Pierrick Bourrat

This thesis examines the theoretical and philosophical underpinnings of the concept of natural selection which is pervasively invoked in biology and other ‘evolutionary’ domains. Although what constitutes the process of natural selection appears to be very intuitive (natural selection results from entities exhibiting differences in fitness in a population), this conceals a number of theoretical ambiguities and difficulties. Some of these have been pointed out numerous times; others have hardly been noticed. One aim of this work is to unpack these difficulties and ambiguities; another is to provide new solutions and clarifications to them using a range of philosophical and conceptual tools. The result is a concept of natural selection stripped down from its biological specificities. I start by revisiting the entangled debates over whether natural selection is a cause of evolutionary change as opposed to a mere statistical effect of other causes, at what level this putative cause operates and whether it can be distinguished from drift. Borrowing tools from the causal modelling literature, I argue that natural selection is best conceived as a causal process resulting from individual level differences in a population. I then move to the question of whether the process of natural selection requires perfect transmission of types. I show that this question is ambiguous and can find different answers. From there, I distinguish the process of natural selection from some of its possible products, namely, evolution by natural selection and complex adaptation. I argue that reproduction and inheritance are conceptually distinct from natural selection, and using individual-based models, I demonstrate that they can be conceived as evolutionary products of it. This ultimately leads me to generalise the concepts of heritability and fitness used in the formal equations of evolutionary change. Finally, I argue that concepts of fitness and natural selection crucially depend on the grains of description at and temporal scales over which evolutionary explanations are given. These considerations reveal that the metaphysical status of the process of natural selection is problematic and why neglecting them can lead to flawed arguments in the levels of selection debate.

Reputation monitoring and the punishment of cheats are thought to be crucial to the viability and maintenance of human cooperation in large groups of non-kin. However, since the cost of policing moral norms must fall to those in the group, policing is itself a public good subject to exploitation by free riders. Recently, it has been suggested that belief in supernatural monitoring and punishment may discourage individuals from violating established moral norms and so facilitate human cooperation. Here we use cross-cultural survey data from a global sample of 87 countries to show that beliefs about two related sources of supernatural monitoring and punishment — God and the afterlife —independently predict respondents' assessment of the justifiability of a range of moral transgressions. This relationship holds even after controlling for frequency of religious participation, country of origin, religious denomination and level of education. As well as corroborating experimental work, our findings suggest that, across cultural and religious backgrounds, beliefs about the permissibility of moral transgressions are tied to beliefs about supernatural monitoring and punishment, supporting arguments that these beliefs may be important promoters of cooperation in human groups.

Cooperation for the public good is vulnerable to exploitation by free-riders because it always pays individuals to exploit the social contract for their own benefit. This problem can be resolved if free-riders are punished, but punishment is itself a public good subject to free-riding. The fear of supernatural punishment hypothesis (FSPH) proposes that belief in supernatural punishment might offer a solution to this problem by deflecting the cost of punishment onto supernatural forces and thereby incentivizing cooperation. FSPH is supported empirically by ethnographic data, but this work has so far focused on (1) institutional cooperative traits which may not reflect individual choices on how to behave in everyday social interactions and (2) threat of punishment from all-powerful moralizing high gods rather than other agents capable of supernatural punishment. Here, we consider the FSPH using variables which are linked to individual interaction and expand the number of variables measuring belief in different forms of supernatural punishment. Our findings do not fit these more general FSPH predictions. We suggest there may be something special about the link between moralizing high gods and institutional enforcement of cooperation that is not captured by these other variables.

Evolutionary transitions in individuality (ETIs) involve the formation of Darwinian collectives from Darwinian particles. The transition from cells to multicellular life is a prime example. During an ETI, collectives become units of selection in their own right. However, the underlying processes are poorly understood. One observation used to identify the completion of an ETI is an increase in collective-level performance accompanied by a decrease in particle-level performance, for example measured by growth rate. This seemingly counterintuitive dynamic has been referred to as fitness decoupling and has been used to interpret both models and experimental data. Extending and unifying results from the literature, we show that fitness of particles and collectives can never decouple because calculations of fitness performed over appropriate and equivalent time intervals are necessarily the same provided the population reaches a stable collective size distribution. By way of solution, we draw attention to the value of mechanistic approaches that emphasise traits, and tradeoffs among traits, as opposed to fitness. This trait-based approach is sufficient to capture dynamics that underpin evolutionary transitions. In addition, drawing upon both experimental and theoretical studies, we show that while early stages of transitions might often involve tradeoffs among particle traits, later—and critical—stages are likely to involve the rupture of such tradeoffs. Thus, when observed in the context of ETIs, tradeoff-breaking events stand as a useful marker of these transitions.