We describe an iterated game between two players, in which the payoff is to survive a number of steps. Expected payoffs are probabilities of survival. A key feature of the game is that individuals have to survive on their own if their partner dies. We consider individuals with hardwired, unconditional behaviors or strategies. When both players are present, each step is a symmetric two-player game. The overall survival of the two individuals forms a Markov chain. As the number of iterations tends to infinity, all probabilities of survival decrease to zero. We obtain general, analytical results for n-step payoffs and use these to describe how the game changes as n increases. In order to predict changes in the frequency of a cooperative strategy over time, we embed the survival game in three different models of a large, well-mixed population. Two of these models are deterministic and one is stochastic. Offspring receive their parent’s type without modification and fitnesses are determined by the game. Increasing the number of iterations changes the prospects for cooperation. All models become neutral in the limit (n → ∞). Further, if pairs of cooperative individuals survive together with high probability, specifically higher than for any other pair and for either type when it is alone, then cooperation becomes favored if the number of iterations is large enough. This holds regardless of the structure of pairwise interactions in a single step. Even if the single-step interaction is a Prisoner’s Dilemma, the cooperative type becomes favored. Enhanced survival is crucial in these iterated evolutionary games: if players in pairs start the game with a fitness deficit relative to lone individuals, the prospects for cooperation can become even worse than in the case of a single-step game.

## WELCOME

We are devoted to the study of theoretical population genetics. The goal of population genetics is to identify and understand the forces that produce and maintain genetic variation in natural populations. These forces include mutation (also recombination and gene conversion), natural selection, various kinds of population structure (e.g. subdivision with migration), and the random fluctuations of gene frequencies through time known as genetic drift. We study these forces mathematically, using both analysis and computation. We also develop statistical methods to make inferences about these forces from DNA sequences or other kinds of genetic data. For more information about specific areas of research, follow the leads to lab members.

## RECENT PUBLICATIONS

A non-zero variance of Tajima’s estimator for two sequences even for infinitely many unlinked loci. Theoretical Population Biology. 2018;122 :22-29.Abstract

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Public goods games in populations with fluctuating size. Theoretical Population Biology. 2018;121 :72-84.Abstract

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Population structure and coalescence in pedigrees: Comparisons to the structured coalescent and a framework for inference. Theoretical Population Biology. 2017;115 :1-12.Abstract

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