Concepts of Drift and Selection in “The Great Snail Debate” of the 1950s and Early 1960s

Recently, much philosophical discussion has centered on the best way to characterize the concepts of random drift and natural selection, and, in particular, on the question of whether selection and drift can be conceptually distinguished (Beatty 1984; Brandon 2005; Hodge 1983, 1987; Millstein 2002, 2005; Pfeifer 2005; Shanahan 1992; Stephens 2004). These authors all contend, to a greater or lesser degree, that their concepts make sense of biological practice. So, it should be instructive to see how the concepts of drift and selection were distinguished by the disputants in a high-profile debate; debates such as these often force biologists to take a more philosophical turn, discussing the concepts at issue in greater detail than usual. A prime candidate for just such a case study is what William Provine (1986) has termed “The Great Snail Debate,” that is, the debate over the highly polymorphic land snails Cepaea nemoralis and Cepaea hortensis in the 1950s and early 1960s. This study will reveal that much of the present-day confusion over the concepts of drift and selection is rooted in confusions of the past. Nonetheless, there are lessons that can be learned about nonadaptiveness, indiscriminate sampling, and causality with respect to these two concepts. In particular, this paper will shed light on the following questions: 1) What is “drift”? Is “drift” a purely mathematical construct, a physical process analogous to the indiscriminate sampling of balls from an urn, or the outcome of a sampling process? 2) What is “nonadaptiveness,” and is a proponent of drift committed to claims that organisms’ traits are nonadaptive? 3) Can disputes concerning selection and drift be settled by statistics alone, or is causal information essential? If causal information is essential, what does that say about the concepts of “drift” and “selection” themselves

Understanding Leopold’s Concept of ‘Interdependence’ for Environmental Ethics and Conservation Biology

Aldo Leopold’s Land Ethic, an extremely influential view in environmental ethics and conservation biology, is committed to the claim that interdependence between humans, other species, and abiotic entities plays a central role in our ethical responsibilities. Thus, a robust understanding of “interdependence” is necessary for evaluating the viability of the Land Ethic and related views, including ecological ones. I characterize and defend a Leopoldian concept of “interdependence,” arguing that it ought to include both negative and positive causal relations. I also show that strength and type of interdependence can vary with time, space, and context

Evolution

This paper is an overview of the philosophy of evolution past, present, and future to be published in Blackwell's Guide to the Philosophy of Science, edited by P.K. Machamer and M. Silberstein. It surveys the following topics: the neutralist/selectionist debate, the adapationist programme and its challenges, sociobiology, contingency, laws of biology, the species category problem, the species taxon problem, the tautology problem, fitness, units of selection

Environmental Ethics

A number of areas of biology raise questions about what is of value in the natural environment and how we ought to behave towards it: conservation biology, environmental science, and ecology, to name a few. Based on my experience teaching students from these and similar majors, I argue that the field of environmental ethics has much to teach these students. They come to me with pent-up questions and a feeling that more is needed to fully engage in their subjects, and I believe some exposure to environmental ethics can help focus their interests and goals. I identify three primary areas in which environmental ethics can con- tribute to their education. The first is an examination of who (or what) should be considered to be part of our moral community (i.e., the community to whom we owe direct duties). Is it humans only? Or does it include all sentient life? Or all life? Or ecosystems considered holistically? Often, readings implicitly assume one or more of these answers; the goal is to make the student more sensitive to these implicit claims and to get them to think about the different reasons that support them. The second area, related to the first, is the application of the different answers concerning the extent of the ethical community to real environmental issues and problems. Students need to be aware of how the different answers concerning the moral community can imply conflicting answers for how we should act in certain cases and to think about ways to move toward conflict resolution. The third area in which environmental ethics can contribute is a more conceptual one, focusing on central concepts such as biodiversity, sustainability, species, and ecosystems. Exploring and evaluating various meanings of these terms will make students more reflective and thoughtful citizens and biologists, sensitive to the implications that different conceptual choices make

Functions and Functioning in Aldo Leopold’s Land Ethic and in Ecology

I examine the use of the term function in Aldo Leopold's land ethic, invoked as: 1) the healthy functioning of the land community, which is dependent on 2) the maintenance of the characteristic functions of populations that are parts of the land community. The latter can be understood as referring to interactions between species that are the products of coevolution (such as parasite-host, predator-prey, etc.), and thus, in terms of the “selected effect” account of function. The performance of these functions under certain conditions maintain what Leopold took to be the healthy functioning of a land community

Probability in Biology: The Case of Fitness

I argue that the propensity interpretation of fitness, properly understood, not only solves the explanatory circularity problem and the mismatch problem, but can also withstand the Pandora’s box full of problems that have been thrown at it. Fitness is the propensity (i.e., probabilistic ability, based on heritable physical traits) for organisms or types of organisms to survive and reproduce in particular environments and in particular populations for a specified number of generations; if greater than one generation, “reproduction” includes descendants of descendants. Fitness values can be described in terms of distributions of propensities to produce varying number of offspring and can be modeled for any number of generations using computer simulations, thus providing both predictive power and a means for comparing the fitness of different phenotypes. Fitness is a causal concept, most notably at the population level, where fitness differences are causally responsible for differences in reproductive success. Relative fitness is ultimately what matters for natural selection

Evolution

This paper is an overview of the philosophy of evolution - past, present, and future - to be published in The Blackwell Guide to the Philosophy of Science, edited by P.K. Machamer and M. Silberstein. It surveys the following topics: the neutralist/selectionist debate, the adapationist programme and its challenges, sociobiology, contingency, laws of biology, the species category problem, the species taxon problem, the tautology problem, fitness, units of selection

Types of Experiments and Causal Process Tracing: What Happened on the Kaibab Plateau in the 1920s

In a well-cited book chapter, ecologist Jared Diamond characterizes three main types of experiment performed in community ecology: laboratory experiment, field experiment, and natural experiment. Diamond argues that each form of experiment has strengths and weaknesses, with respect to, for example, realism or the ability to follow a causal trajectory. But does Diamond’s typology exhaust the available kinds of cause-finding practices? Some social scientists have characterized something they call “causal process tracing.” Is this a fourth type of experiment or something else? I examine Diamond’s typology and causal process tracing in the context of a case study concerning the dynamics of wolf and deer populations on the Kaibab Plateau in the 1920s, a case that has been used as a canonical example of a trophic cascade by ecologists but which has also been subject to controversy. I argue that ecologists have profitably deployed causal process tracing together with other types of experiment to help settle questions of causality in this case. It remains to be seen how widespread the use of causal process tracing outside of the social sciences is (or could be), but there are some potentially promising applications, particularly with respect to questions about specific causal sequences

Debunking Myths About Aldo Leopold’s Land Ethic

Aldo Leopold’s land ethic has been extremely influential among people working in conservation biology, environmental ethics, and related fields. Others have abandoned the land ethic for purportedly being outdated or ethically untenable. Yet, both acceptance of the land ethic and rejection of the land ethic are often based on misunderstandings of Leopold’s original meaning – misunderstandings that have become so entrenched as to have the status of myths. This essay seeks to identify and then debunk six myths that have grown up around the land ethic. These myths include misunderstandings about how we should understand key terms like “stability” and “biotic community” as well as the scope and main message of the land ethic. Properly understanding Leopold’s original meaning, a meaning derived from ideas he developed after a lifetime of scientific theorizing and hands-on practical knowledge, prevents hasty rejection and provides a sounder basis for conservation policy

Thinking about populations and races in time

Biologists and philosophers have offered differing concepts of biological race. That is, they have offered different candidates for what a biological correlate of race might be; for example, races might be subspecies, clades, lineages, ecotypes, or genetic clusters. One thing that is striking about each of these proposals is that they all depend on a concept of population. Indeed, some authors have explicitly characterized races in terms of populations. However, including the concept of population into concepts of race raises three puzzles, all having to do with time. In this paper, I extend the causal interactionist population concept (CIPC) by introducing some simple assumptions about how to understand populations through time. These assumptions help to shed light on the three puzzles, and in the process show that if we want to understand races in terms of populations, we will need to revise our concept(s) of race