The mechanics of acoustic signal evolution in field crickets.

Field crickets (Family Gryllidae, Subfamily Gryllinae) typically produce tonal calls with carrier frequencies in the range 3-8 kHz. In this study, we explored the use of a finite element model (FEM) of the stridulatory apparatus of a field cricket, Gryllus bimaculatus, based on experimental measurements of resonator geometry and mechanical properties, to predict the measured call carrier frequencies of eight other field cricket species, ranging between 3 and 7 kHz. The model allowed accurate predictions of carrier frequencies for all eight species to within a few hundred hertz from morphological measurements of their resonators. We then used the model to explore the plausible evolutionary design space for field cricket call carrier frequency along the axes of resonator size and thickness, and mapped the locations of the nine experimentally measured species in this design space. Although the nine species spanned the evolutionarily conserved spectrum of carrier frequency and body size in field crickets, they were clustered in a small region of the available design space. We then explored the reasons for this apparent evolutionary constraint on field cricket carrier frequencies at both the lower and higher limit. We found that body size and sound radiation efficiency were the main constraints at the lower limits, whereas the energetics of stridulation using the clockwork mechanism may pose a constraint at higher frequencies.

A novel acoustic-vibratory multimodal duet.

The communication strategy of most crickets and bushcrickets typically consists of males broadcasting loud acoustic calling songs, while females perform phonotaxis, moving towards the source of the call. Males of the pseudophylline bushcricket species Onomarchus uninotatus produce an unusually low-pitched call, and we found that the immediate and most robust response of females to the male acoustic call was a bodily vibration, or tremulation, following each syllable of the call. We hypothesized that these bodily oscillations might send out a vibrational signal along the substrate on which the female stands, which males could use to localize her position. We quantified these vibrational signals using a laser vibrometer and found a clear phase relationship of alternation between the chirps of the male acoustic call and the female vibrational response. This system therefore constitutes a novel multimodal duet with a reliable temporal structure. We also found that males could localize the source of vibration but only if both the acoustic and vibratory components of the duet were played back. This unique multimodal duetting system may have evolved in response to higher levels of bat predation on searching bushcricket females than calling males, shifting part of the risk associated with partner localization onto the male. This is the first known example of bushcricket female tremulation in response to a long-range male acoustic signal and the first known example of a multimodal duet among animals.