Zootaxa 4079 (3): 345–356 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2016 Magnolia Press ISSN 1175-5334 (online edition) http://doi.org/10.11646/zootaxa.4079.3.2 http://zoobank.org/urn:lsid:zoobank.org:pub:300B9A6C-F5FF-43A4-A8D2-82C9646F367D One new species and one newly recorded species of Cyllodes Erichson from China (Coleoptera: Nitidulidae: )

MEIKE LIU1, XINGKE YANG2 & MIN HUANG1,3 1Key Laboratory of Plant Protection Resources and Pest Management of Ministry of Education, Entomological Museum, Northwest A&F University, Yangling, Shaanxi, 712100, China 2Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China 3Corresponding author. E-mail: [email protected]

Abstract

One new species, Cyllodes quinquemaculatus Liu, Yang & Huang, sp. nov., one newly recorded species, Cyllodes accen- tus Kirejtshuk, 1985, from China and one previously known species, Cyllodes bifascies (Walker, 1859), are described. Photos of their habitus, antenna, mouthparts, and genitalia are provided. A species checklist of Cyllodes from China is presented.

Key words: Polyphaga, Cucujoidea, , morphology, sap

Introduction

The genus Cyllodes Erichson, 1843 belongs to the subfamily Nitidulinae of the family Nitidulidae. Herbst (1792) erected the genus Strongylus for his species Strongylus ater Herbst, 1792. However, Erichson (1843) noted the existence of Strongylus Müller, 1780 and proposed the name Cyllodes Erichson, 1843 to replace Strongylus Herbst, 1792. Sharp (1891) erected Mecyllodes based on the characters of the spacing between meso- and metacoxae and the posterior edge of metaventrite, which differs from the genus Cyllodes. However, Kirejtshuk (2008) considered that these differences were not sufficient to establish a new genus, therefore he treated Mecyllodes as a synonym of Cyllodes and synonymized the species Mecyllodes nigropictus Sharp, 1891 with Cyllodes jucundus Reitter, 1873. In addition, Grouvelle (1892, 1897, 1908, 1914), Nakane & Hisamatsu (1955), Kirejtshuk (1985, 1987, 1990, 2005), and Cline & Skelley (2013) described a large number of new species in the genus Cyllodes. Moreover, Kirejtshuk (1992, 2008) and Jelínek & Audisio (2009) proposed some specific synonymies in this genus. of the genus Cyllodes are mycophagous. Cyllodes indicus Grouvelle, 1894 has been reported on oyster mushrooms from India (Shivaramu et al. 1993). In Gironde, France, Cyllodes ater (Herbst, 1792) has been found on an oak cortical fungus Polyporus on the Leyre river bank (Thomas 2007). In Guangxi Province of China, Cyllodes bifascies (Walker, 1859) has been identified as an important pest on oyster mushroom. The genus Cyllodes has about 60 known species worldwide, in which eight were previously known from China. Herein, three species are studied, of which one new species is described, one is newly recorded for China and one previously known species is redescribed. A species checklist of the genus Cyllodes from China is presented.

Materials and methods

Examined specimens are deposited in the Institute of Zoology of Chinese Academy of Sciences, Beijing, China (IZAS) and the Entomological Museum of Northwest A&F University, Yangling, Shaanxi, China (NWAFU). All measurements were made under an Olympus SZX10 microscope. Photos were made by Leica ZOOM 2000 microscope with an O-image CCD.

Accepted by M. Gimmel: 12 Jan. 2016; published: 12 Feb. 2016 345 Taxonomy

Cyllodes Erichson, 1843

Cyllodes Erichson 1843: 342; Kirejtshuk 2008: 118 (Type species: Strongylus ater Herbst, 1792). Strongylus Herbst 1792: 179 (Type species: Strongylus ater Herbst, 1792). Mecyllodes Sharp 1891: 357 (Type species: Mecyllodes clavicornis Sharp, 1891). Pseudocamptodes Grouvelle 1896: 76 (Type species: Pseudocamptodes africanus Grouvelle, 1896).

Description. Body highly convex dorsally. Dorsal surface smooth and glabrous. Head moderately transverse with surface punctate. Antennae 11-segmented with 3- or 4-segmented loose club. Scape expanded. Pedicel small, shorter than antennomere 3. Antennal grooves distinct, convergent. Labrum short with anterior margin broadly concave with an abbreviated medial notch. Apical edge of mandibles dentate; molar region with large transverse ridges; prostheca with setal brush. Maxillary palpi with three segments, terminal segment elongate and somewhat conical. Labial palpi with terminal segment barrel-shaped. Mentum broad with minute punctures along lateral and posterior margins. Pronotum somewhat transverse, highly convex. Anterior angles evenly rounded; posterior angles rounded to projecting. Anterior margin with deep trapezoidal notch in medial region; lateral margins evenly arcuate to posterior angles; posterior margin almost straight, slightly expanded in medial region. Posterior prosternal wall present and prosternum not prolonged at apex. Mesoventrite oblique and carinate, usually concealed by the meeting of the pro- and metaventrite. Axillary sclerites present on mesoventrite. Metaventrite protuberant, widely separating the middle coxae. Elytral apices rounded separately with well-developed punctate striae. Scutellar shield large and obvious, presenting three kinds of shapes: semicircular, inverted-trapezoidal, subtriangular. Femora enlarged, almost 1.5–2.5 times as wide as corresponding tibiae. Outer apical angle of tibia acute but not spiniform; meso- and metatibia without sublateral line of setae. Anterior tarsi moderately, middle feebly dilated. Tarsomeres 1–3 bilobed and dilated with dense setal pads of ventrally short setae and longer lateral setae. Axillary space present. Claws simple. Male genitalia well sclerotized. Tegmen with an unpaired long lobe ventrally, shallowly excised at apex. Female genitalia with gonocoxites long and apically fused along midline and broadened basally. Styli absent. The genus Cyllodes is similar to the genera Neopallodes Reitter, 1884 and Viettherchnus Kirejtshuk, 1985. From Neopallodes, as well as from most other genera of Cyllodini, Cyllodes differs in having three basal tarsomeres of posterior tarsi dilated to bilobed and dorsoventrally depressed. Viettherchnus differs from Cyllodes in having a modified head capsule and mandibles.

Checklist of known species of the genus Cyllodes from China

1. Cyllodes bifascies (Walker, 1859) Distribution: China (Guangxi, Yunnan, Taiwan), Russian Far East, Japan, Korea. 2. Cyllodes literatus (Reitter, 1878) Distribution: China (Taiwan), Japan. 3. Cyllodes tigrinus Grouvelle, 1914 Distribution: China (Taiwan). 4. Cyllodes multimaculatus Grouvelle, 1914 Distribution: China (Taiwan). 5. Cyllodes punctidorsum Nakane & Hisamatsu, 1955 Distribution: China (Sichuan, Shaanxi), Japan. 6. Cyllodes nakanei Hisamatsu, 1961 Distribution: China (Taiwan), Japan. 7. Cyllodes accentus Kirejtshuk, 1985 Distribution: China (Yunnan) new record, Vietnam. 8. Cyllodes pseudoliteratus Kirejtshuk, 2005 Distribution: China (Taiwan).

346 · Zootaxa 4079 (3) © 2016 Magnolia Press LIU ET AL. 9. Cyllodes scriptum Kirejtshuk, 2005 Distribution: China (Taiwan). 10. Cyllodes quinquemaculatus Liu, Yang & Huang, sp. nov. Distribution: China (Yunnan).

Cyllodes bifascies (Walker, 1859) (Figs. 1–8)

Tritoma bifascies Walker 1859: 259. Chilocorus opponens Walker 1859: 219. Camptodes ornatus Motschulsky 1863: 442. Strongylus notatus Reitter 1873: 129. Strongylus binotatus Reitter 1879: 217. Strongylus dorsalis Reitter 1884: 268. Cyllodes bifascies: Kirejtshuk 1992: 199.

Description. Length 3.2 mm, width 2.4 mm. Body black with reddish-orange spots on lateral posterior region of pronotum and elytra. Surface sculptured on dorsum consisting of minute punctures with microrecticulate interspaces between punctures. Dorsum without setae, entire surface glossy and shining, only pygidium finely pubescent (Figs. 1, 2).

FIGURES 1–2. Cyllodes bifascies (Walker): 1. Male habitus, dorsal view; 2. Male habitus, ventral view. Scale bar = 0.50 mm.

Head with anterior half black and posterior half reddish-orange. Surface punctures irregular, circular or crescent shaped, about 1.5× eye facet width in diameter, approximately 1.5–2.0 puncture diameters apart. Eyes prominent, finely faceted, interfacetal setae absent. Antennal grooves distinctly margined and slightly convergent posteriorly, the smallest distance between them about 1.5× width of mentum. Antenna with funicle yellow and 3- segmented club black, apex of terminal segment yellow. Club about 0.7× length of 1–8 combined segments. Terminal antennomere shorter than other two combined segments. Scape about 2.0 times longer than pedicel. Pedicel small, barrel-shaped, about 0.6× as long as segment 3 (Fig. 3). Apical edge of mandibles with five teeth; molar region with large transverse ridges; prostheca with dense setal brush (Fig. 4). Terminal segment of maxillary

CYLLODES (NITIDULIDAE) FROM CHINA Zootaxa 4079 (3) © 2016 Magnolia Press · 347 palpi elongate, slightly expanded along middle region, longer than 1–2 combined segments; lacinia with long dense brown setal brush apically and anterior half of lateral region (Fig. 5). Terminal segment of labial palpi slightly longer than wide, slightly narrowed apically, nearly conical. Mentum subpentagonal and angulate at medial anterior margin, with minute punctures, interspaces between the punctures smooth.

FIGURES 3–5. Cyllodes bifascies (Walker): 3. Right antenna; 4. Right mandible; 5. Right maxilla. Scale bar = 0.20 mm.

Pronotum somewhat transverse (L:W = 1.0:2.6), widest near posterior angles; punctures similar to punctures on vertex, punctures on anterior and lateral margin denser than other parts, interspaces smooth; sometimes with a few granules along lateral and posterior margin. Scutellar shield semicircular. Prosternum with space between coxae and anterior margin about 1.5× width of procoxal process between coxae. Prosternal process with weak longitudinal carina, expanded posterior to procoxae. Mesoventrite with punctation similar to pronotum, surface granular with some faint microreticulation. Mesoventrite carinate and on more dorsal plane than metaventrite and prosternum. Meso-metaventral junction broadly procurved. Metaventrite transverse (W:L = 1.0:3.0); metaventral

348 · Zootaxa 4079 (3) © 2016 Magnolia Press LIU ET AL. disc sculpturing granular with punctures separated by 1.5–2.0 puncture diameters. Elytra slightly sloping, covering pygidium almost completely with only apex exposed; serially punctate striae visible; sculpturing smooth; punctures similar to those of pronotum. Distance between mesocoxae widest, about 0.8× length of antennal club, 2.0× wider than distance between metacoxae, 4.0× wider than distance between procoxae. All femora canaliculate for reception of tibiae; widest near middle. Profemur 1.5×, mesofemur about 2×, and metafemur 2.5× as wide as corresponding tibiae. Tibia wide and flat (W:L = 1.0:2.0). Protibia slightly narrower than width of antennal club, with distinct stout spines along outer edge and two apical spurs. Mesotibia with two obvious apical spurs; row of short blunt spines along apical margin; stiff seta-like spines along lateral, apical and distal half of inner margin. Metatibia similar to mesotibia, without distinct stout spines along outer edge, as wide as antennal club. Protarsus about 0.75× width of protibia; mesotarsus and metatarsus slightly narrower than corresponding tibia. Claws simple. Tegmen of male genitalia acuminate at apex with shorter setae medio-apically and longer setae laterally. Tegmen slightly elongate (L:W = 2.1:1.0) (Fig. 6). Median lobe with apex slightly protuberant and with paired, widely separated elongate fingerlike projections antero-laterally (Fig. 7). Female genitalia weakly sclerotized, slightly elongate (L:W = 2.3:1.0) (Fig. 8). Notes. Cyllodes bifascies had been inadequately described formerly. Kirejtshuk (1992) moved it from the genus Tritoma to Cyllodes with simple description. Here we redescribe it in detail and provide photos of habitus, antenna, mouthparts, and genitalia. Cyllodes bifascies differs from other members of the genus by possessing the following characters: 1) body with spots on lateral posterior region of pronotum and elytra; 2) apical edge of mandibles with five teeth; 3) prosternal process expanded posteriorly with weak longitudinal carina; 4) mesoventrite carinate, deeply concave for reception of the prosternal process (not be observed easily); 5) apex of male tegmen acuminate, setae not evenly distributed along apical region, shorter medially and longer laterally; 6) male median lobe with two elongate lateral fingerlike projections in apical half, widely separated.

FIGURES 6–8. Cyllodes bifascies (Walker): 6. Male tegmen; 7. Male median lobe of aedeagus; 8. Female ovipositor. Scale bar = 0.10 mm.

CYLLODES (NITIDULIDAE) FROM CHINA Zootaxa 4079 (3) © 2016 Magnolia Press · 349 Material examined. 1♂, China: Yunnan, Xishuangbanna, Damenglong, 650m a.s.l., 10-IV-1958, Shuyong WANG (IZAS); 8♂, 7♀, China: Guangxi, Nanning, 17-I-2015, Ziling LI (NWAFU). Distribution. China (Guangxi, Yunnan, Taiwan), Russian Far East, Japan, Korea, SE Asia.

Cyllodes accentus Kirejtshuk, 1985, new record for China (Figs. 9–16)

Cyllodes accentus Kirejtshuk 1985: 162.

Description. Length 6.6 mm, width 5.1 mm. Body yellowish-brown with three circular black spots on pronotum and three different sizes of black spots on each elytron. Dorsum glossy and shining, surface consisting of dense punctures with distinct microrecticulate interspaces between them (Figs. 9, 10). Head overall yellowish-brown. Surface with circular punctures, about 1.0–1.5× eye facet width in diameter, 1.5–2.0 puncture diameters apart. Eyes slightly prominent, finely faceted, interfacetal setae absent. Antennal grooves distinctly margined and slightly deepened, convergent posteriorly, the shortest distance between them about 2× width of mentum. Antenna with yellowish-brown funicle and 4-segmented club black. Club slightly longer than antennomeres 1–7 combined. Terminal antennomere about half length of other three combined club antennomeres. Scape 4.0× longer than pedicel. Pedicel small, 0.3× as long as antennomere 3 (Fig. 11). Apical edge of mandibles with only two teeth; molar region with large transverse ridges; prostheca with slightly dense setal brush (Fig. 12). Terminal segment of maxillary palpi elongate, nearly 1.5× longer than 1–2 combined segments; lacinia with short dense brown setal brush apically and long dense black setal brush along anterior half of lateral region (Fig. 13). Terminal segment of labial palpi obviously longer than wide, with convex sides and slightly narrowed apically. Mentum nearly square and flattened at anterior margin, with faint minute punctures laterally and posteriorly, interspaces smooth.

FIGURES 9–10. Cyllodes accentus Kirejtshuk: 9. Male habitus, dorsal view; 10. Male habitus, ventral view. Scale bar = 1.00 mm.

Pronotum transverse (L:W = 1.0:2.8), widest near posterior angles; punctures similar to circular punctures on vertex, punctures on posterior half denser than on anterior half, interspaces smooth to finely microreticulate along lateral and posterior margin. Scutellar shield inverted-trapezoidal with posterior margin truncate. Prosternum with space between coxae and anterior margin about 2× width of procoxal process between coxae. Prosternal process slightly expanded posterior to procoxae. Mesoventrite with punctation slightly smaller than on pronotum, surface

350 · Zootaxa 4079 (3) © 2016 Magnolia Press LIU ET AL. sculpturing granular with some faint microreticulation. Mesoventrite distinctly carinate, on slightly more dorsal plane than metaventrite and prosternum; not or only slightly obscured by prosternal process. Meso-metaventral junction distinctly procurved. Metaventrite transverse (W:L = 1.0:2.4); metaventral disc sculpturing granular with dense punctures, space between them about 1–1.5 puncture diameters. Elytral margins parallel in the anterior half, then curved in the posterior half. Elytra covering pygidium completely; some serially punctate striae distinct; sculpturing smooth to slightly granular; punctures similar to those of pronotum but slightly less dense.

FIGURES 11–13. Cyllodes accentus Kirejtshuk: 11. Right antenna; 12. Right mandible; 13. Right maxilla. Scale bar = 0.20 mm.

CYLLODES (NITIDULIDAE) FROM CHINA Zootaxa 4079 (3) © 2016 Magnolia Press · 351 FIGURES 14–16. Cyllodes accentus Kirejtshuk: 14. Male tegmen; 15. Male median lobe of aedeagus; 16. Male internal sac sclerites. Scale bar = 0.20 mm.

Distance between mesocoxae widest, about 0.5× length of antennal club, 1.5× wider than distance between metacoxae, 2.5× wider than distance between procoxae. Each femur canaliculate for reception of tibiae; widest near middle. Profemur 2.5×, mesofemur and metafemur about 2× width of corresponding tibia. Tibia slender and flat (W:L = 1.0:3.5). Protibia slightly narrower than width of antennal club, with distinct stout spines along outer edge and two unequal apical spurs. Mesotibia with row of short blunt spines along apical margin; with two obvious apical spurs; stiff seta-like spines along lateral, apical and distal half of inner margin. Metatibia similar to mesotibia, without distinct stout spines along outer edge, as wide as antennal club. Tarsi slightly narrower than corresponding tibiae. Claws simple. Tegmen of male genitalia elongate (L:W = 2.8:1.0) with apex narrowly rounded and with dense and long setae around apical margin (Fig. 14). Median lobe with apex slightly convex, near middle of lobe with pair of claw-like projections (Fig. 15). Internal sac sclerites with paired ejaculatory rods, proximal to and around the ejaculatory rods with large fields of callosities, each end of ejaculatory rod divergent (Fig. 16). Notes. Cyllodes accentus had been reported from Vietnam. Here, we first report it from Yunnan in China. It differs from other members of this genus by possessing the following characters: 1) body with three circular black spots on pronotum and three different sizes of black spots on each elytron; 2) antennal club comprised of 4 antennomeres while other Cyllodes with only a 3-segmented club; 3) apical edge of mandibles with only two teeth; 4) maxillary lacinia with dense black setal brush along anterior half of the lateral region while other species of Cyllodes with brown setal brush; 5) scutellar shield inverted-trapezoidal with posterior margin truncate; 6) mentum nearly square; 7) mesoventrite distinctly carinate, on slightly more dorsal plane than metaventrite and prosternum; 8) male genitalia with median lobe with pair of claw-like projections near middle part.

352 · Zootaxa 4079 (3) © 2016 Magnolia Press LIU ET AL. Material examined. 1♂, China: Yunnan, Xishuangbanna, Mengzhe, 870m a.s.l., 08-VI-1958, Fuji PU (IZAS); 1♂, China: Yunnan, Xishuangbanna, Mengzhe, 875m a.s.l., 01-VII-1958, Fuji PU (IZAS). Distribution. China (Yunnan), Vietnam.

Cyllodes quinquemaculatus sp. n. (Figs. 17–24)

Description. Length 3.7 mm, width 2.9 mm. Pronotum reddish-brown with three black spots; elytra black with two reddish-brown spots. Dorsum glossy and shining, surface sculpture consisting of minute punctures with microreticulate interspaces (Figs. 17, 18). Head with anterior half black and posterior half reddish-brown. Surface punctures irregular, circular or crescent shaped, about 1.0–1.5× eye facet width in diameter, 1.5 puncture diameters apart. Eyes prominent, finely faceted, interfacetal setae absent. Antennal grooves distinctly margined and slightly deepened, slightly convergent posteriorly, the shortest distance between them about 2× width of mentum. Antenna with funicle reddish-brown and 3-segmented club black. Club about 0.8× length of remaining antennomeres combined. Scape 4.0× longer than pedicel. Pedicel small, 0.7× as long as segment 3 (Fig. 19). Apical edge of mandibles with four teeth; molar region with large transverse ridges; prostheca with dense setal brush (Fig. 20). Terminal segment of maxillary palpi elongate, nearly 2× longer than remaining palpomeres combined; lacinia with short dense brown setal brush apically and along anterior 1/3 laterally (Fig. 21). Terminal segment of labial palpi longer than wide, slightly narrowed apically. Mentum subpentagonal and angulate at medial anterior margin, with only a few faint minute punctures along posteriorly, interspaces smooth.

FIGURES 17–18. Cyllodes quinquemaculatus sp. n.: 17. Male habitus, dorsal view; 18. Male habitus, ventral view. Scale bar = 0.50 mm.

CYLLODES (NITIDULIDAE) FROM CHINA Zootaxa 4079 (3) © 2016 Magnolia Press · 353 FIGURES 19–21. Cyllodes quinquemaculatus sp. n.: 19. Right antenna; 20. Right mandible; 21. Right maxilla. Scale bar = 0.20 mm.

Pronotum somewhat transverse (L:W = 1.0:2.4), widest near posterior angles; punctures slightly smaller than those on vertex, surface sculpture smooth and slightly granular laterally and posteriorly. Scutellar shield semicircular. Prosternum with space between coxae and anterior margin about 1.5× width of procoxal process between coxae. Prosternal process slightly expanded posterior to procoxae, with a longitudinal carina. Mesoventrite with punctation similar to that of pronotum, surface sculpturing granular with some faint microreticulation. Mesoventrite distinctly carinate and on more dorsal plane than metaventrite and prosternum; often obscured by prosternal process. Meso-metaventral junction distinctly procurved. Metaventrite overall transverse (W:L = 1.0:3.0); metaventral disc sculpturing granular with small punctures 2.5× their diameter apart. Elytra steeply sloping, covering pygidium completely; some serially punctate striae visible but other punctures distributed randomly; sculpturing granular to smooth, glabrous; punctures similar to those of pronotum while interspaces between them slightly wider than on pronotum.

354 · Zootaxa 4079 (3) © 2016 Magnolia Press LIU ET AL. FIGURES 22–24. Cyllodes quinquemaculatus sp. n.: 22. Male tegmen; 23. Male median lobe of aedeagus; 24. Female ovipositor. Scale bar = 0.10 mm.

Distance between mesocoxae widest, 0.4× length of antennal club, slightly wider than distance between metacoxae, 3.0× wider than distance between procoxae. Each femora canaliculate for reception of tibiae; widest near middle. Profemur 2.5×, mesofemur 1.5×, and metafemur about 2× width of corresponding tibia. Tibia wide and flat (W:L = 1.0:2.0). Protibia with distinct stout spines along outer edge and two unequal apical spurs, as wide as antennal club. Mesotibia with row of short blunt spines along apical margin and two obvious apical spurs; thick stiff seta-like spines along lateral, apical and distal 2/3 of inner margin. Metatibial armature similar to that of mesotibia, slightly wider than protibia. Protarsus 0.8× width of protibia; mesotarsus as wide as mesotibia; metatarsus slightly narrower than metatibia. Claws simple. Tegmen of male genitalia short and broad (L:W = 1.5:1.0) with apex rounded, setae dense and long around apical margin, with two distinct apico-lateral tufts present (Fig. 22). Median lobe with apex slightly protuberant and with paired elongate fingerlike projections near middle part (Fig. 23). Female genitalia weakly sclerotized, as observed in other Cyllodes species, with gonocoxites fused along midline, broadly widened at base (L:W = 1.2:1.0) (Fig. 24). Diagnosis. The new species is similar to Cyllodes bifascies, but can be distinguished the latter by the following characters: 1) three spots on pronotum and two spots on elytra; 2) apical edge of mandibles present four dentate cuttings; 3) mesoventrite deeply concave for reception of the prosternal process; 4) the ratio of the length of scape and pedicel is 4.0:1.0, larger than other species of Cyllodes; 5) male genitalia with tegmen rounded apically and median lobe present two obvious elongate lateral fingerlike projections touched to each other near middle part. Material examined. Holotype. ♂, China: Yunnan, Xishuangbanna, Jinghong, Xiaomengyang, 850m a.s.l., 03- V-1983, Панфилов (IZAS). Paratypes. (1♂, 1♀). 1♂, China: Yunnan, Xishuangbanna, Menga, 1050–1080m a.s.l., 20-VIII-1958, Shuyong WANG (IZAS); 1♀, China: Yunnan, Xishuangbanna, Damenglong, 650m a.s.l., 17- IV-1983, Fuji PU (IZAS). Etymology. The specific epithet is derived from the combined Latin words “quinque” and “maculatus” referring to the five spots on its body. Distribution. China (Yunnan).

CYLLODES (NITIDULIDAE) FROM CHINA Zootaxa 4079 (3) © 2016 Magnolia Press · 355 Acknowledgements

We are grateful to Dr. X. Yang, Dr. S. Ge and Dr. H. Xue (Chinese Academy of Sciences) who provided us with great assistance and to Dr. P. Audisio (Sapienza Università di Roma) and Dr. J.R. Schrock (Emporia State University) for reviewing the manuscript. We also appreciate Dr. A.G. Kirejtshuk ( Systematics Zoological Institute, Russian Academy of Sciences) for providing us with important literature. We also greatly appreciate the valuable comments on an earlier version of this manuscript received from anonymous reviewers. This study was supported by the Fundamental Research Funds for Chinese Central Universities (Z109021305).

References

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