Wing mechanics and acoustic communication of a new genus of sylvan katydid (Orthoptera: Tettigoniidae: Pseudophyllinae) from the Central Cordillera cloud forest of Colombia.

Stridulation is used by male katydids to produce sound via the rubbing together of their specialised forewings, either by sustained or interrupted sweeps of the file producing different tones and call structures. There are many species of Orthoptera that remain undescribed and their acoustic signals are unknown. This study aims to measure and quantify the mechanics of wing vibration, sound production and acoustic properties of the hearing system in a new genus of Pseudophyllinae with taxonomic descriptions of two new species. The calling behaviour and wing mechanics of males were measured using micro-scanning laser Doppler vibrometry, microscopy, and ultrasound sensitive equipment. The resonant properties of the acoustic pinnae of the ears were obtained via µ-CT scanning and 3D printed experimentation, and numerical modelling was used to validate the results. Analysis of sound recordings and wing vibrations revealed that the stridulatory areas of the right tegmen exhibit relatively narrow frequency responses and produce narrowband calls between 12 and 20 kHz. As in most Pseudophyllinae, only the right mirror is activated for sound production. The acoustic pinnae of all species were found to provide a broadband increased acoustic gain from ~40-120 kHz by up to 25 dB, peaking at almost 90 kHz which coincides with the echolocation frequency of sympatric bats. The new genus, named Satizabalus n. gen., is here derived as a new polytypic genus from the existing genus Gnathoclita, based on morphological and acoustic evidence from one described (S. sodalis n. comb.) and two new species (S. jorgevargasi n. sp. and S. hauca n. sp.). Unlike most Tettigoniidae, Satizabalus exhibits a particular form of sexual dimorphism whereby the heads and mandibles of the males are greatly enlarged compared to the females. We suggest that Satizabalus is related to the genus Trichotettix, also found in cloud forests in Colombia, and not to Gnathoclita.

Effects of age and noise on tympanal displacement in the Desert Locust.

Insect cuticle is an evolutionary-malleable exoskeleton that has specialised for various functions. Insects that detect the pressure component of sound bear specialised sound-capturing tympani evolved from cuticular thinning. Whilst the outer layer of insect cuticle is composed of non-living chitin, its mechanical properties change during development and aging. Here, we measured the displacements of the tympanum of the desert Locust, Schistocerca gregaria, to understand biomechanical changes as a function of age and noise-exposure. We found that the stiffness of the tympanum decreases within 12 h of noise-exposure and increases as a function of age, independent of noise-exposure. Noise-induced changes were dynamic with an increased tympanum displacement to sound within 12 h post noise-exposure. Within 24 h, however, the tone-evoked displacement of the tympanum decreased below that of control Locusts. After 48 h, the tone-evoked displacement of the tympanum was not significantly different to Locusts not exposed to noise. Tympanal displacements reduced predictably with age and repeatably noise-exposed Locusts (every three days) did not differ from their non-noise-exposed counterparts. Changes in the biomechanics of the tympanum may explain an age-dependent decrease in auditory detection in tympanal insects.

Non-invasive characterization of the elastic protein resilin in insects using Raman spectroscopy.

Resilin is an extremely efficient elastic protein found in the moving parts of insects. Despite many years of resilin research, we are still only just starting to understand its diversity, native structures, and functions. Understanding differences in resilin structure and diversity could lead to the development of bioinspired elastic polymers, with broad applications in materials science. Here, to better understand resilin structure, we offer a novel methodology for identifying resilin-rich regions of the insect cuticle using non-invasive Raman spectroscopy in a model species, the desert locust (Schistocerca gregaria). The Raman spectrum of the resilin-rich semilunar process of the hind leg was compared with that of nearby low-resilin cuticle, and reference spectra and peaks assigned for these two regions. The main peaks of resilin include two bands associated with tyrosine at 955-962 and 1141-1203 cm-1 and a strong peak at 1615 cm-1, attributed to the α-Amide I group associated with dityrosine. We also found the chitin skeletal modes at ~485-567 cm-1 to be significant contributors to spectra variance between the groups. Raman spectra were also compared to results obtained by fluorescence spectroscopy, as a control technique. Principal component analysis of these resulting spectra revealed differences in the light-scattering properties of resilin-rich and resilin-poor cuticular regions, which may relate to differences in native protein structure and relative abundance.

An Eocene insect could hear conspecific ultrasounds and bat echolocation.

Hearing has evolved independently many times in the animal kingdom and is prominent in various insects and vertebrates for conspecific communication and predator detection. Among insects, katydid (Orthoptera: Tettigoniidae) ears are unique, as they have evolved outer, middle, and inner ear components, analogous in their biophysical principles to the mammalian ear. The katydid ear consists of two paired tympana located in each foreleg. These tympana receive sound externally on the tympanum surface (usually via pinnae) or internally via an ear canal (EC). The EC functions to capture conspecific calls and low frequencies, while the pinnae passively amplify higher-frequency ultrasounds including bat echolocation. Together, these outer ear components provide enhanced hearing sensitivity across a dynamic range of over 100 kHz. However, despite a growing understanding of the biophysics and function of the katydid ear, its precise emergence and evolutionary history remains elusive. Here, using microcomputed tomography (µCT) scanning, we recovered geometries of the outer ear components and wings of an exceptionally well-preserved katydid fossilized in Baltic amber (∼44 million years [Ma]). Using numerical and theoretical modeling of the wings, we show that this species was communicating at a peak frequency of 31.62 (± 2.27) kHz, and we demonstrate that the ear was biophysically tuned to this signal and to providing hearing at higher-frequency ultrasounds (>80 kHz), likely for enhanced predator detection. The results indicate that the evolution of the unique ear of the katydid, with its broadband ultrasonic sensitivity and analogous biophysical properties to the ears of mammals, emerged in the Eocene.

The chemistry of an insect ear: ionic composition of a liquid-filled ear and haemolymphs of Neotropical katydids.

The purpose of this study is to examine and to compare the ionic composition of the haemolymph and the ear fluid of seven species of katydids (Orthoptera: Tettigoniidae) with the aim of providing from a biochemical perspective a preliminary assessment for an insect liquid contained in the auditory organ of katydids with a hearing mechanism reminiscent of that found in vertebrates. A multi-element trace analysis by inductively coupled plasma optical-emission spectrometry was run for 16 elements for the ear liquid of seven species and the haemolymph of six of them. Based on the obtained results, it can be recognized that the ionic composition is variable among the studied insects, but sodium (Na+), potassium (K+), calcium (Ca2+) and magnesium (Mg2+) are the most prominent of the dissolved inorganic cations. However, the ion concentrations between the two fluids are considerably different and the absence or low concentration of Ca2+ is a noticeable feature in the inner ear liquid. A potential relationship between the male courtship song peak frequency and the total ion (Na+, K+, Mg2+ and Ca2+) concentration of the inner ear liquid is also reported.

Beyond the exponential horn: a bush-cricket with ear canals which function as coupled resonators.

Bush-crickets have dual-input, tympanal ears located in the tibia of their forelegs. The sound will first of all reach the external sides of the tympana, before arriving at the internal sides through the bush-cricket's ear canal, the acoustic trachea (AT), with a phase lapse and pressure gain. It has been shown that for many bush-crickets, the AT has an exponential horn-shaped morphology and function, producing a significant pressure gain above a certain cut-off frequency. However, the underlying mechanism of different AT designs remains elusive. In this study, we demonstrate that the AT of the duetting Phaneropterinae bush-cricket Pterodichopetala cieloi function as coupled resonators, producing sound pressure gains at the sex-specific conspecific calling song frequency, and attenuating the remainder-a functioning mechanism significantly different from an exponential horn. Furthermore, it is demonstrated that despite the sexual dimorphism between the P. cieloi AT, both male and female AT have a similar biophysical mechanism. The analysis was carried out using an interdisciplinary approach, where micro-computed tomography was used for the morphological properties of the P. cieloi AT, and a finite-element analysis was applied on the precise tracheal geometry to further justify the experimental results and to go beyond experimental limitations.

Quantification of bush-cricket acoustic trachea mechanics using Atomic Force Microscopy nanoindentation.

Derived from the respiratory tracheae, bush-crickets' acoustic tracheae (or ear canals) are hollow tubes evolved to transmit sounds from the external environment to the interior ear. Due to the location of the ears in the forelegs, the acoustic trachea serves as a structural element that can withstand large stresses during locomotion. In this study, we report a new Atomic Force Microscopy Force Spectroscopy (AFM-FS) approach to quantify the mechanics of taenidia in the bush-cricket Mecopoda elongata. Mechanical properties were examined over the longitudinal axis of hydrated taenidia, by indenting single fibres using precision hyperbolic tips. Analysis of the force-displacement (F-d) extension curves at low strains using the Hertzian contact model showed an Elastic modulus distribution between 13.9 MPa to 26.5 GPa, with a mean of 5.2 ± 7 GPa and median 1.03 GPa. Although chitin is the primary component of stiffness, variation of elasticity in the nanoscale suggests that resilin significantly affects the mechanical properties of single taenidia fibres (38% of total data). For indentations up to 400 nm, an intricate chitin-resilin response was observed, suggesting structural optimization between compliance and rigidity. Finite-element analysis on composite materials demonstrated that the Elastic modulus is sensitive to the percentage of resilin and chitin content, their location and structural configuration. Based on our results, we propose that the distinct moduli of taenidia fibres indicate sophisticated evolution with elasticity playing a key role in optimization. STATEMENT OF SIGNIFICANCE: In crickets and bush-crickets, the foreleg tracheae have evolved into acoustic canals, which transport sound to the ears located on the tibia of each leg. Tracheae are held open by spiral cuticular micro-fibres called taenidia, which are the primary elements of mechanical reinforcement. We developed an AFM-based method to indent individual taenidia at the nanometre level, to quantify local mechanical properties of the interior acoustic canal of the bush-cricket Mecopoda elongata, a model species in hearing research. Taenidia fibres were immobilized on a hard substrate and the indenter directly approached the epicuticle surface. This is the first characterization of the nano-structure of unfixed tracheal taenidia, and should pave the way for further in vivo mechanical investigations of auditory structures.

Ear pinnae in a neotropical katydid (Orthoptera: Tettigoniidae) function as ultrasound guides for bat detection.

Early predator detection is a key component of the predator-prey arms race and has driven the evolution of multiple animal hearing systems. Katydids (Insecta) have sophisticated ears, each consisting of paired tympana on each foreleg that receive sound both externally, through the air, and internally via a narrowing ear canal running through the leg from an acoustic spiracle on the thorax. These ears are pressure-time difference receivers capable of sensitive and accurate directional hearing across a wide frequency range. Many katydid species have cuticular pinnae which form cavities around the outer tympanal surfaces, but their function is unknown. We investigated pinnal function in the katydid Copiphora gorgonensis by combining experimental biophysics and numerical modelling using 3D ear geometries. We found that the pinnae in C. gorgonensis do not assist in directional hearing for conspecific call frequencies, but instead act as ultrasound detectors. Pinnae induced large sound pressure gains (20-30 dB) that enhanced sound detection at high ultrasonic frequencies (>60 kHz), matching the echolocation range of co-occurring insectivorous gleaning bats. These findings were supported by behavioural and neural audiograms and pinnal cavity resonances from live specimens, and comparisons with the pinnal mechanics of sympatric katydid species, which together suggest that katydid pinnae primarily evolved for the enhanced detection of predatory bats.

Reviving the sound of a 150-year-old insect: The bioacoustics of Prophalangopsis obscura (Ensifera: Hagloidea).

Determining the acoustic ecology of extinct or rare species is challenging due to the inability to record their acoustic signals or hearing thresholds. Katydids and their relatives (Orthoptera: Ensifera) offer a model for inferring acoustic ecology of extinct and rare species, due to allometric parameters of their sound production organs. Here, the bioacoustics of the orthopteran Prophalangopsis obscura are investigated. This species is one of only eight remaining members of an ancient family with over 90 extinct species that dominated the acoustic landscape of the Jurassic. The species is known from only a single confirmed specimen-the 150-year-old holotype material housed at the London Natural History Museum. Using Laser-Doppler Vibrometry, 3D surface scanning microscopy, and known scaling relationships, it is shown that P. obscura produces a pure-tone song at a frequency of ~4.7 kHz. This frequency range is distinct but comparable to the calls of Jurassic relatives, suggesting a limitation of early acoustic signals in insects to sonic frequencies (<20 kHz). The acoustic ecology and importance of this species in understanding ensiferan evolution, is discussed.

Physiological changes throughout an insect ear due to age and noise - A longitudinal study.

Hearing loss is not unique to humans and is experienced by all animals in the face of wild and eclectic differences in ear morphology. Here, we exploited the high throughput and accessible tympanal ear of the desert locust, Schistocerca gregaria to rigorously quantify changes in the auditory system due to noise exposure and age. In this exploratory study, we analyzed tympanal displacements, morphology of the auditory Müller's organ and measured activity of the auditory nerve, the transduction current, and electrophysiological properties of individual auditory receptors. This work shows that hearing loss manifests as a complex disorder due to differential effects of age and noise on several processes and cell types within the ear. The "middle-aged deafness" pattern of hearing loss found in locusts mirrors that found for humans exposed to noise early in their life suggesting a fundamental interaction of the use of an auditory system (noise) and its aging.

Auditory mechanics in the grig (Cyphoderris monstrosa): tympanal travelling waves and frequency discrimination as a precursor to inner ear tonotopy.

Ensiferan orthopterans offer a key study system for acoustic communication and the process of insect hearing. Cyphoderris monstrosa (Hagloidea) belongs to a relict ensiferan family and is often used for evolutionary comparisons between bushcrickets (Tettigoniidae) and their ancestors. Understanding how this species processes sound is therefore vital to reconstructing the evolutionary history of ensiferan hearing. Previous investigations have found a mismatch in the ear of this species, whereby neurophysiological and tympanal tuning does not match the conspecific communication frequency. However, the role of the whole tympanum in signal reception remains unknown. Using laser Doppler vibrometry, we show that the tympana are tonotopic, with higher frequencies being received more distally. The tympana use two key modalities to mechanically separate sounds into two auditory receptor populations. Frequencies below approximately 8 kHz generate a basic resonant mode in the proximal end of the tympanum, whereas frequencies above approximately 8 kHz generate travelling waves in the distal region. Micro-CT imaging of the ear and the presented data suggest that this tonotopy of the tympana drive the tonotopic mechanotransduction of the crista acustica (CA). This mechanism represents a functional intermediate between simple tuned tympana and the complex tonotopy of the bushcricket CA.

A numerical approach to investigating the mechanisms behind tonotopy in the bush-cricket inner-ear.

Bush-crickets (or katydids) have sophisticated and ultrasonic ears located in the tibia of their forelegs, with a working mechanism analogous to the mammalian auditory system. Their inner-ears are endowed with an easily accessible hearing organ, the crista acustica (CA), possessing a spatial organisation that allows for different frequencies to be processed at specific graded locations within the structure. Similar to the basilar membrane in the mammalian ear, the CA contains mechanosensory receptors which are activated through the frequency dependent displacement of the CA. While this tonotopical arrangement is generally attributed to the gradual stiffness and mass changes along the hearing organ, the mechanisms behind it have not been analysed in detail. In this study, we take a numerical approach to investigate this mechanism in the Copiphora gorgonensis ear. In addition, we propose and test the effect of the different vibration transmission mechanisms on the displacement of the CA. The investigation was carried out by conducting finite-element analysis on a three-dimensional, idealised geometry of the C. gorgonensis inner-ear, which was based on precise measurements. The numerical results suggested that (i) even the mildest assumptions about stiffness and mass gradients allow for tonotopy to emerge, and (ii) the loading area and location for the transmission of the acoustic vibrations play a major role in the formation of tonotopy.

Differentiation between left and right wing stridulatory files in the field cricket Gryllus bimaculatus (Orthoptera: Gryllidae).

Male crickets produce acoustic signals by wing stridulation, attracting females for mating. A plectrum on the left forewing's (or tegmen) anal margin rapidly strikes along a serrated vein (stridulatory file, SF) on the opposite tegmen as they close, producing vibrations, ending in a tonal sound. The tooth strike rate of the plectrum across file teeth is equal to the sound frequency produced by the cricket (i.e., ∼5k teeth/s for ∼5 kHz in field crickets) and is specific to the forewing's resonant frequency. Sound is subsequently amplified using specialised wing cells. Anatomically, the forewings appear to mirror each other: both tegmina bear a SF and plectrum; however, most cricket species stridulate using right-over-left wing overlap making the stridulatory mechanism asymmetrical by default, rendering the left tegmen's SF unused. Therefore, we hypothesised structural differences between functional and unfunctional SFs. Three-dimensional mapping was used to accurately measure SF structures in Gryllus bimaculatus wings. We found that the left SF shows significantly greater variation in inter-tooth distance than the right, but less variation within the first sixty teeth (the functional part) than the right file. The left SF's slow evolutionary change over millions of years is discussed considering modern molecular phylogenies and fossil records.

A narrow ear canal reduces sound velocity to create additional acoustic inputs in a microscale insect ear.

Located in the forelegs, katydid ears are unique among arthropods in having outer, middle, and inner components, analogous to the mammalian ear. Unlike mammals, sound is received externally via two tympanic membranes in each ear and internally via a narrow ear canal (EC) derived from the respiratory tracheal system. Inside the EC, sound travels slower than in free air, causing temporal and pressure differences between external and internal inputs. The delay was suspected to arise as a consequence of the narrowing EC geometry. If true, a reduction in sound velocity should persist independently of the gas composition in the EC (e.g., air, [Formula: see text]). Integrating laser Doppler vibrometry, microcomputed tomography, and numerical analysis on precise three-dimensional geometries of each experimental animal EC, we demonstrate that the narrowing radius of the EC is the main factor reducing sound velocity. Both experimental and numerical data also show that sound velocity is reduced further when excess [Formula: see text] fills the EC. Likewise, the EC bifurcates at the tympanal level (one branch for each tympanic membrane), creating two additional narrow internal sound paths and imposing different sound velocities for each tympanic membrane. Therefore, external and internal inputs total to four sound paths for each ear (only one for the human ear). Research paths and implication of findings in avian directional hearing are discussed.

The Ander's organ: a mechanism for anti-predator ultrasound in a relict orthopteran.

The use of acoustics in predator evasion is a widely reported phenomenon amongst invertebrate taxa, but the study of ultrasonic anti-predator acoustics is often limited to the prey of bats. Here, we describe the acoustic function and morphology of a unique stridulatory structure - the Ander's organ - in the relict orthopteran Cyphoderris monstrosa (Ensifera, Hagloidea). This species is one of just eight remaining members of the family Prophalangopsidae, a group with a fossil record of over 90 extinct species widespread during the Jurassic period. We reveal that the sound produced by this organ has the characteristics of a broadband ultrasonic anti-predator defence, with a peak frequency of 58±15.5 kHz and a bandwidth of 50 kHz (at 10 dB below peak). Evidence from sexual dimorphism, knowledge on hearing capabilities and assessment of local predators, suggests that the signal likely targets ground-dwelling predators. Additionally, we reveal a previously undescribed series of cavities underneath the organ that probably function as a mechanism for ultrasound amplification. Morphological structures homologous in both appearance and anatomical location to the Ander's organ are observed to varying degrees in 4 of the 7 other extant members of this family, with the remaining 3 yet to be assessed. Therefore, we suggest that such structures may either be more widely present in this ancient family than previously assumed, or have evolved to serve a key function in the long-term survival of these few species, allowing them to outlive their extinct counterparts.

On the tympanic membrane impedance of the katydid Copiphora gorgonensis (Insecta: Orthoptera: Tettigoniidae).

Katydids (bush-crickets) are endowed with tympanal ears located in their forelegs' tibiae. The tympana are backed by an air-filled tube, the acoustic trachea, which transfers the sound stimulus from a spiracular opening on the thorax to the internal side of the tympanic membranes (TM). In katydids the sound stimulus reaches both the external and internal side of the membranes, and the tympanal vibrations are then transferred to the hearing organ crista acustica (CA) that contains the fluid-immersed mechanoreceptors. Hence the tympana are principally involved in transmitting and converting airborne sound into fluid vibrations that stimulate the auditory sensilla. Consequently, what is the transmission power to the CA? Are the TM tuned to a certain frequency? To investigate this, the surface normal acoustic impedance of the TM is calculated using finite-element analysis in the katydid Copiphora gorgonensis. From this, the reflectance and transmittance are obtained at the TM. Based on the impedance results obtained from the pressure recordings at TM and the velocity field calculations in the AT, in the frequency range 5-40 kHz, it is concluded that the tympana have considerably higher transmission around 23 kHz, corresponding to the dominant frequency of the male pure-tone calling song in this species.

The Auditory Mechanics of the Outer Ear of the Bush Cricket: A Numerical Approach.

Bush crickets have tympanal ears located in the forelegs. Their ears are elaborate, as they have outer-, middle-, and inner-ear components. The outer ear comprises an air-filled tube derived from the respiratory trachea, the acoustic trachea (AT), which transfers sound from the mesothoracic acoustic spiracle to the internal side of the ear drums in the legs. A key feature of the AT is its capacity to reduce the velocity of sound propagation and alter the acoustic driving forces of the tympanum (the ear drum), producing differences in sound pressure and time between the left and right sides, therefore aiding the directional hearing of the animal. It has been demonstrated experimentally that the tracheal sound transmission generates a gain of ∼15 dB and a propagation velocity of 255 ms-1, an approximately 25% reduction from free-field propagation. However, the mechanism responsible for this change in sound pressure level and velocity remains elusive. In this study, we investigate the mechanical processes behind the sound pressure gain in the AT by numerically modeling the tracheal acoustic behavior using the finite-element method and real three-dimensional geometries of the tracheae of the bush cricket Copiphora gorgonensis. Taking into account the thermoviscous acoustic-shell interaction on the propagation of sound, we analyze the effects of the horn-shaped domain, material properties of the tracheal wall, and the thermal processes on the change in sound pressure level in the AT. Through the numerical results obtained, it is discerned that the tracheal geometry is the main factor contributing to the observed pressure gain.

Testing the role of trait reversal in evolutionary diversification using song loss in wild crickets.

The mechanisms underlying rapid macroevolution are controversial. One largely untested hypothesis that could inform this debate is that evolutionary reversals might release variation in vestigial traits, which then facilitates subsequent diversification. We evaluated this idea by testing key predictions about vestigial traits arising from sexual trait reversal in wild field crickets. In Hawaiian Teleogryllus oceanicus, the recent genetic loss of sound-producing and -amplifying structures on male wings eliminates their acoustic signals. Silence protects these "flatwing" males from an acoustically orienting parasitoid and appears to have evolved independently more than once. Here, we report that flatwing males show enhanced variation in vestigial resonator morphology under varied genetic backgrounds. Using laser Doppler vibrometry, we found that these vestigial sound-producing wing features resonate at highly variable acoustic frequencies well outside the normal range for this species. These results satisfy two important criteria for a mechanism driving rapid evolutionary diversification: Sexual signal loss was accompanied by a release of vestigial morphological variants, and these could facilitate the rapid evolution of novel signal values. Widespread secondary trait losses have been inferred from fossil and phylogenetic evidence across numerous taxa, and our results suggest that such reversals could play a role in shaping historical patterns of diversification.

Complex wing motion during stridulation in the katydid Nastonotus foreli (Orthoptera: Tettigoniidae: Pseudophyllinae).

Male Katydids (Orthoptera: Tettigoniidae) rub together their specialised forewings to produce sound, a process known as stridulation. During wing closure, a lobe on the anal margin of the right forewing (a scraper), engages with a tooth-covered file on the left forewing. The movement of the scraper across the file produces vibrations which are amplified by a large wing cell adjacent to the scraper, the mirror. Katydids are known to stridulate with either sustained or interrupted sweeps of the file, generating resonant pure-tone (narrowband frequency) or non-resonant (broadband frequency) calls. However, some species can conserve some purity in their calls despite incorporating discrete pulses and silent intervals. This mechanism is exhibited by many Pseudophyllinae, such as Nastonotus spp., Cocconotus spp., Triencentrus spp. and Eubliastes spp. This study aims to measure and quantify the mechanics of wing stridulation in Nastonotus foreli, a Neotropical katydid that can produce, relatively narrowband calls at ≈20 kHz. It was predicted that this species will use a stridulatory mechanism involving elastic energy whereby the scraper bends and flicks along the file in periodic bursts. The calling behaviour and wing mechanics of seven males were studied using a combination of technologies (e.g. micro-scanning laser Doppler vibrometry, advanced microscopy, ultrasound-sensitive equipment and optical motion detectors) to quantify wing mechanics and structure. Analysis of recordings revealed no clear relationship between wing velocity and carrier frequency, and a pronounced distinction between wing velocity and scraper velocity during wing closure, suggesting that the scraper experiences considerable deformation. This is characteristic of the elastic scraper mechanism of stridulation. Curiously, N. foreli might have evolved to employ elastic energy to double the duration of the call, despite possessing muscles that can reach velocities high enough to produce the same frequency without the help of elastic energy.