Insect defense sounds have been reported for centuries. Yet, aside from the well-studied anti-bat sounds of tiger moths, little is understood about the occurrence, function, and evolution of these sounds. We define a defense sound as an acoustic signal (air- or solid-borne vibration) produced in response to attack or threat of attack by a predator or parasitoid and that promotes survival. Defense sounds have been described in 12 insect orders, across different developmental stages, and between sexes. The mechanisms of defensive sound production include stridulation, percussion, tymbalation, tremulation, and forced air. Signal characteristics vary between species, and we discuss how morphology, the intended receiver, and specific functions of the sounds could explain this variation. Sounds can be directed at predators or non-predators, and proposed functions include startle, aposematism, jamming, and alarm, although experimental evidence for these hypotheses remains scant for many insects. The evolutionary origins of defense sounds in insects have not been rigorously investigated using phylogenetic methodology, but in most cases it is hypothesized that they evolved from incidental sounds associated with non-signaling behaviors such as flight or ventilatory movements. Compared to our understanding of visual defenses in insects, sonic defenses are poorly understood. We recommend that future investigations focus on testing hypotheses explaining the functions and evolution of these survival sounds using predator-prey experiments and comparative phylogenetics.

Synopsis Insects have a diversity of hearing organs known to function in a variety of contexts, including reproduction, locating food, and defence. While the role of hearing in predator avoidance has been extensively researched over the past several decades, this research has focused on the detection of one type of predator- echolocating bats. Here we reassess the role of hearing in antipredator defence by considering how insects use their ears to detect and avoid the wide range of predators that consume them. To identify the types of sounds that could be relevant to insect prey, we first review the topic of hearing-mediated predator avoidance in vertebrates. Sounds used by vertebrate prey to assess predation risk include incidental sound cues (e.g. flight sounds, rustling vegetation, splashing) produced by an approaching predator or another escaping prey, as well as communication signals produced by a predator (e.g. echolocation calls, songs) or non-predator (e.g. alarm calls). We then review what is known, and what is not known, about such sounds made by the main predators and parasitoids of insects (i.e. birds, bats, terrestrial vertebrates, invertebrates) and how insects respond to them. Three key insights emerged from our review. First, there is a lack of information on how both vertebrate and insect prey use passive sound cues produced by predators to avoid being captured. Second, while there are numerous examples of vertebrate prey eavesdropping on the calls and songs of predators and non-predators to assess risk, there are currently no such examples for eared insect prey. Third, the hearing sensitivity of many insects, including those with ears considered to be dedicated to detecting bats or mates, overlaps with both sound cues and signals generated by non-bat predators. Sounds of particular relevance to insect prey include the flight sounds and calls of insectivorous birds, the flight sounds of insect predators and parasitoids, and rustling vegetation sounds of birds and terrestrial predators. We conclude that research on the role of insect hearing in predator avoidance has been disproportionally focused on bat-detection, and that acoustically-mediated responses to other predators may have been overlooked because the responses of prey may be subtle (e.g. ceasing activity, increasing vigilance). We recommend that researchers expand their testing of hearing-mediated risk assessment in insects by considering the wide range of sounds generated by predators, and the varied responses exhibited by prey to these sounds.

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