Several taxa of insects evolved a tympanate ear at different body positions, whereby the ear is composed of common parts: a scolopidial sense organ, a tracheal air space, and a tympanal membrane. Here, we analyzed the anatomy and physiology of the ear at the ventral prothorax of the sarcophagid fly, Emblemasoma auditrix (Soper). We used micro-computed tomography to analyze the ear and its tracheal air space in relation to the body morphology. Both tympana are separated by a small cuticular bridge, face in the same frontal direction, and are backed by a single tracheal enlargement. This enlargement is connected to the anterior spiracles at the dorsofrontal thorax and is continuous with the tracheal network in the thorax and in the abdomen. Analyses of responses of auditory afferents and interneurons show that the ear is broadly tuned, with a sensitivity peak at 5 kHz. Single-cell recordings of auditory interneurons indicate a frequency- and intensity-dependent tuning, whereby some neurons react best to 9 kHz, the peak frequency of the host’s calling song. The results are compared to the convergently evolved ear in Tachinidae (Diptera).

1. Acoustically guided movement in a three-dimensional space is a complex behavioural task performed notably by birds, bats, and some insect species. The precision of acoustic orientation depends on the directionality of the hearing system as well as on auditory behaviour.2. The fly Emblemasoma auditrix Diptera (Sarcophagidae) is a parasitoid of the cicada Okanagana rimosa Auchenorrhyncha (Cicadidae) and locates its host in the complex habitat of a forest. The phonotactic behaviour of the fly was analysed experimentally with emphasis on the vertical domain in the field. Different experimental setups allowed discriminating subsequent steps in the phonotactic behaviour of E. auditrix.3. During the phonotactic flight, flies first landed on landmarks, which were used to re-adjust to the elevation of the sound source. Acoustic targets were located from these resting positions. The sound source elevation was detected at the start of the flight as the longitudinal body axis was adjusted to the inclination of the target sound source.4. Flies usually did not land directly upon the sound source, but landed nearby, and most often above the target. Within the target area, types of movement for the final approach differed in respect to target position; flies walked predominantly if the final target was located above or below, but for horizontally located targets much of the distance was covered by flight.5. In conclusion, E. auditrix can locate the acoustic target in complex habitats and uses a flexible multi-step approach for short-range phonotaxis.

The parasitoid fly Emblemasoma auditrix locates its hosts using acoustic cues from sound producing males of the cicada Okanagana rimosa. Here, we experimentally analysed the flight path of the phonotaxis from a landmark to the target, a hidden loudspeaker in the field. During flight, the fly showed only small lateral deviations. The vertical flight direction angles were initially negative (directed downwards relative to starting position), grew positive (directed upwards) in the second half of the flight, and finally flattened (directed horizontally or slightly upwards), typically resulting in a landing above the loudspeaker. This phonotactic flight pattern was largely independent from sound pressure level or target distance, but depended on the elevation of the sound source. The flight velocity was partially influenced by sound pressure level and distance, but also by elevation. The more elevated the target, the lower was the speed. The accuracy of flight increased with elevation of the target as well as the landing precision. The minimal vertical angle difference eliciting differences in behaviour was 10°. By changing the elevation of the acoustic target after take-off, we showed that the fly is able to orientate acoustically while flying.

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