispute on the nature of ecology. Lastly, I show that proposals for a new paradigm of biogeography appear to maintain the norms of mechanistic explanation implicit in The Theory of Island Biogeography. (shrink)

Abstract Recent criticisms of intuition from experimental philosophy and elsewhere have helped undermine the authority of traditional conceptual analysis. As the product of more empirically informed philosophical methodology, this result is compelling and philosophically salutary. But the negative critiques rarely suggest a positive alternative. In particular, a normative account of concept determination—how concepts should be characterized—is strikingly absent from such work. Carnap's underappreciated theory of explication provides such a theory. Analyses of complex concepts in empirical sciences illustrates and supports this (...) claim, and counteracts the charge explication is only suitable for highly mathematical, axiomatic contexts. Explication is also defended against the influential criticism it is “philosophically unilluminating”. Content Type Journal Article Category Original paper in Philosophy of Science Pages 1-19 DOI 10.1007/s13194-011-0027-5 Authors James Justus, Philosophy Department, Florida State University and University of Sydney, Tallahassee, FL 32306, USA Journal European Journal for Philosophy of Science Online ISSN 1879-4920 Print ISSN 1879-4912. (shrink)

To generate explanatory theory, ecologists must wrestle with how to represent the extremely many, diverse causes behind phenomena in their domain. Early twentieth-century plant ecologists Frederic E. Clements and Henry A. Gleason provide a textbook example of different approaches to explaining vegetation, with Clements allegedly committed, despite abundant exceptions, to a law of vegetation, and Gleason denying the law in favor of less organized phenomena. However, examining Clements's approach to explanation reveals him not to be expressing a law, and instead (...) to be developing an explanatory structure without laws, capable of progressively integrating causal complexity. Moreover, Clements and Gleason largely agree on the causes of vegetation; but, since causal understanding here underdetermines representation, they differ on how to integrate recognized causes into general theory---that is, in their methodologies. Observers of the case may have mistakenly assumed that scientific representation across the disciplines typically aims at laws like Newton's, and that representations always reveal scientists' metaphysical commitments. Ironically, in the present case, this assumption seems to have been made even by observers who regard Clements as naı¨ve for his alleged commitment to an ecological law. (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)

Beginning in the nineteenth century, scientists speculated that the Pleistocene megafauna—species such as the giant ground sloth, wooly mammoth, and saber-tooth cat—perished because of rapid climate change accompanying the end of the most recent Ice Age. In the 1950s, a small network of ecologists challenged this view in collaboration with archeologists who used the new tool of radiocarbon dating. The Pleistocene overkill hypothesis imagined human hunting, not climate change, to be the primary cause of megafaunal extinction. This article situates the (...) Pleistocene overkill hypothesis in a broader history of the emergence of historical ecology as a distinct sub-discipline of paleoecology. Tracing the work of the Yale Geochronometric Laboratory and an interdisciplinary research network that included Paul Sears, Richard Foster Flint, Edward Deevey, Kathryn Clisby, and Paul S. Martin, it reveals how both the methods and the meaning of studying fossil pollen shifted between the 1910s and 1960s. First used as a tool for fossil fuel extraction, fossil pollen became a means of envisioning climatic history, and ultimately, a means of reimagining global ecological history. First through pollen stratigraphy and then through radiocarbon dating, ecologists reconstructed past biotic communities and rethought the role of humans in these communities. By the 1980s, the discipline of historical ecology would reshape physical environments through the practice of ecological restoration. (shrink)

In “The Origin of Species,” Darwin describes a hypothetical example illustrating that large, slowly reproducing mammals such as the elephant can reach very large numbers if population growth is not affected by regulating factors. The elephant example has since been cited in various forms in a wide variety of books, ranging from educational material to encyclopedias. However, Darwin’s text was changed over the six editions of the book, although some errors in the mathematics persisted throughout. In addition, full details of (...) the problem remained hidden in his correspondence with readers of the Origin. As a result, Darwin’s example is very often misinterpreted, misunderstood or presented as if it were a fact. We show that the population growth of Darwin’s elephant population can be modeled by the Leslie matrix method, which we generalize here to males as well. Darwin’s most often cited figure, about 19 million elephants after 750 years is not a typical outcome, actually a very unlikely result under more realistic, although still hypothetical situations. We provide a recursion formula suggesting that Darwin’s original model corresponds to a tribonacci series, a proof showing that sex ratio is constant over all age classes, and a derivation of a generating function of the sequence. (shrink)

In this contribution, I comment on Raffaella Campaner’s defense of explanatory pluralism in psychiatry (in this volume). In her paper, Campaner focuses primarily on explanatory pluralism in contrast to explanatory reductionism. Furthermore, she distinguishes between pluralists who consider pluralism to be a temporary state on the one hand and pluralists who consider it to be a persisting state on the other hand. I suggest that it would be helpful to distinguish more than those two versions of pluralism – different understandings (...) of explanatory pluralism both within philosophy of science and psychiatry – namely moderate/temporary pluralism, anything goes pluralism, isolationist pluralism, integrative pluralism and interactive pluralism. Next, I discuss the pros and cons of these different understandings of explanatory pluralism. Finally, I raise the question of how to implement or operationalize explanatory pluralism in scientific practice; how to structure the “genuine dialogue” or shape “the pluralistic attitude” Campaner is referring to. As tentative answers, I explore a question-based framework for explanatory pluralism as well as social-epistemological procedures for interaction among competing approaches and explanations. (shrink)

During the 1930s, Aleksandr Promptov—a student of the founder of Russian population genetics Sergei Chetverikov—developed an elaborate concept of speciation in birds. He conducted field investigations aimed at giving a naturalistic content to the theoretical formulations and laboratory models of evolutionary processes advanced within the framework of population genetics, placing particular emphasis on the evolutionary role of bird behavior. Yet, although highly synthetic in combining biogeographical, taxonomic, genetic, ecological, and behavioral studies, Promptov's speciation concept was ignored by the architects of (...) the 1930s and 1940s evolutionary synthesis, including Theodosius Dobzhnasky, Ernst Mayr, and Julian Huxley. In this article, I argue that the story of Promptov's concept and its reception by other evolutionists challenges the traditional presentation of the synthesis as a singular, international process of the unification of biology, which led to the creation of a universal synthetic theory of evolution. It suggests that during the same time period, within largely the same theoretical framework, there were multiple, intrinsically local, attempts at creating synthetic evolutionary concepts. These concepts were often quite particular—in their taxonomic applicability, in their explanations of various evolutionary factors, and in the range of disciplines unified in the synthesis. Apparently, these concepts ran contrary to the universal aspirations of the synthesis architects, and as a result, they were disregarded, first by the architects and later by historians of the evolutionary synthesis. (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.

Ricotta argues against the existence of a unique measure of biodiversity by pointing out that no known measure of α-diversity satisfies all the adequacy conditions that have traditionally been set for it. While that technical claim is correct, it is not relevant in the context of defining biodiversity which is most usefully measured by β-diversity. The concept of complementarity provides a closely related family of measures of biodiversity which can be used for systematic conservation planning. Moreover, these measures cannot be (...) replaced by summary statistics but must rely on inventories of biodiversity surrogates at candidate sites for conservation. (shrink)

This article examines adaptive resource management as it has been applied to the US horseshoe crab fishery over the past decade. As a critical yet constructive exercise, I have three goals: to suggest how adaptive management, for all its promise, can still be improved; to add a nuanced case study to the literatures on the quantification of nature and environmental decision-making; and to use the example of ARM to make certain temporal aspects of contemporary natural resource management more salient to (...) science and technology studies scholars—that is, to show the ways in which time matters in environmental science, policy, and the analysis thereof. I draw attention to the time-related aspects of adaptive management by developing the notions of temporal orientation and chronological accountability. Temporal orientation refers to the time-based perspectives and epistemological commitments—that is, past-facing empiricism versus future-oriented modeling—that scientists of different types bring to bear on environmental problems. Chronological accountability refers to the missing link in adaptive forms of environmental governance: firm time lines and commitments to reflexively revisit management decisions. The time-related aspects of natural resource management deserve greater attention among both environmental managers and analysts of environmental policy. (shrink)

The representation of a complex but stable, self-regulated and, finally, harmonious nature penetrates the whole history of Ecology, thus contradicting the core of the Darwinian evolution. Originated in the pre-Darwinian Natural History, this representation defined theoretically the various schools of early ecology and, in the context of the cybernetic synthesis of the 1950s, it assumed a typical mathematical form on account of α positive correlation between species diversity and community stability. After 1960, these two aforementioned concepts and their positive correlation (...) were proposed as environmental management tools, in the face of the ecological crisis arising at the time. In the early 1970s, and particularly after May’s evolutionary arguments, the consensus around this positive correlation collapsed for a while, only to be promptly restored for the purpose of attaching an ecological value on biodiversity. In this paper, we explore the history of the diversity–stability hypothesis and we review the successive terms that have been used to express community stability. We argue that this hypothesis has been motivated by the nodal ideological presuppositions of order and harmony and that the scientific developments in this field largely correspond to external social pressures. We conclude that the conflict about the diversity–stability relationship is in fact an ideological debate, referring mostly to the way we see nature and society rather than to an autonomous scientific question. From this point of view, we may understand why Ecology’s concepts and perceptions may decline and return again and again, forming a pluralistic scientific history. (shrink)

During its centennial celebrations in 2008, the Museum of Vertebrate Zoology (MVZ) at the University of California, Berkeley paid homage to its founding director, Joseph Grinnell. Recognized as a leading scientific institution, the MVZ managed to grow throughout the twentieth century, a period often characterized by the decline of natural history. To understand how and why research flourished at the MVZ, this paper looks closely at Grinnell's undergraduate course, the Natural History of the Vertebrates (NHV). Taught by MVZ affiliates since (...) 1914, the NHV offers an important window on Grinnell's approach and legacy. This paper argues that the NHV contributed to the MVZ's long-term success by acting as, first, a gateway to natural history; second, a vector for the MVZ's research programme; and third, a shared faculty responsibility. Grinnell's significance in the history of science is understated, in part because his writing style de-emphasized the importance of his theoretical contributions, including his development of the niche concept, his emphasis on population thinking and geographic isolation in studies of evolution, and his effort to integrate speciation questions and genetics. Studying the NHV highlights these contributions because Grinnell freely communicated his ideas to his students. An analysis of Grinnell's course material shows that his theoretical and methodological approach pre-dated the evolutionary synthesis and inspired natural-history research throughout the past century. (shrink)

In biology, the ‘ecological orientation' rests on a commitment to examining systems, and the conceptual challenge of defining that system now employs techniques and concepts adapted from diverse disciplines (i.e., systems philosophy, cybernetics, information theory, computer science) that are applied to biological simulations and model building. Immunology has joined these efforts, and the question posed here is whether the discipline will remain committed to its theoretical concerns framed by the notions of protecting an insular self, an entity demarcated from its (...) environment, or will shift its focus of interest to a wider context. An ecological perspective emphasizes the interchange between the organism and its environment, the processing of information, and the regulation arising from responses to this larger context. Moving from the first attempts at modeling the immune system as a closed network, immunologists have joined the general interest in systems analysis, and that move might portend a significant shift to an open, more holistic consideration of immune regulation. *Received December 2006; revised August 2007. †To contact the author, please write to: Center for Philosophy and History of Science, Boston University, 745 Commonwealth Ave., Boston, MA 02215; e-mail: [email protected]. (shrink)

By comparison with the Atlantic Ocean, the physical oceanography of the Pacific was poorly known as late as the end of the 1930s. International collaboration to study the Pacific, attempted by oceanography committees of the Pacific Science Association, was a failure, owing to the scale of the enterprise, the low scientific abilities of the Pacific nations, and the lack of a compelling need. Even in the U.S.A., where the Scripps Institution of Oceanography was active, lack of good ships and personnel (...) reduced the effectiveness of Pacific oceanography. Scripps's physical oceanographer George F. McEwen was more physicist than physical oceanographer; he also became side-tracked into climatology. Only with the arrival of H. U. Sverdrup at Scripps in 1936 did Pacific physical oceanography begin to prosper, but its greatest success followed World War II, when techniques and approaches transplanted from the Atlantic became established. (shrink)

Ecology is the study of the interactions of organisms and their environments. The methods of ecology fall roughly into three categories: descriptive surveys of patterns of species and resource distribution and abundance, theoretical modeling, and experimental manipulations. Ecological systems are “open” systems, and patterns and processes are products of a huge number of interacting forces. Ecology and the environmental sciences have made enormous advances since the mid-twentieth century in the understanding of ecological systems, as well as in the human impact (...) on the environment. Theory in ecology usually centers on the development of models. Environmental outcomes are uncertain and when making decisions under uncertainty, there are a variety of options available. One option is to carry out a cost benefit analysis based upon expected utilities and other is to adopt the precautionary principle. Uncertainty and under determination of theory by evidence is a fact of life in science. (shrink)

The Swiss-born medical researcher Karl Friedrich Meyer is best known as a ‘microbe hunter’ who pioneered investigations into diseases at the intersection of animal and human health in California in the 1920s and 1930s. In particular, historians have singled out Meyer’s 1931 Ludwig Hektoen Lecture in which he described the animal kingdom as a ‘reservoir of disease’ as a forerunner of ‘one medicine’ approaches to emerging zoonoses. In so doing, however, historians risk overlooking Meyer’s other intellectual contributions. Developed in a (...) series of papers from the mid-1930s onwards, these were ordered around the concept of latent infections and sought to link microbial behavior to broader bio-ecological, environmental, and social factors that impact hostpathogen interactions. In this respect Meyer—like the comparative pathologist Theobald Smith and the immunologist Frank Macfarlane Burnet—can be seen as a pioneer of modern ideas of disease ecology. However, while Burnet’s and Smith’s contributions to this scientific field have been widely acknowledged, Meyer’s have been largely ignored. Drawing on Meyer’s published writings and private correspondence, this paper aims to correct that lacuna while contributing to a reorientation of the historiography of bacteriological epidemiology. In particular I trace Meyer’s intellectual exchanges with Smith, Burnet and the animal ecologist Charles Elton, over brucellosis, psittacosis and plague—exchanges that not only showed how environmental and ecological conditions could ‘tip the balance’ in favor of parasites but which transformed Meyer thinking about resistance to infection and disease. (shrink)

The ArgumentThe development of modern mathematical biology took place in the 1920s in three main directions: population dynamics, population genetics, and mathematical theory of epidemics. This paper focuses on the first trend which is considered the most significant. Modern mathematical theory of population dynamics is characterized by three aspects (the first two being in a somewhat critical relationship): the emergence of the mathematical modeling approach, the attempt at establishing it in a reductionist-mechanist conceptual framework, and the revival of Darwinism. The (...) first section is devoted to the analysis of the concept of mathematical model and the second one presents an example of a mathematical model (Van der Pol's model of heartbeat) which is a good prototype of that concept. In section 3 the main trends of mathematization of biology and the cultural and scientific contexts in which they found their development are discussed. Sections 4 and 5 are devoted to the contributions of V. Volterra and A. J. Lotka, to the analysis of the differences of their scientific conceptions, and to a discussion of a case study: the priority dispute concerning the discovery of the Volterra-Lotka equations. The historical analysis developed in this paper is also intended to detect the roots of some recent trends of mathematization of biology. (shrink)

Ecology has had a lower profile in Biology & Philosophy than one might expect on the basis of the attention ecology is given in public discussions in relation to environmental issues. Our tentative explanation is that ecology appears theoretically redundant within biology and, consequently, philosophically challenging problemsrelated to biology are commonly supposed to be somewhere else, particularly in the molecular sphere. Richard Levins has recognized the genuine challenges posed by ecology for theoretical and philosophical thinking in biology. This essay sets (...) the stage for appreciating his work; it was preceded by four articles published in Biology & Philosophy 15(2),and is followed by a personal reminiscent. (shrink)

This work applies the competitive exclusion principle and the concept of potential competitors as simple axiomatic tools to generalized situations in ecology. These tools enable apparent competition and its dual counterpart to be explicitly evaluated in poorly understood ecological systems. Within this set-theory framework we explore theoretical symmetries and invariances, De Morgan’s laws, frozen evolutionary diversity and virtual processes. In particular, we find that the exclusion principle compromises the geometrical growth of the number of species. By theoretical extending this principle, (...) we can describe interspecific depredation in the dual case. This study also briefly considers the debated situation of intraspecific competition. The ecological consequences of our findings are discussed; particularly, the use of our framework to reinterpret coupled mathematical differential equations describing certain ecological processes. (shrink)