The concept of fitness is central to evolutionary biology. Models of evolutionary change typically use some quantity called “fitness” which measures an organism’s reproductive success. But what exactly does it mean that fitness is such a measure? In what follows, we look at the interplay between abstract evolutionary models and quantitative measures of fitness and develop a measurement-theoretic perspective on fitness in order to explore what makes certain measures of fitness significant.

The origins of human social cooperation confound simple evolutionary explanation. But from Darwin and Durkheim onward, theorists (anthropologists and sociologists especially) have posited a potential link with another curious and distinctively human social trait that cries out for explanation: religion. This dissertation explores one contemporary theory of the co-evolution of religion and human social cooperation: the signalling theory of religion, or religious signalling theory (RST). According to the signalling theory, participation in social religion (and its associated rituals and sanctions) acts (...) as an honest signal of one’s commitment to a religiously demarcated community and its way of doing things. This signal would allow prosocial individuals to positively assort with one another for mutual advantage, to the exclusion of more exploitative individuals. In effect, the theory offers a way that religion and cooperation might explain one another, but which that stays within an individualist adaptive paradigm. My approach is not to assess the empirical adequacy of the religious signalling explanation or contrast it with other explanations, but rather to deal with the theory in its own terms – isolating and fleshing out its core commitments, explanatory potential, and limitations. The key to this is acknowledging the internal complexities of signalling theory, with respect to the available models of honest signalling and the extent of their fit (or otherwise) with religion as a target system. The method is to take seriously the findings of formal modelling in animal signalling and other disciplines, and to apply these (and methods from the philosophy of biology more generally) to progressively build up a comprehensive picture of the theory, its inherent strengths and weaknesses. The first two chapters outline the dual explanatory problems that cooperation and religion present for evolutionary human science, and surveys contemporary approaches toward explaining them. Chapter three articulates an evolutionary conception of the signalling theory, and chapters four to six make the case for a series of requirements, limitations, and principles of application. Chapters seven and eight argue for the value of formal modelling to further flesh out the theory’s commitments and potential and describe some simple simulation results which make progress in this regard. Though the inquiry often problematizes the signalling theory, it also shows that it should not be dismissed outright, and that it makes predictions which are apt for empirical testing. (shrink)

Inclusive fitness has been under intense scrutiny in recent years, with many critics claiming the framework leads to incorrect predictions. We consider one particularly influential heuristic for estimating inclusive fitness in the context of the very case that motivated reliance on it to begin with: the Sir Philip Sidney signalling game played with relatives. Using a neighbour-modulated fitness model, we show when and why this heuristic is problematic. We argue that reliance on the heuristic rests on a misunderstanding of what (...) it means for two organisms to be related and perpetuates a mischaracterization of the role of the ‘relatedness’ parameter in inclusive fitness. _1_ Introduction _2_ Heuristic Inclusive Fitness _3_ The Sir Philip Sidney Game _4_ Model _5_ Results _6_ Conclusion Appendix. (shrink)

I distinguish two roles for a fitness concept in the context of explaining cumulative adaptive evolution: fitness as a predictor of gene frequency change, and fitness as a criterion for phenotypic improvement. Critics of inclusive fitness argue, correctly, that it is not an ideal fitness concept for the purpose of predicting gene-frequency change, since it relies on assumptions about the causal structure of social interaction that are unlikely to be exactly true in real populations, and that hold as approximations only (...) given a specific type of weak selection. However, Hamilton took this type of weak selection, on independent grounds, to be responsible for cumulative assembly of complex adaptations. In this special context, I argue that inclusive fitness is distinctively valuable as a criterion for improvement and a standard for optimality. Yet to call inclusive fitness a criterion for improvement and a standard for optimality is not to make any claim about the frequency with which inclusive fitness optimization actually occurs in nature. This is an empirical question that cannot be settled by theory alone. I close with some reflections on the place of inclusive fitness in the long running clash between ‘causalist’ and ‘statisticalist’ conceptions of fitness. (shrink)

Altruistic deception (or the telling of “white lies”) is common in humans. Does it also exist in non-human animals? On some definitions of deception, altruistic deception is impossible by definition, whereas others make it too easy by counting useful-but-ambiguous information as deceptive. I argue for a definition that makes altruistic deception possible in principle without trivializing it. On my proposal, deception requires the strategic exploitation of a receiver by a sender, where “exploitation” implies that the sender elicits a behaviour in (...) the receiver that is beneficial in a different type of situation and is expressed only because the signal raises the probability, from the receiver’s standpoint, of that type of situation. I then offer an example of a real signal that is deceptive in this sense, and yet potentially altruistic (and certainly cooperative): the purr call of the pied babbler. Fledglings associate purr calls with food, and adults exploit this learned association, in the absence of food, to lead fledglings away from predators following an alarm call. I conclude by considering why altruistic deception is apparently so rare in non-human animals. (shrink)