Lab Publications

You might also want to check out the Erol’s Google Scholar profile, or that of some current and past lab members: Alexandra, Amiyaal, Andrew, Bryce, Chai, Haoran, Eeman, Elliot, Emerson, Marco, Rafael, Slimane

Working papers and preprints

Morsky, B., Plotkin, J. B., & Akçay, E. Indirect reciprocity with Bayesian reasoning and biases. [on SocArxiv, doi:10.31235/]

Akcay, E., & Ohashi, R. The floating duck syndrome: biased social learning leads to effort-reward imbalances [on SocArxiv, doi:10.31235/].

Pandula, N., Akcay, E., & Morsky, B. Indirect reciprocity with abductive reasoning. [on SocArxiv, doi: 10.31235/].

Abbasi, E., & Akçay, E. Host control and species interactions jointly determine microbiome community structure. [on bioRxiv, doi:10.1101/2022.03.03.482885].

Tilman, A. R., Vasconcelos, V. V., Akçay, E., & Plotkin, J. B. The evolution of forecasting for decision making in dynamic environments. [on arXiv:2108.00047].

Zhou, H., Akçay, E., Edwards, E., & Helliker, B. The legacy of C4 evolution in the hydraulics of C3 and C4 grasses. [on bioRxiv, doi:10.1101/2020.05.14.097030].

Brown, A., & Akçay, E. Evolution of symbiont transmission in conditional mutualisms in spatially and temporally variable environments. [on bioRxiv, doi:10.1101/2020.05.05.079103].

Brown, A. and Akçay, E. Phylogenetic analysis suggests joint control of transmission mode in a grass-endophyte symbiosis [on bioRxiv, doi:10.1101/085746].

Akçay, E., Roughgarden, J., Fearon, J., Ferejohn, J. & Weingast, B. Biological institutions: the political science of animal cooperation. [on SSRN].


Morsky, B., Magpantay, F., Day, T., & Akçay, E. (2023). The impact of threshold decision mechanisms of collective behavior on disease spread. Proceedings of the National Academy of Sciences, 120(19), e2221479120. [link to journal] [pdf].

Carlson, C., Akçay, E., & Morsky, B. (2023). The evolution of partner specificity in mutualisms. Evolution, 77(3), 881-892. [link to journal] [pdf].

Smolla, M., & Akçay, E. (2023). Pathways to cultural adaptation: The coevolution of cumulative culture and social networks. Evolutionary Human Sciences, 5, E26. [link to open access article]

Zhou, H., Akçay, E., & Helliker, B. (2023). Optimal coordination and reorganization of photosynthetic properties in C4 grasses. Plant, Cell & Environment, 46(3), 796-811. [link to journal] [pdf]


Ventura, R., & Akcay, E. (2022). A Cognitive-evolutionary Model for the Evolution of Teaching. Journal of Theoretical Biology, 533, 110933 [link to journal] [pdf].

MacDonald, H., Akçay, E., & Brisson, D. (2022). Host phenology can drive the evolution of intermediate virulence strategies in some obligate-killer parasites. Evolution, 76(6), 1183-1194. [link to journal] [pdf]

McGlothlin, J. W., Akçay, E., Brodie III, E. D., Moore, A. J., & Van Cleve, J. (2022). A synthesis of game theory and quantitative genetic models of social evolution. Journal of Heredity, 113(1), 109-119. [link to journal] [pdf]


Ilany, A., Holekamp, K. E., & Akçay, E. (2021). Rank-dependent social inheritance determines social network structure in spotted hyenas. Science, 373(6552), 348-352. [link to journal] [pdf]

Perry, S., Carter, A., Smolla, M., Akçay, E., Nöbel, S., Foster, J. G., & Healy, S. D. (2021). Not by transmission alone: the role of invention in cultural evolution. Philosophical Transactions of the Royal Society B, 376(1828), 20200049. [link to journal] [pdf]

Akçay, E., & Hirshleifer, D. (2021). Social finance as cultural evolution, transmission bias, and market dynamics. Proceedings of the National Academy of Sciences, 118(26), e2015568118. [link to open access article]

MacDonald, H., Akçay, E., & Brisson, D. (2021). The role of host phenology for parasite transmission. Theoretical Ecology, 14, 123-143. [link to journal] [pdf]

Morsky, B., & Akçay, E. (2021). False beliefs can bootstrap cooperative communities through social norms. Evolutionary Human Sciences, 3, e36. [link to open access article]

Akçay, E. and Van Cleve, J. (2021) Internalizing cooperative norms in group-structured populations. to appear in Social Cooperation and Conflict: Biological Mechanisms at the Interface, edited by W. Wilczynski and S.F. Brosnan. Cambridge University Press [on bioRxiv, doi:10.1101/722439] [pdf]


Morsky, B., Smolla, M., & Akçay, E. (2020). Evolution of contribution timing in public goods games. Proceedings of the Royal Society B, 287(1927), 20200735. [link to journal]  [on bioRxiv, doi: 10.1101/2020.01.09.900670]

Molina, C., Akçay, E., Dieckmann, U., Levin, S.A., and Rovenskaya, E.A. (2020) Combating climate change with matching-commitment agreements. Scientific Reports. [link to open access article]

Qian, J. J. and Akçay, E. (2020) The balance of interaction types determines the assembly and stability of ecological communities. Nature Ecology and Evolution [link to journal] [on bioRxiv, doi: 10.1101/643478]

Tilman, A. R., Plotkin, J. B., and Akçay, E. (2020) Evolutionary games with environmental feedbacks 11:915 Nature Communications [link to open access article][on bioRxiv, doi:10.1101/493023]

Akçay, E. (2020) Deconstructing evolutionary game theory: coevolution of social behaviors with their evolutionary setting. The American Naturalist 195:315-330 doi: 10.1086/706811 [pdf] [link to journal]


Smolla, M., and Akçay, E. (2019) Cultural selection shapes network structure. Science Advances. 5:eaaw0609 doi: 10.1126/sciadv.aaw0609 [link to open access article]

Morsky, B. and Akçay, E. The blind choreographer: evolution of social norms and correlated equilibria. PNAS. 116 (18) 8834-8839. doi: 10.1073/pnas.1817095116 [pdf] [link to journal]

Zhou, H., Akçay, E., and Helliker, B. (2019) Deriving C4 photosynthesis parameters by fitting intensive A/Ci curves. Photosynthesis Research. 141: 181-194 doi: 10.1007/s11120-019-00619-8 [pdf] [link to journal]

Brown, A. and Akçay, E. (2019) Evolution of transmission mode in conditional mutualisms with spatial variation in symbiont quality. Evolution. 73: 128-144. doi:10.1111/evo.13656 [pdf] [link to journal] [on bioRxiv, doi:


Zhou, H., Helliker, B., Huber, M., Dicks, A. Akçay, E. (2018) C4 photosynthesis and climate through the lens of optimality. PNAS 115: 12057-12062 [pdf] [link to journal][previous version on bioRxiv, doi:10.1101/048900]

Akçay, E. (2018) Collapse and rescue of cooperation in evolving dynamic networks. Nature Communications 9: 2692 [on bioRxiv, doi:10.1101/226563] [link to open access article]

Akçay, E., Meirowitz, A., and Ramsay, K. (2018) Two-sided Unobservable Investment, Bargaining, and Efficiency. Review of Economic Design doi: 10.1007/s10058-018-0213-4 [pdf] [link to journal pdf]

Aguilar, E. and Akçay, E. (2018) Gene-culture co-inheritance of a behavioral trait. The American Naturalist [pdf] [link to journal] [on bioRxiv, doi:10.1101/069286]

Qian, J. and Akçay, E. (2018) Competition and niche construction in a model of cancer metastasis. PLoS One 13:e0198163 [link to open access article] [on bioRxiv, doi:10.1101/228585] [Penn Today Article]

Dridi, S. and Akçay, E. (2018) Learning to cooperate: The evolution of social rewards in repeated interactions. The American Naturalist 191: 58-73 [pdf] [link to journal] [on bioRxiv, doi:10.1101/074096]


Bienvenu, F., Akçay, E., Legendre, S., and McCandlish, D.M. (2017) The genealogical decomposition of a matrix population model with applications to the aggregation of stages. Theoretical Population Biology 115: 69-80 [ungated link to journal] [on bioRxiv, doi:10.1101/067793]

Akçay, E. (2017) Population structure reduces the benefits from partner choice in mutualism. Proceedings of the Royal Society B: 20162317 [link to journal] [bioRxiv version, doi:10.1101/068445]


Ilany, A. and Akçay, E. (2016) Personality and social networks: a generative model approach. Integrative and Comparative Biology doi: 10.1093/icb/icw068  [ungated link to journal]

Ilany, A. and Akçay, E. (2016) Social inheritance can explain the structure of animal social networks. Nature Communications 7:12084 [link to open access article] [Penn Today Article]

Lehmann, L., Mullon, C., Akçay, E., and Van Cleve, J. (2016) Invasion fitness and inclusive fitness in heterogenous subdivided populations. Evolution (doi: 10.1111/evo.12980) [on bioRxiv] [link to journal]

Akçay, E. and Van Cleve, J (2016) There is no fitness but fitness, and the lineage is its bearer. Phil. Trans. R. Soc. B 371: 20150085 [pdf] [link to journal]


Akçay, E., Linksvayer, T.A., and Van Cleve, J. (2015) Bridging social evolution theory and emerging empirical approaches to social behavior. Current Opinion in Behavioral Sciences 6:59–64 [pdf] [link to journal]

Jones, E.I., Afkhami, M.E., Akçay, E., Bronstein, J.L., Bshary, R., Frederickson, M.E., Heath, K.D., Hoeksema, J.D., Ness, J.H., Pankey, M.S., Porter, S.S., Sachs, J.L., Scharnagl, K. and Friesen, M.L. (2015) Cheaters must prosper: reconciling theoretical and empirical perspectives on cheating in mutualism. Ecology Letters 18:1270–1284 [pdf] [link to journal]

Akçay, E. (2015) Evolutionary models of mutualisms.  in Mutualism, edited by J. Bronstein, Oxford University Press [pdf] [link to book]

Bewick, S., Chisholm, R.A., Akçay, E., Godsoe, W. (2015) A stochastic biodiversity model with overlapping niche structure. Theoretical Ecology: 8: 81-109. [pdf] [link to journal]


Van Cleve, J. and Akçay, E. (2014) Pathways to social evolution: reciprocity, relatedness, and synergy. Evolution. 68:2245-2258 [link to journal] [final version on arXiv, bioarXiv]

Shen, S.-F., Akçay, E., Rubenstein, D.R. (2014). Does social conflict increase with group size? The American Naturalist. 183:301-310 [pdf] [link to journal]

2013 and before

Pringle, E. G., Akçay, E., Raab, T. K., Gordon, D. M., Dirzo, R. (2013) Availability of limiting resources determines context-dependent strength of an ant-plant-hemipteran mutualism. PLoS Biol 11(11): e1001705. doi:10.1371/journal.pbio.1001705 [link to open access article]

Akçay, E., Meirowitz, A., Ramsay, K., Levin, S. A. (2012) Evolution of cooperation and skew under imperfect information. PNAS 109: 14936-14941 [pdf] [link to journal]

Akçay, E.*, J. Van Cleve*. (2012) Behavioral dynamics in structured populations pave the way to group optimality. The American Naturalist; 179:257-269 [pdf] [link to journal]

Akçay, E. (2012) Incentives in the family II: Behavioral dynamics and the evolution of non-costly signaling. Journal of Theoretical Biology;294:9-18 [pdf] [Supplementary Material] [link to journal]

Şekercioğlu, Ç.H., S. Anderson, E. Akçay, et al. (10 other authors). ‘Turkey’s globally important biodiversity in crisis’ Biological Conservation;144: 2752-2769 [pdf] [link to journal] [Highlighted by NY Times Green Blog]

Akçay, E. and E. Simms. 2011. ‘Negotiation, sanctions and context dependency in the legume-rhizobium mutualism.‘ The American Naturalist;178(1):1-14.  [pdf] [link to journal]

Mesterton-Gibbons, M., S. Gavrilets, J. Gravner and E. Akçay. 2011. ‘Models of coalition or alliance formation.’ Journal of Theoretical Biology;274:187-204. [pdf] [link to journal]

Akçay, E. and J. Roughgarden. 2011. ‘The evolution of payoff matrices: providing incentives to cooperate.’ Proc. R. Soc. B; 278:2198-2206.  [pdf] [Supplementary Material]  [link to journal] [Recommended by Faculty of 1000]

Roughgarden, J. and E. Akçay. 2010. ‘Do we need a sexual selection 2.0?’ Animal Behaviour;79(3):e1-e4 [link]
(a commentary on Clutton-Brock 2009 [link], and Carranza 2009 [link])

Roughgarden, J. and E. Akçay. 2010. ‘Final response: sexual selection needs an alternative’ Animal Behaviour;79(3):e18-e23 [link]  (our response to Shuker 2010 [link], who joined the exchange)

[pdf of the whole exchange]

Akçay, E.*, J. Van Cleve*, M. W. Feldman and J. Roughgarden. 2009. ‘A theory for the evolution of other-regarding motivations integrating proximate and ultimate perspectives’ PNAS;106: 19061-19066 [link] [pdf w/ SI]

Akçay, E. and J. Roughgarden. 2009. ‘The perfect family: biparental care in animals.’ PLoS One;4:(10)e7345 [link] [pdf]

Akçay, E. and J. Roughgarden. 2007. ‘Extra-pair parentage: a new theory based on transactions in a cooperative game.’ Evolutionary Ecology Research; 9:1223-1243 [link to journal] [pdf]

Akçay, E. and J. Roughgarden. 2007. ‘Extra-pair paternity in birds: Review of the genetic benefits.’ Evolutionary Ecology Research; 9:855-868 [link to journal] [pdf]

Akçay, E. and J. Roughgarden. 2007. ‘Negotiation of mutualism: rhizobia and legumes.’ Proc. R. Soc. B; 274:25 – 32. [link to journal] [pdf]

Roughgarden, J., M. Oishi and E. Akçay. 2006. ‘Reproductive social behavior: Cooperative games to replace sexual selection.’ Science; 311:965-970 [link to journal][pdf]
(also see the letters criticizing the article, and our response)

*: equal contributions

Letters, commentary, etc.

Kovaka, K., Santana, C., Patel, R., Akçay, E., Weisberg, M. (2016) Agriculture increases individual fitness. Behavioral and Brain Sciences 39:e103 [link to journal]

Akçay, E. An apology for inclusive fitness. (review of Marshall, J. 2015. Social evolution and inclusive fitness theory.) Evolution (forthcoming) [pdf]

Şekercioğlu, Ç.H., S. Anderson, E. Akçay and R. Bilgin ‘Turkey’s rich natural heritage under assault’ Science; 334: 1637-1638 [link] [pdf] (letter detailing recent conservation issues in Turkey)

Akçay, E. and J. Roughgarden. 2007. ‘Neither should competition be assumed.’ Trends in Ecology and Evolution; 22:4 [link to journal] [pdf]