Nesamblyops Jeannel 1937: 279

Sokolov, Igor M., 2023, Fourteen new species of the genus Nesamblyops Jeannel (Coleoptera: Carabidae: Anillini) from the South Island of New Zealand with redescription of the genus and description of a new subtribe, Zootaxa 5375 (2), pp. 151-192 : 164-165

Nesamblyops Jeannel 1937: 279 .

With character states of the subtribe and following combination of additional diagnostic characters:

Fixed setae. Primary head setae include a pair of clypeal, a pair of frontal, and two pairs of supraorbital setae. Postorbital setae lacking. Mentum with two pairs of long primary (paramedial and lateral) setae. Pronotum with two long primary lateral setae (midlateral and basilateral) on each side. Elytra with eight setae in umbilical series of pores, one pair of discal setae in ed6 position ( Figs 6A–B , ed6), with scutellar (ed2) and apical (ed8) setae.

Head. Anterior margin of clypeus (cl) straight. Clypeal “setulation” regular, consists of two pairs of setulae: one lateral (lse) and 1 apical (mse) on each side ( Figs 1A–B ). Apical setulae situated at the margin of clypeus close to its anterior angles. Frontal area flat without tubercle medially near frontoclypeal suture. Fronto-lateral carinae distinct and long.

Comments. The pattern of distribution of setulae (“setulation”) on clypeus might be of taxonomic importance in Anillini . Patterns can be irregular (small setulae randomly scattered across the clypeus) or regular (a few relatively long setulae whose positions follow bilateral symmetry). In the latter case, the position of particular setula is also important (at the edge or shifted interior to the middle of clypeus). Altogether, the pattern and positions of setulae may characterize lineages or groups of genera (cf. the irregular pattern of related Anillinus and Serranillus (not illustrated) in Fig. 1D with the regular pattern of related Pelodiaetodes Moore and Pelodiaetus Jeannel with setulae in mid-clypeal positions, Figs 1H–I ), certain genera (the irregular pattern of Zeanillus Jeannel , Fig. 1G , with the regular pattern of Geocharidius with setulae in mid-clypeal positions, Fig. 1F ), or particular species—some species of Geocharidius have mid-clypeal setulae even longer than fixed clypeal setae, thus having four long setae on clypeus like Bembidarenas Erwin and Tasmanitachoides Erwin of Bembidarenini ( Maddison et al. 2019).

Eyes. Each eye is either presented by one ommatidium with supported black pigment underneath or eyes absent ( Figs 1A–C , om, trom).

Antennae. From filiform to submoniliform, 11-segmented, extended to about posterior margin of pronotum.

Labrum. Labrum transverse and straight, entire anterior margin with six setae apically, increasing in size from the central pair outwards.

Labium. Labium with short acute mental tooth and acute epilobes, mentum and submentum split, with mentalsubmental suture. Glossal sclerite with shortly lobed paraglossae with two setae medio-apically.

Prothorax ( Figs 3A–B ). Pronotum cordiform, moderately convex, rectilinear constricted posteriorly, with narrow marginal gutter. Posterior margin of pronotum varies from rectilinear to slightly convex. Anterior angles wanted. Posterior angles rounded, nearly effaced, bearing basolateral seta at or far anterior to angles.

Elytra ( Figs 6A–B ).Elytra slightly subdepressed along suture, without visible interneurs and without longitudinal grooves. Humeri rounded, to form oblique angle with longitudinal axis of body. Elytral basal margination lacking. Apical half of elytra without subapical sinuation. Sutural angle of elytron shortly rounded, making apices of elytra very slightly dehiscent.

Hind wings. Absent.

Legs. Legs of moderate length, not elongate. Prothoracic legs of males with first two tarsomeres dilated apicolaterally with 1–3 rows of oval articulo-setae (as) ( Stork 1980) on the ventral surface ( Figs 7A–B ).

Abdominal ventrites ( Figs 5A–D ). Five visible abdominal ventrites: 2 nd ventrite longest, 2.8–3.5 times longer than 3 rd or 4 th, 3 rd and 4 th equal in length; the last, 5 th, 1.4–1.6 times longer than 4 th. Abdomen with intercoxal process of the ventrite 2 of an acute triangular shape.

Male genitalia ( Figs 14–16 ). Median lobe of aedeagus anopic, elongate, slightly twisted and moderately arcuate. Apex of median lobe various, often with enlarged tip. Copulatory sclerites of median lobe characterized by weak development of sclerotization and heavily sclerotized only narrowly, presumably along folds or invaginations of the internal sac. In the most common case, weakly sclerotized fields of copulatory sclerites are observed in lateral view as one formation at basal half of the shaft. Typically, two components of this formation are more conspicuous than others, and can be seen in many species. The first component, more dorsal in position, is represented by few elongate and strongly sclerotized structures forming an isosceles triangle, which stretches along the longitudinal axis of the shaft. Congruent sides of this triangle directed apically and, thus, forming a V-shaped contour (further V-contour, Fig. 14L , Vc). The second component, more medial in position, is represented by a weakly to moderately sclerotized field with a narrowly and strongly sclerotized edge having the reversed C-contour (further rC-sclerite, Fig. 14L , rCs). Both components vary in their relative sizes, proportions, and degree of sclerotization giving a particular species a unique pattern of sclerotized contours. In some cases, components are merged and form species specific plates. Some species have only one sclerotized field: in these cases, presumably the rC-sclerite is lacking. Finally, some species develop additional sclerotized structures in apical half of the shaft. Spines of internal sac absent in all examined species. Parameres tri- or bisetose. Ring sclerites trianguloid with an ovoid basal part, either acute or rounded handle-like extension varying in length among species.

Female internal genitalia ( Fig. 18 ). Spermatheca sclerotized, in a form of the light bulb shape, with ball-like distal part and tube-like proximal part representing the continuation of the unsclerotized and short spermathecal duct. Proximal part of spermatheca bears a long spermathecal gland. Both gland and duct slightly longer than the length of spermatheca.

Geographical distribution. Endemic of New Zealand including the Antipodes Islands.

Included taxa. At present the genus includes 24 species, fourteen of which are described below.

Figures

FIGURE 1. SEM illustrations of structural features of heads of Anillini: A–C—subtribe Nesamblyopina; A—Nesamblyops oreobius (Broun); B—Nesamblyops subcaecus (Sharp); C—Nesamblyops magnificus, n.sp.; D—Anillinus alleni Sokolov and Carlton; E—Anillinus acutipennis Sokolov and Reddell; F—Geocharidius jalapensis Sokolov and Kavanaugh; G—Zeanillus punctigerus (Broun); H—Pelodiaetodes nunni Sokolov; I—Pelodiaetus sulcatipennis Jeannel. Legend: cl—clypaeus, lse— lateral clypeal setula, mse—medial clypeal setula, om—ommatidium, trom—trace of disappeared ommatidium. Scale = 0.1 mm.

FIGURE 5. Digital illustrations of structural features of pterothorax of Anillini, ventral aspect: A–D—subtribe Nesamblyopina; A—Nesamblyops oreobius (Broun); B—Nesamblyops subcaecus (Sharp); C—Nesamblyops brouni Sokolov; D—Nesamblyops viator n.sp.; E—Anillinus langdoni Sokolov and Carlton; F—Anillinus Cherokee Sokolov and Carlton; G—Geocharidius longinoi Sokolov and Kavanaugh; H—Geocharidius disjunctus Sokolov and Kavanaugh; I—Zapotecanillus longinoi Sokolov; J—Zeanillus pellucidus Sokolov; K—Zeanillus pallidus (Broun); L—Zeanillus nunni Sokolov; M—Pelodiaetodes insularis Sokolov; N—Pelodiaetodes nunni Sokolov; O—Pelodiaetodes moorei Sokolov; P—Pelodiaetus sulcatipennis Jeannel.Legend: mtes—metendosternite. Scales intentionally omitted.

FIGURE 6. SEM illustrations of structural features of elytra of Anillini, left lateral aspect: A–B—subtribe Nesamblyopina; A—Nesamblyops oreobius (Broun); B—Nesamblyops subcaecus (Sharp); C—Anillinus carolinae Casey; D—Anillinus alleni Sokolov and Carlton; E—Serranillus donovani (Jeannel); F—Geocharidius jalapensis Sokolov and Kavanaugh; G—Zeanillus punctigerus (Broun); H—Pelodiaetodes nunni Sokolov; I—Pelodiaetus nunni Sokolov. Legend: ed2—scutellar seta; ed6— discal seta, ed8—apical seta; eo1…eo9—setae 1–9 of umbilical series of pores. Scale = 0.2 mm.

FIGURE 7. SEM illustrations of structural features of protibia of Anillini, dorsal (E) and ventral (A–D, F–H) aspects: A– B—subtribe Nesamblyopina; A—Nesamblyops oreobius (Broun); B—Nesamblyops subcaecus (Sharp); C—Anillinus lescheni Sokolov and Carlton; D—Geocharidius comayaguanus Sokolov and Kavanaugh; E—Geocharidius zullini Vigna Taglianti; F—Zeanillus brouni Sokolov; G—Pelodiaetodes nunni Sokolov; H—Pelodiaetus sulcatipennis Jeannel. Legend: as—articulosetae; psr—posterior setal row. Scale = 0.05 mm.

FIGURE 14. Line drawings of male genitalia of New Zealand Nesamblyops species. N. canaanensis (Canaan, Nelson, SO): A—left paramere, left lateral aspect, B—right paramere, right lateral aspect, C—median lobe, right lateral aspect. N. disjunctus (Fletchers Creek, Buller, SO): D—left paramere, left lateral aspect, E—right paramere, right lateral aspect, F—median lobe, right lateral aspect. N. hobbit (Mount Domett, Nelson, SO): G—left paramere, left lateral aspect, H—right paramere, right lateral aspect, I—median lobe, right lateral aspect. N. karamea (Karamea River Gorge, Nelson, SO): J—left paramere, left lateral aspect, K—right paramere, right lateral aspect, L—median lobe, right lateral aspect. N. kuscheli (Flora Hut, Mount Arthur, Nelson, SO): M—left paramere, left lateral aspect, N—right paramere, right lateral aspect, O—median lobe, right lateral aspect. N. magnificus (Allison Reserve, Akatore, Dunedin, SO): P—left paramere, left lateral aspect, Q—right paramere, right lateral aspect, R—median lobe, right lateral aspect. Legend: rCs—reversed C-contour; Vs—V-contour. Scale bar = 0.1 mm.

FIGURE 15. Line drawings of male genitalia of New Zealand Nesamblyops species. N. montanus (Relax Shelter, Mount Robert Ridge, Nelson, SO): A—left paramere, left lateral aspect, B—right paramere, right lateral aspect, C—median lobe, right lateral aspect. N. moorei (Kangaroo Creek, Mawhera Forest, Buller, SO): D—left paramere, left lateral aspect, E—right paramere, right lateral aspect, F—median lobe, right lateral aspect. N. ovipennis (Mount Arthur, Nelson, SO): G—left paramere, left lateral aspect, H—right paramere, right lateral aspect, I—median lobe, right lateral aspect. N. rotundicollis (Onekaka, Nelson, SO): J—left paramere, left lateral aspect, K—right paramere, right lateral aspect, L—median lobe, right lateral aspect. N. solitarius (Italian Creek, Capleston, Buller, SO): M—left paramere, left lateral aspect, N—right paramere, right lateral aspect, O—median lobe, right lateral aspect. N. subacecus (Margarets Tarn Lake, Arthur’s Pass, Mid Canterbury, SO): P—left paramere, left lateral aspect, Q—right paramere, right lateral aspect, R—median lobe, right lateral aspect. Scale bar = 0.1 mm.

FIGURE 16. Line drawings of male genitalia of New Zealand Nesamblyops species. N. subrufus (Dublin Terrace, Upper Buller Gorge, Nelson, SO): A—left paramere, left lateral aspect, B—right paramere, right lateral aspect, C—median lobe, right lateral aspect. N. viator (Secretary Island, Grono Bay, Fjordland, SO): D—left paramere, left lateral aspect, E—right paramere, right lateral aspect, F—median lobe, right lateral aspect. N. victoriae (Capleston, Victoria Range, Buller, SO): G—left paramere, left lateral aspect, H—right paramere, right lateral aspect, I—median lobe, right lateral aspect. Scale bar = 0.1 mm.

FIGURE 18. Line drawings of spermatheca of New Zealand Nesamblyops species. A—N. magnificus (Allison Reserve, Akatore, Dunedin, SO), B—N. rotundicollis (Parahamoi, Nelson, SO), C—N. subcaecus (Lake Kaniere Scenic Reserve, Mount Tuhua, Westland, SO). Scale = 0.1 mm.

FIGURE3. SEM illustrationsof structuralfeatures of pronota ofAnillini:A–B—subtribe Nesamblyopina;A—Nesamblyops oreobius (Broun); B—Nesamblyops subcaecus (Sharp); C—Anillinus alleni Sokolov and Carlton; D—Anillinus forthoodensis Sokolov andReddell;E—Serranillusjeanneli Barr; F—Geocharidius balini Sokolov and Kavanaugh; G—Zeanillus punctigerus (Broun); H—Pelodiaetodes nunni Sokolov; I—Pelodiaetus nunni Sokolov. Legend: ast—setae of anterior margin of pronotum, bst—setae of basal margin of pronotum. Scale = 0.1 mm.