Coilodera dives ( Gory & Percheron, 1833 )

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 : 516-517

Coilodera dives ( Gory & Percheron, 1833) , revised status

( Figs. 4–9 , 34–41 , 49, 59, 67 , 73–74, 85 , 86–93 )

Macronota dives Gory & Percheron, 1833: 314 (type locality: Himalaya), plate 61, fig. 6 ♀; Burmeister 1842: 321 (synonym of C. penicillata ); Harold 1869: 1298 (in synonymy); Arrow 1910: 44 (in synonymy); Schenking 1921: 133 (in synonymy); Paulian 1960: 13 (in synonymy); Mikšić 1972: 700 (in synonymy); Nakane 1972: 112 (in synonymy); Mikšić 1976: 268 (in synonymy); Krajčík 1998: 88 (in synonymy); Smetana 2006: 306 (in synonymy); Bezděk 2016: 401 (in synonymy). Synonymized with C. penicillata Hope by Westwood 1842: 104.

Coelodera mearsi (nec Westwood): Mikšić 1972: 699 (part, specimens form Assam).

Coelodera miksici Antoine, 1986: 18 (type locality: Shillong, Assam, India), fig. 10a, 10c ♂. New synonym.

Coilodera miksici Antoine : Krajčík 1998: 88; Krajčík 2012: 76.

Diagnosis. Body length 20.6–23.0 mm; width 8.8–10.0 mm. Tomentum on body surface orange-red. The length of male antennal club about twice as the length of antennomeres 2–7 combined. Posterior margin of the median tomentose area of pronotum notched or slightly curved. Scutellum fully clad with tomentum ( Fig. 49 ). Mesometasternal suture deeply engraved with dense, long setae; apex of mesometasternal process slightly curved; base of mesometasternal process distinctly constricted ( Fig. 59 ). Median portion of abdomen usually glabrous ( Figs. 5, 7 , 38 ), at most sparsely clad with a few setae ( Fig. 9 ). Antennal club of male distinctly longer. Outer teeth of metatibia shorter than ventral teeth in male ( Fig. 67 ). Parameres small, basal piece short and flat in lateral view ( Figs. 73–74 ).

This species can be readily separated from C. penicillata by the scutellum completely tomentose (usually partly tomentose in C. penicillata ), the outer teeth of male metatibia shorter (much longer in C. penicillata ), and the basal piece of the parameres flatter (distinctly convex in C. penicillata ).

Type material. Lectotype of Coilodera dives ( Gory & Percheron, 1833) is here designated to fix the namebearing type: dives G. & P. / penicillata . H. / Himalaya [handwritten on pale yellow label with a black border] // Gory / TYPE [red label with a black border] // LECTOTYPUS [purple label] // Lectotype / Coilodera dives / ( Gory & Percheron, 1833) / des. QIU & XU, 2017 [red label] // Maronota dives Gory-Percheron = Coelodera mearsi West., doch hätte der Name dives G. P. die Priorität! Mikšić 1971 [the name dives G. & P. has priority] // MHNG / ENTO / 0 0 0 0 8926 (female, MHNG, Figs. 34–36 ).

Type series of Coilodera miksici Antoine, 1986 : ♂, Chilong / 1906 // Shillong // T. R. Bell. / B. M. 1934-394 / / HOLOTYPE / COELODERA miksici ♂ / Ph. ANTOINE det 85 (Holotype, BMNH, Figs. 37–39 ); ♀, Chilong / 1906 // Shillong // T. R. Bell. / B. M. 1934-394 // ALLOTYPE / COELODERA miksici ♀ / Ph. ANTOINE det 85 (Allotype, BMNH); other paratypes: 1♂ (MNHN), Shillong [brown label] / T. B. Bell. / B. M. 1934-394 / PARATYPE / COELODERA miksici / Ph. ANTOINE det 85 / COELODERA miksici n. sp. / ex coll. B.M. (N.H.) / MNHN / EC / 7043; 1♀ (BMNH), Shillong // Chilong / 1906 // T. R. Bell. / B. M. 1934-394 // PARATYPE, COELODERA miksici / Ph. ANTOINE det 85; 1♂ (BMNH), Silhet / 47.57 // M. penicillata Hope // PARATYPE / COELODERA miksici / Ph. ANTOINE det 85; 1♀ (BMNH), Coll. Nonfried. / Bengalen // Bourgoin Coll. / B. M. 1938-252 // PARATYPE / COELODERA miksici / Ph. ANTOINE det 85; 1♀ (BMNH), Khasia Hills / Assam // Nevinson Coll. / 1918-14 // PARATYPE, COELODERA miksici / Ph. ANTOINE det 85; 1♀ (MNHN), Khas Hills [brown label] / Nevinson Coll. / 1918-14. / PARATYPE / COELODERA miksici / Ph. ANTOINE det 85 / MNHN EC 7043; 1♀ (BMNH, Figs. 40–41 ), Upper Burma: / Seinghku Valley. / 4,500’ / 28.5 N, 97.35 E. / 13.V.1926. F. Keingdon Ward. // Coilodera penicillata Hp. / det. H. Schein 1955 // Brit. Mus. / 1926-400 // PARATYPE / COELODERA miksici / Ph. ANTOINE det 85.

Comments on type material. We considered “Himalaya” as the correct type locality of Coilodera dives , which we wrote on the lectotype label, even though the collecting site of lectotype was cited as the Malabar coast by Gory & Percheron (1833). Kapur (1954) also mentioned that specimen(s) of this genus were from Nilgiri hills, southwestern of India. There are many mislabeled cases of Indian species in 19th century and in the flower beetle collection of MHNG ( Arrow 1910; Prof. Giulio Cuccodoro, personal communication, November 2015), and all species within the species complex are known from the Indo-China subregion ( Fig. 85 ).

Four paratypes (1 male and 2 females from Shillong , northeastern India, and 1 female from Mount Victoria, Myanmar) of Coilodera miksici were not found in BMNH and MNHN . We assume that the depository is probably Antoine’s personal collection as stated in the original publication ( Antoine 1986). The female paratype without locality actually is a Coilodera mearesii (BMNH, Figs. 13–14 ).

O ther material examined (15♂♂, 15♀♀). 1♂ ( BMNH, Bourgoin Coll.), Coll. Nonfried, Hymalaya; INDIA: 2♀♀ ( BMNH, Fry Coll.), eastern India , Manipur // Doherty ; 1♂ ( MFNB), Assam Cachar ; 1♂ ( MFNB), Jindia, Assam ; 2♂♂, 3♀♀ ( MNHN, Oberthür Coll.), Assam Reg. de Shillong ; 1♀ ( MNHN), Assam Reg. de Shillong ; 1♂ ( MNHN), Assam Naga, Aout 1889, W. Doherty ; 2♀♀ ( IRSNB), Shellong Assam ; 1♂ ( IRSNB), Assam Shillong, ex Coll. F. Schneider. CHINA: Yunnan: 1♂ ( QCCC), Tongbiguan National Nature Reserve , Yingjiang County, 1,200 m, Jian-Yue Qiu & Hao Xu, 2nd instar 6.II.2016, adult emerged X.2016 ; 1♂ ( QCCC), 24.I.2017, Tongbiguan National Nature Reserve, Yingjiang County, Jian-Yue Qiu & Hao Xu; Xizang (Tibet) : 4♂♂, 5♀♀ ( QCCC), Gelin, Beibeng , Mêdog County, 1,600 m, Jian-Yue Qiu & Hao Xu, 3nd instar in rotten wood in 13–16.VII.2016, adults emerged IX–XI.2016 ; 1♂ ( QCCC), Gelin, Beibeng , Mêdog County, 1,600 m, Jian-Yue Qiu & Hao Xu, 3nd instar in rotten wood 13–16.VII.2016, adults emerged IV.2017 ; 1♀ ( QCCC), 22.VIII.2017, Gelin, Beibeng, Mêdog County, 1,600 m, Jian-Yue Qiu & Hao Xu. MYANMAR : 1♂ ( QCCC), Chudu Razi , northern Kachin, IX.2006 ; 1♀ ( BMNH), Burma, Mount Victoria, Chinhills 2,400–2,800 m, V.1938, G. Heinrich / Brit. Mus., 1951-337 / Coelodera mearesi Westwood, Ph. ANTOINE det 85.

Distribution. China: Xizang (new record), Yunnan (new record); Myanmar; northeastern India.

Natural history. Coilodera dives inhabits tropical areas at the elevations between 1,500–2,000 m ( Figs. 91– 93 ). This species was reported (species misidentified) as commensal species in the wood-feeding cockroach galleries of Salganea raggei and S. matsumotoi (Blattaria: Blaberidae : Panesthiinae ) in Myanmar ( Kon et al. 2004; Maekawa et al. 2005). We also found several larvae in the galleries of S. raggei , Panesthia agustipennis cognate Bey-Bienko, 1969 (Blattaria: Blaberidae : Panesthiinae ) and Leptaulax sp. ( Coleoptera : Passalidae ) in southeastern Tibet in July 2016 ( Figs. 88–89 ), and then brought them back for breeding under artificial environment at room temperature. Most became adults in pupal cells from September to October ( Fig. 90 ). A female was found visiting flower of Hydrangea sp. (Rosales: Saxifragaceae ) in August 2017 ( Figs. 86–87 ).

Remarks. Coilodera dives has long been regarded as a synonym of C. penicillata ( Westwood 1842; Arrow 1910; Schenking 1921; Paulian 1960; Mikšić 1972, 1976), but it can readily separated by the scutellum, the distal tooth of metatibia, and the parameres; threrfore the species status of C. dives is here reinstated. Mikšić examined the lectotype in 1971 and identified it as C. mearesii ( Figs. 34–36 ), but this was not reported in his taxonomic works on the genus ( Mikšić 1972, 1976). Mikšić (1972) did mentioned that he saw two male specimens from Assam he identified as C. penicillata with orange tomentum and the mesometasternal suture distinct with long setae (similar to C. mearesii ). Later, Antoine (1986) treated them as a new species, Coilodera miksici , without comparing with the type series of C. dives .

Figures

FIGURES 4 – 14. Habitus of Coilodera. 4 – 9, C. dives: 4 – 5, northern Kachin, Myanmar (male); 6 – 7, Mêdog Xizang (female); 8 – 9, Mêdog, Xizang (male). 8, Mêdog, Xizang (male). 10 – 14, C. mearesii (male): 10 – 11, Tiger Hill, northern India; 12, Kurseong, northern India; 13 – 14, female paratype of C. miksici.

FIGURES 34 – 48. Types and labels of Coilodera. 34 – 36, C. dives, female lectotype (photograph by Giulio Cuccodoro). 37 – 41, C. miksici: 37 – 39, male holotype; 40 – 41, female paratype. 42 – 44, C. mearesii, female syntype (provided by Amoret Spooner). 45 – 46, C. formosana, female syntype (photograph by Joachim Willers). 47 – 48, C. nigroscutellaris, male syntype (photograph by Joachim Willers).

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 86 – 90. Habitat of Coilodera dives: 86, habitat (Brahmaputra Valley, Mêdog, Xizang); 87, host plant Hydrangea sp.; 88 – 89, larvae inhabit in rotten wood with Salganea raggei (Blattodea) and Leptaulax sp. (Passalidae); 90, newly emerged adult in artificial environment.

FIGURES 91 – 94. Habitat of Coilodera. 91 – 93, C. dives: 91, habitat (Tongbiguan Nature Reserve, Yingjiang, Yunnan); 92, closer view of habitat; 93, larva in rotten wood. 94, Host plant (Zanthoxylum sp.) of C. grandimaculata in Mêdog, Xizang.