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

Abstract

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. The tribes Notiophilini, Opisthiini, Pelophilini, and 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 & Roux, 1989 = Boreonebria Jeannel, 1937; Patrobonebria Bänninger, 1923, Paranebria Jeannel, 1937 and Barbonebriola Huber & Schmidt, 2017 = Epinebriola Daniel & 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 & 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, particularly within subgenera Leistus and Evanoleistus of genus Leistus and the Nebria Complex of genus Nebria.

Evidence for niche conservatism in alpine beetles under a climate‐driven species pump model

“…In previous work using mitochondrial sequence data, Kavanaugh and Schoville (2009) estimated that the divergence of species in the N. vandykei complex ranged from 1.46% pairwise divergence at COI (N. v. vandykei to N. praedicta) to 0.24% (N. v. vandykei to N. v. wyeast), suggesting relatively recent evolutionary diversification. Although they are co-distributed, the two species complexes are not closely related to one another, comprising ~7.5% pairwise COI sequence divergence (Kavanaugh et al, 2021;Slatyer & Schoville, 2016). S1 for sample sizes).…”

“…The likelihood of ecological speciation can thus be contrasted to the alternative of mutation-order speciation (reproductive isolation by chance; Mani & Clarke, 1990;Schluter, 2009). The species rich ground beetle genus Nebria (Coleoptera: Carabidae) provides a useful system to explore the influence of the climate-driven species pump on micro-and macroevolutionary processes in a montane setting (Kavanaugh, 1978;Kavanaugh et al, 2021;Schoville et al, 2012). Previous studies have supported rapid ecological divergence in Nebria species complexes (Clarke et al, 2001;Palestrini et al, 2012;Schoville et al, 2012).…”

Molecular phylogeny and revision of species groups of Nearctic bombardier beetles (Carabidae, Brachininae, Brachinus ( Neobrachinus))

“…Bootstrap analyses for the four trees were conducted with 1000 replicates using IQ-TREE v. 1.6.10 (Nguyen et al 2015), as orchestrated by the CIPRES Science Gateway (Miller et al 2010). Support for and against clades were calculated for each species group and the subgenus Neobrachinus in Mesquite using its "Clade Frequencies in Trees" feature and bootstrap trees generated by IQ-TREE Kavanaugh et al 2021).…”