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ESS: Evolutionary Stable Strategies Game Theory Module

This excel workbook illustrates a number of ESS examples. Each example allows the user to manipulate the initial population for each strategy and the fitness payoffs and costs for each strategy. The workbook projects population proportion and fitness over 100 generations, and includes both symmetrical and asymmetrical examples.


John R. Jungck, Beloit College
Jennifer Spangenberg

Published by: BioQUEST Curriculum Consortium

OS: win98:win2000:winXP:os9:osx:linux

User Manuals and Curricular Materials
Popular Text Citations

Weibull, Jörgen. (1995). Evolutionary Game Theory. Cambridge, MA: The M.I.T. Press; Cloth: ISBN 0-262-23181-6; Paper: ISBN 0-262-73121-5.

Dugatkin, L. A.; Hudson, K. R. 2000. Game Theory and Animal Behavior. Oxford University Press: USA.

Maynard Smith, John. (1989; 1992). "DID DARWIN GET IT RIGHT ? Essays on Games, Sex and Evolution". Smith Chapman & Hall, cloth 1989; paperback 1992; ISBN 0-412-03821-8.

Research Articles

Foster, D.; Young, P. 1990. Stochastic evolutionary game dynamics. Theoretical Population Biology 38:219-232.

Sinervo, B.; Lively, C. M. 1996. The rock-paper-scissors game and the evolution of alternative strategies. Nature 380:240-243.

Hines, W. G. 1987. Evolutionarily [corrected] stable strategies: a review of basic theory. Theoretical Population Biology 31:114.

Kokko, H. 1997. Evolutionarily stable strategies of age-dependent sexual advertisement. Behavioral Ecology and Sociobiology 41:99-107.

Education Research & Pedagogical Materials

Donovan T. and Welden C. (2002). Spreadsheet Exercises in Ecology and Evolution. Sinaver Associates: Sunderland, MA. 510-519.

Tutorial & Background materials

Bob Wyttenbach's GameBug (a very powerful Evolutionary Game Theory simulation)

John Maynard Smith Detailed biography

John Maynard Smith Obituary

General Game Theory Links

John Maynard Smith Obituary and biography overview

Kenneth Prestwich's Game Theory and Biology website and applets

Jungck John R., Spangenberg Jennifer A () ESS: Evolutionary Stable Strategies Game Theory Module. A module of the Biological ESTEEM Collection, published by the BioQUEST Curriculum Consortium. URL: