Parena (Bothynoptera) dorsigera ( Schaum, 1863 )

Shi, Hongliang & Liang, Hongbin, 2023, Taxonomic revision of the genus Parena Motschulsky, 1860 (Coleoptera, Carabidae, Lebiini, Metallicina), Zootaxa 5286 (1), pp. 1-144 : 51-55

[11] Parena (Bothynoptera) dorsigera ( Schaum, 1863)

Habitus: Figs 27A–F . Male genitalia: Figs 28 , 29 . Gonocoxites of ovipositor: Fig. 11B .

Schaum, 1863: 76 (original: Bothynoptera ; type locality: N. India; holotype in MNHU); Chaudoir, 1872: 181 ( Bothynoptera ); Jacobson, 1907: 403 ( Bothynoptera ); Andrewes, 1927a: 108 ( Bothynoptera ); Andrewes, 1930: 47 ( Bothynoptera ; Tonkin); Jedlička, 1951: 59 ( Bothynoptera ); Jedlička, 1963: 445 ( Bothynoptera ); Xie & Yu, 1993: 190 (Yunnan); Kirschenhofer, 2006: 88; Park et al., 2006: 103 ( Vietnam).

Bothynoptera perforata Bates, 1873: 313 (type locality: Japan: Hiogo; lectotype in MNHN); Jacobson, 1907: 403 ( Bothynoptera ); Jedlička, 1951: 59 ( Bothynoptera ); Jedlička, 1963: 445 ( Bothynoptera ); Habu, 1967: 161; Habu, 1982: 115; Lafer, 1989: 213 ( Russia: Far East ); Xie & Yu, 1993: 191 ( China: Sichuan, Yunnan); Kryzhanovskij et al. 1995: 162; Park & Kwon, 1998: 36 ( South Korea); Kirschenhofer, 2006: 89. Syn. nov.

Parena nepalensis Kirschenhofer, 1994: 1021 (type locality: Nepal: Kosi; holotype in NMW: Naturhistorisches Museum Wien, Vienne, Austria). Syn. nov.

Parena kunmingensis Kirschenhofer, 1996: 770 (type locality: Yunnan: Kunming ; holotype in NMW); Kirschenhofer, 2006: 88. Syn. nov.

Type material examined. Bothynoptera dorsigera Schaum : Holotype (MNHU, Fig. 27A ): female, body length = 10.4 mm, pin mounted, "42351", " Bothynoptera / dorsigera m. / Ind. bor. " [yellow label]; " Bothynoptera / dorsigera Schaum " [yellow box label].

Bothynoptera perforata Bates : Lectotype (MNHN, Fig. 27B ), designated herein: male, body length = 11.0 mm, pin mounted, " Hiogo / Japan ", "TYPE" [red label], " Bothynoptera / perforata / Bates ", "Ex Musaeo / H.W. Bates / 1892", "MUSEUM PARIS / 1952 / Coll. R. OBERTHUR", " LECTOTYPE / Bothynoptera perforata / Bates, 1873 / des. SHI H.L. 2011" [red label]. Paralectotype (NHML): 1 ex, "Type" [round label with red circle]; "HIOGO", " Japan./ G. Lewis./ 1910-320", " perforata / Bates ".

Notes on types and synonyms. Bothynoptera dorsigera Schaum : This species was originally described from a single specimen from Northern India. The female we examined in the collection of MNHU in accord with the original description is the holotype.

Bothynoptera perforata Bates : This species was originally described from three specimens from Hiogo. We found two specimens in the collection of MNHN and NHML in accord with the original description. We herein designate the male deposited in MNHN bearing Bates's handwriting label ( Fig. 27B ) as lectotype.

As indicated in the original description ( Bates, 1873), B. perforata is different from B. dorsigera in having the pronotum wider, elytra without yellow patch and the dorsum a lighter color. However, as discussed below, all these differences are considered as geographical variations, and specimens with intermediate features occurs in several localities of China. So, B. perforata is treated as a junior synonym of P. dorsigera herein.

Parena nepalensis Kirschenhofer : In the original description ( Kirschenhofer, 1994), this species was compared only with P. andrewesi Jedlička but not with any species previously placed in Bothynoptera . We did not examine the type of this species. However, from the original description and illustration, it is identical to the holotype of P. dorsigera and other specimens examined from Nepal ( Fig. 27D ). So, we synonymize it with P. dorsigera (Schaum) herein.

Parena kunmingensis Kirschenhofer : The original description (Kirschenhofer, 1996) pointed out that this species is very similar to P. nepalensis , and has small differences in pronotum shape, elytral pore size, and shape of elytral apices. As discussed below, these differences are considered as geographical variations. Thus, we synonymize this species with P. dorsigera (Schaum) herein.

Non-type material examined. Japan: 2 ex (MNHN), "leg. A. Habu, Mt. Hiko, Fukuoka P., IX.15,1952". 1 ex (MNHN), "(Pref. Tokyo), Mt Tukao, Jun.14.1951". 1 ex (MNHN), "Yatsugatake, 4.8.52". 1 ex (MNHN), "Japan; Janson, Acq. 1884". 1 ex (MNHN), "Japan". 1 female (IZAS), "[Japan: Honshu] Nagano, Koumi, Toyosato, Yamanokami (1602-1608 m), 36°02'51.9''N~ 36°02'54.9''N, 138°24'16.3''E ~ 138°24'20.83''E 7.VIII.2007 Junhao Huang" < Fig. 5J >. China: 1 female (IZAS), " Beijing, Changping, Huyu; 1997.VII.14, Zhou Haisheng lgt." < Fig. 5I >. 1 female (HBUM), " Henan, Dengfeng, Shaolinsi, 2002.VII.16, Yang Xiujuan lgt.". 1 ex (IZAS), " Shaanxi, Foping, Shangshawo, on vegetation, N33.59716 E108.01366, 1107 m, 2007.VIII.15, Shi Hongliang & Yang Ganyan lgt.". 1 female (IZAS), " Shaanxi, Foping, Longcaoping, on vegetation, N33. 67662 E107.97975, 1369 m, 2007.VIII.17, Shi Hongliang & Yang Ganyan lgt.". 1 female (IZAS), " Shaanxi, Ningshan, Huoditang, on vegetation, N33.43368, E108.44747, 1538 m, 2007.VIII.18, Shi Hongliang & Yang Ganyan lgt." < Fig. 11B >. 1 ex (IZAS), " T’ienmu Shan, 1936.VII.27, O. Piel coll.". 1 male, 1 ex (IZAS), " Zhejiang, Tianmushan, Xianrending, light trap, 1998.VII.29, Wang Xue lgt." < Figs 10C , 27F , 28A >. 1 ex (IZAS), " Zhejiang, Tianmushan, Xianrending, light trap, 1998.VIII.17, Wu Hong lgt.". 1 ex (IZAS), " Zhejiang, Lin'an, W. Tianmushan, Tianmushuyuan-S.Gate, 2006.VI.30, Huang Junhao lgt.". 1 ex (IZAS), " Zhejiang, Lin'an, W. Tianmushan, 2008.VI, Huang Hao lgt.". 1 ex (IZAS), " Zhejiang, Lin'an, W. Tianmushan, Laodian-Xianrending, 1020 m, N30°18' E119°25', 2009.VII.3, Zhu Yijin lgt.". 1 male (CRS), "China, Hangchow/ 30°18’ N 120°07’ E/ 10.vi.1921 / Eigin Suenson leg.". 1 ex (IZAS), " Hunan, Cili, 1988.IX.3-6 ". 3 ex (IZAS), " Sichuan, Luding, Moxi, Hailuogou, light trap, N29.60261, E102.06402, 2130 m, 2007.IX.11 N, Liang Hongbin lgt.". 1 ex (CAS), " Sichuan, Luding, Moxi, Hailuogou, light trap, N29.60261, E102.06402, 2130 m, 2007. IX.12 N, Liang Hongbin lgt.". 1 ex (CAS), " Sichuan, Luding, Hailuogou, 2007.IX.12, light trap, David Kavanaugh lgt.". 1 ex (IZAS), " Sichuan, Emeishan, Baoguosi, 550-750 m, 1957.VI.4, Lu Youcai, lgt.". 1 ex (HBUM), " Sichuan, Jiulong, Hongba, Shi Fuming lgt., 2008.9.23 ". 1 ex (IZAS), " Guangxi, Guilin, Longsheng, Huaping, 2008.V.2, Cao Liangming, Li Hu lgt.". 1 male (CCCC), " Guangxi, Jinxiu, Dayaoshan, Luoyingou, 1250 m, 2019.VIII-06, leg. J.- T. Zhao" < Fig. 4E >. 1 female (CCCC), " Guangxi, Jinxiu, Dayaoshan, Pingbantun, 1250 m, 2019.IV-19, leg. J.-T. Zhao". 1 female (CCCC), " Guangxi, Jinxiu, Dayaoshan, Laoshanlinchang, 1500 m, 2019.V-31, leg. J.-T. Zhao" < Figs 2C , 3C >. 1 male (CCCC), " Guangxi, Jinxiu, Dayaoshan, Laoshanlinchang, 1599 m, 2017.IX-15, leg. J.-T. Zhao". 1 female (CAU), "Lijiang, Yulong, Mingyin, Jiazi vill., 2818 m, 2020.06-17, 100.26927 E 27.1281 N, Huang Weidong lgt. 5307211NSMT0008". 1 ex (IZAS), "Yunna, Pingbian, Yuping, Shuiweicheng, 22.91278 103.69760 2058 m, 2021.5.18. Wang Yong coll.". 1 ex (NHMB), " Yunnan, 8-10 Jun, Yipinglang, 1800 m, 25.05 N 101.53 E, Bolm lgt. 1993". 1 ex (IZAS), " Yunnan, Menglong Banna, Mengsong, 1600 m, 1958.IV.26, Zhang Yiran lgt.". 1 ex (IZAS), " Yunnan, Menglong Banna, Mengsong, 1600 m, 1958.IV.26, Wang Shuyong lgt.". 1 female (CCCC), "China, Yunnan, Gongshan, Galabo, 2680 m, 2019.VIII-11, Leg. Y.-B. LI" < Fig. 5H >. 2 females (CCCC), " Yunnan, Kunming, Xishan, 2017.X-6, alt. 2122 m, Yang Xiaodong leg." < Figs 12B , 27C >. 1 female (IZAS), " Yunnan, Weixi, Pantiange, 2750 m, 1981.VII.22, Wang Shuyong lgt.". 1 ex (HBUM), " Xizang, Mêdog, Hanmi, 2380 m, 2003.VIII.9, Ren Guodong lgt.". 1 male (CCCC), "Taiwan, Nantou county, Chushan, Shanlinhsi, light trap, 2006. IX.27-29, C.J.K & C.C.R lgt." < Figs 27E , 28B >. Vietnam: 1 ex (NHML), "Tonkin. Chapa., May 1916., R.V. de Salvaza". Laos: 1 ex (NHMB), "Lao-NE, Hua Phan prov. ~20°12’ N 104°01’ E, Phu Phan Mt. 1500-1900 m, 17.v.- 3.vi.2007, M. Brancucci leg.", "NHMB Basel expediton to Laos, 2007". India: 1 ex (MNHM), " dorsigera Schaum, Ind. Orient bor., Bacon. C. Laferte". 4 ex (MNHN), "Harmand, Sikkim, 1890" < Fig. 29B >. 1 ex (NHML), "Gopaldhara, Br. Sikkim. H. Stevens.". 1 ex (MNHN), "Kurseong, R.P. Decoly, 1898". 1 ex (NHML), "Inde Angalise, Pedong, Region de Darjeeling., Chasseurs indigenes, 1935". 1 ex (NHMB). "NE India, Meghalaya; 1999, 3km E Tura; 1150 m; 25°30’ N 90°14’ E; 4.v.; Dembicky & Pacholaiko leg.". 1 ex (NHMB). "NE India, Megalaya, SW of Shillong; 1600 m, 25°34’ N91°51’20’’ E; L. Dembicky leg. 14.v.2004 ". 1 ex (NHML), "Allahabad". Nepal: 1 ex (NHMB). "Nepal, Kosi-#4, Pakhribas 27°03’N/ 87°18’E to Mangmaya 27°07’N/ 87°15’E, 1700- 300 m, 29.v.01", "NHMB Basel, expedition to Nepal 2001". 1 ex (NHMB). "Nepal: Kosi-#13, Depitar 27°27’N/ 87°17’E to Barabishe 27°26’N/ 87°18’E, 1250- 560 m, 12.vi.01", "NHMB Basel, expedition to Nepal 2001". 1 male (CRS), "Nepal Bagmati / Nuwakot", "Pati Bhanjyang/ 1900 m 16-18.VI.89/ M. Brancucci" < Figs 27D , 29A >. 1 ex (CMB), "Nep: Mahakali / Darchula, env. Batar, slope above, Chamliya Khola, 1900 m, 29°50’54’’N, 80°54’03’’E, 12.VI.2005, leg. A. Weigel".

Comparisons. P. dorsigera is easily distinguishable from all other species of the genus in having four (occasionally five or six) very large setigerous pores on elytral interval 3. In the subgenus Bothynoptera , P. kurosai and P. tesari are somewhat similar to P. dorsigera in their relatively large body size and dorsum dark brown without a patch, but differ in having elytral interval 3 with only three discal pores. P. tesari also different from P. dorsigera in having the elytral apices evenly rounded.

Description. Body length 9.2–11.0 mm. Dorsum reddish brown to piceous, head with a triangular light patch on frons in many specimens, elytral interval 1 often lighter, elytra disc sometimes with a vague light patch. Tempora short, abruptly narrowed behind eyes; postgenae with a pair of long suborbital setae. Pronotum nearly quadrate, PW/PL = 1.20–1.40, usually slightly narrower than head, PW/HW = 0.90–1.02; lateral explanations narrow; widest at anterior third, slightly rounded at anterior half, and then somewhat sinuate before posterior angles; posterior angles nearly rectangular. Elytra without microsculpture; striae not incised, replaced by rows of fine punctures; intervals completely flat, with very sparse fine punctures; lateral intervals sometimes slightly uneven; interval 3 with four setigerous pores, with five or six in some specimens, first two pores adjacent to stria 3, the others adjacent to stria 2, pores very large and strongly concave, forming fovea slightly wider than interval width; interval 5 with one pore near middle in a few specimens; apical truncation rather distinct, bisinuate, straight, or concave inward, outer apical angles prominent but apically rounded; sutural angles indistinct or with very faint denticles. Apex of abdominal sternite VII with two setae on each side in both sexes, occasionally three. Males with biseriate adhesive setae on apical half of mesotarsomere 2 and full length of mesotarsomere 3. Median lobe of aedeagus moderately stout (AL/AW = 4.0–4.3), apical lamella short, coniform. Endophallus with large primary sclerite, apical flagellum reaching middle of median lobe; apical sclerite well defined, basal core ovate; basal sheath small, finely scaled, apical sheath heavily scaled; squamate sac not divided, large, near middle of median lobe, right to squamate sheath. Gonocoxite II of ovipositor nearly quadrate, inner angle slightly pointed to apex, apex with six or seven ensiform setae, subequally arranged.

Distribution ( Map 6 , blue). China (Beijing, Henan, Shaanxi, Zhejiang, Hunan, Guangxi, Sichuan, Yunnan, Xizang, Taiwan), Russia (Primorye), Japan, South Korea, Vietnam, Laos, Nepal, North India.

Geographical variation. This species has a wide distributional range in East Asia, northeast to Japan and southwest to Nepal ( Map 6 ). Its external features vary geographically in the following aspects:

(1) Coloration: the dorsum is gradually darkened from north to south. In northern localities such as Japan, Russia, Korea, and North China, the dorsum is always reddish brown ( Figs 27B, 27F ); in southern localities such as Vietnam, Laos, and Taiwan, the dorsum is often piceous ( Fig. 27E ), but dark brown individuals occur as well. In the western localities such as Yunnan, Nepal, and Sikkim, the dorum is usually dark brown to reddish brown ( Figs 27C, 27D ).

(2) Elytra patch: the reddish yellow elytra central patch only presents in specimens from Nepal and North India ( Fig. 27D ), such as the types of P. dorsigera and P. nepalensis . However, the background color of the elytra is light brown in some specimens, so that the patch is somewhat indistinct ( Fig. 27A ).

(3) Elytral apex: the elytral apical truncation is very distinct with prominent outer apical angles in this species. However, the shape of elytra apices varies slightly from northeast to southwest. In specimens from Japan and Primorye, the elytra apical truncation is bisinuate with the inner half curved outward and the outer half slightly concave inward ( Fig. 5J ), as in the type of P. perforata . In specimens from northern and eastern China, the elytra apical truncation is often nearly straight ( Fig. 5I ). In southern localities, such as Nepal, Vietnam, and southernmost China, the elytra apical truncation is strongly concave inward ( Fig. 5H ). However, the degree of curvature gradually increases from north to south in China.

Although these above differences are evident, we apply a wide species concept of P. dorsigera , without dividing it into subspecies, because these characters gradually vary by distance, and there is no significant difference in male genitalia among specimens from different localities ( Figs 28 , 29 ).

Remarks. Xie & Yu (1993) recorded P. dorsigera from Yunnan, and indicated that this species is different from P. perforata in having five setigerous pores on interval 3 and one pore on interval 5. After the examination of their specimen, we found the description of P. dorsigera was based on an aberrant individual.

Figures

FIGURE 2. Mandibles of Parena spp.: left mandible, dorsum (ld); right mandible, dorsum (rd); left mandible, venter (lv); right mandible, venter (rv). A P. (Crossoglossa) cavipennis (Bates), Shaanxi. B P. (Parena) latecincta (Bates), Zhejiang. C P. (Bothynoptera) dorsigera (Schaum), Guangxi. D P. (Bothynoptera) taiwana Hua, Taiwan. E P. (Bothynoptera) tripunctata (Bates), Beijing. Scale bar = 0.5mm. Abbreviations: rr retinacular ridge of right mandible; vg ventral groove.

FIGURE 3. A–D Labia of Parena spp.: A P. (Crossoglossa) cavipennis (Bates), Beijing. B P. (Parena) latecincta (Bates), Zhejiang. C P. (Bothynoptera) dorsigera (Schaum), Guangxi. D P. (Bothynoptera) tripunctata (Bates), Beijing. E–F Heads of Parena spp. in lateral view: E P. (Bothynoptera) taiwana Hua, Taiwan, suborbital setae present. F P. (Parena) latecincta (Bates), Yunnan, suborbital seta absent. Scale bars = 0.5mm.

FIGURE 4. Elytral base (left) of Parena spp. A P. (Parena) bicolor Motschulsky, Sumatra. B P. (Parena) africana (Alluaud), Cameroon. C P. (Crossoglossa) laesipennis (Bates), Taiwan. D P. (Bothynoptera) tripunctata (Bates), Beijing. E P. (Bothynoptera) dorsigera (Schaum), Guangxi. F P. (Bothynoptera) heteronycha sp. n., Laos, paratype. Scale bars = 0.5mm (a for fig. D, b for figs A B E F, c for fig. C). Abbreviations: bp elytral basal pore; sa the apex of scutellum; d1 the first discal setigerous pore.

FIGURE 5. Elytral apex (left) of Parena spp. A P. (Parena) africana (Alluaud), Cameroon. B P. (Parena) ruficornis sp. n., Central Africa, paratype. C P. (Parena) valeriae Facchini, holotype. D P. (Parena) madagascariensis (Alluaud), Madagascar. E P. (Bothynoptera) tesari (Jedlička), Taiwan. F P. (Bothynoptera) heteronycha sp. n., Laos, paratype. G P. (Bothynoptera) kurosai Habu, Japan. H P. (Bothynoptera) dorsigera (Schaum), Yunnan. I P. (Bothynoptera) dorsigera (Schaum), Beijing. J P. (Bothynoptera) dorsigera (Schaum), Japan. Scale bars = 0.5mm

MAP 6. Distributions for species in the Parena (Bothynoptera) dorsigera group.

FIGURE 10. Parameres of male genitalia for Parena spp. A P. (Crossoglossa) cavipennis (Bates), Beijing. B P. (Crossoglossa) testacea (Chaudoir), Shanghai. C P. (Bothynoptera) dorsigera (Schaum), Zhejiang. D P. (Bothynoptera) taiwana Hua, Taiwan. E P. (Bothynoptera) tripunctata (Bates), Shaanxi. F P. (Parena) rubripicta Andrewes, Xizang. G P. (Parena) fasciata (Chaudoir), Taiwan. H P. (Parena) nigrolineata (Chaudoir), Taiwan. I P. (Parena) latecincta (Bates), Yunnan. Scale bar = 0.2 mm.

FIGURE 11. Gonocoxites (left side) of ovipositor for Parena spp., ventral side (dorsal side for fig. Q). A–J subgenus Bothynoptera: A P. (Bothynoptera) heteronycha sp.n., Laos, paratype. B P. (Bothynoptera) dorsigera (Schaum), Shaanxi. C P. (Bothynoptera) kurosai Habu, Guangdong. D P. (Bothynoptera) tesari (Jedlička), Taiwan. E P. (Bothynoptera) taiwana Hua, Taiwan. F P. (Bothynoptera) tripunctata (Bates), Shaanxi. G P. (Bothynoptera) malaisei (Andrewes), Taiwan. H P. (Bothynoptera) quadrisignata Mateu, Yunnan. I P. (Bothynoptera) emarginata sp. n., holotype. J P. (Bothynoptera) triguttata sp. n., Sichuan, paratype. K–X subgenus Parena: K P. (Parena) bicolor Motschulsky, Sulawesi. L P. (Parena) rubripicta Andrewes, Yunnan. M P. (Parena) fasciata (Chaudoir), Taiwan. N P. (Parena) nigrolineata (Chaudoir), Taiwan. O P. (Parena) picipes sp. n., Borneo, paratype. P P. (Parena) andrewesi Jedlička, Philippine. Q P. (Parena) politissima (Chaudoir), Vanuatu. R P. (Parena) latecincta (Bates), Guizhou. S P. (Parena) circumdata Shibata, Taiwan. T P. (Parena) pendleburyi Andrewes, Borneo. U P. (Parena) fulva sp. n., Zimbabwe, paratype. V P. (Parena) madagascariensis (Alluaud), Madagascar. W P. (Parena) ruficornis sp.n., CentralAfrica, paratype. X P. (Parena) africana (Alluaud), Uganda. Y–γ subgenus Crossoglossa: Y P. (Crossoglossa) cavipennis (Bates), Taiwan. Z P. (Crossoglossa) laesipennis (Bates), Yunnan. α P. (Crossoglossa) testacea (Chaudoir), Zhejiang. β P. (Crossoglossa) sulawesiensis Kirschenhofer, Sulawesi. γ P. (Crossoglossa) sciakyi sp. n., holotype. Scale bar = 0.2mm.

FIGURE 12. Female internal reproductive tracts of Parena spp. A P. (Crossoglossa) cavipennis (Bates), Beijing. B P. (Bothynoptera) dorsigera (Schaum), Yunnan. C P. (Parena) nigrolineata (Chaudoir), Yunnan. Scale bar = 0.5 mm.

FIGURE 27. Habitus of Parena (Bothynoptera) dorsigera species group. A Holotype of Parena dorsigera (Schaum). B Lectotype of Bothynoptera perforata Bates [= Parena dorsigera (Schaum)]. C P. dorsigera (Schaum), female from Yunnan (Kunming). D P. dorsigera (Schaum), male from Nepal. E P. dorsigera (Schaum), male from Taiwan. F P. dorsigera (Schaum), male from Zhejiang. G Holotype of Parena wrasei Kirschenhofer [= Parena kurosai Habu]. H P. kurosai Habu, male from Nepal. I P. kurosai Habu, male from Guangdong. Scale bars = 2 mm.

FIGURE 28. Median lobe of male genitalia for Parena (Bothynoptera) dorsigera (Schaum), right-lateral (r), ventral (v), leftlateral (l), and dorsal (d) views. A Zhejiang. B Taiwan. Scale bar = 0.5 mm.

FIGURE 29. Median lobe of male genitalia for Parena (Bothynoptera) dorsigera (Schaum) from different localities, left-lateral (l) and dorsal (d) views: A Nepal. B India. C Yunnan. Scale bar = 0.5 mm.