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Airborne sound signals function as key mediators of mate-choice, aggression and other social interactions in a wide range of vertebrate and invertebrate animals. Calling animals produce more than sound, however. When displaying on or near a solid substrate, such as vegetation or soil, they also unavoidably excite substrate vibrations because of the physics of sound production and of acoustic propagation, and these vibrations can propagate to receivers. Despite their near ubiquity, these vibrational signal components have received very little research attention and in vertebrates it is unknown whether they are relevant to mate-choice, an important driver of evolutionary divergence. Here, we show that female red-eyed treefrogs are more than twice as likely to choose a male mating call when airborne sound is paired with its corresponding substrate vibrations. Furthermore, males of the same species are more aggressive towards and display a greater range of aggressive behaviors in response to bimodal (sound and vibration) versus unimodal (sound or vibration alone) calls. In aggressive contexts, at least, air- and substrate-borne signal components function non-redundantly. These results are a clear demonstration that vibrations produced by a calling animal can function together with airborne sound to markedly enhance the function of a signal. If this phenomenon proves widespread, this finding has the potential to substantially influence our understanding of the function and evolution of acoustic signals., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

A study was undertaken to determine whether Scolypopa australis, the passionvine hopper, communicates using substrate-borne vibrations, as its use of such signals for communication is currently unknown. This insect is a costly pest to the kiwifruit industry in New Zealand, where few pest management tools can be used during the growing season. Vibrations emitted by virgin females and males of S. australis released alone on leaves of Griselinia littoralis were recorded with a laser vibrometer to identify and characterise potential spontaneous calling signals produced by either sex. In addition to single-insect trials, preliminary tests were conducted with female–male pair trials to determine whether individuals exchanged signals. The signal repertoire of S. australis includes a male calling signal and two female calling signals. However, no evidence of duetting behaviour that is potentially necessary for pair formation has been found to date. Our outcome suggests that a deeper understanding of the role of vibrational communication employed by S. australis is needed, and by disclosing the pair formation process, a new residue-free pest management tool against this pest may be developed. In addition, this vibration-based tool could contribute to future biosecurity preparedness and response initiatives. [ABSTRACT FROM AUTHOR]

Simple Summary: Plant-dwelling stink bugs communicate with chemical and plant-borne vibratory signals that are altered when transmitted through the substrate and air. Mates are attracted to the same plant by pheromones, where they exchange information through mechanical and close-range chemical signals. Plants absorb odor molecules from insects and produce kairomones that can enhance or reduce insect responses to pheromones. Long-range communication between stink bug mates in the field occurs exclusively via pheromones. The species specificity of the sex-pheromones guarantees their success in finding mates in the plant environment. Substrate-borne communication occurs in a narrow frequency range that is tuned to the mechanical properties of the plant. Vibratory signals are transmitted with low attenuation and with altered frequency, amplitude, and temporal characteristics. The frequency sensitivity of the subgenual organ is tuned to the low-frequency resonant properties of the plants. Recognition is encoded in the vibratory signal species- and sex-specific temporal parameters and directionality in the time delay between signals arriving from different directions. The characteristics of behaviorally described multimodal close-range communication on a plant are under-investigated. Studies of neuronal processing of multimodal sensory signals in the stink bug brain are needed to understand how their integration affects behavioral responses. Plants influenced the evolution of plant-dwelling stink bugs' systems underlying communication with chemical and substrate-borne vibratory signals. Plant volatiles provides cues that increase attractiveness or interfere with the probability of finding a mate in the field. Mechanical properties of herbaceous hosts and associated plants alter the frequency, amplitude, and temporal characteristics of stink bug species and sex-specific vibratory signals. The specificity of pheromone odor tuning has evolved through highly specific odorant receptors located within the receptor membrane. The narrow-band low-frequency characteristics of the signals produced by abdomen vibration and the frequency tuning of the highly sensitive subgenual organ vibration receptors match with filtering properties of the plants enabling optimized communication. A range of less sensitive mechanoreceptors, tuned to lower vibration frequencies, detect signals produced by other mechanisms used at less species-specific levels of communication in a plant environment. Whereas the encoding of frequency-intensity and temporal parameters of stink bug vibratory signals is relatively well investigated at low levels of processing in the ventral nerve cord, processing of this information and its integration with other modalities at higher neuronal levels still needs research attention. [ABSTRACT FROM AUTHOR]

The Hemiptera, with approximately 98 000 species, is one of the largest insect orders. Most species feed by sucking sap from plant tissues and are thus often vectors for economically important phytopathogens. Well known within this group are the large cicadas (Cicadomorpha: Cicadoidea: Cicadidae) because they produce extremely loud airborne sounds. Less well known are their mostly tiny relatives, the leafhoppers, spittlebugs, treehoppers and planthoppers that communicate by silent vibrational signals. While the generation of these signals has been extensively investigated, the mechanisms of their perception are poorly understood. This study provides a complete description and three-dimensional reconstruction of a large and complex array of mechanoreceptors in the first abdominal segments of the Rhododendron leafhopper Graphocephala fennahi (Cicadomorpha: Membracoidea: Cicadellidae). Further, we identify homologous organs in the spittlebug Philaenus spumarius (Cicadomorpha: Cercopoidea: Aphrophoridae) and the planthopper Issus coleoptratus (Fulgoromorpha: Fulgoroidea: Issidae). Such large abdominal sensory arrays have not been found in any other insect orders studied so far. This indicates that these sense organs, together with the signal-producing tymbal organ, constitute a synapomorphy of the Tymbalia (Hemiptera excl. Sternorrhyncha). Our results contribute to the understanding of the evolution from substrate-borne to airborne communication in insects.

Substrate-borne vibratory signals are thought to be one of the most ancient and taxonomically widespread communication signals among animal species, including Drosophila flies. 1–9 During courtship, the male Drosophila abdomen tremulates (as defined in Busnel et al. 10) to generate vibrations in the courting substrate. 8,9 These vibrations coincide with nearby females becoming immobile, a behavior that facilitates mounting and copulation. 8,11–13 It was unknown how the Drosophila female detects these substrate-borne vibratory signals. Here, we confirm that the immobility response of the female to the tremulations is not dependent on any air-borne cue. We show that substrate-borne communication is used by wild Drosophila and that the vibrations propagate through those natural substrates (e.g., fruits) where flies feed and court. We examine transmission of the signals through a variety of substrates and describe how each of these substrates modifies the vibratory signal during propagation and affects the female response. Moreover, we identify the main sensory structures and neurons that receive the vibrations in the female legs, as well as the mechanically gated ion channels Nanchung and Piezo (but not Trpγ) that mediate sensitivity to the vibrations. Together, our results show that Drosophila flies, like many other arthropods, use substrate-borne communication as a natural means of communication, strengthening the idea that this mode of signal transfer is heavily used and reliable in the wild. 3,4,7 Our findings also reveal the cellular and molecular mechanisms underlying the vibration-sensing modality necessary for this communication. [Display omitted] • Courting male abdominal tremulations signal via substrate-borne, not air-borne, vibrations • Substrate-borne vibrations propagate through natural food substrates • Neurons of the leg fCHO mediate female immobility response to these vibrations • Mechanically gated ion channels Nan and Piezo mediate response in these neurons McKelvey et al. find that courting male D. melanogaster abdominal tremulations signal via substrate-borne, not air-borne, vibrations. These vibrations propagate through natural food substrates, promoting female immobility. Neurons and their mechanically gated ion channels within the leg fCHO mediate female response to the vibrations. [ABSTRACT FROM AUTHOR]

Simple Summary: Stingless bees visit flowers to collect pollen and nectar which is stored in their nest. These stores are highly valuable and help the colony to overcome times of resource scarcity. The bees protect the collected nectar and pollen against robbers of all kind. It should be advantageous for a colony if it can recognize intruders and chase them away. In this context, the communication with nestmates might be crucial. The most important signals that stingless bees use are chemicals and vibrations. Communication between nestmates has been mostly studied in the context of foraging. Little is known about communication in the context of defense. We tested if vibrational signals play a role in nest defense and nestmate recognition of the African stingless bee Axestotrigona ferruginea. Stingless bees produce distinct vibrations in the context of foraging and guarding. Foraging vibrations most likely contain food source information whereas guarding vibrations might be used to alarm nestmates. Flower visiting stingless bees store collected pollen and nectar for times of scarcity. This stored food is of high value for the colony and should be protected against con- and heterospecifics that might rob them. There should be high selective pressure on the evolution of mechanisms to discriminate nestmates from non-nestmates and to defend the nest, i.e., resources against intruders. Multimodal communication systems, i.e., a communication system that includes more than one sensory modality and provide redundant information, should be more reliable than unimodal systems. Besides olfactory signals, vibrational signals could be used to alert nestmates. This study tests the hypothesis that the vibrational communication mode plays a role in nest defense and nestmate recognition of Axestotrigona ferruginea. Substrate vibrations induced by bees were measured at different positions of the nest. The experiments show that guarding vibrations produced in the entrance differ in their temporal structure from foraging vibrations produced inside the nest. We show that guarding vibrations are produced during non-nestmate encounters rather than nestmate encounters. This further supports the idea that guarding vibrations are a component of nest defense and alarm communication. We discuss to whom the vibrations are addressed, and what their message and meaning are. [ABSTRACT FROM AUTHOR]

Signalling via substrate vibration represents one of the most ubiquitous and ancient modes of insect communication. In crickets (Grylloidea) and other taxa of tympanate Ensifera, production and detection of acoustic and vibrational signals are closely linked functionally and evolutionarily. Male stridulation produces both acoustic and vibrational signal components, the joint perception of which improves song recognition and female orientation towards the signaller. In addition to stridulation, vibrational signalling mainly through body tremulation and/or drumming with body parts on the substrate has long been known to be part of crickets' close-range communication, including courtship, mate guarding and aggression. Such signalling is typically exhibited by males, independently or in conjunction with stridulation, and occurs literally in all cricket lineages and species studied. It is further also part of the aggressive behaviour of females, and in a few cricket groups, females respond vibrationally to acoustic and/or vibrational signals from males. The characteristics and function of these signals have remained largely unexplored despite their prevalence. Moreover, the communication potential and also ubiquity of cricket vibrational signals are underappreciated, limiting our understanding of the function and evolution of the cricket signalling systems. By providing a concise review of the existing knowledge of cricket perception of vibrations and vibrational signalling behaviour, we critically comment on these views, discuss the communication value of the emitted signals and give some methodological advice respecting their registration and control. The review aims to increase awareness, understanding and research interest in this ancient and widespread signalling mode in cricket communication., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Simple Summary: In this work, we investigated the effects of conspecific female rival signals in vibratory communication and mating behavior of three species of stink bugs. In the presence of rival female signals, as noisy background vibrations, couples (a male and a female) of the three species showed negative effects in their sexual vibratory communication that resulted in reduced mating and copulation in relation to pairs not exposed to rival signals. The results suggest that female rival signals could be used to disrupt mating and may be a tool for stink bug management by reducing their population increase. Stink bugs are major pests in diverse crops around the world. Pest management strategies based on insect behavioral manipulation could help to develop biorational management strategies of stink bugs. Insect mating disruption using vibratory signals is an approach with high potential for pest management. The objective of this work was to investigate the effect of conspecific female rival signals on the mating behavior and copulation of three stink bug species to establish their potential for mating disruption. Previously recorded female rival signals were played back to bean plants where pairs of the Neotropical brown stink bug, Euschistus heros, and two green stink bugs, Chinavia ubica and Chinavia impicticornis were placed. Vibratory communication and mating behavior were recorded for each pair throughout the experimental time (20 min). Female rival signals show a disrupting effect on the reproductive behavior of three conspecific investigated stink bug species. This effect was more clearly expressed in E. heros and C. ubica than in C. impicticornis. The likelihood of copulating in pairs placed on control plants, without rival signals, increased 29.41 times in E. heros, 4.6 times in C. ubica and 1.71 times in C. impicticornis. However, in the last case, the effect of female rivalry signals in copulation was not significant. The effect of mating disruption of female rival signals of the three stink bug species may originate from the observed reduction in specific vibratory communication signals emitted, which influences the duet formation and further development of different phases of mating behavior. Our results suggest that female rival signals have potential for application in manipulation and disruption of mating behavior of stink bugs. Further work needs to focus on the effects of female rival signals used in long duration experiments and also their interactions with chemical communication of stink bugs. [ABSTRACT FROM AUTHOR]

Cicadas and many of their relatives (Hemiptera: Cicadomorpha) generate vibroacoustic signals using tymbal organs located on their first two abdominal segments. Although tymbals are well-studied in Cicadidae, their systematic distribution in other Cicadomorpha and their possible homologies to the vibroacoustic mechanisms of other Hemiptera have been debated for more than a century. In the present study, we re-examine the morphology of the musculoskeletal system of cicadomorphan vibroacoustic organs, and we document their systematic distribution in 78 species drawn from across the phylogeny of Cicadomorpha. We also compare their morphology to the recently-described snapping organ of planthoppers (Fulgoromorpha). Based on the structure and innervation of the metathoracic and abdominal musculoskeletal system, we find that several key elements of cicadomorphan vibroacoustic organs that have previously been assigned to the first abdominal segment in fact belong to the second. We find that tymbal organs are nearly ubiquitous in Cicadomorpha, and conclude based on their phylogenetic distribution, that they are likely to be synapomorphic. The unusual tymbal-like organs of the Deltocephalinae and Typhlocybinae, represent derived modifications. Finally, we propose a standardised terminology for sternal components of the cicadomorphan vibrational organs, which can be used in future taxonomic descriptions. • The segmental affinities of the tymbal musculoskeletal system are re-evaluated. • Tymbals and tymbal-like organs are present across Cicadomorpha. • The possession of tymbal-like organs is likely synapomorphic for Cicadomorpha. • The homologies between tymbal organs and snapping organs require further study. • We propose a new terminology for the cicadomorphan sternal sclerites and their apodemes. [ABSTRACT FROM AUTHOR]

Mechanosensory-cued hatching (MCH) is widespread, diverse and important for survival in many animals. From flatworms and insects to frogs and turtles, embryos use mechanosensory cues and signals to inform hatching timing, yet mechanisms mediating mechanosensing in ovo are largely unknown. The arboreal embryos of red-eyed treefrogs, Agalychnis callidryas , hatch prematurely to escape predation, cued by physical disturbance in snake attacks. When otoconial organs in the developing vestibular system become functional, this response strengthens, but its earlier occurrence indicates another sensor must contribute. Post-hatching, tadpoles use lateral line neuromasts to detect water motion. We ablated neuromast function with gentamicin to assess their role in A. callidryas ' hatching response to disturbance. Prior to vestibular function, this nearly eliminated the hatching response to a complex simulated attack cue, egg jiggling, revealing that neuromasts mediate early MCH. Vestibular function onset increased hatching, independent of neuromast function, indicating young embryos use multiple mechanosensory systems. MCH increased developmentally. All older embryos hatched in response to egg jiggling, but neuromast function reduced response latency. In contrast, neuromast ablation had no effect on the timing or level of hatching in motion-only vibration playbacks. It appears only a subset of egg-disturbance cues stimulate neuromasts; thus, embryos in attacked clutches may receive unimodal or multimodal stimuli. Agalychnis callidryas embryos have more neuromasts than described for any other species at hatching, suggesting precocious sensory development may facilitate MCH. Our findings provide insight into the behavioral roles of two mechanosensory systems in ovo and open possibilities for exploring sensory perception across taxa in early life stages., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Web spiders rely on vibrations propagated via their web to identify, locate and capture entangled prey. Here, we experimentally tested the robustness of the orb weaver's predation strategy when webs are severely distorted and silk tensions are drastically altered throughout the web, a common occurrence in the wild. We assessed prey identification efficiency by comparing the spider's initial reaction times towards a fruit fly trapped in the web, we measured location efficiency by comparing times and number of tugging bouts performed, and we determined capture efficiency by comparing capture times. It emerged that spiders are capable of identifying, locating and capturing prey in distorted webs, albeit taking somewhat longer to do so., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

The production of biotremors has been described in veiled chameleons ( Chamaeleo calyptratus ), but the mechanism by which they are produced is unknown. We gathered muscle activation data via electromyography (EMG), with simultaneous recordings of biotremors using an accelerometer, to test for the role of hyoid muscles in biotremor production. We recorded a mean biotremor frequency of 150.87 Hz for females and 136.01 Hz for males. The durations of activity and the latencies to onset and offset for the M. sternohyoideus profundus (SP), M. sternohyoideus superficialis (SS), Mm. mandibulohyoideus (MH) and M. levator scapulae (LS) were all significantly correlated with biotremor durations and biotremor onset and offset, respectively. Linear mixed-effect regression model comparisons of biotremor duration indicated that models containing either the MH and/or the SP and LS account for the most variation in biotremor duration. Twitch times for the SP (100 ms) and the SS (132 ms) at field active body temperature, however, were individually too slow to produce the biotremors at the observed frequency without alteration after production by other anatomical structures. These results implicate the SP, SS, MH and LS in the production of biotremors, but the exact mechanism of production requires further study., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Multimodal signaling systems are shaped not only by a signaler's physical abilities but also by external factors such as the position of signal receivers and the properties of the medium through which the signals are transmitted. To fully understand the evolution and function of multimodal communication, it is essential to investigate the behavior in the wild. Here, I present evidence that socially monogamous songbirds perform complex courtship displays that can produce multimodal and multicomponent signals in wild conditions. Cordon-bleus (Uraeginthus spp.) are socially monogamous songbirds from East Africa. Both sexes of cordon-bleus perform multimodal courtship displays by holding a piece of nest material, bobbing up and down, and singing. My previous laboratory study using high-speed video cameras revealed that courtship bobbing includes multiple rapid steps similar to human tap-dancing, which presumably contributes to producing non-vocal sounds and/or vibrations in addition to visual signals. As a result of field observation and behavioral analysis, I found that wild cordon-bleus perform tap-dance like displays just as captive cordon-bleus. I also observed that wild cordon-bleus produced non-vocal sounds and shook branches during courtship, which can contribute to multimodal signal production (i.e., visual, acoustic, and vibrational signals). My findings imply that the courtship displays of cordon-bleus are an ideal candidate for investigating the role and function of multimodal communication in animals, and demonstrate the importance of further quantitative studies in both laboratory and field.

Vibrations play an important role in insect behaviour. In bees, vibrations are used in a variety of contexts including communication, as a warning signal to deter predators and during pollen foraging. However, little is known about how the biomechanical properties of bee vibrations vary across multiple behaviours within a species. In this study, we compared the properties of vibrations produced by Bombus terrestris audax (Hymenoptera: Apidae) workers in three contexts: during flight, during defensive buzzing, and in floral vibrations produced during pollen foraging on two buzz-pollinated plants ( Solanum , Solanaceae). Using laser vibrometry, we were able to obtain contactless measures of both the frequency and amplitude of the thoracic vibrations of bees across the three behaviours. Despite all three types of vibrations being produced by the same power flight muscles, we found clear differences in the mechanical properties of the vibrations produced in different contexts. Both floral and defensive buzzes had higher frequency and amplitude velocity, acceleration and displacement than the vibrations produced during flight. Floral vibrations had the highest frequency, amplitude velocity and acceleration of all the behaviours studied. Vibration amplitude, and in particular acceleration, of floral vibrations has been suggested as the key property for removing pollen from buzz-pollinated anthers. By increasing frequency and amplitude velocity and acceleration of their vibrations during vibratory pollen collection, foraging bees may be able to maximise pollen removal from flowers, although their foraging decisions are likely to be influenced by the presumably high cost of producing floral vibrations., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

• Variability of vibrational courtship signals underlies both natural and sexual selection. • Male vibratory signals in Mantophasmatodea vary with temperature, condition and age. • Females were selective towards fine scale signal patterns but not towards coarser signal pattern combinations. • Heelwalker vibrational signals are used to recognize and localize mates but less important in precopulatory mate selection. Communication systems that involve substrate vibrations are increasingly a focus of research since this communication mode - recently termed biotremology - has been found to be remarkably widespread in the animal kingdom. Vibrational signals are often used during courtship and therefore underlie both natural and sexual selection. Mantophasmatodea use species- and sex-specific substrate vibrational signals during courtship. We explored whether male vibrational signals of the South African heelwalker Karoophasma biedouwense vary with temperature, body condition and age, and tested female preference towards various signal pattern combinations. We recorded male signals under varying temperatures and over 3.5 weeks after onset of signaling. Our results show that the temporal structure of male signals is modified by changes in temperature, and changes with male age. Other characteristics, especially duty cycles, are less affected, but correlate with body condition. Females responded along a broad spectrum of signaling patterns, indicating that they do not favor signals of males of a certain age or condition. They were selective towards the fine structure of vibratory signals, suggesting that pulse repetition times carry species-specific information. Mantophasmatodea thus use vibrational signals to identify and localize a mating partner, but presumably not for precopulatory mate selection. [ABSTRACT FROM AUTHOR]

Summary Living in the African savanna is dangerous, especially for plants. Many plants therefore engage in mutualism with ants, in which plants provide food and shelter in exchange for protection against herbivores. Ants become alarmed when the plant takes on some sort of damage. They immediately emerge from their plant shelter and aggressively defend the plant. Mammalian herbivores can have devastating effects on trees by browsing, breaking tree branches, stripping bark, and pushing over entire trees. However, mutualistic ants substantially reduce the amount of damage. To efficiently protect the tree, ants need to rapidly react together when the tree is under attack. Here, we show that the acacia ant Crematogaster mimosae defends its host tree by exploiting plant-borne vibrations caused by browsers feeding on the tree. Experiments with controlled vibrations show that ants discriminate browser-induced vibrations from those induced by wind, become alarmed, and patrol on the branches. Browser-induced vibrations serve as a long-distance alarm cue. The vibrations propagate through the whole acacia tree and trigger ants' defensive behavior, even on the other side of the tree. Furthermore, the ants make use of tropotactic directional vibration sensing to orient to the attacked part of the tree and fight back the attacker. Highlights • Acacia ants discriminate browser-induced vibrations from wind-induced vibrations • Browser-induced vibrations serve as a long-distance alarm cue for ants • Acacia ants make use of tropotactic directional vibration sensing Hager and Krausa show that the acacia ant Crematogaster mimosae defends its host tree by exploiting plant-borne vibrations caused by browsers feeding on the tree. Browser-induced vibrations serve as a long-distance alarm cue, and the ants make use of tropotactic directional vibration sensing to orient to the attacked part of the tree. [ABSTRACT FROM AUTHOR]

The widespread ability to alter timing of hatching in response to environmental cues can serve as a defense against threats to eggs. Arboreal embryos of red-eyed treefrogs, Agalychnis callidryas , can hatch up to 30% prematurely to escape predation. This escape-hatching response is cued by physical disturbance of eggs during attacks, including vibrations or motion, and thus depends critically on mechanosensory ability. Predator-induced hatching appears later in development than flooding-induced, hypoxia-cued hatching; thus, its onset is not constrained by the development of hatching ability. It may, instead, reflect the development of mechanosensor function. We hypothesize that vestibular mechanoreception mediates escape-hatching in snake attacks, and that the developmental period when hatching-competent embryos fail to flee from snakes reflects a sensory constraint. We assessed the ontogenetic congruence of escape-hatching responses and an indicator of vestibular function, the vestibulo-ocular reflex (VOR), in three ways. First, we measured VOR in two developmental series of embryos 3-7 days old to compare with the published ontogeny of escape success in attacks. Second, during the period of greatest variation in VOR and escape success, we compared hatching responses and VOR across sibships. Finally, in developmental series, we compared the response of individual embryos to a simulated attack cue with their VOR. The onset of VOR and hatching responses were largely concurrent at all three scales. Moreover, latency to hatch in simulated attacks decreased with increasing VOR. These results are consistent with a key role of the vestibular system in the escape-hatching response of A. callidryas embryos to attacks., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)