Cooperation in organisms, whether bacteria or primates, has been a difficulty for evolutionary theory since Darwin. On the assumption that interactions between pairs of individuals occur on a probabilistic basis, a model is developed based on the concept of an evolutionarily stable strategy in the context of the Prisoner's Dilemma game. Deductions from the model, and the results of a computer tournament show how cooperation based on reciprocity can get started in an asocial world, can thrive while interacting with a wide range of other strategies, and can resist invasion once fully established. Potential applications include specific aspects of territoriality, mating, and disease.

/pdf/the-evolution-of-cooperation-pwc4krac7e.pdf

The Evolution of Cooperation

Cooperation is needed for evolution to construct new levels of organization. Genomes, cells, multicellular organisms, social insects, and human society are all based on cooperation. Cooperation means that selfish replicators forgo some of their reproductive potential to help one another. But natural selection implies competition and therefore opposes cooperation unless a specific mechanism is at work. Here I discuss five mechanisms for the evolution of cooperation: kin selection, direct reciprocity, indirect reciprocity, network reciprocity, and group selection. For each mechanism, a simple rule is derived that specifies whether natural selection can lead to cooperation.

/pdf/five-rules-for-the-evolution-of-cooperation-2bpocasc6i.pdf

Five Rules for the Evolution of Cooperation

http://osenberglab.ecology.uga.edu/wp-content/uploads/2015/09/Charnov-1976-TPB.pdf

Optimal foraging, the marginal value theorem.

Every form of behavior is shaped by trial and error. Such stepwise adaptation can occur through individual learning or through natural selection, the basis of evolution. Since the work of Maynard Smith and others, it has been realized how game theory can model this process. Evolutionary game theory replaces the static solutions of classical game theory by a dynamical approach centered not on the concept of rational players but on the population dynamics of behavioral programs. In this book the authors investigate the nonlinear dynamics of the self-regulation of social and economic behavior, and of the closely related interactions among species in ecological communities. Replicator equations describe how successful strategies spread and thereby create new conditions that can alter the basis of their success, i.e., to enable us to understand the strategic and genetic foundations of the endless chronicle of invasions and extinctions that punctuate evolution. In short, evolutionary game theory describes when to escalate a conflict, how to elicit cooperation, why to expect a balance of the sexes, and how to understand natural selection in mathematical terms.

 Comprehensive treatment of ecological and game theoretic dynamics
 Invasion dynamics and permanence as key concepts
 Explanation in terms of games of things like competition between species

/pdf/evolutionary-games-and-population-dynamics-pax1uk4ywa.pdf

Evolutionary games and population dynamics

Despite the ubiquity with which patterns of sperm utilization have been studied, the mechanisms underlying fertilization in insects are far from clear. One well‐studied system is the yellow dung fly, in which the last male's ejaculate is thought to displace rival sperm from the female's sperm stores. Here we follow the movement of the copulating male's ejaculate through the female's reproductive tract using males labeled with different radioisotopes. We find that males ejaculate into the bursa copulatrix and that male‐1 sperm are displaced from the spermathecae during copulation. The increase in male‐2 ejaculate in the spermathecae matches the pattern of male‐2 fertilization gain, indicating that only spermathecal sperm are utilized at fertilization. Previously we have analyzed this system with a direct model of sperm displacement in which the male displaces rival sperm from the spermathecae. The data, and morphology of the female, clearly preclude such a mechanism. Here we contrast this model w...

https://www.jstor.org/stable/10.1086/303171

Sperm Displacement in the Yellow Dung Fly, Scatophaga stercoraria: An Investigation of Male and Female Processes

The relationship between continuous input and interference models of ideal free distributions with unequal competitors

MALE mate choice is predicted to occur when males transfer nutrients to females during copulation, provide paternal care, or are otherwise limited in mating capacity1–7. Most commonly, field studies have revealed male choice to be focused on females offering high fecundity8–10. Here we describe male mating selectivity that appears to be driven by sperm competition: male thirteen-lined ground squirrels (Spermophilus tridecemlineatus) avoid copulating with previously mated females once the fertilization gain rate drops below the average obtainable from alternative matings. Although this behaviour seems advantageous to males, it limits the duration of female sexual activity.

Geoff A. Parker

Papers

Sperm Displacement in the Yellow Dung Fly, Scatophaga stercoraria: An Investigation of Male and Female Processes

The relationship between continuous input and interference models of ideal free distributions with unequal competitors

Male mate choice as predicted by sperm competition in thirteen-lined ground squirrels

Arms races in evolution—An ESS to the opponent-independent costs game

Seasonal incidence: Adaptive variation in the timing of life history stages