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Wagner Networks and Ancestors. Syst. Zool. 21:398–413.—Differences between Wagner Networks and Wagner Trees, and, the general problems encountered in estimating ancestral character states are briefly reviewed. The relation between Wagner Network structure and probable ancestral state combinations is illustrated with a hypothetical case and a real case of seven species of bullhead catfishes (genus Ictalurus, subgenus Amiurus). The relation between Wagner Networks and paraphyletic classifications is also noted.
John G. Lundberg (Fri,) studied this question.