Anthony A. Echelle, Alice F. Echelle, Michael H. Smith, Loren G. Hill, Analysis of Genic Continuity in a Headwater Fish, Etheostoma radiosum (Percidae), Copeia, Vol. 1975, No. 2 (May 23, 1975), pp. 197-204
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Echelle et al. (1975) studied this question.