Phylogeny of the supertribe Nebriitae (Coleoptera: Carabidae) based on analyses of DNA sequence data

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.