Adaptations to divert the attacks of visually guided predators have evolved repeatedly in animals. Using high-speed infrared videography, we show that luna moths (Actias luna) generate an acoustic diversion with spinning hindwing tails to deflect echolocating bat attacks away from their body and toward these nonessential appendages. We pit luna moths against big brown bats (Eptesicus fuscus) and demonstrate a survival advantage of ∼47% for moths with tails versus those that had their tails removed. The benefit of hindwing tails is equivalent to the advantage conferred to moths by bat-detecting ears. Moth tails lured bat attacks to these wing regions during 55% of interactions between bats and intact luna moths. We analyzed flight kinematics of moths with and without hindwing tails and suggest that tails have a minimal role in flight performance. Using a robust phylogeny, we find that long spatulate tails have independently evolved four times in saturniid moths, further supporting the selective advantage of this anti-bat strategy. Diversionary tactics are perhaps more common than appreciated in predator-prey interactions. Our finding suggests that focusing on the sensory ecologies of key predators will reveal such countermeasures in prey.antipredator defense | bat-moth interactions | Lepidoptera | Saturniidae P redators are under pressure to perform incapacitating initial strikes to thwart prey escape. It is thought that prey, in turn, have evolved conspicuous colors or markings to deflect predator attack to less vulnerable body regions (1, 2). Eyespots are a wellknown class of proposed deflection marks (3), which are found in a variety of taxa, including Lepidoptera (3) and fishes (4), but only recently have experiments convincingly demonstrated that these color patterns redirect predatory assault. Eyespots on artificial butterfly (5) and fish (4) prey draw strikes of avian and fish predators. Eyespots on the wing margins of woodland brown butterflies (Lopinga achine) lure the attacks of blue tits (Cyanistes caeruleus) (6). Brightly colored lizard tails also divert avian predator attacks to this expendable body region (7).Deflection coloration is unlikely to be an effective strategy against echolocating bats, as these predators have scotopic vision and poor visual acuity unsuited for prey localization and discrimination (8). Most bats rely on echoes from their sonar cries to image prey and other objects in their environment-they live in an auditory world (9). Thus, we would expect a deflection strategy, effective against bats, to present diversionary acoustic signatures to these hearing specialists. Weeks (10) proposed that saturniid hindwing tails might serve to divert bat attacks from essential body parts. We hypothesized that saturniid tails, spinning behind a flying moth (Movie S1) and reflecting sonar calls, serve as either a highly contrasting component of the primary echoic target or as an alternative target. We predicted that bats would aim their attacks at moth tails, instead of the wings or body, durin...
We studied the effects of limestone sand additions in Bear and Rock runs, two chronically and episodically acidified streams in southwestern Pennsylvania, U.S.A. Linn Run, a nearby episodically acidified stream, served as a reference stream. Our objectives were to evaluate the effects of doubling recommended limestone sand amounts on water quality and macroinvertebrates on Bear and Rock runs and to assess substrate changes resulting from limestone sand inundation. Approximately 23 and 6 tonnes of limestone sand were added annually to the headwaters of Bear and Rock runs, respectively, from 1999 to 2001. In 2002, amounts were doubled. Macroinvertebrate communities were assessed from 1999 to 2003 at points above and below the sand additions on Bear Run and Rock Run. Small, plastic substrate samplers were used to assess sand substrate effects. Doubling annual limestone sand amounts resulted in significantly improved pH and acidneutralizing capacity; however, total dissolved aluminum increased significantly downstream (a • 0.05). Macroinvertebrate density and diversity were not significantly affected, but an increase in acid-sensitive taxa was observed at a site 3,500 m downstream. Substrate sampler data indicated a significant negative relationship between amount of sand deposited and density of macroinvertebrates. The mixed water quality and benthic macroinvertebrate results were reasonably consistent with earlier work and call into question the use of limestone sand in the restoration of chronically and episodically acidified waters.
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