The genus Pterostichus in China II: the subgenus Circinatus Sciaky, a species revision and phylogeny (Carabidae, Pterostichini)

Abstract All of the known species of the Chinese endemic subgenus Pterostichus (Circinatus) are revised, keyed, and illustrated. Eleven new species and one new subspecies are described: Pterostichus adelphus sp. n. (Sichuan: Meigu, N28.66°, E103.06°); Pterostichus ailaoicus sp. n. (Yunnan: Xinping, N23.94°, E101.50°); Pterostichus camelus sp. n. (Sichuan: Mianning, N28.97°, E102.16°); Pterostichus dimorphus sp. n. (Yunnan: Dayao, N26.08°, E101.03°); Pterostichus maitreya sp. n. (Guizhou: Fanjingshan, N27.90°, E108.70°); Pterostichus miao sp. n. (Guangxi: Maoershan, N25.87°, E110.41°); Pterostichus tumulus sp. n. (Guizhou: Fanjingshan, N27.90°, E108.70°); Pterostichus wangjiani sp. n. (Yunnan: Dongchuan, N26.08°, E102.87°); Pterostichus yan sp. n. (Hubei: Shennongjia, N31.47°, E110.39°); Pterostichus yuxiaodongi sp. n. (Sichuan: Wolong, N30.99°, E103.15°); Pterostichus zhygealu sp. n. (Sichuan: Meigu, N28.67°, E103.05°); and Pterostichus cavazzutianus mianningensis subsp. n. (Sichuan: Mianning, N28.97°, E102.16°). Pterostichus cavazzutianus is proposed as a replacement name for Pterostichus cavazuttii Allegro and Sciaky 2010, preoccupied by Pterostichus (Sinosteropus) barbarae cavazuttii Sciaky and Facchini 2003. A lectotype is designated for Pterostichus baenningeri Jedlička 1931. Two species, Pterostichus schuelkei Sciaky & Wrase and Pterostichus wenxianensis Allegro & Sciaky, are moved from the subgenus Circinatus to Morphohaptoderus. An infra-subgeneric taxonomy is proposed for the subgenus Circinatus with four species groups. The male endophallus characters for most species of Circinatus were well studied, with three types of endophallus defined. A phylogenetic analysis based on adult morphological characters confirmed the infra-subgeneric classification and clarified some of the relationships among species. Two main lineages within Circinatus were identified from the phylogenetic analyses. Three of the four species groups were monophyletic, whereas the fourth group was paraphyletic.


Introduction
In 2011, we began conducting a comprehensive investigation and taxonomic study of the Pterostichus of China. Our first contribution to the Chinese Pterostichus was the publication two years ago on the endemic subgenus Wraseiellus (Shi et al. 2013). In this paper, we focus on the subgenus Circinatus of the genus Pterostichus, which is also endemic to China. Following an expedition in 2012, a review of Circinatus was planned to describe some new species of the subgenus that were discovered in the provinces of Sichuan, Yunnan and Guizhou. During the expedition, most of the eleven previously described species were also collected. In this paper, eleven new species and one new subspecies are described. We expect to discover more new species of Circinatus in the near future, since the study of the Chinese Pterostichus fauna is still in its early stages compared with its high diversity.
The taxonomic value of the everted endophallus of Carabidae has been recognized in recent decades and was early on applied in some works on the genus Carabus (e.g., Ishikawa 1978, Imura et al. 1998). In the genus Pterostichus, the endophallic characters were also of value for both higher systematics (Sasakawa and Kubota 2007) and separation of similar species (Sasakawa 2009). However, in the Chinese Pterostichus, the endophallus was very poorly studied but it is now known for most of the Circinatus species (16 of 22). Information on this structure improves our understanding of the Chinese Pterostichus fauna and also provides solid support for the species determinations and the relationships among species described in the present paper.
The primary purposes of this paper are to (1) describe 12 new taxa of Circinatus; (2) provide a key for species determinations in this subgenus; (3) describe and illustrate the endophallus of most known species of Circinatus and estimate the rate of evolution of the endophallus in this subgenus; and (4) conduct a primary analysis of the relationships among species and classify the subgenus Circinatus into species groups. males with secondary sexual structures on terminal or penultimate sternum, or without such structures on sterna. Mesofemur with two setae on posterior margin, with single short spine subapically; metacoxa with two setae; metatrochanter with one seta; first metatarsomere with distinct carina on outer surface, more or less shallower in second and third metatarsomeres; fifth tarsomeres usually glabrous beneath, rarely setose. Male genitalia with apical orifice placed on dorsal side of aedeagus or somewhat twisted to left side; right paramere straight, slightly elongate, ratio length / width 2.5-4, apex rounded or obtuse. Endophallus variable, located on dorsal or ventral side of aedeagus. Female genitalia with spermatheca markedly elongate, seminal canal and receptaculum differentiated; receptaculum capitate or clavate; spermathecal gland inserted near base of receptaculum. Stylomere II saber-shaped, elongate, strongly bent outwards; outer margin with two (rarely three) ensiform setae, inner margin with one (rarely two) ensiform seta; two short nematiform setae located in a furrow near apex (Fig. 138). Female sternum VIII with transparent region in middle, V-shaped, triangular or quadrate, apical margin with fine setae or spines; female tergum VIII with apical half evenly chitinized.
Comparison. The majority of the species in the subgenus Circinatus (16 of 22 known species) have a rounded pronotal hind angle and the pronotal posterior seta is distant from the hind angle. In the Chinese fauna of Pterostichus, six subgenera have this type of pronotum: Eosteropus Tschitschérine, Oreolyperus Tschitschérine, Circinatus Sciaky, Gutta Wrase & Schmidt, Tubuliphallus Sciaky & Allegro, and Sinosteropus Sciaky. Two of these subgenera have distributions that are distant from the others: Eosteropus is widely distributed in the Palearctic realm, and Oreolyperus is restricted to middle Asia. The distributions of the other four subgenera are close in range in southern China. In comparison with Eosteropus and Gutta, Circinatus differs in having the elytral third interval usually with two setigerous pores, all pores adjacent to the second stria (the third interval usually with three pores, the first one adjacent to the third stria in the other two subgenera). Compared to Oreolyperus, Circinatus differs in having the elytral basal pore present (elytral basal pore absent in Oreolyperus). Compared to Tubuliphallus, Circinatus differs in having the elytral length more than 0.6 times the body length, apical orifice of aedeagus opened to the dorsal or left side (in Tubuliphallus, elytral length approximately 0.5 times the body length, apical orifice of aedeagus opened to the apex). Compared to Sinosteropus, Circinatus differs in: body size large (9.5-16.5 mm); body form less convex and relatively slender; pronotal basal fovea with the outer area of the inner groove usually flat or concave; right paramere straight and only slightly elongate. In Sinosteropus, the body size is small (6-10 mm); body form strongly convex and relatively stout; pronotal basal fovea with the outer area of the inner groove convex; right paramere usually strongly elongate, apex more or less bent.
Some species of the subgenera Morphohaptoderus Tschitschérine and Neohaptoderus Tschitschérine may have similar rounded pronotal hind angles. However, in these two subgenera, the posterior seta of the pronotum is always located very close to the hind angle, which distinguishes these subgenera from most species of Circinatus. Some species of Morphohaptoderus have pronota shaped similarly to Circinatus, but have a different number of setae on the metacoxa and male genitalia (for details see discussion under P. schuelkei).
Description. Body form relatively stout, body length 9.9-11.1 mm; dorsal side black, shining; elytron with faint iridescent shine; mouthparts, antenna and tarsus yellowish brown; ventral side almost black. Both sexes with similar elytral microsculpture, very faint and linear. Head. Frons without punctures; antenna reaching elytron basal fifth; gena approx same length as eye, briefly tumid behind eye. Pronotum round, widest before middle, PW/PL = 1.21-1.32; usually four (sometimes three) mid-lateral setae present, first one close to anterior angle, last one near middle of lateral margin, a little distant from other three; posterior seta far distant from hind angle, distance between seta and hind angle approx same as distance between hind angle and inner basal foveal groove; hind angle completely rounded; basal fovea shallow, faintly defined; inner groove subparallel to median line, slightly curved outwards; outer groove completely vanished, outer area of inner groove flat; basal foveal area usually impunctate, sometimes with very sparse punctures on inner area of inner groove. Elytron oviform, basal ridge slightly oblique; elytral shoulder slightly narrowed, basal ridge and lateral margin forming obtuse angle, humeral tooth indistinct; intervals feebly convex; striae moderately deep, without punctures; scutellar stria short but complete; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 15-16 pores (6, 1, 8-9). Ventral side. Proepisternum impunctate or slightly punctate near posterior margin; mesepisternum densely punctate; metepisternum impunctate; terminal or penultimate sternum of males not modified. Legs. Fifth tarsomeres glabrous beneath; males with apical half of mesotibia slightly widened, inner margin slightly crenulate; first metatarsomere with distinct carina on outer surface, such carina on second and third metatarsomeres superficial. Male genitalia. Median lobe of aedeagus bent approx 90 degrees, apex slightly bent ventrally (Fig. 34A); ventral margin straight in middle, dorsal margin gradually curved; apical orifice large, slightly turned to left side, not opened on ventral side; apical lamella short, approx one fourth length of apical orifice, laminate, not thickened; in dorsal view, apical lamella located at middle of aedeagal apex, approx triangular with rounded apex, wide and short, length approx equal to basal width (Fig. 34B). Right paramere straight and slender, length approx 3.5 times greatest width, apex rounded (Fig. 34C). Endophallus (Fig. 34D, E, F) short, bent to ventral side, across apical lamella, and then turned to aedeagal base; gonopore (gp) located at approx same level as apical lamella. Six distinct lobes recognized: ventral-basal lobe I (vb-I) very small, close to base of apical lamella, membranous with its upper surface scaled; ventral-basal lobe II (vb-II) large and strongly chitinized, arcuate, strongly pointed, its upper surface with a membranous structure connected with vb-III; ventral-basal lobe III (vb-III) small, rounded with apex pointed, located on left side of vb-II, evenly decorated with very fine scales; ventral-apical lobe (va) short and wide, almost covered by vb-II, heavily pigmented, decorated with coarse scales, located on right side of vb-II; pre-apical lobe (pa) narrow and long, tubiform, without decoration, located on right lateral side; right lobe (rl) large, much thicker and a little longer than pa in ventral view, apex truncate, evenly decorated with fine scales. Female genitalia. Spermatheca ( Fig. 53) with seminal canal approx three times as long as receptaculum; receptaculum capitate, club approx half length of receptaculum; spermathecal gland slightly expanded; seminal canal inserted at base of common oviduct, base of seminal canal sclerotized. Stylomere II with two ensiform setae at outer margin and one at middle of inner margin, two very short nematiform setae located in furrow near the apex. Female sternum VIII ( Fig. 71B) with dense and fine spines on posterior margin; posterior margin rounded, shallowly notched in middle; posterior region chitinized, anterior region semi-chitinized, middle transparent region V-shaped, not adjacent to posterior notch in middle. Female tergum VIII ( Fig. 71A) with anterior third well chitinized, posterior two thirds semi-chitinized and pigmented with dense spots.
Distribution. This species was known only from the border between Meigu and Ebian counties (Sichuan, Liangshan Yi Autonomous Prefecture). The two collecting localities are approx 2 km apart. (Map 2) Etymology. The name adelphus derives from the Greek epithet "adelph" which means brother, referring to the similarity of this new species to a sympatric species P. cavazzutianus.
Affinities. This new species seems to be close to P. cavazzutianus in their similarities in the elytral microsculpture and short apical lamella of aedeagus.
Habitat. This species was collected in mixed forest with dominant large pines, and rich in dead logs. Most specimens were collected by pitfall trap, and also one was found under or in dead logs. In Yizi pass, P. adelphus sp. n. was found together with other three Circinatus species (P. bullatus, P. cavazzutianus s. str., P. zhygealu sp. n.), but seems rarer than other species. This locality has the richest diversity of the subgenus Circinatus.
Variation. Two paratypes have three setigerous pores on one or both elytra. Diagnosis. Pronotum with single mid-lateral seta; posterior seta located almost at hind angle; hind angle forming indistinct obtuse angle, lateral margin straight before hind angle; elytral microsculpture almost invisible; elytral shoulder angle distinct; males with two large tubercles on terminal sternum; tubercles close to each other, distance between tubercles less than distance between setae on penultimate sternum; fifth tarsomeres glabrous beneath. Apical lamella of aedeagus long, its length approx 2.1 times basal width, apex widened, with a small tooth on right margin. Comparisons between similar species are given in Table 1 under P. maitreya sp. n.
This new species is similar to P. baenningeri in body size and form, pronotal shape, and male terminal sternum. Despite the significant differences in male genitalia (shape of apical lamella, see Figs 29B and 40B), they also differ in: (1) P. ailaoicus sp. n. have the fifth tarsomeres glabrous beneath, but these are setose in P. baenningeri; (2) P. ailaoicus sp. n. have the two tubercles on the male terminal sternum close to each other, distance between tubercles less than the distance between setae on penultimate sternum; but in P. baenningeri, distance between tubercles on terminal sternum is greater than the distance between setae on penultimate sternum.
Description. Body form fairly elongate, body length 10.9 mm; dorsal side almost black, moderately shining, elytron with faint iridescent shine; mouthparts, antenna, tarsus, tibia, and apex of femur reddish brown; ventral side blackish. Elytral microsculpture weak, barely visible, linear. Head. Frons without punctures; antenna reaching elytron basal sixth; gena shorter than length of eye, briefly tumid behind eye. Pronotum round, lateral margin curved in middle, nearly straight before hind angle, widest at approx anterior two fifths; posterior margin a little narrower than anterior margin; PW/PL = 1.10; one mid-lateral seta present, located a little before greatest width; posterior seta close to but a little anterior to hind angle; hind angle forming indistinct obtuse angle; basal fovea shallow, faintly defined; inner groove subparallel to median line, not reaching posterior margin; outer groove completely vanished; outer area of inner groove flat; basal foveal area finely punctate along inner groove. Elytron oviform, with basal ridge almost straight; elytral shoulder moderately widened, shoulder angle between basal ridge and lateral margin forming obtuse angle, humeral tooth very small, not jutting out; intervals feebly convex; striae moderately deep, without punctures; scutellar stria short, apex free; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 16-17 pores (6, 1, 9-10). Ventral side. Proepisternum without punctures, mesepisternum and metepisternum finely punctate; penultimate sternum not modified; terminal sternum with two large tubercles vaguely defined, approx one third length of terminal sternum; tubercles located a little before middle of sternum, distance between tubercles less than distance between two setae on penultimate sternum; region between tubercles not depressed. Legs. Fifth tarsomeres glabrous beneath; males with apical half of mesotibia slightly widened, inner margin crenulate; first metatarsomere with distinct carina on outer surface, such carina superficial on full length of second metatarsomere. Male genitalia. Median lobe of male genitalia bent approx 90 degrees, apex gradually bent ventrally (Fig. 29A); ventral margin slightly curved in middle, weakly sinuate before apex; dorsal margin gradually curved; apical orifice large, slightly turned to left side, opened on ventral side; in lateral view, apical lamella very long, laminate with base slightly thickened, sinuate before apex, apex pointing apical-ventrally, its length approx one third length of apical orifice; in dorsal view, apical lamella located on right side of aedeagal apex, its base not grooved on dorsal surface; apical lamella long with rounded apex, length approx 2.1 times basal width, distinctly widened at approx apical third, with a small tooth on right margin (Fig. 29B). Right paramere straight and stout, apical half fusiform, slightly enlarged and then narrowed to apex, inner margin evenly curved before apex; length approx three times greatest width; apex obtuse ( Fig.  29C). Endophallus not studied. Female genitalia unknown.
Distribution. This species is known only from the holotype collected from Ailaoshan Mt. in Yunnan Province (Map 1). The altitude is 2351 m.
Etymology. This new species is named for its type locality, Ailao Mountain. Affinities. Pterostichus ailaoicus sp. n. is close to P. baenningeri, P. maitreya sp. n., P. miao sp. n., and P. yan sp. n. in their similarities of male terminal sternum and male genitalia. Diagnosis. Body rather elongate; pronotum with three or four mid-lateral setae; hind angle completely rounded; basal fovea almost impunctate; fifth tarsomeres setose beneath; males with two large tubercles on penultimate sternum. This species can be easily distinguished from all other Circinatus species of agilisgroup (with multi-setae on pronotal lateral margin) by the setose fifth tarsomeres, and by the very special male secondary sexual character on penultimate sternum.

Pterostichus
Description. Body form fairly slender, body length 12.7-13.5 mm; dorsal side dark brown, moderately shining; elytron without iridescent shine; mouthparts, antenna, tarsus, and apex of tibia yellowish brown; ventral side brownish. Both sexes with distinct isodiametric microsculpture on elytron. Head. Frons without punctures; antenna reaching elytron basal fifth; right mandible without distinct tooth; left mandible with a tooth near base; gena a little shorter than length of eye, very briefly tumid behind eye. Pronotum distinctly narrowed to base, lateral margin almost straight before hind angle, widest before middle, PW/PL = 1.13-1.20; three or four mid-lateral setae present, first one close to anterior angle, last one near middle of lateral margin, setae usually evenly separated; posterior seta far distant from hind angle, distance between seta and hind angle approx same as distance between hind angle and inner basal foveal groove; hind angle completely rounded; basal fovea shallow, faintly defined; inner groove subparallel to median line, curved outwards; outer groove completely vanished, outer area of inner groove faintly depressed; basal foveal area more or less punctate around inner groove. Elytron oviform, with basal ridge almost horizontal; elytral shoulder distinctly narrowed, basal ridge and lateral margin forming obtuse angle, humeral tooth indistinct; intervals feebly convex; striae moderately deep, with faint punctures inside; scutellar stria short and complete; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 15-17 pores (6, 1-2, 8-9). Ventral side. Proepisternum and mesepisternum finely punctate; metepisternum impunctate; penultimate sternum of males ( Fig. 118) with two large tubercles in middle, occupying posterior half of sternum, primary setae of penultimate sternum inserted on outer surface of each tubercle; male terminal sternum not modified. Legs. Fifth tarsomeres with three to four pairs of setae beneath; males with apical half of mesotibia not widened, inner margin not crenulate; first metatarsomere with distinct carina on outer surface, such carina on second and third metatarsomeres superficial. Male genitalia. Median lobe of male genitalia bent less than 90 degrees, apex gradually bent ventrally (Fig. 28A); ventral margin straight in middle, dorsal margin gradually curved; apical orifice large, slightly turned to left side, not opened on ventral side; apical lamella long, approx one third length of apical orifice, laminate with apex slightly thickened; in dorsal view, apical lamella slightly inclined right, nearly triangular with rounded apex, length approx equal to its basal width (Fig. 28B). Right paramere straight and stout, inner margin slightly ex-panded near middle, length approx 2.5 times greatest width, apex rounded (Fig. 28C). Endophallus not studied. Female genitalia. Spermatheca with seminal canal approx five times as long as receptaculum; receptaculum capitate ( Fig. 60), club approx half length of receptaculum; seminal canal inserted at base of common oviduct, base of seminal canal sclerotized. Stylomere II with two ensiform setae at outer margin and one at basal third of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 76B) with dense and fine spines on posterior margin; posterior margin curved, deeply notched in middle; posterior region chitinized, anterior region semi-chitinized, middle transparent region V-shaped, adjacent to posterior notch in middle; three transparent patches present on each side. Female tergum VIII ( Fig. 76A) with major portion chitinized, mid-posterior region semi-chitinized with denser spots.
Distribution. This species is known only from a single locality in Yele Reserve, Sichuan, Liangshan Yi Autonomous Prefecture. (Map 2) Etymology. The scientific name camelus comes from the generic name of the camel, referring to the two large tubercles on the male penultimate sternum of this new species, which are reminiscent of the Bactrian camel (Camelus bactrianus).
Affinities. This new species seems to be allied to P. agilis in that both species have male secondary sexual characters on the penultimate sternum and similar aedeagi.
Habitat. Pterostichus camelus sp. n. was collected in mixed forest with dominant large pines, rich in dead logs. Two specimens were collected by pitfall trap, and the third was found under or in dead logs. In Yele Reserve, P. camelus sp. n. was found together with two other Circinatus species, P. cavazzutianus mianningensis ssp. n., and P. zhygealu sp. n.
Variation. One female paratype has the elytral third and fourth striae briefly fused at approx basal third. Diagnosis. Pronotum with single mid-lateral seta; hind angle completely rounded; basal fovea almost impunctate; each sex with different elytral microsculpture: granular in females (elytron luster dull), and isodiametric as usual in males; terminal sternum of male weakly depressed in middle, with two sharp teeth near posterior margin pointing to apex of sternum.

Pterostichus
The females of this new species can be rapidly distinguished from all the other Circinatus species by the special granular elytral microsculpture. The males are similar to P. liciniformis. In addition to the different sexual characters on the male terminal sternum, P. liciniformis also differs from the new species by its smaller and stouter body form.
Description. Body form fairly slender, body length 11.5-11.9 mm; dorsal side almost black, moderately shining, female elytron dull; elytron without iridescent shine; mouthparts, antenna, tarsus, and apex of tibia dark brown; ventral side blackish. Microsculpture isodiametric on vertex, transverse on pronotum. Elytral microsculpture different in each sexes: isodiametric as usual in males; granular in females, making elytral luster much duller in females. Head. Frons without punctures; antenna hardly reaching elytron base; gena approx same length as eye, briefly tumid behind eye. Pronotum narrowed to base, posterior margin a little narrower than anterior margin; lateral margin evenly rounded, widest a little before middle, PW/PL = 1. 16-1.21; one mid-lateral seta present, located a little before greatest width; posterior seta far distant from hind angle, distance between seta and hind angle greater than distance between hind angle and inner basal foveal groove; hind angle completely rounded; basal fovea shallow, faintly defined; inner groove subparallel to median line, basal half oblique outwards; outer groove completely vanished, outer area of inner groove flat; basal foveal area with sparse and fine punctures on inner side of inner groove. Elytron oviform, with basal ridge slightly curved; elytral shoulder slightly narrowed, basal ridge and lateral margin forming obtuse angle, humeral tooth small but distinct; intervals almost even; striae moderately deep, with faint punctures inside; scutellar stria short, apex free; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 14-16 pores (5-6, [1][2][7][8][9]. Ventral side. Proepisternum impunctate, mesepisternum and metepisternum finely punctate; male terminal sternum weakly depressed in middle, with two sharp teeth close to posterior margin pointing to apex of sternum (Fig. 123). Legs. Fifth tarsomeres glabrous beneath; males with apical half of mesotibia slightly widened, inner margin not crenulate; only first metatarsomere with carina on outer surface, fairly distinct. Male genitalia. Median lobe of male genitalia bent approx 90 degrees, apex nearly straight, a little bent ventrally (Fig. 44A); ventral margin straight in middle, dorsal margin gradually curved; apical orifice large, slightly turned to left side, largely opened on ventral side (Fig. 133); in lateral view, apical lamella laminate, slightly twisted, apex not thickened, its length approx one fifth length of apical orifice; in dorsal view, apical lamella narrow and small, located on right side, pointing apically, length approx 1.5 times basal width, apex rounded (Fig. 44B). Right paramere straight and stout, inner margin slightly expanded near middle, length approx 3.5 times greatest width, apex rounded (Fig. 44C). Endophallus (Figs 44D, E, F) very thick, bent to ventral side from ventral opening of apical orifice, major portion of endophallus located on left-ventral side of aedeagus; gonopore (gp, gonopore lobe folded in Fig. 44) located close to aedeagal base, pointing to right side. Four distinct lobes recognized: left lobe (lf) large, apex bifid in left view, decorated with fine scales, located on left side of endophallus; right lobe (rl) large, pointed to right side with truncate apex in ventral view, located on ventral side of endophallus, adjacent to vl, with a small pigmented patch near base; ventral lobe (vl) smaller than rl, composed of two sub-lobes, with a narrow pigmented band across its full width; pre-apical lobe (pa) indistinct, weakly pointed, just before gp; basal lobe absent. Ventral piece (vp, only partly shown in Fig. 44F) is a short and arcuate chitinized piece, located just behind gp. Female genitalia. Spermatheca with seminal canal approx two times as long as receptaculum; receptaculum clavate ( Fig. 65); seminal canal inserted at base of common oviduct, base of seminal canal with a very fine sclerotized spine. Stylomere II with two ensiform setae at outer margin and one at middle of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII ( Fig. 81B) with dense and fine setae on posterior margin; posterior margin curved, deeply notched in middle; posterior region chitinized, anterior region semi-chitinized, pigmented with sparse denser areas; anterior margin notched in middle; middle transparent region arrow-shaped, adjacent to posterior and anterior notches in middle. Female tergum VIII ( Fig. 81A) with most of region semi-chitinized, pigmented with dense spots, only lateral-anterior region with two chitinized patches.
Distribution. This species is known only from Xiaobaicaoling Mts., Dayao County of north Yunnan (Map 1). The two collecting localities are very close, approx 1 km apart. The altitude range is 2881-3105 m.
Etymology. The scientific name "dimorphus" of the new species comes from Greek meaning "two forms", referring to the different elytral luster between male and female.
Affinities. This new species appears close to P. liciniformis and has the following similarities: the pronotal hind angle completely rounded, males with the terminal sternum distinctly concave, and the median lobe of the aedeagus with the apical orifice opened ventrally. These two species are the only members of the baenningeri-group that have the typical Circinatus pronotum (posterior seta distant from hind angle; hind angle completely rounded). However, based on aedeagal characters (apical orifice largely opened on the ventral side and apical lamella abruptly narrowed after the apical orifice), P. dimorphus sp. n. could also be close to P. wangjiani.
Habitat. Pterostichus dimorphus sp. n. was collected in mixed forest with dominant large pines and rhododendron trees approx 3 -5 m tall. Some individuals were found inside dead logs in day time, others were found running along tree trunks during night or were taken by pitfall traps. Diagnosis. Pronotum with single mid-lateral seta; posterior seta located almost at hind angle; hind angle forming an indistinct obtuse angle; elytron with linear microsculpture; males with two small tubercles on terminal sternum; fifth tarsomeres glabrous beneath.
There are five Circinatus species (P. maitreya, P. baenningeri, P. yan, P. ailaoicus and P. miao) similar in the following external and genital characters: (1) male terminal sternum with two tubercles; (2) pronotal posterior seta very close to hind angle; (3) aedeagus with median lobe slender, apical lamella located on the right side of apex, narrower than half width of apical orifice apex; (4) elytral microsculpture linear, sometimes hardly visible. These five species are considered to be closely allied with each other. The differences among them are listed in Table 1. These five species have relatively similar median lobe of aedeagus, but significant differences are expected to be found in the endophallus. We studied only P. baenningeri and P. maitreya sp. n. Most endophallic lobes of these two species can be recognized as homologous, but their shapes and positions are all different (Figs 40, 41).
Description. Body form slightly elongate, relatively large species within subgenus, body length 13.0-13.8 mm; dorsal side almost black, moderately shining, elytron with iridescent shine; mouthparts, antenna, tarsus, tibia, and apex of femur reddish brown; ventral side blackish. Both sexes with elytral microsculpture weak and linear. Head. Frons without punctures; antenna reaching elytron basal fifth; gena shorter than length of eye, briefly tumid behind eye. Pronotum nearly round, lateral margin strongly curved, widest a little before middle; posterior margin a little narrower than anterior margin; PW/PL = 1.13-1.17; one mid-lateral seta present, located a little before greatest width; posterior seta very close to hind angle; hind angle forming an indistinct obtuse angle; basal fovea shallow, faintly defined; inner groove subparallel to median line, basal half oblique outwards; outer groove very faint, indistinct but present, close to hind angle, approx one fourth length of inner one, outer area of inner groove flat; basal foveal area with fine punctures on inner side of inner groove. Elytron oviform, with basal ridge slightly curved; elytral shoulder moderately widened, shoulder angle between basal ridge and lateral margin forming obtuse angle, humeral tooth very small, not pointed; intervals feebly convex; striae moderately deep, without punctures; scutellar stria short, apex free; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 16-17 pores (6, 1-2, 9-10). Ventral side. Proepisternum and mesepisternum impunctate, metepisternum finely punctate; male terminal sternum with two small tubercles, located a little before middle of sternum, region between tubercles slightly depressed. (Fig. 120). Legs. Fifth tarsomeres glabrous beneath; males with apical half of mesotibia widened, inner margin crenulate; first two metatarsomeres with distinct carina on outer surface, such carina on basal half of third metatarsomere superficial. Male genitalia. Median lobe of male genitalia bent approx 90 degrees, apex abruptly bent ventrally (Fig. 41A); ventral margin straight in middle, strongly bent ventrally near apex; dorsal margin gradually curved; apical orifice large, slightly turned to left side, opened on ventral side; in lateral view, apical lamella short, laminate with base slightly thickened, not twisted, its length approx one eighth length of apical orifice; in dorsal view, apical lamella short and rounded, located on right side of median lobe apex, pointing apical-ventrally, its base distinctly grooved on dorsal surface; length of apical lamella approx equal to its basal width (Fig. 41B). Right paramere straight and stout, subtriangular, apical half slightly enlarged, apex strongly narrowed and slightly hooked, length approx three times greatest width, apex pointed (Fig. 41C). Endophallus (Fig. 41D, E, F) bent to ventral side across left side of aedeagus, major parts of endophallus located on ventral side of aedeagus (in lateral view); gonopore (gp, gonopore lobe folded in Fig. 41) located at level a little before apical lamella, pointing to aedeagal base. Six distinct lobes recognized: basal lobe (bl) small and compressed, close to base of apical lamella, lower surface strongly chitinized; dorsal lobe (dl) small and rounded, close to apex of apical lamella, membranous without decoration; ventral lobe (vl) large, completely rounded, located on ventral side of endophallus, decorated with very fine scales; pre-apical lobe (pa) close to lf, evenly rounded; right lobe (rl) small and helicon-shaped, located on base of vl, decorated with fine scales; left lobe (lf) largest, on left side of endophallus, just before gp, elongated and slightly bent, decorated with large spines, gradually changing to fine scales from lobe apex to gp. Endophallus without any chitinized piece. Female genitalia. Spermatheca with seminal canal approx five times as long as receptaculum; receptaculum capitate ( Fig. 63), club approx half length of receptaculum; seminal canal inserted at base of common oviduct, base of seminal canal not sclerotized. Stylomere II with two ensiform setae at basal half of outer margin, and one near middle of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII ( Fig. 79B) with sparse fine setae on posterior margin; posterior margin almost straight, slightly notched in middle; posterior region chitinized; anterior region semi-chitinized, without denser pigmented spots, deeply notched in middle; middle transparent region triangular, adjacent to anterior and posterior notches in middle. Female tergum VIII ( Fig. 79A) with major portion semi-chitinized, without denser pigmentation; lateral-anterior region with two chitinized patches; anterior margin slightly notched in middle, posterior margin arcuate.
Distribution. This species is known only from Fanjingshan Mt. in Guizhou Province (Map 1). The altitude range is 1778-2300 m.
Etymology. This new species is named for Maitreya Buddha. Its type locality Fanjingshan Mountain (= Mount Fanjing) is one of the sacred Buddhist mountains of China, and is traditionally regarded as the bodhimanda of bodhisattva Maitreyabuddha (Mi Le Fo in Chinese).
Affinities. Pterostichus maitreya sp. n. is close to P. baenningeri, P. yan sp. n., P. ailaoicus sp. n. and P. miao sp. n. in their similarities of male terminal sternum and male genitalia (see diagnosis above).
Habitat. Specimens of P. maitreya sp. n. were collected from mid-high altitude mixed forest at Fanjingshan Mountain. They were found running along living tree trunks during the night. It is presumed that this species hides under bark in the day, and hunts on living tree trunks at night. Diagnosis. Pronotum usually with three mid-lateral setae; hind angle completely rounded; basal fovea slightly punctate; lateral margin of elytron narrow and deep; male terminal sternum slightly depressed in middle, faintly rugose in depression; apical lamella of aedeagus located on right side of aedeagal apex, basal width a little greater than length.

Pterostichus
P. cavazzutianus mianningensis ssp. n. is sympatric with P. camelus sp. n. The latter can be readily distinguished by its larger body size and fifth tarsomeres setose beneath.
Compared to the nominotypical subspecies, in addition to their allopatric distributions, these two subspecies differ in the following four aspects. (1) Male terminal sternum: both subspecies have terminal sternum slightly depressed in males, but in P. cavazzutianus s. str., apex of terminal sternum slightly bending downwards, and almost even in P. cavazzutianus mianningensis ssp. n.; inside the depression, P. cavazzutianus mianningensis ssp. n. has faint wrinkles, such wrinkles lacking in P. cavazzutianus s. str.
(2) Median lobe of aedeagus: P. cavazzutianus mianningensis ssp. n. has the apical lamella wider, length / basal width approx 0.8; this ratio in P. cavazzutianus s. str. is approx 1.0. (3) Endophallus with four major lobes slightly different: vb-I with its apex pointed in P. cavazzutianus s. str., and completely rounded in P. cavazzutianus mianningensis ssp. n.; vb-II more chitinized and less capitate in P. cavazzutianus mianningensis ssp. n.; va with its piece much wider and shorter in P. cavazzutianus mianningensis ssp. n.; rl less distinct in P. cavazzutianus s. str.; dorsal surface of endophallus slightly angulate in P. cavazzutianus s. str., but evenly curved in P. cavazzutianus mianningensis ssp. n. (4) female sternum VIII: the transparent region small, approx semicircular in P. cavazzutianus s. str., but V-shaped with vague extensions in P. cavazzutianus mianningensis subsp. n.
Description. Body length 10.4-11.6 mm; dorsal side almost black, moderately shining; elytron with faint iridescent shine; mouthparts, antenna, tarsus, and apex of tibia yellowish brown; ventral side brownish. Both sexes with faint linear elytral microsculpture. Head. Frons without punctures; antenna reaching elytron basal sixth; gena approx same length as eye, briefly tumid behind eye. Pronotum round, widest before middle, PW/ PL = 1.10-1.17; usually three (occasional four) mid-lateral setae present, first one close to anterior angle, last one near middle of lateral margin, a little distant from rest ones; posterior seta distant from hind angle, distance between seta and hind angle approx same as distance between hind angle and inner basal foveal groove; hind angle completely rounded; basal fovea shallow, faintly defined; inner groove subparallel to median line, slightly curved outwards; outer groove completely vanished, outer area of inner groove flat; basal foveal area usually sparsely punctate on inner side of inner groove. Elytron oviform, basal ridge not oblique; elytral shoulder moderately narrowed, basal ridge and lateral margin forming obtuse angle, humeral tooth very small; intervals feebly convex; striae moderately deep, with fine punctures in basal half; scutellar stria short, complete or not; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 15-16 pores (6, 1-2, 8-9). Ventral side. Proepisternum impunctate or slightly punctate near anterior margin; mesepisternum densely punctate; metepisternum sparsely punctate; male terminal sternum slightly depressed, faintly rugose in depression, depression occupying posterior two thirds length of terminal sternum, apex of terminal sternum nearly flat (Fig. 115). Legs. Fifth tarsomeres glabrous beneath; males with apical half of mesotibia not widened, inner margin slightly crenulate; first metatarsomere with distinct carina on outer surface, such carina on second metatarsomere superficial. Male genitalia. Median lobe of male genitalia bent approx 90 degrees, apex not bent ventrally (Fig. 36A); ventral margin straight before apex, dorsal margin completely curved; apical orifice large, slightly turned to left side, not opened on ventral side; apical lamella short, approx one fourth length of apical orifice, laminate, apex not thickened; in dorsal view, apical lamella distinctly inclined right, nearly triangular with rounded apex, its length approx 0.8 times basal width (Fig. 36B). Right paramere straight and stout, inner margin slightly expanded near middle, length approx 3.5 times greatest width, apex rounded (Fig. 36C). Endophallus (Fig. 36D, E, F) short, bent to ventral side across apical lamella, and then turned to aedeagal base; gonopore (gp) located at approx same level as apical lamella, pointing to aedeagal base; gonopore lobe (gpl) bent to right side of aedeagus. Four distinct lobes recognized: ventral-basal lobe I (vb-I) small, close to base of apical lamella, compressed, its upper surface heavily scaled; ventral-basal lobe II (vb-II) large and long, close to apical lamella, pointing to ventral face of median lobe, apex a little capitate, upper surface strongly chitinized, lower surface membranous, apex scaled; ventral-apical lobe (va) at right side of vb-II, upper surface strongly chitinized, forming a transverse sinuate piece, lower surface membranous; right lobe (rl) small and membranous, on right surface of endophallus, weakly pointed. Female genitalia. Spermatheca with seminal canal approx three times as long as receptaculum; receptaculum capitate (Fig. 58), club approx half length of receptaculum; seminal canal inserted at base of common oviduct, base of seminal canal sclerotized. Stylomere II with one or two ensiform setae at outer margin, and one at basal third of inner margin; two very short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 73B) nearly evenly chitinized; posterior margin nearly straight, with fine setae, notched in middle; middle transparent region V-shaped, its anterior margin vaguely defined, adjacent to anterior and posterior notches in middle. Female tergum VIII (Fig. 73A) semi-chitinized, posterior region without denser pigmentation, anterior margin weakly notched in middle, posterior margin evenly arcuate.

Distribution. This species is known only from a single locality in Yele Reserve in Liangshan Yi Autonomous Prefecture, Sichuan province (Map 2).
Etymology. The scientific name comes from the type locality, Mianning County. Affinities. This new subspecies is proposed for the small but stable differences from the nominotypical subspecies in the male terminal sternum, the median lobe of the aedeagus and the endophallus (see diagnosis above). They were not treated as two distinct species because of their allopatric distributions, identical external appearances, and only very small differences in genital characters. The type localities of these two subspecies are approximately 100 km apart and are at almost the same altitude.
Habitat. Pterostichus cavazzutianus mianningensis subsp. n. was collected in mixed forest with dominant giant pines, and rich in dead logs. Most specimens were collected by pitfall trap, and others were found under or in dead logs. In Yele Reserve, P. cavazzutianus mianningensis subsp. n. was found together with two larger-sized Circinatus species, P. camelus sp. n. and P. zhygealu sp. n. But P. cavazzutianus mianningensis subsp. n. is much more common than the other two. Diagnosis. Pronotum with single mid-lateral seta; posterior seta very close to hind angle; hind angle nearly rounded, lateral margin slightly sinuate before hind angle; elytral microsculpture linear and faint; elytral shoulder angle completely rounded; males with two small tubercles on terminal sternum; fifth tarsomeres glabrous beneath. For comparisons between similar species see table 1 under P. maitreya.
Description. Body form fairly elongate, body length 12.5 mm; dorsal side blackish, moderately shining, elytron with distinct iridescent shine; mouthparts, antenna, tarsus, tibia, and apex of femur reddish brown; ventral side blackish. Elytral microsculpture very faint, linear. Head. Frons without punctures; antenna reaching elytron basal fifth; gena shorter than length of eye, briefly tumid behind eye. Pronotum a little elongate, lateral margin curved in middle, slightly sinuate before hind angle, widest at approx anterior third; posterior margin a little narrower than anterior margin; PW/PL = 1.12; one mid-lateral seta present, a little before greatest width; posterior seta close to hind angle; hind angle nearly rounded, not forming obtuse angle; basal fovea shallow, faintly defined; inner groove subparallel to median line, not reaching posterior margin; outer groove completely vanished; outer area of inner groove flat; basal foveal area finely punctate along inner groove. Elytron oviform, with basal ridge almost straight; elytral shoulder moderately widened, shoulder angle between basal ridge and lateral margin completely rounded, not forming obtuse angle, without humeral tooth; intervals feebly convex; striae moderately deep, without punctures; scutellar stria short, apex free; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 15 pores (6, 1, 8). Ventral side. Proepisternum and mesepisternum finely punctate near anterior margin, metepisternum heavily punctate; male terminal sternum with two very small tubercles, tubercles indistinctly defined, at approx anterior third of sternum, region between tubercles shallowly depressed (Fig. 121). Legs. Fifth tarsomeres glabrous beneath; males with apical half of mesotibia slightly widened, inner margin crenulate; first two metatarsomeres with distinct carina on outer surface, such carina on basal half of third metatarsomere superficial. Male genitalia. Median lobe of male genitalia bent approx 90 degrees, apex gradually bent ventrally (Fig. 30A); ventral margin slightly curved in middle; dorsal margin gradually curved; apical orifice large, slightly turned to left side, opened on ventral side; in lateral view, apical lamella laminate with base slightly thickened, not sinuate or twisted, its length approx one fifth length of apical orifice; in dorsal view, apical lamella located on right side of aedeagal apex, pointing apical-ventrally, not oriented to left side; apical lamella narrow, longish oviform, slightly widened to apex, apex rounded, its base not grooved on dorsal surface; length of apical lamella approx 1.7 times its basal width (Fig. 30B). Right paramere straight and stout, nearly triangular, apical half slightly enlarged and then narrowed to apex, inner margin slightly sinuate before apex; length approx 2.5 times greatest width; apex obtuse, wider than P. baenningeri (Fig.  30C). Endophallus not studied. Female genitalia unknown.
Distribution. This species is known only from Maoershan Mt. in Guangxi Province (Map 1). The altitude range is 1650-2100 m.
Etymology. This new species is named for its type locality, Maoershan Mt. In Chinese, the mountain name means "mountain of the cat". A cat, as a common pet, is always given the nickname "Miao" in Chinese.
Affinities. Pterostichus miao sp. n. is close to P. baenningeri, P. maitreya sp. n., P. ailaoicus sp. n., and P. yan sp. n. because of the similarity of the male terminal sternum and the male genitalia. Among these five species, P. miao sp. n. is similar to P. ailaoicus sp. n. in male genitalia; both species have relatively long apical lamella, 1.7 or 2.1 times as long as its basal width (approximately the same length as the basal width in the other three species), and the apical lamella of the aedeagus without a groove on the dorsal side (with a distinct groove near the base of the dorsal side in the other three species). However, based on external characters, P. miao sp. n. appears to be unique among these five species: the elytral shoulder angle rounded (forming obtuse angle in the other species), and the male terminal sternum with two tubercles located at approximately the basal third (near the middle in the other species). Diagnosis. Pronotum with single mid-lateral seta; posterior seta distant from hind angle; pronotum strongly narrowed to base, hind angle forming obtuse angle, lateral margin slightly sinuate before hind angle; elytral microsculpture distinct, approx isodiametric; males with one elongate large tubercle on terminal sternum; fifth tarsomeres glabrous beneath.
The sp. n. differs from other species of Circinatus in having the pronotal hind angle not completely rounded, forming an obtuse angle, except for four species (P. baenningeri, P. maitreya sp. n., P. ailaoicus sp. n. and P. wangjiani sp. n.), which also have the hind angle more or less obtuse. But P. tumulus sp. n. can be easily distinguished from latter four species by: (1) pronotal posterior seta distant from hind angle (very close to hind angle in latter four species); (2) elytral microsculpture distinct, almost isodiametric (very faint and linear in latter four species except for P. wangjiani sp. n.); (3) males with one large tubercle on terminal sternum (different characters in latter four species).
In Fanjingshan Mt., P. tumulus sp. n. is sympatric with and almost the same size as P. maitreya sp. n., but they can be easily separated by characters listed above.
Description. Body form a little elongate, body length = 13.7-13.8 mm; dorsal side blackish, moderately shining, elytron without iridescent shine; mouthparts, antenna, tarsus, apex of tibia dark brown; ventral side blackish. Elytral microsculpture distinct, isodiametric or slightly transverse. Head. Frons without punctures; antenna reaching elytron basal eighth; gena shorter than length of eye, briefly tumid behind eye. Pronotum strongly narrowed to base, lateral margin curved in middle, slightly sinuate before hind angle, widest at approx anterior third; posterior margin much narrower than anterior margin; PW/PL = 1.17-1.18. One mid-lateral seta present, located at greatest width; posterior seta distant from hind angle, distance between seta and hind angle approx equal to distance between hind angle and inner basal foveal groove; hind angle distinct, forming obtuse angle; lateral margin slightly elevated near hind angle. Basal fovea moderately deep but not well defined; inner groove subparallel to median line, almost straight near posterior margin; outer groove faint but present, shallowly engraved near hind angle, approx one third length of inner one; outer area of inner groove deeply depressed; basal foveal area without punctures. Elytron oviform, with basal ridge almost straight; elytral shoulder a little widened, shoulder angle between basal ridge and lateral margin forming indistinct obtuse angle, humeral tooth small but present; intervals feebly convex; striae moderately deep, without punctures; scutellar stria long, apex free; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 16-17 pores (6, 1, 9-10). Ventral side. Proepisternum almost impunctate, very sparsely punctate near posterior margin; mesepisternum and metepisternum densely and finely punctate. Male terminal sternum with an elongate tubercle on middle, tubercle apex rounded and impunctate; tubercle gradually turned into a short ridge at approx apical two fifths of sternum; ridge not reaching apex of sternum, region after it with fine longitudinal wrinkles (Fig. 125). Legs. Fifth tarsomeres glabrous beneath; males with apical half of mesotibia slightly widened, inner margin crenulate; first metatarsomere with distinct carina on outer surface, such carina on basal half of second metatarsomere superficial. Male genitalia. Median lobe of male genitalia bent approx 90 degrees, apex abruptly bent ventrally (Fig. 32A); ventral margin straight in middle, then sinuate; dorsal margin gradually curved; apical orifice large, slightly turned to left side, opened on ventral side; in lateral view, apical lamella laminate with base slightly thickened, distinctly sinuate and turned dorsally, its length approx one sixth length of apical orifice; in dorsal view, apical lamella on right side of aedeagal apex, pointing apically, slightly oriented to left side; apical lamella short, roundish, widened to apex, apex rounded, its base grooved on dorsal surface; length of apical lamella approx 1.2 times its basal width (Fig. 32B). Right paramere straight and narrow, nearly digitiform, apical half only slightly enlarged and then gradually narrowed to apex, apex narrowly rounded; length approx 3.7 times greatest width (Fig. 32C). Endophallus not studied. Female genitalia. Spermatheca ( Fig. 55) with seminal canal approx two times as long as receptaculum; receptaculum clavate, gradually expanded to apex; seminal canal inserted at base of common oviduct, base of seminal canal strongly sclerotized. Stylomere II with two ensiform setae near middle of outer margin, and a smaller one near middle of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 83B) short and wide, approx evenly chitinized, without denser pigmentation; posterior margin with sparse fine setae, notched in middle; anterior margin deeply notched in middle; middle transparent region wide and V-shaped, adjacent to anterior notch in middle. Female tergum VIII (Fig. 83A) with major portion well chitinized, with irregular fine spots, a narrow region along posterior margin less chitinized; anterior margin weakly notched in middle, posterior margin slightly arcuate.
Distribution. This species is known only from Fanjingshan Mt. in Guizhou Province (Map 1). The altitude range is approx 1778-1973 m.
Etymology. The scientific name comes from a Latin noun "tumulus", which means small hill, referring the male of new species with a tubercle on terminal sternum.
Affinities. In P. tumulus sp. n., the aedeagus is very similar to P. baenningeri, and the new species is assumed to be close to this and related species (listed in Table 1). These species all have the small apical lamella located on the right side of the median lobe apex and grooved on the dorsal surface. However, P. tumulus sp. n. differs from the other species because of the position of posterior pronotal seta, elytral microsculpture, male terminal sternum character, and shape of the receptaculum. These characters suggest an intermediate position for P. tumulus between P. baenningeri and some species from northern Yunnan, such as P. dimorphus sp. n.
Habitat. Pterostichus tumulus sp. n. was collected together with P. maitreya sp. n. from mid-high altitude mixed forest in Fanjingshan Mountain, but seems rarer.
All three paratypes were collected by pitfall trap in one locality (N26.08383, E102.88794, 3367 m). The holotype was collected under a stone along the way to the highest peak of Jiaozishan (3405-4223 m). It is presumed that the holotype was collected from middle-high altitude (ca. 3400-3700 m). Diagnosis. Pronotum approx quadrate, with single mid-lateral seta; posterior seta close to hind angle, distance between seta and hind angle less than half distance between hind angle and inner basal foveal groove; hind angle forming obtuse angle; elytron with distinct microsculpture, transverse or isodiametric; males with terminal sternum strongly concave in middle, concavity occupying four fifths length of terminal sternum.
The new species can be distinguished from most other Circinatus species by the position of the pronotal hind seta (very close to hind angle), except for P. baenningeri and its related four species (listed in table 1). P. wangjiani sp. n. can be distinguished from them by different elytral microsculpture, as well as by the different male terminal sternum. P. baenningeri and its related species all have two tubercles on male terminal sternum, but in P. wangjiani sp. n., the terminal sternum is strongly concave and without such tubercles.
Description. Body form relatively stout, relatively smaller species within subgenus, body length 9.5-10.5 mm; dorsal side blackish, moderately shining, elytron without iridescent shine; mouthparts, antenna, tarsus, and apex of tibia reddish brown; ventral side blackish. Both sexes with similar elytral microsculpture, isodiametric on disc, gradually turned to transverse on elytral apex and outer intervals. Head. Frons without punctures; antenna reaching elytron basal sixth; gena approx same length as eye, briefly tumid behind eye. Pronotum approx quadrate, lateral margin weakly curved, widest approx in middle; posterior margin a little narrower than anterior margin; PW/PL = 1.20-1.21; one mid-lateral seta present, a little before middle; posterior seta close to but a little distant from hind angle, distance between seta and hind angle much shorter than distance between hind angle and inner basal foveal groove; hind angle forming indistinct obtuse angle (Fig. 104); basal fovea shallow, faintly defined; inner groove subparallel to median line, basal half oblique outwards; outer groove indistinct but present, close to hind angle, approx one fourth length of inner one, outer area of inner groove slightly convex; basal foveal area almost impunctate. Elytron oviform, with basal ridge slightly curved; elytral shoulder moderately widened, shoulder angle between basal ridge and lateral margin completely rounded, humeral tooth absent; intervals feebly convex; striae moderately deep, faintly punctate before middle; scutellar stria short, apex free; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 15-16 pores (6, 1-2, 8-9). Ventral side. Proepisternum impunctate, mesepisternum and metepisternum impunctate; terminal sternum of male strongly concave in middle, concavity occupying four fifths length of terminal sternum, border of concavity carinate at anterior half, apex of terminal sternum flat. (Fig. 124). Legs. Fifth tarsomeres glabrous beneath; males with apical half of mesotibia slightly widened, inner margin slightly crenulate; first metatarsomere with distinct carina on outer surface, such carina on second metatarsomere superficial. Male genitalia. Median lobe of male genitalia bent less than 90 degrees, apex gradually bent ventrally (Fig. 43A); ventral margin evenly curved in middle, turned ventrally near apex; apical orifice large, slightly turned to left side, largely opened on ventral side; in lateral view, apical lamella very short, slightly twisted, laminate, a little thickened, its length approx one eighth length of apical orifice; in dorsal view, apical lamella short and wide, located on right side of aedeagal apex, pointing apically, length approx 0.8 times its basal width, apex rounded (Fig. 43B). Right paramere straight and stout, apical half slightly enlarged, length approx 2.5 times greatest width, apex rounded (Fig. 43C). Endophallus (Fig. 43D, E, F) bent to ventral side across left side of aedeagus, major portion of endophallus located on ventral side of aedeagus (in lateral view); gonopore (gp, gonopore lobe folded in Fig. 43) located at level much before apical lamella, pointing to right side. Six distinct lobes recognized: basal lobe (bl) small, close to apical lamella, decorated with very fine scales; right lobe (rl) small and compressed, decorated with fine spines near apex; left lobe I (lf-I) larg-est, basal half decorated with fine scales, apex coniform and well chitinized; left lobe II (lf-II) long and coniform, apex chitinized; ventral lobe (vl) long and slender, covered by lf-I, apex sharp, strongly prolonged and bent, its upper surface chitinized; pre-apical lobe (pa) just before gp, small and coniform, apex clearly chitinized. Besides those six lobes, a patch present close to gp, decorated with dense spines. Female genitalia. Spermatheca with seminal canal approx four times as long as receptaculum; receptaculum tubiform ( Fig. 66), apical half slightly clavate; seminal canal inserted at base of common oviduct, base of seminal canal sclerotized. Stylomere II with two ensiform setae at outer margin, and one near middle of inner margin; two very short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 82B) with dense spines on posterior margin; posterior margin almost straight, slightly notched in middle; posterior region chitinized, anterior region semi-chitinized, without denser pigmentation, deeply notched in middle; middle transparent region V-shaped, adjacent to posterior notch but not to anterior notch in middle. Female tergum VIII ( Fig. 82A) with major portion semi-chitinized, sparsely pigmented with irregular small spots, only lateralanterior region with two chitinized patches; anterior margin deeply notched in middle.
Distribution. This species is known only from Jiaozishan Mountain located on the border between Dongchuan and Luquan counties of north Yunnan (Map 1).
Etymology. The new species is named for Dr Wang Jian (Honghe University, Mengzi, Yunnan), a specialist in reptiles and close friend of the first author. With his company and help, the first author explored Jiaozishan Mountain and discovered this interesting new species.
Affinities. At first examination, we did not recognize this species as a member of Circinatus because of the unique pronotal shape (pronotum approximately quadrate with an obtuse hind angle). However, after careful study, it was found that this species is very similar to P. liciniformis based on the male terminal sternum and the aedeagus. Moreover, in spite of the pronotal shape, the following important characters are all consistent with Circinatus: the pronotal basal foveal grooves faintly defined, the pronotal posterior seta located before the hind angle (although only a short distance before), the third elytral interval with two setigerous pores, the metacoxa with two setae, and the right paramere short and straight. Some species of Circinatus also have a somewhat well-defined pronotal hind angle, such as in P. maitreya sp. n. Thus, this new species is included in the subgenus Circinatus.
From the position of the pronotal posterior seta and the male secondary sex characters on the terminal sternum, this new species shows an intermediate status between the other two species from northwestern Yunnan (P. liciniformis and P. dimorphus sp. n.) and the other five species (listed in Table 1) of the baenningeri-group (vide infra) from the eastern provinces. For detailed discussions, see the section on infrasubgeneric taxonomy.
Habitat. Three paratypes of P. wangjiani sp. n. were collected by pitfall trap in mixed forest with dominant Picea and Rhododendron trees approx 3-5 m tall. Description. Body form slightly elongate, relatively larger-sized species within subgenus, body length 12.8 mm; dorsal side blackish, moderately shining, elytron with distinct iridescent shine; mouthparts, antenna, tarsus, tibia, and apex of femur reddish brown; ventral side blackish. Elytral microsculpture very faint, linear. Head. Frons without punctures; antenna reaching elytron basal fifth; gena shorter than length of eye, briefly tumid behind eye. Pronotum nearly round, lateral margin strongly curved, widest a little before middle; posterior margin narrower than anterior margin; PW/PL = 1.15; one mid-lateral seta present, at approx anterior third; posterior seta very close to hind angle; hind angle nearly rounded, not forming obtuse angle; basal fovea shallow, faintly defined; inner groove subparallel to median line, basal half slightly oblique outwards; outer groove indistinct; outer area of inner groove slightly depressed; basal foveal area distinctly punctate. Elytron oviform, with basal ridge almost straight; elytral shoulder moderately widened, shoulder angle between basal ridge and lateral margin forming obtuse angle, humeral tooth very small, a little pointed; intervals feebly convex; striae moderately deep, without punctures; scutellar stria short, apex free; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 16-17 pores (6-7, 2, 8). Ventral side. Proepisternum impunctate, mesepisternum and metepisternum densely punctate; male terminal sternum with two small tubercles, tubercles indistinctly defined, a little before middle of sternum, region between tubercles slightly depressed. Legs. Fifth tarsomeres glabrous beneath; males with apical half of mesotibia widened, inner margin crenulate; first two metatarsomeres with distinct carina on outer surface, such carina on basal half of third metatarsomere superficial. Male genitalia. Median lobe of male genitalia bent approx 90 degrees, apex slightly bent ventrally (Fig. 31A); ventral margin straight in middle, turned ventrally near apex; dorsal margin gradually curved; apical orifice large, slightly turned to left side, opened on ventral side; in lateral view, apical lamella short and slightly sinuate, laminate with base slightly thickened, not twisted, its length approx one eighth length of apical orifice; in dorsal view, apical la-mella located on right side of aedeagal apex, pointing apical-ventrally, slightly oriented to left side; apical lamella nearly triangular, gradually narrowed to apex, apex rounded, its base distinctly grooved on dorsal surface; length of apical lamella approx 1.1 times its basal width (Fig. 31B). Right paramere straight and stout, nearly triangular, apical half slightly enlarged, strongly narrowed and slightly hooked to apex; length approx 2.8 times greatest width, apex pointed (Fig. 31C). Endophallus not studied. Female genitalia unknown.
Distribution. This species is known only from the holotype collected from Shennongjia in Hubei Province (Map 1). The altitude is approx 2000 m.
Etymology. This new species is named for Emperor Yan, also called Shennong, who was a mythical emperor in ancient China, and generally regarded as one of the ancestors of all Chinese nations. The place Shennongjia (a mountain in west Hubei province, the type locality of this new species) is named for Emperor Yan as well.
Affinities. Pterostichus yan sp. n. is close to P. baenningeri, P. maitreya sp. n., P. ailaoicus sp. n., and P. miao sp. n. in their similarities of male terminal sternum and male genitalia. Diagnosis. Pronotum with single mid-lateral seta; posterior seta distant from hind angle, hind angle completely rounded; basal foveal grooves completely fused, forming simple and deep basal fovea; elytron with faint transverse microsculpture; basal ridge strongly oblique; shoulder angle obtuse, humeral tooth short and obtuse; males with terminal sternum not modified; median lobe of aedeagal apex strongly bent to right side, apical lamella a little shorter than its basal width.
This new species is very similar to P. dentifer and P. subtilissimus. They are all relatively large sized within the subgenus, and have the aedeagal apex strongly bent to the right side. P. yuxiaodongi sp. n. differs from the latter two species by: (1) body form relatively stout, pronotum less narrowed to base (body form a little slenderer, pronotum strongly narrowed to base in the other two species); (2) pronotal basal foveal inner and outer grooves completely fused, forming simple deep fovea (basal fovea bifid anteriorly, inner and outer grooves not fused at anterior part in the other two species); (3) elytral humeral tooth less distinct than the other two species; (4) apical lamella much longer than the other two species.
Description. Body form somewhat elongate, body length 15.5 mm; dorsal side blackish, moderately shining, elytron without iridescent shine; mouthparts, antenna, tarsus, apex of tibia dark brown; ventral side blackish. Elytral microsculpture shallow, transverse. Head. Frons without punctures; antenna reaching elytron basal sixth; gena a little shorter than length of eye, briefly tumid behind eye. Pronotum a little narrowed to base, lateral margin fairly curved in middle, almost straight before hind angle, widest at approx anterior third; posterior margin a little narrower than anterior margin; PW/ PL = 1.17. One mid-lateral seta present, at greatest width. Posterior seta distant from hind angle, distance between seta and hind angle a little shorter than distance between hind angle and inner basal foveal groove; hind angle completely rounded, weakly extended backward; posterior margin slightly emarginate in middle. Basal foveal inner and outer grooves completely fused together, region between them deeply depressed, so that full basal foveal region forms a simple and deep depression; outer margin of depression approx four fifths as long as inner margin; region between pronotal lateral margin and basal fovea approx flat; basal foveal area without punctures, slightly rugose in depression. Elytron oviform, with basal ridge strongly oblique lateral-posteriorly; elytral shoulder strongly narrowed, shoulder angle between basal ridge and lateral margin forming indistinct obtuse angle, humeral tooth small and obtuse, but distinct, pointing to lateral-posterior side; intervals moderately convex; striae somewhat deep, without punctures; scutellar stria moderately long, apex conjunct to first stria; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 17-18 pores (6, 3, 8-9). Ventral side. Proepisternum impunctate; mesepisternum and metepisternum very sparsely punctate near anterior margin. Terminal and penultimate sterna of males not modified. Legs. Fifth tarsomeres glabrous beneath; males with apical half of mesotibia not widened, inner margin simple; first two metatarsomeres with distinct carina on outer surface, such carina absent on third metatarsomere. Male genitalia. Median lobe of male genitalia bent approx 90 degrees, apex not turned ventrally (Fig. 46A); in lateral view, ventral margin straight in middle, and then straight before apex; dorsal margin slightly curved; apical orifice large, located on dorsal side, not turned to left, not opened on ventral side; apical lamella laminate, not turn dorsally or ventrally. In dorsal view, median lobe strongly bent to right side and gradually narrowed after middle of apical orifice; apical orifice not constricted in middle; apical lamella approx quadrate, its length approx 0.8 times its basal width, pointing right-apically; apex truncate; left margin of apical lamella straight; right margin shorter and sinuate, forming an indistinct obtuse angle near apex (Fig. 46B). Right paramere straight, shorter than other species in subgenus; apical half distinctly enlarged, nearly rounded, very faintly bent to its right side; apex completely rounded; length approx 2.4 times greatest width (Fig. 46C). Endophallus (Figs 46D, E, G) without any chitinized piece; bent to dorsal-apical side of aedeagus, all parts of endophallus located on dorsal side of aedeagus; gonopore (gp, gonopore lobe folded in Fig. 46) located at level a little beyond apical lamella, pointing to ae-deagal base; dense and heavy spines present on dorsal surface of endophallus close to gp. Three major lobes recognized: basal lobe (bl) large and rounded, located at level of apical lamella, on right side of endophallus; middle lobe (ml) approx same size as bl, apex finely scaled, located adjacent to dorsal face of median lobe; pre-apical lobe (pa) small, tubiform, between ml and gp, without decoration. Female genitalia unknown.
Distribution. This species is known only from the holotype collected in Wolong, Sichuan Province (Map 2). The altitude is 2535 m.
Etymology. The new species is dedicated to our friend, Dr Yu Xiaodong (IZAS), who conducted a series of fruitful expeditions in West Sichuan, and collected a large number of interesting Pterostichus specimens, including this new species.
Affinities. Pterostichus yuxiaodongi sp. n. is closest to P. dentifer and P. subtilissimus. These three species are considered to form a species group within the subgenus (For morphology definition see the discussion under infra-subgeneric taxonomy).
Habitat. The new species was collected by pitfall trap in mixed forest. Along with a very large number of Pterostichus specimens of other species, only one specimen of P. yuxiaodongi sp. n. was collected in the same batch. So this new species is presumed to be very rare at its type locality, Wolong. Diagnosis. Relatively large-sized species within subgenus; pronotum with three or four mid-lateral setae; hind angle completely rounded; outer basal foveal groove faintly defined, approx half length of inner one, outer area of inner groove distinctly concave; elytron with distinct transverse or isodiametric microsculpture; humeral tooth indis-tinct, not pointed; males with terminal sternum unmodified, penultimate sternum weakly tumid on each side; apical lamella of aedeagus long, slightly hooked to left.

Pterostichus
This new species can be easily distinguished from all other species of the agilisgroup by its distinct larger size. Including the new species, a total of three known Circinatus species (the other two are P. agilis and P. camelus sp. n.) have their male secondary sexual characters on the penultimate sternum. P. camelus sp. n. is distinguishable from the others by its fifth tarsomeres setose, and different male penultimate sternum. P. agilis is the species most similar to P. zhygealu sp. n. in their similar male secondary sexual characters on penultimate sternum (weakly tumid on each side). These two species can be separated by: (1) P. zhygealu sp. n. is much larger than P. agilis; (2) P. zhygealu sp. n. have the outer groove of the pronotal basal foveal present, but completely vanished in P. agilis; (3) elytral humeral tooth almost invisible in P. zhygealu sp. n., but sharp and distinct in P. agilis; (4) apical lamella of aedeagus with obtuse hook in P. zhygealu sp. n., but simple in P. agilis.
Among its sympatric species, P. zhygealu sp. n. is most similar to P. adelphus sp. n. In addition to the difference in size, they also differ in elytral microsculpture: isodiametric or transverse in P. zhygealu sp. n., very faint and linear in P. adelphus sp. n.
Description. Body form stout, body length 13.5-14.5 mm; dorsal side black, shining; elytron without iridescent shine; mouthparts, antenna and tarsus dark brown; ventral side blackish. Both sexes with distinct elytral microsculpture, isodiametric on disc, gradually turned to transverse on elytral apex and outer intervals. Head. Frons without punctures; antenna reaching elytron basal fifth; gena approx same length as eye, briefly tumid behind eye. Pronotum round, widest at approx anterior third, PW/ PL = 1.22-1.28; usually four (sometimes three) mid-lateral setae present, first one close to anterior angle, last one near middle of lateral margin, a little distant from others; posterior seta distant from hind angle, distance between seta and hind angle approx same as distance between hind angle and inner basal foveal groove; hind angle completely rounded; basal fovea shallow, faintly defined; inner groove subparallel to median line, slightly curved outwards; outer groove faint, approx half length of inner one, outer area of inner groove distinctly concave; basal foveal area usually impunctate, sometimes with sparse punctures on inner side of inner groove. Elytron oviform, with basal ridge weakly oblique; elytral shoulder slightly narrowed, basal ridge and lateral margin forming obtuse angle, humeral tooth indistinct, not pointed; intervals feebly convex; striae moderately deep, without punctures; scutellar stria short, usually incomplete; third interval with two setigerous pores adjacent to second stria; umbilical pore series on ninth interval sparse in middle, composed of 15-18 pores (6, 1-3, 7-9). Ventral side. Proepisternum impunctate; mesepisternum finely punctate on anterior half; metepisternum impunctate or very sparsely punctate; penultimate sternum of males very faintly tumid on each side, tumidities close to primary setae, middle region between tumidities faintly depressed (Fig. 117); terminal sternum of males without special structure. Legs. Fifth tarsomeres without setae beneath; males with apical half of mesotibia slightly widened, inner margin slightly crenulate; first metatarsomere with distinct carina on outer surface, such carina on second metatarsomere superficial. Male genitalia. Median lobe of male genitalia bent approx 90 degrees, apex slightly bent ventrally (Fig. 38A); ventral margin straight in middle, slightly sinuate before apex; dorsal margin gradually curved; apical orifice large, slightly turned to left side, not opened on ventral side; apical lamella long, approx half length of apical orifice, laminate, not thickened; in dorsal view, apical lamella distinctly inclined to right, nearly triangular with rounded apex, apex a little enlarged, slightly hooked to left; apical lamella length approx 1.5 times basal width (Fig. 38B). Right paramere straight and fine, length approx 3.5 times greatest width, apex rounded (Fig. 38C). Endophallus (Fig. 38 D, E, F) bent to ventral side across apical lamella; gonopore (gp, gonopore lobe folded in Fig.  38) located at level of apical lamella, pointing to aedeagal base. Four distinct lobes recognized, the three (vb-I, vb-II, va) located on ventral side of endophallus strongly chitinized: ventral-basal lobe I (vb-I) close to base of apical lamella, very small, covered by vb-II, upper surface strongly chitinized, lower surface membranous; ventral-basal lobe II (vb-II) larger than vb-I, upper surface strongly chitinized, lower surface membranous, apex of lower surface distinctly projected; ventral-apical lobe (va) close to apex of apical lamella, approx same length as vb-II, upper surface strongly chitinized, its apex strongly prolonged and sinuate, lower surface membranous; right lobe (rl) small and rounded, decorated with fine scales. Female genitalia. Spermatheca (Fig. 54) with seminal canal approx 5.5 times as long as receptaculum; receptaculum capitate, club approx half length of receptaculum; spermathecal gland slightly expanded; seminal canal inserted at base of common oviduct, base of seminal canal sclerotized. Stylomere II with two ensiform setae at outer margin, and one near middle of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 75B) with dense and fine spines on posterior margin; posterior margin rounded, shallowly notched in middle; posterior region well chitinized, anterior region also, middle transparent region V-shaped, almost adjacent to posterior notch in middle. Female tergum VIII ( Fig. 75A) with anterior half chitinized, posterior half semi-chitinized pigmented with sparse spots, without transparent region in middle.
Distribution. Most specimens of this species were collected from the type locality (Yizi pass) on the border between Meigu and Ebian County (Sichuan, Liangshan Yi Autonomous Prefecture). One female, slightly different from other specimens, was collected in Yele Reserve in Mianning County. (Map 2) Etymology. The scientific name of the new species comes from Zhygealu, an honored hero in the Yi people's legend. Yi is the local nation living in Liangshan Yi Autonomous Prefecture (Sichuan Province), where all known species of the agilis-group are distributed. This species deserves a hero's name as the largest-sized of all known agilis-group species.
Affinities. This new species is closest to P. agilis. Despite the difference in size, these two species are very similar in the following characters: (1) elytral microsculpture transverse or isodiametric; (2) male secondary sexual characters on the penultimate sternum; and (3) apical lamella of the aedeagus long and bent to the right. The type locality of P. agilis is Luojishan (Sichuan), a mountain approximately 150 km southwest of the Yizi Pass, which is the type locality of P. zhygealu sp. n.
Habitat. Pterostichus zhygealu sp. n. was collected in mixed forest with dominant large pines, and rich in dead logs. In daytime, they hide in dead logs, usually under bark. In nighttime, they come out, and can be trapped by pitfalls. P. zhygealu sp. n. lives together with three other Circinatus species. Among them, P. zhygealu sp. n. and P. adelphus sp. n. are rarer than the other two species (P. bullatus and P. cavazzutianus).
Variation. One female collected from Mianning County is slightly different from the holotype and the paratypes in the following characters: pronotum narrower, strongly narrowed to base, and body size small with a body length of 12.7 mm. Because of these morphological differences and a different locality (approximately 100 km NW of the type locality), we suspected that it might represent a different subspecies and excluded it from the type series. Diagnosis. Pronotum with three to five mid-lateral setae; hind angle completely rounded; outer basal foveal groove vanished, outer area of inner groove completely flat; elytron with distinct isodiametric microsculpture; elytral humeral tooth distinct, short and sharp, pointed backwards; fifth tarsomeres glabrous beneath; males with terminal sternum unmodified, penultimate sternum slightly tumid on each side; apical lamella of aedeagus long, slightly bent to left.

Pterostichus (Circinatus) agilis Allegro & Sciaky, 2010
Based on elytral microsculpture and male sternum, P. agilis is most similar to P. camelus sp. n. and P. zhygealu sp. n. For comparisons between them see the diagnosis of P. zhygealu sp. n.
Supplemental description. Male sternum. Penultimate sternum of males faintly tumid on each side, tumidities close to primary setae, middle region between tumidities slightly depressed (Fig. 116); terminal sternum of males without special structures. Male genitalia. Median lobe of male genitalia bent approx 90 degrees, apex gradually and slightly bent ventrally (Fig. 37A); ventral margin straight in middle, faintly sinuate before apex; dorsal margin gradually curved; apical orifice large, slightly turned to left side, not opened on ventral side; apical lamella long, approx half length of apical orifice, laminate, not thickened; in dorsal view, apical lamella somewhat inclined right, approx triangular with rounded apex, apex a little prolonged, slightly bent to left, not hooked; apical lamella length approx 1.5 times its basal width (Fig. 37B). Right paramere straight and slender, length approx three times greatest width, apex rounded (Fig. 37C). Endophallus (Fig. 37D, E, F) bent to ventral side across apical lamella; gonopore (gp) located at level a little beyond apical lamella, pointing to aedeagal base; area around left face of gp decorated with fine scales. Four distinct lobes recognized, the three (vb-I, vb-II, va) located on ventral side of endophallus strongly chitinized: ventral-basal lobe I (vb-I) close to left margin of apical lamella base, very small, upper surface decorated with heavy spines and strongly chitinized, lower surface membranous; ventral-basal lobe II (vb-II) larger than vb-I, half chitinized half membranous, apex of chitinized right surface spined and slightly enlarged, membranous left surface with elongate apex, forming a digitiform extension on left side (in dorsal view); ventral-apical lobe (va) close to apex of apical lamella, evenly chitinized, without membranous region, approx same length as vb-II, broadly triangular with long apex; right lobe (rl) small and rounded, decorated with fine scales, located on right side of endophallus, close to gp. Female genitalia. Spermatheca with seminal canal approx 3.5 times as long as receptaculum; receptaculum capitate ( Fig. 59), club approx two fifths length of receptaculum; spermathecal gland slightly expanded; seminal canal inserted at base of common oviduct, base of seminal canal sclerotized. Stylomere II with two or three ensiform setae at outer margin, and two near middle of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII ( Fig. 74B) with dense and fine spines on posterior margin; posterior margin slightly rounded, shallowly notched in middle; posterior region well chitinized, anterior region also; middle transparent region V-shaped, almost adjacent to posterior notch in middle. Female tergum VIII (Fig. 74A) evenly chitinized except for darker patches near anterior corner, without spots.
Distribution. So far, this species is known only from Luojishan Mountain in Puge County of south Sichuan (Map 2). The highest peak of Luojishan is 4359 m high. We explored this mountain in 2012. This species was found from altitudes ranging from 2549-3859 m, but most specimens were collected above 3400 m.
Affinities. Pterostichus agilis is closest to P. zhygealu sp. n. For details see the discussion under P. zhygealu sp. n. and agilis-group below.
Remarks. The holotype is teneral. Its genitalia were distorted when mounting in eupral film. The illustration of the genitalia in the original literature is inadequate, so a redescription is provided of the male genitalia herein based on mature specimens. Notes on types. According to the original description, this species was described based on 11 males, but a holotype was not designated in the original Description. In the collection of NMPC, a total of four syntypes was found, with the first one (male) bearing a pink determination label and the others bearing white labels. We designate the syntype bearing the pink label, with a serial label no. 22374, as the lectotype.
Diagnosis. Pronotum with single mid-lateral seta; posterior seta close to hind angle; hind angle forming indistinct obtuse angle; elytron with microsculpture almost invisible; males with two large tubercles on terminal sternum; fifth tarsomeres setose beneath.
This species differs from all other known Circinatus species in having the fifth tarsomeres setose beneath, except for P. camelus sp. n. These two species are so different that they can be separated by several characters, such as the different number of pronotal mid-lateral setae, pronotal hind angle shape and so on. P. maitreya sp. n., P. baenningeri, P. yan sp. n., P. ailaoicus sp. n. and P. miao sp. n. are considered close to each other. For comparisons see Table 1 under P. maitreya.
Supplemental description. Male sternum. Penultimate sternum not modified; terminal sternum with two large tubercles with indistinct borders, covering approx one third length of sternum; tubercles a little before middle of sternum, distance between tubercles a little greater than distance between setae on penultimate sternum; region between tubercles strongly depressed (Fig. 119). Endophallus (Fig. 40D, E, F) bent to ventral side across left side of aedeagus, major portion of endophallus located on ventral side of aedeagus (in lateral view); gonopore (gp, gonopore lobe folded in Fig.  40) located at level a little beyond apical lamella, pointing to aedeagal apex. Six distinct lobes recognized: basal lobe (bl) close to base of apical lamella, its main portion membranous, decorated with sparse scales, covered by a narrow chitinized piece; ventral lobe (vl) very large, bearing two small sub-lobes (vl-I, vl-II), major portion of vl membranous, sub-lobes with apex strongly scaled; vl-I with heavy and dense scales, its apex darkest in full endophallus; vl-II on right side of vl-I, wider than vl-I; left lobe (lf) located on left side of endophallus, a little larger than vl-I, apex rounded and scaled, distant from vl; right lobe (rl) small and strongly pointed, clavate with rounded apex, apex scaled; pre-apical lobe (pa) very small, on ventral face of endophallus, just before gp, without decoration. Apical patch (ap) is a scaled region near gp, not pointed, on right side of endophallus. Endophallus without any chitinized piece. Female genitalia. Spermatheca with seminal canal approx two times as long as receptaculum; receptaculum capitate (Fig. 62), club approx half length of receptaculum; seminal canal inserted at base of common oviduct, base of seminal canal not sclerotized. Stylomere II with two ensiform setae at basal half of outer margin, and a small one near middle of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 78B) with setae on posterior margin, setae on middle region heavier; posterior margin almost straight, slightly notched in middle; posterior region chitinized, anterior region semi-chitinized, without denser pigmentation, deeply notched in middle; middle transparent region arrow-shaped, adjacent to anterior and posterior notches in middle. Female tergum VIII (Fig. 78A) semi-chitinized, with sparse and very fine spots; lateral-anterior region with darker patches; anterior margin slightly notched in middle, posterior margin evenly arcuate.
Distribution. So far, this species is known only from the type locality, Jinfoshan Mt. of Chongqing, China. (Map 1) The altitude range is 1700-2185 m.

Pterostichus (Circinatus) beneshi
This species is very similar to P. zoiai. They can be easily separated by their allopatric distribution. But morphologically, these two species can be separated only with difficulty by: (1) P. beneshi with apical lamella longer, length approx 3/4 of the basal width; P. zoiai with apical lamella shorter, length approx half the basal width; (2) P. zoiai with pronotum relatively narrower as indicated by Allegro and Sciaky (2010: 18, key). These two species are very different in endophallus: P. zoiai with endophallus strongly bent dorsal-basally, the dorsal margin of endophallus closely adjacent to the apical orifice of aedeagus; dl and ll completely fused together. In P. beneshi, the endophallus fully expanded on the dorsal face of the aedeagus, the dorsal margin of the endophallus distinctly separate from the apical orifice; dl and ll well separated. (Figs 49,50,51) Supplemental description. Male sternum. Penultimate and terminal sterna without special structures. Endophallus (Fig. 50D, E, G) extended in dorsal direction of aedeagus, main portion of endophallus located on dorsal side of aedeagus; gonopore (gp) located at level near middle of aedeagus, pointing to right (gonopore lobe folded in Fig.  50). Six distinct lobes recognized: dorsal lobe (dl) large with rounded apex, bent backward, close to apical orifice of aedeagus, dorsal half decorated with heavy scales; left lobe (ll) a little larger than dl, with finer scales, also bent backward; ventral lobe I (vl-I) small, tubiform, pointing in apical direction of aedeagus; ventral lobe II (vl-II) small, tubiform, a little longer than vl-I, apex slightly bent to right in dorsal view; right lobe (rl) shorter and thicker than vl-II; pre-apical lobe (pa) located on right surface, close to gp, apex enlarged, with very fine scales; basal-dorsal region on endophallus decorated with fine and dense scales; endophallus without chitinized piece. Female genitalia. Spermatheca with seminal canal approx 3.5 times as long as receptaculum; receptaculum capitate ( Fig. 68), club approx half length of receptaculum; seminal canal inserted at base of common oviduct, base of seminal canal with sclerotized region long and thick. Stylomere II with two ensiform setae at basal half of outer margin, and a small one near middle of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 87B) with fine setae on posterior margin, setae on middle region thicker; posterior margin narrowly notched in middle; posterior region well chitinized; anterior region weakly chitinized, deeply notched in middle; middle transparent region V-shaped, adjacent to anterior and posterior notches in middle. Female tergum VIII (Fig. 87A) short, posterior region semichitinized, with sparse and irregular spots; middle of anterior region with a chitinized large patch; anterior margin notched in middle, posterior margin evenly arcuate.
Affinities. This species is very close to P. zoiai in their similarities of external appearance and male genitalia. The endophalli are also similar, and all lobes can be recognized as homologous.
Geographical variation. We studied males from three localities: Wolong (N31.03°, E103.18°), Erlangshan (N29.88°, E102.30°), and Baoxing (N30.86°, E102.77°). We found that the Erlangshan population is slightly different from the other two in male genitalia: (1) in the Erlangshan population, the apical lamella is slightly shorter and wider than that in the other two populations and is more similar to that in P. zoiai; (2) in the Erlangshan population, the endophallus with ll is strongly and abruptly narrowed to the apex, the upper margin is strongly sinuate, and the apex of ll is much narrower than that in the other two localities; and (3) in the Erlangshan population, vl-I and vl-II are much thicker than those in the other two localities (Figs 50, 51).

Pterostichus (Circinatus) bullatus Allegro & Sciaky, 2010
Chinese common name: 蟾通缘步甲 (Chán Tōng Yuán Bù Jiă) Figures 10, 39, 61 Diagnosis. Pronotum with three to four mid-lateral setae; hind angle completely rounded; elytron with faint transverse microsculpture; even intervals of elytra strongly interrupted, forming large bumps, odd intervals also interrupted but less than even ones; fifth tarsomeres glabrous beneath; males with terminal and penultimate sterna unmodified; apical lamella of aedeagus long, slightly bent to left. This species can be rapidly distinguished from all other Circinatus species by its modified elytral intervals.
Supplemental description. Male sternum. Penultimate and terminal sterna of males without special structures. Endophallus (Fig. 39D, E, F) bent to ventral side across apical lamella; gonopore (gp) located approx at level of apical lamella, pointing to aedeagal base. Four distinct lobes recognized, the three (vb-I, vb-II, va) located on ventral side of endophallus strongly chitinized: ventral-basal lobe I (vb-I) close to left margin of apical lamella base, small, upper surface decorated with heavy spines and strongly chitinized, lower surface membranous; ventral-basal lobe II (vb-II) with a large T-shaped piece, strongly chitinized, apex truncate and wide, base narrow and long; ventral-apical lobe (va) between apex of apical lamella and gp, compressed, strongly chitinized for one half, and membranous for other half; right lobe (rl) very small, present as a small hump, not chitinized or decorated, close to gp. Female genitalia. Spermatheca with seminal canal approx three times as long as receptaculum; receptaculum capitate ( Fig. 61), club approx half length of receptaculum; spermathecal gland slightly expanded; seminal canal inserted at base of common oviduct, base of seminal canal sclerotized. Stylomere II with two or three ensiform setae at outer margin, and two at basal third of inner margin; two very short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 77B) with dense and fine setae on posterior margin; posterior margin almost straight, strongly notched in middle; anterior margin deeply notched in middle; major part of sternum well chitinized, anterior third semi-chitinized; middle transparent region approx quadrate or semicircular, small, adjacent to posterior notch in middle. Female tergum VIII (Fig. 77A) with anterior region strongly chitinized, posterior half less chitinized than anterior region, posterior half pigmented with fine and irregular spots.
Distribution. So far, this species is known only from mountains between Meigu and Ebian County in south Sichuan (Map 2). Large numbers of specimens were collected from several localities in this area. The altitude range is between 2541-2923 m.
Affinities. With the unique bumps on elytral intervals, this species is one of the most special species in the subgenus. However, because of the similarities in elytral microsculpture and apical lamella of the aedeagus, this species could be close to P. agilis and P. zhygealu sp. n.
Variation. We examined two very special females (Fig. 128 Unlike the other individuals, these two specimens have elytral striae without bumps, only slightly tangled near the base in one female and completely regular in the other female. Nevertheless, these two females are in perfect accord with P. bullatus in the shape of the pronotum, elytral microsculpture and female sternum VIII characters. These two females are likely individual aberrants instead of a distinct species. For a definitive determination of the taxonomic status of this "form", male specimens are required.  Sciaky and Facchini, 2003. Diagnosis. Pronotum usually with three mid-lateral setae; hind angle completely rounded; basal fovea punctate; lateral margin of elytron narrow and deep; males with terminal sternum slightly depressed in middle, not rugose in depression; apical lamella of aedeagus located on right side of aedeagal apex, basal width approx same as length.
For comparisons between P. cavazzutianus s. str. and the subspecies P. cavazzutianus mianningensis subsp. n., see the diagnosis of P. cavazzutianus mianningensis ssp. n. This species is similar to and sympatric with P. adelphus sp. n. For their comparisons see the diagnosis of P. adelphus sp. n.
Supplemental description. Male sternum. Penultimate sternum unmodified; terminal sternum slightly depressed in middle, apex of sternum weakly but distinctly bending downwards, almost smooth inside sternum depression (Fig. 114). Endophallus (Figs 35D, E, F) short, bent to ventral side across apical lamella, and then turned to aedeagal base; gonopore (gp) located at approx same level as apical lamella. Three distinct lobes recognized: ventral-basal lobe I (vb-I) small, close to base of apical lamella, compressed, upper surface heavily scaled, apex pointed; ventral-basal lobe II (vb-II) large and long, close to apical lamella, pointing to ventral face of median lobe, upper surface strongly chitinized, lower surface membranous, apex capitate and scaled; ventral-apical lobe (va) at right side of vb-II, its main body strongly chitinized, forming a large piece with notched apex; right lobe (rl) membranous, very small and indistinct. Female genitalia. Spermatheca with seminal canal approx three times as long as receptaculum; receptaculum capitate ( Fig. 57), club approx half length of receptaculum; seminal canal inserted at base of common oviduct, base of seminal canal sclerotized. Stylomere II with two ensiform setae at outer margin, and one at basal third of inner margin; two very short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 72B) evenly chitinized; posterior margin with fine and sparse setae, almost straight, notched in middle; middle transparent region small, approx semicircular, without oblique extensions, adjacent to anterior and posterior notches in middle. Female tergum VIII (Fig. 72A) semi-chitinized, posterior region without denser pigmentation, anterior margin without notch in middle, posterior margin evenly arcuate.
Distribution. The nominotypical subspecies is only recorded from localities near the border between Meigu and Ebian County in south Sichuan (Map 2). It is sympatric with three other Circinatus species, P. bullatus, P. adelphus sp. n. and P. zhygealu sp. n.
Etymology. The name Pterostichus cavazzutii is preoccupied by a subspecies Pterostichus (Sinosteropus) barbarae cavazzutii Sciaky & Facchini, 2003. We therefore propose a replacement name Pterostichus cavazzutianus for this species, respecting the authors' original notion by changing only the suffix.
Affinities. This species seems to be close to P. adelphus sp. n. in their similarities of elytral microsculpture and short apical lamella of aedeagus. Diagnosis. Pronotum with single mid-lateral seta; posterior seta distant from hind angle, hind angle completely rounded; basal foveal grooves not completely fused, anterior part of outer groove distinctly separated from inner one, so that basal fovea is a little bifid anteriorly; elytron with faint transverse microsculpture; basal ridge strongly oblique lateral-posteriorly; shoulder angle forming obtuse angle, humeral tooth large, strongly pointed; males with terminal sternum not modified; median lobe of aedeagus strongly bent to right side, apical lamella very short, less than one fourth its basal width. P. dentifer is very similar to P. subtilissimus and P. yuxiaodongi sp. n. They are all relatively large sized species within the subgenus, and have the aedeagus strongly bent to the right side. P. dentifer differs from the latter two species in: (1) elytral humeral tooth much larger; (2) pronotal basal fovea with more punctures; (3) median lobe of aedeagus with shorter apical lamella (than P. yuxiaodongi), and relatively simpler apex (than P. subtilissimus).
Distribution. So far this species is known only from the holotype from Sichuan Prov., Dayi County, Chadiping, 1500 m. (Map 2) Affinities. This species is close to P. subtilissimus and P. yuxiaodongi sp. n. For details see discussions under P. yuxiaodongi sp. n. and subtilissimus-group in the section of infra-subgeneric taxonomy.

Pterostichus (Circinatus) liciniformis Csiki, 1930
Chinese common name: 盘胸通缘步甲 (Pán Xīong Tōng Yuán Bù Jiă) Figures 16, 42, 52, 64, 80, 102, 122 Csiki 1930: 669, replacement  Type locality. In the original literature there was no detailed locality indicated, but only "Yunnan" was mentioned. According to our knowledge of the type material of the other Pterostichini species described in the same paper (see Shi et al., 2013: 118 for a detailed discussion) and of the further specimens of P. liciniformis collected in recent years, we infer that the type locality of this species could be in northeast Dali in Yunnan Province. Notes on types. The syntype was in Fairmaire's collection, and then should have been transferred to the collection of MNHN. But despite a careful examination of the Carabidae collection in MNHN, we failed to find the type.
Distribution. This species is relatively widely distributed in northwest Yunnan. So far it has been recorded from several localities in Lijiang, Shangri-La (=Zhongdian), and Deqin (Map 1).
Affinities. As indicated by Allegro and Sciaky (2010: 7), P. liciniformis appears to be isolated in the species group. This species could be close to the other two species from north Yunnan, P. wangjiani sp. n. and P. dimorphus sp. n. Despite differences in the pronotal hind angle, elytral shoulder angle and microsculpture, these species are very similar in the depressed male terminal sternum and aedeagus. Nevertheless, P. liciniformis could represent an early branch in the baenningeri-group. A detailed discussion is provided below.
Geographical variation. We studied males from two localities, Lijiang (Yulongxueshan Mt.; N27.20°, E100.27°) and Shangri-La (= Zhongdian; pass S. of Jiulong village; N27.67°, E100.03°). The specimens from these two localities have no differences in the external or aedeagal (endophallus excluded) characters. However, for the endophallus, we found one difference: in a specimen from Lijiang, the apex of lf-II is distinctly enlarged in dorsal view; in the specimen from Shangri-La, lf-II is smaller than that from Lijiang, and the apex less enlarged in ventral view. ( Type locality. The original literature mentioned the locality "Tatsienlu", without further details. But the label of the holotype indicates that the type locality is "Kiulung" (= Jiulong), a county west of Gonggashan Mountain. Type material examined. Holotype of Pterostichus pohnerti Jedlička (NMPC), male, body length = 10.9 mm, board mounted, with left foreleg, right hind tarsomeres (apical four segments), left antenna (apical ten antennomeres) and right antenna (apical six antennomeres) missing, genitalia dissected and stored in micro vial pinned under specimen, " Tatsienlu Diagnosis. Pronotum with single mid-lateral seta; posterior seta distant from hind angle, hind angle completely rounded; outer basal foveal groove distinct, approx half length of inner one, forming a short ridge close to hind angle; basal fovea area finely punctate; elytron with sharp humeral tooth; elytral with faint linear microsculpture; fifth tarsomeres glabrous beneath; terminal sternum of male not modified.
This species is very similar to P. xilingensis. They are difficult to separate by external appearance (the only difference is the punctures in the pronotal basal fovea (see the key), but this does not always distinguish them), but they can be easily determined by their allopatric Distribution. In male and female genitalia, these two species are different. In P. pohnerti: median lobe of aedeagus stouter; apical lamella triangular, strongly narrowed to apex; endophallus extending in apical direction of aedeagus; female transparent region on sternum VIII V-shaped, with very long and narrow extensions. In P. xilingensis: median lobe of aedeagus much slenderer; apical lamella wide, slightly narrowed to apex; endophallus extending in apical-dorsal direction of aedeagus; female transparent region on sternum VIII nearly quadrate, with very short extensions.
Supplemental description. Male sternum. Penultimate and terminal sterna without special structures. Endophallus (Fig. 47D, E, G) short and straight, extending in apical direction of aedeagus, major portion of endophallus located beyond apical lamella; gonopore (gp) located at level near aedeagal apex, pointing to ventral face of aedeagus, gonopore lobe (gpl, fully everted in Fig. 47) bent in basal direction of aedeagus. Four distinct parts recognized: basal lobe (bl) close to base of apical lamella, very small, approx triangular; main piece (mp) large and galericulate, with three branches, covering gpl, apical opening of gpl pointing out from ventral notch between mp branches; apical band (ab) close to gp, also covered by mp, half chitinized, and the rest half scaled and extended to inner margin of pa; pre-apical lobe (pa) small and compressed, ear-like, with fine and dense scales. Female genitalia. Spermatheca (Fig.  56) with seminal canal approx 4.5 times as long as receptaculum; receptaculum capitate, club approx one third as long as receptaculum; seminal canal inserted at base of common oviduct, base of seminal canal with sclerotized region long. Stylomere II with two ensiform setae at basal half of outer margin, and a small one near middle of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 85B) with fine and dense setae on posterior margin; posterior margin almost straight, strongly notched in middle; anterior region semi-chitinized, deeply notched in middle; middle transparent region V-shaped, adjacent to anterior and posterior notches in middle. Female tergum VIII (Fig. 85A) short, major portion chitinized, apical fourth semi-chitinized, with very sparse spots; anterior margin weakly notched in middle, posterior margin evenly arcuate.
Distribution. Pterostichus pohnerti is recorded from four counties of central Sichuan province: Luding (Gonggashan range), Jiulong (type locality), Baoxing (south slope of Jiajinshan), and Tianquan (east slope of Erlangshan). This species is very common and the only known Circinatus species in the Gonggashan range. A large number of specimens was collected from several localities around this mountain. Outside the Gonggashan range, it is also distributed in Baoxing and Tianquan, where it overlaps the range of P. beneshi. (Maps 2, 3) Affinities. Pterostichus pohnerti could be closest to P. xilingensis based on the similarity in external characters. However, in the stout aedeagus and endophallus extending to the aedeagal apex, P. pohnerti could be allied with some species in the agilis-group. Moreover, P. pohnerti has the southernmost distribution in the pohnerti-group, which is the closest distribution to the range of the agilis-group. Based on these distributions, we infer that the systematic position of P. pohnerti could be between P. xilingensis and the agilis-group.

Pterostichus (Circinatus) subtilissimus
Pterostichus subtilissimus is very similar to P. dentifer and P. yuxiaodongi sp. n. They are all relatively large-sized species within the subgenus, and have the aedeagus strongly bent to the right side. P. subtilissimus differs from the latter two species by: (1) elytral basal ridge almost horizontal, humeral tooth pointing laterally (basal ridge strongly oblique, humeral tooth pointing lateral-posteriorly in the latter two species); (2) body form slenderer, elytron narrower than the latter two species; (3) median lobe of aedeagus strongly constricted in the middle part of apical orifice (not constricted in the latter two species).
Supplemental description. Male sternum. Penultimate and terminal sterna of males without special structures. Endophallus (Fig. 45D, E, G) completely membranous, without any chitinized piece or heavy scale; bent to dorsal side of aedeagus, all parts of endophallus located on dorsal side of aedeagus; in dorsal view, endophallus rolled counter-clockwise a full circle, gonopore (gp, gonopore lobe folded in Fig. 45) located close to left margin of apical orifice, pointing to aedeagal apex, region around gp finely pigmented with spots; two major lobes recognized: basal lobe (bl) large and rounded, located at level of apical lamella; middle lobe (ml) narrower and longer than bl, apex finely stained, located on right margin of apical orifice, just supporting constriction on right face of aedeagus (Fig. 45D). Female genitalia. Spermatheca with seminal canal approx 4.5 times as long as receptaculum; receptaculum capitate ( Fig.  67), club approx half length of receptaculum; seminal canal inserted at base of common oviduct, base of seminal canal sclerotized. Stylomere II with two ensiform setae at outer margin, and one near middle of inner margin; two very short nematiform seta located in a furrow near apex. Female sternum VIII (Fig. 84B) with fine and sparse spines on posterior margin; posterior margin slightly curved, strongly notched in middle; posterior region chitinized; anterior region semi-chitinized, deeply and widely notched in middle; middle transparent region V-shaped, adjacent to anterior and posterior notches, oblique extensions of V-shaped transparent region with sparse spots. Female tergum VIII (Fig. 84A) well chitinized, a narrow area close to posterior margin lighter, with irregular small spots; anterior margin deeply notched in middle.
Distribution. So far this species is known only from the type locality, Emei mountain in Sichuan province. The altitude range is between 2420-2800 m. (Map 2) Affinities. This species is close to P. dentifer and P. yuxiaodongi sp. n. For details see discussions under P. yuxiaodongi sp. n. and subtilissimus-group.
Supplemental description. Male sternum. Penultimate and terminal sterna without special structures. Endophallus (Fig. 48D, E, F) short, extending in dorsal-right direction of aedeagus, major portion of endophallus located on dorsal side of aedeagus; gonopore (gp, gonopore lobe folded in Fig. 48) located at level near apical lamella apex, pointing to right, with very fine scales around gp. Two distinct parts recognized: basal lobe absent; dorsal lobe (dl) large and coniform, pointing to dorsal face of aedeagus, apex strongly chitinized, with scales; ventral lobe (vl) with a strongly chitinized piece, narrow and long, pointing to ventral face of aedeagus, apical half with a membranous region on inner surface, apex with fine scales. Female genitalia. Spermatheca with seminal canal approx four times as long as receptaculum; receptaculum capitate ( Fig. 70), club approx one third length of receptaculum; seminal canal inserted at base of common oviduct, base of seminal canal with distinct sclerotized region. Stylomere II with two ensiform setae at basal half of outer margin, and a small one near middle of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 86B) with fine and dense setae on posterior margin; posterior margin slightly curved, narrowly notched in middle; posterior region well chitinized; anterior region semi-chitinized, of inner groove completely flat; basal fovea area finely punctate between two inner grooves; elytron with humeral tooth almost invisible; elytral microsculpture faint, linear; fifth tarsomeres glabrous beneath; terminal sternum of male not modified. This species is very similar to P. beneshi. Their comparison has been provided in the diagnosis under P. beneshi.
Supplemental description. Male sternum. Penultimate and terminal sterna without special structures. Endophallus (Fig. 49D, E, G) extended in dorsal-basal direction of aedeagus, major portion of endophallus located on dorsal side of aedeagus; gonopore (gp, gonopore lobe folded in Fig. 49) located at level near middle of aedeagus, pointing to right. Five distinct lobes recognized: left lobe (ll) largest, located on left face near gp, its apex branched, forming two rounded sub-lobes, upper half of ll decorated with scales, scales near apex of ll very coarse; ventral lobe I (vl-I) tubiform, pointing to dorsal side of aedeagus, apex a little enlarged; ventral lobe II (vl-II) much smaller than vl-I; right lobe (rl) longer and thicker than vl-II, tubiform, straight; pre-apical lobe (pa) located on right face, close to gp, apex enlarged and branched, with very fine scales; basal-dorsal region on endophallus heavily pigmented; endophallus without chitinized piece. Female genitalia. Spermatheca with seminal canal approx three times as long as receptaculum; receptaculum capitate ( Fig. 69), club approx half length of receptaculum; seminal canal inserted at base of common oviduct, base of seminal canal with sclerotized region short and thick. Stylomere II with two ensiform setae at basal half of outer margin, and a small one near middle of inner margin; two short nematiform setae located in a furrow near apex. Female sternum VIII (Fig. 88B) with fine setae on posterior margin, setae on middle region a little thicker; posterior margin almost straight, notched in middle; posterior region well chitinized; anterior region semi-chitinized, deeply notched in middle; middle transparent region small, roughly diamond-shaped, with two short oblique extensions; middle transparent region adjacent to anterior and posterior notches in middle. Female tergum VIII (Fig. 88A) short, posterior region semi-chitinized, without denser pigmentation; anterior region evenly chitinized; anterior margin notched in middle, posterior margin evenly arcuate.
Distribution. Known only from Emei mountain in central Sichuan province (Map 2). The label of the holotype indicates that this species is from low altitude (600-1050 m) on Emei Mt. As a result of our expedition to Emei Mt. in 2012, we found that this species is very common at relatively high altitudes (more than 2000 m) of Emei Mountain.
Affinities. This species is close to P. beneshi in similarities of external appearance and male genitalia. Their endophalli are also similar, all lobes being recognizable as homologous.
Diagnosis. Third antennomere without secondary setae; submentum with one seta on each side; pronotal hind angle completely rounded; posterior seta close to hind angle; basal foveal area finely punctate, outer groove short and superficial; third interval of elytra with two setigerous pores, all adjacent to second stria; metacoxa with three setae; fifth tarsomeres setose beneath; males with terminal and penultimate sterna unmodified. Median lobe of aedeagus carinate ventrally; apical lamella simple, pointing apical-ventrally; apical orifice opened dorsally; right paramere short, length approx 1.6 times greatest width, apical portion completely rounded.
Distribution. This species was recorded from several localities (within 100 km of each other) in Qinling mountain (Shaanxi province).
Discussion. Pterostichus schuelkei was originally placed in the subgenus Circinatus because of its typical Circinatus-like pronotum, namely, the almost rounded hind angle. However, after a careful morphological study, we found that P. schuelkei differs from the other members of Circinatus in some important characters, which leads us to move it from Circinatus to Morphohaptoderus. In the present study, we emphasize the following two characters: (1) number of setae on the metacoxa, and (2) shape of the right paramere of the aedeagus.
Pterostichus schuelkei has three setae on the metacoxa (Fig. 137), but in the true Circinatus species, only two are present. In most Pterostichus species, the metacoxa has only two setae, one close to the outer-posterior angle and the other seta close to anterior margin of the metacoxa (1 and 2 in Fig. 137). In some Pterostichus subgenera, one additional seta is present near the inner angle of the metacoxa (3 in Fig. 137), which is typically finer and shorter than the other two setae. The number of setae on the metacoxa is important in the subgeneric classification of the genus Pterostichus. In the Chinese fauna of Pterostichus (35 total subgenera recorded according to our unpublished results), only three subgenera have three setae present on the metacoxa: Cryobius, Morphohaptoderus, and Tschitscherinea. These three subgenera are closely allied to one another, and all have similar body forms and pronotal basal foveae.
The basal plan of the right paramere in the subgenus Circinatus is straight and fusiform such as in P. baenningeri (Fig. 40C), which is modified to be stouter or triangular in some species such as P. camelus sp. n. (Fig. 28C). The ratio of the right paramere length to its greatest width is between 2.5 and 4.0 in Circinatus. However, in P. schuelkei, the right paramere is much shorter (length/width = 1.6) and the apical portion is approximately round (Fig. 126D), which is consistent with the typical form in the subgenus Morphohaptoderus. In most species of Morphohaptoderus, the right parameres are almost identical, with the apical portions very short and approximately round; there are only two exceptions in the described species (P. dundai and P. muellermotzfeldi). In P. dundai, the apical portion of the right paramere is strongly elongated and bent, and in P. muellermotzfeldi, the right paramere is approximately fusiform. Moreover, this type of short and round right paramere is present only in the subgenus Morphohaptoderus.
Pterostichus schuelkei is surely the closest species to P. wenxianensis, as indicated by Allegro & Sciaky (2010). When sorting for Morphohaptoderus specimens in the collection of the IZAS, we found that two other Morphohaptoderus species could also be related to P. schuelkei and P. wenxianensis, which are P. janatai Sciaky & Wrase and P. hubeicus Facchini & Sciaky. All four of these species are found in or close to the Qinling Mountain range in central China. They share the following characters and may form a species group in subgenus Morphohaptoderus: (1) submentum with only one seta on each side; (2) third interval with two (occasionally three) setigerous pores, all adjacent to the second stria; (3) median lobe of the aedeagus carinate ventrally; and (4) apical lamella of the aedeagus simple, pointing apical-ventrally, and not hooked or twisted. The combination of these four characters is unique to the subgenus Morphohaptoderus and may support a relationship among these four species, although P. janatai and P. hubeicus differ from the other two because of a distinct pronotal hind angle.
Based on all considerations, P. schuelkei is assigned to the subgenus Morphohaptoderus in the present paper. Diagnosis. This species is very similar to P. schuelkei, but differs from the latter by: (1) hind angle forming indistinct obtuse angle (completely rounded in P. schuelkei); (2) body larger and wider than P. schuelkei; (3) ventral carina of median lobe of aedeagus stronger and longer than in P. schuelkei.
Distribution. Known only from the holotype, collected in Wenxian, Gansu, without further detailed locality.
Discussion. This species is very close to P. schuelkei. We assign it to Morphohaptoderus for the same reason as P. schuelkei. Allegro and Sciaky (2010) revised and divided the subgenus Circinatus into three species groups. However, their "group of P. schuelkei", which included two species, actually did not belong in the subgenus Circinatus, as discussed above. The rest of the eleven species were divided into two species groups based primarily on male genitalia characters, namely, the length of the apical lamella and the presence of chitinized structures in the folded endophallus.

Infra-subgeneric taxonomy
In the present study, we found that the endophallic characters have important value in the infra-subgeneric taxonomy of Circinatus. The endophalli of the 22 known species (the endophallus studied in 16 species) in the subgenus are classified into three types: Type I (Fig. 34) has the endophallus bent to the ventral side of the aedeagus across the apex of the apical lamella, with the major portion of endophallus located on the ventral side of aedeagus, and the gonopore oriented to the aedeagus base. All species of the agilis-group have endophallus type I. Type II (Fig. 40) has the apical orifice opened on the ventral face of the median lobe, with the endophallus bent to the ventral side of the aedeagus from the ventral opening (Figs 133,134), the major portion of the endophallus located on the ventral side of the aedeagus, and the gonopore typically oriented to the aedeagus apex. All species of the baenningeri-group have endophallus type II. Type III (Fig. 50) has the endophallus bent to the dorsal side of the aedeagus, with the major portion of the endophallus located on the dorsal side of the aedeagus, and the gonopore orientation variable. All species of the subtilissimus-group and the two species of the pohnerti-group (P. beneshi and P. zoiai) have endophallus type III. The other two species of the pohnerti-group (P. pohnerti and P. xilingensis) have endophallus types intermediate between types I and III; P. xilingensis is closer to type III, and P. pohnerti is closer to type I. The character evolution of the endophallus types is discussed in the section on phylogenetic analyses and is illustrated in Fig. 3.
In addition to the endophallus, the following external or genital characters are also emphasized: (1) the number of pronotal mid-lateral setae; (2) the pronotal hind angle shape and position of the posterior seta; (3) elytral microsculpture; (4) modification of the male terminal and penultimate sterna; and (5) the shape of the median lobe of the aedeagus, including the apical orifice opening and the apical lamella shape and orientation.
Three types of characters are defined in the discussion of the morphology of each species group. Definitive characters of a species group are those characters that are present in all members of the species group but that are not present in any other species group. In general, definitive characters define a species group. Exclusive characters of a species group are those characters that are present only in some members of the species group and that are not present in any other species group. Common characters of a species group are those characters that are present in all members of the species group but are also present in some members of other species groups. The characters that occur in all species of the subgenus are not considered common characters of a species group.
As a result of the analysis of the morphological characters, the following four species groups in the subgenus Circinatus are recognized. Morphological characters, in addition to their geographical distributions, support each species group. (Maps 1, 2)

1) agilis-group
Geographical distribution. This species group contains six species and one subspecies, including two previously described species, four new species and one new subspecies described in the present work. The distributions of all members of the group are in southern Sichuan. The range of the agilis-group is approximately the same as the range of the Liangshan Yi Autonomous Prefecture (Map 2). Because the carabid fauna of Liangshan has been poorly studied, the discovery of more new species is expected on future expeditions. The range of the agilis-group does not overlap with any other species group.
Endophallus. In all species of the agilis-group, the endophallus is bent to the ventral side across the apical lamella, the major portion of the endophallus is located on the ventral side of the median lobe, and the gonopore is oriented to the aedeagal base (endophallus type I). The basal plan of the endophallus in the agilis-group has four lobes (Figs 35-39): the ventral-basal lobe I (vb-I) is small, more or less compressed, with the outer face chitinized, and is close to the left side of the apical lamella base; the ventralbasal lobe II (vb-II) is large and prolonged and contains a strongly chitinized piece and a membranous region; the ventral-apical lobe (va) is more or less chitinized and is smaller than or equal to vb-II; and the right lobe (rl) is membranous, typically weak, scaled or not, and is on the right side of the endophallus and close to the gp. Additional lobes are present only in P. adelphus sp. n. (Fig. 34), with a pre-apical lobe (pa) that may derive from the rl and a ventral-basal lobe III (vb-III) that may derive from the vb-II.
Characters. Definitive characters. Pronotum with two or more mid-lateral setae; endophallus as type I. Exclusive characters. Male penultimate sternum modified; median lobe of aedeagus relatively stout, bent less than 90 degrees; apical lamella of aedeagus with apex hooked to left; inner margin of stylomere II with two ensiform setae near middle. Common characters. Pronotal hind angle completely rounded, posterior seta distant from hind angle; apical lamella of aedeagus with length more than half its basal width, gradually narrowed to apex; apical orifice of aedeagus not opened on ventral side; receptaculum capitate.
Monophyly. The monophyly of the agilis-group appears clearly and is suggested by two characters (supposedly apomorphic), the numerous pronotal mid-lateral setae and the endophallic characters (see details above). Moreover, the isolated distribution of the agilis-group is also consistent with its monophyly.
Affinities. P. pohnerti could be the species with the most affinity to the agilis-group because its endophallus is most similar to the agilis-group (Fig. 47). The endophallus of P. pohnerti is also slightly bent to the ventral side of the aedeagus across its apical lamella. However, unlike the agilis-group, P. pohnerti has the major portion of its endophallus located beyond the apical lamella, and therefore, the endophallus is not completely bent to the venter (also, see the above discussion on endophallus types). The affinity between P. pohnerti and the agilis-group is also suggested by their geographical distributions; the range of P. pohnerti is very close to the range of the agilis-group (Map 2).
Interspecific relationships. P. adelphus sp. n. and P. cavazzutianus appear to be close to one another based on their similarities in linear elytral microsculpture and unmodified male penultimate sternum. However, the unique endophallus of P. adelphus may suggest otherwise; the endophallus has two additional lobes (vb-III, pa), which makes the endophallus more complex than any other species in the agilis-group.
The other four species (P. agilis, P. zhygealu sp. n., P. camelus sp. n., and P. bullatus) are considered close because of their similarities in elytral microsculpture (transverse or isodiametric) and apical lamella of the aedeagus (longer than one-third of the apical orifice length). P. bullatus could be relatively isolated from the other three because of its unmodified male penultimate sternum and different endophallus (vb-II is much longer than the va in P. bullatus, and vb-II is approximately the same length as va in P. agilis and P. zhygealu sp. n.). A close relationship between P. zhygealu sp. n. and P. agilis is suggested by their similar endophalli and by the similar apical lamella and male penultimate sternum.
The peculiar species P. camelus sp. n. has some special characters in the subgenus: the fifth tarsomeres setose; the male penultimate sternum with two large tubercles; and the female sternum VIII with small transparent regions isolated from the major one. Although we did not study the endophallus of P. camelus sp. n., a close relationship between P. camelus sp. n. and P. agilis is suggested by their modified penultimate sterna and relatively slender body forms.

2) baenningeri-group
Geographical distribution. This species group contains nine species, including two previously described species and seven new species described in the present paper. The range of the baenningeri-group is wider than the sum of all other species group distributions, but it does not overlap with any other species group (Map 1). Species of the baenningeri-group have been recorded in five provinces: Yunnan, Guangxi, Guizhou, Chongqing, and Hubei. Yunnan has the richest fauna with four species. In the baenningeri-group, most species are strictly allopatric, except for the two sympatric species (P. maitreya sp. n. and P. tumulus sp. n.) in the Fanjingshan Mountains (Guizhou). Most species of this group have been recorded from a single locality, except P. liciniformis, which has a relatively wide distribution in northwestern Yunnan.
Endophallus. In all species of the baenningeri-group, the apical orifice opens on the ventral side of the median lobe, the endophallus is bent to the ventral side of the aedeagus from the ventral opening, the major portion of the endophallus is located on the ventral side of the aedeagus, and the gonopore orientation is variable (endophallus type II). The basal plan of the endophallus in the agilis-group has five lobes (Figs 40-44): the basal lobe (bl) is small and close to the ventral side of the apical lamella base; the ventral lobe (vl) is large and highly variable (simple, chitinized, or divided into two parts); the right lobe (rl) is small and membranous with scales; the left lobe (lf) is large and variable and has one or two parts; and the pre-apical lobe (pa), which is typically indistinct, weakly pointed or absent. The endophallus in the baenningeri-group is more variable than in the agilis-group, and the variability is primarily represented in the trend of the endophallus main body and gonopore orientation. However, concerning the homology of the endophallic lobes, the endophalli of the five species studied in the present study are generally consistent with the basal plan, with the exception of P. dimorphus sp. n. with the bl absent; P. wangjiani sp. n. with the pa chitinized on the apex; and P. maitreya sp. n. with one additional small lobe (dl).
Characters. Definitive characters. Apical orifice opened on ventral side of median lobe; endophallus of type II. Exclusive characters. Pronotal hind angle forming obtuse angle; posterior seta close to hind angle, distance between seta and hind angle much shorter than the distance between hind angle and inner basal foveal groove; elytral shoulder angle completely rounded, without humeral tooth; female elytral microsculpture granular; receptaculum clavate. Common characters. Pronotum with one mid-lateral seta; male terminal sternum modified; male penultimate sternum not modified; apical lamella of aedeagus with length equal to or greater than basal width.
Monophyly. The monophyly of baenningeri-group appears clear, as suggested by the consistent and exclusive endophallus type (for details, see above) and by the distinctly modified male terminal sternum. Outside of the baenningeri-group, only one species (P. cavazzutianus in the agilis-group) has the male terminal sternum modified, but less distinctly than all members of the baenningeri-group. Moreover, the isolated distribution of the baenningeri-group is also consistent with its monophyly.
Affinities. It is difficult to estimate the affinities of the baenningeri-group because the male genitalia in this species group are different from all other species groups and no intermediate forms are found. However, the modified male terminal sternum might indicate a relationship to P. cavazzutianus.
Interspecific relationships. From the shape of the receptaculum and the male terminal sternum character, the nine species in the baenningeri-group are divided into two subgroups.
Subgroup I contains five species (P. baenningeri, P. maitreya, P. yan, P. miao, and P. ailaoicus) that have the following characters: (1) male terminal sternum with two tubercles; (2) female spermathecal receptaculum capitate; (3) median lobe of the aedeagus slenderer than other species; and (4) elytral microsculpture linear. Although it is difficult to estimate the evolutionary polarity of each character before a comprehensive cladistic study, we suggest a monophyletic clade for this subgroup based on their accordant male sterna and aedeagi. In subgroup I, the interspecific relationships are difficult to estimate.
Subgroup II contains four species (P. liciniformis, P. dimorphus, P. wangjiani, and P. tumulus) that have the following characters: (1) male terminal sternum concave or with one elongate tubercle; (2) female spermathecal receptaculum clavate; (3) median lobe of the aedeagus relatively stout; and (4) elytral microsculpture variable. The clavate receptaculum suggests the monophyly of subgroup II because all other species in Circinatus have capitate receptaculum. The three species from northern Yunnan appear to be allied because of their similarity in the male terminal sternum (concave in the middle). However, the median lobe of the aedeagus in P. tumulus sp. n. is very similar to some species in subgroup I (Fig. 32), and therefore, subgroup II may not be a monophyletic group. Under this supposition, the capitate receptaculum and the concave male terminal sternum should be considered plesiomorphic states (comparisons with allied subgenera also suggest these states), and subgroup II may contain early branches of the baenningeri-group.

3) subtilissimus-group
Geographical distribution. This species group contains three species, including two previously described species and one new species described in the present paper. The three species in the subtilissimus-group are strictly allopatric from one another and are known from only a single locality for each. However, each of these three species is sympatric with one or two species from the pohnerti-group. Moreover, the species of the subtilissimus-group are significantly rarer and larger in body size than their sympatric species from the pohnerti-group. All species of the subtilissimus-group and the pohnerti-group are distributed in central Sichuan Province. However, the range of the subtilissimus-group is narrower than that of the pohnerti-group (Map 2).

Endophallus.
We studied the endophallus of two species (P. subtilissimus and P. yuxiaodongi sp. n.) in the subtilissimus-group. The endophallus is bent to the dorsal side of the aedeagus, the major portion is located on the dorsal side of the aedeagus (endophallus type III), and the gonopore points to the aedeagal base or apex. The endophallus decoration is simpler than in the other three species groups: without chitinized pieces, with at least two lobes (ml and bl in Figs 45, 46) present, and with lobes glabrous or with very fine scales.
Characters. Definitive characters. Median lobe of aedeagus strongly bent to right side at approximately apical third; body size largest in the subgenus, 14.4-16.5 mm in length (in the other species groups, the largest record is a 14.5 mm specimen of P. zhygealu sp. n.). Exclusive characters. Elytral basal ridge strongly oblique lateral-posteriorly. Common characters. Pronotum with one mid-lateral seta; elytral microsculpture transverse or isodiametric; male terminal and penultimate sterna not modified; endophallus type III, without chitinized pieces.
Monophyly. The three species in the subtilissimus-group are closely allied and their monophyly appears clear, suggested by their relatively large body size and the median lobe apex strongly bent to the right side. Moreover, based on the proposition of a close relationship between the subtilissimus-group and P. beneshi + P. zoiai (vide infra), the monophyly of the subtilissimus-group is also inferred from the following characters: elytral microsculpture transverse or isodiametric, pronotal outer basal foveal groove present, and area between inner and outer basal foveal grooves depressed.
Affinities. From the similarities of the endophalli, the subtilissimus-group could be closest to P. beneshi and P. zoiai in the pohnerti-group. These species are similar because the endophallus is bent to the dorsal side of the aedeagus, without chitinized piece.
Interspecific relationships. Because of the similar aedeagi (median lobe of the aedeagus relatively slender, not constricted near the apex), the strongly oblique elytral basal ridges, and the close distributions, P. dentifer and P. yuxiaodongi sp. n. could be close to one another.

4) pohnerti-group
Geographical distribution. This species group contains four species, which are all previously described and are all distributed in central Sichuan Province. The range of the pohnerti-group, although wider, overlaps the distribution of the subtilissimus-group (Map 2). Except for P. zoiai, which is known from only a single locality, the other three species are more widely distributed. P. pohnerti and P. xilingensis are strictly allopatric, as are P. beneshi and P. zoiai. The range of P. beneshi overlaps those of P. pohnerti and P. xilingensis (Map 3).
Endophallus. The endophallus is highly variable in this species group. The endophallus in P. beneshi and P. zoiai is located on the dorsal side of the aedeagus and is strongly bent to the aedeagal base (endophallus type III). The endophallus has scales, without chitinized pieces, and the gonopore points to the dorsal side of the aedeagus. In P. pohnerti, the endophallus is weakly bent to the ventral side across the apical lamella, the major portion of the endophallus is located beyond the apical lamella, and the endophallus has scales and chitinized pieces. The gonopore points to the aedeagal base. In P. xilingensis, the endophallus is located on the dorsal side of the aedeagus and extends to the right side of the aedeagus and has scales and chitinized pieces. The gonopore points to the right side of the aedeagus.
Characters. Common characters. Pronotum with one mid-lateral seta; hind angle completely rounded; posterior seta distant from hind angle; elytron with iridescent shine; elytral microsculpture linear; male terminal and penultimate sterna not modified.
Monophyly. The four species in the pohnerti-group have similar external characters and adjacent ranges, but their male genitalia are very different from one another, primarily in the endophallus (location and decoration, see above). Based on the diversity of the endophallus, we suppose that these four species form at least a paraphyletic group and may represent relatively basal clades in the subgenus. We propose this species group not for its monophyly but for its morphological accordance and similarity, and not for the purpose of systematics but for convenient taxa cognition in classification. Neither dividing the present pohnerti-group into two or three species groups nor combining all or some of the species to other species groups will provide a better infra-subgeneric system.
Affinities. For the endophallus characters, we propose the following affinities: P. pohnerti has relative affinity to the agilis-group; P. beneshi + P. zoiai have affinity to the subtilissimus-group; and the position of P. xilingensis is relatively isolated or closer to P. pohnerti.
Interspecific relationships. P. beneshi and P. zoiai are surely closely allied with one another in their accordant external and genitalia characters; they are two allopatric sister species. However, the relationship between P. pohnerti and P. xilingensis is ambiguous.

Phylogenetic analysis of Circinatus
A phylogenetic analysis was conducted to provide more support for the above taxonomic distinctions. A total of 36 characters was selected, which included ten external characters, eleven male genital characters (including male secondary sexual characters on the sterna), nine male endophallus characters, and six female genital characters. Two of the characters were multistate, whereas the others were binary. Although all characters were unordered in the phylogenetic analysis (without demonstration of character polarities), the supposed plesiomorphic states were coded "0". All 23 of the species and subspecies in Circinatus were included in the phylogenetic analysis, but some of them lacked female genitalia or male endophallus characters. Two species of the Pterostichus subgenera Gutta and Sinosteropus were selected as out-groups. The information on the matrix, character coding, and examined material of the out-groups is found in the Appendix.

Phylogeny reconstruction
The phylogeny reconstruction was performed using WIN-PAUP* Version 4.0b10 with the following parameters: optimality criterion = parsimony; all characters were unordered; starting tree(s) was obtained via stepwise addition; addition sequence: random; number of replicates = 1000; number of trees held at each step during stepwise addition = 10; branch-swapping algorithm: TBR; steepest descent option not in effect; initial 'MaxTrees' setting = 100; 'MulTrees' option in effect; topological constraints not enforced; trees unrooted; bootstrap method with heuristic search; and number of bootstrap replicates = 1000. Both the equal weighting method (EW) and the successive weighting method (SW) were used in the phylogeny reconstruction. Branches with bootstrap values greater than 50% were maintained.
The two cladograms from our heuristic analyses are shown in Fig. 1. In the EW analysis, the two most parsimonious trees were obtained with the same tree length and indices: tree length = 100; consistency index (CI) = 0.380; homoplasy index (HI) = Figure 1. Cladograms of Circinatus species relationships with bootstrap values greater than 50%. EW: cladogram obtained by the equal weighting method; SW: cladogram obtained by the successive weighting method. Different colors show the four species groups: red, baenningeri-group; green, subtilissimus-group; purple, pohnerti-group; and blue, agilis-group. 0.620; retention index (RI) = 0.675; and rescaled consistency index (RC) = 0.257. In the SW analysis, the two most parsimonious trees were obtained. For Tree I, the length and indices were as follow: tree length = 25.27; consistency index (CI) = 0.570; homoplasy index (HI) = 0.430; retention index (RI) = 0.846; and rescaled consistency index (RC) = 0.482. For Tree II, the length and indices were as follow: tree length = 24.39; consistency index (CI) = 0.591; homoplasy index (HI) = 0.409; retention index (RI) = 0.859; and rescaled consistency index (RC) = 0.507. The two most parsimonious trees were obtained for each of the EW and the SW analyses, and the topological configurations of the strict consensus of the two trees from the EW and SW methods were the same.
The cladograms generated with the EW and SW methods had no topological conflicts but did have different reliabilities. The SW tree had higher bootstrap values than the EW tree on most branches. Two monophyletic lineages were supported with relatively high reliabilities (bootstrap values >50% in EW and >80% in SW) in both the EW and SW analyses: Lineage I = baenningeri-group; Lineage II = subtilissimus-group + pohnerti-group + agilis-group. However, in the EW analysis, most species relationships in both lineages were unresolved, except for the monophyly of the subtilissimusgroup, which was supported by a relatively high bootstrap value (= 85).
In the SW analysis, most species relationships were resolved with relatively high reliabilities. Three of the four species groups were suggested as monophyletic, with the exception of the pohnerti-group. In the baenningeri-group, a monophyletic clade composed of five species was supported (P. miao + P. ailaoicus + P. baenningeri + P. yan + P. maitreya, namely, subgroup I of the baenningeri-group, see above). The other four species formed early branches with the earliest branch of P. liciniformis. The monophyly of the subtilissimus-group was well supported (= 95), with a close relationship between P. yuxiaodongi and P. dentifer. In the agilis-group, a monophyletic clade composed of four species was supported, but the relationships of the other two species were unresolved. In the pohnerti-group, the only paraphyletic species group, the relationships among these four species and those among the four species and the other monophyletic clades remained unresolved. As indicated in the previous section, the proposal for the pohnerti-group was only an expedient arrangement for these four species. In the SW analysis, neither the clade of P. pohnerti + P. xilingensis + agilis-group nor that of P. zoiai + P. beneshi + subtilissimus-group was supported by relatively high bootstrap values.
In the present paper, the species relationships of Circinatus were discussed based on evolutionary systematics and phylogenetic systematics methods in separate sections. The major results (e.g., monophyly of the main clades) are in accord for both systematics approaches. However, conflicts are also present, such as some of the species relationships in each species group. We noticed flaws in both systematics analyses in the present work, which did not tend to resolve the species relationships further.

Character evolution
Character evolution was reconstructed using WinClada and NONA software on one of the most parsimonious trees obtained with the SW analysis (tree I). The ancestral states were reconstructed considering only unambiguous transformations. The unambiguous character evolutions are shown with circles on the branches, and the apomorphies, parallelisms and reversals are shown in separate marks.