The early evolution of cooperation in humans. On cheating, group identity and group size

in Behaviour
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The evolution of cooperation is difficult to understand, because cheaters — individuals who profit without cooperating themselves — have a benefit in interaction with cooperators. Cooperation among humans is even more difficult to understand, because cooperation occurs in large groups, making cheating a bigger threat. Restricting cooperation to members of one’s own group based on some tag-based recognition of non-group members (allorecognition) has been shown to stabilise cooperation. We address how spatial structure and group size affect the opportunities for cheating such tag-based cooperation in a spatially explicit simulation. We show that increased group diversity, under conditions of limited dispersal, reduces the selective opportunities for cheaters. A small number can already be sufficient to keep cheating at a low frequency. We discuss how marginal additional benefits of increased group size, above the benefits of local cooperation, can provide the selective pressure to reduce the number of group identities and discuss possible examples.

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Figures
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    Simulations with the basic parameter setting as specified: f, cooperation fitness advantage parameter; c, cooperation fitness cost; μ=μg=μs, mutation rates; D, local dispersal distance; g, proportion of globally dispersed offspring; N, maximum number of clans. (A) The dynamics of clan and strategy distributions. Grey bands indicate proportion of cooperators, black bands above grey ones the proportion of cheaters of the same clan as the cooperators below. (B) The spatial pattern of cooperation and cheating in the 10 000th generation. Shades of green indiacte cooperators of different clans, shades of red cheaters of different clans. (C) The average number of clans (Avg_Groups) and the average cumulated proportion of cheaters (Avg_Cheats) of the last 1000 generations in 10 replicate runs of the simulation program, replicates differing only in random number seeds. Vertical and horizontal bars indicate the ranges of change in the corresponding variable during the last 1000 generations. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1568539x.

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    Parameter settings of the simulations performed. Oval boxes, the basic parameter set; rectangular boxes, alternative values of the parameter in the corresponding oval box. Parameters have been changed one at a time, except for the two mutation rates which have been changed both simultaneously and separately.

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    The role of the maximum number of clan identities for the average cumulated proportion of cheaters (Avg_Cheats) during the last 1000 generations of the simulations. Vertical and horizontal bars indicate the ranges of change in the corresponding variable during the last 1000 generations.

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    The role of the fitness cost of cooperation for the average cumulated proportion of cheaters (Avg_Cheats) during the last 1000 generations of the simulations. Vertical and horizontal bars indicate the ranges of change in the corresponding variable during the last 1000 generations.

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    The role of mutation rates for the average cumulated proportion of cheaters (Avg_Cheats) during the last 1000 generations of the simulations. (A) The role of the mutation rate to new clans; (B) the role of the rate of strategy mutations. Vertical and horizontal bars indicate the ranges of change in the corresponding variable during the last 1000 generations.

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    The role of dispersal for the average cumulated proportion of cheaters (Avg_Cheats) during the last 1000 generations of the simulations. (A) The role of global dispersal (g); (B) the role of local dispersal (D). Vertical and horizontal bars indicate the ranges of change in the corresponding variable during the last 1000 generations.

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