Selective phonotaxis in Neoconocephalus nebrascensis (Orthoptera: Tettigoniidae): call recognition at two temporal scales.

The calls of many Orthopteran species are comprised of a simple trill of pulses, the temporal pattern of which is important for call recognition. Male Neoconocephalus nebrascensis produce pulses with a temporal structure typical for the genus. However, they modify this pattern by grouping their pulses into verses, thereby creating a higher order temporal structure. The importance of the pulse pattern and verse structure for call recognition in N. nebrascensis was determined using a walking compensator. Females required the conspecific pulse pattern for call recognition, responding only when the intervals between pulses were short or absent. Females also required the verse structure for call recognition, and recognized the verse structure only when the amplitude modulation depth between verses and pauses exceeded 18 dB. We discuss that the verse recognition mechanism is a derived trait adapted for pre-mating isolation. We hypothesize that the unusually large amplitude modulation required for verse recognition forces males to synchronize their calls in order to preserve an attractive pattern. Call synchrony appears to be the outcome of cooperation, rather than competition, in this species.

Spectral selectivity during phonotaxis: a comparative study in Neoconocephalus (Orthoptera: Tettigoniidae).

The calls of male Neoconocephalus have most energy concentrated in a relatively narrow low-frequency band. In N. robustus this low-frequency band is centered around 7 kHz, whereas calls of N. nebrascensis and N. bivocatus have center frequencies close to 10 kHz. The importance of the position of the low-frequency band for female phonotaxis in these three species was determined using a walking compensator. Female N. robustus showed significant phonotaxis towards call frequencies from 5 to 10 kHz, and spectral selectivity towards higher frequencies did not change with stimulus amplitude. Significant responses in N. nebrascensis and N. bivocatus occurred at significantly higher frequency ranges than in N. robustus. In these species, spectral selectivity changed with stimulus amplitude; at 68 dB sound pressure level (SPL), upper cut-off frequency was significantly lower than at 80 dB SPL in both species. Adding a higher harmonic to the conspecific carrier frequency had a strong inhibitory effect on phonotaxis in N. robustus: at higher relative amplitudes of the harmonic, phonotaxis was completely suppressed. Adding a higher harmonic to the conspecific carrier frequency had a much weaker but significant inhibitory effect in N. nebrascensis and little, if any, effect in N. bivocatus. The processing of song spectrum in the sensory system is discussed with regard to the differences in spectral selectivity among the three species. The sharp spectral selectivity of N. robustus is interpreted as an adaptation for species isolation.

Recognition of calls with exceptionally fast pulse rates: female phonotaxis in the genus Neoconocephalus (Orthoptera: Tettigoniidae).

Male Neoconocephalus robustus and Neoconocephalus bivocatus produce remarkably fast calls, with pulse rates of approximately 175-200 Hz. The temporal call patterns differ significantly between the two species. Male N. robustus produce calls with a single pulse rate of 200 Hz. In N. bivocatus, pulses are repeated with alternating periods, resulting in distinct pulse pairs: approximately 175 pulses s(-1) are grouped into 87 pulse pairs s(-1). In order to identify the temporal parameters used to recognize calls with such fast pulse rates, female call recognition in both species was tested during phonotaxis on a walking compensator. Female N. robustus were attracted to calls without amplitude modulation. Amplitude-modulated signals were equally attractive, as long as the silent intervals were short enough. The maximally tolerated interval duration varied with pulse duration. Female N. bivocatus did not require the paired-pulse pattern but were attracted to call models in which each pulse pair was merged into one long pulse. Females used the pulse rate to recognize such signals: pulse rates close to 87 Hz were attractive, largely independent of the duty cycle. Thus, females of the sibling species N. robustus and N. bivocatus used qualitatively different call recognition mechanisms.