Cambridge University
Department of History and Philosophy of Science
PhD, 2013
London, London, City of, United Kingdom of Great Britain and Northern Ireland
PhilPapers Editorships
Evolutionary Biology
  •  130
    Inclusive Fitness as a Criterion for Improvement
    Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences 76 101186. 2019.
    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 hol…Read more
  •  329
    Robust processes and teleological language
    European Journal for Philosophy of Science 2 (3): 299-312. 2012.
    I consider some hitherto unexplored examples of teleological language in the sciences. In explicating these examples, I aim to show (a) that such language is not the sole preserve of the biological sciences, and (b) that not all such talk is reducible to the ascription of functions. In chemistry and biochemistry, scientists explaining molecular rearrangements and protein folding talk informally of molecules rearranging “in order to” maximize stability. Evolutionary biologists, meanwhile, often s…Read more
  •  224
    Agents and Goals in Evolution, by Samir Okasha (review)
    Mind 128 (512): 1408-1416. 2019.
    In this essay review of Samir Okasha's Agents and Goals in Evolution, I reflect on the rationale for agential thinking in biology, and consider whether the rationale is the same for genes as for organisms. I also discuss Okasha's ingenious examples of the evolution of irrational behaviour, and in particular the evolution of violations of the "independence axiom" of rational choice theory. These examples rely on a crucial distinction between aggregate and idiosyncratic risk.
  •  276
    Altruistic Deception
    Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences 74 27-33. 2019.
    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 “exploitat…Read more
  •  255
    Kin Selection: A Philosophical Analysis
    Dissertation, University of Cambridge. 2013.
    This PhD dissertation examines the conceptual and theoretical foundations of the most general and most widely used framework for understanding social evolution, W. D. Hamilton's theory of kin selection. While the core idea is intuitive enough (when organisms share genes, they sometimes have an evolutionary incentive to help one another), its apparent simplicity masks a host of conceptual subtleties, and the theory has proved a perennial source of controversy in evolutionary biology. To move towa…Read more
  •  142
    Irretrievably confused? Innateness in explanatory context
    Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences 40 (4): 296-301. 2009.
    The hunt for a biologically respectable definition for the folk concept of innateness is still on. I defend Ariew’s Canalization account of innateness against the criticisms of Griffiths and Machery, but highlight the remaining flaws in this proposal. I develop a new analysis based on the notion of environmental induction. A trait is innate, I argue, iff it is not environmentally induced. I augment this definition with a novel analysis of environmental induction that draws on the contrastive nat…Read more
  •  264
    Joint know-how
    Philosophical Studies 176 (12). 2019.
    When two agents engage in a joint action, such as rowing together, they exercise joint know-how. But what is the relationship between the joint know-how of the two agents and the know-how each agent possesses individually? I construct an “active mutual enablement” account of this relationship, according to which joint know-how arises when each agent knows how to predict, monitor, and make failure-averting adjustments in response to the behaviour of the other agent, while actively enabling the ot…Read more
  •  54
    We should distinguish between the apparent source of a norm and the scope of the norm's satisfaction conditions. Wide-scope social norms need not be externalised, and externalised social norms need not be wide in scope. Attending to this distinction leads to a problem for Stanford: The adaptive advantages he attributes to externalised norms are actually advantages of wide-scope norms.
  •  268
    The Philosophy of Social Evolution
    Oxford University Press. 2017.
    From mitochondria to meerkats, the natural world is full of spectacular examples of social behaviour. In the early 1960s W. D. Hamilton changed the way we think about how such behaviour evolves. He introduced three key innovations - now known as Hamilton's rule, kin selection, and inclusive fitness - and his pioneering work kick-started a research program now known as social evolution theory. This is a book about the philosophical foundations and future prospects of that program. [Note: only the…Read more
  •  422
    Animal Cognition and Human Values
    Philosophy of Science 85 (5): 1026-1037. 2018.
    Animal welfare scientists face an acute version of the problem of inductive risk, since they must choose whether to affirm attributions of mental states to animals in advisory contexts, knowing their decisions hold consequences for animal welfare. In such contexts, the burden of proof should be sensitive to the consequences of error, but a framework for setting appropriate burdens of proof is lacking. Through reflection on two cases—pain and cognitive enrichment—I arrive at a tentative framework…Read more
  •  227
    Fitness Maximization
    In Richard Joyce (ed.), The Routledge Handbook of Evolution and Philosophy, Routledge. pp. 49-63. 2018.
    Is there any way to reconcile the adaptationist’s image of natural selection as an engine of optimality with the more complex image of its dynamics we get from population genetics? This has long been an important strand in the controversy surrounding adaptationism, yet debate has been hampered by a tendency to conflate various different ways of thinking about maximization. Here I distinguish four varieties of maximization principle. I then discuss the logical relations between these varieties, a…Read more
  •  457
    In debates about animal sentience, the precautionary principle is often invoked. The idea is that when the evidence of sentience is inconclusive, we should “give the animal the benefit of the doubt” or “err on the side of caution” in formulating animal protection legislation. Yet there remains confusion as to whether it is appropriate to apply the precautionary principle in this context, and, if so, what “applying the precautionary principle” means in practice regarding the burden of proof for a…Read more
  •  168
    The Inclusive Fitness Controversy: Finding a Way Forward
    Royal Society Open Science 4 (170335): 170335. 2017.
    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 u…Read more
  •  312
    Kin Selection, Group Selection, and the Varieties of Population Structure
    British Journal for the Philosophy of Science 71 (1): 259-286. 2020.
    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 provi…Read more
  •  374
    On the 'Simulation Argument' and Selective Scepticism
    Erkenntnis 78 (1): 95-107. 2013.
    Nick Bostrom’s ‘Simulation Argument’ purports to show that, unless we are confident that advanced ‘posthuman’ civilizations are either extremely rare or extremely rarely interested in running simulations of their own ancestors, we should assign significant credence to the hypothesis that we are simulated. I argue that Bostrom does not succeed in grounding this constraint on credence. I first show that the Simulation Argument requires a curious form of selective scepticism, for it presupposes tha…Read more
  •  576
    The Negative View of Natural Selection
    Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences 43 (2): 569-573. 2012.
    An influential argument due to Elliott Sober, subsequently strengthened by Denis Walsh and Joel Pust, moves from plausible premises to the bold conclusion that natural selection cannot explain the traits of individual organisms. If the argument were sound, the explanatory scope of selection would depend, surprisingly, on metaphysical considerations concerning origin essentialism. I show that the Sober-Walsh-Pust argument rests on a flawed counterfactual criterion for explanatory relevance. I fur…Read more
  •  243
    Social revolution (review)
    Biology and Philosophy 27 (4): 571-581. 2012.
    Andrew Bourke’s Principles of Social Evolution identifies three stages that characterize an evolutionary transition in individuality and deploys inclusive fitness theory to explain each stage. The third stage, social group transformation, has hitherto received relatively little attention from inclusive fitness theorists. In this review, I first discuss Bourke’s “virtual dominance” hypothesis for the evolution of the germ line. I then contrast Bourke’s inclusive fitness approach to the major tran…Read more
  •  300
    How Cooperation Became the Norm (review)
    Biology and Philosophy 29 (3): 433-444. 2014.
    Most of the contributions to Cooperation and Its Evolution grapple with the distinctive challenges presented by the project of explaining human sociality. Many of these puzzles have a ‘chicken and egg’ character: our virtually unparalleled capacity for large-scale cooperation is the product of psychological, behavioural, and demographic changes in our recent evolutionary history, and these changes are linked by complex patterns of reciprocal dependence. There is much we do not yet understand abo…Read more
  •  276
    Hamilton’s rule and its discontents
    British Journal for the Philosophy of Science 65 (2): 381-411. 2014.
    In an incendiary 2010 Nature article, M. A. Nowak, C. E. Tarnita, and E. O. Wilson present a savage critique of the best-known and most widely used framework for the study of social evolution, W. D. Hamilton’s theory of kin selection. More than a hundred biologists have since rallied to the theory’s defence, but Nowak et al. maintain that their arguments ‘stand unrefuted’. Here I consider the most contentious claim Nowak et al. defend: that Hamilton’s rule, the core explanatory principle of kin …Read more
  •  340
    Propositional Content in Signalling Systems
    Philosophical Studies 171 (3): 493-512. 2014.
    Skyrms, building on the work of Dretske, has recently developed a novel information-theoretic account of propositional content in simple signalling systems. Information-theoretic accounts of content traditionally struggle to accommodate the possibility of misrepresentation, and I show that Skyrms’s account is no exception. I proceed to argue, however, that a modified version of Skyrms’s account can overcome this problem. On my proposed account, the propositional content of a signal is determined…Read more
  •  129
    Michael Tomasello: A Natural History of Human Morality (review)
    BJPS Review of Books 2017. 2017.
    Working together creates mutual obligations. For example, the members of a football team owe it to each other to work hard for the good of the team. A player who doesn’t try hard enough, or who makes a costly mistake, lets the side down. When the team loses, feelings of responsibility, guilt, and shame ensue. Players ought to feel committed to the team and responsible for its failures. If they don’t, they deserve to be dropped. The idea at the heart of Michael Tomasello’s ‘natural history’ of hu…Read more
  •  143
    Has Grafen Formalized Darwin?
    Biology and Philosophy 29 (2): 175-180. 2014.
    One key aim of Grafen’s Formal Darwinism project is to formalize ‘modern biology’s understanding and updating of Darwin’s central argument’. In this commentary, I consider whether Grafen has succeeded in this aim
  •  193
    Kin Selection and Its Critics
    with Samir Okasha
    BioScience 65 (1): 22-32. 2015.
    Hamilton’s theory of kin selection is the best-known framework for understanding the evolution of social behavior but has long been a source of controversy in evolutionary biology. A recent critique of the theory by Nowak, Tarnita, and Wilson sparked a new round of debate, which shows no signs of abating. In this overview, we highlight a number of conceptual issues that lie at the heart of the current debate. We begin by emphasizing that there are various alternative formulations of Hamilton’s r…Read more
  •  112
    Queller’s separation condition explained and defended
    with James A. R. Marshall
    American Naturalist 184 (4): 531-540. 2014.
    The theories of inclusive fitness and multilevel selection provide alternative perspectives on social evolution. The question of whether these perspectives are of equal generality remains a divisive issue. In an analysis based on the Price equation, Queller argued (by means of a principle he called the separation condition) that the two approaches are subject to the same limitations, arising from their fundamentally quantitative-genetical character. Recently, van Veelen et al. have challenged Qu…Read more
  •  247
    Collective Action in the Fraternal Transitions
    Biology and Philosophy 27 (3): 363-380. 2012.
    Inclusive fitness theory was not originally designed to explain the major transitions in evolution, but there is a growing consensus that it has the resources to do so. My aim in this paper is to highlight, in a constructive spirit, the puzzles and challenges that remain. I first consider the distinctive aspects of the cooperative interactions we see within the most complex social groups in nature: multicellular organisms and eusocial insect colonies. I then focus on one aspect in particular: th…Read more
  •  162
    Hamilton’s Two Conceptions of Social Fitness
    Philosophy of Science 83 (5): 848-860. 2016.
    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, bu…Read more
  •  577
    Natural Selection and the Maximization of Fitness
    Biological Reviews 91 (3): 712-727. 2016.
    The notion that natural selection is a process of fitness maximization gets a bad press in population genetics, yet in other areas of biology the view that organisms behave as if attempting to maximize their fitness remains widespread. Here I critically appraise the prospects for reconciliation. I first distinguish four varieties of fitness maximization. I then examine two recent developments that may appear to vindicate at least one of these varieties. The first is the ‘new’ interpretation of F…Read more
  •  298
    Samir Okasha and Ken Binmore (eds), Evolution and rationality: Decisions, cooperation, and strategic behaviour (review)
    British Journal for the Philosophy of Science 64 (3): 669-673. 2013.
    Evolution and Rationality marks the end of a three-year project, ‘Evolution, Cooperation, and Rationality’, directed at the University of Bristol by the book’s editors, Samir Okasha and Ken Binmore. The collection draws together the editors’ pick of the papers delivered at the conferences the project hosted, and covers a wide range of topics at the intersection of evolutionary theory and the social sciences. It is a splendid anthology: timely, interdisciplinary, thematically cohesive, and full o…Read more
  •  338
    Gene Mobility and the Concept of Relatedness
    Biology and Philosophy 29 (4): 445-476. 2014.
    Cooperation is rife in the microbial world, yet our best current theories of the evolution of cooperation were developed with multicellular animals in mind. Hamilton’s theory of inclusive fitness is an important case in point: applying the theory in a microbial setting is far from straightforward, as social evolution in microbes has a number of distinctive features that the theory was never intended to capture. In this article, I focus on the conceptual challenges posed by the project of extendi…Read more
  •  118
    The sense of fairness is a central aspect of human moral psychology. Intuitions about fairness lead to many widespread moral beliefs, such as the belief that the punishment should fit the crime or the belief that one deserves a fair share of what one has earned. In The Origins of Fairness, Nicolas Baumard sets out to shed light on the evolutionary origin of these intuitions. He argues that the human sense of fairness is innate and universal, and he offers an account of its evolution that highlig…Read more