Ceruchus niger Boucher & Kral 1997

Huang, Hao, Imura, Yûki & Chen, Chang-Chin, 2011, Notes on the genus Ceruchus MacLeay from China, with the description of a new species (Coleoptera: Scarabaeoidea: Lucanidae), Zootaxa 2906, pp. 1-42 : 5-9

4. Ceruchus niger Boucher & Kral 1997

( Figs. 17–18 , 27 , 37–39 , 49 , 60, 65 , 162–193 , 215–218 , 229 , 240, 243–248 , 254 , 264–267 , 284–287 )

Specimens examined. CHINA: Yunnan Province: 83 & 6ƤƤ (CCCC), Lijiang, Laojunshan, 3500m, 13.VI.2009, C.-H. Shan & X.-Y. Zhu leg.; 23 & 1Ƥ (CHH), Lijiang, Yunlong-xueshan, Yunshanping, 3200m, 31.X.2007, B. Liu leg.; 13 (CHH), North Weixi, 3000m, 2.VI.2006, A.-M. Chen leg.; 23 & 1Ƥ (CYI), 14km east of Weixi, 3400m, 5.X.2005, Y. Imura & Y. Nagahata leg.; 13 & 1Ƥ (CHH), Dali City, Binchuan County, Jizushan, 2500m, 17.VII.2010, X.-B. Song leg.; Sichuan Province: 13 & 1Ƥ (CCCC), Muli City, Muli County, Xi-qiu-yakou Pass, 16.VIII.2008, W.-X. Lin leg. (63 & 3ƤƤ from different localities dissected).

Identification. The following male characters were described or illustrated in the original description ( Boucher & Kral 1997): 1) dorsal tooth of the mandible in lateral view with anterior margin and posterior margin nearly equally concave, causing tip of the tooth to not point forwards at all; 2) ventral tooth of the mandible in lateral view (for large and medium-sized males only) present; 3) anterior margin of the mentum noticeably protruding medially; 4) yellow setae on posterior half of the mesofemur in ventral view sparse and moderately long; 5) yellow setae on inner apex of the mesotibia short, narrowly distributed, dense and forming a thin brush; 6) caudal end of the penis not beyond the paramere in natural shape; 7) central split on caudal margin of the basal piece of the aedeagus in ventral view minute, with two sides protruding posteriorly, forming an m-shaped caudal margin; 8) endophallus narrow and short, not longer than half of the penis; 9) penis in ventral view medium in width, wider than in Ceruchus katerinae and Ceruchus deuvei but thinner than in Ceruchus sinensis and Ceruchus minor ; 10) penis in ventral view noticeably swollen in middle; 11) dorsal surface of the penis in lateral view slightly incurved, without wings along the margins. All the specimens examined matched with all these characters.

Only three useful female characters were found using the original description and Araya’s (2002) photographs of the female paratype: 1) anterior angle of the pronotum not an acute angle; 2) flattened marginal area of the pronotum relatively narrower than in Ceruchus minor ; 3) pronotum with a pair of the isolated protuberances outside of the transversal carina. Several female specimens examined were collected together with males from within the same dead trunks at three localities (Laojunshan, Yunshanping, and Muli). They share all the three above-mentioned characters with the female paratype.

Taxonomic notes. The species with the most characters in common with Ceruchus niger are Ceruchus yangi and Ceruchus reginae . Ceruchus yangi possesses nine different characters (one in the male genitalia and four in the female genitalia) from Ceruchus niger .

Male characters li l m d k y r n s t Dorsal tooth of mandible in larger males with both anterior and posterior margins ?? * * nearly equally concave in lateral view

Dorsal tooth of mandible in larger males with anterior margin more concave than? *? * * * * posterior margin in lateral view

Dorsal tooth of mandible in larger males with anterior margin convex *??

Ventral tooth of mandible more or less marked in lateral view * * * * * * * Ventral tooth of mandible entirely absent in lateral view * * * Setae widely distributed and relatively sparse on antennomeres 8–10 * * *? *? * * * Setae hardly seen at base of antennomeres 8 and 9 and relatively dense? *? Anterior margin of mentum noticeably protruding medially * * * Anterior margin of mentum flat or slightly concave or gently convex * * * * * * * Lateral margins of the mentum parallel *? *

Lateral margins of the mentum more or less convergent posteriorly * *? * * * * * Mentum rather smooth, only with a few punctures? *? Mentum coarsely punctate and irregularly sculptured in central part * * *? *? * * * Emargination on anterior margin of head in larger males shallow *?? Emargination on anterior margin of head in larger males a little deep * *? *? * * * Emargination on anterior margin of head in larger males deep? *? Depression on vertex of head extended towards posterior margin of head? *? Depression on vertex of head restricted to anterior half of head and shallow? *? * Depression on vertex of head restricted to anterior half of head and deep * *?? * * Depression on vertex of head nearly absent *

Dorsal surface of head in lateral view strongly convex in larger males *? *?

Dorsal surface of head in lateral view slightly convex in larger males ?? * * * * * Dorsal surface of head in lateral view nearly straight in larger males ?? *

Pronotum markedly wider than head *

Pronotum slightly wider than head * * * * * * * * * Lateral margins of pronotum strongly convex *

Lateral margins of pronotum straight or weakly convex * * * * * * * * * Two pairs of depressions on pronotum clearly marked * * * * * * * * Two pairs of depressions on pronotum obscure or obsolete * * * Depressed longitudinal striations on elytra relatively deep * * * * * * * Depressed longitudinal striations on elytra relatively shallow * * * Second apical tooth of protibia noticeably longer than half width of protibia, sometimes * * * * * equaling width of protibia

Second apical tooth of protibia not longer than half width of protibia * * * * * * * Yellow setae on posterior half of mesofemur dense and long * * * Yellow setae on posterior half of mesofemur dense and short * * * Yellow setae on posterior half of mesofemur sparse and short * * * * Male characters li l m d k y r n s t Yellow setae on inner apex of mesotibia forming a broad brush of long setae along inner * * * margin

Yellow setae on inner apex of mesotibia forming a broad brush of short setae along *

inner margin

Yellow setae on inner apex of mesotibia forming a thin brush of short setae along inner * * * * margin

Yellow setae on inner apex of mesotibia forming a small tuft of short setae confined to * terminal end

Yellow setae on inner apex of mesotibia very sparse, not forming a brush or tuft * Caudal end of penis noticeably protruding beyond parameres in natural shape? *? Caudal end of penis slightly protruding beyond parameres in natural shape *? * *? Caudal end of penis not protruding beyond parameres in natural shape? * * * *? Central split on caudal margin of basal piece of aedeagus in ventral view wide and *

deep, forming a large V-shaped gap

Central split on caudal margin of basal piece of aedeagus in ventral view narrow and * * shallow, forming a small Y-shaped gap, with two sides flat and not protruding

Central split on caudal margin of basal piece of aedeagus in ventral view minute, with * * * two sides protruding posteriorly, forming an m-shaped caudal margin

Central split on caudal margin of basal piece of aedeagus in ventral view minute or * * * * obsolete, with two sides flat and not protruding posteriorly

Caudal margin of paramere strongly depressed inwards just after dorsal caudal corner, ?? * * forming a fold that is visible in dorsal view

Caudal margin of paramere not forming a fold * * *? * *? * Inner sclerite near dorsal margin of paramere well developed ?? *?? Inner sclerite near dorsal margin of paramere weakly developed or obsolete *? *? *? * *? Endophallus wide and long, nearly as long as penis *?? Endophallus narrow and short, not longer than half of penis? * * * * * * *? Penis in ventral view markedly wide * * * * Penis in ventral view slightly wide * * * * Penis in ventral view thin * * Penis in ventral view swollen in middle and narrowed to both ends * * * Penis in ventral view swollen near cephalic end and narrowed to caudal end * * * Penis in ventral view even in width or evenly narrowed from cephalic end to caudal end * * * * Caudal end of penis in ventral view more or less with lateral shoulders * * * * Caudal end of penis in ventral view without lateral shoulders * * * * * * Penis in lateral view relatively wide * *?? Penis in lateral view relatively thin *? * *? * * * Dorsal surface of penis in lateral view slightly incurved, without wings along margin * *? * *? * * * Dorsal surface of penis in lateral view strongly incurved at cephalic half, with marked *?? wings along margin

Female characters li l m d k y r n s t Inner mola of mandible without clear central tooth *?

Inner mola of mandible with single central tooth * * * *? * * * Inner mola of mandible with double central teeth *?

Mentum longer and less transverse *?

Mentum shorter and more transverse * * * * *? * * * Pronotum hardly transverse *?

Pronotum slightly transverse * * * *? * * * Pronotum markedly transverse *?

Flattened marginal area of pronotum relatively wide *?

Flattened marginal area of pronotum relatively narrow * * * * *? * * * Transversal carina on pronotum obsolete or absent *?

Transversal carina on pronotum conspicuous and usually interrupted at sides * * * * *? * * * Pronotum with lateral protuberances outside of transversal carina * *? * * * Pronotum with lateral pairs of depressions outside of transversal carina *

Pronotum smooth outside of transversal carina * * *?

Depressed longitudinal striations on elytra relatively deep * * * * *? *

Depressed longitudinal striations on elytra relatively shallow *? * * Sclerotized part of hemisternite with the inner lateral margin relatively long * *? * * * Sclerotized part of hemisternite with the inner lateral margin relatively short * * * *?

Stylus nearly three times as long as wide *?

Stylus nearly twice as long as wide *?

Stylus nearly as long as wide, with outer lateral corner weakly protruding * * *?

Stylus nearly as long as wide, with outer lateral corner noticeably protruding *? * * * Bursa copulatrix extended well beyond its connection with accessory gland *??? Bursa copulatrix hardly beyond its connection with accessory gland? * * * *? * *? Spermathecal duct not longer than width of hemisternite *? * * *? * *? Spermathecal duct markedly longer than width of hemisternite? *?? Entrance of spermatheca sclerotized? *?? Entrance of spermatheca not sclerotized *? * * *? * *? Spermatheca (straightened) much longer than stylus *? * *? *? Spermatheca (straightened) nearly as long as stylus? * *? *? Spermathecal gland nearly as long as spermatheca? *? *? Spermathecal gland nearly 1.5 – 2.0 times as long as spermatheca? * * *? *? Spermathecal gland more than twice as long as spermatheca *? *?? Spermatheca nearly as wide as hemisternite *?? * *? Spermatheca markedly thinner than hemisternite? * * * *??

Distribution and geographical variation ( Fig. 293 ). This species was described from Zhongdian (type locality), Lijiang and Deqin areas of northwest Yunnan, and was subsequently recorded from south Sichuan ( Araya 2002) and Weixi area of Yunnan ( Imura & Araya 2006). The specimens from the Weixi area examined by Imura & Araya (2006) were said to be somewhat different from “the topotypical specimens” “in having constantly developed apical inner tooth of the male mandibles and differently shaped pronotum with the lateral sides slightly but markedly convergent towards the front angles, which are a little more sharply pointed at the apices”. However, a male from Lijiang ( Fig. 246 ) examined had a more developed apical inner tooth of the mandible than a male from Weixi ( Fig. 247 ). The shape of the pronotum does not vary by locality.

Figures

FIGURE 229. Female genitalia of Ceruchus niger (same specimen as Fig. 264) consisting of sclerotized part in both ventral and dorsal views at top, the last abdominal tergite and ventrite, and the whole female genitalia at bottom. Scale bar 1 mm for all. Abbreviations: mo = median oviduct; ag = accessory gland; sd = spermathecal duct; bc = bursa copulatrix; s = spermatheca; es = entrance of spermatheca; sg = spermathecal gland.

FIGURE 293. Distribution of Ceruchus species in China

FIGURES 10 – 24. Mandible and head of male in lateral view. 10 — Ceruchus lignarius lignarius (same specimen as Fig. 233); 11 — Ceruchus lignarius monticola (same specimen as Fig. 234); 12 — Ceruchus lignarius nodai (same specimen as Fig. 235); 13 — Ceruchus yangi (same specimen as Fig. 237, holotype); 14 — Ceruchus yangi (same specimen as Fig. 236, paratype); 15 — Ceruchus tabanai (holotype); 16 — Ceruchus sinensis (same specimen as Fig. 239); 17 — Ceruchus niger (same specimen as Fig. 240); 18 — Ceruchus niger (same specimen as Fig. 245); 19 — Ceruchus luojishanensis (holotype); 20 — Ceruchus minor (same specimen as Fig. 249); 21 — Ceruchus minor (specimen not figured elsewhere); 22 — Ceruchus minor (same specimen as Fig. 252); 23 — Ceruchus minor (same specimen as Fig. 250); 24 — Ceruchus katerinae (same specimen as Fig. 253). Scale bar 4 mm for all. Abbreviations: d = dorsal tooth; v = ventral tooth.

FIGURES 25 – 28. Antenna of male. 25 — Ceruchus minor; 26 — Ceruchus yangi; 27 — Ceruchus niger; 28 — Ceruchus katerinae.

FIGURES 29 – 45. Mentum of male in ventral view. 29 — Ceruchus lignarius lignarius (same specimen as Fig. 233); 30 — C. luojishanensis (holotype); 31 — C. tabanai (holotype); 32 — C. yangi, Leigongshan (same specimen as Fig. 236, paratype); 33 — C. yangi, Fanjingshan (same specimen as Fig. 237, holotype); 34 — C. yangi, Leigongshan (same specimen as Fig. 242, paratype); 35 — C. sinensis (same specimen as Fig. 238); 36 — C. sinensis (same specimen as Fig. 234); 37 — C. niger, Laojunshan (same specimen as Fig. 240); 38 — C. niger, Yulongxueshan (same specimen as Fig. 245); 39 — C. niger, Laojunshan (same specimen as Fig. 243); 40 — C. minor, Qinling (same specimen as Fig. 249); 41 — C. minor, Dabashan (corresponding to Fig. 250); 42 — C. minor, Diebu of Gansu (specimen not figured elsewhere); 43 — C. minor, Diebu of Gansu (same specimen as Fig. 252); 44 — C. minor, Zhouqu of Gansu (same specimen as Fig. 256); 45 — C. katerinae (same specimen as Fig. 253). Scale bar 3 mm for all.

FIGURES 46 – 53. Mesofemur and mesotibia in ventral view. 46 — Ceruchus katerinae; 47 — Ceruchus lignarius monticola; 48 — Ceruchus minor; 49 — Ceruchus niger; 50 — Ceruchus sinensis; 51 — Ceruchus yangi; 52 — Ceruchus luojishanensis; 53 — Ceruchus tabanai.

FIGURES 54 – 67. Last abdominal tergite and ventrite of males. 54 — Ceruchus yangi, both from Leigongshan; 55 — Ceruchus yangi, both from Fanjingshan; 56 — Ceruchus minor, Qinling; 57 — Ceruchus minor, Dabashan; 58 — Ceruchus minor, Zhouqu of Gansu; 59 — Ceruchus minor, Diebu of Gansu; 60 — Ceruchus niger, Weixi; 61 — Ceruchus sinensis, Baoshan; 62 — Ceruchus sinensis, Baoshan; 63 — Ceruchus lignarius monticola, Japan; 64 — Ceruchus yangi, Leigongshan; 65 — Ceruchus niger, Weixi; 66 — Ceruchus minor, Diebu of Gansu; 67 — Ceruchus sinensis, Baoshan. Scale bar 1 mm for all.

FIGURES 162 – 169. Male genitalia of Ceruchus niger, Laojunshan, Yunnan (same specimen as Fig. 244). 162 — Ninth abdominal segment in ventral view; 163 — Ninth abdominal segment in dorsal view; 164 — whole genitalia in ventral view; 165 — whole genitalia in dorsal view; 166 — whole genitalia in lateral view; 167 — penis detached in lateral view; 168 — penis detached in lateral view; 169 — penis detached in dorsal view. Scale bar 1 mm for all.

FIGURES 170 – 177. Male genitalia of Ceruchus niger, Yunshanping, Yulongxueshan, Yunnan (same specimen as Fig. 245). 170 — Ninth abdominal segment in ventral view; 171 — Ninth abdominal segment in dorsal view; 172 — whole genitalia in ventral view with penis removed; 173 — whole genitalia in dorsal view with penis removed; 174 — whole genitalia in lateral view with penis removed; 175 — penis detached in lateral view; 176 — penis detached in lateral view; 177 — penis detached in dorsal view. Scale bar 1 mm for all.

FIGURES 178 – 185. Male genitalia of Ceruchus niger, Weixi, Yunnan (same specimen as Fig. 254). 178 — Ninth abdominal segment in ventral view; 179 — Ninth abdominal segment in dorsal view; 180 — whole genitalia in ventral view with penis removed; 181 — whole genitalia in dorsal view with penis removed; 182 — whole genitalia in lateral view with penis removed; 183 — penis detached in lateral view; 184 — penis detached in lateral view; 185 — penis detached in dorsal view. Scale bar 1 mm for all.

FIGURES 186 – 193. Male genitalia of Ceruchus niger, Muli, Sichuan (same specimen as Fig. 248). 186 — Ninth abdominal segment in ventral view; 187 — Ninth abdominal segment in dorsal view; 188 — whole genitalia in ventral view; 189 — whole genitalia in dorsal view; 190 — whole genitalia in lateral view; 191 — penis detached in lateral view; 192 — penis detached in ventral view; 193 — penis detached in dorsal view. Scale bar 1 mm for all.

FIGURES 210 – 224. Detached structures of females, consisting of mandibles in dorsal view, mentum in ventral view, pronotum in dorsal view, protibia in dorsal view, and mesofemur in ventral view. 210 — Ceruchus lignarius lignarius (same specimen as Fig. 259); 211 — C. luojishanensis (paratype); 212 — C. tabanai (paratype); 213 — C. yangi (same specimen as Fig. 262); 214 — C. yangi (same specimen as Fig. 263); 215 — C. niger (same specimen as Fig. 264); 216 — C. niger (same specimen as Fig. 265); 217 — C. niger (same specimen as Fig. 266); 218 — C. niger (same specimen as Fig. 267); 219 — C. deuvei (same specimen as Fig. 268); 220 — C. minor (same specimen as Fig. 269); 221 — C. minor (same specimen as Fig. 270); 222 — C. minor (specimen not figured elsewhere); 223 — C. sinensis (same specimen as Fig. 272); 224 — C. katerinae (same specimen as Fig. 273). Abbreviations: md = mandibles; me = mentum; p = pronotum; pt = protibia; mf = mesofemur; l = lateral protuberance; n = no central tooth on mola; d = double central teeth on mola; s = single central tooth on mola; t = transversal carina; f = flattened marginal area of the pronotum.

FIGURES 233 – 248. Habitus of male Ceruchus in dorsal view. 233 — Ceruchus lignarius lignarius (Hokkaido, Japan); 234 — Ceruchus sinensis (Yaojiaping, Lushui, W. Yunnan); 235 — Ceruchus tabanai (holotype); 236 — Ceruchus yangi, paratype (Leigongshan, Guizhou); 237 — Ceruchus yangi, holotype (Fanjingshan, Guizhou); 238 — Ceruchus sinensis (Baoshan, Yunnan); 239 — Ceruchus sinensis (Baoshan, Yunnan); 240 — Ceruchus niger (Laojunshan, Yunnan); 241 — Ceruchus yangi, paratype (Leigongshan); 242 — Ceruchus yangi, paratype (Leigongshan); 243 — Ceruchus niger (Laojunshan, Yunnan); 244 — Ceruchus niger (Laojunshan, Yunnan); 245 — Ceruchus niger (Yulongxueshan, Yunnan); 246 — Ceruchus niger (Yulongxueshan, Yunnan); 247 — Ceruchus niger (Weixi, Yunnan); 248 — Ceruchus niger (Muli, Sichuan). Scale bar 5 mm for all.

FIGURES 249 – 258. Habitus of male Ceruchus in dorsal view. 249 — Ceruchus minor (Pingheliangding, Qinling, Shaanxi); 250 — Ceruchus minor (Dabashan, Chongqing); 251 — Ceruchus minor (Dabashan, Chongqing); 252 — Ceruchus minor (Diebu, Gansu); 253 — Ceruchus katerinae (Hailuogou, Sichuan); 254 — Ceruchus niger (Weixi, Yunnan); 255 — Ceruchus minor (Micangshan, Sichuan); 256 — Ceruchus minor (Zhouqu, Gansu); 257 — Ceruchus luojishanensis (holotype); 258 — Ceruchus minor (Diebu, Gansu). Scale bar 5 mm for all.

FIGURES 259 – 273. Habitus of female Ceruchus in dorsal view. 259 — Ceruchus lignarius lignarius (Hokkaido, Japan); 260 — Ceruchus luojishanensis (paratype); 261 — Ceruchus tabanai (paratype); 262 — Ceruchus yangi, paratype (Leigongshan, Guizhou); 263 — Ceruchus yangi, paratype (Leigongshan, Guizhou); 264 — Ceruchus niger (Laojunshan, Yunnan); 265 — Ceruchus niger (Yulongxueshan, Yunnan); 266 — Ceruchus niger (Weixi, Yunnan); 267 — Ceruchus niger (Muli, Sichuan); 268 — Ceruchus deuvei (Lixian, Sichuan); 269 — Ceruchus minor (Dabashan, Chongqing); 270 — Ceruchus minor (Diebu, Gansu); 271 — Ceruchus minor (Micangshan, Sichuan); 272 — Ceruchus sinensis (Yaojiaping, Lushui, Yunnan); 273 — Ceruchus katerinae (Hailuogou, Sichuan). Scale bar 5 mm for all.

FIGURES 274 – 292. Field observations of Chinese Ceruchus. 274 — habitat of Ceruchus yangi (Mount Leigongshan, 1856 m, 6 September 2009, by X. - D. Yang); 275 — a standing dead tree inhabited by Ceruchus yangi (by X. - D. Yang); 276 – 277 — newly emerged male adult of Ceruchus yangi in pupal cell; 278 — female adult of Ceruchus yangi found in white wood; 279 – 280 — habitat of Ceruchus katerinae (Hailuogou, Sichuan, 3100 m, 30 July 2006, by L. Tang); 281 – 283 — male of Ceruchus katerinae crawling on living stem of Abies tree (by L. Tang); 284 — habitat of Ceruchus niger (Mount Laojunshan, 3500 m, 13 June 2009, by J. - Q. Zhu); 285 – 286 — fallen dead coniferous tree inhabited by Ceruchus niger (by J. - Q. Zhu); 287 — male adult of Ceruchus niger found in the powdery wood (by J. - Q. Zhu); 288 — habitat of Ceruchus minor (Zhouqu, 2340 m, 18 October 2009, by X. - D. Yang); 289 – 290 — a living broadleaf inhabited by Ceruchus minor (by X. - D. Yang); 291 — adult of Ceruchus minor found in white wood of the tree (by X. - D. Yang); 292 — newly emerged male adult of Ceruchus minor in white wood taken from the habitat at Diebu (by H. Huang).