Phylogeny of the family Sialidae (Insecta: Megaloptera) inferred from morphological data, with implications for generic classification and historical biogeography.

Sialidae (alderflies) is a family of the holometabolous insect order Megaloptera, with ca. 75 extant species in eight genera distributed worldwide. Alderflies are a group of "living fossils" with a long evolutionary history. The oldest fossil attributed to Sialidae dates back to the Early Jurassic period. Further, the global distribution of modern-day species shows a remarkably disjunctive pattern. However, due to the rareness of most species and scarcity of comprehensive taxonomic revisions, the phylogeny of Sialidae remains largely unexplored, and the present classification system is in great need of renewal. Here we reconstruct the first phylogeny for Sialidae worldwide based on the most comprehensive sampling and broadest morphological data ever presented for this group of insects. All Cenozoic alderflies belong to a monophyletic clade, which may also include the Early Jurassic genus †Dobbertinia, and the Late Jurassic genus †Sharasialis is their putative sister taxon. Two subfamilies of Sialidae are proposed, namely †Sharasialinae subfam. nov. and Sialidinae. Austrosialis is the sister of all other extant genera, an assemblage which comprises three monophyletic lineages: the Stenosialis lineage, the Ilyobius lineage, and the Sialis lineage. The revised classification of Sialidae is composed of 12 valid genera and 87 valid species. Ilyobius and Protosialis are recognized as valid generic names, while Nipponosialis is treated as a synonym of Sialis. Reconstruction of the ancestral area proposes a global distribution of alderflies in Pangaea before their diversification. The generic diversification of alderflies might have occurred before the breakup of Pangaea, but the divergence of some lineages or genera was probably promoted by the splitting of this supercontinent.