Richard M. McCourt, Robert W. Hoshaw, Noncorrespondence of Breeding Groups, Morphology, and Monophyletic Groups in Spirogyra (Zygnemataceae: Chlorophyta) and the Application of Species Concepts, Systematic Botany, Vol. 15, No. 1 (Jan. - Mar., 1990), pp. 69-78
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McCourt et al. (1990) studied this question.
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