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List of Publications
- Grafen, A. 2020. The Price equation and reproductive value. Philosophical Transactions of the Royal Society B, 375. doi: 10.1098/rstb.2019.0356
- 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
- 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
- Grafen, A. 2018. Editorial: The fundamental theorem of natural selection. Journal of Animal Breeding and Genetics 135, 393-394. doi: 10.1111/jbg.12361
- 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
- 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.
- 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.
- 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.
- Grafen, A. 2015. Biological fitness and the Fundamental Theorem of Natural Selection. American Naturalist 186, 1-14. doi: 10.1086/681585.
- 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.
- 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.
- 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.
- 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.
- Grafen, A. 2014. The formal darwinism project in outline. Biology and Philosophy, 29, 155-174. doi: 10.1007/s10539-013-9414-y.
- Abbot, P. and over 100 others. 2011. Inclusive fitness theory and eusociality. Nature 471, E1-E4. doi: 10.1038/nature09831.
- 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.
- 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
- 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.
- Grafen, A. 2008. The simplest formal argument for fitness optimisation. Journal of Genetics, 87, 421-433.
- Grafen, A, and M. Archetti. 2008. Natural selection of altruism in inelastic viscous homogeneous population. Journal of theoretical Biology, 252, 694-710.
- Grafen, A. 2007. An inclusive fitness analysis of altruism on a cyclical network. Journal of evolutionary Biology, 20, 2278-2283.
- 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
- Grafen, A. 2007. Detecting kin selection at work using inclusive fitness. Proceedings of the Royal Society (London) B 274, 713-719.
- Grafen, A. 2006. Various remarks on Lehmann and Kellerís article. Journal of evolutionary Biology 19, 1397-1399.
- Grafen, A. 2006. A theory of Fisher’s reproductive value. Journal of mathematical Biology 53, 15-60.
- Grafen A, 2006. Optimization of inclusive fitness, Journal of theoretical Biology 238, 541-563
- 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.
- 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).
- Grafen, A. 2004. Willam Donald Hamilton. Biographical Memoirs of Fellows of the Royal Society, 50, 109-132.
- 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.
- 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.
- 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.
- Grafen, A. 2003. Fisher the evolutionary biologist. Journal of the Royal Statistical Society: Series D (The Statistician), 52, 319-329.
- Grafen, A. 2002. A First Formal Link between the Price Equation and an Optimization Program. Journal of theoretical Biology, 217, 75-91.

- Grafen A and R. Hails. 2002. Modern Statistics for the Life Sciences, Oxford University Press.
- Grafen, A. 2002. A state-free optimization model for sequences of behaviour. Animal Behaviour 63, 183-191. Download Software Package
- Grafen, A. 2000. Developments of the Price equation and natural selection under uncertainty. Proceedings of the Royal Society (London) B, 267, 1223-1227.
- Grafen, A. 2000. A biological approach to economics through fertility. Economics Letters, 66, 241-248.
- Grafen, A. 1999. Formal Darwinism, the individual-as-maximising-agent analogy, and bet-hedging. Proceedings of the Royal Society (London) B, 266, 799-803.
- Grafen, A. 1999. A note on errors in Grafen’s strategic handicap models - A note in response. Journal of theoretical Biology, 195, 416-417.
- 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.
- Grafen, A. 1998. Fertility and labour supply in Femina economica. Journal of theoretical Biology, 194, 429-455.
- Grafen, A. 1998. Evolutionary biology - Green beard as death warrant. Nature, 394, 521-522.
- Grafen, A and M. Ridley. 1997. Non-independence in statistical tests for discrete cross-species data. Journal of theoretical Biology, 188, 507-514.
- 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.
- Grafen, A and M. Ridley. 1997. A new model for discrete character evolution. Journal of theoretical Biology, 184, 7-14. Download Software Package
- Grafen, A and M. Ridley. 1996. Statistical tests for discrete cross-species data. Journal of theoretical Biology, 183, 255-267.
- 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.
- 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.
- 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.
- Johnstone, R.A. and A. Grafen. 1993. Dishonesty and the handicap principle. Animal Behaviour, 46, 759-764.
- Grafen, A. and M.E.J. Woolhouse. 1993. Does the Negative Binomial Distribution add up? Parasitology Today, 9, 475-477.
- Hurst, LD and A. Grafen. 1993. Technical Comment: Unusual Mutational Mechanisms and Evolution. Science, 260, 1959.
- Grafen, A. and R.A. Johnstone. 1993. Why we need ESS signalling theory. Philosophical Transactions of the Royal Society (London) B 340, 245-250.
- Grafen, A. 1992. Of mice and the MHC. Nature, 360, 530.
- Johnstone, R.A. and A. Grafen. 1992. Error-prone signalling. Proceedings of the Royal Society (London) B, 248, 229-233.
- 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.
- 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.
- 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.
- Grafen, A. 1992. The uniqueness of the phylogenetic regression. Journal of theoretical Biology, 156, 405-423.
- Boomsma, J.J. and A. Grafen. 1991. Colony-level sex ratio selection in the eusocial Hymenoptera. Journal of evolutionary Biology, 4, 383-407.
- Haig, D.A. and A. Grafen. 1991. Genetic scrambling as a defence against meiotic driveí. Journal of theoretical Biology, 153, 531-558.
- Grafen, A. 1991. Kin vision?: a reply to Stuart. Animal Behaviour, 41, 1095-1087.
- Grafen, A. 1991. A reply to Byers and Bekoff. Animal Behaviour, 41, 1091-1092.
- Grafen, A. 1991. A reply to Blaustein et al. Animal Behaviour, 41, 1085-1087.
- 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.
- 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.
- Hurst, L.D. and A. Grafen 1990. Sex and flagellation. Trends in Ecology and Evolution, 5, 419-422.
- 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.
- Grafen, A. 1990. Biological signals as handicaps. Journal of theoretical Biology, 144, 517-546. See also Siller’s note and paper for mathematical corrections.
- Boomsma, J.J. and A. Grafen. 1990. Intra-specific variation in ant sex ratios and the Trivers-Hare hypothesis. Evolution, 44, 1026-1034.
- Grafen, A. 1990. Do animals really recognize kin? Animal Behaviour, 39, 42-54.
- Grafen, A. (editor) 1989. Evolution and its Influence: the Herbert Spencer lectures 1986, Clarendon Press, Oxford.
- Grafen, A. 1989. The phylogenetic regression. Philosophical Transactions of the Royal Society B, 326, 119-157. Download Software Package
- 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.
- Grafen, A. 1988. Origin of mutants disputed. Nature, 336, 525-526.
- Godfray, H.C.J. and A. Grafen 1988. ëUnmatedness and the evolution of eusocialityí. American Naturalist, 131, 303-305.
- 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.
- 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.
- 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.
- Grafen, A. 1987. The logic of divisively asymmetric contests: respect for ownership and the desperado effect. Animal Behaviour, 35, 462-467.
- Grafen, A. 1986. Split sex ratios and the evolutionary origins of eusociality. Journal of theoretical Biology, 122, 95-121.
- Sutherland, W.J., A. Grafen and P.H. Harvey 1986. Life history correlations and demography. Nature, 320, 88.
- Grafen, A. 1985. Hamilton’s rule OK. Nature, 318, 310-311.
- Grafen, A. 1985. A geometric view of relatedness. Oxford Surveys in evolutionary Biology, 2, 28-90.
- Lucas, J.R. and A. Grafen 1985. Partial prey consumption by ambush predators. Journal of theoretical Biology, 113, 455-473.
- 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.
- 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.
- Grafen, A. and M. Ridley 1983. A model of mate guarding. Journal of theoretical Biology, 102, 549-567.
- Grafen, A. 1983. Review of “Evolution and the theory of games” by J. Maynard Smith. Times Higher Educational Supplement, 18th February.
- Grafen, A. 1982. How not to measure inclusive fitness. Nature, 298, 425-426.
- Ridley, M and A. Grafen 1981. Are green beard genes outlaws? Animal Behaviour, 29, 954-955.
- Grafen, A. 1980. Review of “The natural selection of populations and communities” by D.S. Wilson. Nature, 284, 494.
- Grafen, A. 1980. Opportunity cost, benefit and degrees of relatedness. Animal Behaviour, 28, 967-968.
- Grafen, A. 1979. The hawk-dove game played between relatives. Animal Behaviour, 27, 905-907.
- Brockmann, H.J., A. Grafen and R. Dawkins 1979. Evolutionarily stable nesting strategy in a digger wasp. Journal of theoretical Biology, 77, 473-496.
- Grafen, A. and R. Sibly 1978. A model of mate desertion. Animal Behaviour, 26, 645-652.
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