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)

This paper attempts to reconcile critics and defenders of inclusive fitness by constructing a synthesis that does justice to the insights of both. I argue that criticisms of the regression-based version of Hamilton’s rule, although they undermine its use for predictive purposes, do not undermine its use as an organizing framework for social evolution research. I argue that the assumptions underlying the concept of inclusive fitness, conceived as a causal property of an individual organism, are unlikely to be exactly true (...) in real populations, but they are approximately true given a specific type of weak selection that Hamilton took, on independent grounds, to be responsible for the cumulative assembly of complex adaptation. Finally, I reflect on the uses and limitations of “design thinking” in social evolution research. The debate about the foundations of inclusive fitness theory that has followed in the wake of Nowak, Tarnita and Wilson’s (1) critique has been remarkably polarizing. After several rounds of rebuttals and replies, there is still little evidence of any serious reconciliation between the theory’s critics (2-10) and its defenders (11-24). It doesn’t have to be this way. I believe that, on the main points of disagreement, it is possible to find a way forward that does justice to the insights of both camps. My aim in this paper is to find that way forward. (shrink)

Various results show the ‘formal equivalence’ of kin and group selectionist methodologies, but this does not preclude there being a real and useful distinction between kin and group selection processes. I distinguish individual- and population-centred approaches to drawing such a distinction, and I proceed to develop the latter. On the account I advance, the differences between kin and group selection are differences of degree in the structural properties of populations. A spatial metaphor provides a useful framework for thinking about these (...) differences: kin and group selection may be conceptualized as large, overlapping regions of K - G space. I then consider some implications of the account, defend it from possible objections, and further argue that the structural features characteristic of both kin and group selection may recur at multiple levels of biological organization. (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)

Hamilton introduced two conceptions of social fitness, which he called neighbour-modulated fitness and inclusive fitness. Although he regarded them as formally equivalent, a re-analysis of his own argument for their equivalence brings out two important assumptions on which it rests: weak additivity and actor's control. When weak additivity breaks down, neither fitness concept is appropriate in its original form. When actor's control breaks down, neighbour-modulated fitness may be appropriate, but inclusive fitness is not. Yet I argue that, despite its more (...) limited domain of application, inclusive fitness provides a distinctively valuable perspective on social evolution. (shrink)

While the existence of human cognitive and behavioral diversity is now widely recognized, it is not yet well established how to explain this diversity. In particular, it is still unclear how to determine whether any given instance of human cognitive and behavioral diversity is due to a common psychology that is merely “triggered” differently in different bio-cultural environments, or whether it is due to deeply and fundamentally different psychologies. This paper suggests that, to answer this question, we need to employ (...) subtle theoretical considerations of theory choice—especially the consideration of the complexity-weighted differential predictive successes of the two accounts. To make this clearer, the paper develops a novel analysis of the observed differences in human sharing dispositions. (shrink)

The puzzle of how altruism can evolve has been at the center of recent debates over Hamilton’s Rule, inclusive fitness, and kin-selection. In this paper, I use recent debates over altruism and Hamilton’s legacy as an example to illustrate a more general problem in evolutionary theory that has philosophical significance; I attempt to explain this significance and to draw a variety of conclusions about it. The problem is that specific behaviours and general concepts of organism agency and intentionality are defined (...) in terms of concepts of evolutionary “costs” and “benefits,” and these terms have determined the role that agency should play in evolutionary explanation. However, costs, benefits, and agency are not only or even best conceived through evolutionary effects in a biological context. The paper proceeds as follows: first, I explain how the issue of agency relates to the evolutionary puzzle of altruism. Next, I discuss how questions about agency have figured in recent debates over Hamilton’s legacy. In the final section, I argue that Denis Walsh’s “situated Darwinism,” which attempts to return the organism to central status in biological explanation, offers a more productive route for thinking about different forms of costs, benefits, and agency. Finally, I argue that the upshot of all this is that there may be many different, and equally valid, ways to express what organisms are doing and how they are behaving based on different currencies of cost and benefit—even if these may stand in some tension. I illustrate this through returning to the case of altruism and using examples to show that even in non-humans there can be many forms of altruism, even if they are not all biological altruism as defined in the conventional evolutionary terms. (shrink)

This paper examines the points of disagreement between Neo-Darwinian and recent Neo-Aristotelian discussions of the status of purposive language in biology. I discuss recent Neo-Darwinian “evolutionary” treatments and distinguish three ways to deal with the philosophical status of teleological language of purpose: teleological error theory, methodological teleology, and Darwinian teleological realism. I then show how “non-evolutionary” Neo-Aristotelian approaches in the work of Michael Thompson and Philippa Foot differ from these by offering a view of purposiveness grounded in life-cycle patterns, rather (...) than in long-term evolutionary processes or natural selection. Finally, I argue that the crucial difference between Neo-Darwinian and Neo-Aristotelian approaches regards the question of whether or not reproduction deserves the status of an “ultimate” aim of organisms. I offer reasons to reject the concept of an “ultimate” aim in evolutionary biology and to reject the notion that reproduction serves a purpose. I argue that evolutionary biology is not in the position to determine what the “ultimate” explanation of natural purpose is. (shrink)

The concept of creativity is a central one in folk psychological explanation and has long been prominent in philosophical debates about the nature of art, genius, and the imagination. The scientific investigation of creativity in humans is also well established, and there has been increasing interest in the question of whether the concept can be rigorously applied to non-human animals. In this paper, I argue that such applications face serious challenges of both a conceptual and methodological character, reflecting deep controversies (...) within both philosophy and psychology concerning how to define and apply the concept of creativity. After providing a brief review of some of the leading theories of creativity and discussing some of the strongest putative cases of creative intelligence in non-human animals, I examine some of the more worrisome difficulties faced by attempts to use these theories to answer the question of whether animals are truly creative. I conclude by examining how we might overcome them, and suggest that one approach worth taking seriously is to adopt what I term a Strong Rejectionist view of creativity, eschewing use of the term entirely in the scientific study of comparative cognition. (shrink)

I distinguish two versions of kin selection theory—a purely genetic version and a version that also appeals to cultural forms of cooperation —and present an argument in favor of using the former when it comes to accounting for the evolution of cooperation in non-human organisms. Specifically, I first show that both GKST and WKST are equally mathematically coherent—they can both be derived from the Price equation—but not necessarily equally empirically plausible, as they are based on different assumptions about the inheritance (...) system underlying the cooperative phenotype. Given this, I then, second, present a model selection theoretic argument in favor of GKST over WKST. This argument is based on the fact that, in non-human cases, the former theory is likely to be as empirically successful as WKST, while containing fewer degrees of freedom. I end by defending both the intrinsic importance of this argument and its relevance to the discussion surrounding the “gene’s eye view of evolution.”. (shrink)

Hamilton’s theory of inclusive fitness is a widely used framework for studying the evolution of social behavior, but controversy surrounds its status. Hamilton originally derived his famous rb > c rule for the spread of a social gene by assuming additivity of costs and benefits. However, it has recently been argued that the additivity assumption can be dispensed with, so long as the −c and b terms are suitably defined, as partial regression coefficients. I argue that this way of generalizing (...) Hamilton’s rule to the nonadditive case, while formally correct, faces conceptual problems. (shrink)

Social evolution theory provides a wide array of successful evolutionary explanations for cooperative traits. However and surprisingly, a number of cases of unexplained cooperative behaviour remain. Shouldn’t they cast doubt on the relevance of the theory, or even disconfirm it? This depends on whether the theory is akin to a research programme such as adaptationism, or closer to a theory – a set of compatible, confirmable hypotheses. In order to find out, we focus on the two main tenets of social (...) evolution theory, namely reciprocity explanations and kin selection. Reciprocity-based explanations are extremely hard to confirm. This is due, first to the multiple realisability of explanatory processes, factors and strategies, despite apparent reasons to the contrary; second, to the high quantity, and limited availability of data needed to eliminate or back up such explanations. One of our target cases vividly illustrates these limitations. Moreover, kin selection, while relatively easy to disconfirm in particular cases, seems to enjoy a more limited explanatory scope than previously thought. Overall, social evolution theory turns out to be neither a research programme nor a theory, but a heterogeneous scientific entity, composed of parts that are amenable to confirmation and others barely so. (shrink)

The theory of evolution by natural selection is, perhaps, the crowning intellectual achievement of the biological sciences. There is, however, considerable debate about which entity or entities are selected and what it is that fits them for that role. This article aims to clarify what is at issue in these debates by identifying four distinct, though often confused, concerns and then identifying how the debates on what constitute the units of selection depend to a significant degree on which of these (...) four questions a thinker regards as central. (shrink)

I consider two attempts to combine realism with pluralism about the units of selection: Sober and Wilson’s combination of “model” and “unit” pluralism, and Sterelny and Griffiths’ “local pluralism”. I argue that both of these attempts fail to show that realism and pluralism are compatible. Sober and Wilson’s pluralism turns out, on closer inspection, to be a kind of monism in disguise, while Sterelny and Griffiths’ local pluralism involves a combination of realism and anti-realism about interactors, and the units of (...) selection, that is fundamentally unstable. My conclusion is that one must choose whether to be a realist or a pluralist in this area: one cannot be both. The question of which we should choose is a further question that I do not take a stand on. (shrink)

The role of genetic relatedness in social evolution has recently come under critical attention. These arguments are here critically analyzed, both theoretically and empirically. It is argued that when the conceptual structure of the theory of natural selection is carefully taken into account, genetic relatedness can be seen to play an indispensable role in the evolution of both facultative and advanced eusociality. Although reviewing the empirical evidence concerning the evolution of eusociality reveals that relatedness does not play a role in (...) the initial appearance of helper phenotypes, this follows simply from the fact that natural selection – of which relatedness is a necessary component – does not play a causal role in the origin of any traits. Further, separating two logically distinct elements of causal explanation – necessity and sufficiency – explains why the debate lingers on: although relatedness plays a necessary role in the evolution of helping and advanced eusociality, relatedness alone is not sufficient for their appearance. Therefore, if the relatedness variable in a given data set is held at a uniformly high value, then it indeed may turn out that other factors occupy a more prominent role. However, this does not change the fact that high relatedness functions as a necessary background condition for the evolution of advanced eusociality. (shrink)