title: Species delimitation in northern European water scavenger beetles of the genus Hydrobius (Coleoptera, Hydrophilidae).
abstract: The chiefly Holarctic Hydrobius species complex (Coleoptera, Hydrophilidae) currently consists of Hydrobius arcticus Kuwert, 1890, and three morphological variants of Hydrobius fuscipes (Linnaeus, 1758): var. fuscipes, var. rottenbergii and var. subrotundus in northern Europe. Here molecular and morphological data are used to test the species boundaries in this species complex. Three gene segments (COI, H3 and ITS2) were sequenced and analyzed with Bayesian methods to infer phylogenetic relationships. The Generalized Mixed Yule Coalescent (GMYC) model and two versions of the Bayesian species delimitation method BPP, with or without an a priori defined guide tree (v2.2 & v3.0), were used to evaluate species limits. External and male genital characters of primarily Fennoscandian specimens were measured and statistically analyzed to test for significant differences in quantitative morphological characters. The four morphotypes formed separate genetic clusters on gene trees and were delimited as separate species by GMYC and by both versions of BPP, despite specimens of Hydrobius fuscipes var. fuscipes and Hydrobius fuscipes var. subrotundus being sympatric. Hydrobius arcticus and Hydrobius fuscipes var. rottenbergii could only be separated genetically with ITS2, and were delimited statistically with GMYC on ITS2 and with BPP on the combined data. In addition, six or seven potentially cryptic species of the Hydrobius fuscipes complex from regions outside northern Europe were delimited genetically. Although some overlap was found, the mean values of six male genital characters were significantly different between the morphotypes (p < 0.001). Morphological characters previously presumed to be diagnostic were less reliable to separate Hydrobius fuscipes var. fuscipes from Hydrobius fuscipes var. subrotundus, but characters in the literature for Hydrobius arcticus and Hydrobius fuscipes var. rottenbergii were diagnostic. Overall, morphological and molecular evidence strongly suggest that Hydrobius arcticus and the three morphological variants of Hydrobius fuscipes are separate species and Hydrobius rottenbergii Gerhardt, 1872, stat. n. and Hydrobius subrotundus Stephens, 1829, stat. n. are elevated to valid species. An identification key to northern European species of Hydrobius is provided.
authors.affiliation: Department of Biology, Centre for Biodiversity Dynamics, NTNU Norwegian University of Science and Technology, 7491 Trondheim, Norway; Department of Natural History, NTNU University Museum, 7491 Trondheim, Norway.
authors.forename: Erlend I
authors.initials: EI
authors.lastname: Fossen
authors.affiliation: Department of Natural History, NTNU University Museum, 7491 Trondheim, Norway.
authors.forename: Torbjørn
authors.initials: T
authors.lastname: Ekrem
authors.affiliation: Department of Ecology and Environmental Science, University of Umeå, S-901 87 Umeå, Sweden.
authors.forename: Anders N
authors.initials: AN
authors.lastname: Nilsson
authors.affiliation: Department of Zoology, Swedish Museum of Natural History, Box 50007, SE-10405 Stockholm, Sweden.
authors.forename: Johannes
authors.initials: J
authors.lastname: Bergsten
keywords: BPP; Bayesian; DNA barcoding; Fennoscandia; GMYC; checklist; cryptic species; genitalia; guide tree; identification key; integrative taxonomy; molecular phylogeny; morphology; morphometrics; species boundaries; species complex; taxonomy