Research Article |
Corresponding author: Charles Huber ( charles.huber@nmbe.ch ) Academic editor: Christoph Germann
© 2020 Charles Huber, Peer Hajo Schnitter.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Huber C, Schnitter PH (2020) Nebria (Pseudonebriola) tsambagarav sp. nov., a new alpine species from the Mongolian Altai (Coleoptera, Carabidae). Alpine Entomology 4: 29-38. https://doi.org/10.3897/alpento.4.50408
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Nebria (Pseudonebriola) tsambagarav sp. nov. is described from an alpine altitude in the Mongolian Altai. The new species is separated from other two Pseudonebriola species from the Mongolian Altai Mountain range, N. kerzhneri and N. medvedevi, by morphometric and morphological analyses. The new species and its habitat are illustrated, the subgeneric key is amended, and a distribution map is given.
carabids, morphometry, multivariate ratio analysis, taxonomy, distribution, high altitude
The subgenus Pseudonebriola Ledoux & Roux, 1989, genus Nebria Latreille, 1802 comprises 13 species and two subspecies (
Up to now four Pseudonebriola species are known from the Altai Mountains, a border mountain range common to Kazakhstan, Russia, China, and Mongolia: N. kaszabi Shilenkov, 1982 from the Kazakhstan and Russian part, N. stanislavi Dudko & Matalin, 2002, an endemic species in the Russian Altai, N. medvedevi Shilenkov, 1982 from the Mongolian (and Russian) Altai, and N. kerzhneri Shilenkov, 1982, endemic to the Gobi Altai, the Southeastern continuation of the Mongolian Altai (
Pseudonebriola specimens were collected by the junior author on the occasion of an expedition in 2016 to the Tsambagarav uul, Mongolian Altai, Northwestern Mongolia. The Tsambagarav Mountain is part of the central Mongolian Altai bordering the Great Lakes Depression and is separated from the main ridges of the Mongolian Altai by a straight tectonogenic hollow (
The edeagi and gonocoxae were dissected and dehydrated in alcohol and xylene, and finally embedded in Fluka DPX Mountant on transparent mounting cards.
For a morphometric analysis we measured eleven characters (Table
Characters measured in the Nebria kerzhneri species complex for use in a morphometric analysis.
Abbreviation | Character | Definition | Conversion factor |
eye.l | eye length | length of eye | 0.074 |
ant.w | antennomere 1 width | maximum width of antennomere 1 | 0.074 |
ant.l | antennomere 1 length | length of antennomere 1 | 0.074 |
ely.w | elytra width | maximum width of elytra | 0.01 |
ely.l | elytron length | length of elytron from basal margin to apex | 0.01 |
fro.w | frons width | width of frons between the eyes | 0.026 |
hea.w | head width | maximum width of the head (over eyes) | 0.026 |
pra.w | pronotum anterior width | width between the anterior angles of the pronotum | 0.026 |
prm.w | pronotum maximum width | maximum width of the pronotum | 0.026 |
prm.l | pronotum median length | median length of the pronotum | 0.026 |
prp.w | pronotum posterior width | width between the posterior angles of the pronotum | 0.026 |
The photographs were taken with a digital camera Leica MC 190 HD using a motorised focusing drive. The pictures are composites processed using the Leica Application Suite X (LAS X) 3.6.0.20104 software, and were retouched using Adobe Photoshop version 10.0.1. The median lobes of the type specimens of both N. kerzhneri and N. medvedevi were originally glue-fixed on mounting cards. The photographs of the median lobes were taken in that dried situation. The habitus photograph was taken by a digital microscope Keyence VHX-2000 with the objective VH-Z20R.
The distribution map (Fig.
Collections examined with acronyms:
cBK private collection of I.A. Belousov and I.I. Kabak, St Petersburg, Russia.
cSCHN collection Peer Schnitter, Halle, Germany.
For abbreviations of morphological characters see Tab.
Type material examined [with authors’ remarks or supplements in square brackets]:
N. medvedevi
: Holotype ♂ and 1 paratype ♂: [red label] Holotype [and Paratype respectively; printed, in Latin letters], Nebria medvedevi sp. nov. Shilenkov det. 1981 [handwritten, in Latin letters] // [white label] 7. 7. 1980; MNR Kobdosk. aim. Mongolsk. Altai, per. Ulan-Daba [47°23.96’N; 91°12.23’E], G. Medvedev [handwritten, in Cyrillic letters] // 3000–3200 m, под камнями у снежниов [handwritten, in Cyrillic letters] (
N. kerzhneri
: Holotype ♂ [red label]: Holotypus [printed, in Latin letters], Nebria kerzhneri sp. nov. Shilenkov det. 1980 [handwritten, in Latin letters] // [white label] 28. 6. 1973, Gobi-Altai Aimak, Dschargalan [46°58.59’N; 95°55.07’E], в старыx ходах рогоxвоста [handwritten, in Cyrillic letters]. V. Yanovskiy [printed, in Cyrillic letters] (
Additional material examined:
N. kaszabi
: 2 ♀ Vostochno-Kasachst., Marka-Kul, above Urunkhaika, 2400 m, 25. 6. 1986, leg. Shilenkov [in Cyrillic letters] // Nebria kaszabi Shil., det Shilenkov 1994 (
Additional localities of non-examined specimens used for the distribution map (Fig.
N. medvedevi : In June 1998 A. Matalin and D. Fedorenko collected N. medvedevi specimens for the first time in a Russian area: Ukok Highland, Teplyy Klyuch, 2640 m [49°07.703’N; 87°26.605’E].
N. kerzhneri
: Paratypes: Hasagt-Khairkhan, 15 km South of Jargalan, 14.8.1970, leg. Kerzhner [46°47.42’N; 95°48.23’E]. Taishirin-Ula, 15 km Southeastern of Altai [Altai = village in the Northwestern part of the Gobi Altai: 46°12.96’N; 96°27.66’E], 20.8.1970, leg. Emelyanov (
N. stanislavi
: Central Altai, Eastern slope of Mt. Kyzylart [50°32.20’N; 87°12.28’E], spring of river Achin, 2100–2300 m (
The scatterplots of the first scape PC against isosize of the four OTUs do not overlap and show no sign of allometry (Fig.
The LDA ratio extractor provides the best ratios to separate the groups. The best two ratios to discriminate the tsambagarav unit from the sample of the two kerzhneri and medvedevi units (ker+med in Fig.
Furthermore, we examined eight specimens of N. (P.) kaszabi, a member of the sajanica species group, as a geographical and taxonomic outgroup. The LDA extractor provides the same ratio (the antennal scape length/eye length) as best ratio to separate the tsambagarav and kaszabi units, supported by the second best ratio of elytral length/head width (Fig.
Best and second-best diagnostic ratios by linear discriminant analysis (LDA) ratio extractor for separating groups: A, in Nebria (Pseudonebriola) tsambagarav sp. nov. and the two other species of the Mongolian Altai N. (P.) medvedevi Shilenkov, 1982 and N. (P.) kerzhneri Shilenkov, 1982 (= ker+med); B, in N. (P.) tsambagarav sp. nov. as a member of the kerzhneri species group and N. (P.) kaszabi Shilenkov, 1982 as a member of the sajanica species group of the Kazakhstan and Russian Altai.
Holotype ♂: 48°41’27.1”N, 90°40’33.4”E; Mongolei, Bajan Ulgij Aimak, Tsambagarav uul; alpine Zone: Bachufer; 3168 üNN; 1. 7. 2016; 20 MG 2016 [internal abbreviation of the collector: location No. 20, Mongolia expedition 2016]; leg. Schnitter (
Paratypes: 31 ♂, 22 ♀, same data as holotype (
Body size: 7.5–9 mm.
Colour: Dorsal surface black, head black, without lightened spots on the vertex; appendages of the head brownish lightened, always lighter than the head. Antennal scape black, antennomeres 2–11 brown with darkened apices. Femora black, tibiae and tarsomeres brown, apices of tarsomeres usually black.
Head: Mandibles short; stipes flat, smooth, with 4–6 setae. Anterior margin of the labrum straight, bearing six setae. Anterior margin of the clypeus straight or slightly concave; clypeus laterally unisetose. Frontoclypeal suture concave. Frons with transverse wrinkles, which roundly merge into the lateral and longitudinal supraorbital wrinkles. Eyes protruding, temples present, oblique. Head supraorbitally unisetose. Disc impunctate with an isodiametric microreticulation. Antennae long and thin, extending to the end of the basal third of the elytra. Antennal scape as long as the eye’s diameter (ratio antennal scape length/eye length = 0.98±0.03 (0.93–1.04), slightly conical to suboval, distinctly narrowed basally and slightly apically, with one dorsoapical seta (Fig.
Pronotum (Fig.
Elytra: Silhouette suboval, apically faintly expanded, maximum width at two thirds. Ratio length/width of the elytra = 1.53 ± 0.02 (1.51–1.57). Elytral width = 1.59 ± 0.04 (1.51–1.66) times the pronotal width. Lateral margin faintly rounded, apex acutely rounded. Basal margination slightly curved, joined at an obtuse angle with the lateral margination. Shoulder flatly curved. Hindwings vestigial. Humeral and apical carinae little pronounced. Striae 1–7 distinct, punctate, stria 8 as a row of punctures. Striae 1–3 reaching the apex, the other striae obliterate slightly in front of the apex. Intervals on disc flat, interval 3 generally asetose (76%) on disc, disregarding the always present apical seta, occasionally with one seta on the disc (24%). Scutellar seta absent. Microsculpture isodiametric.
Ventral surface: Mes- and Metepisterna smooth and impunctate. Metacoxa basally and apically unisetose. Sternum II laterally faintly and widely dispersed punctate. Sternum III medially asetose. Sterna IV–VI each with one posterior paramedial seta. Anal sternum paralaterally unisetose in the male, bisetose in the female.
Legs: All tarsomeres dorsally glabrous; metatarsomere 4 ventrally with a short projecting tooth. Metafemur dorsoapically with a few short setae.
Male genitalia: Edeagus (Fig.
Female genitalia: Gonocoxa (Fig.
Median lobes; A–B dried lobes, C–D embedded lobes, A–C with partially everted endophallus: A, Nebria (Pseudonebriola) kerzhneri Shilenkov, 1982, holotype; B, N. (P.) medvedevi Shilenkov, 1982, paratype; C, N. (P.) tsambagarav sp. nov., holotype; D, N. (P.) kaszabi Shilenkov, 1982, Urunkhaika, Marakol Lake, Kazakhstan. Scale bar: 1 mm.
(Figs
Hydrophilic, ripicol, cold-preferring, alpine level.
The specific epithet refers to the type locality, the Tsambagarav uul; noun in apposition.
(Fig.
The Mongolian Pseudonebriola species are restricted to the main ridge of the Mongolian Altai (N. medvedevi), to the Tsambagarav mountain, the easternmost ridge bordering the Great Lakes Depression (N. tsambagarav sp. nov.), and to the Northwestern part of the Gobi Altai (N. kerzhneri).
Antennae: Antennal scape (in relation to the eye length) in N. tsambagarav sp. nov. longer than in N. kerzhneri and in N. medvedevi, the ratio scape length/eye length distinctly different: 0.98 in N. tsambagarav sp. nov., 0.89 in N. kerzhneri, and 0.82 in N. medvedevi (see also fig. 2 in
Second antennomere of N. tsambagarav sp. nov. oligosetose with (2)3–4 setae (Fig.
Pronotum: In N. tsambagarav sp. nov. the lateral groove of the pronotum at the anterior angle is wider than in N. kerzhneri and N. medvedevi. Lateral margin of the pronotum at the anterior angle distinctly edged as in N. medvedevi; such an edged anterolateral margin is absent in N. kerzhneri. In N. tsambagarav sp. nov. the pronotum is distinctly concave in front of the posterior angles (Fig.
Edeagus: The tip of the edeagus in N. tsambagarav sp. nov. is longer (Fig.
Gonocoxa: Gonocoxite 1 of N. tsambagarav sp. nov. (Fig.
N. tsambagarav sp. nov. is clearly identifiable by its oligosetose 2nd antennomere. In general the 2nd antennomere of all Pseudonebriola species has a single ventral seta. According to
The members of the sajanica species group are distributed in the Kazakhstan and Russian part of the Altai Mountain range, in the Sajan Mountains as far as to the Khamar Daban South of Baikal Lake. The members of the sajanica species group of the Kazakhstan and Russian Altai (N. sajanica, N. kaszabi, N. stanislavi) are characterized by the long antennal scape, which is bi- or oligosetose, and by the tarsomeres with a few dorsal setae. In contrast, the members of the kerzhneri species group (N. kerzhneri, N. medvedevi) are characterized by tarsomeres which are glabrous dorsally, and by the unisetose and short antennal scape. Due to its unisetose and short antennal scape, the dorsally glabrous tarsomeres, and due to the geographic position within the Mongolian Altai the new species N. tsambagarav sp. nov. is assigned to the kerzhneri species group within the subgenus Pseudonebriola.
The determination key of the subgenus Pseudonebriola in
8 | Lateral margin of the pronotum faintly concavely sinuate in front of the posterior angles, posterior angles rectangular, not protruding outwards. 2nd antennomere unisetose. Mongolia, Russia; main ridge of the Mongolian Altai | N. medvedevi Shilenkov, 1982 |
– | Lateral margin of the pronotum distinctly concavely sinuate in front of the posterior angles, posterior angles acute and protruding outwards. 2nd antennomere bi- or oligosetose. | 8a |
8a | Colour brown. Lateral margin of the posterior angles short, slightly protruding outwards. 2nd antennomere generally bisetose, occasionally unisetose. Elytra elliptic with lateral margin straight. Ratio antennal scape length/eye length < 0.9. Mongolia, Gobi Altai. | N. kerzhneri Shilenkov, 1982 |
– | Colour black. Lateral margin of the posterior angles long, distinctly protruding outwards. 2nd antennomere generally tri- or quadrisetose. Elytra suboval with lateral margin rounded. Ratio antennal scape length/eye length > 0.9. Mongolia, Mongolian Altai, Tsambaragav uul. | N. tsambagarav sp. nov. |
The publication on hand is a result of the German-Mongolian Expedition of the Martin-Luther-University Halle, Germany (em. Prof Michael Stubbe), in collaboration with the State University Ulan-Bataar, Mongolia (Dr Lkhagvaa Lkhagvasuren). We are grateful to Dr Volker Neumann und Karl Blaue for their tireless help in fieldwork. We thank Dr Boris Kataev (