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Document Type

Peer-Review Article

Abstract

Visual and phonotactic orientation often occur simultaneously in diurnal cicadas. and these animals generally have their largest sensory elaboration in eyes and hearing organs. Phonotactic orientation occurs principally during flight. Males and females of Magicicada cassini commonly perform low- altitude « 5 m) and short-distance « 15 m) flights in their natural habitat at flight speeds of 3 to 6 m/s. During flight, the long body axis is tilted 10° to Q , head upward. Wing beat frequencies of tethered animals at 24° to 26°C averaged 28.8 Hz. Body temperature in the field for flying individuals aver- aged 4.6°C above ambient.

Compound eyes of females possess about 7% more facets than males, and the binocular field of view for both is especially expanded dorsa-frontally, frontally, and fronto-ventrally. The role of vision for phonoresponses, and in flight and landing behavior. was studied in nature by comparing controls with cicadas with eyes partly to completely covered with aluminum paint. Cicadas with their three ocelli covered behaved like controls and exhibited low-altitude and short-distance flights with landings on neighboring shrubs, as did cicadas with only both caudal halves or both dorsal halves of the compound eyes covered. Those with both compound eyes covered completely (with or without additionally covering the three ocelli) flew to higher altitudes and for longer distances. Higher and longer flight courses were also seen in cicadas (A) with only one compound eye covered. which in addition circled during walking and flight toward the side of unrestricted vision, (B) with both frontal or both ventral halves of their compound eyes covered. and (C) with either the binocular or monocular fields of the eyes covered. Thus, the paired fronta-antero-ventral regions of the compound eyes provide visual information for habitat-dependent low-altitude flights and landings.

Females with intact compound eyes and ocelli responded to playbacks of just the frequency/intensity sweep at the end of the buzz in calling songs of a male by flying within 1.2 m above the ground and landing on a nylon screen- covered small bush directly above the loudspeaker from distances of 2 to 8 m. mostly from lower vegetation. Males that were blinded, or blinded and deafened, sang less and flew less than normal males. However. they performed all of those behaviors, and all also walked and fed.

Periodical cicadas (Magicicada, Tibicininae) are known for synchronized adult emergence and noisy aggregations of millions of individuals of three intermingled species in each brood population (Alexander and Moore 1962). Broods are isolated geographically and chronologically, such that in some years no periodical cicada adults emerge, and most areas of the eastern United States have only one brood population appearing as adults at intervals of either 13 or 17 years. In all Magicicada species, daily flights affect spacing and aggregation of both sexes during feeding, chorusing, mating, and ovipositing. Flights are mediated by both acoustical and visual cues. Each species in these aggregations establishes mating leks. These aggregations continue to mix, every day and unpredictably, during the emergence period. Toward the end of the reproductive season, males die sooner than females, leading to little or no chorusing, and then females disperse progressively further from the lek sites. The cohesive effect of the acoustical cues of chorusing males on these cicada populations is obvious.

Both sexes of all six species of periodical cicadas live and feed on shrubs and trees of different species, sizes, and shapes, and females lay eggs in their living twigs. Their niches overlap almost completely, the three species of 13­ year or 17-year cicadas being separated principally by diurnal acoustic behavior leading to aggregation sites that change every day and are seldom exclusive to a single Magicicada species. Adults frequently change location in these complex visual environments by short-distance and low-altitude flights. which we call bush-hopping. These flights are associated with sound communication and reproductive activities and are most commonly observed during bright sunlight and at ambient temperatures above 25°C with little wind (Alexander and Moore 1958,1962; Dunning et al. 1979). Otte (1990) and Toms (1992) discuss the common correlation between hearing and flying in orthop­teroid insects, interactions basically similar to those found in cicadas. The present paper describes the interaction of vision (compound eyes and ocelli) and phonoresponses of males and females of Magicicada cassini (Fisher) in walking, but especially in flight and landing behavior, within a natural habitat.

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