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4689The evolutionary arms race between insectivorous echolocating bats and moths has long fascinated biologists (Roeder, 1967;Fullard, 1998;Miller and Surlykke, 2001;Jones and Rydell, 2003;Waters, 2003). The primary purpose of the moth's simple ear -to detect bat echolocation callshas made this a particularly useful model for study (Fullard, 1988;Waters, 2003). The ears of moths have evolved as a direct result of selective pressure by bats; they are broadly tuned to the frequency ranges of sympatric insectivorous bat communities and inform the central nervous system to initiate erratic flight behaviours and/or stop flying (Fullard, 1988;Hoy et al., 1989). As part of the acoustic startle response, some species of tiger moths (family Arctiidae) produce clicks using thoracic tymbals that appear to deter attacks from nearby bats (Dunning and Roeder, 1965; Hirstov and Conner, 2005a). Many arctiid species contain sequestered or synthesized defensive chemicals (Rothschild et al., 1970;Weller et al., 1999;Nishida, 2002) unpalatable to a variety of predators, including bats (Dunning, 1968;Coutts et al., 1973;Goss, 1979;Boppré, 1990;Hristov and Conner, 2005b).The clicks of arctiid moths have been proposed to function as acoustic aposematic signals (Blest et al., 1963;Dunning and Roeder, 1965;Dunning, 1968;Hristov and Conner, 2005a) and as signals that interfere with information processing facilitating escape, by startling the bat (deimatism) and/or jamming echolocation (Bates and Fenton, 1990;Fullard et al., 1979Fullard et al., , 1994Miller, 1991;Masters and Raver, 1996;Tougaard et al., 1998Tougaard et al., , 2004. These functional hypotheses are not mutually exclusive although they are often portrayed as such (e.g. Surlykke and Miller, 1985;Fullard et al., 1994;Waters, 2003; Hristov and Miller, 2005a). Until recently (Hristov and Conner, 2005a), all laboratory studies had used (1) synthetic bat echolocation calls and stationary moths, (2) synthetic arctiid clicks and stationary bats or (3) free-flying bats capturing mealworms while synthetic clicks were broadcast >1·m away from the prey trajectory (see Waters, 2003 and We studied the efficiency and effects of the multiple sensory cues of tiger moths on echolocating bats. We used the northern long-eared bat, Myotis septentrionalis, a purported moth specialist that takes surface-bound prey (gleaning) and airborne prey (aerial hawking), and the dogbane tiger moth, Cycnia tenera, an eared species unpalatable to bats that possesses conspicuous colouration and sound-producing organs (tymbals). This is the first study to investigate the interaction of tiger moths and wild-caught bats under conditions mimicking those found in nature and to demand the use of both aerial hawking and gleaning strategies by bats. Further, it is the first to report spectrograms of the sounds produced by tiger moths while under aerial attack by echolocating bats. During both aerial hawking and gleaning trials, all muted C...