A. M. Jarosz, J. J. Burdon, Host-Pathogen Interactions in Natural Populations of Linum marginale and Melampsora lini: II. Local and Regional Variation in Patterns of Resistance and Racial Structure, Evolution, Vol. 45, No. 7 (Nov., 1991), pp. 1618-1627
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Jarosz et al. (1991) studied this question.