DOI: 10.5061/dryad.6wwpzgmxn
title: Phylogeny of the supertribe Nebriitae (Coleoptera: Carabidae) based on analyses of DNA sequence data

description: The phylogeny of the carabid beetle supertribe Nebriitae is inferred from
 analyses of DNA sequence data from eight gene fragments including one
 nuclear ribosomal gene (28S), four nuclear-protein coding genes (CAD,
 topoisomerase 1, PEPCK and wingless) and three mitochondrial gene
 fragments (16S + tRNA-Leu + ND1, COI (“barcode” region) and COI (“Pat/Jer”
 region)). Our taxon sample included 264 exemplars representing 241 species
 and subspecies (25% of the known nebriite fauna), 39 of 41 currently
 accepted genera and subgenera (all except Notiokasis and
 Archileistobrius), and eight outgroup taxa. Separate maximum likelihood
 (ML) analyses of individual genes, combined ML analyses of nuclear,
 nuclear protein-coding and mitochondrial genes, and combined ML and
 Bayesian analyses of the eight-gene-fragment matrix resulted in a
 well-resolved phylogeny of the supertribe, with most nodes in the tree
 strongly supported. Within Nebriitae, 167 internal nodes of the tree (out
 of the maximum possible 255) are supported by maximum-likelihood bootstrap
 values of 90% or more. Tribes Notiophilini, Opisthiini, Pelophilini,
 Nebriini are well supported as monophyletic but relationships among these
 are not well resolved. Nippononebria is a distinct genus more closely
 related to Leistus than Nebria. Archastes, Oreonebria, Spelaeonebria, and
 Eurynebria, previously treated as distinct genera by some authors, are all
 nested within a monophyletic genus Nebria. Within Nebria, four major
 clades are recognized: (1) the Oreonebria Series, including eight
 subgenera arrayed in two subgeneric complexes (the Eonebria and Oreonebria
 Complexes); (2) the Nebriola Series, including only subgenus Nebriola; (3)
 the Nebria Series, including ten subgenera arrayed in two subgeneric
 complexes, the Boreonebria and Nebria Complexes, with the latter further
 subdivided into three subgeneric subcomplexes (the Nebria, Epinebriola and
 Eunebria Subcomplexes)); and (4) the Catonebria Series, including seven
 subgenera arrayed in two subgeneric complexes (the Reductonebria and
 Catonebria Complexes). A strong concordance of biogeography with the
 inferred phylogeny is noted and some evident vicariance patterns are
 highlighted. A revised classification, mainly within the Nebriini, is
 proposed to reflect the inferred phylogeny. Three genus-group taxa
 (Nippononebria, Vancouveria and Archastes) are given revised status and
 seven are recognized as new synonymies (Nebriorites Jeannel, 1941 and
 Marggia Huber, 2014 = Oreonebria Daniel, 1903; Pseudonebriola Ledoux and
 Roux, 1989 = Boreonebria Jeannel, 1937; Patrobonebria Bänninger, 1923,
 Paranebria Jeannel, 1937 and Barbonebriola Huber and Schmidt, 2017 =
 Epinebriola Daniel and Daniel, 1904; and Asionebria Shilenkov, 1982 =
 Psilonebria Andrewes, 1923). Six new subgenera are proposed and described
 for newly recognized clades: Parepinebriola Kavanaugh subgen. nov. (type
 species: Nebria delicata Huber and Schmidt, 2017), Insulanebria Kavanaugh
 subgen. nov. (type species: Nebria carbonaria Eschscholtz, 1829),
 Erwinebria Kavanaugh subgen. nov. (type species Nebria sahlbergii Fischer
 von Waldheim, 1828), Nivalonebria Kavanaugh subgen. nov. (type species:
 Nebria paradisi Darlington, 1931), Neaptenonebria Kavanaugh subgen. nov.
 (type species: Nebria ovipennis LeConte, 1878) and Palaptenonebria
 Kavanaugh subgen. nov. (type species: Nebria mellyi Gebler, 1847). Future
 efforts to better understand relationships within the supertribe should
 aim to expand the taxon sampling of DNA sequence data, particulary within
 subgenera Leistus and Evanoleistus of genus Leistus and the Nebria Complex
 of genus Nebria.
description: DNA was sequenced using both PCR/Sanger sequencing and Illumina
 sequencing.  Phylogenetic trees included in the submission were inferred
 using maximum likelihood in RAxML.