Summary
Vibrational communication in hook-tip moth caterpillars is thought to be widely used and highly variable across species, but this phenomenon has been experimentally examined in only two species to date. The purpose of this study is to characterize and describe the function of vibrational signaling in a species, Oreta rosea Walker 1855 (Lepidoptera: Drepanidae), that differs morphologically from previously studied species. Caterpillars of this species produce three distinct types of vibrational signals during territorial encounters with conspecifics -mandible drumming, mandible scraping and lateral tremulation. Signals were recorded using a laser-doppler vibrometer and characterized based on temporal and spectral components. Behavioural encounters between a leaf resident and a conspecific intruder were staged to test the hypothesis that signaling functions as a territorial display. Drumming and scraping signals both involve the use of the mandibles, being hit vertically on, or scraped laterally across, the leaf surface. Lateral tremulation involves quick, short, successive lateral movements of the anterior body region that vibrates the entire leaf. Encounters result in residents signaling, with the highest rates observed when intruders make contact with the resident. Residents signal significantly more than intruders and most conflicts are resolved within 10 minutes, with residents winning 91% of trials. The results support the hypothesis that vibrational signals function to advertise leaf occupancy. Signaling is compared between species, and evolutionary origins of vibrational communication in caterpillars are discussed.
Methodology
Animals
Oreta rosea Walker 1855 (Lepidoptera: Drepanidae) moths were collected from the wild at ultraviolet collecting lights between May and August 2007 in Dunrobin, Ontario, Canada. Females oviposited on cuttings of viburnum (Viburnum lentago) and larvae were reared indoors on V. lentago or V. opulus under a LD 18:6 photoperiod at 21-26°C. Early- (first and second) and late- (third to fifth) instar larvae were used for life-history and behavioural observations. Late-instars were further used for morphological analysis of sound-producing structures, laser vibrometry recordings and behavioural trials.
General behaviour and life-history
Behavioural observations relevant to communication and spacing were recorded daily. These included the position on the leaf, presence of silk on the leaf, mode of feeding, and interactions between individuals. Photographs of eggs, larvae and adults were obtained with an Olympus dissection microscope (SZX12; www.olympus.com) equipped with a Zeiss camera (AxioCam MRc5;www.zeiss.com), or with a Nikon Digital SLR camera (D80; www.nikon.com).
Signal characteristics
Vibrational signals were monitored and characterized using two recording methods - a microphone and laser-doppler vibrometer (LDV). Both methods involved recording late-instar larvae with a videocamera and a microphone or LDV during encounters with conspecific intruders (see below). Vibrations measured using a Polytec LDV (PDV 100; www.polytec.com) were digitized and recorded onto a Marantz Professional portable solid state recorder (PMD 671; www.marantzpro.com; 44.1 kHz sampling rate). Vibrations perpendicular to the leaf surface were measured at the location of a circular piece of reflective tape (2.0 mm in diameter) positioned 1 - 2.5 cm from the resident caterpillar. All recordings were made in an acoustic chamber (Eckel Industries, www.eckelacoustic.com). These recordings were used to determine the types of signals produced and to measure temporal and spectral characteristics of signaling. Temporal characteristics, including mean signaling bout duration, mean interval duration between signaling bouts and number of signals per bout were measured using Raven Bioacoustics Research Program (Cornell Laboratory of Ornithology; www.birds.cornell.edu/brp/). Bouts were defined as any combination of signals that was preceded and followed by feeding, walking or at least 1 s of inactivity. Durations of each Scott et al. signal type were calculated from 20 individuals. Power spectra were made using a 512-point Fourier transform (DFT, Hann window) in Raven Bioacoustics Research Program. Signals were not filtered and a power spectrum of background noise was included for comparison.