Nebria Nivalonebria Kavanaugh,, 2021

Kavanaugh, David H., Maddison, David R., Simison, W. Brian, Schoville, Sean D., Schmidt, Joachim, Faille, Arnaud, Moore, Wendy, Pflug, James M., Archambeault, Sophie L., Hoang, Tinya & Chen, Jei-Ying, 2021, Phylogeny of the supertribe Nebriitae (Coleoptera, Carabidae) based on analyses of DNA sequence data, ZooKeys 1044, pp. 41-152 : 41

Subgenus Nebria Nivalonebria Kavanaugh, subgen. nov.

Nebria ovipennis = " Nebria ovipennis group" (in part) sensu Lindroth 1961

Nakanebria Ledoux & Roux, 2005 (in part)

Type species.

Nebria paradisi Darlington, 1931:24, by present designation.

Diagnosis.

Body size small to medium, SBL = 8.7 to 11.3 mm. Head markedly broadened, not constricted behind eyes; vertex with a single medial pale area and a single pair of supraorbital setae. Eyes moderate in size, moderately convex. Antennal scape with one subapicodorsal seta; antennomeres 3 and 4 not laterally compressed, without extra setae. Labrum with three pairs of apical setae. Maxillary stipes typical for genus, with setae inserted flush on smooth surface. Penultimate labial palpomere with three setae. Pronotum with one midlateral and one basolateral seta present on each side. Elytral intervals smooth, without macrosculpture, interval 3 with one to seven setiferous pores, interval 5 with zero to two and interval 7 with zero to two or more setiferous punctures, intervals with setae only faintly catenate; elytral medial margins diverge slightly and arcuate from the midline apically. Hindwings reduced to short, slender strap-like vestiges. Metepisterna smooth, impunctate. Protarsomeres 1-3 expanded in males; mesotarsomeres 2-4 longer than their apical width; tarsi dorsally glabrous. Abdominal sternites IV-VI with one to four pairs of posterior paramedial setae, without paralateral setae. Median lobe of male aedeagus sclerotized dorsally at least to midlength on shaft, symmetrical in dorsal aspect; basal bulb expanded, quadrate, broadly open basally and closed dorsally, without a sagittal aileron present at base or with only a lightly sclerotized collar; mid-shaft parallel-sided in lateral aspect, slightly compressed in cross-section, with right lateral face unmodified; apical orifice markedly deflected right. Right paramere slender and very long. Female valvifers without vestiture; gonopods VIII fused to dorsomedial bases of gonocoxae; gonocoxae with ventral diagonal row of setiform setae and mediodorsal row of setae present. Bursa copulatrix with a pair of large dorsal paramedial sclerotized plates in vestibular chamber, with its longitudinal axis moderately sigmoid dorsally in lateral aspect; spermathecal chamber broadly cordate in dorsal aspect, without dorsal sclerites (in some N. paradisi ) or with a small midline plate (other N. paradisi ) or large and broad dorsal plate (in N. turmaduodecima ); spermathecal duct very long and of uniform diameter throughout or nearly so, inserted basodorsally on spermathecal chamber; spermathecal reservoir of medium length.

Etymology.

The subgeneric epithet is a noun of feminine gender and a combination of the Latin word, Nebria nivalis , meaning snow, and the genus name, Nebria , in reference to the occurrence of all known members of this clade in the vicinity of montane snowfields and glaciers.

Remarks.

Members of this subgenus can be distinguished from those of subgenus Nivalonebria Nakanebria , to which they were assigned by Ledoux and Roux (2005), by several external and internal morphological features. Externally, the head is markedly broad, the medial margins of the elytra are slightly and arcuately divergent apically, and the ventroapical margin of metatarsomere 4 is markedly lobate laterally in members of Nivalonebria species, whereas the head is of moderate width, the medial margins of the elytra are sinuately divergent apically, and the ventroapical margin of metatarsomere 4 is extremely lobate laterally in Nakanebria members. Internally, the apical orifice of the median lobe of male genitalia is markedly deflected right in Nivalonebria members, only slightly so in Nakanebria members. Females of Nivalonebria species have valvifers without vestiture, a bursa copulatrix with a pair of large dorsal paramedial sclerotized plates in the vestibular chamber and with its longitudinal axis moderately sigmoid dorsally in lateral aspect and a spermathecal duct markedly long and inserted basodorsally on the chamber. In contrast, Nakanebria females have valvifers with vestiture, a bursa copulatrix without sclerotized plates in the vestibular chamber and with its longitudinal axis only slightly deflected apicodorsally, and a spermathecal duct slightly to markedly short and inserted medioventrally on the chamber. Nivalonebria members are distinguished from those of the other subgenera in the Catonebria complex externally in having a single medial pale area on the vertex (as opposed to a distinct pair of paramedial pale spots as seen in the other subgenera) and the medial margins of the elytra slightly and arcuately divergent apically. In members of Neaptenonebria , Palaptenonebria and all Catonebria (except Nebria baumanni Kavanaugh, 2015) the medial margins of the elytra diverge apically and a distinct angle. Internally, they are distinguished in having males with the right paramere of the aedeagus very long and slender and females with a bursa copulatrix with a pair of large dorsal paramedial sclerotized plates in the vestibular chamber and a very long spermathecal duct. Males of Neaptenonebria , Palaptenonebria and Catonebria have the right paramere slightly or moderately shorter, and females have a bursa without sclerites in the vestibular chamber and with a shorter spermathecal duct. The right lateral face of the mid-shaft of the median lobe of Nivalonebria males is smoothly convex and lacks any of the depressions or invaginations seen in this region in Neaptenonebria , Palaptenonebria , and some Catonebria males.

Known distribution and diversity.

The geographical range of this clade includes two disjunct areas of western North America, each with a single endemic species: (1) the northern Cascade Range of Washington and northern Oregon, from North Cascades National Park in the north to Mount Hood in the south (with N. paradisi endemic to that area); and (2) the Trinity Alps of northwestern California (with N. turmaduodecima endemic there).