The genus Anaedus Blanchard, 1842 in China with description of two new species (Coleoptera: Tenebrionidae: Goniaderini)

The genus Anaedus Blanchard, 1842 in China with description of two new species (Coleoptera: Tenebrionidae: Goniaderini)

Journal of Asia-Pacific Entomology 23 (2020) 91–97 Contents lists available at ScienceDirect Journal of Asia-Pacific Entomology journal homepage: www...

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Journal of Asia-Pacific Entomology 23 (2020) 91–97

Contents lists available at ScienceDirect

Journal of Asia-Pacific Entomology journal homepage: www.elsevier.com/locate/jape

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The genus Anaedus Blanchard, 1842 in China with description of two new species (Coleoptera: Tenebrionidae: Goniaderini) Zhonghua Wei, Guodong Ren

T



The Key Laboratory of Zoological Systematics and Application, College of Life Sciences, Hebei University, 071002 Baoding, Hebei Province, China

A R T I C LE I N FO

A B S T R A C T

Keywords: Darkling beetles Lagriinae Taxonomy Palaearctic Oriental

The genus Anaedus Blanchard, 1842 from China is revised. Anaedus serratus sp. nov. (Yunnan) and Anaedus nonedgus sp. nov. (Anhui, Hubei, Zhejiang) are described and illustrated. In addition to, a key to Chinese Anaedus species is given, and their geographical locations have been mapped. www.zoobank.org/urn:lsid:zoobank.org:pub:479055B1-08F6-4A35-8D45-999D46B0AEC9

Introduction Anaedus Blanchard, 1842 is distributed in the tropical and subtropical regions of Asia, and the tropical Africa, North and South America. A total of 81 species have been described by Champion (1886, 1892), Pic (1917a, b, c, 1921, 1922, 1938), Ardoin, 1973, 1976, Kaszab, 1973, 1979, Makhan, 2013, 2017; Makhan and Saeizad, 2013; Schawaller, 1994; Wang and Ren, 2007; Ardoin, 1969; Gebien, 1911; Gebien, 1920; Makhan, 2011. Ardoin (1969) synonymised two species Anaedus diversithorax Pic (1917a, b, c) and Anaedus niger Pic, 1932. Furthermore, Kaszab (1979, 1983) transferred Luprops marginicollis Fairmaire, 1896 to Anaedus and Anaedus orientalis Motschulsky, 1868 to the genus Luprops Hope, 1833. Similarly, Schawaller (2007) transferred Anaedus andohahelae Ferrer, 1998 to the genus Antennoluprops Schawaller, 2007. Arndt (1993) described the larva of A. camerunus Gebien, 1920. Ferrer (1998), Schawaller (2011) and Ferrer et al. (2016) placed Anaedus into the tribe Lupropini Ardoin (1958). Aalbu et al. (2017) constructed the phylogeny Lagriinae assigning, Anaedus to the tribe Goniaderini. In this paper, we followed this last systematic placement. Nine species of Anaedus have been recorded in China by Pic (1938) 1 species; Kaszab (1968, 1973) 2 species; Wang and Ren, 2007 4 species; Makhan and Saeizad (2013) 2 species. Most species are found in south China, except Anaedus mroczkowskii Kaszab, 1968 which was recorded in the Liaoning Province. Recent study of the Lagriinae specimens from China in the HBUM and IZCAS collection allowed the identification of two species new to science, which are here described and illustrated: A. serratus sp. nov. (Yunnan) and A. nonedgus sp. nov. (Anhui, Hubei, Zhejiang). A new identification key to Chinese Anaedus ⁎

is provided in this paper. It does not include A. aschnae Makhan, 2013 (possibly belonging to the genus Sphingocorse, and widely distributed in the Chongqing, Henan, Sichuan, Yunnan Province of China) and A. maryame Makhan, 2013 since the validity of these two species needs further confirmation and the authors did not revised the types. The geographical locations of the Chinese Anaedus species are mapped (Fig. 51). Materials and methods All specimens were examined, illustrated, photographed and measured using a Leica M205A stereomicroscope equipped with a drawing tube and Leica DFC450 camera. All measurements are in millimetres. The following abbreviations used in the text represent the studied collections. DMPC

Dewanand Makhan private collection, Nieuwegein, Netherlands

HBUM HNHM IZCAS SNUC

Hebei University Museum, Baoding, China Hungarian Natural History Museum, Budapest, Hungary Institute of Zoology, Chinese Academy of Sciences, Beijing, China Shanghai Normal University, Shanghai, China

Taxonomy Genus Anaedus Blanchard, 1842 Aspisoma Duponchel and Chevrolat, 1841: 210. Type species: Aspisoma fulvipenne Duponchel and Chevrolat, 1841. Anaedus Blanchard, 1842: pl. 14. Type species: Anaedus

Corresponding author. E-mail address: [email protected] (G. Ren).

https://doi.org/10.1016/j.aspen.2019.10.010 Received 6 March 2019; Received in revised form 26 September 2019; Accepted 17 October 2019 1226-8615/ © 2019 Korean Society of Applied Entomology. Published by Elsevier B.V. All rights reserved.

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margin of pronotum widely arcuate in middle A. basilatilus

punctatissimus Blanchard, 1842, monotypy. Synonymy: Lacordaire (1859: 396). Anaedes Agassiz, 1846: 36. Unjustifed emendation of Anaedus Blanchard, 1842, not in prevailing usage. Aspidosoma Agassiz, 1846: 36. Unjustifed emendation of Aspisoma Duponchel and Chevrolat, 1841, not in prevailing usage. Diagnosis. Body flat, brown to black. Clypeus convex; frontoclypeal suture distinctly deep; frons with irregular punctures. Antennomeres cylindrical. Pronotum with irregular punctures; anterior margin emarginate; lateral margins in most species flat and laterally expanded (expect A. nonedgus sp. nov., A. striatus Gebien, 1921 and some North American species); posterior angles acute. Elytral base slightly wider than pronotal base. Elytral surface with irregular punctures, lateral margins serrated at the base. Basal metatarsus elongated, longer than the following tarsomeres combined together. In the tribe Goniderini, the genera Pseudolyprops and Anaedus are distinguishable from other goniderine genera by the following caharcters: body wide, flat, pubesent; head prognathous; antenna filform, antennomere IX to XI not enlarged; tentorium with the transverse bridge present; pronotum strongly transverse, lateral margins of pronotum simple, often laterally expanded; lateral margins of elytra finely serrate in basal portion; disc of pronotum and elytra not sculptured; tibiae simple; basal metatarsus distinctly elongated. However, The Anaedus species can be separated from Pseudolyprops species using the following characters: body larger; anterior margin of pronotum distinctly emarginated; lateral margins of pronotum smooth and laterally expanded; basal metatarsus distinctly longer than the following tarsomeres combined together; coxite lobe IV of ovipositor much shorter than that of Pseudolyprops.

1 Anaedus basilatilus Wang & Ren, 2007 (Figs. 1–5) Anaedus basilatilus Wang & Ren, 2007: 37. Type locality: Guandu, Chishui, Guizhou, China. Type depository: HBUM. Specimens examined. Holotype: ♂, China, Guizhou, Chishui, Guandu, 23.IX.2000, Guodong Ren leg., HBUM. Paratype: 1♂, China, Guizhou, Libo, 16.VIII.2000, Fuming Shi leg., HBUM; 2♀, China, Guizhou, Xishui, Dabaitang, 25–29.IX.2000, Guodong Ren leg., HBUM. Distribution. China (Guizhou). 2 Anaedus maryame Makhan, 2013 Anaedus maryame Makhan, 2013: 2. Type locality: Jinyun mountains, China. Type depository: DMPC. Specimens examined. 1♀, China, Anhui, Yuexi, Yaoluoping, 17–21.VII.2007, Yibin Ba, Juntong Lang and Fengyan Wang leg., HBUM; ♀, China, Hubei, Tianluo, Qingtaiguan, 31.18941°N, 115.69388°E, alt. 602 m, 26.VI.2014, Yingbin Ba & Shenyan Tang leg., HBUM; ♀, China, Hubei, Yingshan, Wujiashan, 16–19.VII.2013, Caixia Yuan, Ying Tian & Ling Bai leg., HBUM; 1♀, China, Hubei, Longtangu, 31.08283°N, 115.81796°E, alt. 314 m, 24.VI.2014, Guodong Ren leg., HBUM. Distribution. China (Anhui, Chongqing, Hubei). Comments. The holotype of A. maryame Makhan, 2013 is a female. The females of A. mroczkowskii Kaszab, 1968 from Anhui and Hubei are similar to those of A. maryame Makhan, 2013. We retain that A. maryame Makhan, 2013 and A. mroczkowskii Kaszab, 1968 could be potentially synonyms, but since our deduction is based only on photos and original description and not type revision we prefer to remand this conclusion to another paper. 3 Anaedus mroczkowskii Kaszab, 1968 (Figs. 6–15) Anaedus mroczkowskii Kaszab, 1968: 10; Wang & Ren, 2007: 34. Type locality: Phjǒngan-namdo, Korea. Type depository: HNHM. Specimens examined. 1♂1♀, China, Hubei, Tianluo, Qingtaiguan, 31.18941°N, 115.69388°E, alt. 602 m, 26.VI.2014, Yingbin Ba & Shenyan Tang leg., HBUM; 1♂5♀, China, Hubei, Yingshan, Wujiashan, 16–19.VII.2013, Caixia Yuan, Ying Tian & Ling Bai leg., HBUM; 2♀, Hubei, Yingshan, Taohuachong, 11–15.VII.2013, Caixia Yuan, Ying Tian & Ling Bai leg., HBUM; 2♂, China, Henan, Tongbaishan, 25.VII.2010, Yibing Ba & Zhenxing Zhang leg., HBUM; 3♀, China, Guizhou, Libo, Maolan, alt. 750–850 m, 20.VII.2015, Jiayaohu Hu leg., HBUM; 1♀, China, Henan, Luoshan, Dongzhai, 26.VI.1999, Yao Niu leg., HBUM; 1♀, China, Hubei, Longtangu, 31.08283°N, 115.81796°E, alt. 314 m, 24.VI.2014, Guodong Ren leg., HBUM; 1♂, China, Liaoning, Qianshan, alt. 360 m, 26.VI.1990, Guodong Ren leg., HBUM; 2♂, China, Hainan, Ledong, Jianfengling, 21.V.2007, Yibin Ba & Langjun Tong leg., HBUM; 1♀, China, Shaanxi, Ningqiang, Qingmuchuan, alt. 1600–1900 m, 11–13.VII.2010, Mingyi Tian leg., HBUM. Distribution. China (Guizhou, Hainan, Henan, Hubei, Liaoning, Shaanxi); South Korea; North Korea. 4 Anaedus pluridentatus Wang & Ren, 2007 (Figs. 16–23) Anaedus basilatilus Wang & Ren, 2007: 39. Type locality: Jinyun Mountains, Chongqing, China. Type depository: HBUM. Specimens examined. Holotype: ♀, China, Chongqing, Jinyun Mountain, 8.VII.2002, Ming Bai & Fengyan Wang leg., HBUM. 2♂2♀, China, Guizhou, Libo, Maolan, alt.750 m, 19.VII.2015, Cheng & Zhao leg., HBUM; 2♀, China, Guizhou, Libo, Maolan, Wengang, alt. 800 m, 22.VII.2015, Jiayao Hu leg., HBUM. Distribution. China (Chongqing, Guizhou). Comments. The male of A. pluridentatus was firstly found in Guizhou Province from China. This species does not show sexual dimorphism. 5 Anaedus serratus sp. nov. (Figs. 24–28) Type material. Holotype: ♂, China, Yunnan, Jinghong, Jinghong farm, 22.10607°N, 100.68500°E, alt. 759 m, 6.IV.2009, Lingzeng Meng Leg., IZCAS. Paratype: ♀, same data as holotype, HBUM. Diagnosis. This new species can be easily separated from the other

Key to known species of Anaedus from China 1 Lateral margins of pronotum serrate, aedeagal apex truncated A. serratus sp. nov. – Lateral margins of pronotum not serrate, aedeagal apex not truncated 2 2 Lateral margins of pronotum not expanded laterally (Fig. 43) A. nonedgus sp. nov. – Lateral margins of pronotum distinctly expanded laterally (Figs. 3, 8, 14, 17, 21, 26, 31, 37, 48) 3 3 Lateral margins of elytra serrated through most of their length A. pluridentatus – Lateral margins of elytra serrated in the basal fourth 4 4 Legs sexually dimorphic, meso- and metatibiae serrated in the apical half in males (Figs. 9, 32, 38); dorsal integuments covered with short and sparse setae 5 – Legs without sexual dimorphism, meso- and metatibiae not serrated in the apical half in both males and females; dorsal integuments covered with long and dense setae 7 5 Antennae without sexual dimorphism; flattened lateral parts of pronotum somewhat upturned A. tibiodentatus – Antennae with sexual dimorphism, males antennomere V elongated and spiny; flattened lateral parts of pronotum not upturned 6 6 Base of scutellum with punctures; mesotibiae slightly excavated near apex on inner sides, antennomere V large and almost lamellate (Fig. 30) A. spinicornis – Base of scutellum without punctures; mesotibiae not excavated near apex at inner sides, antennomere V small and slightly thick (Fig. 7) A. mroczkowskii 7 Lateral margin of pronotum bearing one small tooth 8 – Lateral margin of pronotum without any tooth A. substriatus 8 The vertical posterior surface of prothorax with a pair of teeth which bent downwards, basal margin of pronotum comparatively narrowly arcuate in middle A. unidentatus – The vertical posterior surface of prothorax without teeth, basal 92

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Figs. 1–18. Anaedus species. 1–5. A. basilatilus Wang and Ren, 2007, paratype, male, 1. Habitus, in dorsal view, 2. Antennae, 3. Pronotum, 4–5. Aedeagus, in dorsal and lateral views. 6–15. A. mroczkowskii Kaszab, 1968, male and female, 6. Habitus, male, dorsal view, 7. Antennae, male, 8. Pronotum, male, 9. Mesotibia, male, 10–11, Aedeagus, in dorsal and lateral views; 12. Habitus, female, 13. Antennae, female, 14. Pronotum, female, 15, Mesotibia, female. 16–18, A. pluridentatus Wang and Ren, 2007, female, holotype; 16. Habitus, in dorsal view, 17. Pronotum, 18. Lateral margin of right elytron.

genae strongly raised, without punctures; frons covered with coarse punctures, larger medially. Eyes reniform, transverse, slightly divided by genal canthus; eye diameter shorter than length of temple in dorsal view; width at narrowest point of eye equal to 3–4 ommatidia in lateral view. Antennae (Fig. 25) robust, extending to the base of pronotum if bent backwards, terminal antennomere 1.4 times longer than width, ratio of length of antennomeres II–XI as follows: 9.6: 17.8: 22.6: 22.6: 22.6: 22.6: 23.9: 22.6: 24.2: 37.7. Pronotum (Fig. 26) strongly transverse, 1.8 times as wide as long, widest in middle; anterior margin shallowly emarginated, wide and straight in middle; lateral margins flat, serrated and unevenly expanded laterally, strongly convergent in apical half, and strongly sinuous before posterior angles; posterior margin gently arcuate in middle, posteriorly produced; anterior angles acute; posterior angles acute and pointing outwards. Pronotal surface between lateral margins distinctly convex, with moderately dense and deep punctures on disc, and longitudinal carina in middle. Prothoracic hypomere glossy, with punctuations

Chinese species on account of its distinctly serrated lateral margins of pronotum, while the other species present smooth or toothed lateral margins (A. unidentatus Wang and Ren, 2007 with one tooth on each lateral margin of pronotum). A. serratus sp. nov. resembles A. buricus Schawaller, 1994 due to the serrated lateral margins of pronotum, but the new species differs from the later in the following characters: (1) basal margin of the pronotum without teeth, (2) male anterior tibiae without oval excavation on inner side. Etymology. The specific name is derived from “serrate”, referring to the serrate lateral margins of pronotum. Description. Body (Fig. 24) elongate-oval, flat; head and pronotum black, elytra brown, antennae and legs blackish brown; body dorsum covered with dense and long setae, erect on head and pronotum, while those on elytra and abdomen posteriorly recumbent. Male. Labrum transverse, transversely produced in the middle, with coarse punctures; clypeus strongly convex, anterior margin straight, surface with coarse punctures; frontoclypeal suture straight and deep; 93

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Figs. 19–40. Anaedus species. 19–23. A. pluridentatus Wang and Ren, 2007, male; 19. Habitus, in dorsal view, 20. Antennae, 21. Pronotum, 22–23. Aedeagus, in dorsal and lateral views. 24–28. Anaedus serratus sp. nov., male, holotype, 24. Habitus, in dorsal view, 25. Antennae, 26. Pronotum, 27–28. Aedeagus, in dorsal and lateral views. 29–34. Anaedus spinicornis Kaszab, 1973, male, 29. Habitus, in dorsal view, 30. Antennae, 31. Pronotum, 32. Mesotibia, 33–34. Aedeagus, in dorsal and lateral views. 35–40. A. tibiodentatus Wang and Ren, 2007, male, holotype, 35. Habitus, in dorsal view, 36. Antennae, 37. Pronotum, 38. Mesotibia, 39–40. Aedeagus, in dorsal and lateral views.

Distribution. China (Yunnan). 6 Anaedus spinicornis Kaszab, 1973 (Figs. 29–34) Anaedus spinicornis Kaszab, 1973: 39. Type locality: Leovaret and Salsali, Nepal. Type depository: HNHM. Specimens examined. 2♂, China, Hainan, Ledong, Jianfengling, 17–21.V.2007, Yibin Ba & Juntong Lang leg., HBUM; ♂, China, Taiwan, Gaoxiong County, Nanhengmeishan, 17.V.2008, Changqing Chen leg., HBUM; ♂, China, Yunnan, Lvchun county, 27.VII.2004, Caixia Yuan & Jing Li leg., HBUM; 1♂2♀, China, Xizang, Chayu, Chawalong, Dahuotong, 28.57343°N, 98.325383°E, alt. 2331 m, 28.VII.2017, Xinglong Bai, Zhonghua Wei, Xianlei Shao leg., HBUM; 2♀, China, Xizang, Chayu, Zhuwagen, Ria, 28.57343°N, 98.325383°E, alt. 2436 m, 29.VII.2017, Xinglong Bai, Zhonghua Wei & Xianlei Shao leg., HBUM. Distribution. China (Hainan, Taiwan, Xizang, Yunnan); Nepal, Vietnam. Diagnosis. This species is similar to A. mroczkowskii in the triangular shape of antennomere V. However, A. spinicornis has much longer and more erect pubescence and the dentiform process on antennomere V is larger and almost lamellate. 7 Anaedus substriatus Pic, 1938

sparser and smaller than those on disc. Prosternal process with shallow punctuations and sparse setae, widest in middle, apex convex downward. Scutellum glossy, triangular and without punctures. Elytra elongate-oval, 1.5 times as long as wide, and 1.1 times as wide as pronotum, lateral margins finely serrate in basal fourth, rounded at humeri. Elytral surface with dense and irregular punctures, disc convex. Visible ventrites I–IV with dense setae on lateral sides, ventrite V with denser setae. Legs short. Tibiae straight, inner apical of protibiae with setae shorter and denser than outer one. Length ratio of pro-, meso- and metafemora is 10.8: 12.8: 14.9, and that of corresponding tibiae is 10.1: 11.5: 13.0. Aedeagus (Figs. 27–28): length 1.29 mm, width 0.29 mm, curved in basal third in lateral view. Apex of the aedeagus triangular in dorsal view, longer than its width. Female. Body slightly longer and wider than male. Without distinct sexual dimorphism. Measurements. Body length 7.5–7.7 mm, body width 3.0–3.1 mm. Habit. A. serratus sp. nov. was collected in grasses around the Jinghong farm in Jinghong City. 94

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Figs. 41–50. Anaedus species. 41–45. Anaedus nonedgus sp. nov., male, holotype, 41. Habitus, in dorsal view, 42. Antennae, 43. Pronotum, 44–45. Aedeagus, in ventral and dorsal views. 46–50. A. unidentasus Wang and Ren, 2007, male, holotype, 46. habitus, in dorsal view, 47. Antennae, 48. Pronotum, 49–50. Aedeagus, in dorsal and lateral views.

Fig. 51. Distribution of known Chinese Anaedus species. ? type locality of Anaedus substriatus Pic, 1938 (distributed in Sichuan Province) without detail 95

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lateral view (probably because the adult has just emerged and aedeagal ossification is low). Apicale longer than width with rounded apex, lateral margins nearly paralleled. Female. Body size slightly larger than male; apex of ventrite V rounded; abdominal ventrites III and IV more convex than male. Measurements. Body length 7.4–8.1 mm, body width 3.5–3.6 mm. Distribution. China (Anhui, Hubei, Hunan, Sichuan, Zhejiang). 10 Anaedus unidentasus Wang & Ren, 2007 (Figs. 46–50) Anaedus unidentasus Wang & Ren, 2007: 34. Type locality: Gongshan, Yunnan, China. Type deposition: HBUM. Specimens examined. Holotype: ♂, China, Yunnan, Gongshan, 4.V.2004, Xiujuan Yang & Yushuang Liu leg., HBUM. 2♂1♀, China, Yunan, Mengla, bubang, 21.60138°N, 101.58611°E, alt. 680 m, 20.V.2017, Zhaoxu Li & Qiaoqiao Ji leg., HBUM; 2♂1♀, China, Shaanxi, Chushui, Zhonghecun, 33.56861°N, 109.40055°E, alt. 920 m, 25.VI.2015, Cixia Yuan & Ying Tian leg., HBUM; 1♂, China, Guangdong, Nanling, 8.V.2008, Xiaoyu Zhu leg., HBUM; 1♂, China, Anhui, Guichi, Xiaokeng, 5.VIII.2010, Yibin Ba & Zhenxing Zhang leg., HBUM; 1♀, China, Guizhou, Maolan, Dongtangxiang, 19.VII.2015, alt. 710 m, Caixia Yuan leg., HBUM; 2♂4♀, China, Zhejiang, Linan, Qingliangfeng, Shunxiwu, alt. 360 m, 9.VIII.2008, Junhao Huang leg., HBUM. Distribution China (Anhui, Guangdong, Guizhou, Shaanxi, Yunnan, Zhejiang).

Anaedus substriatus Pic, 1938: 17; Wang & Ren, 2007: 34 (in key). Type locality: Sichuan, China. Specimens examined. None. Distribution. China (Sichuan). 8 Anaedus tibiodentatus Wang & Ren, 2007 (Figs. 35–40) Anaedus tibiodentatus Wang & Ren, 2007: 36. Type locality: Yingjiang, Yunnan, China. Type depository: HBUM. Specimens examined. Holotype: ♂, China, Yunnan, Yingjiang, 20.VI.1995, Zhenghui Xu leg., HBUM; paratype: ♀, China, Yunnan, Mengling, alt. 2000 m, 16.IV.2002, Jinxin Song leg., HBUM. Distribution. China (Yunnan). 9 Anaedus nonedgus sp. nov. (Figs. 41–45) Type material. Holotype: ♂, China, Hubei, Yingshan, longtangu, 31.08283°N, 115.81796°E, alt. 314 m, 24.VI.2014, Guodong Ren leg., HBUM. Paratypes: ♂2♀, China, Zhejiang, Longquan, Fengyangshan, 25.VII.2007, Haoyu Liu & Zhenhua Gao leg., HBUM; 4♂, same date as holotype, HBUM; ♂, China, Anhui, Yuexi, Yaoluoping, Jingangling, 27–29.VII.2007, Yibin Ba, Juntong Lang & Fengyan Wang leg., HBUM; 1♂2♀, Sichuan, Fengtongzhai, Dashuigou, 31.VII.2016, Caixia Yuan, Saihong Dong & Shanshan Liu leg., HBUM; 1♂, Hunan, Dongshun, Huangshan, 3.X.2004, Jianhua Huang leg., HBUM. Diagnosis. Pronotal lateral part of this species without distinctly extended laterally, unique character among all Chinese species. It can be separated from the others using the following characters: (1) body with indistinctly short and sparse setae in dorsal view; (2) apex of the parameres truncated and lateral margins of apicale nearly paralleled; (3) each elytron with four indistinct longitudinal carinae. Etymology. The specific name is derived from “non”and “edge”, referring to pronotum without flat, edged lateral margins. Description. Body (Fig. 41) elongate-oval, flat; body blackish brown with shining, maxillary palpus brown; doral surface of body with indistinctly short and sparse setae. Labrum subquadrate, transversely produced in the middle, with coarse punctures and erect setae; clypeus strongly convex, with straight anterior margin, apical portion without punctures, basal portion with dense coarse punctures; frontoclypeal suture straight and deep; genae strongly raised, without punctures; frons covered with coarse punctures. Eyes transversely reniform, slightly divided by genal canthus; in dorsal view, temple length about 2 times as long as eye diameter; width at narrowest point of eye equal to 3–4 ommatidia in lateral view. Antennae (Fig. 42) robust, extending base of pronotum, terminal antennomere 1.8 times longer than width, antennomeres III–IV longer than antennomeres V–X, ratio of length of antennomeres II–XI as follows: 12.8: 28.6: 34.3: 28.4: 27.9: 29.1: 26.6: 26.6: 26.6: 40.9. Pronotum (Fig. 43) strongly transverse, 1.6 times as wide as long, widest in middle; anterior margin shallowly emarginated; lateral margins slightly flat and not expanded laterad, strongly convergent in apical half, strongly sinuous before posterior angles; posterior margin arcuate in middle, posteriorly produced; anterior angle obtuse and protruding; posterior angles acute and point outwards. Pronotal surface between lateral margins convex, with moderately dense and deep punctures on disc. Prothoracic hypomere glossy, with punctuations smaller than those on disc. Prosternal process with shallow punctuations, and slightly expanded apex. Scutellum triangular, without punctures. Elytra elongate-oval, 1.4 times as long as wide, and 1.2 times as wide as pronotum, lateral margins finely serrated in basal fifth and rounded at humeri. Elytral surface with dense and irregular punctures, disc convex, each elytron with four indistinct longitudinal carinae. Epipleura with shallow punctures. Visible abdominal ventrites I–V with sparse punctures with long setae, apex of ventrite V triangular. Legs short and thin. Tibiae straight, gradually broadening towards apex, inner apex with setae denser than outer one. Length ratio of pro-, meso- and metafemora is 1.6: 1.9: 2.2, and that of corresponding tibiae is 1.4: 1.5: 2.0. Aedeagus (Figs. 44–45): length 1.61 mm, width 0.36 mm, flat in

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