SOUND PRODUCTION BY AQUATIC INSECTS

1. Sound production by aquatic insects is found in four orders — Trichoptera, Odonata, Heteroptera and Coleoptera. 2. Immature aquatic insects that produce sound are rare, stridulation being present in one family of Trichoptera (Hydropsychidae) and one genus and species in a relic suborder of Odonata (Anisozygoptera) - Epiophlebia superstes. Hydropsychid larvae produce sound with a head/fore femur mechanism and use sound as part of aggressive behaviour for defence of feeding nets. Larval E. superstes use a hind femur/abdominal mechanism to dissuade predators. 3. Sound production has been documented in adults of all families of aquatic Heteroptera except Helotrephidae. In corixids and notonectids, acoustic signals play a role in mating. Members of the genus Buenoa (Notonectidae) are unique in having two stridulatory mechanisms in the same individual. Sound production has been most intensively studied in the Corixidae. Although sounds are used in mating by all singing corixids, their use seems to be facultative in some species and obligatory in others. Recent experiments by Theiss (1982) have shown that the air stores carried by corixids are used for both sound radiation and reception. 4. The adephagan beetle families Hygrobiidae, Dytiscidae and Haliplidae have all been shown to produce sound. Mechanisms of sound production have been established for haliplids and hygrobiids but have yet to be for most dytiscids. Sound production is used by beetles as part of sequences of aggressive/defensive and reproductive behaviour. 5. Sound production is especially well documented in the Hydrophilidae (Polyphaga). Hydrophilids use an abdominal/elytral mechanism and sound appears to be used in the same contexts as in adephagans. 6. Insects that produce sound under water must contend with the physical problems of sound transmission in a relatively dense, viscous medium with sharp boundaries. Because of potential distortion of the frequency components in a signal by reflection from the air/water interface in very shallow water, frequency is unreliable for encoding information. Aquatic insects use instead amplitude modulation and temporal patterning of signals. 7. For aquatic invertebrates, sound fields are different than those in air because the extent of the near field is approximately four times greater in water. This near field, a region in which displacement waves are predominant over pressure waves, extends to a greater distance than most aquatic insects communicate over. Such displacement waves could have important but as yet unconsidered effects. 8. The mass and viscosity of the water dictates that sound producing structures of aquatic insects should be heavier and more massive than those of terrestrial insects. A survey of stridulatory organs of aquatic insects reveals this to be true and reveals that the relatively fragile, membranous stridulatory organs of some terrestrial insects (especially Orthoptera) are absent. 9. The elaboration of sound producing structures in aquatic insects probably occurred at the family or subfamily level and for Heteroptera, Trichoptera and Odonata evolved after the invasion of the water. Acoustic signals used reproductively would probably be more closely associated with the emergence of new taxa. 10. Stridulatory structures have been derived from either structures devoted to some other function or from structures involved in the behaviour currently enhanced by sound production.