The Extended Phenotype

by Richard Dawkins

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  White, M. J. D. (1978). Modes of Speciation. San Francisco: W. H. Freeman.

  Whitham, T. G. & Slobodchikoff, C. N. (in press). Evolution of individuals, plant-herbivore interactions, and mosaics of genetic variability: the adaptive significance of somatic mutations in plants. Oecologia.

  Whitney, G. (1976). Genetic substrates for the initial evolution of human sociality. I. Sex chromosome mechanisms. American Naturalist 110, 867–875.

  Wickler, W. (1968). Mimicry. London: Weidenfeld & Nicolson.

  Wickler, W. (1976). Evolution-oriented ethology, kin selection, and altruistic parasites. Zeitschrift für Tierpsychologie 42, 206–214.

  Wickler, W. (1977). Sex-linked altruism. Zeitschrift für Tierpsychologie 43, 106–107.

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  Williams, G. C. (1966). Adaptation and Natural Selection. Princeton, N.J.: Princeton University Press.

  Williams, G. C. (1975). Sex and Evolution. Princeton, N.J.: Princeton University Press.

  Williams, G. C. (1979). The question of adaptive sex ratio in outcrossed vertebrates. Proceedings of the Royal Society of London, B 205, 567–580.

  Williams, G. C. (1980). Kin selection and the paradox of sexuality. In Sociobiology: Beyond Nature/Nurture? (eds G. W. Barlow & J. Silverberg), pp. 371–384. Boulder: Westview Press.

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  Wilson, E. O. (1971). The Insect Societies. Cambridge, Mass.: Harvard University Press.

  Wilson, E. O. (1975). Sociobiology: the New Synthesis. Cambridge, Mass.: Harvard University Press.

  Wilson, E. O. (1978). On Human Nature. Cambridge, Mass.: Harvard University Press.

  Winograd, T. (1972). Understanding Natural Language. Edinburgh: Edinburgh University Press.

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  Wright, S. (1980). Genic and organismic selection. Evolution 34, 825–843.

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  Zahavi, A. (1979). Parasitism and nest predation in parasitic cuckoos. American Naturalist 113, 157–159.

  Further Reading

  Allaby, M. (1982). Animal Artisans. London: Weidenfeld & Nicolson.

  Barkow, J. H., Cosmides, L. & Tooby, J. (1992). The Adapted Mind. New York: Oxford University Press.

  Basalla, G. (1988). The Evolution of Technology. Cambridge: Cambridge University Press.

  Blackmore, S. (1999). The Meme Machine. Oxford: Oxford University Press.

  Bonner, J. T. (1988). The Evolution of Complexity. Princeton: N.J.: Princeton University Press.

  Brandon, R. N. (1990). Adaptation and Environment. Princeton, N.J.: Princeton University Press.

  Brandon, R. N. & Burian, R. M. (1984). Genes, Organisms, Populations: Controversies over the Units of Selection. Cambridge, Mass.: MIT Press.

  Buss, L. W. (1987). The Evolution of Individuality. Princeton, N.J.: Princeton University Press.

  Clayton, D. & Harvey, P. (1993). Hanging nests on a phylogenetic tree. Current Biology 3, 882–883.

  Cronin, H. (1991). The Ant and the Peacock. Cambridge: Cambridge University Press.

  Cziko, G. (1995). Without Miracles. Cambridge, Mass.: MIT Press.

  Davies, N. B. (1992). Dunnock Behaviour and Social Evolution. Oxford: Oxford University Press.

  Davis, B. D. (1986). Storm over Biology. Buffalo: Prometheus Books.

  Dawkins, R. (1982). Universal Darwinism. In Evolution from Molecules to Men (ed. D. S. Bendall), pp. 403–425. Cambridge: Cambridge University Press.

  Dawkins, R. (1985). Review of Not in our Genes (S. Rose, L. J. Kamin & R. C. Lewontin). New Scientist 105, 59–60.

  Dawkins, R. (1987). Universal parasitism and the extended phenotype. In Evolution and Coadaptation in Biotic Communities (eds S. Kawano, J. H. Connell & T. Hidaka), pp. 183–197. Tokyo: University of Tokyo Press.

  Dawkins, R. (1989a). The evolution of evolvability. In Artificial Life (ed. C. Langton). Santa Fe, N.M.: Addison Wesley.

  Dawkins, R. (1989b). The Selfish Gene, 2nd edn. Oxford: Oxford University Press.

  Dawkins, R. (1990). Parasites, desiderata lists, and the paradox of the organism. In The Evolutionary Biology of Parasitism (eds A. E. Keymer & A. F. Read). Cambridge: Cambridge University Press.

  Dawkins, R. (1991). Darwin triumphant: Darwinism as a universal truth. In Man and Beast Revisited (eds M. H. Robinson & L. Tiger), pp. 23–39. Washington: Smithsonian Institution.

  Dawkins, R. (1996). Climbing Mount Improbable. New York: Norton.

  Dawkins, R. (1998). Unweaving the Rainbow. London: Penguin.

  Dennett, D. (1995). Darwin’s Dangerous Idea. New York: Simon & Schuster.

  Depew, D. J. & Weber, B. H. (1996). Darwinism Evolving. Cambridge, Mass.: MIT Press. de Winter, W. (1997). The beanbag genetics controversy: towards a synthesis of opposing views of natural selection. Biology and Philosophy 12, 149–184.

  Durham, W. H. (1991). Coevolution: Genes, Culture and Human Diversity. Stanford, Calif.: Stanford University Press.

  Eldredge, N. (1995). Reinventing Darwin: The Great Debate at the High Table of Evolutionary Theory. New York: John Wiley.

  Endler, J. A. (1986). Natural Selection in the Wild. Princeton, N.J.: Princeton University Press.

  Ewald, P. (1993). Evolution of Infectious Diseases. Oxford: Oxford University Press.

  Fletcher, D. J. C. & Michener, C. D. (1987). Kin Recognition in Humans. New York: Wiley.

  Fox Keller, E. and Lloyd, E. A. (1992). Keywords in Evolutionary Biology. Cambridge, Mass.: Harvard University Press.

  Futuyma, D. J. (1998). Evolutionary Biology, 3rd edn. Sunderland, Mass.: Sinauer.

  Goodwin, B. and Dawkins, R. (1995). What is an organism? A discussion. Perspectives in Ethology 2, 47–60.

  Gould, S. J. & Eldredge, N. (1993). Punctuated equilibrium comes of age. Nature 366, 222–227.

  Grafen, A. (1984). Natural selection, kin selection and group selection. In Behavioural Ecology: An Evolutionary Approach (eds J. R. Krebs & N. B. Davies). Oxford: Blackwell Scientific Publications.

  Grafen, A. (1985). A geometric view of relatedness. Oxford Surveys in Evolutionary Biology 2, 28–89.

  Grafen, A. (1990a). Biological signals as handicaps. Journal of Theoretical Biology 144, 517–546.

  Grafen, A. (1990b). Do animals really recognize kin? Animal Behaviour 39, 42–54.

  Grafen, A. (1991). Modelling in behavioural ecology. In Behavioural Ecology: An Evolutionary Ap
proach (eds J. R. Krebs & N. B. Davies). Oxford: Blackwell Scientific Publications.

  Guilford, T. & Dawkins, M. S. (1991). Receiver psychology and the evolution of animal signals. Animal Behaviour 42, 1–14.

  Haig, D. (1993). Genetic conflicts in human pregnancy. Quarterly Review of Biology 68, 495–532.

  Hamilton, W. D. (1996). Narrow Roads of Gene Land: The Collected Papers of W. D. Hamilton, i: Evolution of Social Behaviour. Oxford: W. H. Freeman/Spektrum.

  Hansell, M. H. (1984). Animal Architecture and Building Behaviour. London: Longman.

  Harvey, P. H. & Pagel, M. D. (1991). The Comparative Method in Evolutionary Biology. Oxford: Oxford University Press.

  Hecht, M. K. & Hoffman, A. (1986). Why not neo-Darwinism? A critique of paleobiological challenges. In Oxford Surveys in Evolutionary Biology 3, 1–47. Oxford: Oxford University Press.

  Hepper, P. G. (1986). Kin recognition: functions and mechanisms. A review. Biological Reviews 61, 63–93.

  Hoffman, A. (1989). Arguments on Evolution. New York: Oxford University Press.

  Hölldobler, B. & Wilson, E. O. (1990). The Ants. Berlin: Springer-Verlag.

  Hull, D. L. (1988a). Interactors versus vehicles. In The Role of Behaviour in Evolution (ed. H. C. Plotkin), pp. 19–50. Cambridge, Mass.: MIT Press.

  Hull, D. L. (1988b). Science as a Process. Chicago: University of Chicago Press.

  Keymer, A. & Read, A. (1989). Behavioural ecology: the impact of parasitism. In Parasitism: Coexistence or Conflict? (eds C. A. Toft & A. Aeschlimann). Oxford: Oxford University Press.

  Kimura, M. (1983). The Neutral Theory of Molecular Evolution. Cambridge: Cambridge University Press.

  Kitcher, P. (1985). Vaulting Ambition. Cambridge, Mass.: MIT Press.

  Koch, W. A. (1986). Genes vs. memes. Hagen: Druck Thiebes.

  Krebs, J. R. & Dawkins, R. (1984). Animal signals: mind-reading and manipulation. In Behavioural Ecology: An Evolutionary Approach (eds J. R. Krebs & N. B. Davies). Oxford: Blackwell Scientific Publications.

  Lloyd, E. A. (1988). The Structure and Confirmation of Evolutionary Theory. New York: Greenwood Press.

  Maynard Smith, J. (1982). Evolution and the Theory of Games. Cambridge: Cambridge University Press.

  Maynard Smith, J. (1986a). The Problems of Biology. Oxford: Oxford University Press.

  Maynard Smith, J. (1986b). Structuralism versus selection—is Darwinism enough? In Science and Beyond (eds S. Rose & L. Appignanesi). Oxford: Basil Blackwell.

  Maynard Smith, J. (1988). Did Darwin Get it Right? London: Penguin.

  Maynard Smith, J., Burian, R., Kauffman, S., Alberch, P., Campbell, J., Goodwin, B., Lande, R., Raup, D. & Wolpert, L. (1985). Developmental constraints and evolution. Quarterly Review of Biology 60, 265–287.

  Maynard Smith, J. & Szathmáry, E. (1995). The Major Transitions in Evolution. Oxford: W. H. Freeman/Spektrum.

  Mayr, E. (1983). How to carry out the adaptationist program. American Naturalist 121, 324–334.

  Minchella, D. J. (1985). Host life-history variation in response to parasitism. Parasitology 90, 205–216.

  Nesse, R. & Williams, G. C. (1995). Evolution and Healing: The New Science of Darwinian Medicine. London: Weidenfeld & Nicolson.

  Nunney, L. (1985). Group selection, altruism, and structured-deme models. American Naturalist 126, 212–229.

  Otte, D. & Endler, J. A. (eds) (1989). Speciation and its Consequences. Sunderland, Mass.: Sinauer.

  Oyama, S. (1985). The Ontogeny of Information. Cambridge: Cambridge University Press.

  Parker, G. A. & Maynard Smith, J. (1990). Optimality theory in evolutionary biology. Nature 348, 27–33.

  Pinker, S. (1997). How the Mind Works. London: Allen Lane, Penguin Press.

  Queller, D. C. (1992). Quantitative genetics, inclusive fitness, and group selection. American Naturalist 139, 540–558.

  Ridley, M. (1986). The Problems of Evolution. Oxford: Oxford University Press.

  Ridley, M. (1996). Evolution. Oxford: Blackwell Scientific.

  Ruse, M. (1996). Taking Darwin Seriously. Oxford: Basil Blackwell.

  Segerstråle, U. (1999). Defenders of the Truth. The Battle for ‘Good Science’ in the Sociobiology Debate and Beyond. Oxford: Oxford University Press.

  Sherman, P. W., Jarvis, J. U. M. & Alexander, R. D. (1991). The Biology of the Naked Mole-Rat. Princeton, N.J.: Princeton University Press.

  Sober, E. (1984). The Nature of Selection. Cambridge, Mass.: MIT Press.

  Sober, E. & Wilson, D. S. (1998). Unto Others. Cambridge, Mass.: Harvard University Press.

  Sterelny, K. & Kitcher, P. (1988). The return of the gene. Journal of Philosophy 85, 339–361.

  Sterelny, K., Smith, K. C. & Dickison, M. (1996). The extended replicator. Biology and Philosophy 11, 377–403.

  Stone, G. N. & Cook, J. M. (1998). The structure of cynipid oak galls: patterns in the evolution of an extended phenotype. Proceedings of the Royal Society of London B 265, 979–988.

  Trivers, R. L. (1985). Social Evolution. Menlo Park, N.J.: Benjamin/Cummings.

  Vermeij, G. J. (1987). Evolution and Escalation: An Ecological History of Life. Princeton, N.J.: Princeton University Press.

  Vollrath, F. (1988). Untangling the spider’s web. Trends in Ecology and Evolution 3, 331–335.

  Weiner, J. (1994). The Beak of the Finch. London: Cape.

  Werren, J. H., Nur, U. & Wu, C.-I. (1988). Selfish genetic elements. Trends in Ecology & Evolution 3, 297–302.

  Williams, G. C. (1985). A defence of reductionism in evolutionary biology. In Oxford Surveys in Evolutionary Biology (eds R. Dawkins & M. Ridley), pp. 1–27. Oxford: Oxford University Press.

  Williams, G. C. (1992). Natural Selection: Domains, Levels and Challenges. Oxford: Oxford University Press.

  Williams, G. C. (1997). Plan and Purpose in Nature. London: Weidenfeld & Nicolson.

  Wills, C. (1990). The Wisdom of the Genes. New York: Basic Books.

  Zahavi, A. & Zahavi, A. (1997). The Handicap Principle. Oxford: Oxford University Press.

  Glossary

  This book is primarily intended for biologists who will have no need of a glossary, but it has been suggested to me that it would be worth explaining a few technical terms to make the book more widely accessible. Many of the terms are well defined in other places (e.g. Wilson 1975; Bodmer & Cavalli-Sforza 1976). My definitions are certainly no improvement on those already available, but I have added personal asides on controversial words, or on matters of particular relevance to the thesis of this book. I have tried to avoid cluttering up the glossary with excessive numbers of explicit cross-references, but many of the words used in the definitions will be found to have their own definitions elsewhere in the glossary.

  adaptation A technical term which has evolved somewhat away from its common usage as a near synonym of ‘modification’. From sentences like ‘cricket wings are adapted (modified from their primary function of flying) for singing’ (and by implication are well designed for singing), ‘an adaptation’ has come to mean approximately an attribute of an organism that is ‘good’ for something. Good in what sense?, and good for what or for whom?, are difficult questions which are discussed at length in this book.

  alleles (short for allelomorphs) Each gene is able to occupy only a particular region of chromosome, its locus. At any given locus there may exist, in the population, alternative forms of the gene. These alternatives are called alleles of one another. This book emphasizes that there is a sense in which alleles are competitors of each other, because over evolutionary time successful alleles achieve numerical superiority over others at the same locus, in all the chromosomes of the population.

  allometry A disproportionate relationship between size of a body part and size of the whole body, the comparisons being made either across individuals or across different life stages in the same individual. For example, large ants (but small humans) tend to have relatively very large heads; the head grows at a different rate from the body as a whole. Mathemati
cally, the size of the part is usually taken as being related to the size of the whole raised to a power, which may be fractional.

  allopatric theory of speciation The widely supported view that the evolutionary divergence of populations into separate species (which no longer interbreed) takes place in geographically separate places. The alternative, sympatric theory gives rise to difficulties in understanding how the incipient species can separate if they are continuously in a position to interbreed with each other, and therefore to mix their gene-pools (q.v.).

  altruism Biologists use the word in a restricted (some would say misleadingly so) sense, only superficially related to common usage. An entity, such as a baboon or a gene, is said to be altruistic if it has the effect (not purpose) of promoting the welfare of another entity, at the expense of its own welfare. Various shades of meaning of ‘altruism’ result from various interpretations of ‘welfare’ (see page 57). Selfish is used in exactly the opposite sense.

  anaphase That phase of the cycle of cell division during which the paired chromosomes move apart. In meiosis (q.v.) there are two successive divisions and correspondingly two anaphases.

  anisogamy A sexual system in which fusion takes place at fertilization between a large (female) and a small (male) gamete. Contrast with isogamy in which there is sexual fusion but no male/female separation: all gametes are of roughly the same size.

  antibodies Protein molecules, produced in the immune response of animals, which neutralize invading foreign bodies (antigens).

  antigens Foreign bodies, usually protein molecules, which provoke the formation of antibodies.

  aposematism The phenomenon whereby distasteful or dangerous organisms like wasps ‘warn’ enemies by bright colours or equivalent strong stimuli. These are presumed to work by making it easy for the enemies to learn to avoid them, but there are (not insuperable) theoretical difficulties over how the phenomenon might evolve in the first place.

  assortative mating The tendency of individuals to choose mates that resemble (positive assortative mating or homogamy) or specifically do not resemble (negative assortative mating) themselves. Some people use the word only in the positive sense.

  autosome A chromosome that is not one of the sex chromosomes.