Sound production and hearing in the blue cracker butterfly Hamadryas feronia (Lepidoptera, nymphalidae) from Venezuela.

Certain species of Hamadryas butterflies are known to use sounds during interactions with conspecifics. We have observed the behaviour associated with sound production and report on the acoustic characteristics of these sounds and on the anatomy and physiology of the hearing organ in one species, Hamadryas feronia, from Venezuela. Our observations confirm previous reports that males of this species will take flight from their tree perch when they detect a passing conspecific (male or female) and, during the chase, produce clicking sounds. Our analyses of both hand-held males and those flying in the field show that the sounds are short (approximately 0.5 s) trains of intense (approximately 80-100 dB SPL at 10 cm) and brief (2-3 ms) double-component clicks, exhibiting a broad frequency spectrum with a peak energy around 13-15 kHz. Our preliminary results on the mechanism of sound production showed that males can produce clicks using only one wing, thus contradicting a previous hypothesis that it is a percussive mechanism. The organ of hearing is believed to be Vogel's organ, which is located at the base of the forewing subcostal and cubital veins. Vogel's organ consists of a thinned region of exoskeleton (the tympanum) bordered by a rigid chitinous ring; associated with its inner surface are three chordotonal sensory organs and enlarged tracheae. The largest chordotonal organ attaches to a sclerite positioned near the center of the eardrum and possesses more than 110 scolopidial units. The two smaller organs attach to the perimeter of the membrane. Extracellular recordings from the nerve branch innervating the largest chordotonal organ confirm auditory sensitivity with a threshold of 68 dB SPL at the best frequency of 1.75 kHz. Hence, the clicks with peak energy around 14 kHz are acoustically mismatched to the best frequencies of the ear. However, the clicks are broad-banded and even at 1-2 kHz, far from the peak frequency, the energy is sufficient such that the butterflies can easily hear each other at the close distances at which they interact (less than 30 cm). In H. feronia, Vogel's organ meets the anatomical and functional criteria for being recognized as a typical insect tympanal ear.