Nebria Palaptenonebria 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 Palaptenonebria Kavanaugh, subgen. nov.

Catonebria Shilenkov, 1975 (in part); = " Nebria mellyi " group sensu Ledoux and Roux 2005.

Type species.

Nebria mellyi Gebler, 1847:312, by present designation.

Diagnosis.

Body size small, SBL = 7.2 to 9.5 mm. Head width medium or slightly broadened, not constricted behind eyes; vertex with a pair of paramedial pale spots on vertex and one pair of supraorbital setae. Eyes slightly to moderately reduced in size, moderately convex. Antennae medium length to slightly elongate; antennal scape with one subapicodorsal seta (or two in Nebria roddi Dudko & Shilenkov, 2001 and Nebria baenningeri korgonica Dudko & Shilenkov, 2001); 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 moderately cordate in dorsal aspect, with lateral margination shallowly to moderately impressed and markedly narrow at middle; one midlateral seta and one basolateral seta present on each side. Elytral intervals smooth, slightly to moderately flattened, without macrosculpture, elytral interval 3 with one to seven setiferous pores, intervals 5 and 7 with zero to two setiferous pores; umbilicate series comprised of eight to ten setae; intervals bearing setae faintly to moderately catenate. Hindwings reduced to short, slender strap-like vestiges. Metasternum moderately to markedly short. Metepisterna smooth, impunctate. Protarsomeres 1-3 expanded in males; mesotarsomeres 2-4 longer than their apical width; tarsi dorsally glabrous (or with few sparse and fine setae in Nebria mellyi Gebler, 1847 specimens). Abdominal sternites IV-VI with from two to five or more pairs of posterior paramedial setae and with zero to three pairs of anterior paralateral setae; sternite VII with one or two pairs of paramedial apical setae in males, two to three or more pairs in females. Median lobe of male aedeagus sclerotized dorsally at least to midlength on shaft, symmetrical to slightly deflected right 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 to moderately compressed in cross-section, right lateral face with a shallow longitudinal groove or shallow indentation; apical orifice moderately (in Nebria lyubechanskii Dudko, 2008 specimens only) or extremely deflected right. Right paramere narrow and slightly shortened. Female hemisternites VIII with basal apodeme faintly to moderately emarginated. Female valvifer without vestiture; gonopods VIII fused to dorsomedial bases of gonocoxae; gonocoxae with ventral diagonal row of setiform setae and mediodorsal row of setae present, mediodorsal row oriented parallel to longitudinal axis of gonocoxa. Bursa copulatrix without dorsal sclerites in vestibular chamber and without a posterodorsal extension from its longitudinal axis in lateral aspect; spermathecal chamber broadly cordate in dorsal aspect, without dorsal sclerites (in most species) or with a simple, broad and convoluted plate anterior to spermathecal duct insertion (in Nebria sajana sarlyk Dudko & Shilenkov, 2001, N. s. dubatolovi Dudko & Shilenkov, 2001 and N. s. sitnikovi Dudko & Shilenkov, 2001); spermathecal duct slightly short in length and of uniform diameter throughout or nearly so, inserted basodorsally on spermathecal chamber or on a dorsal accessory lobe of that chamber (in Nebria sajana sajana Dudko & Shilenkov, 2001); spermathecal reservoir of medium length.

Etymology.

The subgeneric epithet is a noun of feminine gender and combination of the Greek word, palaios, meaning old, in reference to the Palearctic (Old World) distribution of the group, aptenos, meaning unable to fly, and the genus name, Nebria , in reference to observation that all known members of this clade have extremely reduced hindwings incapable of supporting flight.

Remarks.

As for members of subgenus Catonebria Neaptenonebria , those of this group are easily identified as members of the Catonebria Complex using the Ledoux and Roux’s (2005) key to subgenera. Features that distinguish them from members of subgenus Catonebria Nivalonebria have been discussed above for that taxon. We cannot suggest any external morphological feature that consistently and satisfactorily distinguishes all members of all species of Palaptenonebria from all Catonebria members, and there are few uniquely distinguishing internal features evident as well. The hindwings are more highly reduced (to short, slender strap-like vestiges) in all Palaptenonebria members than in any Catonebria member, but are similar in size to those of Neaptenonebria species. Internally, Palaptenonebria males have the right paramere of the aedeagus narrowed and slightly shorter than those of Catonebria males, but, again, similar in form to those in Neaptenonebria males. All males of Palaptenonebria species have the right lateral face of the mid-shaft region of the aedeagal median lobe modified to some extent. There is at least a shallow longitudinal depression or groove present in this region, and in N. mellyi that groove is deeply impressed and deepest and most sharply delineated basally. However, in no males of this clade is there an invaginated, basally-directed pouch as seen in all Neaptenonebria males. As noted above, a groove or depression similar to that seen in Palaptenonebria males is found also in Catonebria males of the Nebria gebleri group, as well as of N. metallica and Nebria labontei . In Palaptenonebria females, the mediodorsal row of setae on the gonocoxae is present and oriented parallel to the longitudinal axis of the gonocoxa rather than obliquely in relation to the longitudinal axis as in Neaptenonebria females. See the Discussion section above for Neaptenonebria Neaptenonebria for a few additional distinguishing features or trends.

Known distribution and diversity.

The geographical range of this clade is restricted to the Altai-Sayan region of southern Siberia, including the Altai, Western Sayan and Tannu-Ola Mountain systems, and presently includes six taxa ranked as species and another eight ranked as subspecies. Dudko and Shilenkov (2001) provided an excellent revision of this group.