We also review auditory-based defences in other insect groups. Here, we concentrate on the traits of lepidopteran species that reduce their risk of being attacked by bats, emphasizing moths' ears, auditory-evoked evasive flight and defensive sounds. Dawkins and Krebs (1979) suggest that the rarity of co-evolution between predators and prey is due to the ‘life–dinner principle’ – the cost of a mistake being death for the prey versus only a lost meal for the predator – resulting in more specific antipredator defences than predator counter-strategies. Co-evolutionary relationships are most common between host and parasite species or mutualistic partners such as plants and pollinators, but are rarely observed between predator and prey. The interactions of bats and moths do not meet a strict definition of co-evolution in which two species demonstrate reciprocal adaptations ( Janzen, 1980 Jones and Rydell, 2003 Jacobs et al., 2008 Ratcliffe, 2009). Likewise, some moths use their ears to detect not only bats but also insect-eating birds or mates ( Conner, 1999 Nakano et al., 2015). Most adaptations that make bats better moth hunters also make them more effective hunters of other insects, and such adaptations are therefore not moth specific. Although the phrase ‘co-evolutionary arms race’ is often used to describe the moths' anti-bat defences (which include bat-detecting ears, sound-evoked defensive flight and sound production), there is no evidence of reciprocal adaptations between bats and moths at the species level, nor are the causes and effects of this arms race exclusive to the interactions between bats and moths. In turn, some moths listen for bat echolocation calls to avoid being captured. Insect-eating bats detect and track moths using echolocation and attempt to capture and consume these insects in flight. Specific predator counter-adaptations include calling at frequencies outside the sensitivity range of most eared prey, changing the pattern and frequency of echolocation calls during prey pursuit, and quiet, or ‘stealth’, echolocation.Īs a well-known predator–prey model system, the interactions of bats and moths have long fascinated scientists ( Roeder, 1967), university students ( Alcock, 2013) and children alike ( Oppel, 1997). We suggest two levels of support for classifying bat traits as counter-adaptations: traits that allow bats to eat more eared prey than expected based on their availability in the environment provide a low level of support for counter-adaptations, whereas traits that have no other plausible explanation for their origination and maintenance than capturing defended prey constitute a high level of support. It is assumed that these prey adaptations would select for counter-adaptations in predatory bats. We also review hypotheses on the neural basis for anti-predator behaviours (such as evasive flight and sound production) in moths. We consider life history variables such as size, sex, circadian and seasonal activity patterns, geographic range and the composition of sympatric bat communities. Although cause and effect are difficult to determine, correlations between hearing and life history strategies in moths provide evidence for how these two variables influence each other. Ears evolved in a remarkable number of body locations across insects, with the original selection pressure for ears differing between groups. Here, we review the evolutionary history of bats and eared insects, focusing on the insect order Lepidoptera, and consider the evidence for antipredator adaptations and predator counter-adaptations. Echolocation in bats and high-frequency hearing in their insect prey make bats and insects an ideal system for studying the sensory ecology and neuroethology of predator–prey interactions.
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