Feeding responses of adult butterflies, Nymphalis xanthomelas, Kaniska canace and Vanessa indica, to components in tree sap and rotting fruits: synergistic effects of ethanol and acetic acid on sugar responsiveness

Exuded tree sap and rotting fruits are important feeding sources for adults of many butterfly species. However, the chemical composition of such rotting foods and the effects of these constituents on butterfly feeding behaviour have rarely been investigated. Chemical analyses revealed that these food sources contain several major components, including fructose, glucose, ethanol and acetic acid. Moreover, 15 amino acids were identified from the exuded tree sap. The gustatory responsiveness to these components was examined in the proboscis of nymphalid adults of three species (Nymphalis xanthomelas, Kaniska canace and Vanessa indica). The first two species primarily feed on sap and rotting fruits, while V. indica frequently visits flowers. Fructose elicited larger feeding responses from these saprophagous butterflies than glucose, which had a feeding-stimulatory effect only on N. xanthomelas at a concentration equivalent to that of the food. Ethanol, acetic acid and the five major amino acids identified in tree sap did not elicit feeding responses by themselves. However, ethanol and/or acetic acid, when mixed with these sugars at equivalent concentrations, evoked feeding and enhanced probing responses in K. canace and V. indica. These results suggest that ethanol and acetic acid, together with sugars at low concentrations, synergistically stimulate butterfly feeding behaviour.