Mechanisms of high-frequency song generation in brachypterous crickets and the role of ghost frequencies

Sound production in crickets relies on stridulation, the well-understood rubbing together of a pair of specialised wings. As the file of one wing slides over the scraper of the other, a series of rhythmic impacts cause harmonic oscillations, usually resulting in the radiation of pure tones delivered at low frequencies (2-8 kHz). In the short winged crickets of the Lebinthini tribe, acoustic communication relies on signals with remarkably high frequencies (> 8 kHz) and rich harmonic content. Using several species of the subfamily Eneopterinae, we characterise the morphological and mechanical specialisations supporting the production of high frequencies, and demonstrate that higher harmonics are exploited as dominant frequencies. These specialisations affect the structure of the stridulatory file, the motor control of stridulation and the resonance of the sound radiator. We place these specialisations in a phylogenetic framework and show that they serve to exploit high frequency vibrational modes pre-existing in the phylogenetic ancestor. In Eneopterinae, the lower frequency components are harmonically related to the dominant peak, suggesting they are relicts of ancestral carrier frequencies. Yet, such ghost frequencies still occur in the wings' free resonances, highlighting the fundamental mechanical constraints of sound radiation. These results support the hypothesis that such high frequency songs evolved stepwise, by a form of punctuated evolution which could be related to functional constraints, rather than by the progressive increase of the ancestral fundamental frequency

Biomechanics of hearing in katydids

Animals have evolved a vast diversity of mechanisms to detect sounds. Auditory organs are used to detect intraspecific communicative signals and environmental sounds relevant to survival. To hear, terrestrial animals must convert the acoustic energy contained in the airborne sound pressure waves into neural signals. In mammals, spectral quality is assessed by the decomposition of incoming sound waves into elementary frequency components using a sophisticated cochlear system. Some neotropical insects like katydids (bushcrickets) have evolved biophysical mechanisms for auditory processing that are remarkably equivalent to those of mammals. Located on their front legs, katydid ears are small, yet are capable of performing several of the tasks usually associated with mammalian hearing. These tasks include air-to-liquid impedance conversion, signal amplification, and frequency analysis. Impedance conversion is achieved by a lever system, a mechanism functionally analogous to the mammalian middle ear ossicles, yet morphologically distinct. In katydids, the exact mechanisms supporting frequency analysis seem diverse, yet are seen to result in dispersive wave propagation phenomenologically similar to that of cochlear systems. Phylogenetically unrelated, katydids and tetrapods have evolved remarkably different structural solutions to common biophysical problems. Here, we discuss the biophysics of hearing in katydids and the variations observed across different species

Wing mechanics, vibrational and acoustic communication in a new bush-cricket species of the genus Copiphora (Orthoptera: Tettigoniidae) from Colombia

Male bush-crickets produce acoustic signals by wing stridulation to call females. Several species also alternate vibratory signals with acoustic calls for intraspecific communication, a way to reduce risk of detection by eavesdropping predators. Both modes of communication have been documented mostly in neotropical species, for example in the genus Copiphora. In this article, we studied vibratory and acoustic signals and the biophysics of wing resonance in C. vigorosa, a new species from the rainforest of Colombia. Different from other Copiphora species in which the acoustic signals have been properly documented as pure tones, C. vigorosa males produce a complex modulated broadband call peaking at ca. 30 kHz. Such a broadband spectrum results from several wing resonances activated simultaneously during stridulation. Since males of this species do rarely sing, we also report that substratum vibrations have been adopted in this species as a persistent communication channel. Wing resonances and substratum vibrations were measured using a μ-scanning Laser Doppler Vibrometry. We found that the stridulatory areas of both wings exhibit a relatively broad-frequency response and the combined vibration outputs fits with the calling song spectrum breadth. Under laboratory conditions the calling song duty cycle is very low and males spend more time tremulating than singing

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

Male grigs, bush-crickets and field crickets produce mating calls by tegminal stridulation: the scraping together of modified forewings functioning as sound generators. Bush- (Tettigoniidae) and field-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 field-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 specialisation on grig forewings, through identification of sound radiating area 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 unspecialised forewings and derived conditions observed in modern species

Sound Analysis and Synthesis with R

Book Revie

Community detection in complex networks by dynamical simplex evolution

We benchmark the dynamical simplex evolution (DSE) method with several of the currently available algorithms to detect communities in complex networks by comparing correctly identified nodes for different levels of fuzziness of random networks composed of well-defined communities. The potential benefits of the DSE method to detect hierarchical substructures in complex networks are discussed

Structural biomechanics determine spectral purity of bush-cricket calls

Bush-crickets (Orthoptera: Tettigoniidae) generate sound using tegminal stridulation. Signalling effectiveness is affected by the widely varying acoustic parameters of temporal pattern, frequency and spectral purity (tonality). During stridulation, frequency multiplication occurs as a scraper on one wing scrapes across a file of sclerotized teeth on the other. The frequency with which these tooth–scraper interactions occur, along with radiating wing cell resonant properties, dictates both frequency and tonality in the call. Bush-cricket species produce calls ranging from resonant, tonal calls through to non-resonant, broadband signals. The differences are believed to result from differences in file tooth arrangement and wing radiators, but a systematic test of the structural causes of broadband or tonal calls is lacking. Using phylogenetically controlled structural equation models, we show that parameters of file tooth density and file length are the best-fitting predictors of tonality across 40 bush-cricket species. Features of file morphology constrain the production of spectrally pure signals, but systematic distribution of teeth alone does not explain pure-tone sound production in this family

Endless forms most hidden: katydids that masquerade as moss

In the cloud forests of the central range of the Colombian Andes, we discovered a species of katydid (Orthoptera: Tettigoniidae) that imitates mosses to an uncanny degree and is exceedingly difficult to detect. The camouflage exhibited by this particular katydid seems quite specific. We discuss the evolutionary consequences of this sort of specialization. Selection to maintain effective disguises can result in reproductive isolation between populations specialized for different microhabitats, which makes it reasonable to speculate that camouflage may increasing diversification rates. Camouflage could also come at the price of elevated extinction risk. This possibility must be considered because although antipredator defenses are often thought of as leading to “escape-and-radiate” dynamics where diversification follows innovation that allows expansion into new niches, recent work has shown unexpected extinction risk associated with some antipredator adaptations. Highly specialized camouflage would seem an ambiguous case because of its obvious benefits, but also potential costs such as inhabiting habitats with low carrying capacities, vulnerability to predators at high densities if predators form search images, or metabolic trade-offs with thermoregulation. Groups such as the Tettigoniidae provide a tantalizing opportunity for their exceptional diversity, wide geographic distribution, and striking array of disguises suggest that many independent evolutionary experiments have already taken place

Lack of correlation between vertical distribution and carrier frequency, and preference for open spaces in arboreal katydids that use extreme ultrasound, in Gorgona, Colombia (Orthoptera: Tettigoniidae)

Male Tettigoniidae emit sound to attract conspecific females. The sound is produced by stridulation. During stridulation the forewings open and close, but it is during the closing stroke that the scraper contacts the file teeth to generate the predominant sound components, which are amplified by adjacent wing cells specialized in sound radiation. The sounds usually exceed the sonic boundary and might occur above 40 kHz, reaching extreme ultrasonic frequencies of 150kHz in some species. Here we test the hypothesis that Tettigoniidae species should prefer microhabitats that favour efficient signal transmission, i.e. that there is a relationship of sound frequency with the vertical distribution of the species (from ground to canopy) at Gorgona National Natural Park, Colombia. We sampled 16 trees and four different altitudinal levels between 1 and 20m above the understory vegetation. We placed collecting blankets separated by vertical distances of 5m, and knocked insects down using the technique known as fogging. We found no correlation between vertical distribution and carrier frequency, but there was a preference for open spaces (below the canopy and above the understory) in species using extreme ultrasound. This is the first quantitative description of the vertical distribution in neotropical species of the family Tettigoniidae and its relationship to the calling song frequency

Shrinking wings for ultrasonic pitch production: hyperintense ultra-short-wavelength calls in a new genus of neotropical katydids (Orthoptera: tettigoniidae)

This article reports the discovery of a new genus and three species of predaceous katydid (Insecta: Orthoptera) from Colombia and Ecuador in which males produce the highest frequency ultrasonic calling songs so far recorded from an arthropod. Male katydids sing by rubbing their wings together to attract distant females. Their song frequencies usually range from audio (5 kHz) to low ultrasonic (30 kHz). However, males of Supersonus spp. call females at 115 kHz, 125 kHz, and 150 kHz. Exceeding the human hearing range (50 Hz–20 kHz) by an order of magnitude, these insects also emit their ultrasound at unusually elevated sound pressure levels (SPL). In all three species these calls exceed 110 dB SPL rms re 20 µPa (at 15 cm). Males of Supersonus spp. have unusually reduced forewings (<0.5 mm2). Only the right wing radiates appreciable sound, the left bears the file and does not show a particular resonance. In contrast to most katydids, males of Supersonus spp. position and move their wings during sound production so that the concave aspect of the right wing, underlain by the insect dorsum, forms a contained cavity with sharp resonance. The observed high SPL at extreme carrier frequencies can be explained by wing anatomy, a resonant cavity with a membrane, and cuticle deformation