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List of Publications

94 Grafen, A. 2018. Editorial: The fundamental theorem of natural selection. Journal of Animal Breeding and Genetics 135, 393-394. doi: 10.1111/jbg.12361

93 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

92 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.

91 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.

90 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.

89 Grafen, A. 2015. Biological fitness and the Fundamental Theorem of Natural Selection. American Naturalist 186, 1-14. doi: 10.1086/681585.

88 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.

87 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.

86 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.

85 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.

84 Grafen, A. 2014. The formal darwinism project in outline. Biology and Philosophy, 29, 155-174. doi: 10.1007/s10539-013-9414-y.

83 Abbot, P. and over 100 others. 2011. Inclusive fitness theory and eusociality. Nature 471, E1-E4. doi: 10.1038/nature09831.

82 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.

81 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

80 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. View Text in PDF

79 Grafen, A. 2008. The simplest formal argument for fitness optimisation. Journal of Genetics, 87, 421-433. View Text in PDF

78 Grafen, A, and M. Archetti. 2008. Natural selection of altruism in inelastic viscous homogeneous population. Journal of theoretical Biology, 252, 694-710. View Text in PDF

77 Grafen, A. 2007. An inclusive fitness analysis of altruism on a cyclical network. Journal of evolutionary Biology, 20, 2278-2283. View Text in PDF

76 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 View Text in PDF

75 Grafen, A. 2007. Detecting kin selection at work using inclusive fitness. Proceedings of the Royal Society (London) B 274, 713-719. View Text in PDF

74 Grafen, A. 2006. Various remarks on Lehmann and Keller’s article. Journal of evolutionary Biology 19, 1397-1399 View Text in PDF

73 Grafen, A. 2006. A theory of Fisher’s reproductive value. Journal of mathematical Biology 53, 15-60. View Text in PDF

72 Grafen A, 2006. Optimization of inclusive fitness, Journal of theoretical Biology 238, 541-563 View Text in PDF

71 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. View Text in PDF

70 Nee, S, N. Colegrave, S.A. West and A. Grafen. 2005. The illusion of invariant quantities in life histories. Science 309: 1236-1239 View Text in PDF (Profiled in: de Jong, Is Invariance Across Animal Species Just an Illusion?, Science 309: 1193-1195)

69 Grafen, A. 2004. ‘Willam Donald Hamilton’. Biographical Memoirs of Fellows of the Royal Society, 50, 109-132. View Text in PDF

68 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. View Text in PDF

67 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. View Text in PDF

66 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. View Text in PDF

65 Grafen, A. 2003. ‘Fisher the evolutionary biologist’. Journal of the Royal Statistical Society: Series D (The Statistician), 52, 319-329. View Text in PDF

64 Grafen, A. 2002. ‘A First Formal Link between the Price Equation and an Optimization Program’. Journal of theoretical Biology, 217, 75-91. View Text in PDF

DustJacket of Book by Grafen and Hails Buy (UK or Europe or rest of world); Buy (US); Companion website with teaching resources
63 Grafen A and R. Hails. 2002. ‘Modern Statistics for the Life Sciences’, Oxford University Press.

62  Grafen, A. 2002. ‘A state-free optimization model for sequences of behaviour’. Animal Behaviour 63, 183-191. View Text in PDF  Download Software Package

61  Grafen, A. 2000. ‘Developments of the Price equation and natural selection under uncertainty’. Proceedings of the Royal Society (London) B, 267, 1223-1227. View Text in PDF

60  Grafen, A. 2000. ‘A biological approach to economics through fertility’. Economics Letters, 66, 241-248. View Text in PDF

59  Grafen, A. 1999. ‘Formal Darwinism, the individual-as-maximising-agent analogy, and bet-hedging’. Proceedings of the Royal Society (London) B, 266, 799-803. View Text in PDF

58  Grafen, A. 1999. ‘A note on errors in Grafen's strategic handicap models - A note in response’. Journal of theoretical Biology, 195, 416-417. View Text in PDF

57  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.

56  Grafen, A. 1998. ‘Fertility and labour supply in Femina economica’. Journal of theoretical Biology, 194, 429-455. View Text in PDF

55  Grafen, A. 1998. ‘Evolutionary biology - Green beard as death warrant’. Nature, 394, 521-522. View Text in PDF

54  Grafen, A & M. Ridley. 1997. ‘Non-independence in statistical tests for discrete cross-species data’. Journal of theoretical Biology, 188, 507-514. View Text in PDF

53  Newman, J.R., J. Bergelson & A. Grafen. 1997. ‘Blocking factors and hypothesis tests in ecology: Is your statistics text wrong?’ Ecology, 78, 1312-1320. View Text in PDF

52  Grafen, A & M. Ridley. 1997. ‘A new model for discrete character evolution’. Journal of theoretical Biology, 184, 7-14. View Text in PDF  Download Software Package

51  Grafen, A & M. Ridley. 1996. ‘Statistical tests for discrete cross-species data’. Journal of theoretical Biology, 183, 255-267. View Text in PDF

50  Ridley, M. & 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. View Text in PDF

49  Quinnell, R.J., A. Grafen & M.E.J. Woolhouse. 1995. ‘Changes in parasite aggregation with age - a discrete infection model’. Parasitology, 111, 635-644. View Text in PDF

48  Kell, D.B., A.S. Kapreylants & A. Grafen. 1995. ‘Pheromones, social behaviour and the functions of secondary metabolism in bacteria’. Trends in Ecology and Evolution, 10, 126-129. View Text in PDF

47  Johnstone, R.A. & A. Grafen. 1993. ‘Dishonesty and the handicap principle’. Animal Behaviour, 46, 759-764. View Text in PDF

46  Grafen, A. & M.E.J. Woolhouse. 1993. ‘Does the Negative Binomial Distribution add up?’ Parasitology Today, 9, 475-477.

45  Hurst, LD & A. Grafen. 1993. ‘Technical Comment: Unusual Mutational Mechanisms and Evolution’. Science, 260, 1959.

44  Grafen, A. & R.A. Johnstone. 1993. ‘Why we need ESS signalling theory’. Philosophical Transactions of the Royal Society (London) B 340, 245-250. View Text in PDF

43  Grafen, A. 1992. ‘Of mice and the MHC’. Nature, 360, 530.

42  Johnstone, R.A. & A. Grafen. 1992. ‘Error-prone signalling’. Proceedings of the Royal Society (London) B, 248, 229-233.

41  Johnstone, R.A. & A. Grafen. 1992. ‘The continuous Sir Philip Sidney game: a simple model of biological signalling’. Journal of theoretical Biology, 156, 215-234. View Text in PDF

40  Grafen, A. 1992. ‘New Light on Modern Darwinism’ (review of G.C. Williams’ "Natural Selection: Domains, Levels and Challenges") Quarterly Review of Biology, 67, 343-345.

39  Brockmann, H.J. & 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.

38  Grafen, A. 1992. ‘The uniqueness of the phylogenetic regression’. Journal of theoretical Biology, 156, 405-423. View Text in PDF

37  Boomsma, J.J. & A. Grafen. 1991. ‘Colony-level sex ratio selection in the eusocial Hymenoptera’. Journal of evolutionary Biology, 4, 383-407. View Text in PDF

36  Haig, D.A. & A. Grafen. 1991. ‘Genetic scrambling as a defence against meiotic drive’. Journal of theoretical Biology, 153, 531-558. View Text in PDF

35  Grafen, A. 1991. ‘Kin vision?: a reply to Stuart’. Animal Behaviour, 41, 1095-1087. View Text in PDF

34  Grafen, A. 1991. ‘A reply to Byers & Bekoff’. Animal Behaviour, 41, 1091-1092. View Text in PDF

33  Grafen, A. 1991. ‘A reply to Blaustein et al.’ Animal Behaviour, 41, 1085-1087. View Text in PDF

32  Grafen, A. & 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. View Text in PDF

31  Grafen, A. 1991. ‘Modelling in behavioural ecology’. Chapter 1 of Behavioural Ecology, 3rd edition (ed.s J.R. Krebs & N.B. Davies), 5-31. Blackwell Scientific Publications, Oxford. View Text in PDF

30  Hurst, L.D. & A. Grafen 1990. ‘Sex and flagellation’. Trends in Ecology and Evolution, 5, 419-422.

29  Grafen, A. 1990. ‘Sexual selection unhandicapped by the Fisher process’. Journal of theoretical Biology, 144, 473-516. View Text in PDF. See also Siller's note and paper 58 for mathematical corrections.

28  Grafen, A. 1990. ‘Biological signals as handicaps’. Journal of theoretical Biology, 144, 517-546. View Text in PDF. See also Siller's note and paper 58 for mathematical corrections.

27  Boomsma, J.J. & A. Grafen. 1990. ‘Intra-specific variation in ant sex ratios and the Trivers-Hare hypothesis’. Evolution, 44, 1026-1034. View Text in PDF

26  Grafen, A. 1990. ‘Do animals really recognize kin?’ Animal Behaviour, 39, 42-54. View Text in PDF

25  Grafen, A. (editor) 1989. Evolution and its Influence: the Herbert Spencer lectures 1986, Clarendon Press, Oxford.

24  Grafen, A. 1989. ‘The phylogenetic regression’. Philosophical Transactions of the Royal Society B, 326, 119-157.  View Text in PDF (5.8Mb, 720dpi images) Download Software Package

23  Brockmann, H.J. & A. Grafen 1989. ‘Mate conflict and male behaviour in a solitary wasp Trypoxylon (Trypargilum) politum (Hymenoptera: Sphecidae)’. Animal Behaviour, 37, 232-255. View Text in PDF

22  Grafen, A. 1988. ‘Origin of mutants disputed’. Nature, 336, 525-526.

21  Godfray, H.C.J. & A. Grafen 1988. ‘Unmatedness and the evolution of eusociality’. American Naturalist, 131, 303-305. View Text in PDF

20  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. View Text in PDF

19  Grafen, A. 1988. ‘A centrosomal theory of the short term evolutionary maintenance of sexual reproduction’. Journal of theoretical Biology, 131, 163-173. See also Erratum, 132, 379. View Text in PDF

18  Grafen, A. 1987. ‘Measuring sexual selection: why bother?’ In Sexual Selection: Testing the Alternatives (ed.s J.W. Bradbury & M.B. Andersson), 221-233, Wiley, New York. View Text in PDF

17  Grafen, A. 1987. ‘The logic of divisively asymmetric contests: respect for ownership and the desperado effect’. Animal Behaviour, 35, 462-467. View Text in PDF

16  Grafen, A. 1986. ‘Split sex ratios and the evolutionary origins of eusociality’. Journal of theoretical Biology, 122, 95-121. View Text in PDF

15  Sutherland, W.J., A. Grafen & P.H. Harvey 1986. ‘Life history correlations and demography’. Nature, 320, 88.

14  Grafen, A. 1985. ‘Hamilton’s rule OK’. Nature, 318, 310-311.

13  Grafen, A. 1985. ‘A geometric view of relatedness’. Oxford Surveys in evolutionary Biology, 2, 28-90. View Text in PDF (6.8 Mb file, 600dpi images).

12  Lucas, J.R. & A. Grafen 1985. ‘Partial prey consumption by ambush predators’. Journal of theoretical Biology, 113, 455-473.

11  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.

10  Grafen, A. 1984. ‘Natural selection, kin selection and group selection’. Chapter 3 of Behavioural Ecology, 2nd edition (ed.s J.R. Krebs & N.B. Davies), 62-84. Blackwell Scientific Publications, Oxford. View Text in PDF

9  Grafen, A. & M. Ridley 1983. ‘A model of mate guarding’. Journal of theoretical Biology, 102, 549-567.

8  Grafen, A. 1983. Review of "Evolution and the theory of games" by J. Maynard Smith. Times Higher Educational Supplement, 18th February.

7  Grafen, A. 1982. ‘How not to measure inclusive fitness’. Nature, 298, 425-426. View Text in PDF

6  Ridley, M & A. Grafen 1981. ‘Are green beard genes outlaws?’ Animal Behaviour, 29, 954-955. View Text in PDF

5  Grafen, A. 1980. Review of "The natural selection of populations and communities" by D.S. Wilson. Nature, 284, 494.

4  Grafen, A. 1980. ‘Opportunity cost, benefit and degrees of relatedness’. Animal Behaviour, 28, 967-968. View Text in PDF

3  Grafen, A. 1979. ‘The hawk-dove game played between relatives’. Animal Behaviour, 27, 905-907. View Text in PDF

2  Brockmann, H.J., A. Grafen & R. Dawkins 1979. ‘Evolutionarily stable nesting strategy in a digger wasp’. Journal of theoretical Biology, 77, 473-496. View Text in PDF

1  Grafen, A. & R. Sibly 1978. ‘A model of mate desertion’. Animal Behaviour, 26, 645-652. View Text in PDF

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