Stealth echolocation in aerial hawking bats reflects a substrate gleaning ancestry

Predator-prey co-evolution can escalate into an evolutionary arms race.1 Examples of insect countermea-sures to bat echolocation are well-known,2 but presumptive direct counter strategies in bats to insect anti-bat tactics are rare. The emission of very low-intensity calls by the hawking Barbastella barbastellus to circumvent high-frequency moth hearing is the most convincing countermeasure known.2,3 However, we demonstrate that stealth echolocation did not evolve through a high-intensity aerial hawking ancestor becoming quiet as previously hypothesized2-4 but from a gleaning ancestor transitioning into an obligate aerial hawker. Our ancestral state reconstructions show that the Plecotini ancestor likely gleaned prey using low-intensity calls typical of gleaning bats and that this ability-and associated traits-was subsequently lost in the barbastelle lineage. Barbastelles did not, however, revert to the oral, high-intensity call emission that other hawking bats use but retained the low-intensity nasal emission of closely related gleaning plecotines despite an extremely limited echolocation range. We further show that barbastelles continue to emit low -in-tensity calls even under adverse noise conditions and do not broaden the echolocation beam during the terminal buzz, unlike other vespertilionids attacking airborne prey.5,6 Together, our results suggest that bar-bastelles' echolocation is subject to morphological constraints prohibiting higher call amplitudes and beam broadening in the terminal buzz. We suggest that an abundance of eared prey allowed the co-opting and maintenance of low-intensity, nasal echolocation in today's obligate hawking barbastelle and that this unique foraging behavior7 persists because barbastelles remain a rare, acoustically inconspicuous predator to eared moths.