Vachinius holzschuhi (Casale, 1984) (type locality: East Nepal, Tashigaon 2100 m; holotype in NHMB), by original designation.

Sphodromimus can be distinguished from other subgenera in Chlaenius by the following character combinations: body large, BL 19–26 mm; body black or metallic colored, luster matt or strongly shiny, antennae, mandibles usually dark brown, elytra black, ventral side black; head finely punctate; penultimate labial palpomere with 5–7 setae, apex truncate; antennae long, antennomere 3 longer than 4; mentum tooth stout, bifid; pronotum long, with sides usually sinuate before posterior angles, posterior lateral seta situated before posterior angles, anterior lateral seta absent; proepisterna sparsely punctate and pubescent; elytral intervals flat or convex, not ribbed, densely punctate and pubescent, basal margin reaching the scutellum; hind wings reduced in all species except Chlaenius flavofemoratus Laporte, 1834 and Chlaenius peterseni (Louwerens, 1967); prosternal process unbordered at apex; metepisterna wider than long in all species except C. flavofemoratus and C. peterseni, coarsely punctate, pubescent; legs sparsely pubescent, tarsi nearly smooth dorsally, claws simple, protibiae sulcate on dorsal side; abdominal sternites finely punctate laterally; apical lamella of aedeagus denticulate on dorsal side; apical gonocoxite without ensiform setae; receptaculum very short to absent.

This subgenus is most similar to subgenera Haplochlaenius Lutshnik and Vachinius Casale, but differs in having elytra with intervals flat or slightly convex, densely punctate and pubescent, and with basal margin complete, connected with scutellum (intervals strongly ribbed in Haplochlaenius, basal margin obsolete near scutellum; intervals smooth in Vachinius; cfr. Azadbakhsh and Kirschenhofer 2019).

Subgenus is composed of 16 species distributed in the Oriental Region (China, Indonesia, Laos, Myanmar, Nepal, Philippines, Thailand, Vietnam), including the two new species described below.

China, Hunan, Guidong: Qiyun Shan (25.9010°N, 114.0068°E), altitude 1299.12 m.

Holotype. Male (IZAS) [genitalia dissected and glued on plastic film pinned under specimen], Hunan, Guidong, Qiyun Shan, 25.9010°N, 114.0068°E, 1299.12 m, 2017.XI.12–14, S.P. Yu, Y.Z. Liu leg., Institute of Zoology, IZAS/Holotype Chlaenius (Sphodromimus) caperatus sp. nov. des. by Y.Y. Qin, 2022 [red label].

Figure 1. 

A, B Chlaenius (Sphodromimus) caperatus sp. nov. (holotype, Hunan, Guidong) C, D C. (Sphodromimus) caperatus sp. nov. (female paratype, Hunan, Guidong). Scale bars: 5.0 mm.

Paratypes. Total 8 specimens: 2 ♂♂ and 3 ♀♀ (IZAS), same data as holotype; 1 ♀ (IZAS), Hunan, Guidong, Qiyun Shan, 25.9007°N, 114.01318°E, 1487.17 m, 2017.XI.12–14, S.P. Yu, Y.Z. Liu leg., Institute of Zoology, IZAS; 1 ♂ and 1 ♀ (SCAU) “Hunan, Guidong, Dongluo, Chishuixian, 1350–1450 m, 2011.XII.1, M.Y. Tian, Q. Gao, F.F. Sun leg., SCAU. All paratypes also bear the following label: Paratype. Chlaenius (Sphodromimus) caperatus sp. nov. des. by Y.Y. Qin, 2022 [red label].

Dorsum black. PW/PL = 1.18–1.21; PAW/PBW = 0.99–1.00 (Fig. 17); pronotum with anterior angles rounded, moderately projected forward; disc sparsely punctate and pubescent with deep transverse rugosities, but with a glabrous area in the middle. Elytral intervals flat, densely punctate and pubescent. Hind wings reduced. Metepisterna short, MW/ML = 1.1–1.3 (Fig. 11). Apex of apical lamella rounded (Fig. 25A–E).

This new species is most similar to Chlaenius (Sphodromimus) hunanus (Morvan, 1997) (Fig. 2A, B), sharing the large size, shape of pronotum, black elytra, and reduced hind wings, but can be distinguished from the latter by: (1) PAW/PBW = 0.99–1.00 (Fig. 17), (0.87–0.97 in C. hunanus, Fig. 18); (2) apex of lamella of median lobe rounded (apex of lamella triangular in C. hunanus, Fig. 26A–E); (3) in female genitalia, receptaculum longer (shorter in C. hunanus, Fig. 35A–C), and seminal canal shorter (longer in C. hunanus, Fig. 35A–C).

Figures 2, 3. 

2A, B Chlaenius (Sphodromimus) hunanus (Morvan, 1997) (male, Guangdong, Nanling) 3A, B C. (Sphodromimus) deuvei (Morvan, 1997) (male, Guangxi, Huaping). Scale bars: 5.0 mm.

BL = 20.3–22.0 mm, BW = 7.6–8.9 mm [BL = 21.0 mm, BW = 7.0 mm in holotype], PAW = 3.6–3.9 mm, PBW = 3.6–3.9 mm, PW = 5.0–5.5 mm, PL = 4.3–4.5 mm, MW = 1.9–2.0 mm, ML = 1.5–1.8 mm. Head, pronotum, elytra, legs, and venter black; antennae, labial and maxillary palpi, apex of mouthparts, and tarsomeres dark brown.

Head. Vertex punctate and pubescent with a glabrous and rugose area in the middle; antennae long, reaching middle of elytra; antennomere 3 ~ 1.5× longer than antennomere 4.

Pronotum cordiform, PW/PL = 1.18–1.21 (Fig. 17), widest at apical third; anterior margin slightly concave, its width equal to its basal width, PAW/PBW = 0.99–1.00; lateral margins rounded before middle, then distinctly narrowed to base, faintly sinuate before posterior angles; anterior angles rounded, moderately projected forward; posterior angles obtuse, slightly sharp at tips; disc gently convex, sparsely punctate and pubescent, with deep transverse rugosities, with a small glabrous area in the middle; median line distinct, not reaching anterior margin and base; basal foveae deep and long, punctate, pubescent.

Elytra elongate, EL/BW = 1.45–1.61; gently convex, widest near posterior third, rounded at apex in males, subtruncate in females; striae with fine punctures; parascutellar striae well developed; parascutellar pores present; intervals flat, densely punctate and pubescent; sutural angles sharp; hind wings reduced.

Venter densely punctate, pubescent, metepisterna (Fig. 11) short, MW/ML = 1.25–1.33; abdominal sternites III–VI with one setiferous puncture each side, sternite VII with one pair of setiferous punctures in males, two pairs in females; all abdominal sternites with distinct impressions laterally.

Legs long and slender; tarsi nearly smooth dorsally.

Male genitalia. Median lobe (Fig. 25B–E) long, strongly bent to ventral side; apical orifice opened dorsally, long and wide, not reaching basal bulb, slightly turned to left side; in dorsal view, apical lamella (Fig. 25A) slightly bent to left side, length nearly equal to its basal width, apex rounded; in left lateral view, apical portion distinctly bent into a hook ventrally (Fig. 25B, E), basal orifice ~ 90 ° relative to preapical shaft; left paramere large and round; right paramere helical carved; endophallus with coiled flagellum; basal part of flagellum strongly bent with an oval sclerite facing the right; apical part of flagellum distinctly bent with a triangular bursa.

Female genitalia. Spermatheca (Fig. 34A–C) with seminal canal ~ 15× as long as receptaculum; receptaculum short linear; seminal canal inserted at base of common oviduct; spermathecal gland rounded and inserted near apex of seminal canal; villous canal long, tortuously contorted, adhered to common oviduct.

(Fig. 40) This species is known only from Guidong, Hunan. Its distribution seems isolated by a mountain barrier from C. hunanus.

The new species caperatus is named for its rugosity on the vertex, pronotal disc, and abdominal sternites.

We have not examined the types of C. (S.) hunanus, but Dr. Deuve (MNHN) kindly helped us to compare types and sent us illustrations of the apical lamella of holotype. More than 60 specimens collected from Nanling, Guangdong fit Morvan’s descriptions and illustration (1997: 17, fig. 13), and we determined them as C. (S.) hunanus. Nanling is located ~ 100 km north of the type locality of C. (S.) hunanus Jiuyi Shan, Hunan (Fig. 40).

China, Hainan, Yinggeling.

Holotype. Male (IZAS) [genitalia dissected and glued on plastic film pinned under specimen], Hainan, Yinggeling, 2009.V.11, Xinlei Huang leg., Institute of Zoology, IZAS/Holotype Chlaenius (Sphodromimus) yinggelingensis sp. nov. des. by Y.Y. Qin, 2022 [red label].

Paratypes. Total 4 specimens: 1 ♂ (IZAS), same data as holotype; 2 ♀♀ (IZAS), Hainan, Jianfengling, Mingfenggu, 947 m, 2015.I.23, Deyao Zhou leg., Institute of Zoology, IZAS; 1♀ (IZAS), Hainan, Wuzhishan, 18°54'N, 109°41'E, 1000–1600 m, 2012.IV.18, PAN & LI leg. All paratypes also bear the following label: Paratype. Chlaenius (Sphodromimus) yinggelingensis sp. nov. des. by Y.Y. Qin, 2022 [red label].

Pronotum metallic coppery to green. PW/PL = 1.12–1.21; PAW/PBW = 0.84–0.91 (Fig. 21); pronotum cordate with posterior angles right angled, rounded at tips; disc sparsely punctate and pubescent with shallow, transverse rugosities. Elytral intervals distinctly convex, with a row of setae laterally and sparse setae centrally. Hind wings reduced. Metepisterna short or width nearly equal to length; MW/ML = 1.03–1.17 (Fig. 12). Distal half of femora red-brown, the rest of legs black.

This new species is similar to Chlaenius (Sphodromimus) flavofemoratus (Figs 7A–C), in having a large size, coloration of the pronotum and femora, and the absence of a spermatheca, but can be distinguished from the latter by: (1) pronotum cordate (subquadrate in C. flavofemoratus); (2) metepisterna wider than or nearly equal to long (longer than wide in C. flavofemoratus as in Fig. 14); (3) hind wings reduced (developed in C. flavofemoratus); (4) interval convex throughout (convex basally, flat apically in C. flavofemoratus).

It is also similar to Chlaenius (Sphodromimus) peterseni (Louwerens, 1967) from the Philippines in having pronotum with green metallic luster and elytral intervals slightly convex, but differs in having meso- and metafemora with yellow coloration in the middle, pronotum with lateral margins sinuate before posterior angles, hind wings reduced (femoral black, pronotum with lateral margins straight, and hind wings developed in C. peterseni, Fig. 5A, B)

Figure 4. 

A, B Chlaenius (Sphodromimus) yinggelingensis sp. nov. (holotype, Hainan, Yinggeling) C, D C. (S.) yinggelingensis sp. nov. (female paratype, Hainan, Jianfengling). Scale bars: 5.0 mm.

Figures 5, 6. 

5A, B Chlaenius (Sphodromimus) peterseni (Louwerens, 1967) (female paratype, Philippine, photos by Shi Hongliang) 6A, B C. (Sphodromimus) pilosus (Casale, 1984) (male, Yunnan, Pingbian). Scale bar: 5.0 mm.

BL = 20.9–21.6 mm, BW = 7.5–7.9 mm [BL = 21.3 mm, BW = 7.7 mm in holotype], PAW = 3.4–3.6 mm, PBW = 3.7–4.0 mm, PW = 4.8–5.3 mm, PL = 4.1–4.5 mm, MW = 1.7–1.8 mm, ML = 1.5–1.6 mm. Head, elytra, and venter black; pronotum metallic green to metallic coppery; antennae, labial and maxillary palpi, apex of mouthparts, and tarsomeres dark brown; distal half of femora red-brown, the rest of legs black.

Head. Vertex sparsely, finely punctate and pubescent; antennae long, reaching middle of elytra; antennomere 3 ~ 1.5× longer than antennomere 4.

Pronotum cordiform, PW/PL = 1.12–1.21 (Fig. 21), widest at apical third; anterior margin slightly concave, PAW/PBW = 0.84–0.91; lateral margins rounded before middle, then distinctly narrowed to base, straight before posterior angles; anterior angles rounded, moderately projected forward; posterior angles almost right angled, slightly sharp at tips; disc sparsely punctate and pubescent, with shallow transverse rugosities, without glabrous area in the middle; median line distinct, not reaching anterior margin and base; basal foveae deep and long, punctate, pubescent.

Elytra elongate, EL/BW = 1.67–1.73, gently convex, widest near posterior third, rounded at apex in males, subtruncate in females; basal margin sinuate, reaching the scutellum (but slightly obsolete on one side in a female); striae with deep punctures; parascutellar striae well developed; parascutellar pores present; intervals distinctly convex, with a row of setae laterally and sparse setae centrally; sutural angles obtuse; hind wings reduced.

Venter sparsely pubescent, punctate; metepisterna (Fig. 12) short or width nearly equal to length, MW/ML = 1.03–1.17; abdominal sternites III–VI with one setiferous puncture each side, sternite VII with one pair of setiferous punctures in males, two pairs in females; all abdominal sternites with a few impressions laterally.

Legs long and slender; tarsi nearly smooth dorsally.

Male genitalia. Median lobe (Fig. 29B–E) long, strongly bent to ventral side; apical orifice opened dorsally, long and wide, not reaching basal bulb; in dorsal view, apical lamella (Fig. 29A) wide and short, wider than long, apex subtruncate, each side distinctly widened and thickened; in left lateral view, apex with a denticulation bent to back; in right lateral view, each side convex into a denticulation, right side smaller than left side, basal orifice ~ 90° relative to preapical shaft; left paramere large and oval; right paramere helically curved; endophallus with flagellum thick and straight; basal part of flagellum with a disciform sclerite facing the left.

Female genitalia. Bursa copulatrix (Fig. 37A–C) asymmetric, base with a bifid irregular protrusion; villous canal long, tortuously contorted, adhered to common oviduct; spermatheca and spermathecal gland absent.

(Fig. 40) China (Hainan).

The new species yinggelingensis is named for the type locality Yinggeling, Hainan.

We dissected two females in C. yinggelingensis, four in C. davidi and nine in C. flavofemoratus. As a result, we could not find either spermatheca or spermathecal gland. The absence of spermatheca is uncommon in Carabidae and only occasionally found in Trechini (Deuve 1993: fig. 250). They are also absent at least in other two species of the subgenus, C. (Sphodromimus) davidi and C. (Sphodromimus) flavofemoratus (see female genitalia descriptions below).

Vietnam, Chapa, Tonkin, Coll. J. Clermont.

China – Yunnan Prov.: 4 ♂♂ (IZAS), Pingbian, Dawei Shan, 2100 m, 2010.V.23, X.D. Yang, X.Y. Zhu leg.

Dorsum black. PW/PL = 1.02–1.07; PAW/PBW = 0.98–1.00 (Fig. 20); pronotum subquadrate with anterior angles rounded, slightly projected forward; disc densely and completely rugose, punctate. Elytral intervals flat and densely punctate and pubescent. Hind wings reduced. Metepisterna short, MW/ML = 1.3–1.4 (Fig. 13). Apical lamella of median lobe truncated, right side with a large denticulation, left side with a small denticulation (Fig. 28A–E), sometimes, such denticulation absent.

Figures 7, 8. 

7A, B Chlaenius (Sphodromimus) flavofemoratus Laporte, 1834 (male, Yunnan, Menglun) C C. (Sphodromimus) flavofemoratus Laporte, 1834 (female, Yunnan, Menglun) 8 C. (Sphodromimus) enleensis Mandl, 1992 (holotype, Indo Chine) Scale bars: 5.0 mm.

BL = 19.6–21.9 mm, BW = 7.3–7.6 mm, PAW = 3.33–3.55 mm, PBW = 3.40–3.55 mm, PW = 4.35–4.55 mm, PL = 4.05–4.45 mm, MW = 1.65–1.80 mm, ML = 1.20–1.35 mm. Head, pronotum, elytra, legs, and venter black; antennae, labial and maxillary palpi, apex of mouthparts and tarsomeres dark brown.

Head. Vertex punctate and pubescent with a rugose area; antennae long, reaching middle of elytra; antennomere 3 ~ 1.5× longer than antennomere 4.

Pronotum subquadrate, PW/PL = 1.02–1.07 (Fig. 20), widest at apical third; anterior margin slightly concave, its width equal to width of basal margin, PAW/PBW = 0.98–1.00; lateral margins slightly bent before middle, then gently narrowed to base, straight before posterior angles; anterior angles rounded, slightly projected forward; posterior angles right angled, slightly rounded at tips; disc gently convex, sparsely punctate, pubescent, with a few transverse rugosities, without glabrous area in the middle; median line distinct, deep, reaching anterior margin and base; basal foveae deep, short, broad, punctate, and pubescent.

Elytra elongate, EL/BW = 1.64–1.77; gently convex near anterior third, widest near posterior third, rounded at apex in males; striae with deep punctures; parascutellar striae well developed; parascutellar pores present; intervals flat, densely punctate and pubescent; sutural angles sharp; hind wings reduced.

Venter densely punctate, pubescent, metepisterna (Fig. 13) short, MW/ML = 1.30–1.38; abdominal sternites III–VI with one setiferous puncture each side, sternite VII with one pair of setiferous punctures in males, two pairs in females; all abdominal sternites with distinct impressions laterally.

Legs long and slender; tarsi nearly smooth dorsally.

Male genitalia. Median lobe (Fig. 28B–E) stout, strongly bent to ventral side; apical orifice opening dorsally, long and wide, not reaching basal bulb; in dorsal view, apical lamella (Fig. 28A) wide and short, wider than long, apex truncated; in left lateral view, apical right side with a large denticulation towards the base and outside, basal left side only thickened and convex or convex into a small denticulation, basal orifice ~ 90 ° relative to preapical shaft; left paramere large and round; right paramere helically curved; endophallus with flagellum coiled; basal part of flagellum with a disciform sclerite facing the right; apical part of flagellum with strip of sclerite.

Female genitalia unknown.

China (Yunnan), Vietnam.

The type locality is situated in the mountains of the Black River, northern Vietnam, not far from the Chinese frontier. Our identification is based on the original description and illustration of the male genitalia of the holotype by Casale (1984: 379, figs 11, 13).

Guangdong, Xinyi, Datianding.

Total 10 specimens. China – Guangdong: Holotype female (DWC, photo), China, Guangdong, 1500 m, Xinyi: Datianding (22.16/111.15) -VIII-1997 – leg. Li/Holotypus, Chlaenius (Sphodromimus) wrasei sp. nov. det. Kirschenhofer, 2001[red label]/COLL WRASE, BERLIN; 1 ♂ and 4 ♀♀ (IZAS), Guangdong, Xinyi, Yunkai Shan, 1508.21m/22.291317°N, 111.209888°E, 2017.V.30, Y.Z. Liu, S.P. Yu leg., Inst. of Zoology; 2 ♀♀ (IZAS), Guangdong, Xinyi, Yunkai Shan, 1250.55 m/22.292692°N, 111.203833°E, 2017.V.31, Y.Z. Liu, S.P. Yu leg., Inst. of Zoology; Guangxi: 2 ♂♂ (IZAS), Guangxi Prov, Daming Shan, Tianping Protect Station, N 23.49811, E 108.43715/1230 m, 2011.V.27 N, Xinlei Huang Coll., Inst. of Zoology.

Figures 9, 10. 

9A, B Chlaenius (Sphodromimus) davidi nom. nov. (male, Guangdong, Xinyi) 10A, B C. (Sphodromimus) enleensis (male, Vietnam, Tam Dao). Scale bars: 5.0 mm.

Pronotum fully metallic purple, or greenish purple. PW/PL = 1.09–1.11; PAW/PBW = 0.83–0.93 (Fig. 23); pronotum cordate with posterior angles nearly right angled, rounded at tips; disc sparsely punctate and pubescent, with dense, shallow, transverse rugosities, without glabrous area in the middle. Elytral intervals convex basally, flat apically, densely punctate and pubescent. Hind wings reduced. Metepisterna short, MW/ML = 1.11–1.15 (Fig. 15). Legs totally black.

This species is most similar to Chlaenius (Sphodromimus) enleensis Mandl, 1992 (Figs 8, 10A, B), sharing the large size, general shape of apical lamella, metepisterna short (Figs 15, 16), and reduced hind wings, but can be distinguished from the latter by: (1) pronotum purple or greenish purple (Fig. 23) (green to coppery, but not purple in C. enleensis, Fig. 24); (2) femora black (middle of mesofemora and metafemora brown in C. enleensis); (3) elytral intervals convex basally, flat apically (interval convex throughout in C. enleensis); (4) apical lamella wider at base (narrower in C. enleensis, Figs 32A, C, D, 33A, C, D)

BL = 21.3–24.4 mm, BW = 8.0–8.7 mm. PL = 4.5–5.0 mm, PW = 5.0–5.5 mm, MW = 2.0–2.1 mm, ML = 1.7–1.9 mm. Head, elytra, venter, and legs dark and black; pronotum fully purple or greenish purple; antennae, labial and maxillary palpi, apex of mouthparts and tarsomeres dark brown.

Head. Vertex finely punctate, pubescent, without a distinct glabrous area; antennae long, reaching middle of elytra; antennomere 3 ~ 1.5× longer than antennomere 4.

Pronotum cordiform, PW/PL = 1.09–1.11 (Fig. 23), widest at apical third; anterior margin slightly concave, PAW/PBW = 0.83–0.93; lateral margins distinctly narrowed from middle to base, distinctly sinuate before posterior angles; anterior angles rounded, slightly projected forward; posterior angles nearly right angled, rounded at tips; disc gently convex, sparsely punctate and pubescent, with dense shallow transverse rugosities, without glabrous area in the middle; median line distinct, fine, not reaching anterior margin and base; basal foveae deeply arcuate, punctate and pubescent.

Elytra elongate, EL/BW = 1.61–1.75, gently convex near anterior third, widest near posterior third, rounded at apex in males, subtruncate in females; striae with deep punctures; parascutellar striae well developed; parascutellar pores present; intervals convex at base, flat from middle to apex, densely punctate and pubescent; sutural angles obtuse and right; hind wings reduced.

Venter densely punctate, pubescent, metepisterna (Fig. 15) short, MW/ML = 1.11–1.15; abdominal sternites III–VI with one setiferous puncture each side, sternite VII with one pair of setiferous punctures in males, two pairs in females; all abdominal sternites with distinct impressions laterally.

Legs long and slender; tarsi nearly smooth dorsally.

Male genitalia. Median lobe (Fig. 31B–E) long, strongly bent to ventral side; apical orifice opened dorsally, long and wide, not reaching basal bulb; in dorsal view, apical lamella triangular (Fig. 31A), distinctly bent to right side, longer than basal width, each side of middle apical lamella with a denticulation or absent on right side; in left lateral view, apical portion slightly bent dorsally at apex, basal orifice ~ 90° relative to preapical shaft; left paramere larger than right paramere, both helically curved; endophallus with flagellum thick and straight; basal part of flagellum with irregular bursa; apical part of flagellum with drop–shaped sclerite.

Female genitalia. Bursa copulatrix (Fig. 39A–C) round, base with a distinct long digitiform protrusion; villous canal long, tortuously contorted, adhered to common oviduct; spermatheca and spermathecal gland absent.

(Fig. 40) China (Guangdong; Guangxi).

Chlaenius (Sphodromimus) wrasei was originally described as a member of the genus Vachinius. Azadbakhsh and Kirschenhofer (2019) moved it to the genus Chlaenius. However, in a result of this treatment, it became a junior homonym of Chlaenius (Lithochlaenius) wrasei (Kirschenhofer, 1997). Herein, we propose a new replacement name for the former – Chlaenius (Sphodromimus) davidi nom. nov., based on the first name of the well-known specialist on the ground beetles, David W. Wrase.

This species was described based on a single female from Datianding, Xinyi, Guangdong, China. Recently, one male and six females were collected from the type locality, and those specimens fit well with the original description and illustration of C. davidi nom. nov. Two more males were collected in Guangxi, which are identical to the holotype.

Indonesia, Java.

Total 83 specimens. China – Fujian: 1 ♀ (IZAS), Fujian, Nanping, 1985.VI.7; 1 ♀ (IZAS), Fujian, Nanjing, 1991.V.15; Guangdong: 3 ♂♂ (IZAS), Guangdong, Haifeng, Jimingsi, 23.037605°N, 115.25343°E, 178.09m/2017.V.22, Y.Z. Liu, S.P. Yu leg., Inst. of Zoology; Guangxi: 1 ♂ (IZAS), Guangxi, Longsheng, Liluo, 1985. IV Jun Li leg.; 1 ♂ (IZAS), Guangxi, Guilin, Longsheng, 2003.VII.7 Jianxin Cui leg.; 1 ♀ (IZAS), Guangxi, Nanning, 1980.IV.21, Rongquan Cai leg.; 1 ♂ (IZAS), Guangxi, Jingxi, 840 m, 1998.IV.1 Chunsheng Wu leg.; Guizhou: 1 ♂ and 1 ♀ (IZAS), Guizhou, Luodian, 420 m, 1979.IV.16, Qingqiang Li leg.; 1 ♀ (IZAS), Guizhou, Luodian, 1979.V.1; Hainan: 1 ♂ (IZAS), Hainan, Baisha, Yinggeling, 2011.V.1, 600 m, light trap, Wenxin Lin leg., Inst. of Zoology; Yunnan: 4 ♂♂ and 1 ♀ (IZAS), Yunnan, Jinghong, Virgin Forest Park, Peacock Villa, 22.0304°N, 100.8763°E, 682 m/2021.VIII.6 N, along road Pingzhou Zhu leg., Inst. of Zoology; 1 ♂ (IZAS), Yunnan, Xishuangbanna, Botanical Garden, Chao Wu leg.; 1 ♀ (IZAS), Yunnan, Hekou, Qiaotou, Bajiaotian village/ 22.85348°N, 104.14211°E, 886 m, 2021.IV.21 N, Y. Xu, Z.Q. Yan coll., Inst. of Zoology; 1 ♂ (CAS), China, Yunnan Province, Tengchong, Shangying, Longwenqiao, field, beach, 25°01'19.9"N, 98°40'40.4"E/1290 m, 2003.X.20 D., H.B. Liang, X.C. Shi Coll., Institute of Zool., CAS & California Acad. Sciences; 1 ♂ and 3 ♀♀ (IZAS), Yunnan, Xishuangbanna, Menglun, Botanical Garden, 549 m, 21°56.035'N, 101°15.154'E /2005.V.25, Guo Zheng leg, Inst. of Zoology, CAS; 4 ♂♂ and 6 ♀♀ (CAS), Yunnan, Xishuangbanna, Menglun, Botanical Garden, 558 m, 21°55.035'N, 101°16.500'E /2007.V.20, Guo Zheng leg, Inst. of Zoology, CAS; 3 ♂♂ and 8 ♀♀ (IZAS), Yunnan, Xishuangbanna, Menglun, Botanical Garden, 558 m, 21°55.035'N, 101°16.500'E /2007.VII.10, Guo Zheng leg, Inst. of Zoology, CAS; 1 ♂ and 4 ♀♀ (CAS), Yunnan, Xishuangbanna, Menglun, Botanical Garden, 572 m, 21°54.646'N, 101°16.257'E /2007.I.10, Guo Zheng leg, Inst. of Zoology, CAS; 2 ♀♀ (IZAS), Yunnan, Xishuangbanna, Menglun, Botanical Garden, 2009.XII.1, Guo Tang leg, Inst. of Zoology; 2 ♀♀ (IZAS), China, Yunnan Prov., Nabanhe N.R. Guomenshan, alt.1150 m, 2009.V.6, Jiayao Hu, Ziwei Yin leg.; 1 ♂ (IZAS), Yunnan, Jinghong, Menghai, Nabanhe N.R. Guomenshan, Forest, 2009.VI.26, 1114 m/22.24644°N, 100.60610°E, pitfall, L.Z. Meng leg., Inst. of Zoology; 1 ♂ (IZAS), China, Yunnan, Hekou, Longpu, 240 m, 2011.IV.13D, 22.65404°N, 103.98193°E/Xinlei Huang leg., Inst. of Zoology, CAS; 1 ♂ (IZAS), Yunnan, Xishuangbanna, Menglun, Botanical Garden, 540 m, 21.92987°N, 101.24820°E, 2011.IV.21N/Hongbin Liang, Kaiqing Li leg., Institute of Zool., CAS; 1 ♂ (CAS), Yunnan, Xishuangbanna, Menglun, Botanical Garden, 540 m, 21.92987°N, 101.24820°E, 2011.IV.22/Yan Li leg., Institute of Zool., CAS; 1 ♂ (IZAS), Yunnan Prov. Menglun, Botanical Garden, vegetation, 21.91175°N, 101.28163°E /650 m, 2009.XI.15, Guo Tang Coll., Inst. of Zoology; 1 ♀ (IZAS), Yunnan, Xishuangbanna, Menglun, Botanical Garden, 2011.V.04, 560 m/ 21°55'39.66"N, 101°15'18.09"E, Jingxin Liu leg., Inst. of Zoology; 1 ♀ (IZAS), Yunnan, Xishuangbanna, 29 km NW, Jinghong, vic. Da Nuo You/ 22°12.41'N, 100°38.29'E, 790 m, 2009.V.16 leg. L. Meng, rice follow; 1 ♀ (IZAS), Yunnan, Xishuangbanna, 20km NW, Jinghong, vic. Man Dian (NNNR)/22°07.80'N, 100°40.05'E, 740 m, 2008.V.13 leg. A. Weigei rubb. plant; 1 ♂ and 1 ♀ (CAS), Yunnan, Lushui, Pianma, Gangfang, Xuetang 26.12218°N, 98.57546°E/1625 m,2005.V.16N, D. Kavanaugh, D.Z. Dong leg., Inst. of Zoology, CAS; 1 ♂ (CAS), China, Yunnan, Mengla, Biodiversity Corridor, 660 m, 2011.IV.25D, 21.40482°N, 101.63035°E /Xinlei Huang leg., G213 1999 km, Inst. of Zoology, CAS; 1 ♂ (IZAS), Yunnan, Yangbi, Pingpo, 1422 m, 25°35'33"N, 100°2'56"E/2002.VI.27, Min Wu leg.; 1 ♀ (CAS), China, Yunnan Province, Tengchong, Qushi Town, Xiaojiangqiao, riverside, 25°14'22.2"N, 98°37'38.0"E/1445 m, 2003.X.21, night, H.B. Liang, X.C. Shi Coll., Institute of Zool., CAS & California Acad. Science; 1 ♀ (CAS), Yunnan, Gaoligongshan, Nujiang Prefecture, 1500 m, 26°07.3'N, 98°34.5'E, 1998.X.14, D.H. Kavanaugh leg.; 2 ♂♂ (IZAS), Yunnan, Xishuangbanna, Menglun Town NO.55, 2009.V.04, Hu Li leg.; 4 ♀♀ (IZAS), Yunnan, Xishuangbanna, Menglun, Botanical Garden, 2012.V.16, L.Z. Meng leg., Inst. of Zoology; 1 ♂ (IZAS), Yunnan, Xishuangbanna, Menglun, 1982.IV.26, Linyao Wang leg.; 1 ♂ (IZAS), Yunnan, 1980.VI.5, Fen Liu leg.; 1 ♀ (IZAS), Yunnan, Gaoligong, Cikai Town, Pulahe joint of Nujiang, 27.74843°N, 98.66498°E/1530 m 2004.X.23, night, D. Kavanaugh, D.Z. Dong leg. Inst. of Zoology, CAS; 1 ♂ (IZAS), Yunnan, Lancang, Mafang By pitfall traps, 22.57925°N, 99.99849°E/1723 m, 2004.VI.16, W.B. Gu coll, Inst. of Zoology, CAS; 1 ♀ (CAS), Yunnan, Lushui, Liuku, Gaoligong Shan, 25°51'20"N, 98°50'58"E/800 m, 2002.IX.19, H.B. Liang, night, Sino-American Exped., Inst. of Zoology, CAS; 1 ♂ (CAS), China, Yunnan, Gaoligongshan, Nujiang Prefecture, Gangfang, Sancha Lukou/26 07.3'N, 98 34.5'E, 1500 m, 1998.X.12, D.H. Kavanaugh collector.; Myanmar: 1 ♂ (NHML), Carin Cheba, 900–1100 m, L. Fea/Fry Coll., 1905.100./Chlaenius flavofemoratus Cast. = nigricoxis Mots., comp with type, H.E. Andrewes det.; Vietnam: 1 ♀ (IZAS), Tonkin, Hoa Binh, leg: A. de Cooman.

Figures 11–16. 

Metepisternum features of Sphodromimus spp. 11 Chlaenius (Sphodromimus) caperatus sp. nov., holotype 12 C. (S.) yinggelingensis sp. nov., holotype 13 C. (S.) pilosus (Casale, 1984), male 14 C. (S.) flavofemoratus Laporte, 1834, male 15 C. (S.) davidi nom. nov.; male 16 C. (S.) enleensis Mandl, 1992, male.

Pronotum metallic green to metallic coppery. PW/PL = 1.14–1.26; PAW/PBW = 0.76–0.92 (Fig. 22); pronotum subquadrate with anterior angles rounded, not projected forward; disc gently convex, sparsely punctate. Elytral intervals convex basally, flat apically, densely punctate and pubescent. Hind wings developed. Metepisterna long, MW/ML = 0.75–0.92 (Fig. 14). Distal half of femora red-brown, the rest of legs black. This species is similar to Chlaenius (Haplochlaenius) costiger Chaudoir, 1856, but intervals interval convex basally and flat apically, not costulate.

Figures 17–24. 

Pronotum features of Sphodromimus spp. 17 Chlaenius (Sphodromimus) caperatus sp. nov.; holotype 18 C. (S.) hunanus (Morvan, 1997) male 19 C. (S.) deuvei (Morvan, 1997); male 20 C. (S.) pilosus (Casale, 1984), male 21 C. (S.) yinggelingensis sp. nov.; holotype 22 C. (S.) flavofemoratus Laporte, 1834, male 23 C. (S.) davidi nom. nov., male 24 C. (S.) enleensis Mandl, 1992, male. Scale bars: 1.0 mm.

BL = 22.3–25.3 mm, BW = 8.5–10.4 mm, PAW = 3.5–4.0 mm, PBW = 4.3–4.6 mm, PW = 5.5–5.8 mm, PL = 4.5–4.9 mm, MW = 1.9–2.5 mm, ML = 2.4–3.0 mm. Head, elytra, and venter black; pronotum metallic green to metallic coppery; antennae, labial and maxillary palpi, apex of mouthparts and tarsomeres dark brown; distal half of femora red-brown, the rest of legs black.

Head. Vertex finely punctate and pubescent, with a glabrous area in the middle; antennae long, reaching middle of elytra; antennomere 3 ~ 1.7× longer than antennomere 4.

Pronotum subquadrate, PW/PL = 1.14–1.26 (Fig. 22), widest at apical four-ninth; anterior margin slightly concave, PAW/PBW = 0.76–0.92; lateral margins rounded or straight before posterior angles; anterior angles rounded, not projected forward; posterior angles obtuse; disc gently convex, sparsely punctate, without glabrous area in the middle; median line distinct, fine, reaching anterior margin and base; basal foveae deep, punctate and pubescent.

Figures 25–27. 

Aedeagus of Sphodromimus spp. 25 Chlaenius (Sphodromimus) caperatus sp. nov., holotype 26 C. (Sphodromimus) hunanus (Morvan, 1997) (Guangdong, Nanling) 27 C. (Sphodromimus) deuvei (Morvan, 1997) (Guangxi, Huaping) A apical lamella B left lateral view C dorsal view D ventral view E right lateral view. Left scale bars: 0.5 mm (A); right scale bars: 1.0 mm (B–E).

Elytra elongate, EL/BW = 1.47–1.83; slightly convex, widest near posterior third, rounded at apex in males, subtruncate in females; parascutellar striae well developed; parascutellar pores present; striae with deep punctures; interval convex basally, flat apically, densely punctate and pubescent; sutural angles obtuse; hind wings developed.

Venter densely punctate, pubescent; metepisterna (Fig. 14) long, MW/ML = 0.75–0.92; abdominal sternites III-VI with one setiferous puncture each side, sternite VII with one pair of setiferous punctures in males, two pairs in females.

Legs long and slender; tarsi nearly smooth dorsally.

Male genitalia. Median lobe (Fig. 30B–E) large, long, strongly bent to ventral side; apical orifice opened dorsally, long and wide, not reaching basal bulb; in dorsal view, apical lamella (Fig. 30A) slightly bent to left side, width slightly longer than length, apex rounded; in left lateral view, apical portion slightly bent ventrally at apex, left side near apical lamella with a large denticulation; internal sac with flagellum slender, apex with a helical sclerite, not reaching to apical orifice; left paramere larger than right paramere, both rounded; endophallus with flagellum fine and slightly bent; basal part of flagellum without a sclerite; apical part of flagellum with a drop–shaped bursa.

Figures 28–30. 

Aedeagus of Sphodromimus spp. 28 Chlaenius (Sphodromimus) pilosus (Casale, 1984) (Yunnan, Pingbian) 29 C. (Sphodromimus) yinggelingensis sp. nov, holotype 30 C. (Sphodromimus) flavofemoratus Laporte, 1834 (Yunnan, Menglun) A apical lamella B left lateral view C dorsal view D ventral view E right lateral view. Left scale bars: 0.5 mm (A); right scale bars: 1.0 mm (B–E).

Female genitalia. Bursa copulatrix (Fig. 38A–C) very long, base with trapeziform protrusion; villous canal long, tortuously contorted, adhered to common oviduct; spermatheca and spermathecal gland absent.

(Fig. 40) China (Fujian; Guangdong; Guangxi; Guizhou; Hainan; Hong Kong; Yunnan), Indonesia, Laos, Myanmar, Vietnam.

Due to the fully developed hind wings and the shape of pronotum, this species is very special among species of Sphodromimus. It also has a wider distribution than other species. But the morphological characteristic of the apical lamella of the aedeagus, denticulate on the dorsal side, and the mentum with a bifid tooth show that the species belongs to Sphodromimus. The subspecies C. (Sphodromimus) flavofemoratus enleensis Mandl, 1992 was upgraded as a valid species (see below).

Indo Chine.

Total 5 specimens. Vietnam: Holotype, Male (NHMB), Indo Chine coll. Dussault/ En-Le 1908/Chl (Macrochlaeniles) flavofemoratus ssp. enleensis Dr. K. MANDL det. 1978/Holotype [red label]. Paratype (DWC, photo), 1 ♂ (IZAS), Vietnam, Tam Dao, 20–28.VI.1990, Dr. Blazicek lgt./Paratypus, Chlaenius (Haplochlaenius) tamdaoensis mihi det. Kirschenhofer 2001[red label]/COLL WRASE, BERLIN; 1 ♂ (IZAS), Vietnam, Tam Dao, 60 km NW Hanoi, 900 m, 1997 May-June, S. Ryabov [internal sac fully everted]; 1 ♂ (IZAS), Vietnam, Cao Bang, Nguen Binh, 800 m, 2003.V.13, S. Ryabov [internal sac partially everted]; 1 ♂ (IZAS), Tonkin, Hoa-Binh, leg. A. de Cooman [genitalia damaged by dermestes beetle].

Pronotum green to coppery. PW/PL = 1.06–1.12; PAW/PBW = 0.82–0.95 (Fig. 24); pronotum cordate with anterior angles rounded, not projected forward; disc sparsely punctate and pubescent, with shallow transverse rugosities. Elytral intervals gently convex throughout; densely punctate and pubescent. Hind wings reduced. Metepisterna short; MW/ML = 1.11–1.17 (Fig. 16). Apex of femora dark brown or yellow-brown, the rest of legs black.

BL = 21.7–24.1 mm, BW = 8.2–8.7 mm. PL = 4.5–4.9 mm, PW = 5.0–5.3 mm, MW = 2.0–2.1 mm, ML = 1.7–1.8 mm. Head, elytra, venter dark and black; pronotum green to coppery; antennae, labial and maxillary palpi, apex of mouthparts and tarsomeres dark brown; apex of femora dark brown or yellow-brown, the rest of legs black.

Head. Vertex finely punctate, pubescent, without a distinct glabrous area; antennae long, reaching middle of elytra; antennomere 3 ~ 1.5× longer than antennomere 4.

Pronotum cordiform, PW/PL = 1.06–1.12 (Fig. 24), widest at apical third; anterior margin slightly concave, PAW/PBW = 0.82–0.95; lateral margins distinctly narrowed from middle to base, slightly sinuate before posterior angles; anterior angles rounded, not projected forward; posterior angles nearly right angled, rounded at tips; disc gently convex, sparsely punctate and pubescent, with shallow, transverse rugosities, without glabrous area; median line distinct, fine, not reaching anterior margin and base; basal foveae deeply arcuate, punctate and pubescent.

Elytra elongate, EL/BW = 1.58–2.22, gently convex near anterior third, widest near posterior third, rounded at apex in males; striae with deep punctures; parascutellar striae well developed; parascutellar pores present; intervals gently convex throughout, densely punctate and pubescent; sutural angles sharp at tips; hind wings reduced.

Venter densely punctate, pubescent, metepisterna (Fig. 16) short, MW/ML = 1.11–1.17; abdominal sternites III–VI with one setiferous puncture each side, sternite VII with one pair of setiferous punctures in males; all abdominal sternites with distinct impressions laterally.

Legs long and slender; tarsi nearly smooth dorsally.

Male genitalia. Median lobe (Figs 32B–E, 33B–E) long, strongly bent to ventral side; apical orifice opened dorsally, long and wide, not reaching basal bulb; in dorsal view, apical lamella linear, longer than basal width (Figs 32A, 33A), distinctly bent to right side, with a denticulation laterally in the middle, left denticulation distinctly larger than the right one; in left lateral view, apical portion slightly bent dorsally at apex, basal orifice ~ 90° relative to preapical shaft; left paramere larger than right paramere, both helically curved; endophallus (Fig. 32F) with flagellum helically thick and straight; basal part of flagellum with irregular bursa; apical part of flagellum with triangular sclerite.

Figures 31–33. 

Aedeagus of Sphodromimus spp. 31 Chlaenius (Sphodromimus) davidi nom. nov. (Guangdong, Xinyi) 32 C. (Sphodromimus) enleensis Mandl, 1992 (Vietnam, Tam Dao) 33 C. (Sphodromimus) enleensis Mandl, 1992 (holotype, “Indo Chine”) A apical lamella B left lateral view C dorsal view D ventral view E right lateral view F endophallus. Left scale bars: 0.5 mm (A); right scale bars: 1.0 mm (B–F).

Figures 34–36. 

Internal reproductive system of females 34A–C Chlaenius (Sphodromimus) caperatus sp. nov., paratype 35A–C C. (Sphodromimus) hunanus (Morvan, 1997) 36A–C C. (Sphodromimus) deuvei (Morvan, 1997) A spermatheca B dorsal view C ventral view. Vertical scale bars: 0.5 mm (A); horizontal scale bars: 1.0 mm (B, C)

Figures 37–39. 

Internal reproductive system of females 37 Chlaenius (Sphodromimus) yinggelingensis sp. nov., paratype 38 C. (Sphodromimus) flavofemoratus Laporte, 1834 39 C. (Sphodromimus) davidi nom. nov. A bursa copulatrix B dorsal view C ventral view. Scale bars: 1.0 mm.

Female genitalia unknown.

(Fig. 40) Vietnam (Indo Chine). We mark En-le, Yunnan on the map with a question mark ‘?’.

Figure 40. 

Distribution of Chlaenius (Sphodromimus) species in China and adjacent areas (a possible distribution of C. enleensis, En-le, Yunnan, represented by a question mark).

Lorenz (1998) and Lorenz (2005) proposed C. flavofemoratus enleensis Mandl, 1992 as a synonym of C. flavofemoratus, but Kirschenhofer (2017) treated it as a distinct subspecies. Based on its original description, its pronotum longer than C. flavofemoratus, and black femora indicated it probably represented a different species. After the examination of the holotype and its dissected genitalia, we find that it has the same apical lamella as C. tamdaoensis. As a consequence of this surprising discovery, the locality should also be critically revised. The labels of the holotype contain two localities (“En-le” and “Indo Chine”), the latter including today’s Vietnam. Based on the type locality of C. tamdaoensis, it is very unlikely that C. flavofemoratus enleensis also occurs in Yunnan’s En-le. Hence, we think that the label of En-le is likely to be the wrong one and may have been erroneously added, as it rarely happened in the historical collections of the NHMB (e.g., Caldara et al. 2022). We upgrade C. enleensis as a valid species and consequently treat C. tamdaoensis as synonym of C. enleensis.