Drumming behavior of twelve North American stonefly (Plecoptera) species: First descriptions in Peltoperlidae, Taeniopterygidae and Chloroperlidae1

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Abstract
Previously unreported drumming signals of 11 spp. in the families Peltoperlidae, Taeniopterygidae, Capniidae, Perlodidae and Chloroperlidae are described, and signals of a 12th species, Isogenoids zionensis, are further described. Drumming in 4 Soliperla spp. [S. fenderi, S. quadrispinula, S. sierra and S. thyra] was of 2 basic types; the diphasic male calls and long 1st-phase beat intervals of S. fenderi were in marked contrast to monophasic call and shorter beat intervals of other males. Females of 3 Soliperla would drum without a male call. The 3-way Taeniopteryx nivalis signals were generally similar to 2 other known Taeniopteryx in the monophasic nature of the male call, and the number of beats in the male call, female answer and male response. Beat intervals were distinctly different from other species at comparable temperatures. Calliperla luctuosa male calls were arranged in a series of 1- or 2-beat groupings, a pattern basically similar to several other Perlodidae. Drumming of I. zionensis is the most complex known in the Plecoptera. Male-female exchanges began as interspersed signals, then changed to sequenced 3-way signals. During the interspersed portion male calls consisted of 11 beats in 3-4 groupings, within which the female answer was spaced. Both chloroperlids, Kathroperla perdita and Paraperla frontalis, had di-phasic male calls, with 6.1 .+-. 0.7 and 74 .+-. 10.6 beats, respectively, in the 1st phases, and with distinctly different beat intervals. Males of other species analyzed included: Isocapnia grandis, Helopicus nalatus and Isogenoides elongatus.