Localization of heart poisons in the monarch butterfly

The cardiac glycosides that monarch butterflies sequester from milkweed plants during the larval stage differ remarkably in their emetic potency and are concentrated to different degrees in the various parts of the body as well as in the two sexes (Fig. 1). The very high concentrations of these compounds in the wings probably facilitate learned taste rejection in predators and account for the relatively high frequency of Danaid butterflies with beak-marked wings in natural populations. The cardiac glycosides in the abdomen have a much higher emetic potency than those in the rest of the body. Consequently, naive, extremely hungry, or forgetful birds which capture and peck off the wings but eat the abdomen discard the least emetic glycosides and ingest the most emetic, and thus again experience emesis. The nonrandom distribution of cardenolides in the wings, abdomen, and thorax, together with the fact that monarch males not only contain lower concentrations of cardiac glycosides than females but also contain cardenolides that are overall less emetic than those in females, is interpreted as evidence that these poisons are incorporated at a physiological cost. This cost, balanced against the benefits of protection from predation, provides a selective basis for the occurrence of both emetic and nonemetic individuals in natural populations. Since birds can discriminate emetic from nonemetic monarchs on the basis of taste, it is not necessary to invoke theories of kind of group selection to explain the evolution of this kind of unpalatability.