Benedict D. Chivers

Ultrasonic reverse stridulation in the spider-like katydid Arachnoscelis (Orthoptera: Listrosceledinae)

This paper illustrates the biomechanics of sound production in the neotropical predaceous katydid Arachnoscelis arachnoides (Insecta: Orthoptera: Tettigoniidae). Described and previously known from only one male specimen, this genus of predaceous katydids resembles spiders in their general body appearance. To call distant females, male katydids produce songs by stridulation where one forewing possessing a sclerotized file rubs against a row of teeth (scraper) on the other wing. In most katydid species, the songs are produced during the wing-closing phase of the stridulation. Morphological comparative studies of the stridulatory apparatus of the type specimen of Arachnoscelis arachnoides and males of other closely related species suggest that this insect sings with a frequency of ca. 80 kHz to attract conspecific females. We found an abundant population of A. arachnoides in Central Northeast of Colombia and undertook a complete analysis of the biomechanics of stridulation in this species. Using ultrasound-sensitive equipment and high-speed video, we determined that male A. arachnoides sing at ca. 74 kHz and use elastic energy and wing deformation to reach such ultrasonic frequencies. In contrast to most katydids, males of this species produce their calls during the opening phase of the wing; this form of stridulation is discussed.

Functional morphology of tegmina-based stridulation in the relict species Cyphoderris monstrosa (Orthoptera: Ensifera: Prophalangopsidae)

ABSTRACT Male grigs, bush crickets and crickets produce mating calls by tegminal stridulation: the scraping together of modified forewings functioning as sound generators. Bush crickets (Tettigoniidae) and crickets (Gryllinae) diverged some 240 million years ago, with each lineage developing unique characteristics in wing morphology and the associated mechanics of stridulation. The grigs (Prophalangopsidae), a relict lineage more closely related to bush crickets than to crickets, are believed to retain plesiomorphic features of wing morphology. The wing cells widely involved in sound production, such as the harp and mirror, are comparatively small, poorly delimited and/or partially filled with cross-veins. Such morphology is similarly observed in the earliest stridulating ensiferans, for which stridulatory mechanics remains poorly understood. The grigs, therefore, are of major importance to investigate the early evolutionary stages of tegminal stridulation, a critical innovation in the evolution of the Orthoptera. The aim of this study is to appreciate the degree of specialization on grig forewings, through identification of sound radiating areas and their properties. For well-grounded comparisons, homologies in wing venation (and associated areas) of grigs and bush crickets are re-evaluated. Then, using direct evidence, this study confirms the mirror cell, in association with two other areas (termed ‘neck’ and ‘pre-mirror’), as the acoustic resonator in the grig Cyphoderris monstrosa. Despite the use of largely symmetrical resonators, as found in field crickets, analogous features of stridulatory mechanics are observed between C. monstrosa and bush crickets. Both morphology and function in grigs represents transitional stages between unspecialized forewings and derived conditions observed in modern species. Summary: Using laser Doppler vibrometry in a relict species, evolutionary origins of sound generator features are investigated by comparison of functional morphology between distinct lineages of acoustically calling orthopterans.

Chamber music - An unusual Helmholtz resonator for song amplification in a Neotropical bush-cricket (Orthoptera, Tettigoniidae).

ABSTRACT Animals use sound for communication, with high-amplitude signals being selected for attracting mates or deterring rivals. High amplitudes are attained by employing primary resonators in sound-producing structures to amplify the signal (e.g. avian syrinx). Some species actively exploit acoustic properties of natural structures to enhance signal transmission by using these as secondary resonators (e.g. tree-hole frogs). Male bush-crickets produce sound by tegminal stridulation and often use specialised wing areas as primary resonators. Interestingly, Acanthacara acuta, a Neotropical bush-cricket, exhibits an unusual pronotal inflation, forming a chamber covering the wings. It has been suggested that such pronotal chambers enhance amplitude and tuning of the signal by constituting a (secondary) Helmholtz resonator. If true, the intact system – when stimulated sympathetically with broadband sound – should show clear resonance around the song carrier frequency which should be largely independent of pronotum material, and change when the system is destroyed. Using laser Doppler vibrometry on living and preserved specimens, microcomputed tomography, 3D-printed models and finite element modelling, we show that the pronotal chamber not only functions as a Helmholtz resonator owing to its intact morphology but also resonates at frequencies of the calling song on itself, making song production a three-resonator system. Summary: Laser vibrometry, microcomputed tomography and finite element modelling are used to show that an unusual pronotal inflation covering the wings of a bush-cricket acts as a Helmholtz resonator.

Evidence of auditory insensitivity to vocalization frequencies in two frogs

The emergence and maintenance of animal communication systems requires the co-evolution of signal and receiver. Frogs and toads rely heavily on acoustic communication for coordinating reproduction and typically have ears tuned to the dominant frequency of their vocalizations, allowing discrimination from background noise and heterospecific calls. However, we present here evidence that two anurans, Brachycephalus ephippium and B. pitanga, are insensitive to the sound of their own calls. Both species produce advertisement calls outside their hearing sensitivity range and their inner ears are partly undeveloped, which accounts for their lack of high-frequency sensitivity. If unheard by the intended receivers, calls are not beneficial to the emitter and should be selected against because of the costs associated with signal production. We suggest that protection against predators conferred by their high toxicity might help to explain why calling has not yet disappeared, and that visual communication may have replaced auditory in these colourful, diurnal frogs.

Distribution of sound pressure around a singing cricket: radiation pattern and asymmetry in the sound field

Abstract Male field crickets generate calls to attract distant females through tegminal stridulation: the rubbing together of the overlying right wing which bears a file of cuticular teeth against the underlying left wing which carries a sclerotized scraper. During stridulation, specialized areas of membrane on both wings are set into oscillating vibrations to produce acoustic radiation. The location of females is unknown to the calling males and thus increasing effective signal range in all directions will maximize transmission effectiveness. However, producing an omnidirectional sound field of high sound pressure levels may be problematic due to the mechanical asymmetry found in this sound generation system. Mechanical asymmetry occurs by the right wing coming to partially cover the left wing during the closing stroke phase of stridulation. As such, it is hypothesized that the sound field on the left-wing side of the animal will contain lower sound pressure components than on the right-wing side as a result of this coverage. This hypothesis was tested using a novel method to accurately record a high-resolution, three dimensional mapping of sound pressure levels around restrained Gryllus bimaculatus field crickets singing under pharmacological stimulation. The results indicate that a bilateral asymmetry is present across individuals, with greater amplitude components present in the right-wing side of the animal. Individual variation in sound pressure to either the right- or left-wing side is also observed. However, statistically significant differences in bilateral sound field asymmetry as presented here may not affect signalling in the field.

Non-invasive biophysical measurement of travelling waves in the insect inner ear

Frequency analysis in the mammalian cochlea depends on the propagation of frequency information in the form of a travelling wave (TW) across tonotopically arranged auditory sensilla. TWs have been directly observed in the basilar papilla of birds and the ears of bush-crickets (Insecta: Orthoptera) and have also been indirectly inferred in the hearing organs of some reptiles and frogs. Existing experimental approaches to measure TW function in tetrapods and bush-crickets are inherently invasive, compromising the fine-scale mechanics of each system. Located in the forelegs, the bush-cricket ear exhibits outer, middle and inner components; the inner ear containing tonotopically arranged auditory sensilla within a fluid-filled cavity, and externally protected by the leg cuticle. Here, we report bush-crickets with transparent ear cuticles as potential model species for direct, non-invasive measuring of TWs and tonotopy. Using laser Doppler vibrometry and spectroscopy, we show that increased transmittance of light through the ear cuticle allows for effective non-invasive measurements of TWs and frequency mapping. More transparent cuticles allow several properties of TWs to be precisely recovered and measured in vivo from intact specimens. Our approach provides an innovative, non-invasive alternative to measure the natural motion of the sensilla-bearing surface embedded in the intact inner ear fluid.