Taxonomy and natural history of the myrmecophilous genus Clinterocera Motschulsky, 1858 (Coleoptera: Scarabaeidae: Cetoniinae) from China and adjacent regions: definition of species group and revision of the C. discipennis species group Author Qiu, Jian-Yue Author Xu, Hao text Zootaxa 2016 4127 1 81 104 journal article 38759 10.11646/zootaxa.4127.1.4 797d978f-8112-41fc-a0dd-11a7a89a8a50 1175-5326 265702 66B1FA32-28B6-4CF5-95D7-6DE777B2C233 Clinterocera discipennis Fairmaire, 1889 ( Figs. 10–42 , 52–57 , 64–70 , 78–83 , 101–105 , 107 ) Clinterocera discipennis Fairmaire, 1889 : 32 ( type locality: Nyenhangli, China [ i.e. , Yuankeng, Wuhua, Guangdong, China ]); Schenkling 1921 : 365 ; Mao 1937 : 1097 ; Medvedev 1964 : 340 ; Ma 1992 : 452 (western Hunan, China ), fig. 1384; Ma 1993a : 286 (Leishan, Guizhou, China ); Ma 1993b : 253 (Mount Longqishan, Jiangle, Fujian, China ), figs. 13; Ma 1995 : 166 , figs. 139a-c, plate V, fig. 29; Sakai & Nagai 1998 : 160 , plate 4, fig. 83 (Mount Yuntaishan, Guizhou, China ); Krajčík 1999 : 40 ; Ma 1999 : 148 , plate; Ma 2001 : 332 (Mount Tianmushan, Zhejiang, China ); Ma 2002b : 382 (Jianyang and Jiangle, Fujian, China ); Ma 2004 : 352 (Mount Shiwandashan, Guangxi, China ) fig. 5b; Li 2004 : 8 (Mount Wuzhishan, Hainan, China ), fig. 1; Smetana 2006 : 299 ; Krajčík 2011 : 70 ; Krajčík 2012b : 72 ; Li & Manola 2016 : 9 . Callynomes rufithorax Moser, 1901 : 530 ( type locality: Tonkin : Montes-Mauson [ i.e. , Mount Mẫu SƠn, Lạng SƠn, Vietnam ]); Schenkling 1921 : 365 . New synonym. Callinomes rufithorax Moser, 1901 : Paulian 1961 : 10 . Clinterocera rufithorax ( Moser, 1901 ) : Sakai & Nagai 1998 : 160 , plate 4, figs. 82-1–2 ; Krajčík 1999 : 40 ; Sakai 2008 : 12 ( Laos ), fig. 6 Ƌ ; Krajčík 2012a : 72 . Callynomes cruciatus Moser, 1901 : 530 ( type locality: Tonkin : Montes-Mauson); Schenkling 1921 : 364 . New synonym. Callinomes rufithorax var. cruciatus Moser, 1901 : Paulian 1961 : 10 . Clinterocera cruciata ( Moser, 1901 ) : Krajčík 1999 : 40 ; Krajčík 2012b : 72 . Callinomes rufiventris Fairmaire, 1904 : 86 ( type locality: Laos ); Paulian 1961 : 10 ; Krajčík 2012b : 72 . New synonym. Callynomes rufiventris Fairmaire, 1904 : Schenkling 1921 : 365 . Clinterocera rufiventris ( Fairmaire, 1904 ) : Krajčík 1999 : 40 . Callynomes vitalisi Bourgoin, 1924 : 138 ( type locality: Laos ). New synonym. Callinomes vitalisi ( Bourgoin, 1924 ) : Paulian 1961 : 9 . Clinterocera vitalisi ( Bourgoin, 1924 ) : Krajčík 1999 : 40 ; Sasaki 2007 : 23 (Guangxi, China ), fig. 2 Ƌ ; Krajčík 2012b : 72 ; Li & Manola 2016 : 11 . Clinterocera rubra Ma, 1992 : 453 ( type locality: Mount Mangshan, Hunan, China ), f. 1385 ♀; Ma 1995 : 167 , fig. 140; Krajčík 1999 : 40 ; Smetana 2006 : 299 ; Krajčík 2011 : 72 ; Krajčík 2012b : 72 ; Li & Manola 2016 : 10 . New synonym. Redescription (male). General: Length: 16.6–21.0 mm; width: 6.4–8.3 mm. Body subelliptical, greatly variable in macula color pattern ( Figs. 21–37 ); dorsal and ventral surface with numerous U-shaped, round, or elliptical setiferous punctures; setae short, acicular, fulvous ( Figs. 64–70 ). Head: Disc orange-red; eye canthus, frontolateral area, and margin black. Dorsal surface densely microsculptured; punctures round or elliptical. Antenna and mouthparts as in Figs. 53–55 and Figs. 78–80 respectively. Gula small, orange-red, both sides with sparse, long setae ( Fig. 52 ). Pronotum: Disc mainly orange-red, subcircular; anterior margin curved, concave, black; a small area in front of scutellum black; each side with a round, blurred, black macula. Scutellum: Triangular, black. Elytra: Black and red; disc flat, about 1.45 times as long as wide, widest at humeral umbone; sutural costa sleek, without punctures; disc matt. Each elytron usually with orange-red maculae. Epimeron and episternum: Mesepimeron and metepimeron black, metepisternum red. Metepimeron clad with a piece of compact testaceous tomentum. Sternum: Postprosternum black; medial portion strongly cuspate, and apex rounded with a few, long setae. Both sides of metasternum orange-red, media portion black; a deep longitudinal black groove in the middle. Abdomen: 7 sternites; 6 sternites visible, glossy, black; a longitudinal groove between abdominal sternites 3–5. Parameres as Figs. 81–83 . Pygidium: Arched, glossy, black; with sparse, large, round punctures. Terminal spiracle slightly convex. Leg: Black. Procoxa, trochanter, and basal portion of femur with small tomentum spot. Mesotibia and metatibia with small, blunt spine near middle of outer margin; protibia with blunt tooth on outer margin, apical tooth of inferior ridge long ( Fig. 56 ), curved downwards. Tarsi with 4 tarsomeres; protarsus tiny. Sexual dimorphism. Female body size similar to male; abdomen evenly convex, sternites 2–6 red; pygidium red; the apical tooth of inferior protibial ridge short, extended distad ( Fig. 57 ). Variation. In both sexes, the shape and size of maculae on pronotum, elytra, and postprosternum vary considerably; color patterns ( Figs. 21, 27–37 ) are presented in the plate as comprehensively as is possible. The predominant patterns are shown in Figs. 21, 27–32 , are found on individuals from various places in southern China , such as north Guangdong (Mount Nanling), north Fujian (Mount Wuyishan), and south Guangxi (Mount Shiwandashan). Individuals ( Figs. 33–34 ) were found in Mount Jinyunshan, Chongqing, and other specimens with black area on pronotum were found in Fujian, Hainan Island, and Laos ( Fig. 35 ). The specimen ( Fig. 36 ) closely resembling the holotype of Clinterocera vitalisi ( Bourgoin, 1924 ) ( Fig. 19 ) was collected at Jingxi, Gunagxi. The entirely black form was found at Mount Nanling, Guangdong and Mount Leigongshan, Guizhou, China ( Fig. 37 ). FIGURES 10–20. Holotypes of Clinterocera species. 10–12, Clinterocera cruciata , holotype, female; 13–15, Clinterocera rufithorax , holotype, male; 16–18, Clinterocera rufiventris , holotype, female; 19–20, Clinterocera vitalisi , holotype, male. Type material examined . Clinterocera rufithorax ( Moser, 1901 ) : holotype (by monotypy): “ Tonkin , Montes Mauson, April Mai 2 -3000, H. Fruhstorfer [printed on yellowed label] / rufithorax Mos. [Moser’s handwriting on yellowed label]” (male, ZMHB , Figs. 13–15 ). Clinterocera cruciata ( Moser, 1901 ) : holotype (by monotypy): “ Tonkin , Montes Mauson, April Mai 2 -3000, H. Fruhstorfer [printed on yellowed label] / cruciatus Mos. [Moser’s handwriting on yellowed label]” (female, ZMHB, Figs. 10–12 ). Clinterocera rufiventris ( Fairmaire, 1904 ) : holotype (by monotypy): “Ht. Tonkin , Dong-Van Cap ne Gadel, 1898 [printed on yellowed label] / Callinomes rufiventris Fair. n. sp. [Fairmaire’s handwriting on yellowed label] / TYPE [printed on red label] / MUSEUM PARIS, 1906, Coll. L. Fairmaire [printed on white label] / HOLOTYPE , Clinterocera rufiventris ( Fairmaire, 1904 ) [printed on white label] / HOLOTYPE [printed on red label] / MNHN EC3847 [printed on white label]” (female, MNHN, Figs. 16–18 ). Clinterocera vitalisi ( Bourgoin, 1924 ) : holotype (by monotypy): “ Callynomes vitalisi Bourgoin Type ♂[handwriting on yellowed label] / Bourgoin Coll., B. M. 1938-252 [printed on yellowed label] / Laos , le, 1918, R. Vitalis de Salvaza [printed on yellowed label] / BMNH(E), 1268980 [printed on white label] / TYPE [printed on yellowed label with red circle] ” (male, BMNH, Figs. 19–20 ). Clinterocera rubra Ma, 1992 : holotype : “Pingkeng, Mt. Mangshan, Hunan, VII.1978 , Shi-Cai Yin leg. [handwritten], Chinese Academy of Sciences [printed] / HOLOTYPE [printed] ” (female, IZAS). FIGURES 21–42. Clinterocera discipennis . 21, dorsal, female (Guangdong); 22, ventral, female (Guangdong); 23, lateral, female (Guangdong); 24, ventral, male (Guangxi); 24, ventral, male (Guangxi); 25, lateral, male (Guangxi); 26, male (Chongqing); 27, female (Fujian); 28, female (Guangxi); 29, female (Chongqing); 30, male (Guangdong); 31, female (Guangxi); 32, male (Guangxi); 33, male (Chongqing); 34, female (Chongqing); 35, female (northeast Laos); 36, male (Guangxi); 37, male (Guangdong). 38–41, newly emerged male (Hunan) and label; 42, Illustration of Clinterocera rubra Ma, 1992 from the original description (Ma 1992). FIGURES 43–51. Clinterocera trimaculata . 43, dorsal, male; 44, ventral, male; 45, lateral, male; 46, ventral, female; 47, lateral, female; 48, male; 49, female; 50, female; 51, illustration of Clinterocera trimaculata Ma, 1993 from the original description (Ma 1993a). Comments on type locality . The type locality of Clinterocera discipennis , Nyenhangli , eg. Yuankengli ( Kffi ffl/ aeffiffl ), or Nyenhang (Yuankeng Kffi/ aeffi ), is a small village (near 23°48'11''N , 115°27'30''E ) in Changbu, Wuhua, Meizhou, Guangdong province, and a chapel and a missionary residence of the Basel Mission was established by missionary Charles Piton in 1866 ( Lutz & Lutz 1995 ; Lamarre 2002 ; Tiedemann 2009 ). Nyenhangli is also the type locality of Pyrestes quinquesignata Fairmaire, 1889 ( Coleoptera : Cerambycidae ), Spilopopillia sexmaculata Kraatz, 1892 ( Coleoptera : Scarabaeidae ), and Coraebus larminati Kubáň, 1995 ( Coleoptera : Buprestidae ) ( Fairmaire 1889 ; Kraatz 1892 ; Kubáň 1995 ). The type locality of Callynomes rufithorax and C. cruciata , Montes-Mauson , is Mount Mẫu SƠn, located in Lạng SƠn, Vietnam , near the border between Guangxi, China and Vietnam . According to the original publication ( Fairmaire 1904 ), the type specimen of Clinterocera rufiventris was collected from border of Laos , which far from the locality (Dong Van, Vietnam ) in the “ type ” labels. Other material examined ( 30♂♂, 33♀♀ ) . CHINA : Fujian: 1♀ ( MTD ), China , Province Fokien, G. Siemssen vend., 1.10.1912 / J. Moser, determ. 1914. / 1931, 28 / Clinterocera rufithorax Mos., Det. Mikšić 1985 ; 1♀ ( QCCC ), 12.V.2013 , campus of Sanming Forestry School, Liang Guo leg.; 1♂ ( QCCC ), 2.VII.2013 , Geshikao ( Castanopsis kawakamii ) Nature Reserve, Xinkou, Sanming, Liang Guo leg.; 1♀ ( QCCC ), 15.VI.2015 , Xianrengu National Forest Park, Sanming, Liang Guo leg.; 2♂♂ ( QCCC ), 3.VI.2015 , Daopaiyan, Youxi County, Sanming, Liang Guo leg.; 1♂ ( QCCC ), 20.VII.2012 , Mount Emeifeng, Taining County, Hai-Tian Song leg.; 1♀ ( NWAU ), 20.IX.1990 , Dazhulan, Mount Wuyishan, Tong Chen leg.; 1♂ ( HBUM , 045460), 23.V.2004 , Dazhulan, Mount Wuyishan, Cai-Xia Yuan & Jing Li leg. Hunan: 1♂ ( HFCS ), 1981, Shaoyang. collector unknown; 2♂ ( HFCS ), VII.1978 , Pingkeng, Mount Mangshan, Shi-Cai Yin leg.; 1♂ ( QCCC ), 30.IV.2016 , near Zhongposhan Forest Park, Hecheng, Huaihua, Tie-Gao Zhu leg. Guangdong: 1♀ ( QCCC ), Mount Wuzhishan, Mount Nanling National Nature Reserve, alt. 400m , 11.IV.2016 , Jun-Qiang Xu leg.; 1♂ ( QCCC ), 16.III.2013 , Qujiang County, Shaoguan, Chao-Wu Zheng leg.; 1♀ ( QCCC ), 25.IX.2012 , Xiaokeng, Qujiang County, Shaoguan, Chao-Wu Zheng leg.; 1♀ ( SCAU ), VI.2009 , Mount Nanling National Nature Reserve, Lei Gao leg.; 3♀ ( HLSC ), 3–4.VI.2009 , Nanling National Nature Reserve, 1200 m , Lei Gao leg.; 1♀ ( QCCC ), 14.VII.2015 , Nanling work station, Mount Nanling National Nature Reserve, Zhao-Yang Tang leg.; 1♀ ( SCAU ), 7.VI.2011 , Mount Nanling National Nature Reserve, Qi Gao leg.; 2♂♂ ( QCCC ), 9.VI.2009 , Mount Wuzhishan, Mount Nanling National Nature Reserve, Chen-Hui Zhan leg.; 1♂ ( QCCC ), V.2007 , Mount Wuzhishan, Mount Nanling, Chen-Hui Zhan leg.; 1♂ ( BPHC ), 2.IV.2002 , Mount Nankunshan, Huizhou, Bao-Ping Huang & Ying Yan leg.; 1♂ , 1♀ ( SYSM , En-398672, En-398679), 5.VI.2001 , Mount Dadongshan, Lianxian County, Zheng-Yao Chen leg.; 1♀ ( SYSM , En-398675), 25.V.1997 , Mount Dadongshan, Lianxian County, Zheng-Yao Chen leg.; 1♀ ( SYSM , En-398674), 29.V.1997 , Mount Dadongshan, Lianxian County, Zheng-Yao Chen leg.; 1♀ ( SYSM , En-398676), 26.V.1997 , Mount Dadongshan, Lianxian, Zheng-Yao Chen leg.; 1♀ ( SYSM , En-398677), 27.V.1997 , Mount Dadongshan, Lianxian, Zheng-Yao Chen leg.; 1♀ ( SYSM , En-398673), 26.V.1997 , Mount Dadongshan, Lianxian County, Jian-Hua Li leg.; 1♂ ( SYSM , En-398681), 23.VI.1993 , Mount Dadongshan, Lianxian County, Zhi-Ming Chen leg.; 1♂ ( SYSM , En- 398682), 5.IX.1992 , Mount Dadongshan, Lianxian County, Jing Wang leg.; 1♀ ( SYSM , En-398671), 14.IX.1993 , Mount Dadongshan, Lianxian, Xu-Sheng Zhong leg.; 1♂ ( SYSM , En-398680), 28.XI.1992 , Sanshimen, Zhuhai, Lei Kuang leg.; Hainan: 1♀ ( NSCJ ), Mount Wuzhishan, 25–30.IV.2007 . Guangxi: 1 6 ( QCCC ), Mount Shengtangshan, Jingxiu County, Laibin, 1000 m , 21.III.2015 , Jing-Teng Zhao leg.; 1♀ ( QCCC ), Mount Shengtangshan, Jingxiu County, Laibin, 1000 m , 4.IV.2015 , Jing-Teng Zhao leg.; 1 6 ( QCCC ), Mount Shengtangshan, Jingxiu County, Laibin, 1200 m , 8.IX.2014 , Jing-Teng Zhao leg.; 1♂ ( NSCJ ), Mount Nonghuaishan, Ande, Jingxi County, Baise, 1–17.VII. 2006 , collector unknown; 1♂ ( QCCC , ex GXNU ), 6.VII.2011 , Mount Guposhan, Hezhou, College students leg.; 1♀ ( QCCC ), VII.2012 , Mount Damingshan, Wuming County, native leg.; 1♀ ( GXIP ), 10.VI.1982 , Timberland, Jinxiu County, Zhu-Yin Wang leg.; 1♂ ( GXIP ), 12.IX.1981 , Gonghe, Jinxiu County, 900 m , Zhu-Yin Wang leg.; 1♀ ( CSCS , C00895), 30.IX.1981 , Hechi, Jie Long leg.; 1♀ (SWU), the pass from Nongli to Tengmao, Renzhuang, Jingxi County, Baise, 690 m , 8.VII.2015 , Jian-Yue Qiu leg. Chongqing: 2♂♂, 3♀♀ ( QCCC ), VI.2009 , Mount Jinyunshan National Nature Reserve, Yangjiagou, Shapingba, Yong Zhou leg. Guizhou: 1♂ ( SYSM , En-398678), 27–28.VII.2001 , Heihewan, Mount Fanjingshan, Hong Pang leg.; 1♀ ( GFGY ), 9.V.1985 , Dongjia , Luodian County, Jing-Hua Zhao, Qun Wang & Yan-Xian Liu leg.; 1♂ ( GFGY ), 19.V.1979 , Zhenyuan County, Jing-Hua Zhao leg.; 1♂ ( HLSC ), 16.IX.2005 , Mount Leigongshan, Fangxiang, 900–1360 m , Jun-Rui Zhi leg.; 1♂ ( KLUC ), 16.VI.2014 , Mount Leigongshan, Leishan County, 1300–2178 m , X.-X. Yang leg. Yunnan: 1♀ ( NSCJ ), Mount Laojunshan, Wenshan, 15–27.VIII.2008 ; 1♂ ( NSCJ ), Mount Daxueshan, Lincang, 1–8.VII.2007 . VIETNAM : 1♀ ( QCCC ), Mount Ngoc Linh, Kon Tum Prov., 1900 m , IV.2016 ; 1♂ ( QCCC ), V.2014 , Ha Giang, native leg. LAOS : 2♀♀ ( QCCC ), VI.2014 , Mount Phan, 2060 m , native leg. Distribution. China : Zhejiang, Fujian, Jiangxi, Guangdong, Chongqing ( new record) , Guizhou, Guangxi, Hunan, Hainan, Yunnan ( new record ); northern Vietnam ; northern Laos . Natural history. Clinterocera discipennis is not very rare, and inhabits various landscapes, such as karst topography, meadows, and bushes in subtropical humid forest ( Figs. 101–104 ). But its natural history is still unknown. Adults are occasionally observed moving on the ground, and some crushed individuals were found near pathway and road. The living female from Jingxi County, Guangxi was transferred to a box within rotten wood in laboratory, but oviposition and feeding behaviors were not observed before it died ( Fig. 105 ). Remarks. Clinterocera discipennis is widely distributed in southern China , northern Vietnam , and northern Laos . Smetana (2006) listed this species from Anhui province, but we know of no specimen with reliable collecting data to confirm this record, perhaps indicating that the record is a misinterpretation of an old locality name. The record from Mount Tianmushan in Zhejiang province reported by Ma (2001) was accepted and presented in the distribution map because this species is usually not difficult to recognize. FIGURES 52–63. Characters of Clinterocera species. 52, Clinterocera discipennis venteral view of head; 53, Clinterocera discipennis antenna; 54, Clinterocera discipennis permentum; 55, Clinterocera discipennis punctures on permentum; 56, Clinterocera discipennis male protibia; 57, Clinterocera discipennis female protibia; 58, Clinterocera trimaculata venteral view of head; 59, Clinterocera trimaculata antenna; 60, Clinterocera trimaculata permentum; 61, Clinterocera trimaculata punctures on permentum; 62, Clinterocera trimaculata male protibia; 63, Clinterocera trimaculata female protibia. Although the holotype for Clinterocera discipennis Fairmaire, 1889 was not found, we still could recognize the elytral pattern of the type specimen similar to Fig. 21 from the original description ( Fairmaire 1889 ). Due to the considerable variation in color patterns on elytra and pronotum, C. discipennis is easily identified as different species based on insufficient specimens. Sakai & Nagai (1998) considered the color patterns in Figs. 27 and 31 as C. discipennis ( Fairmaire, 1889 ) and C. rufithorax ( Moser, 1901 ) , respectively. A specimen with a black band on its pronotum was first described by Sakai (2008) from Laos and he identified it as C. rufithorax ( Moser, 1901 ) ; the pattern similar to Fig. 35 but black on anteapical umbone of elytra, and we a similar specimen from Mount Wuyishan, Fujian. Paulian (1961) was aware of the color variations in this species, and indicated C. cruciata ( Moser, 1901 ) as a variety of C. rufithorax ( Moser, 1901 ) . Ma (1992 ; 1993b ; 1995 ; 2004) also regarded three patterns ( Figs. 21, 27, 31 ) as C. discipennis , although no taxonomic action on this issue was taken in her works. Li (2004) recorded a specimen from Mount Wuzhishan on Hainan Island, with the elytral pattern given in Fig. 28 , and we examined a female similar to Fig.21 from the same locality. A male specimen ( Fig. 36 ) from Jingxi County of Guangxi province was identified as C. vitalisi ( Bourgoin, 1924 ) by Sasaki (2007) , which has the same pattern as the holotype ( Fig. 19 ), but a specimen with a different elytral pattern was collected by author (QIU) from nearby place ( Fig. 105 ). The continuous variations of coloration patterns ( Figs. 21–37 ) are displayed by these specimens occur sympatrically, and this is why we propose regard C. rufithorax ( Moser, 1901 ) , C. cruciata ( Moser, 1901 ) , C. rufiventris ( Fairmaire, 1904 ) , and C. vitalisi ( Bourgoin, 1924 ) as junior synonyms of C. discipennis Fairmaire, 1889 . The mysterious species, Clinterocera rubra Ma, 1992 , was discovered from Mount Mangshan National Nature Reserve in Hunan where nearby Mount Nanling National Nature Reserve of Guangdong ( Ma 1992 ). The female holotype was examined in IZAS in February 2012 ; it resembles C. discipennis , but the body is completely red, and each side of the pronotum has an unclear macula. No more specimens was found since the species was described, hence this rare species is only listed in several catalogues ( Krajčík 1999 , 2011 , 2012a ; Smetana 2006 ). Fortunately, we found two conspecific male specimens in HFCS ( Figs. 38–42 ), bearing the same collecting data as the holotype . Apparently these two specimens were collected together with holotype , but were omitted when Ma (1992) was preparing the Cetoniinae fauna of Hunan province. After careful comparison, we believe that they were collected immediately after emergence from the pupa in rotten wood, since the body of newly emerged beetles is usually red, the integument being flexible, and without maculae. Consequently, Clinterocera rubra Ma, 1992 is also regarded as a junior synonym of C. discipennis ( Fairmaire, 1889 ) .