In this paper two models are described which incorporate both temporal and spatial heterogeneity. Both are compared to the spatial model of Levene and the temporal model of Haldane and Jayakar. It was found that both models can allow more variability with respect to fitness values, although one is not necessarily better than the other in this regard. In addition, neither marginal, geometric mean, nor harmonic mean overdominance is necessary for the maintenance of a protected polymorphism, as was described in the discussion.
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Evelyn P. Ewing (1979) studied this question.
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