A new brachypterous Nusalala species from Costa Rica, with comments on the evolution of flightlessness in brown lacewings (Neuroptera: Hemerobiidae)
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A new flightless hemerobiid species, Nusalala brachyptera, collected at high elevation in Costa Rica, is described and illustrated, and a variety of data relevant to the evolution of flightlessness in the family Hemerobiidae are reviewed. Flightlessness due to brachyptery has evolved independently in at least five monophyletic [= holophyletic] lineages of the family Hemerobiidae (brown lacewings). Volant hemerobiids are primarily foliage foraging arboreal predators [presumed ancestral condition], while flightless species are predominantly associated with terricolous-type microhabitats (e.g. ground-litter, epiphytic mosses) [presumed derived condition]. These differences suggest a significant habitat shift for flightless hemerobiid species, and that the parallel evolution of flightlessness and brachyptery in hemerobiids are shared responses to the conditions of a terricolous existence. The restriction of most flightless hemerobiid species to insular and/or montane/alpine land areas may be related to the typically depauperate nature of the faunas of such areas. This faunal characteristic may facilitate transitions from arboreality to terricolousness by presenting ancestrally arboreal predators such as hemerobiids with novel ecological opportunities in terricolous microhabitats.
