Body slightly convex. Elytral striae regular, third interval with two or three setigerous pores. Metepisternum slightly wider than length. Mesofemur with four or more setae on ventral surface. Elytral plica absent or indistinct. Right paramere of male genitalia falciform, with somewhat elongated apex. Spermatheca with seminal canal and receptaculum not differentiated.

Medium size, body length 9.0–15.0 mm. Black, elytra slightly shiny, without metallic luster. Submentum with two long setae on each side. Pronotum cordate or quadrate; basal foveae deep, inner and outer grooves indistinctly separated or outer groove absent; one baso-lateral seta inserted on basal angle. Elytra striae straight and continuous, neither interrupted nor sinuate; interval microsculpture isodiametric, similar in male and female, third interval usually with two or three setigerous pores, fifth interval without pore; ninth interval with umbilical series slightly sparser in the middle than basal and apical areas; elytral plica indistinct; scutellar stria present; basal pore present or not. Metepisternum slightly wider than length. Terminal ventrite of males slightly depressed or without modification. Mesofemur with four or more setae on ventral surface, with a spine near apex; metacoxae with two setae; metatrochanters without seta. Fifth tarsomere with or without setae on ventral side. Apical orifice of aedeagus obviously twisted to left side; apical lamella narrow, short; right paramere falciform, apex more or less elongate and bent; endophallus bent to ventral-left or venter, gonopore opened to the base, with a large cap-like sclerotized gonopore piece (Figs 7–9). Gonocoxite II of ovipositor stout or slightly slender and bent, apex rounded, inner and outer margin each with one ensiform spine, apex with two very short nematiform setae in groove (Figs 60–65). Spermatheca tube-like, surface glabrous, receptaculum not differentiated from seminal canal, base of seminal canal sclerotized; spermathecal gland very fine, atrium and gland duct not differentiated, connected to the middle of spermatheca (Figs 58, 59).

This subgenus is endemic to China. A total of four species are distributed in Qinghai, Sichuan, Gansu, and Ningxia.

In its original description (Berlov, 1998), the subgenus Chinapterus was erected based on type species Pterostichus balthasari Jedlička, which was formerly a member of the subgenus Euryperis Motschulsky (Jedlička, 1962) and was subsequently synonymized with Petrophilus Chaudoir (Kryzhanovskij et al., 1995). Chinapterus is similar to Euryperis in having metepisternum slightly wider than length, metatrochanters without seta, and pronotum posterior angles widely rounded (only for P. singularis). But, Chinapterus is quite different from Euryperis in the following aspects: (1) mesofemur with four or more setae on the ventral side (with two setae in Euryperis); (2) elytral plica absent or indistinct (distinct in Euryperis); (3) spermatheca with seminal canal and receptaculum not differentiated (well differentiated in Euryperis).

Among the Chinese subgenera of Pterostichus, Chinapterus is most similar to the subgenera Sinoreophilus and Metallophilus in external appearance, and having the mesofemur with four or more setae near the hind margin, the metepisternum width subequal to its length, but differs in: (1) right paramere falciform, with the apex more or less elongate and bent; (2) elytral microsculpture isodiametric, similar in both sexes (in the other two subgenera, right paramere rounded triangular, apex not elongate, nor slightly bent; elytral microsculpture granular in females, isodiametric in males). Comparisons with other similar subgenera present in the key to subgenera.

Among all subgenera of Pterostichus from China, Chinapterus is doubtless closely related to other five subgenera (Platysma, Adelosia, Metallophilus, Sinoreophilus, Plectes) and shares the following important characters: (1) mesofemur with four or more setae along posterior margin; (2) elytral plica absent or indistinct; (3) metatrochanters without seta; (4) spermatheca tube-like, seminal canal and receptaculum not differentiated. These six subgenera form a monophyletic group (the Platysma group as defined here) supported by synapomorphic characters 1, 2, and 3 (character polarities discussed in this section follow Bousquet, 1999: 32–36). Except for the Chinese fauna, Myosodus, a subgenus centered in the Caucasus, also belongs to this group.

Among the above four characters, characters 1 is exclusive for the Platysma group in Pterostichus and character 4 is plesiomorphic. The undifferentiated spermatheca is unusual in Pterostichus and may suggest a relatively basal position of the Platysma group in the genus. So far as we know, all species of the Platysma group have a distinctive form of the female reproductive tract: spermatheca tube-like, seminal canal and receptaculum not differentiated, spermathecal canal inserted between the midpoint to the apical third of the spermatheca. In contrast, for most subgenera of Pterostichus, the spermatheca is usually very long with the seminal canal and receptaculum differentiated. If not so distinctly differentiated, the seminal canal is at least a little slenderer than the receptaculum, as in the subgenus Orientostichus. Except the Platysma group, only the subgenus Argutor and its relatives (five subgenera from China) have the undifferentiated spermatheca, but their spermathecae are always very short with the spermathecal canal inserted near the basal fourth of spermatheca.

The relationships among subgenera of the Platysma group are quite unclear, and even the monophyly of several subgenera is questioned. The subgenera Platysma and Sinoreophilus can be differentiated by plesiomorphic characters only, while other subgenera are merely defined by one or two apomorphic characters. Except for Adelosia (monotypic) and Plectes (includes two very closely related species), the monophyly of the other four subgenera are difficult to demonstrate.

We here redefine the subgenus Chinapterus and assign P. przewalskyi together with two new species very closely related to it into the subgenus for the following similarities: metepisternum length subequal to its basal width and right paramere falciform, apex rather elongated and bent. These two apomorphic characters may support the subgenus Chinapterus and can clearly differentiate it from all other related Chinese subgenera. However, the monophyly of Chinapterus is still questioned, because the elongate right paramere is also present in other subgenera of the Platysma group, including part of Platysma and all members of Myosodus. Moreover, P. singularis and P. przewalskyi do not look similar in their general appearances, and simply from the shape of right paramere, the former species is more similar to some species of Platysma while the latter is more like Myosodus. Nevertheless, the present definition of the subgenus Chinapterus is good for convenient taxa recognition at present. Under this, all Chinese Pterostichus species with multisetose mesofemora can be assigned to each subgenus. We expected an in-depth phylogenetic study will propose a better and more objective assignment of subgenera in the future.

When the present study on Chinapterus was conducted, we examined all Pterostichus species belonging to the Platysma group from China. We found that, P. lanista (Tschitschérine), P. militaris (Tschitschérine), and P. peilingi Jedlička should be moved into the subgenus Sinoreophilus for the following characters: mesofemur with four or more setae along posterior margin; elytra striae regular; metepisternum short, length subequal to the width of anterior margin; right paramere short and straight.

Pterostichus singularis Tschitschérine, Amdo, pres du fleuve Tay-tong-che (= Qinghai, Datong River). Pterostichus balthasari Jedlička, Gansu: Liangchow (= Gansu, Wuwei City).

Holotype of Pterostichus singularis Tschitschérine, male (ZIN) [Figs 1, 22], Amdo, 1886, G. Patani / F. singularis Typ. m. Tschitscherin det; Lectotype of Pterostichus balthasari Jedlička (designated herein), male (NMPC) [Figs 3, 23], Liangchow, W. Kansu / Type [red label] / Mus. Nat. Pragae, Inv. 24641 [orange label] / baltharsari, type, sp. n., Det. Ing. Jedlička [pink label]. Paralectotypes of Pterostichus balthasari Jedlička, 2 females (NMPC), Liangchow, W. Kansu / Cotype [red label] / Mus. Nat. Pragae, Inv. 24642/24643 [orange label] / baltharsari, sp. n., det. Ing. Jedlička.

Figures 1–6. 

Habitus of Pterostichus (Chinapterus) singularis Tschitschérine. 1 Holotype of P. singularis Tschitschérine, male (ZIN) 2 specimen determined by Tschitschérine, female, LT: alpes de Sinin (MNHN) 3 Lectotype of P. balthasari Jedlička, male (NMPC) 4 male, LT: Lajishan (IZAS) 5 male, LT: Lenglongling (IZAS) 6 male, LT: Dadongshu pass (IZAS).

1 female (MNHN) [Fig. 2], Thibet., alpes de Sinin [handwritten] / F. singularis, typ. m., Tschitscherin det [handwritten] / Museum Paris, coll. R. Oberthur. 1 male (MSNM), N slope of Xining mountains near Liandzha-sian, before 19–VI–1890, Gr.-Grzhimajlo leg. [in Russian] / F. singularis, typ. m. Tschitscherin det. / Paralectotypus, Pterostichus, Singularis Tschitsch. Vejeschagina 1983 [red label]. 1 female (NMPC), N slope Xining, Liandzha-sian vill., before 19–VI–1890, Grum leg. [in Russian] / F. singularis, typ., m., Tschitscherin det / Type [red label] /singularis, typ. Tsch., Det. Tschitscherin [pink label]. 5 males and 2 females (IZAS) [Figs 10, 11, 21], Gansu, Tianzhu county, G312 Wushaoling pass; alpine shrub, 37.2057N, 102.8799E, 3307 m, under rock; 2017.VII.12, Shi HL, Lu ZB & Zhu PZ lgt. 1 male and 1 female (IZAS) [Figs 5, 20], Gansu Prov. Lenglong Ling, 80 km NNW Honggu, 3392–3900 m, 37°03'50.3"N, 102°39'57.2"E, alpine pasture with Rhododendron, under stones, 2011.VI.30–31, D.W. Wrase.1 male and 1 female (IZAS) [Figs 4, 13], China, Qinghai Prov. Laji Shan pass, 34 km SSE Huangyuan, 36°23'01"–50"N, 101°19'31"–20'06"E, 3850–3880 m, wet plateau with depression, under stones/ clods, 8.VII.2011, D.W.Wrase. 1 female (IZAS), 2006.7.1–10, Qinghai Prov., Laji Shan. 2 males and 2 females (IZAS) [Figs 7–9, 14, 58, 64, 65], Qinghai, Qilian Shan, 3800m, pass Xi-ning-Guide, 9–11.08.10. M. Murzin. 7 males and 8 females (IZAS) [Figs 12, 16], Qinghai, Guide county, Lajishan pass, alpine meadow, N36.3573 E101.4463, 3829 m, under rock, 2017.VII.20, Shi HL, Lu ZB & Zhu PZ lgt. 2 males and 7 females (IZAS), Qinghai, Guide county, N. Slope of Guoshize Mt.; screes; 4007 m; 36.2914, 101.6005, 2017.VII.21, under stone, SHI HL & LU ZB leg. 1 male and 1 female (IZAS), Qinghai, Guide county, 3 km W of Lajishan pass, alpine meadow, 36.3622N, 101.4117E, 3686 m, pitfall trap; 2017.VII.21; Shi HL, Lu ZB & Zhu PZ lgt. 2 males and 1 female (IZAS) [Fig. 15], Qinghai, Guide county, Guoshizeshan pass, Salix shrubs, N36.3082 E101.5973, 3487 m, pitfall trap, 2017.VII.21; Shi HL, Lu ZB & Zhu PZ lgt. 1 male and 2 females (IZAS), Qinghai, Qilian, Datong Shan, 38.020364N, 100.134777E / 3770 m, 2012.V.31, Huang Xinlei leg. 6 males and 1 female (IZAS) [Figs 6, 18, 19], Qinghai, Qilian county, 8 km N of Dadongshu pass, alpine meadow, 38.0531N, 100.2273E, 3541 m, pitfall trap; 2017.VII.18, Shi HL, Lu ZB & Zhu PZ lgt. 1 male (IZAS) [Fig. 17], Qinghai, E. Qilian Shan, 4200 m, 1.8.1992, J.Kalab leg. / Compared with type, SHI HL 2011 / Pterostichus balthasari Jedlička, 1937 / ex coll. Sciaky 2011. 1 male and 1 females (IZAS), Qinghai, Menyuan county, Xianmi, Ningchan pass, alpine meadow, 37.5374N, 101.8704E, 3900 m, under rock, 2017.VII.14, Shi HL, Lu ZB & Zhu PZ lgt. 1 female (IZAS), Qinghai, Menyuan county, Xianmi xiang, Taola, Picea forest, 37.2309N, 102.0134E, 2636 m, under rock; 2017.VII.15, Shi HL, Lu ZB & Zhu PZ lgt. 1 female (IZAS), Qinghai, Menyuan county, Xianmi xiang, Bangusi, Picea forest, 37.2908N, 101.9438E, 2707 m, pitfall trap; 2017.VII.15, Shi HL, Lu ZB & Zhu PZ lgt. 1 male and 4 females (IZAS), Qinghai, Qilian county, Zhamashi, Zhamashixigou; alpine meadow; 3014 m, 38.1590N, 99.9921E, under rock; 2019.VIII.19; YAN Weifeng lgt. 4 males and 2 females (IZAS), Qinghai, Qilian county, Babao town, Lujiaogou; alpine meadow; 3461 m, 38.0991N, 100.4838E, under rock; 2019.VIII.17; YAN Weifeng lgt. 3 females (IZAS), Qinghai, Menyuan county, Xianmi, Qihankaigou; mixed forest; 2683 m, 37.1568N, 102.0278E, under rock; 2019.VIII.12; YAN Weifeng lgt. 2 males (IZAS), Qinghai, Qilian county, Epu town, Jingyangling pass; alpine meadow; 3716 m 37.8387N, 101.1117E, under rock; 2019.VIII.15; YAN Weifeng lgt. 2 females (IZAS), Qinghai, Qilian county, Babao township, Binggou; alpine meadow; 3826 m 38.1171N, 100.1716E, under rock; 2019.VIII.18; YAN Weifeng lgt. 1 male (NMPC), China: Qinghai province, Gangca Dasi [lamasery]., 37°32.4'–33.0'N, 100°05.3'–06.0'E, 3505–3840 m, 11–12.VII.2005, J. Hajek, D.Kral & J.Ruzicka leg; individually under stones, in excrements and on vegetation; spring; alpine meadows and pastures around the lamasery and in the nearby valley. 1 female (NMPC), China: Qinghai province, Yuning Si [lamasery], 2890 m, 36°45.6'N, 102°10.6'E, 16.VII.2005, J. Hajek, D. Kral & J. Ruzicka leg, individually under stones and logs, in excrements, and on vegetation in coniferous forest, on the pastures, and along the path to a village; pool.

Figures 7–23. 

male genitalia of Pterostichus (Chinapterus) singularis Tschitschérine. 7–9 Endophallus, LT: Pass Xining-Guide (IZAS) 7 right lateral view 8 left lateral view 9 ventral view, abbreviations as stated in text 10–11 median lobe of aedeagus, LT: Wushaoling (IZAS) 10 dorsal view 11 left lateral view 12–23 right paramere, inner face 12 LT: Lajishan (IZAS) 13 LT: Lajishan (IZAS) 14 LT: Pass Xining-Guide (IZAS) 15 LT: Guoshizeshan (IZAS) 16 LT: Lajishan (IZAS) 17 LT: E Qilianshan (IZAS) 18 LT: Dadongshu pass (IZAS) 19 LT: Dadongshu pass (IZAS) 20 LT: Lenglongling (IZAS) 21 LT: Wushaoling (IZAS) 22 Holotype of P. singularis (ZIN) 23 Lectotype of P. balthasari (NMPC). Scale bars: 0.5 mm.

Femora black. Pronotum quadrate; lateral margins slightly rounded before middle, nearly straight before basal angles; basal angles rounded, or slightly rectangular with an indistinct denticle; basal fovea depressed between outer and inner grooves; elytral basal pore usually absent, third interval usually with two setigerous pores; fifth tarsomere with one or two pairs of fine setae ventrally; right paramere falciform, apex acicular or triangular.

BL 9.2–10.8 mm, BW 3.9–4.4 mm. Robust, black, elytra shiny. Head large; frons smooth or very sparsely punctate; genae short, less than one-third length of eyes; eyes small, slightly prominent. Pronotum quadrate; widest before middle, PW/HW = 1.26–1.43, PW/PL = 1.37–1.50; lateral margins slightly rounded from apical angles to the middle, nearly straight before basal angles, mid-lateral seta present at apical third; basal margin slightly wider than apical margin, PBW/PAW = 1.07–1.20; basal angles rounded or slightly rectangular with an indistinct denticle, not protruding outward; basal fovea with inner and outer grooves faintly defined and partly fused, forming deep depression between them, outer groove slightly shorter than inner one; basal foveal area coarsely punctate; middle area between two basal foveae smooth or very sparsely punctate; area between outer groove and lateral margin slightly convex, forming a weak carina; disc convex, smooth, transversely rugose in some specimens. Elytra oblong, EL/EW = 1.33–1.47; basal ridge slightly oblique; shoulder rounded, basal ridge and lateral margin forming an obtuse angle, humeral tooth absent; apical plica absent; basal setigerous pores usually absent, occasionally present in one elytron or both elytra; scutellar striae short, apex free or connected to first stria; intervals feebly convex; microsculpture similar in both sexes, finely isodiametric; third interval often with two setigerous pores on posterior half, adjacent to second stria, sometimes one or two additional pores present; umbilical series on ninth interval sparse in middle, composed of 14–17 pores; striae moderately deep, distinctly punctate. Ventral side. Proepisternum sparsely punctate near inner margin, rugose throughout; mesepisternum very sparsely punctate; metepisternum nearly smooth; terminal or penultimate ventrite of males not modified. Tarsomere 5 with one or two pairs of setae ventrally; metatarsomeres without distinct outer furrow. Male genitalia. Ventral margin of median lobe near straight at middle, slightly bent downwards near apex; apical orifice opened left-dorsally (Figs 10, 11); apical lamella narrowed in dorsal view, length near two folds of basal width, apex rounded. Right paramere falciform, strongly curved, the obtuse angle between basal portion and apical portion 105–110°; apex acicular or narrow triangular, narrow, sharp (Figs 12–21). Endophallus bent to the ventral left side, major portion of endophallus located at left side of median lobe apex; gp opened to basal-left; gpl large and coniform; gpp large and cap-like, on the ventral side of gp; three groups of lobes recognized: dl on dorsal surface of endophallus, gradually swollen; rl on ventral-right surface of endophallus, divided into two conjoint rounded sub-lobs; lbl on the ventral-left surface of endophallus, smaller than rl; lal absent (Figs 7–9). Female genitalia. Gonocoxite II of ovipositor stout, length ca. 1.5 times maximum width; inner and outer margins arched, each with one ensiform setae near middle, apex rounded, with two very short nematiform setae in a groove (Figs 64, 65). Spermatheca short and tubiform, length ca. four times maximum width (Fig. 58).

Only known from the eastern section of the Qilian mountains on the border of Qinghai and Gansu provinces (Map 1).

Map 1. 

Distributions for P. singularis.

This species prefers living in open habitat like the alpine meadows of the Qilian mountains between 3000 m and 4000 m, but a few specimens were also found on the edge of Picea forest at ca. 2600 m.

Pterostichus singularis Tschitschérine was described based on a single male from “Amdo, pres du fleuve Tay-tong-che”, referring to what is now the valley of the Datong River in Menyuan County. Thus, the three specimens we examined from different collections are not type specimens but only subsequently determined by Tschitschérine (1898: 181) although labeled as types by the author. The true holotype was deposited in the collection of ZIN.

Pterostichus balthasari Jedlička was described based on five specimens from “Liangchow” without an original fixation for holotype. In the collection of NMPC, we examined three specimens in accordance with the original literature. We designate the male bearing a “type” label as the lectotype herein, for the taxonomic purpose of fixing the species name to a single specimen and preventing further confusion. The type locality “Liangchow” refers to what is now Wuwei City in Gansu Province. The exact type locality could be the north slope of Mt. Qilian Shan in Wuwei territory.

Comparing with specimens of P. singularis determined by Tschitschérine, the types of P. balthasari are nearly identical but the basal angles of the pronotum are more rounded and the type locality is further north. We studied specimens from several localities of the Qilian mountains and found that the northern population (Wushaoling, Lenglongling, Datong Shan) usually has a rounded or a slightly rectangular pronotal basal angle, but the southern population (Laji Shan) generally has a rectangular basal angle of the pronotum;however, a few specimens from Laji Shan also have rounded pronotal basal angles. Moreover, there is no significant difference in the male genitalia between the different localities, both for the median lobe and right paramere. Thus, we synonymize P. balthasari with P. singularis.

This species is widely distributed in the eastern section of the Qilian mountain range and is sometimes locally abundant. We studied many specimens from the five branches of the Qilian mountain range: Wushaoling, Lenglongling, Daban Shan, Datong Shan, and Laji Shan. In addition to the above variations of the basal angle, three more variations exist geographically or individually as follows:

(1) elytral dorsal pores on third interval. For ca. two-thirds of the examined specimens, two setigerous pores are present on the third elytral interval. For the other one-third, one or two additional pores are present anterior to the first pore or between the first and second pores.

(2) elytra basal pore. For most specimens there is no basal pore on the elytra, but occasionally, a basal pore is present on one elytron or on both elytra. The proportion of individuals with elytra basal pores is larger in the population from Laji Shan than in the others.

(3) shape of right paramere. The length of the right paramere varies individually as well as geographically. Statistically, the northeastern populations (Wushaoling, Lenglongling, Datong Shan) have a longer right paramere with the apex usually more elongate and acicular as in Figs 17–21. In contrast, the southern (Laji Shan) populations usually have shorter right paramere with the apex less elongate or narrowly triangular as in Figs 12–14. But, this individual variation with a long acicular right paramere is also present in some specimens from the southern localities (Figs 15, 16).

From the original description, the type localities are “Amdo: près des rivières Tala-tchu et By-tchu, aux sources du Jangtsekiang”. By-tchu refers to the Buqu River in Qumarlêb County of Qinhai Province, and Tala-tchu probably refers to the Derqu River in Chindu County, but the detailed locality was not labeled for any examined type so the precise type locality for the lectotype is unclear.

Lectotype of Pterostichus przewalskyi Tschitschérine (designated herein), male (ZIN) [Figs 24, 30, 31, 47], Amdo / Przewalskii m. Typ. Tschitscherin det / Zoological Institute Russian Academy of Sciences, St. Petersburg [yellow label]. Paralectotypes of Pterostichus przewalskyi Tschitschérine, 1 male (ZIN), N. E. Thibet, 1884, Przewalsky / Przewalskii m. typ. Tschitscherin det / Zoological Institute, Russian Academy of Sciences, St. Petersburg [yellow label]. 1 female (NMPC), N.E. Thibet, 1884, Przewalsky. / przewalskii m., typ., Tschitscherin det / Type [red label] / przewalskii, typ., Tsch., Det. Tschitscherin [pink label].

1 female (IZAS), China, N Sichuan prov., Hongyuan, ca. 4200 m, 21.7–3.8.1991, J. Kalab leg. 1 male (IZAS) [Figs 42, 51], Ch-NW Sichuan, ±3700 m, 32.59N, 98.06E, 3+15/7, SERXÜ env. 1995, alpine meadow, Jaroslav Turna leg. 1 female (IZAS), Qinghai, Baima county, Makehe, Meilanggou, 2013.VI.28, D 3500 m, Chen Jin leg. 2 males (IZAS) [Figs 25, 44, 50], Sichuan, Zoigê county, 8km W. of Hongxing, wetland, 3236 m, 34.1236N, 102.6545E, under rock, 2016.VII.25, Shi HL lgt. 3 males and 1 female (IZAS) [Figs 32, 33, 49], Qinghai, Jigzhi county, E. of Luanshitou pass; alpine meadow; 4091 m; 33.4008N, 101.2537E; 2017.VIII.2, pitfall trap, SHI HL, LU ZB, ZHU PZ & YAN WF leg. 1 male (IZAS) [Figs 43, 48], Qinghai, Jigzhi county, Luanshitou pass; alpine meadow; 4212 m; 33.3986N, 101.2432E; 2017.VIII.2, under stone, SHI HL, LU ZB, ZHU PZ & YAN WF leg. 8 females (IZAS) [Figs 59–61], Qinghai, Chidu county, 2 km S of Zhenqin; alpine wetland, 4329 m, 33.3908N, 97.2977E, 2017.VII.27, under stone, SHI HL, LU ZB, ZHU PZ & YAN WF leg.

Femora reddish brown. Pronotum cordate; lateral margins strongly sinuate before basal angles; basal angles right-angled, clearly pointed outwards. Elytral basal fovea depressed between inner and outer grooves; basal pore present; third interval often with two setigerous pores. Fifth tarsomere glabrous ventrally; right paramere strongly elongate and bent, apex rounded, not declined to dorsum.

BL 12.0–13.4 mm, BW 4.8–5.6 mm. Robust, black, femora reddish brown except black apex; elytra slightly shiny. Head large, frons smooth or very sparsely punctate; genae short, less than one-third length of eyes; eyes prominent. Pronotum cordate; widest slightly before middle, PW/PL = 1.33–1.41; lateral margins largely rounded before the middle, strongly sinuate before basal angles; one mid-lateral seta present at apical third; basal margin slightly wider than apical margin, PBW/PAW = 1.02–1.12; basal angles rectangular, clearly protruding outwards; basal foveae deep, inner and outer grooves faintly defined, inner groove straight, distant from basal margin, outer groove shorter than inner one, extending to basal margin; basal fovea depressed between inner and outer grooves, coarsely punctate in basal fovea; area between outer groove and lateral margin hardly convex; disc convex, smooth, slightly transversely rugose; apical angle rounded, not protruding. Elytra oblong, EL/EW = 1.42–1.45; basal ridge slightly oblique; shoulder rounded, basal ridge and lateral margin forming an obtuse angle, humeral tooth small and obtuse; apical plica indistinct; basal setigerous pores present; scutellar striae complete; intervals slightly convex; microsculpture similar in both sexes, finely isodiametric; third interval usually with two setigerous pores, at basal two-fifths and two-thirds respectively, occasionally with one or three pores on one elytron, all adjacent to second stria; ninth interval with umbilical series regularly arranged, slightly sparser in the middle; striae deep, indistinctly punctate. Fifth tarsomere glabrous ventrally; meso- and metatarsomere I and II with outer groove. Terminal ventrite shallowly depressed and finely rugose in males. Male genitalia. Ventral margin of median lobe nearly straight at middle, gradually bent downwards near apex; apical orifice opening left-dorsally; apical lamella small, rounded triangular, length sub-equal to basal width, apex rounded (Figs 30, 31); apical lamella slightly twisted, forming a continuously curved dorsal-left surface (Figs 42–44). Right paramere strongly elongate and curved, the obtuse angle between basal portion and apical portion 100–105°; apex slightly thick, well rounded, not or weakly bent to dorsum (Figs 47–51). Endophallus bent to the ventral side of aedeagus, major portion of endophallus located at ventral side of median lobe apex; gp opened to basal-dorsum; gpl large and coniform; gpp large and cap-like, on the ventral side of gp; two groups of lobes recognized: rl on ventral-right surface of endophallus, large, rounded or apex a little pointed; lbl on ventral-left surface of endophallus, divided into two clearly separated sub-lobes, the basal one large and nearly rounded, the apical one much smaller than the basal one; lal absent (Figs 32, 33). Female genitalia. Gonocoxite II of ovipositor slightly slender, length ca. 2.5 times maximum width; inner and outer margins near straight, each with one ensiform setae before middle, apex narrowly rounded, with two very short nematiform setae in a groove (Figs 60, 61). Spermatheca moderately long and tube-like, length ca. 15 times maximum width (Fig. 59).

Qinghai (southeast part), Sichuan (northwest part). Probably also in southwest Gansu (Map 2).

Map 2. 

Distributions for P. przewalskyi.

This hygrophilous species tends to live in wet habitats such as wetlands or the banks of seasonal streams on alpine meadows.

This species was described based on an unspecified number of specimens from “Amdo: près des rivières Tala-tchu et By-tchu” collected by Przewalsky in 1884. In the collection of ZIN, we examined two males which matched with the original literature. We designated the male with its genitalia dissected as the lectotype herein, for the taxonomic purpose of fixing the species name to a single specimen and preventing further confusion.

Sciaky (1996) assigned this species to the subgenus Sinoreophilus due to external similarities but this species has its right paramere strongly elongated and the elytral microsculpture is similar in both sexes, supporting a close relationship to P. singularis. So, we herein moved it from the subgenus Sinoreophilus to the subgenus Chinapterus.

Holotype : male (IZAS) [Figs 26, 34, 35, 45, 52], China, Gansu, Kangle, Lianhuashan, Shahetan station 34.93917N, 103.73472E / 2850 m, 2008.VI.1, WANG J. leg.; pit fall; Institute of Zoology. Paratypes: 1 male and 1 female (IZAS), 2012,VI,21 D 2960 m, China, Gansu, Kangle, Lianhuashan, 34.93577N, 103.75054E / Morii leg., Inst. of Zoology, CAS. 1 male and 2 females (IZAS) [Figs 27, 36, 37, 53, 62, 63], 2012,VI,22. D 2960 m, Gansu, Kangle, Lianhuashan, 34.91943N, 103.72860E / Liang Hongbin, Sota leg., Inst. of Zoology, CAS. 1 male and 1 female (IZAS) [Fig. 54], China, Gansu, Kangle, Lianhuashan, Shahetan station 34.93917N, 103.73472E / 2850 m, 2008.VII.6, WANG J. leg.; pitfall trap; Institute of Zoology.

Figures 24–29. 

Habitus of Pterostichus (Chinapterus) spp. 24 Lectotype of P. przewalskyi, male (ZIN) 25 P. przewalskyi, male, LT: Hongxing (IZAS) 26 Holotype of P. lianhuaensis sp. nov. (IZAS) 27 Paratype of P. lianhuaensis sp. nov., male (IZAS) 28 Holotype of P. liupanensis sp. nov. (IZAS) 29 Paratype of P. liupanensis sp. nov., male (IZAS).

Femora black; pronotum cordate, lateral margins strongly sinuate before basal angles, which clearly pointed outwards; basal foveae convex between inner and outer grooves; elytral basal pore present, third interval often with three setigerous pores; fifth tarsomere with one or two pairs of fine setae ventrally; right paramere strongly elongated and bent, apex slightly bent to dorsum, angulate at dorso-apical end.

BL 12.3–13.6 mm, BW 5.0–5.5 mm. Robust, black, femora completely black, elytra slightly shiny. Head large, frons smooth or very sparsely punctate; genae short, less than one-third length of eyes; eyes prominent. Pronotum cordate; widest slightly before middle, PW/PL = 1.31–1.40; lateral margins largely rounded before middle, strongly sinuate before basal angles; one mid-lateral seta present at apical one-third; basal margin slightly wider than apical margin, PBW/PAW = 1.02–1.13; basal angles rectangular, clearly protruding outwards; basal foveae narrow and deep, inner groove well present, apex reaching basal third of pronotum, outer groove obsolete; basal fovea convex between inner and outer grooves, not convex between outer groove and lateral margin, basal foveal area coarsely punctate; disc convex, smooth, finely transversely rugose on basal half; apical angles rounded, not protruding. Elytra oblong, EL/EW = 1.36–1.46; basal ridge slightly oblique; shoulder rounded, basal ridge and lateral margin forming an obtuse angle, humeral tooth small and obtuse; apical plica indistinct; basal setigerous pores present; scutellar striae complete; intervals slightly convex, microsculpture similar in both sexes, finely isodiametric; third interval usually with three setigerous pores, the basal one at basal seventh, adjacent to third stria, the apical two at basal two-fifths and two-thirds respectively, all adjacent to second stria, occasionally with four or five pores on one elytron; ninth interval with umbilical series regularly arranged, slightly sparser in middle; striae deep, indistinctly punctate. Fifth tarsomere with one or two pairs of setae ventrally; meso- and metatarsomere I and II with outer groove. Terminal ventrite shallowly depressed and finely rugose in males. Male genitalia. Ventral margin of median lobe near straight at middle, gradually bent downwards near apex; apical orifice opened left-dorsally; apical lamella small, rounded triangular, length subequal to basal width, apex rounded, weakly bent to left (Figs 34, 35); apical lamella slightly twisted, forming a continuously curved dorsal-left surface (Fig. 45). Right paramere strongly elongate and curved, the acute angle between basal portion and apical portion 85–90°; apex slightly thick, slightly angulate at dorsal-apical end, slightly bent to dorsum (Figs 52–54). Endophallus bent to the ventral side of aedeagus, major portion of endophallus located at ventral side of median lobe apex; gp opened to basal-dorsum; gpl large and coniform; gpp large and cap-like, on ventral side of gp; three groups of lobes recognized: rl on ventral-right surface of endophallus, large, near rounded, without projection; lbl on ventral-left surface of endophallus, large and fully round, not divided into sub-lobes; lal on left surface of endophallus, very small, rounded or divided into two small sub-lobes (Figs 36, 37). Female genitalia same as that of P. przewalskyi (Figs 62, 63).

Only known from the type locality, Lianhua Shan mountain, Kangle County, south of Gansu province (Map 3).

Map 3. 

Distributions for P. lianhuaensis sp. nov. (red) and P. liupanensis sp. nov. (blue).

The name of the new species refers to its type locality, Lianhua mountain.

This species is similar to P. przewalskyi in having a large body size, lateral margins of the pronotum strongly sinuate before basal angles, but differs from the latter in: (1) femora completely black; (2) basal foveae convex between inner and outer grooves; (3) elytra third interval usually with three pores, the basal one adjacent to the third stria; (4) fifth tarsomere with one or two setae ventrally. In P. przewalskyi, femora reddish brown; basal foveae of pronotum depressed between inner and outer grooves; elytral third interval usually with two pores, if with three the basal one adjacent to the second stria; fifth tarsomere glabrous ventrally. The male genitalia of these two species are very similar, with small differences in: (1) apical lamella of aedeagus weakly bent to left in P. lianhuaensis sp. nov. (Fig. 34), almost straight in P. przewalskyi (Fig. 30); (2) apex of right paramere slightly angulate at dorsal-apical end in P. lianhuaensis sp. nov., completely rounded in P. przewalskyi; (3) right paramere more bent in P. lianhuaensis sp. nov., forming an acute angle 85–90°, versus 100–105° in P. przewalskyi; (4) endophallus with lbl not divided and lal present in P. lianhuaensis sp. nov., lbl divided into two sub-lobes and lal absent in P. przewalskyi.

We noticed that this new species only has very minute differences from P. przewalskyi in the male genitalia. However, considering the stable gaps between their features (especially the external characters) and distributions, we decide to establish a new species but not a new subspecies. A similar situation is also present in the next new species, P. liupanensis. Several external differences among these three species are generally of interspecific level in other Pterostichus groups, such as the shape of the pronotum base and basal fovea, chaetotaxy on elytral third interval, and setae on the fifth tarsomere.

Holotype : male (IZAS) [Figs 28, 38, 39, 46, 55], China, Ningxia A.R., Liupanshan, Xixia Forestry Station, 35.51620N, 106.25086E / 2200 m, 2008.6.25–27, Zhao Zongyi Pitfall trap, Institute of Zoology. Paratypes, 1 male (IZAS) [Fig. 56], China, Ningxia A.R., Liupanshan, Longtan Forestry Station, 35.38982N, 106.34508E / 1936 m, 2008.6.23–25, Lou Qiaozhe, pitfall trap, Institute of Zoology. 1 male (IZAS) [Figs 29, 40, 41, 57], China, Ningxia A.R., Liupanshan, Xixia Forestry Station, 35.49673N, 106.31310E / 1994 m, 2008.6.25N, Lou Qiaozhe collector, Institute of Zoology.

Figures 30–41. 

male genitalia of Pterostichus (Chinapterus) spp. 30–33 P. przewalskyi; 30–31 median lobe of aedeagus, dorsal and left lateral view, Lectotype (ZIN) 32, 33 endophallus, right and left lateral view, LT: Jigzhi (IZAS) 34–37 P. lianhuaensis sp. nov. 34, 35 median lobe of aedeagus, dorsal and left lateral view, Holotype (IZAS) 36, 37 endophallus, right and left lateral view, Paratype, LT: Lianhuashan (IZAS) 38–41 P. liupanensis sp. nov. 38, 39 median lobe of aedeagus, dorsal and left lateral view, Holotype (IZAS) 40, 41 endophallus, right and left lateral view, Paratype, LT: Liupanshan, Xixia (IZAS). Scale bars: 0.5 mm, abbreviations as stated in text.

Femora black; pronotum near quadrate, lateral margins near straight before basal angles, which weakly pointed outward and with a small denticle; basal foveae convex between inner and outer grooves; elytral basal pore present, third interval usually with three setigerous pores; fifth tarsomere glabrous ventrally; right paramere strongly elongated and bent, apex rounded, not bent to dorsum.

BL 12.4–13.2 mm, BW 4.8–5.1 mm. Robust, black, femora black, elytra slightly shiny. Head large, frons smooth or very sparsely punctate; genae short, less than one-third length of eyes; eyes prominent. Pronotum near quadrate; widest slightly before middle, PW/PL = 1.33–1.37; lateral margins largely rounded before middle, nearly straight before basal angles; one mid-lateral seta present at apical one-third; basal margin slightly wider than apical margin, PBW/PAW = 1.12–1.16; basal angles rectangular, weakly protruding outwards, with a small denticle; basal foveae narrow and deep, inner groove obviously present, apex reaching basal third of pronotum, outer groove obsolete; basal fovea convex between inner and outer grooves, not convex between outer groove and lateral margin, basal foveal area coarsely punctate; disc convex, smooth, finely transversely rugose at basal half; apical angle rounded, not protruding. Elytra oblong, EL/EW = 1.46–1.52; basal ridge slightly oblique; shoulder rounded, basal ridge and lateral margin forming an obtuse angle, humeral tooth small and obtuse; apical plica indistinct; basal setigerous pores present; scutellar striae complete; intervals slightly convex, microsculpture finely isodiametric in males; third interval with three setigerous pores, the basal one at basal seventh, adjacent to third stria, the apical two at middle and apical fourth respectively, all adjacent to second stria; ninth interval with umbilical series regularly arranged, slightly sparser in middle; striae deep, indistinctly punctate. Fifth tarsomere glabrous ventrally; meso- and meta-tarsomeres I and II with outer groove. Terminal ventrite shallowly depressed and rugose in males. Male genitalia. Ventral margin of median lobe near straight at middle, obviously bent downwards near apex; apical orifice opened left-dorsally; apical lamella small, rounded triangular, length sub-equal to basal width, apex rounded (Figs 38, 39); apical lamella not twisted with its dorsal surface perpendicular to left surface (Fig. 46). Right paramere strongly elongate and curved, the acute angle between basal portion and apical portion near 90°; apex slightly thick, well rounded, not or weakly bent to dorsum (Figs 55–57). Endophallus bent to the ventral side of aedeagus, major portion of endophallus located at ventral side of median lobe apex; gp opened to basal-dorsum; gpp large and cap-like, on ventral side of gp; three groups of lobes recognized: rl on ventral-right surface of endophallus, large and rounded, with a papillary projection to the apex of endophallus; lbl on ventral-left surface of endophallus, divided into two clearly separated sub-lobes, the apical one very small, same size as lal, the basal one large, same size as rl; lal on left surface of endophallus, very small and rounded (Figs 40, 41).

Figures 42–57. 

male genitalia of Pterostichus (Chinapterus) spp. 42–46 Apex of median lobe, dorsal left view 42 P. przewalskyi, LT: Serxü (IZAS) 43 P. przewalskyi, LT: Jigzhi (IZAS) 44 P. przewalskyi, LT: Hongxing (IZAS) 45 P. lianhuaensis sp. nov. Holotype (IZAS) 46 P. liupanensis sp. nov. Holotype (IZAS) 47–57 right paramere, inner face 47–51 P. przewalskyi 52–54 P. lianhuaensis sp. nov. 55–57 P. liupanensis sp. nov. 47 Lectotype (ZIN) 48 LT: Jigzhi (IZAS) 49 LT: Jigzhi (IZAS) 50 LT: Hongxing (IZAS) 51 LT: Serxü (IZAS) 52 Holotype (IZAS) 53 Paratype, LT: Lianhuashan (IZAS) 54 Paratype, LT: Lianhuashan, Shahetan (IZAS) 55 Holotype (IZAS) 56 Paratype, LT: Liupanshan, Longtan (IZAS) 57 Paratype, LT: Liupanshan, Xixia (IZAS). Scale bars: 0.5 mm (42–45, 47–57); 0.2 mm (46).

Only known from the Liupan Shan mountain range, Jingyuan County, Ningxia Huizu Autonomous Region (Map 3).

The name of the new species refers to its type locality, Liupan Shan mountain.

Map 4. 

Distributions for Pterostichus (Chinapterus) spp. P. singularis (green), P. przewalskyi (violet), P. lianhuaensis sp. nov. (red), P. liupanensis sp. nov. (blue); black boxes mark area shown in maps 1–3.

The new species is similar to P. lianhuaensis sp. nov. in having a large body size, black femora, narrow pronotum basal fovea, elytra third interval usually with three pores, but differs from the latter by having: (1) pronotum lateral margins nearly straight before basal angles; and (2) fifth tarsomere glabrous ventrally. In P. lianhuaensis sp. nov., pronotum lateral margins strongly sinuate before basal angles; fifth tarsomere with one or two pairs of setae ventrally. The male genitalia of the new species is very similar to that of P. przewalskyi, with small differences in: (1) ventral margin of aedeagus more distinctly bent downwards near apex in lateral view in P. liupanensis sp. nov. (Fig. 39), evenly and gradually bent downwards in lateral view in P. przewalskyi (Fig. 31); (2) apical lamella with dorsal surface perpendicular to left surface in P. liupanensis sp. nov., slightly twisted, forming a continuously curved dorsal-left surface in P. przewalskyi; (3) right paramere more bent forming an acute angle near 90°, versus 100–105° in P. przewalskyi; (4) endophallus with rl projected to the apex of endophallus and lal present, in P. przewalskyi rl not or slightly projected to the base of endophallus and lal absent.

Figures 58–65. 

female genitalia of Pterostichus (Chinapterus) spp. 58, 59 Female reproductive system 58 P. singularis, LT: Pass Xining-Guide (IZAS) 59 P. przewalskyi, LT: Zhenqin (IZAS). Scale bar: 0.5 mm. 60–65 Female ovipositor 60, 61 P. przewalskyi, LT: Zhenqin (IZAS) 62, 63 P. lianhuaensis sp. nov., Paratype LT: Lianhuashan (IZAS) 64–65 P. singularis, LT: Pass Xining-Guide (IZAS) 60, 62, 64 Gonocoxite II, inner view, 61, 63, 65 Gonocoxite I, II, ventral view. Scale basr: 0.2 mm.