Coilodera penicillata Hope, 1831

Qiu, Jian-Yue, Xu, Hao & Chen, Li, 2017, Taxonomic review of the Oriental flower beetle Coilodera penicillata species complex (Coleoptera: Scarabaeidae: Cetoniinae), Zootaxa 4350 (3), pp. 511-537 : 521-523

Coilodera penicillata Hope, 1831

( Figs. 21–24 , 52–56, 63–64, 70 , 79–80, 85 , 98–102 )

Coilodera penicillata Hope, 1831: 25 (type locality: Nepal); Paulian 1960: 13 (Chapa, northern Vietnam; Xieng Khouang & Lakhon, Laos), fig. 290; Nakane 1972: 112 (Nepal), fig. 1 (parameres), 6 (habitus); Antoine 1986: 18, fig. 10b, 10d ♂; Ma 1987: 573 (Lincang, Yunnan, China), plate 2, fig. 26; Sakai & Ikeda 1996: 35 (northern Thailand); Krajčík 1998: 88; Sakai & Nagai 1998: 357 (Nepal), plate 136, fig. 1625-1 ♀; Smetana 2006: 306; Ek-Amnuay 2008: 246 (northern Thailand), plate 95, figs. C69 ♂ & ♀; Sakai 2008: 20 (Xieng Khuang & Phu Pan, Laos), fig. 60 ♂; Krajčík 2011: 53; Krajčík 2012: 76; Li et al 2013: 32 (Mount Gaoligongshan, Yunnan, China), fig. 37 ♂; Bezděk 2016: 401.

Coelodera penicillata Hope : Burmeister 1842: 321; Schoch 1896a: 378 (Assam, India); Schoch 1896b: 34; Schenkling 1921: 133; Mikšić 1972: 700 (Nepal; northeastern India; northern Myanmar; northern Vietnam), figs. 18–19; Mikšić 1976: 268, fig. 32d; Ma 1995: 41, fig. 33.

Macronota penicillata (Hope) : Harold 1869: 1298; Gestro 1891: 843 (Bhamo, northeastern Myanmar); Arrow 1910: 44 (northeastern India; Keren Hills, Myanmar).

Macronota dives (nec Gory & Percheron): Schaum 1844: 367; Schaum 1845: 43; Schaum 1848: 27. (misapplication of the name)

Diagnosis. Tomentum on body surface orange-red. The length of male antennal club about 1.5 times as the length of antennomeres 2–7 combined. Posterior margin of the median tomentose area of pronotum notched. Scutellum usually with arrow-shaped tomentose area ( Figs. 52–53 ). Sutural maculae and anteapical maculae usually separated. Apex of mesometasternal process slightly curved; mesometasternal suture not engraved, almost without setae ( Fig. 63 ). Median portion of abdomen usually glabrous. Outer teeth of metatibia longer than ventral teeth in both sexes ( Fig. 70 ). Basal piece of male genitalia distinctly convex ( Fig. 80 ).

Variation. Body length 18.5–22.4 mm; width 8.8–10.8 mm. Sutural maculae and anteapical maculae occasionally connected in some individuals from western Yunnan, China ( Fig. 22 ). Posthumeral maculae sometimes absent ( Figs. 21, 23 ). Most individuals have arrow-shaped tomentose area on their scutellum ( Figs. 21– 22 , 52–53 ), but that of the four individuals from Mount Daweishan (Yunnan, China) and central Vietnam is almost glabrous ( Figs. 23 , 54–56 ). A Chinese specimen from western Yunnan is setiferous along the mesometasternal suture ( Fig. 64 ).

Type material. The type series cannot be located, and are perhaps lost ( Arrow 1910).

Other material examined (63♂♂, 109♀♀). NEPAL: 1♂ ( MFNB) , 1♀ ( NHMB), Nepal, Brit. Ind., Coll. Dr. Itzinger ; 2♀♀ ( NHMB), Kosi-#3, Pakhribas , 1,700–1,900m, 27°03′N, 87°18′E , 27–28.V.2001 // NHMB Basel expedition to Nepal May/ June 2001 ; INDIA: Sikkim: 1♂ ( MNHN) , British India, Sikkim, Lachen-Dachung , VIII.1933; 1♀ ( IRSNB), Sikkim ; West Bengal: 1♂ ( SDEI), British Bootan, Padong [Pedong, Darjeeling], L. Durel 1891 ; 2♀♀ (BMNH, Bourgoin Coll.), 3♂♂, 7♀♀ (MNHN), British Bootan, L. Durel 1898; 2♀♀ (BMNH, Bourgoin Coll.), 1♂, 1♀ (IRSNB), British Bootan, Padong, L. Durel 1913; 1♂, 1♀ (BMNH, Bourgoin Coll.), 5♀♀ (MNHN), British Bootan, Padong, L. Durel 1914; 2♂♂, 2♀♀ (BMNH, Bourgoin Coll.), British Bootan, Maria Basti, L. Durel; 1♂, 2♀♀ (BMNH, Bourgoin Coll.), Darjeeling, Mowis 1891; 1♂, 1♀ ( BMNH), India: Kurseong Div. Lat Panchar, 4000 ft. , VI.1934, H.G. Champion; 1♂ (MFNB), Mungphu [Mungpoo]; 1♀ (SDEI), Darjeeling; 1♀ (SDEI), Pedong, Desgodins; 1♂ (MSNG), Mungphu 1890, D. Atkinson; 1♂ (NHMB), Pedong bei, Darjeeling 1935, Slg, R. Oberthür, Eing. Nr.4.1956; 1♀ (NHMB), Darjeeling; 1♂, 7♀♀ ( MNHN), Inde Anglaise, Pedong Region de Darjeeling. Chasseurs indigenes 1933 ; 3♂♂, 3♀♀ ( MNHN), Inde Anglaise, Pedong District de Darjeeling. Chasseurs indigenes 1932 ; 3♀♀ ( MNHN), Inde Anglaise, Pedong Region de Darjeeling. Chasseurs indigenes 1934 ; 1♂, 2♀ ( MNHN), Inde Anglaise, Pedong Region de Darjeeling. Chasseurs indigenes 1937 ; 2♂♂ (MNHN), Maria Basti, Bootang; 1♀ (MNHN), British Bootang, L. Durel, 1899; 5♀♀ (MNHN), British Bootang, Maria Basti, L. Durel; 1♀ (MNHN), Kurseong, R.P. Decoly, 1898; 1♂ ( MNHN), Environs de Kurseong , R.P. Bretaudeau ; Assam: 1♂ , 4♀♀ ( BMNH, Fry Coll.), Assam, Polkai Mount // Doherty ; Meghalaya: 1♀ ( MNHN, Oberthür Coll.) , Assam Reg. de Shillong ; Manipur: 1♀ ( BMNH, G. B. Rogers Coll.), northern Manipur , Asaam, 6,000ft ; 2♀♀ (BMNH, Nevinson Coll.), India; 1♀ ( BMNH), northern Manipur ; 1♀ (BMNH), Manipur; 3♂♂ , 1♀ ( BMNH, Fry Coll.), east India, Manipur // Doherty . MYANMAR: 1♀ ( MSNG), Carin Chebà [near Taungoo], 900– 1,100 m, L. Fea, V– XII-88 ; 1♂ (MSNG), Carin Asciuii Chebà, 1,200–1,300 m, L. Fea, I-88 / penicillata Hope ; 1♀ (BMNH), Burma, 1920-132; 3♂♂, 3♀♀ ( MNHN), H te Brimanie, Mines de Rubis [near Mogok, Mandalay], 1,200–2,300 m, Doherty, 1880 . CHINA: Xizang: 1♀ ( QCCC), VI.2016, Yarlung Tsangpo Grand Canyon Nature Reserve, Hanmi, Beibeng, Mêdog County, 2,200 m, Wei Hu ; Yunnan: 2♂♂ ( BMNH), Sima [Xima, Yingjiang], 1914-145 ; 1♂ (BMNH), Zhehou, Jingdong, 15.V.1978; 1♀ (BMNH), 24.V.1985, Menghun, Xishuangbanna, 1,200–1,400 m; 1870; 1♀ ( SWU), 21.V.2015, Mount Daweishan Nature Reserve , Hekou County, Jian-Yue Qiu ; 1♂, 1♀ ( SWU), Mount Daweishan Nature Reserve , Hekou County, Jian-Yue Qiu, pupae in 21.V.2015, adults emerged in VI.2015 ; 2♂♂, 7♀♀ ( QCCC), V.2014, Manjiu, Manwan, Yunxian County, Lincang, Zi-Chun Xiong ; 3♀♀ (QCCC), F1 generation of the female from Lincang, Zi-Chun Xiong, eggs in V.2014, adult emerged in IX.2014; 1♂ ( QCCC), 10.VIII.2013, Huguo, Longchuan County, Guang-Lin Xie ; 1♂ ( QCCC), 12.VI.2015, Kangjiaba reservoir, Wumulong, Lincang County, Mao Ye ; 1♀ ( QCCC), 18.II.2016, Tongbiguan Nature Reserve, Yingjiang County, Dehong, 2,100 m, Jian-Yue Qiu & Hao Xu ; 1♀ ( QCCC), Bangzhongdazhai (S233 Rd), Nongzhang, Yingjiang County, Dehong, 1,320 m, Jian-Yue Qiu & Hao Xu, 3nd instar in 18.II.2016, adult emerged in V.2016 ; 1♀ ( QCCC), VI.2015, Mount Ailaoshan , Xinping to Gasha, 1,900 m, Huang-Shun Xi ; 1♂ ( NKUM), 12.V.1984, Mount Ailaoshan , Fangjiajing, Jingdong County, Liu ; 3♀♀ ( QCCC), 30.I.2017, Mount Xiaolushan, Cizhuhe, Zhutang, Lancang County, Puer, Jian-Yue Qiu & Hao Xu ; 1♂ ( QCCC), Bulong Nature Reserve, Bulangshan, Menghai County, Xishuanbanna, Jian-Yue Qiu & Hao Xu, mature larva in 31.I.2017, adult emerged in V.2017 ; 1♂ ( QCCC), Xujialiangzi, Manla, Yiwu, Mengla County, Xishuanbanna, Jian-Yue Qiu & Hao Xu, larva in pupal cell in 1.II.2017, adult emerged in V.2017 ; 1♂ (KIZ), 6.V.2017, erfu, Mount Huanglianshan Nature Reserve , Lvchun County, 1,300 m, Zhong-Cheng Tian 2♀♀ ( QCCC) , 1.II.2017, Xujialiangzi, Manla, Yiwu, Mengla County, Xishuanbanna, Jian-Yue Qiu & Hao Xu. THAILAND: 3♂♂ ( NHMB) , Chiang Mai, 7.VI.1990, G. Minet. LAOS: 1♀ ( BMNH, Bourgoin Coll.) , Xieng Khouang, 18 May 1915 // Laos R. Vitalis; 1♂ , 2♀♀ (BMNH, Bourgoin Coll.), Muong Houoc, 10.V.1920; 1♂ (BMNH, Bourgoin Coll.), Pou Bia, 23.I.1919; 1♂ (BMNH, Bourgoin Coll.), Muong Pak, 4.XII.1918; 1♂ (BMNH, Bourgoin Coll.), Pou Mi, 27.XII.1918; 1♂ (BMNH, Bourgoin Coll.), Nam Tha, 10.IV.1918; 2♂♂ ( MNHN), Xieng Khouang, V.1960, Laos (Baudon) ; 1♂, 1♀ (MNHN), Ban Van Eua, Laos, 30.IV.1967; 1♀ ( BMNH, Bourgoin Coll.) , 1♀ ( IRSNB), Nam Long, 28.IV.1918, Laos // Macronota penicillata det. A. Bourgoin 1925 ; 1♀ ( IRSNB), Pakson ( Laos), 1,100 m ; 1♀ ( BMNH, Krajčík Coll.), 26.IV–11.V.2001, Hua Phan, Ban Saluei , Phu Phan Mountain, 20°15′N, 104°02′E, 1,500–2,000m, D. Hauck; 1♀ ( QCCC) , VI.2014, Mount Phan, 2,060 m ; 1♂ ( QCCC), VII.2011, Mount Phu Pane , Hua Phan ; 1♀ ( QCCC), IV.2014, Mount Phan , Hua Phan, 2,026 m; 1♂ , 1♀ (QCCC), 15.V.2008, Phou Pan, Houa Phan; 1♂ , 3♀♀ ( QCCC), V.2013, Mount Pan , Sam Neua . VIETNAM: 3♂♂ , 2♀♀ ( QCCC), 16–22.V.1992, Sapa, Lao Cai Province ; 1♀ ( QCCC), 1.VI.2015, Kon ka Kinh, Gia Lai, Van Dang ; 1♀ ( QCCC), 1.VII.2015, Ngoc Linh, Kon Tum, Van Dang ; 1♂ ( QCCC), IV.2016, Mucangchai, Yen Bai Province, 1,700 m ; 1♀ ( QCCC), VII.2016, Mucangchai, Yen Bai Province, 1,700 m ; 1♂ (BMNH, Bourgoin Coll.), Chapa, VI.1916; 1♂ ( BMNH, Bourgoin Coll.), Tonkin, Lao-Kay.

Distribution. China: Xizang (new record), Yunnan; Vietnam; Laos; northern Thailand; northern Myanmar; northeastern India; Nepal.

Natural history. This species usually inhabit the mountainous area at elevations of 1,000–2,500 m. The label data indicated that the adult activity from May to August. This species overwinter as mature larvae or newly emerged adults: some fresh adults were found in September 2014 (Mr. Zi-Chun Xiong, by artificial breeding), and many larvae with a fresh adult were captured in decayed wood in February 2016 within the cockroach galleries of Salgunea raggei Roth, 1979 ( Blattodea : Blaberidae : Panesthiinae ) ( Figs. 98–99 ), or Passalidae ( Fig. 102 ).

Remarks. Coilodera penicillata is similar to both C. nigroscutellaris and C. formosana , but the scutellum of the latter two species are glabrous (scutellum usually tomentose in C. penicillata ), and the distal maculae and sutural maculae on the elytra of C. formosana always connected (those of the former two species unconnected). These diagnostic characters were adopted when Mikšić (1972) revised the genus, but the latter two were recognized as two subspecies of C. penicillata , and this treatment was widely accepted ( Mikšić 1976; Miyake et al. 1991; Krajčík 1998, 2011, 2012; Sakai & Nagai 1998; Yu et al. 1998; Smetana 2006; Bezděk 2016).

The character of distal maculae connecting with sutural maculae appears in all three species discussed above ( Figs. 22, 25–26, 30 ), but the differences of the scutellum, the mesometasternal suture, and the parameres are evident and consistent. Consequently, we regarded them as three independent species: scutellum usually clad with arrow-shape yellow tomentum in C. penicillata ( Figs. 21–22 ) (except for a few individuals, see variation above), while glabrous in C. nigroscutellaris ( Figs. 29–31 ) and C. formosana ( Figs. 25–26 ); mesometasternal suture not engraved in C. penicillata ( Figs. 63–64 ) and C. nigroscutellaris ( Fig. 66 ), while deeply engraved in C. formosana ( Fig. 65 ); parameres and basal piece distinctly different between the three species as showed in Figs. 79–84 .

The distribution of Coilodera penicillata ranges from Nepal and northeastern India to northern Vietnam ( Arrow 1910; Mikšić 1972; Nakane 1972; Sakai & Ikeda 1996; Krajčík 1998, 2012; Sakai & Nagai 1998; Sakai 2008; Bezděk 2016), and the record from Yunnan, China was reported by Ma (1987). Coilodera nigroscutellaris occurs in northern Vietnam and southern China and C. formosana is endemic to Taiwan Island ( Fig. 85 ).

Figures

FIGURES 21 – 33. Habitus of Coilodera. 21 – 24, C. penicillata (male): 21 & 24, Hua Phan, northeastern Laos; 22, Yunxian, Yunnan; 23, Mount Daweishan, Yunnan. 25 – 28, C. formosana: 25 & 28, Nantou, Taiwan (male); 26, Taitung, Taiwan (female); 27, Taitung, Taiwan (male). 29 – 33, C. nigroscutellaris: 29, Mount Dayaoshan, Guangxi (male); 30, Mount Wuzhishan, Hainan Island (female); 31, Mount Wuyishan, Fujian (female); 32, Mount Shengtangshan, Guangxi (male); 33, Mount Maoershan, Guangxi (male).

FIGURES 49 – 72. Scutellum, mesometasternal process, and male metatibia of Coilodera. 49 – 58, scutellum: 49, C. dives. 50, C. grandimaculata. 51, C. mearesii. 52 – 56, C. penicillata: 52 – 53, Lincang, Yunnan; 54, Gia Lai, central Vietnam; 55 – 56, Mount Daweishan, Yunnan. 57, C. formosana. 58, C. nigroscutellaris. 59 – 66, mesometasternal process. 59, C. dives. 60 – 61, C. grandimaculata: 60, male holotype; 61, female paratype. 62, C. mearesii. 63 – 64, C. penicillata: 63. Hua Phan, northeastern Laos; 64, Jingdong, Yunnan. 65, C. formosana; 66, C. nigroscutellaris. 67 – 72, male metatibia. 67, C. dives. 68, C. grandimaculata. 69, C. mearesii. 70, C. penicillata. 71, C. formosana. 72, C. nigroscutellaris.

FIGURES 73 – 85. Male genitalia and distribution of Coilodera. 73 – 84, Male genitalia: 73 – 74, C. dives. 75 – 76, C. grandimaculata, holotype. 77 – 78, C. mearesii. 79 – 80, C. penicillata. 81 – 82, C. formosana. 83 – 84, C. nigroscutellaris. 85, distribution of Coilodera.

FIGURES 95 – 99. Habitat of Coilodera. 95 – 97, C. mearesii: 95, habitat (Zhangmu, Nyalam, Xizang); 96 – 97, host plant (Hydrangea sp.). 98 – 99, C. penicillata: larva in cockroach galleries of Salganea raggei in rotten wood (Yingjiang, Yunnan).

FIGURES 100 – 106. Habitat of Coilodera. 100 – 102, C. penicillata: 100 – 101, closer view of habitat and newly emerged adult in pupal cell in rotten wood (Tongbiguan Nature Reserve, Yingjiang, Yunnan); 102, newly emerged adult and larva inhabit in rotten wood with Blattodea and Passalidae (Mount Xiaolushan, Lancang, Yunnan). 103 – 106, Host plants of C. nigroscutellaris: 103 – 104. Hydrangea sp. (Mount Dayaoshan, Guangxi, photograph by Lu Qiu); 105 – 106. Ampelopsis sp. (Mount Wuzhishan, Hainan Island).