This dissertation is about the extent to which natural selection has causal power. In Chapter 1, I articulate and defend a version of an interventionist account of causation. Put simply, I claim that some event e causes another event e' in a causal system S just in case a precise manipulation to the value of e, while holding various other events in S fixed, results in a change to the probability of e'. Also in Chapter 1, I endorse a recent constraint on causal relata that James Woodward has endorsed, which requires that it be possible in principle to manipulate each relatum in a causal system while simultaneously holding fixed the other relata in the causal system at arbitrary values. In Chapter 2, I use this account of causation to determine the extent to which natural selection can be treated as a cause of other evolutionary phenomena. Through a variety of examples, I show that natural selection can cause trait evolution, as well as the origin of novel traits. I also show that, given various assumptions, there are cases in which natural selection can cause a particular individual to have one trait, rather than another. In Chapter 3, I analyze the extent to which natural selection can operate at various levels of the biological hierarchy, in particular among individuals within a group and between groups in a group-structured population. For reasons that I describe, while "group selection" is a coherent concept, I am dissatisfied with various accounts of that concept. As I develop my own account, I comment more broadly on the way in which we can understand natural selection and causation in a group-structured population, given the variety of non-causal dependency relationships that exist in such a setting

Recent arguments concerning the nature of causation in evolutionary theory, now often known as the debate between the 'causalist' and 'statisticalist' positions, have involved answers to a variety of independent questions – definitions of key evolutionary concepts like natural selection, fitness, and genetic drift; causation in multi-level systems; or the nature of evolutionary explanations, among others. This Element offers a way to disentangle one set of these questions surrounding the causal structure of natural selection. Doing so allows us to clearly reconstruct the approach that some of these major competing interpretations of evolutionary theory have to this causal structure, highlighting particular features of philosophical interest within each. Further, those features concern problems not exclusive to the philosophy of biology. Connections between them and, in two case studies, contemporary metaphysics and philosophy of physics demonstrate the potential value of broader collaboration in the understanding of evolution.

A comprehensive treatment of the concept of causation in evolutionary biology that makes clear its central role in both historical and contemporary debates. Most scientific explanations are causal. This is certainly the case in evolutionary biology, which seeks to explain the diversity of life and the adaptive fit between organisms and their surroundings. The nature of causation in evolutionary biology, however, is contentious. How causation is understood shapes the structure of evolutionary theory, and historical and contemporary debates in evolutionary biology have revolved around the nature of causation. Despite its centrality, and differing views on the subject, the major conceptual issues regarding the nature of causation in evolutionary biology are rarely addressed. This volume fills the gap, bringing together biologists and philosophers to offer a comprehensive, interdisciplinary treatment of evolutionary causation. Contributors first address biological motivations for rethinking evolutionary causation, considering the ways in which development, extra-genetic inheritance, and niche construction challenge notions of cause and process in evolution, and describing how alternative representations of evolutionary causation can shed light on a range of evolutionary problems. Contributors then analyze evolutionary causation from a philosophical perspective, considering such topics as causal entanglement, the commingling of organism and environment, and the relationship between causation and information. Contributors John A. Baker, Lynn Chiu, David I. Dayan, Renée A. Duckworth, Marcus W Feldman, Susan A. Foster, Melissa A. Graham, Heikki Helanterä, Kevin N. Laland, Armin P. Moczek, John Odling-Smee, Jun Otsuka, Massimo Pigliucci, Arnaud Pocheville, Arlin Stoltzfus, Karola Stotz, Sonia E. Sultan, Christoph Thies, Tobias Uller, Denis M. Walsh, Richard A. Watson

The Nature of Selection is a straightforward, self-contained introduction to philosophical and biological problems in evolutionary theory. It presents a powerful analysis of the evolutionary concepts of natural selection, fitness, and adaptation and clarifies controversial issues concerning altruism, group selection, and the idea that organisms are survival machines built for the good of the genes that inhabit them. "Sober's is the answering philosophical voice, the voice of a first-rate philosopher and a knowledgeable student of contemporary evolutionary theory. His book merits broad attention among both communities. It should also inspire others to continue the conversation."-Philip Kitcher, Nature "Elliott Sober has made extraordinarily important contributions to our understanding of biological problems in evolutionary biology and causality. The Nature of Selection is a major contribution to understanding epistemological problems in evolutionary theory. I predict that it will have a long lasting place in the literature."-Richard C. Lewontin

This illuminating volume explores the effects of chance on evolution, covering diverse perspectives from scientists, philosophers, and historians. The evolution of species, from single-celled organisms to multicellular animals and plants, is the result of a long and highly chancy history. But how profoundly has chance shaped life on earth? And what, precisely, do we mean by chance? Bringing together biologists, philosophers of science, and historians of science, Chance in Evolution is the first book to untangle the far-reaching effects of chance, contingency, and randomness on the evolution of life. The book begins by placing chance in historical context, starting with the ancients and moving through Darwin to contemporary biology. It documents the shifts in our understanding of chance as Darwin’s theory of evolution developed into the modern synthesis, and how the acceptance of chance in Darwinian theory affected theological resistance to it. Other chapters discuss how chance relates to the concepts of genetic drift, mutation, and parallel evolution—as well as recent work in paleobiology and the experimental evolution of microbes. By engaging in collaboration across biology, history, philosophy, and theology, this book offers a comprehensive overview both of the history of chance in evolution and of our current understanding of the impact of chance on life.

No other scientific theory has had as tremendous an impact on our understanding of the world as Darwin's theory as outlined in his Origin of Species, yet from the very beginning the theory has been subject to controversy. The Evolution of Darwinism, first published in 2004, focuses on three issues of debate - the nature of selection, the nature and scope of adaptation, and the question of evolutionary progress. It traces the varying interpretations to which these issues were subjected from the beginning and the fierce contemporary debates that still rage on and explores their implications for the greatest questions of all: Where we come from, who we are and where we might be heading. Written in a clear and non-technical style, this book will be of use as a textbook for students in the philosophy of science who need to become familiar with the background to the debates about evolution.

Does natural selection act primarily on individual organisms, on groups, on genes, or on whole species? Samir Okasha provides a comprehensive analysis of the debate in evolutionary biology over the levels of selection, focusing on conceptual, philosophical and foundational questions. A systematic framework is developed for thinking about natural selection acting at multiple levels of the biological hierarchy; the framework is then used to help resolve outstanding issues. Considerable attention is paid to the concept of causality as it relates to the levels of selection, in particular the idea that natural selection at one hierarchical level can have effects that 'filter' up or down to other levels. Unlike previous work in this area by philosophers of science, full account is taken of the recent biological literature on 'major evolutionary transitions' and the recent resurgence of interest in multi-level selection theory among biologists. Other biological topics discussed include Price's equation, kin and group selection, the gene's eye view, evolutionary game theory, outlaws and selfish genetic elements, species and clade selection, and the evolution of individuality. Philosophical topics discussed include reductionism and holism, causation and correlation, the nature of hierarchical organization, and realism and pluralism.