Integument glabrous; frons usually with two pair of supraorbital setae (one seta in Haptotapinus Reitter, 1886, Haptoderotapinus Jedlička, 1930 and Unitrichus Sciaky, 1997, three or more setae in Rambousekiella Knirsch, 1925 and Ethira Andrewes, 1936); mandibular scrobe without seta; antennae filiform, antennomeres 1–3 glabrous (antennomere 3 pubescent in Molops Bonelli, 1810, Rambousekiella Knirsch, 1925, and a few West Mediterranean genera), scape attached to pedicel centrically; penultimate labial palpomere usually with two median setae, without apical setae (2–5 median setae and 1–3 apical setae in some species of Cyclotrachelus Chaudoir, 1838, cfr. Bousquet 1999: 195, 1–5 apical setae in Steropanus Fairmaire, 1888); pronotum usually with one anterolateral and one posterolateral seta (two or more anterolateral setae in Parapterostichus Desbrochers des Loges, 1906 and Ethira Andrewes, 1936, posterior setae absent in Tapinopterus Schaum, 1858, Speluncarius Reitter, 1886 and Neostomis Bousquet, 1983); procoxal cavities uniperforated and closed; mesocoxal cavities conjunct (mesepimeron does not reach the middle coxal cavity); metepisternum separated from metepimeron by distinct suture, laterally not coadunate with elytral epipleuron; elytron with parascutellar pore (if present) near or inside base of the second stria and apicolateral plica (secondarily lost in several groups); protibia with outer apical angle not produced; posterior margin of mesofemur with at least two long setae; metacoxa usually with two lateral setae (a third seta at medial position in Cryobius Chaudoir, 1838, Tapinopterus Schaum, 1858, Speluncarius Reitter, 1886, Rambousekiella Knirsch, 1925 and Pseudorambousekiella Schweiger, 1967); claws smooth; median lobe of aedeagus tube-like, with basal bulb closed dorsally; parameres without setae, the left paramere mostly conchoid; gonocoxal rami absent.

The lack of synapomorphies, in either adult or larval stages, yet discovered to suggest that the tribe (about 2500 species) forms a monophyletic lineage, arrives at the conclusion that it is an evolutionary grade (Bousquet 2012).

Apophylon schillhammeri sp. n.

The following combination of morphological features distinguishes this taxon from all the other Pterostichini known to date:

1. mentum transverse, weakly emarginate;

2. pronotal basal impressions, lateral margins and basal part of midline, as well as the elytral striae both grooved and widened;

3. elytral microreticulation double, isodiametric within the striae, transverse-mesh on the intervals;

4. apicolateral plica of elytron large and visible;

5. parascutellar striola absent, i.e. striola anastomosing in full with stria 1;

6. angular base of stria 1 absent;

7. stria 7 shallow (punctiform) in anterior half;

8. parascutellar setiferous punctures present at base of stria 2;

9. no discal setiferous punctures in interval 3;

10. remnants of both stria 9 and interval 10 present along penultimate fifth of elytron;

11. meso- and metatarsomeres 1–4 with two dorsolateral grooves divergent distally;

12. metatarsomeres 1–2 (and sometimes metatarsomere 3) setose on medioventral surface, in addition to the lateroventral setae;

13. tarsomere 4 moderately emarginate along apical margin in all legs;

14. abdominal sternite 6 bisinuate and bordered along apical margin in both sexes, flat and smooth in males, impressed and rugose on apical part in females.

15. median lobe of aedeagus with dorsal ostium;

16. left paramere without transverse apophysis;

17. right paramere falcate, with styloid apex, medial part broadened and hasp situated at subbasal position;

18. laterotergite and basal gonocoxite 1 both with trichoid setae;

19. apical gonocoxite narrowed distally, with apex widely rounded, two ensiform setae, two nematiform setae, and a lots of pit pegs on dorsal and ventral surfaces.

Figures 1–3. 

Apophylon gen. n. schillhammeri sp. n. from vil. Fangxiang, Guizhou, China, holotype. 1 Habitus; 2 Head and pronotum; 3 Head, ventral part of mouthparts. Scale lines: 3 mm (Fig. 1); 1 mm (Figs 2–3).

Figures 4–7. 

Apophylon gen. n. schillhammeri sp. n. from vil. Fangxiang, Guizhou, China, holotype. 4 Apex of elytra (a preapical puncture in stria 7; b apical setiferous puncture behind end of stria 7; c fused intervals 2 and 8); 5 Left metatarsus, ventral view; 6 Right mesotarsus, dorsal view; 7 Abdominal sternites 4–6, ventral view. Scale lines: 1 mm (Figs 4–7).

Figures 8–10. 

Apophylon gen. n. schillhammeri sp. n. from vil. Fangxiang, Guizhou, China, holotype. 8–10. Aedeagus (8 left lateral view; 9 right lateral view; 10 dorsal view; a median lobe; b right paramere, internal face; c left paramere, external face; d left paramere, internal face; e right paramere, external face). Scale lines: 1 mm (Figs 4–10).

Figures 11–16. 

Apophylon gen. n. pangu sp. n. from Zhangjiajie national Forest Park, Hunan, China, holotype. 11 Habitus; 12 Head and pronotum; 13 Apex of left elytron (a apicolateral plica; b preapical puncture in stria 7; c apical setiferous puncture at end of stria 7; 1–10 numeration of intervals); 14–16. Aedeagus with attached parameres (14 left lateral view; 15 right lateral view; 16 dorsal view; a left paramere; b right paramere). Scale lines: 3 mm (Fig. 11); 1 mm (Figs 12–16).

Figures 17–21. 

Apophylon gen. n. pangu sp. n. from Zhangjiajie national Forest Park, Hunan, China, female paratype. 17 Abdominal sternites 5–6, ventral view; 18 Postabdominal sternite 7, dorsal view; 19 Gonocoxae, ventral view; 20 Left gonocoxa and left laterotergite, ventral view; 21 Left gonocoxite 2, ventral view. Scale lines: 1 mm (Figs 17–18); 0.3 mm (Figs 19–20); 0.1 mm (Fig. 21).

Figure 22. 

Map of the provinces of Guizhou and Hunan with localities (red circe type locality of Apophylon gen. n. schillhammeri sp. n.; blue circle type locality of Apophylon gen. n. pangu sp. n.).

Habitus. Size medium (ca. 12.5–13.5 mm) for Oriental Pterostichini, shape elongate, subparallel (Figs 1, 11). Tegument. Glabrous throughout (excl. antennomeres 4–11), dorsally smooth; sternal parts partly punctate, namely postgenae, proepisterna, mesosternum and mesosterna, gula of head, prosternum and proepipleura impunctate, abdomen mostly impunctate, slightly wrinkled laterally. Color. Body dark brown to black on dorsal and ventral surface, antennae, palpomeres and legs red-brown to black. Microsculpture. Present (A. pangu) or largely reduced (A. schillhammeri); striae of elytra with shiny isodiametric microreticulation, in contrast to less shiny, transverse-mesh microreticulation on elytral intervals. Head. Eyes large, laterally protruding, tempora minute; frons with two well-developed and relatively deep furrows, divergent backward, reaching anterior supraorbital punctures; two pairs of supraorbital punctures, anterior puncture just before middle of eye, posterior one at posterior fifth of eye; antennae long, filiform, with segments 1–3 and basal fifth of segment 4 glabrous, pubescent from antennomere 4, scape shorter than antennomere 3, with single dorsal seta distally, pedicel with one ventral seta, not eccentrically attached to scape, segment 3 with six apical setae, end of antennomere 8 and antennomeres 9–11 exceeding posterior margin of pronotum; labrum subrectangular, with anterior margin concave in middle and six equidistant setae, middle four shorter than lateral two; clypeus subtrapezoid, with anterior margin straight and with two punctures removed from anterior margin at distance of one and a half diameters of puncture and closer to lateral margins than to anterior margin; mandibles large, elongate, more or less pointed and hooked at apex; maxillae shorter than mandibles, stipes with two long lateroventral setae; ligula with two setae, paraglossae long, glabrous (Fig. 3); labial palpomeres elongate, fusiform, with penultimate segment bisetose; maxillary palpi considerably longer than labial palpi, with glabrous and elongate segments; labium with distinct suture between mentum and submentum, mentum transverse, weakly emarginate, sides of sinus widely divergent, epilobes narrow, median tooth large, weakly emarginate, slightly bifid in front, bordered posteriorly, labial setae long, paramedial pits indistinct; submentum with two medial setae, without lateral setae (Fig. 3). Thorax. Pronotum subcordate, wider than long, widest before middle; midline narrow anteriorly and wide posteriorly, not reaching anterior and posterior borders; anterotransverse sulcus superficial; two basal impressions, linear, wide, deep, and divergent to base; anterior border slightly narrower than basal border, concave medially, finely rimmed laterally, rim obliterated medially; sides rounded anteriorly, sinuate before posterior angles, with wide and grooved lateral margin from each side; basal border slightly convex, oblique towards angles, not rimmed; anterior angles rounded, hardly protruding; posterior angles distinct, obtuse, slightly outward protruded; anterolateral and posterolateral setiferous punctures present. Elytra elongate, not fused, widest after middle, with slightly convex disc and subapical sinuation; apicolateral plica present, large and visible from above and at sides; basal border complete, touching stria 1; shoulders completely rounded, without denticles; striae wide, deep, impunctate; stria 7 shallow along anterior half of elytron, with one or two preapical setiferous punctures (posterior one closer to apex of elytron than to suture) (Figs 4, 13); stria 9 present along penultimate fifth of elytron; parascutellar striola anastomosing with stria 1, angular base of stria 1 absent; intervals smooth, 7–8 almost fused anteriorly; interval 9 beginning from umbilicate puncture III, not reaching apex; interval 10 present at penultimate fifth of elytron; parascutellar setiferous punctures present, small, situated at base of stria 2; no discal setae in interval 3; umbilicate series mostly with 16 pores (sometimes 15, 17, or 18), indistinctly divided into humeral and apical groups by an interruption between pores VII and VIII, interruption as long as double distance between pores VI and VII, humeral group of seven pores, I-III situated before beginning of interval 8, IV-VII within stria 8, III larger than rest, apical group of nine pores, all within stria 8, pores XI and XVI larger than others. Hind wings present. Prosternum smooth, prosternal process unbordered, proepipleura broadened posteriorly. Metasternum smooth, mostly impunctate, finely punctate only laterally, metepisterna finely punctate, twice longer than wide, strongly narrowed posteriorly. Legs moderately long and slender; procoxa without seta; mesocoxa with one lateral and one medial seta; metacoxa with two lateral setae, one anterolateral and one posterolateral seta, without medial seta, with metacoxal sulcus arcuate and ended before lateral end of coxa; protrochanter with one anterior seta at distal position; mesotrochanter with one posterior seta at distal position; metatrochanter without seta (but one female paratype of A. pangu with a seta at proximal position on left metatrochanter); profemur with three long setae on posterior surface, one proximal, one medioventral and one distal seta; mesofemur with three long setae on anterior surface, two closely situated proximal setae and one medioventral seta; metafemur with two long setae on anterior surface, one proximal seta and one medioventral seta; protibia rather dilated distally in both sexes, with two long clip setae; mesotibia nearly as long as mesofemur, with ventral ctenidium well differentiated; metatibia longer than metafemur; all tarsomeres dorsally glabrous (only tarsomere 5 with a pair of subapical setae), with two rows of lateroventral setae, glabrous on medioventral surface, except for metatarsomeres 1–2 (and sometimes metatarsomere 3) which are setose also medioventrally (Fig. 5), tarsomere 4 with apical emargination from above in all legs, meso- and metatarsomeres 1–4 with two dorsolateral grooves, posteriorly divergent (Fig. 6); male protarsomeres 1–3 strongly and symmetrically expanded, biseriate squamose beneath, segment 1 and 2 nearly equal in width, segment 3 flatter and narrower than previous two; tarsal claws smooth, one seta at base of each tarsal claw. Abdomen. Sternites 3–5 each with a pair of ambulatory setae, sternites 4–6 without transverse sulci, terminal sternite 6 in both sexes bisinuate and bordered marginally, in males flat and smooth with two large terminal setiferous punctures (Fig. 7), in females apically impressed and rugose, with four terminal setiferous punctures (Fig. 17). Male genitalia: aedeagus (Figs 8–10, 14–16). Median lobe of aedeagus long, in lateral view strongly arcuate ventrally, with basal bulb large and well developed, basal orifice concave, medial part narrower than bulb, apical lamella slightly bent ventrally; median lobe dorsally elongate, slightly bent to right apically, with left margin straight to slightly concave and right margin convex, apical lamella somewhat narrowed before tip, thereupon widened, subapical orifice (ostium) on dorsal side, elongate, extended over entire dorsum; inner sac of aedeagus with or without chitinized structures, with complex membranous pattern distally. Left paramere conchoid, without transverse apophysis, with small apical denticle on ventral margin. Right paramere falcate, attenuate at apex, broadened medially, with a hasp situated at subbasal position. Female genitalia (Figs 19–21), consisting of bipartite laterotergits and gonocoxae (latter clearly separated into basal gonocoxite 1 and apical gonocoxite 2). Laterotergite subtriangular, articulated to lateral base of gonocoxite 1, with protruding apophysis on anterior margin and 10–12 trichoid setae on posterior margin medially; gonocoxite 1 relatively narrow, elongate, with several long trichoid setae on ventral surface laterodistally; gonocoxite 2 not curved, with sides narrowed distally, apex not tapering, but widely rounded, two broad ensiform setae, one dorsomedial and one dorsolateral, preapical sensory furrow on ventral surface, with a furrow peg bearing two long and thin nematiform setae, and numerous pit pegs on both surfaces.

Apophylon is a compound word, based on the Greek prefix ἀπό [apó] (away from, separate) and φύλον [fýlon] (tribe, clan), alluding to the putative systematic remoteness of this taxon concerning its relatives. It is treated as a Greek neuter.

Holotype ♂, well-preserved (metarsomeres 3 and 5 of the left hind leg were lost after taking Fig. 1), “CHINA, Guizhou, Leishan Co. SE Kali, NE Leishan Leigong Shan, E - slope 26°23.07’N 108°13.03’E” [printed, white] / “above Fangxiang vill. Nanmang riv., 900 m 16.6.2001, leg. Wang & Schillhammer (10)” [printed, white] / “HOLOTYPE Apophylon gen. nov. schillhammeri sp. nov. Guéorguiev & Sciaky des. 2015” [printed, red]. The holotype is preserved in NMW. The genitalia of the holotype are preserved in glycerine inside a plastic vial pinned under the card on which the specimen is mounted.

See Table 1.

(based on male only). Habitus (Fig. 1). Elongate, subparallel, moderately convex. Tegument. Dorsally smooth, largely impunctate, only head in frontal furrows and vertex laterally, as well as pronotum base between impressions coarsely punctate. Measurements and ratios. See Table 2. Color. Body anthracite black dorsally and ventrally, antennomeres 2–11, palpomeres and tarsomeres dark red to fire brick red. Microsculpture. Chiefly reduced (not visible); isodiametric on pronotal lateral margins and basal impressions, elytral basal border and striae, transverse-mesh on labrum and abdominal sterna in middle. Lustre. Dorsal and ventral surfaces moderately shiny, prosternal process, metasternum, and abdominal sterna shinier medially. Head (Fig. 2). Longer than wide, slightly narrower than pronotum; disc smooth; eyes as long as 1.2 times length of antennomere 3, tempora minute, ca. 1/5 of length of eye; frontal furrows wide, deep, roughly and coarsely punctate on inside and marginally; vertex with some large, scattered punctures on each side; clypeal suture partly obliterated; mandibles lightly pointed and hooked at apex. Thorax. Pronotal disc slightly convex, mostly smooth, only internal margin at each basal impression with a few coarse punctures; anterior angles rounded, hardly protruded; posterior angles obtuse, slightly protruded outside (Fig. 2). Elytra elongate, with sides narrowed towards base, widened towards apex; shoulders completely rounded; striae well-impressed, both grooved and wide; stria 7 with one preapical setiferous puncture (Fig. 4); intervals impunctate, smooth, 1–3 flat over most extent, 4–9 subconvex; intervals 2 and 8 fused apically; single apical puncture on lateral margin behind both end of stria 7 and junction of intervals 2 and 8 (Fig. 4); umbilicate series with 16 pores on left elytron and 18 pores on right elytron, humeral group of seven pores on both elytra, apical group consists of nine pores on left elytron and eleven pores on right elytron. Hind wings present. Prosternal process with posterior margin rounded. Legs moderately long and slender; metatarsomeres 1–2 setose medioventrally, in addition to lateroventral setae. Abdomen. Terminal sternite 6 distinctly sinuate and bordered on external margin, with two large marginal punctures (Fig. 7). Male genitalia (Figs 8–10). Median lobe of aedeagus long, ventrally bent at an angle of nearly 90˚, dorsally slightly curved to right apically, with apical lamella narrowed before tip, thereupon widened, subquadrate at tip, with a left-sided lobe prominent. Inner sac of aedeagus with a submedial lamella transversely situated, large and well chitinized. Left paramere conchoid, without transverse apophysis, with apical denticle on ventral margin (Figs 8–9, c–d). Right paramere falcate, with apical part protracted, medial part moderately broadened and a hasp at subbasal position (Figs 8–9, b, e).

A noun in the genitive case for the collector of the holotype of this new species, the prominent specialist of Staphylinidae Harald Schillhammer.

Harald Schillhammer (personal communication) kindly provided us detailed description for the site and time when the holotype of A. schillhammeri sp. n. was collected: “9) SE Kaili, Leishan Co., NE Leishan, E - slope of Leigong Shan, 26°26.59’N, 108°16.53’E, ca. 900 m, Nanmang river at and above Fangxiang village, ca. 15-20 m wide, variably fast flowing, slightly polluted by surrounding farmland. 16.6.2001, leg. Schillhammer & Wang; 10) as 9), gravel banks, large-sized stones (up to 50 cm diameter), with interstitial of fine sand and clay. 16.6.2001, leg. Schillhammer & Wang”.

According to this information, the correct GPS coordinates of the type locality are 26°26.59’N, 108°16.53’E, which differ from the data written on the label pinned under the specimen (see “Type material”). In reality, it was an error in the process of printing the labels (Harald Schillhammer, personal communication).

The small river of Nanmang flows into the upper part of the basin of the Qingshui River. The Qingshui enters the Yuan River, which is one of the main tributaries of the Yangtze River in Hunan. Most probably, the species is a hygrophilous upland dweller that lives adjacent to water. Administratively, the type locality is in Leishan County, Qiandongnan Miao and Dong Autonomous Prefecture, south-eastern part of Guizhou Province, China (Fig. 22).

Holotype ♂, well-preserved, no part missing, “CHINA, NW-Hunan 1993 Wulingyuan, N Dayong Zangjiajie, 30.10., 450m leg. Schillhammer (4)” [printed, white] / “HOLOTYPE Apophylon gen. nov. pangu sp. nov. Guéorguiev & Sciaky des. 2015” [printed, red]. Paratypes 3♀♀, each one supplied with a first label as that of the holotype, and with a second label: “PARATYPE Apophylon gen. nov. pangu sp. nov. Guéorguiev & Sciaky des. 2015” [printed, red]. The holotype and one paratype are preserved in NMW, while the other two paratypes are deposited in NMNHS and cRS, respectivelly. The genitalia of the holotype are preserved in Euparal on a separate piece of plastic pinned under the card on which the specimen is mounted.

See Table 1.

Habitus (Fig. 11). Elongate, subparallel, slightly convex. Tegument. Dorsally smooth, but head, pronotum and elytra with micropunctures visible at higher magnification. Measurements and ratios. See Table 2. Color. Body rufous to black dorsally and ventraly, antennomeres 2–11, palpomeres and tarsomeres lighter. Microsculpture. Isodiametric to slightly transverse-mesh on head and pronotum, distinctly transverse-mesh on elytra and abdominal sterna. Lustre. Body throughout moderately shiny. Head (Fig. 12). Longer than wide, narrower than pronotum; disc moderately convex; eyes as long as 1.1 times length of antennomere 3, tempora minute, ca. 1/4 of length of eye; frontal furrows narrow, with micropunctures; vertex without large punctures; clypeal suture present, fine; mandibles pointed and hooked at apex. Thorax. Pronotal disc slightly convex, with micropunctation throughout and several longitudinal wrinkles between basal impressions; anterior angles rounded, slightly protruded; posterior angles obtuse, slightly protruded outside (Fig. 12). Elytra with sides narrowed towards base, widened towards apex; shoulders completely rounded; striae well-impressed, wide and somewhat grooved; stria 7 with two setiferous punctures, a larger preapical and a smaller apical, as latter on very end of stria (Fig. 13); intervals smooth, with scattered micropunctation, subconvex; intervals 2 and 8 divided apically by stria 7; umbilicate series mostly with 16 pores (sometimes 15 or 17), humeral group of seven pores, apical group of nine pores. Hind wings present. Prosternal process with posterior margin nearly straight. Legs moderately long and slender; metatarsomeres 1–2, and sometimes metatarsomere 3, setose medioventrally, in addition to lateroventral setae. Abdomen. Sternite 6 in both sexes bisinuate and bordered marginally, in males flat and smooth with two large marginal setiferous punctures, in females impressed and rugose apically with four punctures (Fig. 17); postabdominal sternite 7 of female short, consisting of two chitinized plates closely connected with a tight membrane each to other, each plate proximally with well-developed ramus (Fig. 18). Male genitalia (Figs 14–16). Median lobe of aedeagus long, ventrally bent at an angle lesser than 90˚, dorsally slightly curved to right apically, with apical lamella somewhat narrowed before tip, thereupon widened and rounded at tip. Inner sac of aedeagus without chitinized structures. Left paramere conchoid, without transverse apophysis, with apical denticle on ventral margin (Figs 14–15, a). Right paramere broken, but clearly falcate, with apical part shortly attenuate, medial part strongly broadened and a hasp at subbasal position (Figs 14–15, b). Female genitalia (Figs 19–21). Laterotergite subtriangular, with round, protruding apophysis anteriorly and group of 10–12 trichoid setae posteromedially; gonocoxite 1 relatively narrow, elongate, nearly twice as long as gonocoxite 2, with four long trichoid setae on ventral surface; gonocoxite 2 rather wide, about two and a half times longer than wide, with sides narrowed distally, apex widely rounded, two broad ensiform setae, one dorsomedial and one dorsolateral, ensiform setae slightly shorther than both nematiform setae and trichoid setae of laterotergite and basal gonocoxite, preapical sensory furrow on ventral surface, with furrow peg bearing two very long and thin nematiform setae, and numerous pit pegs on both surfaces (some larger pit pegs more or less regularly arranged along dorsal edge spreading between two ensiform setae).

A noun in the nominative for the Chinese deity Pangu or Pan Gu, who is the first living being and the creator of everything in some versions of Chinese mythology.

The field notebook of Harald Schillhammer (personal communication) indicates the following data about the site and time when the type series of A. pangu sp. n. was collected: “Zhangjiajie Forest National Park, Suoxiyu Nature Reserve, Wulingyuan section (ca. 30 km N Zhangjiajie City); ca. 2 km downstream of Shuiraosimen; small branch of Jinbian Xi, ca. 1–2 m wide, slowly flowing, with riffle areas and pools; 30.X.1993; leg. Schönmann, Schillhammer & Ji; [4]”. According to Harald Schillhammer (personal communication), the altitude of the site of collecting is ca. 450 m and approximate GPS coordinates are 29°21.10’N, 110°29.06’E. This site is located near Wulingyuan Town, Wulingyuan District, Zhangjiajie Prefecture, north-western part of Hunan Province, China (Fig. 22).

The rivers of Shuiraosimen and Jin Bian Xi flow into the basin of the Lishui River (or Li River), which is one of the main tributaries of Yangtze River in Hunan. Most probably, as its congener, the species is a hygrophilous lowland dweller that lives adjacent to water.

We note that the description of A. pangu is based on teneral specimens. Hence, some observations, such as micropunctation, microsculpture, color and luster, structure of internal sac of aedeagus, may differ in fully sclerotized adults.