Roberta L. Millstein

The concept of “land community” (or “biotic community”) that features centrally in Aldo Leopold’s Land Ethic has typically been equated with the concept of “ecosystem.” The author argues that we need to rethink Leopold’s concept of land community. First, Leopold’s views are not identical to those of his contemporaries, although they resemble those of some subsequent ecologists. Second, the land community concept does not map cleanly onto the concept of “ecosystem”; it also incorporates elements of the “community” concept in community ecology. Third, the question of whether land communities have boundaries can be addressed by an analysis of land communities as individuals. There are challenges to be worked out, but the author argues that these challenges can be resolved. The result is a defensible land community concept that is ontologically robust enough to be a locus of moral obligation while being consistent with contemporary ecological theory and practice.

Recent work on the heat-shock protein Hsp90 by Rutherford and Lindquist (1998) has been included among the pieces of evidence taken to show the essential role of developmental processes in evolution; Hsp90 acts as a buffer against phenotypic variation, allowing genotypic variation to build. When the buffering capacity of Hsp90 is altered (e.g., in nature, by mutation or environmental stress), the genetic variation is “revealed,” manifesting itself as phenotypic variation. This phenomenon raises questions about the genetic variation before and after what I will call a “revelation event”: Is it neutral, nearly neutral, or non-neutral (i.e., strongly deleterious or strongly advantageous)? Moreover, what kinds of evolutionary processes do we take to be at work? Rutherford and Lindquist (1998) focus on the implications of non-neutral variation and selection. Later works by Queitsch, Sangster, and Lindquist (2002) and Sangster, Lindquist, and Queitsch (2004) raise the possibility that Hsp90 buffering may play the role that was played by drift in Sewall Wright’s shifting balance model, permitting transition from one adaptive peak to another. However, Ohta (2002) suggests that much of this variation may be nearly neutral, which in turn, would imply a strong role for drift as well as selection. The primary goal of this article is to illuminate the alternative scenarios and the processes operating in each. At the end, I raise the possibility of a synthesis between EvoDevo and nearly neutral evolution.

In “‘Population’ is Not a Natural Kind of Kinds,” Jacob Stegenga argues against the claim that the concept of “population” is a natural kind and in favor of conceptual pluralism, ostensibly in response to two papers of mine (Millstein 2009, 2010). Pluralism is often an attractive position in the philosophy of science. It certainly is a live possibility for the concept of population in ecology and evolutionary biology, and I welcome the opportunity to discuss the topic further. However, I argue that the case for conceptual pluralism has not yet been made. In what follows, I first clarify the issues at stake before taking up the topic of conceptual pluralism and responding to Stegenga’s criticisms of the causal interactionist population concept.

The controversy over genetically modified [GM] organisms is often framed in terms of possible hazards for human health. Articles in a previous volume of this Encyclopedia give a general overview of GM crops (Mulvaney 2014) and specifically examine human health (Nordgard, Gronsberg, and Myhr 2014) and labeling (Bruton 2014) issues surrounding GM organisms. This article explores several other aspects of the controversy: environmental concerns, political and legal disputes, and the aim of “feeding the world” and promoting food security. Rather than discussing abstract, hypothetical GM organisms, this article explores the consequences of the GM organisms that have actually been deployed in the particular contexts that they have been deployed, on the belief that there is little point in discussing GM organisms in an idealized or context-independent way

Informed by his experiences as a hunter, forester, wildlife manager, ecologist, conservationist, and professor, Aldo Leopold developed a view he called the land ethic. In a classic essay, published posthumously in A Sand County Almanac, Leopold advocated for an expansion of our ethical obligations beyond the purely human to include what he variously termed the “land community” or the “biotic community”—communities of interdependent humans, nonhuman animals, plants, soils, and waters, understood collectively. This philosophy has been extremely influential in environmental ethics as well as conservation biology and related fields. Using an approach grounded in environmental ethics and the history and philosophy of science, Roberta L. Millstein reexamines Leopold’s land ethic in light of contemporary ecology. Despite the enormous influence of the land ethic, it has sometimes been dismissed as either empirically out of date or ethically flawed. Millstein argues that these dismissals are based on problematic readings of Leopold’s ideas. In this book, she provides new interpretations of the central concepts underlying the land ethic: interdependence, land community, and land health. She also offers a fresh take on of his argument for extending our ethics to include land communities as well as Leopold-inspired guidelines for how the land ethic can steer conservation and restoration policy.

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) 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.

In their book, Defending Biodiversity, Newman, Varner, and Linquist (NVL) cast doubt on whether Leopoldian defenses of biodiversity, in their current form, have been successful. I argue that there is a more accurate interpretation of Leopold that is not subject to the criticisms made by NVL, and that Leopold’s body of work as a whole, including but not limited to the essay “The Land Ethic” in A Sand County Almanac, provides quite a bit of useful guidance and perspective. I begin with a brief summary of some of my own recent work on Leopold. This is intended to orient the reader who is more familiar with J. Baird Callicott’s influential interpretations of Leopold. I then discuss NVL’s Chapter 10 first followed by a discussion of their Chapter 9, responding to their critiques. I then conclude. On the revised interpretation that I have given, Leopold’s land ethic is defended by the method of reflective equilibrium, showing us that the land communities that we are interdependent with have intrinsic value, necessitating preserving their (land) health, which in turn necessitates preserving biodiversity.

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.

Sex and sensibility: The role of social selection Content Type Journal Article DOI 10.1007/s11016-010-9464-6 Authors Erika L. Milam, Department of History, University of Maryland, 2115 Francis Scott Key Hall, College Park, MD 20742, USA Roberta L. Millstein, Department of Philosophy, University of California, Davis, One Shields Avenue, Davis, CA 95616, USA Angela Potochnik, Department of Philosophy, University of Cincinnati, P.O. Box 210374, Cincinnati, OH 45221, USA Joan E. Roughgarden, Department of Biology, Stanford University, Stanford, CA 94305-5020, USA Journal Metascience Online ISSN 1467-9981 Print ISSN 0815-0796.

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.

The controversy over genetically modified [GM] organisms is often framed in terms of possible hazards for human health. Articles in a previous volume of this *Encyclopedia* give a general overview of GM crops [@Mulvaney2014] and specifically examine human health [@Nordgard2014] and labeling [@Bruton2014] issues surrounding GM organisms. This article explores several other aspects of the controversy: environmental concerns, political and legal disputes, and the aim of "feeding the world" and promoting food security. Rather than discussing abstract, hypothetical GM organisms, this article explores the consequences of the GM organisms that have actually been deployed in the particular contexts that they have been deployed, on the belief that there is little point in discussing GM organisms in an idealized or context-independent way.

Evolutionary theory is thoroughly probabilistic, due to such elements as random mutation, random drift, and stochastic models of speciation and extinction. This uncontroversial claim raises a number of contentious issues: Is evolutionary theory probabilistic in any general sense, or only in a variety of particular, distinct senses? Are these processes inherently probabilistic, or does the use of probabilities simply provide a convenient way to camouflage a multitude of unknown causes? Does chance play an explanatory role in evolutionary theory, and if so, how? My research explores the seemingly disparate meanings of biologists' various usages of the concept of chance in order to shed light on the answers to these questions. ;Recently, philosophers of science have looked to quantum mechanics to settle issues concerning the nature of chance. However, evolutionary theory itself has a decidedly probabilistic character, apart from quantum mechanics; evolutionary theory was given a probabilistic formulation prior to and independently of the development of quantum mechanics. Previous accounts of chance in evolutionary theory have focused primarily on one particular aspect of chance. I examine three distinct areas where chance enters into evolutionary theory in order to provide a comprehensive, synthetic account. Random drift has been the subject of much discussion by philosophers of biology, most notably by Beatty, Brandon, Hodge, Rosenberg, Shanahan, and Sober. My dissertation thus focuses on random drift in order to settle a number of unresolved controversies raised by these philosophers. The discussion of random drift provides a template for exploring random mutation and stochastic models of macroevolution. On the basis of these analyses, I argue that evolutionary theory provides reasons to rethink some of our traditional philosophical ideas about chance

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.

Evolution in its contemporary meaning in biology typically refers to the changes in the proportions of biological types in a population over time (see the entry on the concept of evolution to 1872 for earlier meanings). As evolution is too large of a topic to address thoroughly in one entry, the primary goal of this entry is to serve as a broad overview of contemporary issues in evolution with links to other entries where more in-depth discussion can be found. The entry begins with a brief survey of definitions of evolution, followed by a discussion of the different modes of evolution and related philosophical issues, and ends with a summary of other topics in the philosophy of evolution focusing particularly on topics covered in this encyclopedia.

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.

The “land community” (or “biotic community”) that features centrally in Aldo Leopold’s Land Ethic has typically been equated with the concept of “ecosystem.” Moreover, some have challenged this central Leopoldean concept given the multitude of meanings of the term “ecosystem” and the changes the term has undergone since Leopold’s time (see, e.g., Shrader-Frechette 1996). Even one of Leopold’s primary defenders, J. Baird Callicott, asserts that there are difficulties in identifying the boundaries of ecosystems and suggests that we recognize that their boundaries are determined by scientific questions ecologists pose (Callicott 2013). I argue that we need to rethink Leopold’s concept of land community in the following ways. First, we should recognize that Leopold’s views are not identical to those of his contemporaries (e.g., Clements, Elton), although they resemble those of some subsequent ecologists, including some of our contemporaries (e.g., O’Neill 2001, Post et al. 2007, Hastings and Gross 2012). Second, the land community concept does not map cleanly onto the concept of “ecosystem”; it also incorporates elements of the “community” concept in community ecology by emphasizing the interactions between organisms and not just the matter/energy flow of the ecosystem concept. Third, the boundary question can be illuminated by considering some of the recent literature on the nature of biological individuals (in particular, Odenbaugh 2007; Hamilton, Smith, and Haber 2009; Millstein 2009), focusing on concentrations of causal relations as determinative of the boundaries of the land community qua individual. There are challenges to be worked out, particularly when the interactions of community members do not map cleanly onto matter/energy flows, but I argue that these challenges can be resolved. The result is a defensible land community concept that is ontologically robust enough to be a locus of moral obligation while being consistent with contemporary ecological theory and practice.

In Darwin’s Sacred Cause, Adrian Desmond and James Moore contend that “Darwin would put his utmost into sexual selection because the subject intrigued him, no doubt, but also for a deeper reason: the theory vindicated his lifelong commitment to human brotherhood”. Without questioning Desmond and Moore’s evidence, I will raise some puzzles for their view. I will show that attention to the structure of Darwin’s arguments in the Descent of Man shows that they are far from straightforward. As Desmond and Moore note, Darwin seems to have intended sexual selection in non-human animals to serve as evidence for sexual selection in humans. However, Darwin’s account of sexual selection in humans was different from the canonical cases that Darwin described at great length. If explaining the origin of human races was the main reason for introducing sexual selection, and if sexual selection was a key piece of Darwin’s anti-slavery arguments, then it is puzzling why Darwin would have spent so much time discussing cases that did not really support his argument for the origin of human races, and it is also puzzling that his argument for the origin of human races would be so poor.

This paper aims to illustrate one of the primary goals of the philosophy of biology⎯namely, the examination of central concepts in biological theory and practice⎯through an analysis of the concepts of population and metapopulation in evolutionary biology and ecology. I will first provide a brief background for my analysis, followed by a characterization of my proposed concepts: the causal interactionist concepts of population and metapopulation. I will then illustrate how the concepts apply to six cases that differ in their population structure; this analysis will also serve to flesh out and defend the concepts a bit more. Finally, I will respond to some possible questions that my analysis may have raised and then conclude briefly.

Genetic drift (variously called “random drift”, “random genetic drift”, or sometimes just “drift”) has been a source of ongoing controversy within the philosophy of biology and evolutionary biology communities, to the extent that even the question of what drift is has become controversial. There seems to be agreement that drift is a chance (or probabilistic or statistical) element within population genetics and within evolutionary biology more generally, and that the term “random” isn’t invoking indeterminism or any technical mathematical meaning, but that’s about where agreement ends. Yet genetic drift models are a staple topic in population genetics textbooks and research, with genetic drift described as one of the main factors of evolution alongside selection, mutation, and migration. Some claim that genetic drift has played a major role in evolution (particularly molecular evolution), while others claim it to be minor. This article examines these and other controversies. In order to break through the logjam of competing definitions of drift, this entry begins with a brief history of the concept, before examining various philosophical claims about the proper characterization of drift and whether it can be distinguished from natural selection; the relation of drift to debates over statisticalism; whether drift can be detected empirically and if so, how; and the proper understanding of drift as a model and as a (purported) law.

Recently, a number of philosophers of biology have endorsed views about random drift that, we will argue, rest on an implicit assumption that the meaning of concepts such as drift can be understood through an examination of the mathematical models in which drift appears. They also seem to implicitly assume that ontological questions about the causality of terms appearing in the models can be gleaned from the models alone. We will question these general assumptions by showing how the same equation — the simple 2 = p2 + 2pq + q2 — can be given radically different interpretations, one of which is a physical, causal process and one of which is not. This shows that mathematical models on their own yield neither interpretations nor ontological conclusions. Instead, we argue that these issues can only be resolved by considering the phenomena that the models were originally designed to represent and the phenomena to which the models are currently applied. When one does take those factors into account, starting with the motivation for Sewall Wright’s and R.A. Fisher’s early drift models and ending with contemporary applications, a very different picture of the concept of drift emerges. On this view, drift is a term for a set of physical processes, namely, indiscriminate sampling processes

Biologists studying ecology and evolution use the term “population” in many different ways. Yet little philosophical analysis of the concept has been done, either by biologists or philosophers, in contrast to the voluminous literature on the concept of “species.” This is in spite of the fact that “population” is arguably a far more central concept in ecological and evolutionary studies than “species” is. The fact that such a central concept has been employed in so many different ways is potentially problematic for the reason that inconsistent usages (especially when the usage has not been made explicit) might lead to false controversies in which disputants are simply talking past one another. However, the inconsistent usages are not the only, or even the most important reason to examine the concept. If any set of organisms is legitimately called a “population,” selection and drift processes become purely arbitrary, too. Moreover, key ecological variables, such as abundance and distribution, depend on a nonarbitrary way of identifying populations. I sketch the beginnings of a population concept, drawing inspiration from the Ghiselin-Hull individuality thesis, and show why some alternative approaches are nonstarters.

Recent work on the heat-shock protein Hsp90 by Rutherford and Lindquist (1998) has been included among the pieces of evidence taken to show the essential role of developmental processes in evolution; Hsp90 acts as a buffer against phenotypic variation, allowing genotypic variation to build. When the buffering capacity of Hsp90 is altered (e.g., in nature, by mutation or environmental stress), the genetic variation is "revealed," manifesting itself as phenotypic variation. This phenomenon raises questions about the genetic variation before and after what I will call a "revelation event": Is it neutral, nearly neutral, or non-neutral (i.e., strongly deleterious or strongly advantageous)? Moreover, what kinds of evolutionary processes do we take to be at work? Rutherford and Lindquist (1998) focus on the implications of non-neutral variation and selection. Later work by Queitsch, Sangster, and Lindquist (2002) and Sangster, Lindquist, and Queitsch (2004) raises the possibility that Hsp90 buffering may play the role that was played by drift in Sewall Wright's shifting balance model, permitting transition from one adaptive peak to another. However, Ohta (2002) suggests that much of this variation may be nearly neutral, which in turn, would imply a strong role for drift as well as selection. The primary goal of this paper is to illuminate the alternative scenarios and the processes operating in each. At the end, I raise the possibility of a synthesis between evo-devo and nearly neutral evolution.

Recently, much philosophical discussion has centered on the best way to characterize the concepts of random drift and natural selection, and, in particular, 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. Moreover, it is important to consider a debate where the disputants are actually trying to apply the models of population genetics to natural populations; only then can their proper interpretations become fully apparent. (Indeed, I contend that some of the philosophical confusion has arisen because authors have considered only the models themselves, and not the phenomena that the models are attempting to represent). A prime candidate for just such a case study is what Provine (1986) has termed “The Great Snail Debate,” that is, the debates over the highly polymorphic land snails Cepaea nemoralis and C. hortensis in the 1950s and early 1960s. These studies represent one of the best, if not the best, of the early attempts to demonstrate drift in natural populations.

As a number of biologists and philosophers have emphasized, ‘chance’ has multiple meanings in evolutionary biology. Seven have been identified. I will argue that there is a unified concept of chance underlying these seven, which I call the UCC (Unified Chance Concept). I will argue that each is characterized by which causes are consid- ered, ignored, or prohibited. Thus, chance in evolutionary biology can only be under- stood through understanding the causes at work. The UCC aids in comparing the different concepts and allows us to characterize our concepts of chance in probabilistic terms, i.e. provides a way to translate between ‘chance’ and ‘probability’.

Evolutionary theory (ET) is teeming with probabilities. Probabilities exist at all levels: the level of mutation, the level of microevolution, and the level of macroevolution. This uncontroversial claim raises a number of contentious issues. For example, is the evolutionary process (as opposed to the theory) indeterministic, or is it deterministic? Philosophers of biology have taken different sides on this issue. Millstein (1997) has argued that we are not currently able answer this question, and that even scientific realists ought to remain agnostic concerning the determinism or indeterminism of evolutionary processes. If this argument is correct, it suggests that, whatever we take probabilities in ET to be, they must be consistent with either determinism or indeterminism. This raises some interesting philosophical questions: How should we understand the probabilities used in ET? In other words, what is meant by saying that a certain evolutionary change is more or less probable? Which interpretation of probability is the most appropriate for ET? I argue that the probabilities used in ET are objective in a realist sense, if not in an indeterministic sense. Furthermore, there are a number of interpretations of probability that are objective and would be consistent with ET under determinism or indeterminism. However, I argue that evolutionary probabilities are best understood as propensities of population-level kinds.