New descriptions of drumming and evolution of the behavior in North American Perlodidae (Plecoptera)

The drumming behavior of six Plecoptera species, Clioperla clio, Diploperla robusta, Oconoperla innubila, Osobenus yakimae, Perlinodes aurea and Yugus arinus, are described for the first time, and additional data for Isogenoides zionensis are provided. Patterns and evolution of drumming in the family Perlodidae are discussed, and an out-group comparison shows that: (1) a simple call-answer exchange between sexes in seven species is ancestral, and (2) bi-grouped calls of Hydroperla crosbyi and the complex, grouped exchanges of Isogenoides zionensis are derived expressions of drumming. Similarly, grouped calls of some Isoperla and grouped bi-beat calls of Calliperla luctuosa and Kogotus modestus are derived. The grouped drumming pattern is known only in Perlodidae, and the European Leuctra pseudosignifera (Leuctridae).