Geographical history of the central‐western Pacific black fly subgenus Inseliellum (Diptera: Simuliidae: Simulium) based on a reconstructed phylogeny of the species, hot‐spot archipelagoes and hydrological considerations

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Abstract
Aim: With six new species of subgenus Inseliellum Rubtsov recently described, a revised reconstructed phylogeny based on morphology is required. Geological history of islands where Inseliellum occurs, plus a cladistic analysis and hydrological considerations, provide the basis for a reconstructed geographical history of the species.Location: Inseliellum is widely distributed and occurs in Micronesia, Cook Islands and Polynesia. A single specimen is known from Tonga Islands.Methods: Maximum parsimony criteria using PAUP*, plus cytological information, were used to arrive at a preferred phylogenic reconstruction. Island ages of the hot spot archipelagoes involved are well known. The phylogeny was then compared with the palaeogeology. Information on evolution of running water habitats as islands age was incorporated into the biogeography.Results: Cladistic analysis of forty of the forty‐eight known Inseliellum species with Simulium (Nevermannia) neornatipes Dumbleton from New Caledonia and S. (Hebridosimulium) laciniatum Edwards from Fiji as outgroups, shows basal species and clades to be on widely separated older islands. In the Society Islands basal species are widely spread. Derived species, with morphological adaptations to deal with specialized habitats, are on younger islands (e.g. Tahiti), where a major species radiation has taken place. The reconstructed phylogeny indicates dispersal back to older islands, with minor subsequent species radiation.Main conclusions: Palaeogeological evidence provides a basis for postulating that Inseliellum entered the western Pacific area some 20 Ma, with the possibility that it rafted eastwards on proto‐Tonga Islands to the edge of southern‐central Pacific. Older Cook Islands were present at that time. Movement into the Marquesas Islands was not earlier than 6 Ma and into the Society Islands perhaps 8–10 Ma. Basal species with generalized habitat requirements would have found suitable habitats (inferred from hydrological postulates) on leaves in the original, small shaded streams. With erosion and valley development, in particular on Tahiti, species radiated into specialized habitats such as cascades. Rich seston and high velocity probably drove reduction of filtering fans in some clades. With collapse of the caldera and formation of large rivers at c. 0.9 Ma, S. exasperans Craig and S. tahitiense Edwards adapted to deep, swiftly flowing water, all indicative that specialized habitat availability drove species radiation in Inseliellum. In the Society Islands, dispersal of derived species back to the oldest western islands was not possible because erosion has removed suitable habitats.