Jonathan Birch

Department of Philosophy, Logic and Scientific Method
London School of Economics and Political Science
Houghton Street, London, WC2A 2AE



I'm an Associate Professor of Philosophy and Principal Investigator (PI) on the Foundations of Animal Sentience project. I mainly work on (i) animal sentience, cognition and welfare and (ii) the evolution of altruism and social behaviour.

3rd-Person Bio / CV / ”Meet the Faculty” Video

Dr Jonathan Birch is an Associate Professor of Philosophy at the LSE and Principal Investigator (PI) on the Foundations of Animal Sentience project. In addition to his interest in animal sentience, cognition and welfare, he also has a longstanding interest in the evolution of altruism and social behaviour.

He joined the LSE in 2014. Before moving to London, he was a Junior Research Fellow at Christ’s College, Cambridge. He completed his PhD at the University of Cambridge in 2013, with a dissertation entitled Kin Selection: A Philosophical Analysis.

He has published widely on various topics in the philosophy of the biological sciences, in journals such as Current Biology, The American Naturalist, Biological Reviews, Philosophy of Science, and The British Journal for the Philosophy of Science. His first book, The Philosophy of Social Evolution, was published by Oxford University Press in 2017.

In 2014, he was one of four UK philosophers honoured with a Philip Leverhulme Prize, which recognize “the achievement of outstanding researchers whose work has already attracted international recognition and whose future career is exceptionally promising”.


The Philosophy of Social Evolution, Oxford: 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.

"This is philosophy at its best: a clear, informed, and ambitious synthesis of the conceptual, the formal, and the empirical. ... It is lucid, well written, sensible, and intellectually valuable, and the most critical parts are accessible enough for anyone willing to do a bit of homework now and then. ... It is an essential text for philosophy of biology and, more generally, anyone working on the evolution of cooperation." - Carl Brusse and Kim Sterelny, The British Journal for the Philosophy of Science

"Jonathan Birch's recent book The Philosophy of Social Evolution is a superb exploration of philosophical implications of Hamilton's work. ... Birch shows why philosophy will continue to be an integral part of the future of the study of social evolution." - J. Arvid Agren, Studies in History and Philosophy of Biological and Biomedical Sciences

"Birch is extremely clear and careful in the development of his ideas and this makes the breadth and depth of this work even more impressive. The Philosophy of Social Evolution will, I suspect, become required reading in several different fields and, as I have suggested, there are insights here that extend beyond the philosophy of biology and evolutionary theory." - John Thrasher, Metascience

Recent non-academic writing

2019. Naturally Social. An LSE Research Highlight on my work on the evolution of cooperation.

2019. A Conversation [with David Sloan Wilson] About the Equivalence of Theories of Social Evolution.

2019. How Do We Know How to Act Together?, LSE Philosophy Blog.

2018. How to Fix Oxbridge's Biased Admissions System. Letter to The Guardian.

2018. The Elusive Calculus of Insects' Altruism and Kin Selection. A Quanta Magazine article in which I am quoted.

2018. Shell-Shocked. An LSE Research Highlight on my work on animal sentience.

Other activities

1. I’m a trustee of the British Society for the Philosophy of Science. To see what we do, visit us at

2. I’m a Fellow of the Forum for European Philosophy. The Forum is an educational charity which runs public philosophy events in London, and I organize some of these events. To see what’s coming up soon, and to access our huge archive of free audio recordings of past events, visit the Forum’s website.

Key journal articles and book chapters

Find my articles on PhilPapers and Google Scholar. I only update this website every now and then. Trouble accessing these? Email me.

2020. Kin Selection, Group Selection, and the Varieties of Population Structure. British Journal for the Philosophy of Science 71:259-286. 1-Line Summary

A long companion piece to "Are Kin and Group Selection Rivals or Friends?". There is no sharp distinction between kin selection and group selection. Instead, we should think of them as large, overlapping regions of 'K-G space'.

2019. Joint Know-How. Philosophical Studies 176:3329-3352. 1-Line Summary

What is it for two people to know how to do something together? The key idea here is 'actual mutual enablement'.

2019. Are Kin and Group Selection Rivals or Friends? Current Biology 29:R433-R438. 1-Line Summary

There is no sharp distinction between kin selection and group selection. Instead, we should think of them as large, overlapping regions of 'K-G space'.

2019. Inclusive Fitness as a Criterion for Improvement. Studies in History and Philosophy of Biological and Biomedical Sciences 76:101186. 1-Line Summary

What role does the concept of inclusive fitness play in evolutionary theory? It provides a criterion for improvement and a standard for optimality.

2019. Altruistic Deception. Studies in History and Philosophy of Biological and Biomedical Sciences 74:27-33. 1-Line Summary

"Deception" is the strategic exploitation of a receiver by a sender. Sometimes, deception can be altruistic. Pied babblers may provide an example.

2018. Animal Cognition and Human Values. Philosophy of Science 85:1026-1037. 1-Line Summary

When advising on animal welfare policy, a scientific expert should weigh the risks of over-attributing and under-attributing mental states to animals using the principle of expected welfare maximization.

2018. Fitness maximization. In R. Joyce, ed., The Routledge Handbook of Evolution and Philosophy. London: Routledge, pp. 49-63. 1-Line Summary

A critique of fitness maximization, less technical than my 'Natural selection and the maximization of fitness' and more focussed on conceptual issues.

2017. Animal Sentience and the Precautionary Principle. Animal Sentience 2:16(1) (published with 20 commentaries and my responses, Part I and Part II). 1-Line Summary

When there is suggestive but inconclusive evidence of sentience in a particular order of organisms (e.g. decapod crustaceans), give them the benefit of the doubt in a policy-making context.

2017. The Inclusive Fitness Controversy: Finding a Way Forward. Royal Society Open Science 4:170355. 1-Line Summary

My latest attempt to reconcile critics and defenders of inclusive fitness by synthesizing their insights (for a special issue on "Debating Inclusive Fitness").

2016. Hamilton's Two Conceptions of Social Fitness. Philosophy of Science 83:848-860. 1-Line Summary

Inclusive fitness has distinctive advantages over neighbour-modulated fitness for understanding social adaptation.

2016. Natural Selection and the Maximization of Fitness. Biological Reviews 91:712-727. 1-Line Summary

I distinguish four varieties of the idea that fitness is “maximized” by natural selection and argue that none is theoretically supported.

2015 (with Samir Okasha). Kin Selection and Its Critics. BioScience 65:22-32 (Open Access). 1-Line Summary

An overview of recent controversies surrounding the idea of “kin selection” and its relationship to “multi-level selection”.

2014. Gene Mobility and the Concept of Relatedness. Biology and Philosophy 29:445-476. 1-Line Summary

Recent developments in microbiology should cause us to rethink the concept of "genetic relatedness" in microbes.

2014. Hamilton’s Rule and Its Discontents. British Journal for the Philosophy of Science 65:381-411 (Open Access). 1-Line Summary

An analysis of the philosophical assumptions underlying a recent dispute about the status of kin selection.

2014. Propositional Content in Signalling Systems. Philosophical Studies 171:493-512. 1-Line Summary

I argue for an amendment to Skyrms’s account of propositional content in signalling systems, in order to make room for the possibility of false propositional content.

2014 (with James A. R. Marshall). Queller’s Separation Condition Explained and Defended. American Naturalist 184:531-540. 1-Line Summary

We argue for the “formal equivalence” of phenotypic formulations of inclusive fitness theory and multi-level selection theory, exposing some errors in a recent challenge to this equivalence.

2013. On the ‘Simulation Argument’ and Selective Scepticism. Erkenntnis 78:95-107. 1-Line Summary

Bostrom’s notorious “simulation argument” does not constrain one’s rational degree of belief in the proposition that one is simulated, unless one already accepts an implausible form of selective scepticism.

2012. Collective Action in the Fraternal Transitions. Biology and Philosophy 27:363-380. 1-Line Summary

Considerations concerning the task-structure of cooperation can shed light on the evolution of both multicellular organisms and eusocial insect colonies.

2012. Robust Processes and Teleological Language. European Journal for Philosophy of Science 3:299-312. 1-Line Summary

Teleological language occurs in physical as well as biological sciences, and it provides a convenient way of describing the robust convergence of population-level processes on particular end-points.

2012. The Negative View of Natural Selection. Studies in History and Philosophy of Biological and Biomedical Sciences 43:569-573. 1-Line Summary

Given Michael Strevens’s account of explanatory relevance, natural selection is relevant to explaining the traits of individual organisms, at least in some contexts.

2009. Irretrievably Confused? Innateness in Explanatory Context. Studies in History and Philosophy of Biological and Biomedical Sciences 40:296-301. 1-Line Summary

I argue for a contrastive account of innateness, on which innateness ascriptions imply that environmental mechanisms did not make the difference between the actual developmental outcome and a class of relevant alternatives.


Thanks to Bryan Roberts, who designed the template for this website.