Google Scholar profiles of current members: Alexandra, Andrew, Bryce, Chai, Elliot, Erol, Marco

Working papers and preprints

Tilman, A. R., Plotkin, J. B., and Akçay, E. Evolutionary games with environmental feedbacks [on bioRxiv, doi:10.1101/493023]

Molina, C., Akçay, E., Dieckmann, U., Levin, S.A., and Rovenskaya, E.A. Combating climate change with matching-commitment agreements. [on SocArXiv, doi:10.31235/]

Smolla, M., and Akçay, E. Cultural selection shapes network structure. [on bioRxiv, doi:10.1101/464883]

Morsky, B. and Akçay, E. The blind choreographer: evolution of social norms and correlated equilibria. [on bioRxiv, doi:10.1101/435586]

Zhou, H., Akçay, E., and Helliker, B. Deriving C4 photosynthesis parameters by fitting intensive A/Ci curves [on bioRxiv, doi: 10.1101/153072]

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.  (available at SSRN)

2018, in press

Brown, A. and Akçay, E. (2018) Evolution of transmission mode in conditional mutualisms with spatial variation in symbiont quality. Evolution. doi:10.1111/evo.13656 [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 [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]


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]

Some old press:

NIMBioS interview
Another interview (also in Turkish) — Turkish Journal
An interview (in Turkish) — Evrim Çalışkanları
Overthrowing Darwin’s number 2 — Seed Magazine


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