Mating Behaviour and Vibratory Signalling in Non-Hearing Cave Crickets Reflect Primitive Communication of Ensifera

In Ensifera, the lack of well-supported phylogeny and the focus on acoustic communication of the terminal taxa hinders understanding of the evolutionary history of their signalling behaviour and the related sensory structures. For Rhaphidophoridae, the most relic of ensiferans following morphology-based phylogenies, the signalling modes are still unknown. Together with a detailed description of their mating process, we provide evidence on vibratory signalling for the sympatric European species Troglophilus neglectus and T. cavicola. Despite their temporal shift in reproduction, the species' behaviours differ significantly. Signalling by abdominal vibration constitutes an obligatory part of courtship in T. neglectus, while it is absent in T. cavicola. Whole-body vibration is expressed after copulation in both species. While courtship signalling appears to stimulate females for mating, the function of post-copulation signals remains unclear. Mating and signalling of both species were found to take place in most cases on bark, and less frequently on other available substrates, like moss and rock. The signals' frequency spectra were substrate dependent, but with the dominant peak always expressed below 120 Hz. On rock, the intensity of T. neglectus courtship signals was below the species' physiological detection range, presumably constraining the evolution of such signalling in caves. The species' behavioural divergence appears to reflect their divergent mating habitats, in and outside caves. We propose that short-range tremulation signalling in courtship, such as is expressed by T. neglectus, represents the primitive mode and context of mechanical signalling in Ensifera. The absence of high-frequency components in the signals may be related to the absence of the crista acoustica homologue (CAH) in the vibratory tibial organ of Rhaphidophoridae. This indirectly supports the hypothesis proposing that the CAH, as an evolutionary precursor of the ear, evolved in Ensifera along the (more) complex vibratory communication, also associated with signals of higher carrier frequency.