The leopard frog, Rana pipiens, exists as a polymorphic species in an area of about 100,000 square miles in Minnesota, western Wisconsin, northern Iowa, and northeastern South Dakota (Merrell, 1965). In addition to the usual spotted wild type, the populations frequently contain about 1 to 10% of the unspotted burnsi type, shown by Moore (1942) to be due to a single dominant gene. The wide distribution and fairly high frequency of the burnsi gene seem to rule out the possibility that it is being maintained by recurrent mutation. Furthermore, heterozygote advantage, the mechanism most commonly invoked to explain balanced polymorphism, does not seem an adequate explanation in this case. First, the frequency of the heterozygotes in natural populations is lower than would reasonably be expected if heterosis were involved. Also, the results from crosses in the laboratory between wild-type and burnsi individuals showed no evidence of heterozygote advantage (Merrell-unpublished). Crosses between heterozygous burnsi (B/+) and pipiens (+/+) yielded the expected one-to-one burnsi to pipiens ratio, even under crowded conditions. Crosses between heterozygous burnsi frogs gave the expected 3: 1 ratios with no indication that the homozygous burnsi type (B/B) was less vigorous than the B/+ or +/+ genotypes. Finally, crosses between burnsi and kandiyohi heterozygotes gave the four genotypes, burnsi, kandiyohi, double mutant burnsi-kandiyohi, and wild type pipiens, in the expected equal numbers. (Kandiyohi is another dominant polymorphic gene in R. pipiens not linked to burnsi-Volpe 1955, 1956, 1960; Anderson and Volpe, 1958). Not only were the expected ratios obtained, but there were no significant differences in rate of development or size at metamorphosis among the different types of progeny. Therefore, the available evidence gives no support to heterozygote advantage as an explanation for the burnsi polymorphism. For this reason, other mechanisms for maintaining the burnsi polymorphism must be considered. One possibility is a seasonal shift in selection pressure (Merrell, 1965). The data presented below suggest that the burnsi frogs are somewhat better able to survive the winter than the pipiens type.
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