High acoustic diversity and behavioral complexity of katydids in the Mesozoic soundscape.

Acoustic communication has played a key role in the evolution of a wide variety of vertebrates and insects. However, the reconstruction of ancient acoustic signals is challenging due to the extreme rarity of fossilized organs. Here, we report the earliest tympanal ears and sound-producing system (stridulatory apparatus) found in exceptionally preserved Mesozoic katydids. We present a database of the stridulatory apparatus and wing morphology of Mesozoic katydids and further calculate their probable singing frequencies and analyze the evolution of their acoustic communication. Our suite of analyses demonstrates that katydids evolved complex acoustic communication including mating signals, intermale communication, and directional hearing, at least by the Middle Jurassic. Additionally, katydids evolved a high diversity of singing frequencies including high-frequency musical calls, accompanied by acoustic niche partitioning at least by the Late Triassic, suggesting that acoustic communication might have been an important driver in the early radiation of these insects. The Early-Middle Jurassic katydid transition from Haglidae- to Prophalangopsidae-dominated faunas coincided with the diversification of derived mammalian clades and improvement of hearing in early mammals, supporting the hypothesis of the acoustic coevolution of mammals and katydids. Our findings not only highlight the ecological significance of insects in the Mesozoic soundscape but also contribute to our understanding of how acoustic communication has influenced animal evolution.

Tympanal bone fracture in forensic practice.

A tympanal bone fracture is an uncommon complication of head trauma and is mostly associated with a mandibular or petrous bone fracture. Upon reviewing the medicolegal literature, we could not find any publications on this topic. Tympanal bone fracture may lead to chronic complications (including external auditory canal stenosis and conductive hearing loss), with an important impact in both the medical and judiciary fields (e.g., chronic disabilities with loss of income). We decided to investigate the prevalence and mechanisms of tympanal bone fractures by means of a retrospective observational study on living victims who underwent head computed tomography after blunt head trauma and clinical forensic investigation at our center. We selected 159 cases of living victims with blunt head trauma (following an assault, traffic accident, or work accident) between January 2016 and December 2020. Re-examination of head imaging revealed 12 cases of tympanal bone fracture. Seven individuals showed cranial fractures involving the petrous bone (on the same side as the tympanal bone fracture). Three individuals had a temporomandibular fracture after a fall with chin impact. Only two victims exhibited an isolated tympanal bone fracture.

Evolutionary diversification of the auditory organ sensilla in Neoconocephalus katydids (Orthoptera: Tettigoniidae) correlates with acoustic signal diversification over phylogenetic relatedness and life history.

Neoconocephalus Tettigoniidae are a model for the evolution of acoustic signals as male calls have diversified in temporal structure during the radiation of the genus. The call divergence and phylogeny in Neoconocephalus are established, but in tettigoniids in general, accompanying evolutionary changes in hearing organs are not studied. We investigated anatomical changes of the tympanal hearing organs during the evolutionary radiation and divergence of intraspecific acoustic signals. We compared the neuroanatomy of auditory sensilla (crista acustica) from nine Neoconocephalus species for the number of auditory sensilla and the crista acustica length. These parameters were correlated with differences in temporal call features, body size, life histories and different phylogenetic positions. By this, adaptive responses to shifting frequencies of male calls and changes in their temporal patterns can be evaluated against phylogenetic constraints and allometry. All species showed well-developed auditory sensilla, on average 32-35 between species. Crista acustica length and sensillum numbers correlated with body size, but not with phylogenetic position or life history. Statistically significant correlations existed also with specific call patterns: a higher number of auditory sensilla occurred in species with continuous calls or slow pulse rates, and a longer crista acustica occurred in species with double pulses or slow pulse rates. The auditory sensilla show significant differences between species despite their recent radiation, and morphological and ecological similarities. This indicates the responses to natural and sexual selection, including divergence of temporal and spectral signal properties. Phylogenetic constraints are unlikely to limit these changes of the auditory systems.

Sensory evolution of hearing in tettigoniids with differing communication systems.

In Tettigoniidae (Orthoptera: Ensifera), hearing organs are essential in mate detection. Male tettigoniids usually produce calling songs by tegminal stridulation, whereas females approach the males phonotactically. This unidirectional communication system is the most common one among tettigoniids. In several tettigoniid lineages, females have evolved acoustic replies to the male calling song which constitutes a bidirectional communication system. The genus Poecilimon (Tettigoniidae: Phaneropterinae) is of special interest because the ancestral state of bidirectional communication, with calling males and responding females, has been reversed repeatedly to unidirectional communication. Acoustic communication is mediated by hearing organs that are adapted to the conspecific signals. Therefore, we analyse the auditory system in the Tettigoniidae genus Poecilimon for functional adaptations in three characteristics: (i) dimension of sound-receiving structures (tympanum and acoustic spiracle), (ii) number of auditory sensilla and (iii) hearing sensitivity. Profound differences in the auditory system correlate with uni- or bidirectional communication. Among the sound-receiving structures, the tympana scale with body size, whereas the acoustic spiracle, the major sound input structure, was drastically reduced in unidirectional communicating species. In the unidirectional P. ampliatus group, auditory sensilla are severely reduced in numbers, but not in the unidirectional P. propinquus group. Within the P. ampliatus group, the number of auditory sensilla is further reduced in P. intermedius which lost acoustic signalling due to parthenogenesis. The auditory sensitivity correlated with the size of the acoustic spiracle, as hearing sensitivity was better with larger spiracles, especially in the ultrasonic range. Our results show a significant reduction in auditory structures, shaped by the differing sex roles during mate detection.

Moths in space: The below-canopy structure of forest drives abundance and mobility-related traits of moth communities.

The distribution of species is primarily driven by the availability of trophic resources. In a given forest type, insects trophically related to the dominant tree are expected to be evenly distributed due to the abundance of their foodplant. However, their distribution is also influenced by complex relationships with abiotic and biotic parameters such as available space, predatory pressure, and morphometric traits. In this study, we investigated how the three-dimensional structure of space below the canopy may affect the composition of nocturnal lepidoptera communities. To synthesise the complexity of the dispersal behaviour of these insects, we evaluated easily measurable traits such as wingspan and the presence of tympanic organs, both connected to their mobility and thus potentially influenced by the structure of the available flight space. The study was conducted in the Sila National Park (Italy), where 12 sampling sites were selected in pine forests and an additional 12 in beech forests. Forest spatial structure was investigated using a portable terrestrial laser scanner. Moths were sampled monthly using light traps from May to October in both 2019 and 2020. Among measured forest traits, we observed that the space above three meters from the ground is the only factor influencing community composition. Larger species with tympanic organs prefer environments with less space below tree canopies. Our findings could be the starting point for future studies that investigate a potential defence strategy of moths against bats, as tympanate and larger species not only actively avoid chiropter predation but could also choose denser forests because of a lower bat activity. Moths' distribution and community composition thus appear to be significantly shaped by the spatial structure of forests.

Extremely high frequency sensitivity in a 'simple' ear.

An evolutionary war is being played out between the bat, which uses ultrasonic calls to locate insect prey, and the moth, which uses microscale ears to listen for the approaching bat. While the highest known frequency of bat echolocation calls is 212 kHz, the upper limit of moth hearing is considered much lower. Here, we show that the greater wax moth, Galleria mellonella, is capable of hearing ultrasonic frequencies approaching 300 kHz; the highest frequency sensitivity of any animal. With auditory frequency sensitivity that is unprecedented in the animal kingdom, the greater wax moth is ready and armed for any echolocation call adaptations made by the bat in the on-going bat-moth evolutionary war.

Bimodality of auditory receptors in bush-crickets. Сontinued discussion. It's time to experiment.

Continuation of the discussion on the sensitivity of the chordotonal sensilla of the tympanal organ of bush-crickets to vibratory stimuli. We have previously shown that individual receptors registered directly in the tympanal organ perceive vibrations along with sound stimuli. In addition, scolopidia of the crista acustica possess mixed sensitivity, too, as well as receptors of the intermediate organ. The authors of the comment offered their opinion concerning our applied methods as well as our obtained results. In particular, they noted the dissimilarity of our data from the previously obtained data (the 1970s-1990s), mainly in the laboratory of Prof. K. Kalmring, who assumed that only low-frequency receptors, in particular receptors of the intermediate organ, possess mixed sensitivity. At the same time, receptor activity was recorded in the tympanal nerve without morphological identification of receptors (with the exception of one stained neuron in the prothoracic ganglion). We carried out a series of experiments using the method of K. Kalmring and found that it is possible to register several receptors in the tympanal nerve with different reactions during one experiment: to sound only, also both to vibration stimuli and sound. In the latter case, we dealt with low-threshold receptors, which responded to ultrasound, and this with high probability belonged to the crista acustica. Similar data were previously obtained on the bush-cricket Decticus verrucivorus. In this publication, we explain the methodological features of our work and suggest that the loss of sensitivity to vibrations at the level of the tympanal nerve by some auditory receptors may be due to the ephaptic and/or chemical interaction of the tympanal organ receptors with vibroreceptors of the subgenual or other organs. To verify this hypothesis, it is necessary to conduct additional studies, such as physiological, morphological, and immunohistochemical, along the entire vibroacoustic afferent tract, that is, from the peripheral part to the first switches to the corresponding interneurons.

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.

Do the evolutionary interactions between moths and bats promote niche partitioning between bats and birds?

Ecological theory suggests that the coexistence of species is promoted by the partitioning of available resources, as in dietary niche partitioning where predators partition prey. Yet, the mechanisms underlying dietary niche partitioning are not always clear. We used fecal DNA metabarcoding to investigate the diets of seven nocturnal insectivorous bird and bat species. Low diet overlap (2%-22%) supported resource partitioning among all species. Differences in diet corresponded with species identity, prey detection method, and foraging behavior of predators. Insects with ultrasonic hearing capabilities were consumed significantly more often by birds than bats, consistent with an evolved avoidance of echolocating strategies. In turn, bats consumed a greater proportion of noneared insects such as spruce budworms. Overall, our results suggest that evolutionary interactions among bats and moths translate to dietary niche partitioning and coexistence among bats and nocturnal birds.

Discrepancies in the spiking threshold and frequency sensitivity of nocturnal moths explainable by biases in the canonical auditory stimulation method.

The auditory stimulation method used in experiments on moth A cell(s) is generally believed to be adequate to characterize the encoding of bat echolocation signals. The stimulation method hosts, though, several biases. Their compounded effects can explain a range of discrepancies between the reported electrophysiological recordings and significantly alter the current interpretation. To test the hypothesis that the bias may significantly alter our current understanding of the moth's auditory transducer characteristics, papers using the same auditory stimulation method and reporting on either spiking threshold or spiking activity of the moth's A cells were analysed. The consistency of the reported data was assessed. A range of corrections issued from best practices and theoretical background were applied to the data in an attempt to re-interpret the data. We found that it is not possible to apply a posteriori corrections to all data and bias. However the corrected data indicate that the A cell's spiking may (i) be independent of the repetition rate, (ii) be maximum when detecting long and low-intensity pulses and (iii) steadily reduce as the bat closes on the moth. These observations raise the possibility that a fixed action pattern drives the moths' erratic evasive manoeuvres until the final moment. In-depth investigations of the potential bias also suggest that the auditory transducer's response may be constant for a larger frequency range than thought so far, and provide clues to explain the negative taxis in response to the searching bats' calls detection.

The cochlea of the enigmatic pygmy right whale Caperea marginata informs mysticete phylogeny.

The pygmy right whale, Caperea marginata, is the least understood extant baleen whale (Cetacea, Mysticeti). Knowledge on its basic anatomy, ecology, and fossil record is limited, even though its singular position outside both balaenids (right whales) and balaenopteroids (rorquals + grey whales) gives Caperea a pivotal role in mysticete evolution. Recent investigations of the cetacean cochlea have provided new insights into sensory capabilities and phylogeny. Here, we extend this advance to Caperea by describing, for the first time, the inner ear of this enigmatic species. The cochlea is large and appears to be sensitive to low-frequency sounds, but its hearing limit is relatively high. The presence of a well-developed tympanal recess links Caperea with cetotheriids and balaenopteroids, rather than balaenids, contrary to the traditional morphological view of a close Caperea-balaenid relationship. Nevertheless, a broader sample of the cetotheriid Herpetocetus demonstrates that the presence of a tympanal recess can be variable at the specific and possibly even the intraspecific level.

High duty cycle pulses suppress orientation flights of crambid moths.

Bat-and-moth is a good model system for understanding predator-prey interactions resulting from interspecific coevolution. Night-flying insects have been under predation pressure from echolocating bats for 65Myr, pressuring vulnerable moths to evolve ultrasound detection and evasive maneuvers as counter tactics. Past studies of defensive behaviors against attacking bats have been biased toward noctuoid moth responses to short duration pulses of low-duty-cycle (LDC) bat calls. Depending on the region, however, moths have been exposed to predation pressure from high-duty-cycle (HDC) bats as well. Here, we reveal that long duration pulse of the sympatric HDC bat (e.g., greater horseshoe bat) is easily detected by the auditory nerve of Japanese crambid moths (yellow peach moth and Asian corn borer) and suppress both mate-finding flights of virgin males and host-finding flights of mated females. The hearing sensitivities for the duration of pulse stimuli significantly dropped non-linearly in both the two moth species as the pulse duration shortened. These hearing properties support the energy integrator model; however, the threshold reduction per doubling the duration has slightly larger than those of other moth species hitherto reported. And also, Asian corn borer showed a lower auditory sensitivity and a lower flight suppression to short duration pulse than yellow peach moth did. Therefore, flight disruption of moth might be more frequently achieved by the pulse structure of HDC calls. The combination of long pulses and inter-pulse intervals, which moths can readily continue detecting, will be useful for repelling moth pests.

Development of leg chordotonal sensory organs in normal and heat shocked embryos of the cricket Teleogryllus commodus (Walker).

This paper describes the embryonic development of some parts of the sensory peripheral nervous system in the leg anlagen of the cricket Teleogryllus commodus in normal and heat shocked embryos. The first peripheral neurons appear at the 30% stage of embryogenesis. These tibial pioneer neurons grow on a stereotyped path to the central nervous system and form a nerve which is joined by the growth cones of axons that arise later, including those from the femoral chordotonal organ, subgenual organ and tympanal organ. The development of these organs is described with respect to the increase in number of sensory receptor cells and the shape and position of the organs. At the 100% stage of embryogenesis all three organs have completed their development in terms of the number of sense cells and have achieved an adult shape. To study the function of the tibial pioneer neurons during embryogenesis a heat shock was used to prevent their development. Absence of these neurons has no effect on the development of other neurons and organs proximal to them. However, the development of distal neurons and organs guided by them is impaired. The tibial pioneer neurons grow across the segmental boundary between femur and tibia early in development, and the path they form seems to be essential for establishing the correct connections of the distal sense organs with the central nervous system.