Shrinking wings for ultrasonic pitch production: hyperintense ultra-short-wavelength calls in a new genus of neotropical katydids (Orthoptera: Tettigoniidae).
ABSTRACT: 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 mm(2)). 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.
Project description:Guadeloupe, the largest of the Leeward Islands, harbors three species of Pseudophyllinae (Orthoptera: Tettigoniidae) belonging to distinct tribes. This study examined the basic aspects of sound production and acousto-vibratory behavior of these species. As the songs of many Pseudophyllinae are complex and peak at high frequencies, they require high quality recordings. Wild specimens were therefore recorded ex situ. Collected specimens were used in structure-function experiments. Karukerana aguilari Bonfils (Pterophyllini) is a large species with a mirror in each tegmen and conspicuous folds over the mirror. It sings 4-6 syllables, each comprising 10-20 pulses, with several peaks in the frequency spectrum between 4 and 20 kHz. The song is among the loudest in Orthoptera (> 125 dB SPL in 10 cm distance). The folds are protective and have no function in song production. Both mirrors may work independently in sound radiation. Nesonotus reticulatus (Fabricius) (Cocconotini) produces verses from two syllables at irregular intervals. The song peaks around 20 kHz. While singing, the males often produce a tremulation signal with the abdomen at about 8-10 Hz. To our knowledge, it is the first record of simultaneous calling song and tremulation in Orthoptera. Other males reply to the tremulation with their own tremulation. Xerophyllopteryx fumosa (Brunner von Wattenwyl) (Pleminiini) is a large, bark-like species, producing a syllable of around 20 pulses. The syllables are produced with irregular rhythms (often two with shorter intervals). The song peaks around 2-3 kHz and 10 kHz. The hind wings are relatively thick and are held between the half opened tegmina during singing. Removal of the hind wings reduces song intensity by about 5 dB, especially of the low frequency component, suggesting that the hind wings have a role in amplifying the song.
Project description:Male parasitic wasps attract females with a courtship song produced by rapid wing fanning. Songs have been described for several parasitic wasp species; however, beyond association with wing fanning, the mechanism of sound generation has not been examined. We characterized the male courtship song of Cotesia congregata (Hymenoptera: Braconidae) and investigated the biomechanics of sound production.Courtship songs were recorded using high-speed videography (2,000 fps) and audio recordings. The song consists of a long duration amplitude-modulated "buzz" followed by a series of pulsatile higher amplitude "boings," each decaying into a terminal buzz followed by a short inter-boing pause while wings are stationary. Boings have higher amplitude and lower frequency than buzz components. The lower frequency of the boing sound is due to greater wing displacement. The power spectrum is a harmonic series dominated by wing repetition rate ?220 Hz, but the sound waveform indicates a higher frequency resonance ?5 kHz. Sound is not generated by the wings contacting each other, the substrate, or the abdomen. The abdomen is elevated during the first several wing cycles of the boing, but its position is unrelated to sound amplitude. Unlike most sounds generated by volume velocity, the boing is generated at the termination of the wing down stroke when displacement is maximal and wing velocity is zero. Calculation indicates a low Reynolds number of ?1000.Acoustic pressure is proportional to velocity for typical sound sources. Our finding that the boing sound was generated at maximal wing displacement coincident with cessation of wing motion indicates that it is caused by acceleration of the wing tips, consistent with a dipole source. The low Reynolds number requires a high wing flap rate for flight and predisposes wings of small insects for sound production.
Project description:Insects of the order Orthoptera are well-known for their acoustic communication. The structures used for this purpose show a high diversity which obviously relates to differences in song parameters and to the physics of sound production. Here we describe song and morphology of the sound producing organs of a tropical bush-cricket, Ectomoptera nepicauda, from East Africa. It has a very unusual calling song consisting of frequency-modulated, pure-tone sounds in the high ultrasonic range of 80 to 120 kHz and produced by extremely fast wing movements. Concerning morphology, it represents the most extreme state in the degree of left-right fore-wing differentiation found among Orthoptera: the acoustic parts of the left fore-wing consist exclusively of the stridulatory file, comparable in function to the bow of a violin, while the right wing carries only the plectrum (?=? string) and mirror (?=? soundbox).
Project description:The Luneburg lens is a spherically symmetrical gradient refractive index (GRIN) device with unique imaging properties. Its wide field-of-view (FoV) and minimal aberration have lead it to be successfully applied in microwave antennas. However, only limited realizations have been demonstrated in acoustics. Previously proposed acoustic Luneburg lenses are mostly limited to inherently two-dimensional designs at frequencies from 1?kHz to 7?kHz. In this paper, we apply a new design method for scalable and self-supporting metamaterials to demonstrate Luneburg lenses for airborne sound and ultrasonic waves. Two Luneburg lenses are fabricated: a 2.5D ultrasonic version for 40?kHz and a 3D version for 8?kHz sound. Imaging performance of the ultrasonic version is experimentally demonstrated.
Project description:Standing waves can cause measurement errors when sound-pressure level (SPL) measurements are performed in a closed ear canal, e.g., during probe-microphone system calibration for distortion-product otoacoustic emission (DPOAE) testing. Alternative calibration methods, such as forward-pressure level (FPL), minimize the influence of standing waves by calculating the forward-going sound waves separate from the reflections that cause errors. Previous research compared test performance (Burke et al., 2010) and threshold prediction (Rogers et al., 2010) using SPL and multiple FPL calibration conditions, and surprisingly found no significant improvements when using FPL relative to SPL, except at 8 kHz. The present study examined the calibration data collected by Burke et al. and Rogers et al. from 155 human subjects in order to describe the frequency location and magnitude of standing-wave pressure minima to see if these errors might explain trends in test performance. Results indicate that while individual results varied widely, pressure variability was larger around 4 kHz and smaller at 8 kHz, consistent with the dimensions of the adult ear canal. The present data suggest that standing-wave errors are not responsible for the historically poor (8 kHz) or good (4 kHz) performance of DPOAE measures at specific test frequencies.
Project description:With varied, brightly patterned wings, butterflies have been the focus of much work on the evolution and development of phenotypic novelty. However, the chemical structures of wing pigments from few butterfly species have been identified. We characterized the orange wing pigments of female Elymnias hypermnestra butterflies (Lepidoptera: Nymphalidae: Satyrinae) from two Southeast Asian populations. This species is a sexually dimorphic Batesian mimic of several model species. Females are polymorphic: in some populations, females are dark, resemble conspecific males, and mimic Euploea spp. In other populations, females differ from males and mimic orange Danaus spp. Using LC-MS/MS, we identified nine ommochrome pigments: six from a population in Chiang Mai, Thailand, and five compounds from a population in Bali, Indonesia. Two ommochromes were found in both populations, and only two of the nine compounds have been previously reported. The sexually dimorphic Thai and Balinese populations are separated spatially by monomorphic populations in peninsular Malaysia, Singapore, and Sumatra, suggesting independent evolution of mimetic female wing pigments in these disjunct populations. These results indicate that other butterfly wing pigments remain to be discovered.
Project description:Although cockroaches (Blattodea s. str.) exhibit high proportion of species with reduced wings, the underlying evolutionary forces remain unclear. Wing reduction in insects is generally considered advantageous for females and a trade-off between investment into the flying apparatus and reproduction is predicted to explain its evolution. However, what if the wing maintenance is an important issue for males' fitness? Males raise wings during the ritualized courtship which is viewed as an unavoidable movement unveiling the tergal glands for female access. We, however, propose a novel male mating success hypothesis suggesting that male wings are essential for their successful mating. We tested these two competing, but not mutually exclusive hypotheses in the cockroach Eublaberus distanti. We found no effect of female wing loss on any of the measured fecundity characteristics despite that alatectomized females histolyzed flight muscles. On the contrary, alatectomized males did not histolyze wing muscles, but experienced a markedly decreased mating success. Our findings, therefore, provide the first evidence on the crucial mechanical role of wings on male mating success. Consequently, selection for the retention of wings in males rather than for their reduction in females can explain the evolution of sexual wing dimorphism in cockroaches and other insects.
Project description:Probe-microphone measurements are a reliable method of verifying hearing-aid sound pressure level (SPL) in the ear canal for frequencies between 0.25 and 4 kHz. However, standing waves in the ear canal reduce the accuracy of these measurements above 4 kHz. Recent data suggest that speech information at frequencies up to 10 kHz may enhance speech perception, particularly for children. Incident and reflected components of a stimulus in the ear canal can be separated, allowing the use of forward (incident) pressure as a measure of stimulus level. Two experiments were conducted to determine if hearing-aid output in forward pressure provides valid estimates of in-situ sound level in the ear canal. In experiment 1, SPL measurements were obtained at the tympanic membrane and the medial end of an earmold in ten adults. While within-subject test-retest reliability was acceptable, measures near the tympanic membrane reduced the influence of standing waves for two of the ten participants. In experiment 2, forward pressure measurements were found to be unaffected by standing waves in the ear canal for frequencies up to 10 kHz. Implications for clinical assessment of amplification are discussed.
Project description:Most implantable hearing aids currently available were developed to compensate the sensorineural hearing loss by driving middle ear structures (e.g., the ossicles). These devices are successfully used in round window (RW) stimulation clinically, although this was initially not the intended use. Here, a novel microactuator, specifically designed for RW stimulation, was tested in human temporal bones to determine actuator performance and applicability.Stapes footplate response to RW stimulation was determined experimentally in human temporal bones and the obtained sound pressure output level was estimated.The actuator had a flat displacement response between 0.125 and 4?kHz, a resonance between 4 and 7?kHz, and a roll-off above. At increasing contact force, the stapes footplate displacement decreased by 5-10?dB re ?m for forces ? 2?mN. The equivalent sound pressure level between 0.125 and 4?kHz amounted to 87-97?eq?dB SPL and increased to 117?eq?dB SPL for frequencies of 4-7?kHz. The total harmonic distortion (THD) of the actuator ranged within 15-40% for static forces of 5?mN.The feasibility of an electromagnetic actuator that may be placed into the RW niche was demonstrated but requires further optimization in terms of THD and static force sensitivity.
Project description:We examined acoustic masking in a chirping katydid species of the Mecopoda elongata complex due to interference with a sympatric Mecopoda species where males produce continuous trills at high amplitudes. Frequency spectra of both calling songs range from 1 to 80 kHz; the chirper species has more energy in a narrow frequency band at 2 kHz and above 40 kHz. Behaviourally, chirper males successfully phase-locked their chirps to playbacks of conspecific chirps under masking conditions at signal-to-noise ratios (SNRs) of -8 dB. After the 2 kHz band in the chirp had been equalised to the level in the masking trill, the breakdown of phase-locked synchrony occurred at a SNR of +7 dB. The remarkable receiver performance is partially mirrored in the selective response of a first-order auditory interneuron (TN1) to conspecific chirps under these masking conditions. However, the selective response is only maintained for a stimulus including the 2 kHz component, although this frequency band has no influence on the unmasked TN1 response. Remarkably, the addition of masking noise at 65 dB sound pressure level (SPL) to threshold response levels of TN1 for pure tones of 2 kHz enhanced the sensitivity of the response by 10 dB. Thus, the spectral dissimilarity between masker and signal at a rather low frequency appears to be of crucial importance for the ability of the chirping species to communicate under strong masking by the trilling species. We discuss the possible properties underlying the cellular/synaptic mechanisms of the 'novelty detector'.