This article proposes two conditions to assess whether an entity at a level of description is a unit of selection qua interactor. These two conditions make it possible to (1) distinguish biologically relevant entities from arbitrary ones and (2) distinguish units that can _potentially_ enter a selection process from those that have already done so. I show that the classical approaches used in the literature on units and levels of selection do not fare well with respect to either or both (...) of these desiderata. (shrink)

The levels of selection debate is generally taken to be a debate about how natural selection can occur at the various levels of biological organization. In this paper, we argue that questions about levels of selection should be analyzed separately from questions about levels of organization. In the deflationary proposal we defend, all that is necessary for multilevel selection is that there are cases in which particles are nested in collectives, and that both the collectives and the particles that compose (...) them each separately undergo natural selection. We argue that adopting this deflationary account helps to disentangle the levels of selection and the levels of organization, and thereby contributes to advancing the levels of selection debate. (shrink)

It has become customary in multilevel selection theory to use the same terms to denote both two explanatory goals and two explanatory means. This paper spells out some of the benefits that derive from avoiding this terminological conflation. I argue that keeping explanatory means and goals well apart allows us to see that, contrary to a popular recent idea, Price’s equation and contextual analysis—the statistical methods most extensively used for measuring the effects of certain evolutionary factors on the change in (...) the focal individual trait in multilevel selection scenarios—do not come with built-in notions of group selection and, therefore, the efficacy of these methods at analyzing various kinds of cases does not constitute a basis for deciding how group selection should best be defined. Moreover, contrary to another widely accepted idea, I argue that more than one type of group selection may serve as explanatory means when one’s goal is that of explaining the evolution of individual traits in multilevel selection scenarios and I spell out how this explanatory role should be understood. (shrink)

The contextual approach is a prominent framework for thinking about group selection. Here, I highlight ambiguity about what the contextual approach is. Then, I discuss problematic entailments the contextual approach has for what processes count as group selection—entailments more troublesome than typically noted. However, Sober and Wilson’s version of the Price approach, which is the main alternative to the contextual approach, is problematic too: it leads to an underappreciated paradox called the vanishing selection problem and thereby generates the wrong qualitative (...) account of whether group selection is occurring in a certain family of cases. In response, I develop an account of group selection that can deal with the counterexamples to both the contextual approach and the Price approach. I then discuss the role that contextual analysis can continue to play in the discussion of individual fitness and metapopulation evolution. (shrink)

Debates concerning the units and levels of selection have persisted for over fifty years. One major question in this literature is whether units and levels of selection are genuine, in the sense that they are objective features of the world, or merely reflect the interests and goals of an observer. Scientists and philosophers have proposed a range of answers to this question. This Element introduces this literature and proposes a novel contribution. It defends a realist stance and offers a way (...) of delineating genuine levels of selection by invoking the notion of a functional unit. (shrink)

In this paper, I identify two general positions with respect to the relationship between environment and natural selection. These positions consist in claiming that selective claims need and, respectively, need not be relativized to homogenous environments. I then show that adopting one or the other position makes a difference with respect to the way in which the effects of selection are to be measured in certain cases in which the focal population is distributed over heterogeneous environments. Moreover, I show that (...) these two positions lead to two different interpretations—the Pricean and contextualist ones—of a type of selection scenarios in which multiple groups varying in properties affect the change in the metapopulation mean of individual-level traits. Showing that these two interpretations stem from different attitudes towards environmental homogeneity allows me to argue: that, unlike the Pricean interpretation, the contextualist interpretation can only claim that drift or selection is responsible for the change in frequency of the focal trait in a given metapopulation if details about whether or not group formation is random are specified; that the traditional main objection against the Pricean interpretation—consisting in arguing that the latter takes certain side-effects of individual selection to be effects of group selection—is unconvincing. This leads me to suggest that the ongoing debate about which of the two interpretations is preferable should concentrate on different issues than previously thought. (shrink)

In "Some Criticism of the Contextual Approach, and a Few Proposals" in Biological Theory, Brian McLoone discusses some of the points about the contextual approach that I made in a recent paper. Besides offering a reply to McLoone’s comments on my paper, in this article I show why McLoone’s discussion of the two main frameworks for thinking about group selection—the contextual and the Price approach—is partly misguided. In particular, I show that one of McLoone’s main arguments against the contextual approach (...) is missing the target and that one of his (and Elliott Sober’s) arguments in defense of the Price approach is flawed. Criticizing these arguments will help me present an entirely different picture than McLoone’s of the current status of multilevel selection theory. More precisely, I argue that the idea that we are dealing with "multilevel selection" in the type of multigroup cases in which the focal units are the individuals (and their traits) has recently come under threat. Finally, I discuss the ways in which this idea might be salvaged by appealing either to the contextual or to the Price approach. (shrink)

Evolutionary transitions in individuality (ETIs) are events during which individuals at a given level of organization (particles) interact to form higher-level entities (collectives) which are then recognized as new individuals at that level. ETIs are intimately related to levels of selection, which, following Okasha, can be approached from two different perspectives. One, referred to as ‘synchronic’, asks whether selection occurs at the collective level while the partitioning of particles into collectives is taken for granted. The other, referred to as ‘diachronic’, (...) asks about the origins of the partitioning of particles into collectives. After having presented the two perspectives and a classical formalism used to deal with the levels-of-selection question in the literature, namely the multilevel version of the Price equation, I show that because this formalism treats the levels-of-selection question from a synchronic perspective, it is inadequate to explain ETIs. This is because a fundamental aspect of ETIs is the origin of collectives. From there, I develop a framework for levels of selection compatible with the diachronic perspective. This framework relies on the distinction between what I call, on the one hand, a ‘functional aggregative collective trait’, and on the other hand, a ‘functional non-aggregative collective trait’. After having presented this distinction, I implement it in the Price equation, leading to a new statistical partitioning of this equation which, I argue, represents a causal decomposition more relevant for ETIs. Finally, I exploit this partitioning to explain the critical stages of an ETI. In addition to its explanatory power, a measure of the degree of functional non-aggregativity could be used as a proxy for the degree of individuality of a collective. (shrink)