Tschitscherinea Berg, 1898

Yin, Wen-Qi, Chen, Jia-Heng & Shi, Hong-Liang, 2024, Revision of the Chinese endemic subgenus Tschitscherinea Berg, with descriptions of three new species (Coleoptera, Carabidae), Zootaxa 5481 (1), pp. 43-65 : 44-47

Subgenus Tschitscherinea Berg, 1898

Eunema Tschitschérine, 1897: 70 [junior homonym of Eunema Salter, 1859 ( Lophospiridae )]. Type species: Feronia filum Tschitschérine, 1897 , by monotypy.

Tschitscherinea Berg, 1898: 18 . Replacement name for Eunema Tschitschérine, 1897 . Jedlička, 1962: 228. Sciaky, 1997: 166.

Jedlickaia Sciaky, 1997: 162 . Type species: Pterostichus sterbai Jedlička, 1934 , by original designation. New synonymy.

Diagnosis. Species of the subgenus Tschitscherinea can be recognized among Chinese Pterostichus by the following combinations: medium-sized pterostichid beetles; body form relatively slender, more or less parallel-sided; dorsum reddish brown to dark brown; submentum with two setae on each side; elytra interval 3 with one to three setigerous pores; umbilicate pore series on interval 9 discontinuous in middle; metepisternum at most slightly longer than basal width; mesofemora with two setae on posterior-ventral margin; metacoxae with three setae; median lobe of aedeagus relatively small, apical lamella short with rounded apex; right paramere elongated, often curved at anterior third ( Figs 15–24 ).

Subgeneric characters. Body form relatively slender, more or less parallel-sided; body length 9.0– 15.5 mm; dorsum reddish-brown to dark brown, elytra without or with very faint metallic luster. Head small to distinctly thickened, always narrower than pronotum; eyes small, two supraorbital setae present; vertex without puncture or wrinkle; frontal grooves shallow, sometimes indistinct, slightly sinuate, reaching mid-point of eyes. Antennae hardly reaching or slightly exceeding elytra base; scape shorter than combined length of following two segments; antennomere 3 with 6–7 primary long setae forming an apical ring, without short accessory seta. Mandibles long, outer margin nearly straight, curved near apical fourth, apex sharply hooked; terminal labial and maxillary palpomere cylindrical, weakly broadened in males; penultimate labial palpomere with two setae along inner margin, without an extra seta near apex; submentum with two setae on each side. Pronotum usually nearly quadrate, lateral sides nearly paralleled or slightly curved, straight or slightly sinuate before posterior angles, but in P. sterbai , pronotum nearly ovate, lateral margins evenly arched near middle; posterior margin almost straight, sometimes very shallowly emarginate at middle, usually a little narrower than base of elytra between humerus; anterior angles slightly protruded with blunt apex; posterior angles usually rectangular, apex forming faint denticles, except for P. sterbai which with rounded-obtuse posterior angles; lateral margins with one mid-lateral seta near maximum width, with one posterior seta near posterior angle; disc flat to slightly convex, often with transverse wrinkles aside median line. Basal foveae composed of one or two shallow grooves, inner groove about one-third length of pronotum, outer groove short or absent; area between outer groove and lateral margin ridged or not; basal foveal area often finely punctate. Elytra parallel-sided or slightly elliptic, width a little greater than half of length, widest a little behind middle, with similar microsculpture in males and females, isodiametric or transverse. Humerus distinct, humeral angles more or less toothed and projected outward. Striae deeply incised, punctate or not, intervals slightly convex; parascutellar pore absent or present; parascutellar stria joined to stria 1, angular base of stria 1 usually short, free or joined to stria 1; interval 3 with one to three setigerous pores: the basal one (if present) adjacent to stria 3, the rest two adjacent to stria 2, the middle pore always present, the apical pore absent only in P. filum ; interval 5 without setigerous pore; umbilicate pore series on interval 9 discontinuous in middle, often with 5–6 pores in basal group, 2–3 pores in the middle, 9–10 pores in apical group. Metepisternum slightly longer than basal width; sternite VII with one seta on each side in males, two in females; male abdominal sexual modification absent in most species, present in two species ( P. sterbai Jedlička and P. jiuzhaiicus sp. nov.), with simple median depression or depressed and longitudinally ridged on sternite VII ( Figs 48–49 ). Mesofemora with two setae on posterior ventral margin; metacoxae with three setae; metatrochanters with one seta; metatarsus slender or thickened; fifth tarsomere glabrous ventrally; apical half of inner margin of meso- and metatibiae crenulate in males. Male genitalia relatively small in compare to other Pterostichus groups; ventral surface simple, almost smooth, without ridge; apical lamella relatively short with rounded apex, often more or less twisted to right side; apical orifice opened to dorsal-left side. Right paramere strongly elongated, often curved before midpoint, apex thickened or sharp ( Figs 15–24 ). Endophallus simple, left apical lobe (la) larger and spherical, right apical lobe (ra) smaller and sometimes with slightly sclerotized apex, a small sclerotized projection present near gonopore ( Figs 25–30 ). Gonocoxite 2 of ovipositor almost identical across in species: falcate in ventral view, length about four times as basal width; outer margin with two ensiform setae, inner margin without ensiform seta; apex a little sharp in lateral view, with two nematiform setae in a groove ( Fig. 53 ). Spermatheca with seminal canal and receptaculum well differentiated; receptaculum straight and digitate, surface smooth, base without sclerotized projection; seminal canal five to six times length of receptaculum; spermathecal gland inserted on base of receptaculum ( Fig. 52 ).

Habitat. Species of the subgenus Tschitscherinea were usually found on the edge of mixed or coniferous forests in median-high altitudes (2600–4200 m) in southwest China. Adults tend to live under loose bark of fallen or standing dead trees. Different from other groups of Pterostichus , as they are more readily found under or in dead wood, rather than by pitfall traps.

Taxonomical notes. The subgenus Jedlickaia contains only one species P. sterbai distributed in North Yunnan, and was supposed to be related to subgenera Tschitscherinea or Megabea (Sciaky, 1997) . After examining the types of P. sterbai , it is found that this species is in accord with subgenus Tschitscherinea in the following important aspects: submentum with two setae on each side; metacoxae with three setae; elytra with distinct humeral tooth; umbilicate pore series on interval 9 discontinuous in middle; apical lamella of aedeagus very short; right paramere elongated; endophallus simple and strongly bent ventrally, with la and ra small and in similar size. Moreover, P. sterbai has two uncommon features among Chinese Pterostichus : (1) metatarsus distinctly thickened in males ( Figs 14 , 43 ); (2) eyes small with strongly swollen tempora. These two features also present in two other species belonging to the subgenus Tschitscherinea ( P. spissipedes sp. nov. and P. spectabilis sp. nov.) distributed in Yunnan province as well. Thus, these two species could be the closest relatives to P. sterbai ( Figs 13, 14 ). Additionally, P. sterbai is different from most other species of Tschitscherinea in having a straight right paramere of male genitalia ( Fig. 15 ) and sexual modification on sternite VII ( Fig. 49 ). But P. jiuzhaiicus sp. nov. is similar to P. sterbai in these two aspects ( Figs 19 , 48 ). Therefore, despite the circular pronotum which is different from all other species of Tschitscherinea , P. sterbai fits all characters of this subgenus. In conclusion, the subgenus Jedlickaia Sciaky, 1997 is newly proposed as a junior synonym of Tschitscherinea Berg, 1898 .

For most of the Chinese Pterostichus subgenera, the metacoxa has only two setae close to the outer-posterior angle and the anterior margin respectively. Metacoxae have one additional seta near the inner angle (see Shi & Liang, 2015: Fig. 137) only in the following four Chinese subgenera: Tschitscherinea Berg, 1898 , Morphohaptoderus Tschitschérine, 1898, Cryobius Chaudoir, 1838, and Huaius Tian & Huang, 2019 . Based on this synapomorphy (character polarity determined by Bousquet 1999: 35) and phenetic similarities, these four subgenera are considered to be closer to each other than to other subgenera of the Chinese fauna.

Compared to the subgenus Huaius , Tschitscherinea is different in having a parallel-sided body form, subquadrate or ovate pronotum, and discontinuous umbilicate series on interval 9. whereas, In the troglobitic Huaius , pronotum is more cordate with lateral margins distinctly sinuate before posterior angles and the umbilicate pore series on interval 9 continuous. The subgenus Cryobius contains more than a hundred species with a Holarctic distribution. In China, there are only a few species belonging to the subgenus distributed in the Northeast. For all the Chinese species of Cryobius , they are distinguishable from Tschitscherinea by having a smaller and stouter body form, and the fifth tarsomere setose ventrally.

Morphohaptoderus is distributed in South China, North Vietnam, and North Myanmar and has 36 described species, and probably more than that undescribed. The subgenus Tschitscherinea is suspected to be closest to Morphohaptoderus . However, it is difficult to infer the systematic relationship between these two subgenera before the phylogenetic relationship within Pterostichus is better known. In most cases, these two subgenera can be distinguished through the following morphological features but not always: (1) In Tschitscherinea the body form is generally more elongated and parallel-sided; whereas in Morphohaptoderus , if the body is somewhat parallel-sided then the body size is stouter and smaller (less than 9 mm). (2) In Tschitscherinea the fifth tarsomere is always glabrous ventrally; whereas in Morphohaptoderus , the fifth tarsomere is usually setose ventrally with the exception of four described species ( P. geberti Sciaky & Wrase, 1997 , P. glabellus Fedorenko, 2023 , P. primitivus Fedorenko, 2023 , and P. unisetosus Fedorenko, 2023 ), and additionally in some undescribed species. (3) In Tschitscherinea , the apical lamella of aedeagus is small with a simple rounded apex; whereas in Morphohaptoderus , the apical lamella is generally larger, often toothed or twisted. (4) In Tschitscherinea , the right paramere of aedeagus is elongated, with the length of the apical portion (area distal to the articulation) at least fourth as width, curved or nearly straight, apex blunt or gradually attenuate ( Figs 15–24 ); whereas in Morphohaptoderus , the right paramere of aedeagus is usually very short, with length of apical portion at most twice as width, apex widely rounded ( Fig. 33 ). The only exception of Morphohaptoderus is in P. dundai Sciaky, 1994 which has the apical portion of right paramere widely triangular with an abruptly narrowed and elongated apical extension. (5) In Tschitscherinea , the endophallus is relatively small and simple, only with a spherical left apical lobe and a very small right apical lobe; whereas in Morphohaptoderus , the endophallus is diverse in form, usually much larger with several lobes in more complicated shapes ( Figs 56–57 ). Among these characters mentioned above, the shape of the right paramere of aedeagus can be well-define in these two subgenera, because this character is always reliable for all known species in both subgenera.

The characters of the subgenus Huaius were based on Tian et al. (2019). The characters of the Vietnamese species of subgenus Morphohaptoderus were based on Fedorenko (2023). The characters of subgenus Cryobius and other Morphohaptoderus species were based on the following specimen examinations:

Pterostichus (Cryobius) jaechi Krischenhofer : 1 male and 1 female (IZAS), “ Jilin, Changbaishan, 2500m, 1987.VII.20–21, Peiyu Yu leg.”. Pterostichus (Morphohaptoderus) janatai Sciaky & Wrase : 2 males and 1 female (BJFU), “ Shaanxi, Ningshan County, Pingheliang, N33.4718, E108.4961, 2110m, 2017.VII.2–9, Weifeng Yan pitfall trap.”. Pterostichus (Morphohaptoderus) schuelkei Sciaky & Wrase : 1 male and 1 female (BJFU), “ Shaanxi, Ningshan County , Huoditang , 2017.VII.2–9, Weifeng Yan et al. pitfall trap ”. Pterostichus (Morphohaptoderus) lingshanus Sciaky & Wrase : 1 male and 1 female (BJFU), “ Shaanxi, Ningshan , Pingheliang , N33.4718, E108.4961, 2110m, 2017.VII.2–9, Weifeng Yan pitfall trap.”. Pterostichus (Morphohaptoderus) dundai Sciaky : 1 male and 1 female (IZAS), “ Shaanxi, 3km of Houzhenzi to old Foping Township , 2008.V.9, Hao Huang leg.”. Pterostichus (Morphohaptoderus) geberti Sciaky & Wrase : 1 male and 1 female (IZAS), “ Shaanxi, Zhouzhi , Houzhenzi , 2007. V.25, Hongliang Shi lgt.”.

Key to species of subgenus Tschitscherinea Berg, 1898

1 Elytra with isodiametric microsculpture, length of each mesh less than 1.5 times than width ( Figs 44, 46 )............... 2

- Elytra with transverse microsculpture, length of each mesh more than 2 times than width ( Figs 45, 47 ).................. 5

2 Elytral interval 3 with three setigerous pores, the first one adjacent to stria 3, the rest two adjacent to stria 2; eyes large, tempora shorter than eyes; male metatarsus slender, tarsomere 1 with length/width = 3.1–3.5 ( Fig. 41 ); Yunnan (Zhongdian).............................................................................................. P. farkaci Sciaky

- Elytral interval 3 with two setigerous pores, both adjacent to stria 2; eyes small, tempora swollen, longer than eyes; male metatarsus in a stout form, tarsomere 1 with length/width = 1.7–2.2 ( Figs 35–38, 43 )................................ 3

3 Body larger, BL = 14.8–15.5 mm; head very large, HW/PW = 0.78–0.81; basal foveae with inner groove about 1.5 times length as outer groove; Yunnan (Ninglang), Sichuan (Yanyuan)........................................ P. spectabilis sp. n.

- Body smaller, BL = 10.3–11.6 mm; head smaller, HW/PW = 0.64–0.69; basal foveae with inner groove about twice length as outer groove......................................................................................... 4

4 Pronotum more quadrate, lateral margins subparallel near middle; elytra parascutellar pore present; maxim width of head at outer margin of eyes; right paramere of male genitalia curved with apex truncated ( Figs 16–17 ); male sternite VII without modification; Yunnan (Lijiang, Dali, Deqin)................................................. P. spissipedes sp. n.

- Pronotum more ovate, lateral margins distinctly arched near middle; elytra parascutellar pore absent; maxim width of head at outer margin of tempora; right paramere of male genitalia nearly straight with apex relatively sharp ( Fig. 15 ); male sternite VII with a round depression ( Fig. 49 ); Yunnan (Dayao)............................................ P. sterbai Jedlička

5 Elytral parascutellar pore present, occasionally absent on one side; pronotum basal foveae almost impunctate............ 6

- Elytral parascutellar pore absent; pronotum basal foveae distinctly punctate....................................... 7

6 Elytral microsculpture stronger, transverse-isodiametric, length of each mesh usually 2–3 times as width ( Fig. 45 ); elytra humeral tooth smaller ( Figs 7–8 ); right paramere of male genitalia curved with the maximum curvature near apical fourth of apical portion ( Fig. 20 ); Sichuan (Muli)...................................................... P. mulensis Sciaky

- Elytral microsculpture shallower, transverse-linear, length of each mesh usually more than 4 times as width ( Fig. 47 ); elytra humeral tooth larger ( Fig. 9 ); right paramere curved with the maximum curvature near middle of apical portion ( Fig. 21 ); Yunnan ( Haba Mt. )......................................................................... P. krali Sciaky

7 Elytral interval 3 with one setigerous pore; eyes small, tempora strongly swollen and distinctly longer than eyes, head maxim width at outer margin of tempora; pronotum basal foveae without outer groove; Sichuan (Baoxing, Wenchuan)............................................................................................ P. filum (Tschitschérine)

- Elytral interval 3 with three setigerous pores; eyes large, tempora shorter than eyes, head maxim width at outer margin of eyes; pronotum basal foveae with inner and outer grooves well defined; Sichuan (Jiuzhaigou, Abaco), Gansu (Bailongjiang).......................................................................................... P. jiuzhaiicus sp. n.

Figures

FIGURES 13–18. 13–14. Habitus and labels of P. sterbai Jedlička: (13) holotype. (14) a male paratype. Scale bars: 2 mm. 15–18. Male genitalia of Pterostichus (Tschitscherinea) spp., left lateral view, dorsal view of median lobe of aedeagus, and right paramere. (15) P. sterbai Jedlička, Pe-Yen-Tsin. (16) P. spissipedes sp. nov., holotype. (17) P. spissipedes sp. nov., Deqin, paratype. (18) P. spectabilis sp. nov., holotype. Scale bars: 0.5 mm.

FIGURES 19–24. Male genitalia of Pterostichus (Tschitscherinea) spp., left lateral view, dorsal view of median lobe of aedeagus, and right paramere. (19) P. jiuzhaiicus sp. nov., holotype. (20) P. mulensis Sciaky, holotype. (21) P. krali Sciaky, Haba Snow Mountain. (22) P. farkaci Sciaky, west of Zhongdian, paratype. (23) P. farkaci Sciaky, Tianchi, Zhongdian. (24) P. farkaci Sciaky, Hongshan, Zhongdian. Scale bars: 0.5 mm.

FIGURES 48–53. 48–49. Male sternite VII of Pterostichus (Tschitscherinea) spp., (48) P. jiuzhaiicus sp. nov., Jiuzhaigou, paratype. (49) P. sterbai Jedlička, Pe-Yen-Tsin. Scale bars: 0.5 mm. 50–53. Female genitalia of Pterostichus (Tschitscherinea) mulensis Sciaky, Muli. (50) tergite VIII. (51) sternite VIII. (52) female internal reproductive tracts. (53) ventral view of gonocoxites. Abbreviations: g1 gonocoxite 1, g2 gonocoxite 2, es ensiform setae, ns nematiform setae, bc bursa copulatrix, co common oviduct, sc seminal canal, rc receptaculum, sg spermathecal gland, spc spermathecal canal. Scale bars: 0.5 mm.

FIGURES 25–30. Endophallus of Pterostichus (Tschitscherinea) spp., left lateral view and right lateral view. (25) P.spissipedes sp. nov., Jianchuan, paratype. (26) P. jiuzhaiicus sp. nov., holotype. (27) P. mulensis Sciaky, Muli. (28) P. farkaci Sciaky, Hongshan, Zhongdian. (29) P. farkaci Sciaky, Tianchi, Zhongdian. (30) P. sterbai Jedlička, Pe-Yen-Tsin. Abbreviations: gp gonopore, la left apical lobe, ra right apical lobe, va ventral apical lobe. Scale bar: 0.5 mm.

FIGURES 35–47. 35–43. Metatarsi of Pterostichus (Tschitscherinea) spp. (35) P. spissipedes sp. nov., male. (36) P. spissipedes sp. nov., female. (37) P. spectabilis sp. nov., male. (38) P. spectabilis sp. nov., female. (39) P. mulensis Sciaky, male. (40) P. mulensis Sciaky, female. (41) P. farkaci Sciaky, male. (42) P. krali Sciaky, male. (43) P. sterbai Jedlička, male. Scale bar: 1 mm. 44–47. Elytral microsculpture of Pterostichus (Tschitscherinea) spp. (44) isodiametric, P. specillatus sp. nov., holotype. (45) transverse-isodiametric, P. mulensis Sciaky, a male from Muli. (46) isodiametric, P. farkaci Sciaky, a male from Tianchi, Zhongdian. (47) transverse-linear, P. krali Sciaky, a male from Haba Snow Mountain, Zhongdian. Scale bar: 0.2 mm.

FIGURES 31–34. habitus and male genitalia of P. parvicollis Sciaky & Wrase. (31) habitus of holotype. (32–34) a male specimen from Pingheliang, Shaanxi. (32) habitus. (33) male genitalia, left lateral view, dorsal view of median lobe of aedeagus, and right paramere. (34) endophallus. Abbreviations: gp gonopore, gpp gonopore piece, la left apical lobe, ra right apical lobe. Scale bars: 2 mm (31–32), 0.5 mm (33–34).

FIGURES 54–57. 54. Confirmed distribution of the Pterostichus (Tschitscherinea) spp. in China: Red P. spissipedes sp. nov. Blue P. spectabilis sp. nov. Black P. jiuzhaiicus sp. nov. Pink P. filum (Tschitschérine). Purple P. mulensis Sciaky. Green P. fakarci Sciaky. Yellow P. krali Sciaky. Cyan P. sterbai Jedlička. 55–57. Endophallus of Pterostichus spp., right lateral view and left lateral view: (55) Pterostichus (Cryobius) jaechi Krischenhofer, Changbaishan, Jilin. (56) Pterostichus (Morphohaptoderus) janatai Sciaky & Wrase, Pingheliang, Shaanxi. (57) Pterostichus (Morphohaptoderus) dundai Sciaky, Houzhenzi, Shaanxi. Scale bars = 0.5 mm, a (55), b (56–57).

FIGURES 7–12. Habitus of Pterostichus (Tschitscherinea) spp. (7) P. mulensis Sciaky, a male from Muli. (8) P. mulensis Sciaky, holotype. (9) P. krali Sciaky, holotype. (10) P. farkaci Sciaky, a male from Tianchi, Zhongdian. (11) P. farkaci Sciaky, a male from Hongshan mountain, Zhongdian. (12) P. farkaci Sciaky, a male paratype from west of Zhongdian. Scale bars: 2 mm.