Origin of the Red Wolf: Response to Nowak and Federoff and Gardener

The discussion by Nowak and Federoff distills to seven issues pertinent to Roy et al. (1996) to which we would like to respond. We feel, however, that none of the issues they raised are consequential, and we stand by our conclusions. First, Nowak and Federoff criticize our contention that Nowak (1979) argued that the red wolf was ancestral to modern coyotes and gray wolves. Our interpretation was based on Fig. 55 in his treatise, duplicated here as Fig. 1, and the statement from their text (p. 87) that "C. rufus [the red wolf] represents a primitive line of wolves that has undergone less change than C. lupus [the gray wolf], and has thus retained more characters found in the ancestral stock from which both wolves and coyotes arose." We interpreted this to mean that the red wolf lineage diverged first from the same ancestral stock that later gave rise to coyotes and gray wolves (Fig. 1). This apparently was not what Nowak intended, and we apologize. In view of the paucity and fragmentary nature of the canid fossil record (e.g., Nowak 1979; Kurten & Anderson 1980), we believe that such evidence alone cannot adequately support any single view of Pleistocene canid phylogeny, a conclusion clearly supported by all the alternative connections in Nowak's figure (Fig. 1). We disagree strongly with the statement by Nowak and Federoff that "If the red wolf did not exist, we would have to invent it." The phylogenetic transition between coyote and gray wolf, if it occurred, could have progressed rapidly such that intermediate forms would not be preserved or even recognizable as a distinct species. The issue of whether the red wolf was ancestral to either coyote or gray wolf is a red herring; it is simply not relevant. The essential dichotomy, the focus of our paper, concerns whether the red wolf has an ancient origin or was formed recently by hybridization between gray wolves and coyotes. We state that "the evolutionary