The caddisfly subfamily Drusinae BANKS comprises roughly 100 species inhabiting mountain ranges in Europe, Asia Minor and the Caucasus. A 3-gene phylogeny of the subfamily previously identified three major clades that were corroborated by larval morphology and feeding ecologies: scraping grazers, omnivorous shredders and filtering carnivores. Larvae of filtering carnivores exhibit unique head capsule complexities, unknown from other caddisfly larvae. Here we assess the species-level relationships within filtering carnivores, hypothesizing that head capsule complexity is derived from simple shapes observed in the other feeding groups. We summarize the current systematics and taxonomy of the group, clarify the systematic position of Cryptothrix nebulicola, and present a larval key to filtering carnivorous Drusinae. We infer relationships of all known filtering carnivorous Drusinae and 34 additional Drusinae species using Bayesian species tree analysis and concatenated Bayesian phylogenetic analysis of 3805bp of sequence data from six gene regions (mtCOI5-P, mtCOI3-P, 16S mrDNA, CADH, WG, 28S nrDNA), morphological cladistics from 308 characters, and a total evidence analysis. All analyses support monophyly of the three feeding ecology groups but fail to fully resolve internal relationships. Within filtering carnivores, variation in head setation and frontoclypeus structure may be associated with progressive niche adaptation, with less complex species recovered at a basal position. We propose that diversification of complex setation and frontoclypeus shape represents a recent evolutionary development, hypothetically enforcing speciation and niche specificity within filtering carnivorous Drusinae.
Two new species of the genus Drusus (Trichoptera, Limnephilidae, Drusinae) from the Western Balkans are described. Additionally, observations on the biodiversity and threats to the region’s endemic aquatic fauna are discussed. Drusus krpachi sp. n. is a micro-endemic of the Korab Mountains, Macedonia, and Drusus malickyi sp. n. is a micro-endemic of the Prokletije Mountains, Albania. Both new species are most similar to Drusus macedonicus but differ from the latter in the shape of segment IX, the shape of the tips of the intermediate appendages in lateral view, the shape of the inferior appendages, and the form and shape of the parameres. In addition, males of the European species of filtering carnivore Drusinae are diagnosed and illustrated, including Cryptothrix nebulicola McLachlan, Drusus chrysotus Rambur, Drusus discolor Rambur, Drusus macedonicus Schmid, Drusus meridionalis Kumanski, Drusus muelleri McLachlan, Drusus romanicus Murgoci and Botosaneanu, and Drusus siveci Malicky. These additions to the Western Balkan fauna demonstrate the significance of this region for European biodiversity and further highlight the importance of faunistic studies in Europe.
The taxonomy of the genera of Hydropsychinae is revised. The following genera are synonymized: Hydatomanicus Ulmer, 1951 and Hydatopsyche Ulmer, 1926 are synonymized with Hydromanicus Brauer, 1865; Streptopsyche Ross & Unzicker, 1977 is synonymized with Calosopsyche Ross & Unzicker, 1977; Symphitopsyche Ulmer, 1907, Mexipsyche Ross & Unzicker, 1977, Ceratopsyche Ross & Unzicker, 1977 and Herbertorossia Ulmer, 1957 are synonymized with Hydropsyche Pictet, 1834; and Plectropsyche Ross, 1947 is synonymized with Cheumatopsyche Wallengren, 1891. A new genus, Schmidopsyche Oláh & Schefter, new genus is established and the single species in the genus, S. rossi Oláh & Schefter, new species is described and illustrated, based on material collected by Schmid in Sikkim in 1959. The additional 61 new species are described and illustrated: India: Hydromanicus krsamgin Oláh & Barnard, new species; H. sacso Oláh & Johanson, new species; H. naraik Oláh & Barnard, new species; H. palnis Oláh & Barnard, new species; H. topali Oláh & Johanson, new species; Hydropsyche januha Oláh & Barnard, new species; H. kamenga Oláh & Schefter, new species; H. mizora Oláh & Schefter, new species; H. narayana Oláh & Johanson, new species; H. khasigiri Oláh & Barnard, new species; H. igunapali Oláh & Schefter, new species; H. kangra Oláh & Barnard, new species; H. kiogupa Oláh & Schefter, new species; H. minutanga Oláh & Johanson, new species; H. tritiyaha Oláh & Barnard, new species; H. ekaropa Oláh & Schefter, new species; H. yathetima Oláh & Schefter, new species; H. keralana Oláh & Barnard, new species; H. semkala Oláh & Johanson, new species; H. shillonga Oláh & Barnard, new species; Cheumatopsyche barakambra Oláh & Johanson, new species; C. mariannae Oláh & Johanson, new species; C. sandrae Oláh & Johanson, new species; Potamyia assamana Oláh & Schefter, new species. Vietnam: Hydromanicus demden Oláh & Johanson, new species; and Hydropsyche soinha Oláh & Johanson, new species (also from Malaysia); H. tanung Oláh & Johanson, new species; H. lomdom Oláh & Johanson, new species; H. sangbung Oláh & Johanson, new species; H. keoda Oláh & Johanson, new species; H. xenga Oláh & Johanson, new species; and Cheumatopsyche simayorum Oláh & Johanson, new species. Malaysia: Hydropsyche gekilara Oláh & Barnard, new species; H. tanua Oláh & Barnard, new species; and H. namea Oláh & Barnard, new species. Philippines: Hydropsyche meyi Oláh & Johanson, new species; Cheumatopsyche anonima Oláh & Johanson, new species; C. kuranishii Oláh & Johanson, new species; and C. montapo Oláh & Johanson, new species. Papua New Guinea: Hydropsyche flintorum Oláh & Johanson, new species; and H. carolae Oláh & Johanson, new species. Tibet: Hydropsyche gyantsana Oláh & Barnard, new species. Nepal: Hydropsyche nepalarawa Oláh & Johanson, new species; and Cheumatopsyche chitawana Oláh & Johanson, new species. China: Hydropsyche chengdu Oláh & Barnard, new species; and Cheumatopsyche yangmorseorum Oláh & Johanson, new species. Iran: Hydropsyche masula Oláh & Johanson, new species. New Caledonia: Orthopsyche anulmika Oláh & Johanson, new species. Japan: Cheumatopsyche okinawana Oláh & Johanson, new species; and C. tanidai Oláh & Johanson, new species. Taiwan: Cheumatopsyche davisi Oláh & Johanson, new species; and C. tungpa Oláh & Johanson, new species. Sri Lanka: Cheumatopsyche hasalaka Oláh & Johanson, new species. South Africa: Cheumatopsyche krugerana Oláh & Johanson, new species. Madagascar: Cheumatopsyche mafana Oláh & Johanson, new species; C. nondra Oláh & Johanson, new species; C. tiokalamkita Oláh & Johanson, new species; C. fianara Oláh & Johanson, new species; C. masiposa Oláh & Johanson, new species; C. ranoma Oláh & Johanson, new species; and C. rantsoa Oláh & Johanson, new species. The following species are synonymized: Hydropsyche kawamurai Tsuda, 1940 is synonymized with H. kozhantschikovi Martynov, 1924, Hydropsyche belos Malicky & Chantaramongkol, 2000 is synonymized with H. klimai Mey, 1999, Hydropsyche taiwanensis Mey, 1997 is synonymized with H. spinata Kobayashi, 1987, and Cheumatopsyche copiosa Kimmins, 1956 is synonymized with C. columnata Martynov, 1935. Recognized genera are divided into previously established and new species groups, defined by shared diagnostic characters. All Oriental species of Hydropsyche Pictet and Hydromanicus Brauer are grouped into different levels of species groups based on similar morphology in male genitalia.
The following new species are described from the west coast of Sumatra (Indonesia): Dipseudopsis gabiana and Pseudoneureclipsis complexa (Dipseudopsidae); Psychomyia selatana and Tinodes kerinciana (Psychomyiidae); Nyctiophylax anryutah, N. kupirah, and N. tapan (Polycentropodidae); Diplectrona dimagaha, D. tiguna, Hydropsyche pulakita, and Cheumatopsyche urhpirdana (Hydropsychidae); and Oecetis barisan, Setodes pantangulil, S. trikuthia, and S. wirhal (Leptoceridae). In addition, the following species are recorded from Sumatra for the first time: Chimarra briseis Malicky (Philopotamidae); Ecnomus hendersoni Mosely (Ecnomidae); Diplectrona dulitensis Kimmins and Hydropsyche butes Malicky & Chantaramongkol (Hydropsychidae); and Ganonema ochraceellum (McLachlan) (Calamoceratidae). Fourteen more species are recorded, but are previously known from Sumatra.
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