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Department of History and Philosophy of Science


Tim Lewens

The literature on the philosophy of biology is enormous; what follows only scratches the surface. Here I have restricted myself mainly to evolutionary biology, although there are short sections on evolutionary epistemology and evolutionary ethics. I have tried to include a handful of the best, the most interesting or the most often slated books and articles under each topic heading. These selections are intended as starting points for the interested student. Those who want to go further will have to follow their noses. Sterelny and Griffiths (1999) provide useful assistance to nose followers in the form of very up-to-date reading lists at the end of each of their chapters. Their book would also be a good starting point for researchers interested in topics that are not covered here – for example, issues in the philosophy of molecular biology and the philosophy of ecology, or traditional questions for philosophy of biology such as how to define life and how to define fitness.


Online resources

The two leading online encyclopaedias of philosophy have good resources for philosophy of biology, and both are expanding the range of material they have available. They are:


  • Sober, E. (1999) The Philosophy of Biology. Second Edition. Boulder: Westview.
  • Sterelny, K. and P. Griffiths (1999) Sex and Death. Chicago: Chicago University Press. (An invigorating and comprehensive introduction. It also includes a helpful glossary of technical terms from biology.)
  • Lewens, T. (2007) Darwin. London: Routledge.

Those who want to check their biology might benefit from either of the following textbooks, written for biology students.

  • Ridley, M. (2003) Evolution (3rd Edition). Cambridge MA: Blackwell Scientific.
  • Futuyma, D. (2005) Evolutionary Biology (4th Edition). Sunderland, MA: Sinauer Associates.


  • Sober, E. ed (1984) Conceptual Issues in Evolutionary Biology. Cambridge MA: MIT Press.
  • Sober, E. ed (1994) Conceptual Issues in Evolutionary Biology (2nd Edition). Cambridge MA: MIT Press.
  • Sober, E. ed (2006) Conceptual Issues in Evolutionary Biology (3rd Edition). Cambridge MA: MIT Press. (The three editions all contain slightly different material, and all are useful.)
  • Fox-Keller, E. and E. Lloyd eds (1992) Keywords in Evolutionary Biology. Cambridge, Mass: Harvard University Press.
  • Hodge, J. and G. Radick eds. (2003) The Cambridge Companion to Darwin. Cambridge: Cambridge University Press.
  • Ruse, M. and D. Hull, eds. (2007) The Cambridge Companion to the Philosophy of Biology. Cambridge: Cambridge University Press.
  • Matthen, M. and C. Stephens eds. (2007) The Handbook of Philosophy of Biology. Elsevier.
  • Hull, D. and M. Ruse, eds (1998) The Philosophy of Biology. Oxford: Oxford University Press.


Many generalist philosophy and philosophy of science journals (e.g. British Journal for the Philosophy of Science, Philosophy of Science, Journal of Philosophy) also regularly carry articles on the philosophy of biology. The Philosophers' Index will help with literature searches.

Researchers without biological training shouldn't be scared of reading articles in biology journals. Many debates take place across disciplines these days, and you will miss out if you only read philosophy journals. TREE (Trends in Ecology and Evolution) is a good example of a journal that is not too technical and full of interesting bits of real biology to intimidate other philosophers with. It and many other useful biology journals are held in the Central Science Library, only a stone's throw from the Whipple. The Whipple is also near to the Zoology Library, another useful research resource.

Adaptation and adaptationism

Gould and Lewontin (1979) is probably the best place to start. The debate can get confusing and frustrating, as squabbling about whether adaptationism is right or wrong often kicks off before the multiple meanings of 'adaptationism' have been clarified. Sober's (1998) article is an exception, and provides a useful landmark in the post-Spandrels debate. Ron Amundson's contribution to the collection edited by Rose and Lauder is also extremely helpful. Many of the items in the following section on developmental biology also contain challenges to adaptationism.

  • Dawkins, R. (1986) The Blind Watchmaker. New York: Norton.
  • Dennett, D. (1995) Darwin's Dangerous Idea. New York: Simon and Schuster.
  • Gould, S.J. and R. Lewontin (1979) 'The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme' Proceedings of the Royal Society of London B 205: 581-98. (Reprinted in Sober, E. ed [1984])
  • Griffiths, P. (1996) 'The Historical Turn in the Study of Adaptation' British Journal for the Philosophy of Science 47: 511-532.
  • Lewens, T. (2007) 'Adaptation' in The Cambridge Companion to the Philosophy of Biology, edited by M. Ruse and D. Hull. Cambridge: Cambridge University Press.
  • Lewontin, R (1985) 'Adaptation' in The Dialectical Biologist, edited by R. Levins and R.C. Lewontin. Cambridge, MA: Harvard University Press.
  • Orzack, S. and E. Sober eds. (2001) Adaptationism and Optimality. Cambridge: Cambridge University Press. (The paper by Godfrey-Smith is particularly useful.)
  • Rose, M. and G. Lauder eds (1996) Adaptation. San Diego: Academic Press. (See especially articles by Ronald Amundson and George Lauder.)
  • Sober, E (1998) 'Six Sayings about Adaptationism' in D. Hull and M. Ruse (eds) The Philosophy of Biology. Oxford: Oxford University Press.

Developmental biology and developmental systems theory

Developmental biology is a topic attracting more and more interest from philosophers who are bored with thinking about species and the units of selection. Many of these items overlap with the related topics of innateness and the question of whether there is any sense in which genes can be said to carry information.

  • Amundson, R. (1994) 'Two Concepts of Constraint: Adaptationism and the Challenge from Developmental Biology' Philosophy of Science 61: 556-578. (Reprinted in Hull and Ruse, eds, [1998])
  • Amundson, R. (2005) The Changing Role of the Embryo in Evolutionary Thought. Cambridge: Cambridge University Press.
  • Bateson, P (1983) 'Genes, Environment and the Development of Behaviour' In Animal Behaviour: Genes, Development and Learning, P. Slater and T. Halliday (eds) Oxford: Blackwell.
  • Gray, R. (1992) 'Death of the Gene: Developmental Systems Strike Back' in Griffiths, P. (ed) Trees of Life: Essays in the Philosophy of Biology. Dordrecht: Kluwer.
  • Griffiths, P. and R. Gray (1994) 'Developmental Systems and Evolutionary Explanation' Journal of Philosophy 91: 277-304. (Reprinted in Hull and Ruse, eds, [1998])
  • Oyama, S. (2000) The Ontogeny of Information, 2nd Edition. Duke University Press.
  • Oyama, S. (2000) Evolution's Eye. Duke University Press.
  • Oyama, S., P. Griffiths and R. Gray eds. (2001) Cycles of Contingency. Cambridge, MA: MIT Press.

Information in molecular biology

  • Maynard Smith, J. (2000) 'The Concept of Information in Biology' Philosophy of Science 67: 177-194. (Be sure to read all the commentaries to this paper.)
  • Griffiths, P. (2001) 'Genetic Information: A Metaphor in Search of a Theory' Philosophy of Science 68: 394-412.
  • Godfrey-Smith, P. (2001) 'On the Theoretical Role of "Genetic Coding"' Philosophy of Science 67: 26-44.


  • Ariew, A. (1999) 'Innateness is Canalization', in V. Hardcastle (ed.) Where Biology Meets Psychology, Cambridge, MA: MIT Press.
  • Griffiths, P. (2002) 'What is Innateness?', Monist, 85: 70-85.
  • Mameli, M. and Bateson, P. (2006) 'Innateness and the Sciences', Biology and Philosophy, 21: 155-188.

The units of selection

A difficult topic with a huge literature. The questions raised under the units of selection debate are confusing because even the seasoned combatants often cannot decide what the debate is about. John Maynard Smith thinks that Elliott Sober doesn't understand what the units of selection problem is. If Sober doesn't understand it, then there is little hope for the rest of us (but it still makes an exciting topic for research). Items in the section on developmental biology are relevant here, too; developmental systems theorists believe they have powerful resources to undermine genic selectionism.

  • Dawkins, R. (1976) The Selfish Gene. Oxford: Oxford University Press.
  • Dawkins, R. (1982) The Extended Phenotype. Oxford: Oxford University Press. (Dawkins' best book. Although philosophers sometimes don't like to admit it, Dawkins is on fine analytical form here.)
  • Hull, D. (1981) 'Units of Evolution: A Metaphysical Essay' In The Philosophy of Evolution, R. Jensen and R. Harre (eds) Brighton: Harvester.
  • Lloyd, E. (1993) 'Unit of Selection' In Keywords in Evolutionary Biology, E. Fox-Keller and E. Lloyd (eds) Cambridge MA: Harvard.
  • Okasha, S. (2006) Evolution and the Levels of Selection. Oxford: Oxford University Press.
  • Sober, E. (1984) The Nature of Selection: Evolutionary Theory in Philosophical Focus. Cambridge, MA: MIT Press.
  • Sober, E. and R.C. Lewontin (1982) 'Artifact, Cause and Genic Selection' Philosophy of Science 49: 157-180. (Reprinted in Sober, ed, [1984])
  • Sober, E. and D.S. Wilson (1998) Unto Others: The Evolution of Altruism. Cambridge, MA: Harvard University Press.
  • Sterelny, K. and P. Kitcher (1988) 'The Return of The Gene' Journal of Philosophy 85: 339-60. (Reprinted in Hull and Ruse, eds, [1998])
  • Williams, G.C. (1966) Adaptation and Natural Selection. Princeton, NJ: Princeton University Press.

Functions and teleology

Almost every article the teleologist could need is collected in one of these three collections:

  • Allen, C, M. Bekoff and G. Lauder eds (1998) Nature's Purposes: Analyses of Function and Design in Biology. Cambridge, MA: MIT Press.
  • Buller, D. ed (1999) Function, Selection and Design. New York: SUNY Press.
  • Ariew, A, Cummins, R., and Perlman, M. (2002) Functions. Oxford: Oxford University Press.

The first two collections contain many seminal papers in the field. Be sure to look at those of Wright, Cummins, Millikan, Neander, Amundson and Lauder, Godfrey-Smith and Buller.

Interesting and/or important contributions that slipped through these editors' nets include:

  • Matthen, M (1997) 'Teleology and the Product Analogy' Australasian Journal of Philosophy 75: 21-37.
  • Walsh, D.M. (1996) 'Fitness and Function' British Journal for the Philosophy of Science 47: 553-574.

There is also a recent overview of the functions literature by me:

  • Lewens, T. (2007) 'Functions' in M. Matthen and C. Stephens, eds. The Handbook of Philosophy of Biology. Elsevier.


Another topic with an intimidating literature. Dupré's (1992) piece gives a short, friendly introduction. Readers could then move on to Ereschefsky's collection, which contains many of the most important pieces. Wilson's recent collection contains some important newer contributions.

  • Dupré, J. (1981) 'Natural Kinds and Biological Taxa' Philosophical Review 90: 66-90.
  • Dupré, J. (1992) 'Species: Theoretical Contexts' in Keywords in Evolutionary Biology, E. Keller and E. Lloyd (eds) Cambridge MA: Harvard.
  • Dupré, J. (1993) The Disorder of Things. Harvard, MA: Harvard University Press.
  • Ereschefsky, M. ed (1992) The Units of Evolution: Essays on the Nature of Species. Cambridge, MA: MIT Press.
  • Ghiselin, M. (1974) 'A Radical Solution to the Species Problem' Systematic Zoology 23: 536-44.
  • Hull, D. (1978) 'A Matter of Individuality' Philosophy of Science 45: 335-60.
  • Hull, D. (1997) 'The Ideal Species Concept – And Why We Can't Get It.' In Species: The Units of Biodiversity, edited by M.F. Claridge, H.A. Dawah, and M.R. Wilson. New York: Chapman and Hall.
  • Mayr, E. (1982) Systematics and the Origin of Species. Reprint of 1942 edition. New York: Columbia University Press.
  • Mayr, E. (1988) Towards a New Philosophy of Biology. Cambridge, MA: Harvard University Press. (See part seven for discussion of the species problem.)
  • Stamos, D. (2003) The Species Problem. Lexington Books.
  • Wilson, R. ed. (1999) Species: New Interdisciplinary Essays, Cambridge, MA: MIT Press.


The idea that evolution makes progress has tended to get more attention from historians than philosophers; however, the following are more philosophically oriented. This said, the fight in print between Dawkins (1997) and Gould (1997) is of more interest as an entertaining spectacle than as an illuminating discussion. Sterelny offers some useful criticism. Nitecki's (1988) collection is probably the best place to start.

  • Dawkins,R. (1997) 'Human Chauvinism: A Review of S.J. Gould's Full House' Evolution 51(3): 1015-20.
  • Gould, S.J. (1996) Full House: The Spread of Excellence from Plato to Darwin. New York: Harmony Press. (Published in the UK as Life's Grandeur)
  • Gould, S.J. (1997) 'Self-help for a Hedgehog Stuck on a Molehill' Evolution 51(3): 1010-24. (The title is self-explanatory.)
  • McShea, D. (1991) 'Complexity and Evolution: What Everybody Knows' Biology and Philosophy 6: 303-234.
  • Nitecki, M. ed (1988) Evolutionary Progress. Chicago: University of Chicago Press.
  • Ruse, M. (1996) Monad to Man: The Concept of Progress in Evolutionary Biology. Cambridge, MA: Harvard University Press.
  • Sterelny, K. (2001b) Dawkins vs. Gould: Survival of the Fittest, Cambridge: Icon.

Evolutionary epistemology, cultural evolution and memetics

Here follows a mish-mash of different evolutionary approaches to conceptual change, technology change, cultural change and more. Amundson's (1989) paper does a good job of distinguishing the question of whether selection processes are somehow involved in creating a set of phenomena, from what the explanatory import of the selection process may be. Dawkins (1976) presents a first outline of what is now know as memetics, and his views are defended and extended by Dennett (1995) towards the end of his book. Hull (1988) offers a useful examination of the idea that cultural evolution is Lamarckian. Some of the most serious and fecund work in this area has been done by Boyd and Richerson, and their (2005) book is an excellent place to start. My own SEP article attempts an overview.

  • Amundson, R. (1989) 'The Trials and Tribulations of Selectionist Explanations' in K. Hahlweg and C.A. Hooker eds Issues in Evolutionary Epistemology. New York: SUNY Press.
  • Basalla, G. (1988) The Evolution of Technology. Cambridge: Cambridge University Press.
  • Boyd, R. and P. Richerson (1985) Culture and the Evolutionary Process. Chicago: University of Chicago Press.
  • Campbell, D.T. (1974) 'Evolutionary Epistemology'. In P. Schlipp (ed) The Philosophy of Karl Popper, La Salle, IL: Open Court.
  • Cavalli-Sforza, L. and M. Feldman (1981) Cultural Transmission and Evolution: A Quantitative Approach. Princeton, NJ: Princeton University Press.
  • Dawkins, R (1976) The Selfish Gene. Oxford: Oxford University Press.
  • Dennett, D.C. (1995) Darwin's Dangerous Idea. New York: Norton.
  • Hull, D (1988) Science as a Process. Chicago: University of Chicago Press.
  • Lewens, T. (2007) 'Cultural Evolution' Stanford Encylcopedia of Philosophy.
  • Richerson, P. and R. Boyd (2005) Not by Genes Alone: How Culture Transformed Human Evolution. Chicago: Chicago University Press.
  • Sober, E. (1992) 'Models of Cultural Evolution' in Griffiths, P (ed) Trees of Life: Essays in the Philosophy of Biology. Dordrecht: Kluwer.

Evolution and ethics

The items listed below link evolutionary studies to ethics in a variety of ways. Joyce's recent book is a stimulating read, and may be a good place to start. Sober and Wilson's book is fast becoming a modern classic.

  • Binmore, K. (2005) Natural Justice. Oxford: Oxford University Press.
  • Joyce, R. (2006) The Evolution of Morality. Cambridge, MA: MIT Press.
  • Kitcher, P. (1993) 'Four Ways of "Biologicizing" Ethics', in E. Sober (ed.) Conceptual Issues in Evolutionary Biology, Second Edition, Cambridge, MA: MIT Press.
  • Maienshein, J. and M. Ruse eds (1998) Evolution and Ethics. Cambridge: Cambridge University Press.
  • Ruse, M. and E.O. Wilson (1986) 'Moral Philosophy as Applied Science: A Darwinian Approach to the Foundations of Ethics' Philosophy 61: 173-92.
  • Skyrms, B. (1996) Evolution of the Social Contract. Cambridge: Cambridge University Press.
  • Sober, E. (1988) 'What is Evolutionary Altruism?' Canadian Journal of Philosophy suppl. vol. 14: 75-99. Reprinted in Hull and Ruse (eds).
  • Sober, E. (1994a) 'Prospects for an Evolutionary Ethics', in E. Sober, From a Biological Point of View, Cambridge: Cambridge University Press.
  • Sober, E. (1994b) 'Did Evolution make us Psychological Egoists?', in E. Sober, From a Biological Point of View, Cambridge: Cambridge University Press.
  • Sober, E. and D.S. Wilson (1998) Unto Others: The Evolution of Altruism. Cambridge, MA: Harvard University Press.
  • Thompson, P. ed (1995) Issues in Evolutionary Ethics. Albany, NY: SUNY Press.