Effects of sodium puddling on male mating success, courtship and flight in a swallowtail butterfly

In many Lepidoptera species usually only males puddle for sodium. Two explanations have been offered for this: (1) neuromuscular activity: males need increased sodium for flight because they are more active flyers than females; and (2) direct benefits: sodium is a type of direct benefit provided by males to females via ejaculate during mating. Surprisingly, there is little direct experimental evidence for either of these. In this study, we examined both explanations using the pipevine swallowtail butterfly, Battus philenor L. If sodium increases neuromuscular activity, males consuming sodium should be better fliers than males without sodium. If males collect sodium for nuptial gifts that benefit their mates, males consuming sodium may have greater mating success than males without sodium. In that case, females then need an honest cue/signal of the quality of male-provided direct benefits that they can assess before mating. If sodium affects male courtship flight by increasing neuromuscular activity, how a male courts could serve as such a premating cue/signal of male benefit quality. Therefore, sodium may benefit males in terms of obtaining mates by increasing their neuromuscular activity. In this study we found that males that consumed sodium courted more vigorously and had greater mating success than males that consumed water. In addition, the courtship displays of males consuming sodium were significantly different from those of males consuming water, providing a possible honest cue/signal of male benefit quality that females can assess. Interestingly, we did not find evidence that sodium consumption affects male flight outside of courtship. That only aspects of male flight related to mating were affected by sodium, while aspects of general flight were not, is consistent with the idea that sodium may benefit males in terms of obtaining mates via effects on neuromuscular activity.