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List of Publications

  1. Grafen. A. 2024. What distinctions are usefully drawn today between natural and sexual selection? Chapter 6, pp 67-80, in: Sex, Gender, Ethics and the Darwinian Evolution of Mankind (ed. M. Veuille), Routledge. ISBN: 9781003405313
  2. Scott, T. W., Grafen, A. and S.A. West. 2023. Host–parasite coevolution and the stability of genetic kin recognition. Proceedings of the National Academy of Sciences (USA), 120, 1091-6490. doi: 10.1073/pnas.2220761120
  3. Scott, T. W., Grafen, A. and S.A. West. 2022. Multiple social encounters can eliminate Crozier’s paradox and stabilise genetic kin recognition. Nature Communications 13: 3902. doi: 10.1038/s41467-022-31545-4
  4. Levin, S. and A. Grafen 2021. Extending the range of additivity in using inclusive fitness. Ecology and Evolution, 11, 1970-1983. doi: 10.1002/ece3.6935
  5. Grafen, A. 2020. A simple completion of Fisher’s fundamental theorem of natural selection, Ecology and Evolution, 11, 735–742. doi: 10.1002/ece3.6918
  6. Grafen, A. 2020. The Price equation and reproductive value. Philosophical Transactions of the Royal Society B, 375: 20190356. doi: 10.1098/rstb.2019.0356
  7. Grafen, A. 2019. Should we ask for more than consistency of Darwinism with Mendelism? Studies in History and Philosophy of Biological and Biomedical Sciences, 78. doi: 10.1016/j.shpsc.2019.101224
  8. Levin, S., and A. Grafen, A. 2019. Inclusive fitness is an indispensable approximation for understanding organismal design. Evolution, 73, 1066-1076. doi: 10.1111/evo.13739
  9. Grafen, A. 2018. Editorial: The fundamental theorem of natural selection. Journal of Animal Breeding and Genetics 135, 393-394. doi: 10.1111/jbg.12361
  10. Grafen, A. 2018. The left hand side of the Fundamental Theorem of Natural Selection. Journal of theoretical Biology 456, 175-189. doi: 10.1016/j.jtbi.2018.07.022
  11. Biernaskie, J.M., Grafen, A. and J.C. Perry. 2018. A general model of biological signals, from cues to handicaps. Evolution Letters. doi: 10.1002/evl3.57.
  12. Crewe, P., Gratwick, R. and A. Grafen. 2018. Defining fitness in an uncertain world. Journal of mathematical Biology 76, 1059-1099. doi: 10.1007/s00285-017-1164-z.
  13. Grafen, A. 2015. Biological fitness and the Price Equation in class-structured populations. Journal of theoretical Biology 373, 62-72. doi: 10.1016/j.jtbi.2015.02.014.
  14. Grafen, A. 2015. Biological fitness and the Fundamental Theorem of Natural Selection. American Naturalist 186, 1-14. doi: 10.1086/681585.
  15. Grafen. 2014. Total reproductive values for females and for males in sexual diploids are not equal. Journal of theoretical Biology 359, 233-235. doi: 10.1016/j.jtbi.2014.05.021.
  16. Biernaskie, J.M., Grafen, A. and J.C. Perry. 2014. The evolution of index signals to avoid the cost of dishonesty. Proc. R. Roy. Soc. B 281, 20140876. doi: 10.1098/rspb.2014.0876.
  17. Batty, C.J.K., Crewe, P., Grafen, A. and R. Gratwick. 2014. Foundations of a mathematical theory of darwinism. Journal of mathematical Biology 69, 295-334. doi: 10.1007/s00285-013-0706-2.
  18. Grafen, A. 2014. The formal darwinism project in outline: response to commentaries. Biology and Philosophy, 29, 281-292. doi: 10.1007/s10539-013-9413-z.
  19. Grafen, A. 2014. The formal darwinism project in outline. Biology and Philosophy, 29, 155-174. doi: 10.1007/s10539-013-9414-y.
  20. Abbot, P. and over 100 others. 2011. Inclusive fitness theory and eusociality. Nature 471, E1-E4. doi: 10.1038/nature09831.
  21. Taylor, P.D. and A. Grafen. 2010. Relatedness with different interaction configurations. Journal of theoretical Biology, 262, 391-397. doi: 10.1016/j.jtbi.2009.10.012.
  22. Grafen, A. 2009. Formalizing Darwinism and inclusive fitness theory. Philosophical Transactions of the Royal Society B, 364, 3135-3141. doi: 10.1098/rstb.2009.0056
  23. Gardner, A. and A. Grafen. 2009. Capturing the superorganism: a formal theory of group adaptation. Journal of Evolutionary Biology. doi: 10.1111/j.1420-9101.2008.01681.x.
  24. Grafen, A. 2008. The simplest formal argument for fitness optimisation. Journal of Genetics, 87, 421-433.
  25. Grafen, A, and M. Archetti. 2008. Natural selection of altruism in inelastic viscous homogeneous population. Journal of theoretical Biology, 252, 694-710.
  26. Grafen, A. 2007. An inclusive fitness analysis of altruism on a cyclical network. Journal of evolutionary Biology, 20, 2278-2283.
  27. Grafen, A. 2007. The formal Darwinism project: a mid-term report. Journal of evolutionary Biology 20, 1243-1254. doi: 10.1111/j.1420-9101.2007.01321.x
  28. Grafen, A. 2007. Detecting kin selection at work using inclusive fitness. Proceedings of the Royal Society (London) B 274, 713-719.
  29. Grafen, A. 2006. Various remarks on Lehmann and Kellerís article. Journal of evolutionary Biology 19, 1397-1399.
  30. Grafen, A. 2006. A theory of Fisher’s reproductive value. Journal of mathematical Biology 53, 15-60.
  31. Grafen A, 2006. Optimization of inclusive fitness, Journal of theoretical Biology 238, 541-563
  32. Nee S, Colegrave N, West SA, and A. Grafen. 2006 Response to comment on “The illusion of invariant quantities in life histories” Science 312, 5771.
  33. Nee, S, N. Colegrave, S.A. West and A. Grafen. 2005. The illusion of invariant quantities in life histories. Science 309: 1236-1239 (Profiled in: de Jong, Is Invariance Across Animal Species Just an Illusion?, Science 309: 1193-1195).
  34. Grafen, A. 2004. Willam Donald Hamilton. Biographical Memoirs of Fellows of the Royal Society, 50, 109-132.
  35. Goymann W., I.T. Moore, A. Scheuerlein, K. Hirschenhauser, A. Grafen, and J.C. Wingfield. 2004. Testosterone in tropical birds: Effects of environmental and social factors. American Naturalist, 164, 327-334.
  36. Axelrod R., R.A. Hammond, and A. Grafen 2004. Altruism via kin-selection strategies that rely on arbitrary tags with which they coevolve. Evolution, 58, 1833-1838.
  37. Belshaw, R., A. Grafen, and D.L.J. Quicke. 2003. Inferring life history from ovipositor morphology in parasitoid wasps using phylogenetic regression and discriminant analysis. Zoological Journal of the Linnaean Society, 139, 213-228.
  38. Grafen, A. 2003. Fisher the evolutionary biologist. Journal of the Royal Statistical Society: Series D (The Statistician), 52, 319-329.
  39. Grafen, A. 2002. A First Formal Link between the Price Equation and an Optimization Program. Journal of theoretical Biology, 217, 75-91.
  40. Grafen A and R. Hails. 2002. Modern Statistics for the Life Sciences, Oxford University Press.
  41. Grafen, A. 2002. A state-free optimization model for sequences of behaviour. Animal Behaviour 63, 183-191. Download Software Package
  42. Grafen, A. 2000. Developments of the Price equation and natural selection under uncertainty. Proceedings of the Royal Society (London) B, 267, 1223-1227.
  43. Grafen, A. 2000. A biological approach to economics through fertility. Economics Letters, 66, 241-248.
  44. Grafen, A. 1999. Formal Darwinism, the individual-as-maximising-agent analogy, and bet-hedging. Proceedings of the Royal Society (London) B, 266, 799-803.
  45. Grafen, A. 1999. A note on errors in Grafen’s strategic handicap models - A note in response. Journal of theoretical Biology, 195, 416-417.
  46. Grafen, A. 1999. The correspondence of Charles Darwin, vol 10, 1862 - Burkhardt,F, Porter,DM, Harvey,J, Topham,JR. Notes and Records of the Royal Society of London, 53, 175-176.
  47. Grafen, A. 1998. Fertility and labour supply in Femina economica. Journal of theoretical Biology, 194, 429-455.
  48. Grafen, A. 1998. Evolutionary biology - Green beard as death warrant. Nature, 394, 521-522.
  49. Grafen, A and M. Ridley. 1997. Non-independence in statistical tests for discrete cross-species data. Journal of theoretical Biology, 188, 507-514.
  50. Newman, J.R., J. Bergelson and A. Grafen. 1997. Blocking factors and hypothesis tests in ecology: Is your statistics text wrong? Ecology, 78, 1312-1320.
  51. Grafen, A and M. Ridley. 1997. A new model for discrete character evolution. Journal of theoretical Biology, 184, 7-14. Download Software Package
  52. Grafen, A and M. Ridley. 1996. Statistical tests for discrete cross-species data. Journal of theoretical Biology, 183, 255-267.
  53. Ridley, M. and A. Grafen. 1996. How to study discrete comparative methods. In: Phylogenies and the Comparative Method in Animal Behavior (ed. E. P. Martins), 77-104, Oxford University Press.
  54. Quinnell, R.J., A. Grafen and M.E.J. Woolhouse. 1995. Changes in parasite aggregation with age - a discrete infection model. Parasitology, 111, 635-644.
  55. Kell, D.B., A.S. Kapreylants and A. Grafen. 1995. Pheromones, social behaviour and the functions of secondary metabolism in bacteria. Trends in Ecology and Evolution, 10, 126-129.
  56. Johnstone, R.A. and A. Grafen. 1993. Dishonesty and the handicap principle. Animal Behaviour, 46, 759-764.
  57. Grafen, A. and M.E.J. Woolhouse. 1993. Does the Negative Binomial Distribution add up? Parasitology Today, 9, 475-477.
  58. Hurst, LD and A. Grafen. 1993. Technical Comment: Unusual Mutational Mechanisms and Evolution. Science, 260, 1959.
  59. Grafen, A. and R.A. Johnstone. 1993. Why we need ESS signalling theory. Philosophical Transactions of the Royal Society (London) B 340, 245-250.
  60. Grafen, A. 1992. Of mice and the MHC. Nature, 360, 530.
  61. Johnstone, R.A. and A. Grafen. 1992. Error-prone signalling. Proceedings of the Royal Society (London) B, 248, 229-233.
  62. Johnstone, R.A. and A. Grafen. 1992. The continuous Sir Philip Sidney game: a simple model of biological signalling. Journal of theoretical Biology, 156, 215-234.
  63. Grafen, A. 1992. New Light on Modern Darwinismí (review of G.C. Williamsí “Natural Selection: Domains, Levels and Challenges”) Quarterly Review of Biology, 67, 343-345.
  64. Brockmann, H.J. and A. Grafen. 1992. Sex ratios and life-history patterns of a solitary wasp, Trypoxylon (Trypargilum) politum (Hymenoptera: Sphecidae). Behavioural Ecology and Sociobiology, 30, 7-27.
  65. Grafen, A. 1992. The uniqueness of the phylogenetic regression. Journal of theoretical Biology, 156, 405-423.
  66. Boomsma, J.J. and A. Grafen. 1991. Colony-level sex ratio selection in the eusocial Hymenoptera. Journal of evolutionary Biology, 4, 383-407.
  67. Haig, D.A. and A. Grafen. 1991. Genetic scrambling as a defence against meiotic driveí. Journal of theoretical Biology, 153, 531-558.
  68. Grafen, A. 1991. Kin vision?: a reply to Stuart. Animal Behaviour, 41, 1095-1087.
  69. Grafen, A. 1991. A reply to Byers and Bekoff. Animal Behaviour, 41, 1091-1092.
  70. Grafen, A. 1991. A reply to Blaustein et al. Animal Behaviour, 41, 1085-1087.
  71. Grafen, A. and H.C.J. Godfray 1991. Vicarious selection explains some paradoxes in dioecious fig-pollinator systems. Proceedings of the Royal Society (London) B, 245, 73-76.
  72. Grafen, A. 1991 Modelling in behavioural ecology. Chapter 1 of Behavioural Ecology, 3rd edition (ed.s J.R. Krebs and N.B. Davies), 5-31. Blackwell Scientific Publications, Oxford.
  73. Hurst, L.D. and A. Grafen 1990. Sex and flagellation. Trends in Ecology and Evolution, 5, 419-422.
  74. Grafen, A. 1990. Sexual selection unhandicapped by the Fisher process. Journal of theoretical Biology, 144, 473-516. See also Siller’s note and paper for mathematical corrections.
  75. Grafen, A. 1990. Biological signals as handicaps. Journal of theoretical Biology, 144, 517-546. See also Siller’s note and paper for mathematical corrections.
  76. Boomsma, J.J. and A. Grafen. 1990. Intra-specific variation in ant sex ratios and the Trivers-Hare hypothesis. Evolution, 44, 1026-1034.
  77. Grafen, A. 1990. Do animals really recognize kin? Animal Behaviour, 39, 42-54.
  78. Grafen, A. (editor) 1989. Evolution and its Influence: the Herbert Spencer lectures 1986, Clarendon Press, Oxford.
  79. Grafen, A. 1989. The phylogenetic regression. Philosophical Transactions of the Royal Society B, 326, 119-157. Download Software Package
  80. Brockmann, H.J. and A. Grafen 1989. Mate conflict and male behaviour in a solitary wasp Trypoxylon (Trypargilum) politum (Hymenoptera: Sphecidae). Animal Behaviour, 37, 232-255.
  81. Grafen, A. 1988. Origin of mutants disputed. Nature, 336, 525-526.
  82. Godfray, H.C.J. and A. Grafen 1988. ëUnmatedness and the evolution of eusocialityí. American Naturalist, 131, 303-305.
  83. Grafen, A. 1988. On the uses of data on lifetime reproductive success. Chapter 28 of Reproductive Success (ed. T.H. Clutton-Brock) 454-471, University of Chicago Press.
  84. Grafen, A. 1988. A centrosomal theory of the short term evolutionary maintenance of sexual reproductioní. Journal of theoretical Biology, 131, 163-173. See also Erratum, 132, 379.
  85. Grafen, A. 1987. Measuring sexual selection: why bother? In Sexual Selection: Testing the Alternatives (ed.s J.W. Bradbury and M.B. Andersson), 221-233, Wiley, New York.
  86. Grafen, A. 1987. The logic of divisively asymmetric contests: respect for ownership and the desperado effect. Animal Behaviour, 35, 462-467.
  87. Grafen, A. 1986. Split sex ratios and the evolutionary origins of eusociality. Journal of theoretical Biology, 122, 95-121.
  88. Sutherland, W.J., A. Grafen and P.H. Harvey 1986. Life history correlations and demography. Nature, 320, 88.
  89. Grafen, A. 1985. Hamilton’s rule OK. Nature, 318, 310-311.
  90. Grafen, A. 1985. A geometric view of relatedness. Oxford Surveys in evolutionary Biology, 2, 28-90.
  91. Lucas, J.R. and A. Grafen 1985. Partial prey consumption by ambush predators. Journal of theoretical Biology, 113, 455-473.
  92. Grafen, A. 1984. Mathematical appendix to: C.E.J. Kennedy and T.R.E. Southwood “The number of species of insects associated with British trees: a re-analysis”, Journal of Animal Ecology, 53, 455-478.
  93. Grafen, A. 1984. Natural selection, kin selection and group selection. Chapter 3 of Behavioural Ecology, 2nd edition (ed.s J.R. Krebs and N.B. Davies), 62-84. Blackwell Scientific Publications, Oxford.
  94. Grafen, A. and M. Ridley 1983. A model of mate guarding. Journal of theoretical Biology, 102, 549-567.
  95. Grafen, A. 1983. Review of “Evolution and the theory of games” by J. Maynard Smith. Times Higher Educational Supplement, 18th February.
  96. Grafen, A. 1982. How not to measure inclusive fitness. Nature, 298, 425-426.
  97. Ridley, M and A. Grafen 1981. Are green beard genes outlaws? Animal Behaviour, 29, 954-955.
  98. Grafen, A. 1980. Review of “The natural selection of populations and communities” by D.S. Wilson. Nature, 284, 494.
  99. Grafen, A. 1980. Opportunity cost, benefit and degrees of relatedness. Animal Behaviour, 28, 967-968.
  100. Grafen, A. 1979. The hawk-dove game played between relatives. Animal Behaviour, 27, 905-907.
  101. Brockmann, H.J., A. Grafen and R. Dawkins 1979. Evolutionarily stable nesting strategy in a digger wasp. Journal of theoretical Biology, 77, 473-496.
  102. Grafen, A. and R. Sibly 1978. A model of mate desertion. Animal Behaviour, 26, 645-652.

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