Not by Genes Alone: How culture transformed human evolution gives an accessible treatment of my work (with Peter Richerson) on cultural evolution is now available from the University of Chicago Press (webpage).

Mathematical Models of Social Evolution: A guide for the perplexed provides an introduction to the mathematical theory of social behavior for readers with modest mathematical background (with Richard McElreath) Now available from the University of Chicago Press (webpage).

The Origin and Evolution of Cultures is a collection of papers on cultural evolution with Pete Richerson and includes five new essays. Available from Oxford University Press, 2005

Foundations of Human Sociality J. Henrich, R. Boyd, S. Bowles, C. Camerer, E. Fehr & H. Gintis eds. Oxford University Press, Oxford 2004 reports the results of cross cultural project in 15 small scale societies which used games from experimental economics to measure the sources of cooperative behavior. Oxford University Press webpage for this book.

Moral Sentiments and Material Interests
The Foundations of Cooperation in Economic Life

Herbert Gintis, Samuel Bowles, Robert T. Boyd and Ernst Fehr, eds. MIT Press, Cambridge, 2005 presents an synthesis of research in different disciplines to argue that cooperation stems presence of strong reciprocators in a social group. MIT Press website for this book.

Culture and the Evolutionary Process presents some of the theory of cultural evolution that Peter Richerson and I have developed is now back in print. To order visit the University of Chicago Press webpage for this book.

Below are downloadable versions of some of my papers. Most are in Adobe Acrobat (pdf) format. To down load a free copy of the Adobe Acrobat Reader. For a complete publication list see my CV. Last updated 5/1/21

J. Harris, R. Boyd, and B. Wood. The role of causal knowledge in the evolution of
traditional technology, Current Biology,, 2021 (pdf). Commentary by Joe Henrich (pdf)

K. Hill and R. Boyd, Behavioral convergence in humans and animals, Science, DOI: 10.1126/science.abf75722021.2021 (pdf)

R. Boyd and S. Mathew. Arbitration supports reciprocity when there are frequent perception errors, Nature Human Behavior, 2021 (pdf)


P. J. Richerson and R. Boyd The human life history is adapted to exploit the adaptive advantages of culture, Philosophical Transactions of the Royal Society (B),,, 2020. (pdf)

M. Kline, M. Gervais, C. Moya, and R. Boyd, Irrelevant‐action imitation is short‐term and contextual: Evidence from two under‐studied populations, Developmental Science, DOI: 10.1111/desc.12903, 2019.(pdf)

M. Derex, J-F Bonnefon, R. Boyd, and A. Mesoudi Causal understanding is not necessary for the improvement of culturally evolving technology, Nature Human Behavior, 3, 449-452, 2019, (pdf)

M. Derex and R. Boyd Social information can potentiate understanding despite inhibiting
cognitive effort, Scientific Reports, DOI:10.1038/s41598-018-28306-z, 2018 (pdf)

M. Derex, C. Perreault, and R. Boyd Divide and conquer: intermediate levels
of population fragmentation maximize cultural accumulation, Philosophical Transactions of the Royal Society (B), 373,  DOI: 10.1098/rstb.2017.0062, 2018 (pdf)

M. Derex and R. Boyd, Partial connectivity increases cultural accumulation within groups, Proceedings of the National Academy of Sciences (USA), 113, 2982-2987, 2016.(pdf)

R.H. Schonmann and R. Boyd, A simple rule for the evolution of contingent cooperation in large groups, Philosophical Transactions of the Royal Society (B), 317,  DOI: 10.1098/rstb.2015.0099, 2016 (pdf)

J. Henrich, M. Chudek and R. Boyd, The Big Man Mechanism: how prestige fosters cooperation and creates prosocial leaders, Philosophical Transactions of the Royal Society (B), DOI: 10.1098/rstb.2015.0013, 2015. (pdf)

C. Moya and  R. Boyd, Different selection pressures give rise to distinct ethnic phenomena: A functionalist framework with illustrations from the Peruvian Altiplano. Human Nature, 26(1). doi:10.1007/s12110-015-9224-9, 2015. (pdf)

M. Derex and R. Boyd, The foundations of the human cultural niche, Nature Communications, doi:10.1038/ncomms9398, 2015 (pdf)

C. Moya, R. Boyd, and J. Henrich. Reasoning about cultural and genetic transmission: Developmental and cross-cultural evidence from Peru, Fiji, and the United States on how people make inferences about trait transmission, Topics in Cognitive Science, 7, 1-16, 2015. (pdf)

R. Boyd and S. Mathew. Third-party monitoring and sanctions aid the evolution of language, Evolution and Human Behavior, 36, 475-479, 2015.(pdf)

R. Boyd, R. H. Schonmann and R. Vicente, Hunter–Gatherer population structure and the evolution of contingent cooperation, Evolution and Human Behavior, 35, 219–227, 2014.(pdf)

K. Hill, B. M. Wood, J. Baggio, A. M. Hurtado, R. Boyd, Hunter-Gatherer Inter-Band Interaction Rates: Implications for Cumulative Culture, PLoS One, 9, 1-9, 2014.(pdf)

S. Mathew and R. Boyd, The cost of cowardice. Evolution and Human Behavior, 35, 58–64. 2014 (pdf)

S. Mathew, R. Boyd, M. van Veelen. Human cooperation among kin and close associates may require enforcement of norms by third parties. In: Cultural Evolution, ed. P. J. Richerson and M. Christiansen, Strüngmann Forum Report 12,  Cambridge, MA: MIT Press, 2013,  (pdf)   

R. Boyd, P. J. Richerson and J. Henrich, The cultural evolution of technology, facts and theories, Cultural Evolution, Society, Technology, Science and Religion, M. Christiansen and P.J. Richerson, eds., MIT Press, 119–142, 2013 (pdf)

M. Kline, R. Boyd, and J. Henrich, Teaching and the life history of cultural transmission in Fijian villages, Human Nature, 24, 351–374, 2013.(pdf)

C. Perreault, C. Moya, and R. Boyd. A Baysian approach to the evolution of social learning. Evolution and Human Behavior, 33, 449-459,2012 (pdf).

J. Henrich, R. Boyd, and P. J. Richerson. The puzzle of monogamous marriage, Philosophical Transactions of the Royal Society (B), 367, 657–669, 2012 (pdf); Supplementary material (pdf).

R. Boyd, P. J. Richerson, and J. Henrich. The cultural niche: Why social learning is essential for human adaptation, Proceedings of the National Academy of Sciences (USA) 108: 10918–10925, 2011 (pdf). Commentary: Discover,

Mathew, S. and R. Boyd. Punishment sustains large-scale cooperation in prestate warfare. Proceedings of the National Academy of Sciences (USA)108: 11375–11380, 2011, (pdf). Supporting Information (pdf). Commentary: Richard McElreathDiscoverNew ScientistBaumard and Liénard, and our reply to Baumard and Liénard.

Boyd, R., P. J. Richerson, and J. Henrich. Rapid cultural adaptation can facilitate the evolution of large-scale cooperation, Behavioral Ecology and Sociobiology, 65:431–444, 2011 (pdf).

Boyd, R. Primed for reading, a review of Reading and the Brain by Stanislas Dehaene, PLoS Biology, 8:1–2, 2010, (pdf).

Boyd, R. and P. J. Richerson. Transmission coupling mechanisms: cultural group selection. Philosophical Transactions of the Royal Society (B), 365, 3787-3795, 2010, (pdf).


Kline, M. and R. Boyd, Population size predicts technological complexity in oceania, Proceedings of the Royal Society (B), 277, 2559–2564, 2010, (pdf) a Science story about the paper (pdf), and a Matt Ridley column in the Wall Street Journal (pdf).

Richerson, P. J., R. Boyd, and J. Henrich, Gene-culture coevolution in the age of genomics, Proceedings of the National Academy of Sciences (USA), 107, 8985–8992, 2010. (pdf)

Boyd, R., H. Gintis, and S. Bowles, Coordinated punishment of defectors sustains cooperation and can proliferate when rare, Science, 328, 617-620, 2010. (pdf)

Rendell, L., R. Boyd, D. Cownden, M. Enquist, K. Eriksson, M. W. Feldman, L. Fogarty, S. Ghirlanda, T. Lillicrap, and K. N.Laland, Why copy others? Insights from the social learning strategies tournament, Science, 328, 208-213, 2010. (pdf

R. Boyd and P. J. Richerson, Culture and the evolution of human cooperation, Philosophical Transactions of the Royal Society (B), 364:3281-3288, 2009. (pdf)

Richerson, P. J, R. Boyd, and R.L. Bettinger, Cultural Innovations and Demographic Change, Human Biology, 81, 211–235, 2009. (pdf)

Boyd, R. Does an evolutionary perspective help understand environmental degredation? Trends in Ecology and Evolution, 24: 72, 2009. (pdf)

Boyd, R. and P. J. Richerson, Voting with your feet: Payoff biased migration and the evolution of group beneficial behavior, Journal of Theoretical Biology, 257: 331-339, 2009. (pdf)

Mathew, S. and R. Boyd. When does optional participation allow the evolution of cooperation? Proceedings of the Royal Society (B). 276:1167-1174, 2009. (pdf)

P. J. Richerson and R. Boyd, Migration: An Engine for Social Improvement, Nature, 456: 18, 2008. (pdf)

J. Henrich and R. Boyd, Division of Labor, Economic Specialization and the Evolution of Social Stratification, Current Anthropology, 49: 715-724, 2008. (pdf) Online enhancements (pdf).

R. Boyd and P. J. Richerson. Gene-Culture Coevolution and the Evolution of Social Institutions, In: Better than Conscious? Decision Making, the Human Mind, and Implications for Institutions. C. Engel and W. Singer eds, MIT Press, Cambridge. Pp 305-324, 2008. (pdf)

J. Henrich, R. Boyd, and P. J. Richerson. Five Misunderstandings about Cultural Evolution. Human Nature, 19: 119–137, 2008. (pdf)

R. Boyd and S. Mathew. A Narrow Road to Cooperation, Science, 316: 1858–1859, 2007. (pdf).

S. Le and R. Boyd, Evolutionary Dynamics of the Continuous Iterated Prisoner's Dilemma, Journal of Theoretical Biology, Volume 245, 258–267. 2007, (pdf).

R. Boyd. The Puzzle of Human Sociality, Science, 314: 1553 2006. (pdf)

R. Boyd and P. J. Richerson. Culture, Adaptation, and Innateness. In: The Innate Mind: Culture and Cognition, P. Carruthers, S. Stich, & S. Laurence, eds., 2006.(pdf).

R. Boyd. Reciprocity: You have to think different. Journal of Evolutionary Biology, 19, 1380–1382. (pdf).

R. Boyd and P. J. Richerson. Culture and the evolution of the human social instincts. In: Roots of Human Sociality, S. Levinson and N. Enfield, eds., Berg, Oxford. 2006, (pdf)

R. Boyd and P. J. Richerson, Solving the Puzzle of Human Cooperation, In: Evolution and Culture, S. Levinson ed. MIT Press, Cambridge MA, pp 105–132, 2005(pdf).

J. Henrich, R. Boyd, S. Bowles, C. Camerer, E. Fehr, H. Gintis, R. McElreath, M. Alvard, A. Barr, J. Ensminger, K. Hill, F. Gil-White, M. Gurven, F. Marlowe, J. Q. Patton, N. Smith, and D. Tracer, 'Economic Man' in Cross-cultural Perspective: Behavioral Experiments in 15 Small-scale Societies, Behavioral and Brain Sciences, 28: 795–855, 2005. (pdf)

K. Panchanathan and R. Boyd. Indirect reciprocity can stabilize cooperation without the second-order free rider problem. Nature 432: 499–502, 2004. (pdf). Accompanying News & Views commentary by Ernst Fehr (pdf).

K. Panchanathan and R. Boyd. A Tale of Two Defectors: The Importance of Standing for the Evolution of Indirect Reciprocity. Journal of Theoretical Biology, 224:115–126, 2003, (pdf).

H. Gintis, S. Bowles, R. Boyd, & E. Fehr, Explaining Altruistic Behavior in Humans, Evolution and Human Behavior, 24: 153–172, 2003, (pdf).

P. J. Richerson, R. Boyd, and J. Henrich. The Cultural Evolution of Human Cooperation. The Genetic and Cultural Evolution of Cooperation, P. Hammerstein ed. MIT Press, Cambridge MA, 2003, pp 357–388 (pdf). 

R. Boyd, H. Gintis, S. Bowles, and P. J. Richerson. The Evolution of Altruistic Punishment. Proceedings of the National Academy of Sciences (USA) 100: 3531–3535, 2003 (pdf). Visual Basic 5 files used for simulation (.zip)

R. McElreath, R. Boyd and P. J. Richerson. Shared Norms Can Lead to the Evolution of Ethnic Markers. Current Anthropology, 44: 122–130, 2003 (pdf).

J. Henrich and R. Boyd, Culture and Cognition: Why Cultural Evolution Does Not Require Replication of Representations, Culture and Cognition, 2: 87–112, 2002. (pdf).

P. J. Richerson, R. Boyd and B. Paciotti. An Evolutionary Theory of Commons Management, The Drama of the Commons. Elinor Ostrom, Thomas Dietz, Nives Dolsak, Paul C. Stern, Susan Stonich, and Elke U. Weber, Eds., National Academy Press, Washington, D.C. 2002. pp 413–413 (pdf).

R. Boyd and P. J. Richerson, Group Beneficial Norms Spread Rapidly in a Structured Population, Journal of Theoretical Biology, 215: 287–296, 2002. (pdf).

P. J. Richerson and R. Boyd. Built for Speed, Not for Comfort, Darwinian Theory and Human Culture, History and Philosophy of the Life Sciences, 23: 425–465 (pdf).

R. Boyd and P. J. Richerson, Norms and Bounded Rationality. In: The Adaptive Tool Box, G. Gigerenzer and R. Selten, eds. pp 281–296, MIT Press, Cambridge MA, 2001. Abstract (html)

J. Henrich and R. Boyd, Why people punish defectors: Weak conformist transmission can stabilize costly enforcement of norms in cooperative dilemmas. Joural of Theoretical Biology, 208: 79–89. 2001 (pdf).

P. J. Richerson, R. Boyd, and R. L. Bettinger, Was Agriculture Iimpossible During the Pleistocene but Mandatory during the Holocene? A Climate Change Hypothesis, American Antiquity, 66: 387–411, 2001 (pdf).

J. Henrich, R. Boyd, S. Bowles, C. Camerer E. Fehr, H. Gintis, and R. McElreath, Cooperation, Reciprocity and Punishment in Fifteen Small-scale Societies, American Economic Review, 91: 73–78, 2001(pdf).

J. B. Silk, E. Kaldor, and R. Boyd, Cheap Talk When there are Conflict of Interest. Animal Behaviour, 59: 423-432, 2000 (pdf).

R. Boyd and P. J. Richerson, Memes: Universal Acid or a Better Mouse Trap. In: Darwinizing Culture: The Status of Memetics as a Science. R. Aunger ed. pp.143–162, Oxford University Press, Oxford. 2000 (pdf).

P. J Richerson and R. Boyd, The Evolutionary Dynamics of a Crude Super Organism. Human Nature, 10: 253–289, 1999 (pdf).

J. Henrich and R. Boyd, The Evolution of Conformist Transmission and the Emergence of Between-Group Differences,.Evolution and Human Behavior, 19: 215–242. 1998 (pdf).

P. J. Richerson and R. Boyd, The Evolution of Human Ultra-Sociality, In: Ideology, Warfare, and Indoctrinability. I. Eibl-Eibisfeldt and F. Salter, eds. pp. 71–95, Berghan Books, 1998 (pdf).

R. Boyd, P.J. Richerson, M. Borgerhoff-Mulder, and W. H. Durham. Are Cultural Phylogenies Possible? In: Human by Nature, Between Biology and the Social Sciences, P. Weingart, P.J. Richerson, S.D.Mitchell, and S. Maasen, eds. Lawrence Erlbaum Associates: Mahwah, NJ, 1997. Pp. 355–386 (pdf).

R. Boyd and P. J. Richerson, Why Culture is Common but Cultural Evolution is Rare, Proceedings of the British Academy, 88: 73–93, 1996 (pdf).

J. Soltis, R. Boyd and P. J. Richerson, Can Group-functional behaviors evolve by cultural group selection? An empirical test. Current Anthropology, 63: 473–494, 1995 (pdf).

R. Boyd. The evolution of reciprocity when conditions vary. In: Coalitions and Alliances in Humans and Other Animals, A. Harcourt and F DeWaal, eds., Oxford University Press, NY, 1992 (pdf).

R. Boyd and P. J. Richerson, Group Selection Among Alternative Evolutionarily Stable Strategies, Journal of Theoretical Biology, 145: 331-342, 1990. (pdf)

R. Boyd and P. J. Richerson, Why Does Culture Increase Human Adaptibility? Ethology and Sociobiology. 16: 125–143, 1995 (pdf).

R. Boyd and P. J. Richerson, Punishment Allows the Evolution of Cooperation (or Anything Else) in Sizable Groups, Ethology and Sociobiology, 13: 171-195, 1992 (pdf).

R. Boyd, Mistakes allow evolutionary stability in the repeated prisoner's dilemma game, Journal of Theoretical Biology, 136, 47–56, 1989 (pdf).

R. Boyd and P. J. Richerson, The Evolution of Indirect Reciprocity, Social Networks, 11: 213-236, 1989 (pdf).

R. Boyd and P. J. Richerson, The Evolution of Reciprocity in Sizable Groups, Journal of Theoretical Biology, 132: 337-356, 1988.(pdf).

R. Boyd and P. J. Richerson, The Evolution of Ethnic Markers, Cultural Anthropology, 2: 65–79, 1987. (pdf)

R. Boyd and J. P. Lorberbaum, No Pure Strategy is Stable in the Repeated Prisoner's Dilemma Game, Nature, 327: 58–59, 1987, (pdf)

R. Boyd and P. J. Richerson, Why is Culture Adaptive? Quarterly Review of Biology, 58: 209–214, 1983. (pdf)

R. Boyd, Density-dependent Mortality and the Evolution of Social Interactions, Animal Behaviour 30: 972-982, 1982, (pdf).

R. Boyd and P. J. Richerson, Effect of Phenotypic Variation on Kin Selection, Proceedings of the National Academy of Sciences (USA), 77: 7506-7509, 1980. (pdf)