Epiverta Dieke (Coleoptera: Coccinellidae: Epilachnini): A Complex of Species, Not a Monotypic Genus

Epiverta Dieke (Coleoptera: Coccinellidae: Epilachnini): A Complex of Species, Not a Monotypic Genus Rich sampling and modern research techniques, including SEM, revealed that rarely collected epilachnine spe- cies Epiverta chelonia is a complex of four closely related species: E. chelonia (Mader, 1933), E. albopilosa, E. angusta, and E. supinata spp. nov. All Epiverta species are described and illustrated, a key to the species and a distribution map are provided. Lectotype of Solanophila chelonia Mader, 1933 is designated and its type local- ity delimited to Yunnan Province, Deqin County (China). Key words: entomology, taxonomy, Coccinelloidea, Epiverta Epilachnini Mulsant, as currently defined (Szawaryn et al. 2015, formed from the former Epilachna species (Afissa, Diekeana, Tomaszewska and Szawaryn 2016), with its over 1,000 described Ryszardia ¼ Uniparodentata—Szawaryn et al. 2015, Tomaszewska species is one of the largest groups and the only primarily phytopha- and Szawaryn 2016). In morphological analysis, it was placed gous group of the mostly predatory family Coccinellidae. The family closely to the former Epilachna species from Asia or to is classified in the superfamily Coccinelloidea (Robertson et al. Papuaepilachna plus the Papuan species of Henosepilachna 2015)—former Cerylonid Series of the superfamily Cucujoidea (e.g. (Szawaryn et al. 2015). Crowson 1955, Robertson et al. 2008, Lord et al. 2010, Slipinski  The genus Epiverta was established by Dieke (1947) for and Tomaszewska 2010, Bocak et al. 2014). Solanophila chelonia Mader, 1933, described originally from Historically Epilachninae was divided into four tribes: Sichuan and Tibet (Mader 1933). Comprehensive investigations of Epilachnini Mulsant, Madaini Gordon, Epivertini Pang & Mao and the Epilachnini materials from the museum historical collections Eremochilini Gordon & Vandenberg (Jadwiszczak and and recently collected samples of Chinese ladybirds, carried out by We ˛ grzynowicz 2003). Seago et al. (2011) however combined these the authors, revealed in new findings. Apart from the type species E. into a single tribe Epilachnini within a broadly defined subfamily chelonia, three new species have been recognized and are described Coccinellinae. This was confirmed by molecular and morphology here as Epiverta albopilosa, E. angusta, E. supinata spp. nov. This based research by Szawaryn et al. (2015) who reconstructed phylog- genus is known only from China. eny of Epilachnini confirming its monophyly, and proposed a mod- ern classification of this tribe. Szawaryn et al. (2015) made the first attempt to test monophyly Material and Methods of formerly recognized genera of phytophagous Coccinellidae This study was based on examination of material from the following (Jadwiszczak and We ˛ grzynowicz 2003, Szawaryn and museums: Tomaszewska 2013, Tomaszewska and Szawaryn 2013). As a result of that study, 27 genera of the tribe have been recognized. • IOZ—The Institute of Zoology, Chinese Academy of Science, Epiverta Dieke is very distinctive among Epilachnini, easily rec- Beijing, China; ognized by its very widely explanate elytral lateral margins and lack • MIZ—Museum and Institute of Zoology PAS, Warszawa, of metaventral and abdominal postcoxal lines. In the molecular and Poland; combined phylogenetic analyses (Szawaryn et al. 2015) Epiverta • MNHN—Muse ´ um National d’Histoire Naturelle, Paris, France; was recovered as sister taxon to the remaining genera of • BMNH—The Natural History Museum, London, United Epilachnini, as the next lineage after the split of the Asian clades Kingdom; V C The Authors 2017. Published by Oxford University Press on behalf of Entomological Society of America. 1 This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited. For commercial re-use, please contact Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 journals.permissions@oup.com by Ed 'DeepDyve' Gillespie user Version of Record, first published online April 19, 2017 with fixed content and layout in compliance with Art. 8.1.3.2 ICZN. on 17 July 2018 2 Journal of Insect Science, 2017, Vol. 17, No. 2 NHMV—Naturhistorisches Museum, Vienna, Austria; Labels of many specimens written originally in Chinese have SCAU—Department of Entomology, South China Agriculture been translated here to Latin alphabet (by LH) and new labels have University, Guangzhou, China been printed and attached to specimens examined—these labels are marked with asterisk (*) in the “Type Material” and “Other Genitalia of both sexes were dissected, cleared in a 10% solution Material Examined” sections under each species description. of KOH, rinsed with distilled water, transferred to glycerol, and ex- The beetle-specific terminology and classification follow amined on slides. Photographs were taken from slide preparations Slipinski  and Tomaszewska (2010). using a digital camera attached to Leica microscope (www.leica- microsystems.com). After examination the genitalia were transferred to microvials and pinned beneath the specimens. Measurements Nomenclature were made using an ocular micrometer attached to an Olympus This article and the nomenclatural acts it contains have been regis- (SZX 16) (www.olympus-global.com) dissecting microscope. The tered in Zoobank (www.zoobank.org), the official register of the following measurements as shown in Fig. 1a–d were made and are International Commission on Zoological Nomenclature. The LSID used in descriptions: TL—total length, from apical margin of clyp- (Life Science Identifier) number of the publication is: eus to apex of elytra; PL—pronotal length, from the middle of ante- urn:lsid:zoobank.org:pub:6D40CA1B-AE6D-482F-BC40- rior margin to margin of basal foramen; PW—pronotal width at 3480BC1AFCE6. widest part; EL—elytral length across sutural line including scutel- lum; EW—elytral width across both elytra at the widest part; ED— width of elytral disc, across both elytra, measured at the same Results position as EW; BH—body height; ventral surfaces: pl—length of Genus Diagnosis and Species Descriptions prosternum in front of procoxa; pp—width of prosternal process Epiverta Dieke, 1947 medially between procoxae; mp—width of mesoventral process me- Epiverta Dieke, 1947: 169. Type species: S. chelonia Mader, 1933 (by dially between mesocoxae; eew—elytral epipleuron width in widest original designation).—Pang and Mao 1979:159; Jadwiszczak and part; een—elytral epipleuron width in narrowest part; mf— We ˛ grzynowicz 2003:208;Kov ar 2007: 631; Szawaryn et al. 2015: mesofemur length. Scanning electron images were made using a 556, 565; Tomaszewska and Szawaryn 2016: 47, redescription. HITACHI S-3400N machine (www.hitachi.com) in the Electron Microscopy Laboratory of the MIZ. Photographs of total specimens were produced using a digital camera, and enhanced using Helicon Diagnosis. Epiverta is easily recognizable by the following combina- Focus software. The final plates were prepared using Adobe tion of characters: body (Fig. 2a–x) of males weakly smaller, more Photoshop CS. oval and less convex than in females, gular sutures at least as long as The species descriptions start from the type species then are or- half length of gula; antenna longer than head width; scutellum trans- dered alphabetically. verse or as long as wide; epipleura with at least weak foveae for Fig. 1. Measurements. (a, b) E. chelonia (Mader). (c) Epiverta supinata sp. nov. (d) Epiverta albopilosa sp. nov. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 Journal of Insect Science, 2017, Vol. 17, No. 2 3 receiving tips of femora; inner margin of epipleuron with bordering Epiverta chelonia (Mader) line extending at most to level of mid coxa; fore and mid trochanters (Figs. 2m–r, 3, 4a–g, and 5a–d) simple; tergite VIII of both sexes weakly emarginate medially at Solanophila chelonia Mader, 1933: 79. Type locality: Yunnan apex; penis guide on outer edge broadened in apical part in lateral Province, Deqin County, China. view, pointed at apex; ventral surface of coxite with sclerotized Epiverta chelonia: Dieke 1943:169.—Pang and Mao 1979: 159; pocket antero-medially; apodeme of male sternum IX absent; meta- Jadwiszczak and We ˛ grzynowicz 2003: 208; Tomaszewska and ventral and abdominal postcoxal lines absent or sometimes poorly Szawaryn 2016: 47. developed postcoxal lines visible on metaventrite. Differential Diagnosis. Epiverta chelonia most closely resembles E. Distribution. Southwest China (including Tibet) (Fig. 3). supinata in its body shape and color pattern on the elytra, however Fig. 2. Habitus: (a, b, c) Epiverta supinata sp. nov., male—dorsal, lateral and anterior. (d, e, f): Epiverta supinata sp. nov., female—dorsal, lateral and anterior. (g, h, i) Epiverta angusta sp. nov., male—dorsal, lateral and anterior. (j, k, l) Epiverta angusta sp. nov., female—dorsal, lateral and anterior. (m, n, o) E. chelonia (Mader), male—dorsal, lateral and anterior. (p, q, r) E. chelonia (Mader), female—dorsal, lateral and anterior. (s, t, u) Epiverta albopilosa sp. nov., male–dorsal, lat- eral and anterior. (v, w, x) Epiverta albopilosa sp. nov., female—dorsal, lateral and anterior. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 4 Journal of Insect Science, 2017, Vol. 17, No. 2 Fig. 3. Distribution map of the genus Epiverta. E. chelonia can be separated from that species by having humeral subtruncate to weakly emarginate. Tergite  transverse subtruncate margins of the elytra simple (Fig. 4c and f) (at most weakly thick- to scarcely emarginate at apex. In females apical margin of ventrite ened but not upturned) and lateral margins of the pronotum evenly 5 weakly emarginate; ventrite 6 rounded apically; tergite VIII sub- arcuate (Fig. 4b). Moreover, the following combination of charac- truncate to scarcely excised apically. ters distinguish E. chelonia from its congeners: (1) penis guide not Male genitalia. Aedeagus stout. Tegmen (Fig. 5b and c) with par- longer than parameres, (2) tegminal strut more or less distinctly ameres scarcely longer than penis guide; tegminal strut truncate or emarginate at its apex, (3) anterior elytral maculae (near scutellum) emarginate apically; anterior margin of tegminal plate weakly pro- in form of weakly arcuate, most often uniformly yellow stripes or duced, emarginate or simply rounded medially (without pointed tri- sometimes with inner dark marking (Fig. 2m and p). angular projection medially). Penis guide with pointed apex almost straight or weakly curved backwards in lateral view (Fig. 5b). Penis with remnant of inner arm of reduced basal T-shaped capsule in Description. Length 5.55–7.55 mm; TL/EW ¼ 1.07–1.20; PL/ form of regular triangle (Fig. 5a). PW ¼ 0.38–0.42; EL/EW ¼ 0.88–1.01; TL/BH ¼ 2.36–2.70. Female genitalia (Fig. 5d) with proctiger (TX) weakly excised at Body regularly oval (Figs. 2m and p, 4f). Background of elytra apex; coxites with styli well developed; spermatheca weakly curved distinctly brownish black with regular yellow maculae arranged as in about half length; accessory gland about 2 times longer than in Fig. 2m and p. Pubescence moderately dense and moderately spermatheca. long, yellowish. Head (Fig. 4b) with interocular distance 0.68 times as wide as head across eyes. Antenna (Fig. 4a) with antennomere 3 about 1.25 Type Material. Lectotype of S. chelonia Mader, male, YUNNAN times as long as antennomere 4 and about 1.15 times as long as 5; PROVINCE, “Thibet. Atentse [¼Deqin]. (R. P. Goutelle)./Coll. antennomere 4 about 0.90 times as long as antennomere 5. Mus. Vindob./Fu ¨ r diese Art [.. .] jetzt eine neue Gattung gemacht Terminal maxillary palpomere about 1.12 times wider than long worden (hand writing)/S. chelonia Mad., det. Mader” (NHMV). (Fig. 4e). Paralectotype of S. chelonia Mader, female, “Thibet. Atentse Pronotum with lateral margins evenly arcuate (Fig. 4b). [¼Deqin]. (R. P. Goutelle)/Epilachna chelonia Fairm.” (MNHN). Prosternal process 0.62–0.74 times as wide as mesoventral process (pp/mp) (Fig. 4d); 0.80–0.97 times as wide as length of prosternum Other Material Examined. YUNNAN PROVINCE, “Yunnan in front of procoxa (pp/pl). Elytra with lateral flattened margins Province, Deqin County, 3,300 m, 2.IX.1987, Wang Shuyong leg./ 00 00 scarcely narrowing toward elytral apices (Fig. 4f); humeral margins IOZ(E) 626305 (*one: MIZ); same but IOZ(E) 626306 (*one: at most weakly thickened and raised (Fig. 4c); elytral total width IOZ); “Yunnan Province, Zhongdian County, Wegnshui Village, 1.35–1.48 times of elytral disc width (EW/ED); elytral epipleuron in 3,000 m, 10.VII.1982, Zhang Xuezhong leg./IOZ(E) 626243 (*one widest part 2.75–4.00 times as wide as epipleuron in narrowest part female: MIZ); “Yunnan Province, Deqin County, Meili Snow (eew/een), 2.65–3.40 times as wide as mesoventral process (eew/mp) Mountain, 3,350 m, 25.VII.1982, Zhang Xuezhong leg./IOZ(E) and 0.90–1.07 as wide as mesofemur length (Fig. 4g) (eew/mf). 626307 (*one male: IOZ); same but IOZ(E) 626308 (*one female: Abdomen. Hind margin of male ventrite 5 widely emarginate. MIZ); “CHINA: NW-Yunnan, 10 km SW Lijiang, 2,500 m, Hind margin of male ventrite 6 excised. Male tergite VIII 5.7.1994, leg. Schillhammer (13) (one female: MIZ, and two: Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 Journal of Insect Science, 2017, Vol. 17, No. 2 5 Fig. 4. E. chelonia (Mader): (a) antenna, (b) pronotum, (c) antero-lateral margin of elytron, (d) prosternum and mesoventrite, (e) mouthparts, (f) habitus, dorsal view, (g) habitus, ventral view. Fig. 5. E. chelonia (Mader): (a) penis, (b) tegmen, lateral, (c) tegmen, ventral, (d) female genitalia. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 6 Journal of Insect Science, 2017, Vol. 17, No. 2 Fig. 6. Epiverta albopilosa sp. nov.: (a) head and prothorax, ventral view, (b) mouthparts, (c) antenna, (d) pronotum, (e) habitus, dorsal view, (f) habitus, ventral view. NHMV); “Yunnan Province, Dali, Cangshan Mountain, 2,850 m, Differential Diagnosis. Epiverta albopilosa resembles E. angusta in 30.VI.1981, Wang Shuyong leg./IOZ(E) 626310 (*one male: MIZ); its dorsal coloration—background and maculae more irregular (as same but IOZ(E) 626311 (*one female: IOZ); “Yunnan Province, compared with E. chelonia and E. supinata) with background color Lijiang City, 3100 m, 27.V.1980, Wang Linyao leg./IOZ(E) 626285 of the elytra hard to distinguish (Fig. 2a and v). Both species are also (*one male: IOZ). SICHUAN PROVINCE, “China, S.W. Sichuan, similar in having white or greyish elytral pubescence. E. albopilosa, 0 00  0 00 Qingmai 3,000 m, N28 48 46.7 , E099 52 09.2 , 3.7.2006, lgt. however, can be distinguished from that species and also from E. Janata M./BMNH(E), 2006-158, M Janata/Epiverta sp., det. R.G. chelonia and E. supinata by: (1) antennomere 3 much longer than 4, Booth 2006 (female and male: BMNH). (2) dorsal surfaces covered with dense and comparatively long, white pubescence, (3) elytral lateral flattened margins widest in Distribution. Southwest China: NW Yunnan, SW Sichuan (Fig. 3). about 1/3 of elytral length thence narrowing towards elytral apices, (4) penis base of different shape (Fig. 7a). Epiverta albopilosa sp. nov Description. Length 6.10–7.73 mm; TL/EW ¼ 1.18–1.24; PL/ (Figs. 2s–x and 3, 6a–f, 7a–d) PW ¼ 0.41–0.42; EL/EW ¼ 0.99–1.04; TL/BH ¼ 2.40–3.00. (urn:lsid:zoobank.org:act:D00BEA06-441F-4757-A8C8- Body oval in males and elongate oval in females (Figs. 2s and v 8DF76B9FD0AE) and 6e). Dorsum bicoloured: brownish black and orange–yellow but Etymology. The name of the new species refers to white, dense pu- with background of elytra hard to distinguish; maculae irregular as bescence covering the body, especially its dorsal surfaces. in Fig. 2s and v. Pubescence dense, comparatively long, white. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 Journal of Insect Science, 2017, Vol. 17, No. 2 7 Fig. 7. Epiverta albopilosa sp. nov.: (a) penis, (b) tegmen, lateral, (c) tegmen, ventral, (d) female genitalia. Head (Fig. 6d) with interocular distance 0.70 times as wide as Type Material. Holotype, male, YUNNAN PROVINCE, “CHINA, head across eyes. Antenna (Fig. 6a and c)withantennomere 3 Yunnan Prov., Dongchuan; 19.VIII.1982; Xiao Ningnian leg.” about 1.80 times as long as antennomere 4 and about 1.38 times (SCAU). Paratypes, “CHINA, Yunnan Prov., Dongchuan; 19-VIII- as long as 5; antennomere 4 about 0.80 times as long as antenno- 1982; Xiao Ningnian leg.” (two males: SCAU; one male and one fe- mere 5. Terminal maxillary palpomere about as wide as long male: MIZ); “Yunnan Province, Luquan County, Jiaozishan (Fig. 6b). Mountain, 2750m, 23-VIII-2013, Huo Lizhi leg.” (one female: SCAU). Pronotum with lateral margins evenly arcuate (Fig. 6d). Prosternal process 0.62–0.80 times as wide as mesoventral pro- Distribution. Southwest China: NE Yunnan (Fig. 3). cess (pp/mp) (Fig. 6a); 0.82–0.96 times as wide as length of pros- ternum in front of procoxa (pp/pl). Elytra with lateral flattened Epiverta angusta sp. nov margins gradually narrowing from about 1/3 of elytral length to- (Figs. 2g–l, 3, 8a–f, 9a–d) ward their apices (Fig. 6e) humeral margins simple; elytral total (urn:lsid:zoobank.org:act:D32E2A64-A9AB-4362-A7F0- width 1.22–1.32 times of elytral disc width (EW/ED); elytral epi- BFFDA632C62E) pleuron in widest part 3.05–3.85 times as wide as epipleuron in Epiverta chelonia: Bielawski 1960: 441 (misidentification). narrowest part (eew/een), 2.70–3.20 times as wide as mesoventral process (eew/mp) and 0.70–0.82 as wide as mesofemur length Etymology. The name of the new species in Latin means narrow and (eew/mf) (Fig. 6f). refers to flattened lateral margins of the elytra which are compara- Abdomen. Hind margin of male ventrite 5 truncate to weakly tively narrow throughout of their length. emarginate. Hind margin of male ventrite 6 weakly emarginate. Male tergite VIII scarcely emarginate. Tergite  transverse truncate Differential Diagnosis. Epiverta angusta is most similar to E. albopilosa at apex. In females apical margin of ventrite 5 truncate; ventrite 6 in its dorsal coloration with the background and maculae more irregular subtruncate apically; tergite VIII rounded to subtruncate apically. (as compared with E. chelonia and E. supinata) with background color Male genitalia. Aedeagus stout. Tegmen (Fig. 7b and c)with of the elytra hard to distinguish (Fig. 2d and j). Both species are also cov- parameres slightly shorter than penis guide; tegminal strut ered with white or greyish elytral pubescence (yellowish in E. chelonia rounded apically; anterior margin of tegminal plate weakly pro- and E. supinata). E.angusta, however, can be distinguished from E. albo- duced, simply rounded medially or scarcely emarginate medially. pilosa by: (1) lateral elytral flattened margins comparatively narrow Penis guide with pointed apex distinctly upturned backwards (vis- throughout of their length, (2) antennomere 3 about 1.25 times longer ible in lateral view) (Fig. 7b). Penis with remnant of inner arm of than4(1.8times longer in E. albopilosa), (3) antennomeres 3 and 5 sub- reduced basal T-shaped capsule comparatively large, irregularly equal in length (antennomere 3 about 1.38 times longer than 5 in E. albo- triangular (Fig. 7a). pilosa), (4) different shape of penis base and spermatheca. Female genitalia (Fig. 7d) with proctiger (TX) scarcely excised at apex; coxites with styli well developed; spermatheca variable, shorter weakly curved near base or longer curved near mid length; Description. Length 5.62–7.30 mm; TL/EW ¼ 1.11–1.27; PL/ accessory gland about 3 times longer than spermatheca. PW ¼ 0.39–0.42; EL/EW ¼ 1.00–1.10; TL/BH ¼ 2.27–2.74. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 8 Journal of Insect Science, 2017, Vol. 17, No. 2 Fig. 8. Epiverta angusta sp. nov.: (a) pronotum and base of elytra, (b) antero-lateral margin of elytron, (c) mouthparts, (d) pro- and mesothorax, ventral view, (e) habitus, dorsal view, (f) habitus, ventral view. Body oval in males and elongate oval in females (Figs. 2g and j (eew/mp) and 0.83–1.00 as wide as mesofemur length (eew/mf) and 8e). Dorsum bicoloured: brownish black and orange-yellow (Fig. 8f). butwithbackgroundof elytrahardtodistinguish;maculae irreg- Abdomen. Hind margin of male ventrite 5 distinctly emarginate. ular as in Fig. 2g and j. Pubescence moderately dense and long, Hind margin of male ventrite 6, tergite VIII and tergite X with greyish. scarce emargination/excision at apex. In females apical margin of Head (Fig. 8a) with interocular distance 0.70 times as wide as ventrite 5 and tergite VIII subtruncate apically; ventrite 6 subtrun- head across eyes. Antenna (Fig. 8d) with antennomere 3 about 1.25 cate to weakly triangularly angulate (with hind margin weakly proj- times as long as antennomere 4 and about 1.04 times as long as 5; ected backwards medially). antennomere 4 about 0.83 times as long as antennomere 5. Male genitalia. Aedeagus stout. Tegmen (Fig. 9b and c) with par- Terminal maxillary palpomere about as wide as long (Fig. 8c). ameres slightly shorter than penis guide; tegminal strut rounded api- Pronotum with lateral margins evenly arcuate (Fig. 8a). cally; anterior margin of tegminal plate not produced, simply Prosternal process 0.62–0.73 times as wide as mesoventral pro- rounded medially. Penis guide with pointed apex weakly upturned cess (pp/mp) (Fig. 8d); 0.86–1.04 times as wide as length of pros- backwards (visible in lateral view) (Fig. 9b). Penis with remnant of ternum in front of procoxa (pp/pl). Elytra with lateral flattened inner arm of reduced basal T-shaped capsule comparatively small, margins comparatively narrow, scarcely narrowing toward elytral irregularly triangular (Fig. 9a). apices (Fig. 8e); humeral margins simple; elytral total width 1.27– Female genitalia (Fig. 9d) with proctiger (TX) subtruncate or 1.35 times of elytral disc width (EW/ED); elytral epipleuron in sometimes weakly emarginated at apex; coxites with styli well devel- widest part 3.15–3.77 times as wide as epipleuron in narrowest oped; spermatheca curved in mid length; accessory gland 2.5–2.8 part (eew/een), 2.90–3.73 times as wide as mesoventral process times as long as than spermatheca. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 Journal of Insect Science, 2017, Vol. 17, No. 2 9 Fig. 9. Epiverta angusta sp. nov.: (a) penis, (b) tegmen, lateral, (c) tegmen, ventral, (d) female genitalia. Type Material. Holotype, male, SICHUAN PROVINCE, “Ta-Tsien- recognize these specimens as belonging to the new species, described Lu, 30.V.93, Potanin (translated and transcripted here from here as E. angusta. Russian, hand-writting)/E. chelonia (Mader), det. R. Bielawski 1959)/Inst. Zool. P.A.N, Warszawa, 86/60” (MIZ). Paratypes, “Fu- Distribution. Southwest China: SW Sichuan (Fig. 3). dian-xo (translated and transcripted here from Russian, hand writ- ing)/K. A. Semenowa/E. chelonia (Mader), det. R. Bielawski 1958)/ Epiverta supinata sp. nov Inst. Zool. P.A.N, Warszawa, 95/59” (one male: MIZ); “Thibet, (Figs. 2a–f, 3, 10a–g, 11a–d) Chasseurs de, T a-tsien-lou, ^ 1895” (two males, two females and 13: (urn:lsid:zoobank.org:act:0BC7054D-3368-422D-9D97- MNHN; 2: MIZ); “Chasseurs Indige ` nes, des Missionnaires, de Ta- EBAC94345B88) tsien-Lou,  1906” (one male, one female and nine: MNHN); “Sichuan Province, Kangding County, 2,400 m, 1963.VII.13, Mao Jinlong leg.” (one male: IOZ); “Thibet, Tatsienlou, mgr F. Birt/ Etymology. The name of the new species is derived from Latin Museum Paris, ex Coll., R. Oberthu ¨ r” (two: MNHN; 1: MIZ); “Su- “supinatas”, which means upturned and refers to humeral margins Tchuen, Mo-Sy-Mien, 1897” (two males, four females and 12: of the elytra upturned backwards. MNHN; 3: MIZ); “Su-Tchuen, Chass. Thibetains, 1897” (two: MNHN); same but 1899 (one: MNHN; one: MIZ); “CHINA: Differential Diagnosis. Epiverta supinata can be easily separated Sichuan Prov., 1999, Zeng Tao leg.” (two males, two females: from its congeners by having humeral margins of the elytra thick- SCAU; one male and one female: MIZ). ened and upturned backwards (Fig. 10c and f), lateral margins of the pronotum subparallel along basal 2/3 length, antennomere 4 and Comment. Bielawski (1960) studied Epiverta material from “der 5 subequal in length, female genitalia with comparatively short styli Sammlung des Zoologischen Institut der Akademie der and anterior margin of tegminal plate regularly, triangularly pro- Wissenschafte der UdSRR in Leningrad (now Saint Petersburg). He duced medially. studied 14 specimens from Sichuan Province (Ta-Tsien-Lu and Mao-nju-ku-Tal) and treated them as E. chelonia. He also provided Description. Length 6.10–6.30 mm; TL/EW ¼ 1.03–1.17; PL/ drawings of the male genitalia of this species unknown until then. PW ¼ 0.38–0.43; EL/EW ¼ 0.90–0.98; TL/BH ¼ 2.11–2.60. After study of two specimens from that series (retained in the MIZ Body oval (Figs. 2a and d and 10f). Background of elytra brown- collection), his pictures of the male genitalia (Bielawski 1960) (with ish black with regular yellow maculae arranged as in Fig. 2a and d, tegmen with parameres slightly shorter than penis guide, tegminal most of them with inclusions of blackish spots. Pubescence moder- strut rounded apically, anterior margin of tegminal plate not pro- ately dense and moderately long, yellowish. duced and simply rounded medially, and penis guide with pointed Head (Fig. 10b) with interocular distance 0.71 times as wide as apex weakly upturned backwards in lateral view) and the drawings head across eyes. Antenna (Fig. 10e) with antennomere 3 about 1.10 of elytral pattern commented by him, that “Die Flecken auf den times as long as antennomere 4 and about 1.15 times as long as 5; Flu ¨ geldecken verbindensich miteinender auf sehr verschidene Weise antennomere 4 about as long as antennomere 5. Terminal maxillary und bilden eine Mosaik Schwarz und braungrau gef€ arbt”, we palpomere about as wide as long (Fig. 10e). Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 10 Journal of Insect Science, 2017, Vol. 17, No. 2 Fig. 10. Epiverta supinata sp. nov.: (a) pronotum and base of elytra, (b) pronotum, (c) antero-lateral margin of elytron, (d) prosternum and mesoventrite, (e) mouthparts, (f) habitus, dorsal view, (g) habitus, ventral view. Female genitalia (Fig. 11d) with proctiger (TX) scarcely to Pronotum with lateral margins subparallel along basal 2/3 length then arcuate anteriorly (Fig. 10a and b). Prosternal process 0.65– weakly excised medially at apex; coxites with styli small to well de- 0.80 times as wide as mesoventral process (pp/mp) (Fig. 10d); 0.83– veloped; spermatheca curved in mid length; accessory gland 2.5–3.0 1.15 times as wide as length of prosternum in front of procoxa (pp/ times longer than spermatheca. pl). Elytra with lateral flattened margins scarcely narrowing towards elytral apices (Fig. 10f); humeral margins thickened and upturned Type Material. Holotype, male, SICHUAN PROVINCE, “Thibet, backwards (Fig. 10c and f); elytral total width 1.49–1.60 times of Yare ´ gong [Sichuan], P. Soulie ´ , 1900” (MNHN). Paratypes, same elytral disc width (EW/ED); elytral epipleuron in widest part 2.70– data as holotype (one male, two females and two: MNHN; two: 3.70 times as wide as epipleuron in narrowest part (eew/een), 3.90– MIZ); “Sichuan Province, Xiangcheng County, 2900 m, 4.80 times as wide as mesoventral process (eew/mp) and 1.10–1.30 17.VI.1982, Wang Shuyong leg./IOZ(E) 626295” (*one: IOZ); as wide as mesofemur length (eew/mf) (Fig. 10g). same but IOZ(E) 626296 and IOZ(E) 626297” (*one and male, re- Abdomen. Hind margin of male ventrite 5 emarginate. Hind spectively: IOZ); “Sichuan Province, Xiangcheng County, 3200 m, margin of male ventrite 6 weakly excised; tergite VIII and tergite X 26.VI.1982, Chai Huaicheng leg./IOZ(E) 626293” (*one: IOZ); with weakly emarginate at apex. In females apical margin of ventrite same but IOZ(E) 626294 (*one female: MIZ). TIBET, “Tibet, 5 truncate; ventrite 6 subtruncate to weakly rounded apically; tergite Mangkang County, Haitong, 3250 m, 12.VIII.1982, Wang Shuyong VIII subtruncate to scarcely emarginate apically. leg./IOZ(E) 626298” (male: *IOZ). “Tibet, Mangkang County, Male genitalia. Aedeagus stout. Tegmen (Fig. 11b and c) with Haitong 3200 m, 9.VIII.1982, Wang Shuyong leg./IOZ(E) 626301” parameres slightly shorter than penis guide; tegminal strut rounded (*one: MIZ); same data but IOZ(E) 626302” (*one female: MIZ); apically; anterior margin of tegminal plate triangularly produced “Tibet, Mangkang County, Haitong 3250 m, 11.VIII.1982, Wang medially. Penis guide with pointed apex weakly upturned back- Shuyong leg./IOZ(E) 626300” (*one male: IOZ); same data but wards (visible in lateral view) (Fig. 11b). Penis with remnant of inner IOZ(E) 626303” (*one female: IOZ); “Tibet, Mangkang County, arm of reduced basal T-shaped capsule most often trigger like Haitong 3250 m, 12.VIII.1982, Wang Shuyong leg./IOZ(E) (Fig. 11a). 626259” (*one: IOZ); same data but IOZ(E) 626299” (*one: IOZ). Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 Journal of Insect Science, 2017, Vol. 17, No. 2 11 Fig. 11. Epiverta supinata sp. nov.: (a) penis, (b) tegmen, lateral, (c) tegmen, ventral, (d) female genitalia. YUNNAN PROVINCE, “Yunnan Province, Deqin County, Adong – Body covered with greyish pubescence (Fig. 2g–l); elytra 1.00– Village, 2,700–3,000 m, 6.IX.1981, Wang Shuyong leg./IOZ(E) 1.10 times as long as wide; total width of elytra 1.27–1.35 times as 626255” (*one male: IOZ); same but IOZ(E) 626256 and IOZ(E) wide as elytral disc; antennomere 3 about as long as antennomere 5; 626257 (*two: IOZ). penis guide slightly longer parameres (Fig. 9b); tegminal strut rounded apically (Fig. 9c) ................................. . E. angusta sp. nov. Distribution. Southwest China: E Tibet, NW Yunnan, SW Sichuan (Fig. 3). Acknowledgments We thank Roger Booth (BMNH), Antoine Mantilleri (MNHN), Harald Schillhammer and Rudolf Schuh (NHM) and Kuiyan Zhang (IOZ) for the loan of specimens used in this study. Adam Slipinski  (CSIRO) read an early Key to Species of Epiverta draft of this manuscript and provided helpful suggestions. Magdalena 1. Lateral margins of pronotum subparallel along basal 2/3 Kowalewska-Groszkowska (MIZ) helped with SEM images. length (Fig. 10a and b); humeral margins of elytra thickened and up- turned backwards (Fig. 10c, f); total width of elytra 1.49-1.60 times References Cited as wide as elytral disc; elytral epipleuron in widest part 1.10-1.30 Bielawski, R. 1960. Materialien zur Kenntnis der Coccinellidae (Coleoptera). times of mesofemur length ............................... E. supinata sp. nov. Ann. Zool. 18: 435–458. – Lateral margins of pronotum regularly at least weakly arcuate; Bocak, L., C. Barton, A. Crampton-Platt, D. Chesters, D. Ahrens, and A. humeral margins of elytra simple and not upturned backwards; total Vogler. 2014. Building the Coleoptera tree-of-life for >8000 species: com- width of elytra 1.22–1.48 times as wide as elytral disc; elytral epi- position of public DNA data and fit with Linnaean classification. Syst. pleuron in widest part 0.70–1.07 times of mesofemur length ........ 2 Entomol. 39: 97–110. 2. Elytra covered with dense and comparatively long, white pu- Crowson, R. A. 1955. The natural classification of the families of Coleoptera, bescence (Fig. 2s and v); antennomere 3 about 1.8 times as long as 187 pp. Nathaniel Lloyd, London. antennomere 4 (Fig. 6c); elytral epipleuron in widest part 0.70-0.82 Dieke, G. H. 1947. Ladybeetles of the genus Epilachna (sens. lat.) in Asia, Europe and Australia. Smithsonian Miscellaneous Collect. 106: 1–183. times of mesofemur length (Fig. 6f); .............. E. albopilosa sp. nov. Jadwiszczak, A., and P. We˛grzynowicz. 2003. World Catalogue of – Elytra covered with sparser and shorter, yellowish or greyish Coccinellidae. Part I—Epilachninae, 264 pp. Mantis, Olsztyn. pubescence; antennomere 3 about 1.25 times as long as antenno- Kovar, I. 2007. Family Coccinellidae Latreille, 1807, pp. 568–631. In I., Lobl mere 4; elytral epipleuron in widest part 0.83–1.07 times of mesofe- & A., Smetana (Ed.), Catalogue of Palaearctic Coleoptera, Vol. 4. mur length .................................................................................... 3 Stenstrup, Apollo Books, 935 pp. 3. Body covered with yellowish pubescence (Fig. 2m–r); elytra at Lord, N., C. S. Hartley, J. F. Lawrence, J. V. McHugh, M. F. Whiting, and K. most as long as wide; total width of elytra 1.35–1.48 times as wide as B. Miller. 2010. Phylogenetic analysis of the minute brown scavenger bee- elytral disc; antennomere 3 about 1.15 times as long as antennomere 5; tles (Coleoptera: Latridiidae), and recognition of a new beetle family, penis guide at most as long as parameres (Fig. 5b); tegminal strut emar- Akalyptoischiidae fam. n. (Coleoptera: Cucujoidea). Syst. Entomol. 35: ginate or truncate apically (Fig. 5c) ....................... E. chelonia (Mader) 753–763. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 12 Journal of Insect Science, 2017, Vol. 17, No. 2 Mader, L. 1933. Uber bekannte und neue Coccinelliden. Entomologischer Szawaryn, K., and W. Tomaszewska. 2013. Two new genera of Epilachnini Anzeiger, Wien. 13: 79–84. Mulsant from New Guinea and Aru Islands (Coleoptera: Coccinellidae). Pang, X. F., and J. L. Mao. 1979. Economic Insect Fauna of China (XIV) , 170 J. Nat. Hist. doi:10.1080/00222933.2012.763067. pp. Coleoptera: Coccinellidae II. Science Press, Beijing Szawaryn, K., L. Bocak, A. Slipinski,  H. E. Escalona, and W. Tomaszewska. Robertson, J. A., F. Whiting, and J. V. McHugh. 2008. Searching for natural 2015. Phylogeny and evolution of phytophagous ladybird beetles lineages within the Cerylonid Series (Coleoptera: Cucujoidea). Mol. (Coleoptera: Coccinellidae: Epilachnini), with recognition of new genera. Phylogenet. Evol. 46: 193–205. Syst. Entomol. 40: 547–569. Robertson, J., A. Slipinsk  i, M. Moulton, F. W. Shockley, A. Giorgi, N. P. Slipinski,  S. A., and W. Tomaszewska. 2010. Coccinellidae Latreille, 1802, Lord, D. D. McKenna, W. Tomaszewska, J. Forrester, K. B. Miller, et al. pp. 454–72. In: R.A.B., Leschen, R.G, Beutel and J.F. Lawrence (ed.): 2015. Phylogeny and classification of Cucujoidea and the recognition of a Handbook of Zoology, Vol. 2, Coleoptera. Berlin/New York: Walter de new superfamily Coccinelloidea (Coleoptera: Cucujiformia). Syst. Entomol. Gruyter GmbH & Co. KG. XIII þ 786 pp. 40: 745–778. Tomaszewska, W., and K. Szawaryn. 2013. Revision of the Asian species of Seago, A. E., A. Giorgi, J. Li, and A. Slipinski.  2011. Phylogeny, classification Afidentula Kapur, 1958 (Coleoptera: Coccinellidae: Epilachnini). Zootaxa. and evolution of ladybird beetles (Coleoptera: Coccinellidae) based on si- 3608: 26–50. multaneous analysis of molecular and morphological data. Mol. Tomaszewska, W., and K. Szawaryn. 2016. Epilachnini (Coleoptera: Phylogenet. Evol. 60: 137–151. Coccinellidae)—a revision of the world genera. J. Insect Sci. 16: 101. 1–91. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Journal of Insect Science Oxford University Press

Epiverta Dieke (Coleoptera: Coccinellidae: Epilachnini): A Complex of Species, Not a Monotypic Genus

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Abstract

Rich sampling and modern research techniques, including SEM, revealed that rarely collected epilachnine spe- cies Epiverta chelonia is a complex of four closely related species: E. chelonia (Mader, 1933), E. albopilosa, E. angusta, and E. supinata spp. nov. All Epiverta species are described and illustrated, a key to the species and a distribution map are provided. Lectotype of Solanophila chelonia Mader, 1933 is designated and its type local- ity delimited to Yunnan Province, Deqin County (China). Key words: entomology, taxonomy, Coccinelloidea, Epiverta Epilachnini Mulsant, as currently defined (Szawaryn et al. 2015, formed from the former Epilachna species (Afissa, Diekeana, Tomaszewska and Szawaryn 2016), with its over 1,000 described Ryszardia ¼ Uniparodentata—Szawaryn et al. 2015, Tomaszewska species is one of the largest groups and the only primarily phytopha- and Szawaryn 2016). In morphological analysis, it was placed gous group of the mostly predatory family Coccinellidae. The family closely to the former Epilachna species from Asia or to is classified in the superfamily Coccinelloidea (Robertson et al. Papuaepilachna plus the Papuan species of Henosepilachna 2015)—former Cerylonid Series of the superfamily Cucujoidea (e.g. (Szawaryn et al. 2015). Crowson 1955, Robertson et al. 2008, Lord et al. 2010, Slipinski  The genus Epiverta was established by Dieke (1947) for and Tomaszewska 2010, Bocak et al. 2014). Solanophila chelonia Mader, 1933, described originally from Historically Epilachninae was divided into four tribes: Sichuan and Tibet (Mader 1933). Comprehensive investigations of Epilachnini Mulsant, Madaini Gordon, Epivertini Pang & Mao and the Epilachnini materials from the museum historical collections Eremochilini Gordon & Vandenberg (Jadwiszczak and and recently collected samples of Chinese ladybirds, carried out by We ˛ grzynowicz 2003). Seago et al. (2011) however combined these the authors, revealed in new findings. Apart from the type species E. into a single tribe Epilachnini within a broadly defined subfamily chelonia, three new species have been recognized and are described Coccinellinae. This was confirmed by molecular and morphology here as Epiverta albopilosa, E. angusta, E. supinata spp. nov. This based research by Szawaryn et al. (2015) who reconstructed phylog- genus is known only from China. eny of Epilachnini confirming its monophyly, and proposed a mod- ern classification of this tribe. Szawaryn et al. (2015) made the first attempt to test monophyly Material and Methods of formerly recognized genera of phytophagous Coccinellidae This study was based on examination of material from the following (Jadwiszczak and We ˛ grzynowicz 2003, Szawaryn and museums: Tomaszewska 2013, Tomaszewska and Szawaryn 2013). As a result of that study, 27 genera of the tribe have been recognized. • IOZ—The Institute of Zoology, Chinese Academy of Science, Epiverta Dieke is very distinctive among Epilachnini, easily rec- Beijing, China; ognized by its very widely explanate elytral lateral margins and lack • MIZ—Museum and Institute of Zoology PAS, Warszawa, of metaventral and abdominal postcoxal lines. In the molecular and Poland; combined phylogenetic analyses (Szawaryn et al. 2015) Epiverta • MNHN—Muse ´ um National d’Histoire Naturelle, Paris, France; was recovered as sister taxon to the remaining genera of • BMNH—The Natural History Museum, London, United Epilachnini, as the next lineage after the split of the Asian clades Kingdom; V C The Authors 2017. Published by Oxford University Press on behalf of Entomological Society of America. 1 This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited. For commercial re-use, please contact Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 journals.permissions@oup.com by Ed 'DeepDyve' Gillespie user Version of Record, first published online April 19, 2017 with fixed content and layout in compliance with Art. 8.1.3.2 ICZN. on 17 July 2018 2 Journal of Insect Science, 2017, Vol. 17, No. 2 NHMV—Naturhistorisches Museum, Vienna, Austria; Labels of many specimens written originally in Chinese have SCAU—Department of Entomology, South China Agriculture been translated here to Latin alphabet (by LH) and new labels have University, Guangzhou, China been printed and attached to specimens examined—these labels are marked with asterisk (*) in the “Type Material” and “Other Genitalia of both sexes were dissected, cleared in a 10% solution Material Examined” sections under each species description. of KOH, rinsed with distilled water, transferred to glycerol, and ex- The beetle-specific terminology and classification follow amined on slides. Photographs were taken from slide preparations Slipinski  and Tomaszewska (2010). using a digital camera attached to Leica microscope (www.leica- microsystems.com). After examination the genitalia were transferred to microvials and pinned beneath the specimens. Measurements Nomenclature were made using an ocular micrometer attached to an Olympus This article and the nomenclatural acts it contains have been regis- (SZX 16) (www.olympus-global.com) dissecting microscope. The tered in Zoobank (www.zoobank.org), the official register of the following measurements as shown in Fig. 1a–d were made and are International Commission on Zoological Nomenclature. The LSID used in descriptions: TL—total length, from apical margin of clyp- (Life Science Identifier) number of the publication is: eus to apex of elytra; PL—pronotal length, from the middle of ante- urn:lsid:zoobank.org:pub:6D40CA1B-AE6D-482F-BC40- rior margin to margin of basal foramen; PW—pronotal width at 3480BC1AFCE6. widest part; EL—elytral length across sutural line including scutel- lum; EW—elytral width across both elytra at the widest part; ED— width of elytral disc, across both elytra, measured at the same Results position as EW; BH—body height; ventral surfaces: pl—length of Genus Diagnosis and Species Descriptions prosternum in front of procoxa; pp—width of prosternal process Epiverta Dieke, 1947 medially between procoxae; mp—width of mesoventral process me- Epiverta Dieke, 1947: 169. Type species: S. chelonia Mader, 1933 (by dially between mesocoxae; eew—elytral epipleuron width in widest original designation).—Pang and Mao 1979:159; Jadwiszczak and part; een—elytral epipleuron width in narrowest part; mf— We ˛ grzynowicz 2003:208;Kov ar 2007: 631; Szawaryn et al. 2015: mesofemur length. Scanning electron images were made using a 556, 565; Tomaszewska and Szawaryn 2016: 47, redescription. HITACHI S-3400N machine (www.hitachi.com) in the Electron Microscopy Laboratory of the MIZ. Photographs of total specimens were produced using a digital camera, and enhanced using Helicon Diagnosis. Epiverta is easily recognizable by the following combina- Focus software. The final plates were prepared using Adobe tion of characters: body (Fig. 2a–x) of males weakly smaller, more Photoshop CS. oval and less convex than in females, gular sutures at least as long as The species descriptions start from the type species then are or- half length of gula; antenna longer than head width; scutellum trans- dered alphabetically. verse or as long as wide; epipleura with at least weak foveae for Fig. 1. Measurements. (a, b) E. chelonia (Mader). (c) Epiverta supinata sp. nov. (d) Epiverta albopilosa sp. nov. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 Journal of Insect Science, 2017, Vol. 17, No. 2 3 receiving tips of femora; inner margin of epipleuron with bordering Epiverta chelonia (Mader) line extending at most to level of mid coxa; fore and mid trochanters (Figs. 2m–r, 3, 4a–g, and 5a–d) simple; tergite VIII of both sexes weakly emarginate medially at Solanophila chelonia Mader, 1933: 79. Type locality: Yunnan apex; penis guide on outer edge broadened in apical part in lateral Province, Deqin County, China. view, pointed at apex; ventral surface of coxite with sclerotized Epiverta chelonia: Dieke 1943:169.—Pang and Mao 1979: 159; pocket antero-medially; apodeme of male sternum IX absent; meta- Jadwiszczak and We ˛ grzynowicz 2003: 208; Tomaszewska and ventral and abdominal postcoxal lines absent or sometimes poorly Szawaryn 2016: 47. developed postcoxal lines visible on metaventrite. Differential Diagnosis. Epiverta chelonia most closely resembles E. Distribution. Southwest China (including Tibet) (Fig. 3). supinata in its body shape and color pattern on the elytra, however Fig. 2. Habitus: (a, b, c) Epiverta supinata sp. nov., male—dorsal, lateral and anterior. (d, e, f): Epiverta supinata sp. nov., female—dorsal, lateral and anterior. (g, h, i) Epiverta angusta sp. nov., male—dorsal, lateral and anterior. (j, k, l) Epiverta angusta sp. nov., female—dorsal, lateral and anterior. (m, n, o) E. chelonia (Mader), male—dorsal, lateral and anterior. (p, q, r) E. chelonia (Mader), female—dorsal, lateral and anterior. (s, t, u) Epiverta albopilosa sp. nov., male–dorsal, lat- eral and anterior. (v, w, x) Epiverta albopilosa sp. nov., female—dorsal, lateral and anterior. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 4 Journal of Insect Science, 2017, Vol. 17, No. 2 Fig. 3. Distribution map of the genus Epiverta. E. chelonia can be separated from that species by having humeral subtruncate to weakly emarginate. Tergite  transverse subtruncate margins of the elytra simple (Fig. 4c and f) (at most weakly thick- to scarcely emarginate at apex. In females apical margin of ventrite ened but not upturned) and lateral margins of the pronotum evenly 5 weakly emarginate; ventrite 6 rounded apically; tergite VIII sub- arcuate (Fig. 4b). Moreover, the following combination of charac- truncate to scarcely excised apically. ters distinguish E. chelonia from its congeners: (1) penis guide not Male genitalia. Aedeagus stout. Tegmen (Fig. 5b and c) with par- longer than parameres, (2) tegminal strut more or less distinctly ameres scarcely longer than penis guide; tegminal strut truncate or emarginate at its apex, (3) anterior elytral maculae (near scutellum) emarginate apically; anterior margin of tegminal plate weakly pro- in form of weakly arcuate, most often uniformly yellow stripes or duced, emarginate or simply rounded medially (without pointed tri- sometimes with inner dark marking (Fig. 2m and p). angular projection medially). Penis guide with pointed apex almost straight or weakly curved backwards in lateral view (Fig. 5b). Penis with remnant of inner arm of reduced basal T-shaped capsule in Description. Length 5.55–7.55 mm; TL/EW ¼ 1.07–1.20; PL/ form of regular triangle (Fig. 5a). PW ¼ 0.38–0.42; EL/EW ¼ 0.88–1.01; TL/BH ¼ 2.36–2.70. Female genitalia (Fig. 5d) with proctiger (TX) weakly excised at Body regularly oval (Figs. 2m and p, 4f). Background of elytra apex; coxites with styli well developed; spermatheca weakly curved distinctly brownish black with regular yellow maculae arranged as in about half length; accessory gland about 2 times longer than in Fig. 2m and p. Pubescence moderately dense and moderately spermatheca. long, yellowish. Head (Fig. 4b) with interocular distance 0.68 times as wide as head across eyes. Antenna (Fig. 4a) with antennomere 3 about 1.25 Type Material. Lectotype of S. chelonia Mader, male, YUNNAN times as long as antennomere 4 and about 1.15 times as long as 5; PROVINCE, “Thibet. Atentse [¼Deqin]. (R. P. Goutelle)./Coll. antennomere 4 about 0.90 times as long as antennomere 5. Mus. Vindob./Fu ¨ r diese Art [.. .] jetzt eine neue Gattung gemacht Terminal maxillary palpomere about 1.12 times wider than long worden (hand writing)/S. chelonia Mad., det. Mader” (NHMV). (Fig. 4e). Paralectotype of S. chelonia Mader, female, “Thibet. Atentse Pronotum with lateral margins evenly arcuate (Fig. 4b). [¼Deqin]. (R. P. Goutelle)/Epilachna chelonia Fairm.” (MNHN). Prosternal process 0.62–0.74 times as wide as mesoventral process (pp/mp) (Fig. 4d); 0.80–0.97 times as wide as length of prosternum Other Material Examined. YUNNAN PROVINCE, “Yunnan in front of procoxa (pp/pl). Elytra with lateral flattened margins Province, Deqin County, 3,300 m, 2.IX.1987, Wang Shuyong leg./ 00 00 scarcely narrowing toward elytral apices (Fig. 4f); humeral margins IOZ(E) 626305 (*one: MIZ); same but IOZ(E) 626306 (*one: at most weakly thickened and raised (Fig. 4c); elytral total width IOZ); “Yunnan Province, Zhongdian County, Wegnshui Village, 1.35–1.48 times of elytral disc width (EW/ED); elytral epipleuron in 3,000 m, 10.VII.1982, Zhang Xuezhong leg./IOZ(E) 626243 (*one widest part 2.75–4.00 times as wide as epipleuron in narrowest part female: MIZ); “Yunnan Province, Deqin County, Meili Snow (eew/een), 2.65–3.40 times as wide as mesoventral process (eew/mp) Mountain, 3,350 m, 25.VII.1982, Zhang Xuezhong leg./IOZ(E) and 0.90–1.07 as wide as mesofemur length (Fig. 4g) (eew/mf). 626307 (*one male: IOZ); same but IOZ(E) 626308 (*one female: Abdomen. Hind margin of male ventrite 5 widely emarginate. MIZ); “CHINA: NW-Yunnan, 10 km SW Lijiang, 2,500 m, Hind margin of male ventrite 6 excised. Male tergite VIII 5.7.1994, leg. Schillhammer (13) (one female: MIZ, and two: Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 Journal of Insect Science, 2017, Vol. 17, No. 2 5 Fig. 4. E. chelonia (Mader): (a) antenna, (b) pronotum, (c) antero-lateral margin of elytron, (d) prosternum and mesoventrite, (e) mouthparts, (f) habitus, dorsal view, (g) habitus, ventral view. Fig. 5. E. chelonia (Mader): (a) penis, (b) tegmen, lateral, (c) tegmen, ventral, (d) female genitalia. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 6 Journal of Insect Science, 2017, Vol. 17, No. 2 Fig. 6. Epiverta albopilosa sp. nov.: (a) head and prothorax, ventral view, (b) mouthparts, (c) antenna, (d) pronotum, (e) habitus, dorsal view, (f) habitus, ventral view. NHMV); “Yunnan Province, Dali, Cangshan Mountain, 2,850 m, Differential Diagnosis. Epiverta albopilosa resembles E. angusta in 30.VI.1981, Wang Shuyong leg./IOZ(E) 626310 (*one male: MIZ); its dorsal coloration—background and maculae more irregular (as same but IOZ(E) 626311 (*one female: IOZ); “Yunnan Province, compared with E. chelonia and E. supinata) with background color Lijiang City, 3100 m, 27.V.1980, Wang Linyao leg./IOZ(E) 626285 of the elytra hard to distinguish (Fig. 2a and v). Both species are also (*one male: IOZ). SICHUAN PROVINCE, “China, S.W. Sichuan, similar in having white or greyish elytral pubescence. E. albopilosa, 0 00  0 00 Qingmai 3,000 m, N28 48 46.7 , E099 52 09.2 , 3.7.2006, lgt. however, can be distinguished from that species and also from E. Janata M./BMNH(E), 2006-158, M Janata/Epiverta sp., det. R.G. chelonia and E. supinata by: (1) antennomere 3 much longer than 4, Booth 2006 (female and male: BMNH). (2) dorsal surfaces covered with dense and comparatively long, white pubescence, (3) elytral lateral flattened margins widest in Distribution. Southwest China: NW Yunnan, SW Sichuan (Fig. 3). about 1/3 of elytral length thence narrowing towards elytral apices, (4) penis base of different shape (Fig. 7a). Epiverta albopilosa sp. nov Description. Length 6.10–7.73 mm; TL/EW ¼ 1.18–1.24; PL/ (Figs. 2s–x and 3, 6a–f, 7a–d) PW ¼ 0.41–0.42; EL/EW ¼ 0.99–1.04; TL/BH ¼ 2.40–3.00. (urn:lsid:zoobank.org:act:D00BEA06-441F-4757-A8C8- Body oval in males and elongate oval in females (Figs. 2s and v 8DF76B9FD0AE) and 6e). Dorsum bicoloured: brownish black and orange–yellow but Etymology. The name of the new species refers to white, dense pu- with background of elytra hard to distinguish; maculae irregular as bescence covering the body, especially its dorsal surfaces. in Fig. 2s and v. Pubescence dense, comparatively long, white. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 Journal of Insect Science, 2017, Vol. 17, No. 2 7 Fig. 7. Epiverta albopilosa sp. nov.: (a) penis, (b) tegmen, lateral, (c) tegmen, ventral, (d) female genitalia. Head (Fig. 6d) with interocular distance 0.70 times as wide as Type Material. Holotype, male, YUNNAN PROVINCE, “CHINA, head across eyes. Antenna (Fig. 6a and c)withantennomere 3 Yunnan Prov., Dongchuan; 19.VIII.1982; Xiao Ningnian leg.” about 1.80 times as long as antennomere 4 and about 1.38 times (SCAU). Paratypes, “CHINA, Yunnan Prov., Dongchuan; 19-VIII- as long as 5; antennomere 4 about 0.80 times as long as antenno- 1982; Xiao Ningnian leg.” (two males: SCAU; one male and one fe- mere 5. Terminal maxillary palpomere about as wide as long male: MIZ); “Yunnan Province, Luquan County, Jiaozishan (Fig. 6b). Mountain, 2750m, 23-VIII-2013, Huo Lizhi leg.” (one female: SCAU). Pronotum with lateral margins evenly arcuate (Fig. 6d). Prosternal process 0.62–0.80 times as wide as mesoventral pro- Distribution. Southwest China: NE Yunnan (Fig. 3). cess (pp/mp) (Fig. 6a); 0.82–0.96 times as wide as length of pros- ternum in front of procoxa (pp/pl). Elytra with lateral flattened Epiverta angusta sp. nov margins gradually narrowing from about 1/3 of elytral length to- (Figs. 2g–l, 3, 8a–f, 9a–d) ward their apices (Fig. 6e) humeral margins simple; elytral total (urn:lsid:zoobank.org:act:D32E2A64-A9AB-4362-A7F0- width 1.22–1.32 times of elytral disc width (EW/ED); elytral epi- BFFDA632C62E) pleuron in widest part 3.05–3.85 times as wide as epipleuron in Epiverta chelonia: Bielawski 1960: 441 (misidentification). narrowest part (eew/een), 2.70–3.20 times as wide as mesoventral process (eew/mp) and 0.70–0.82 as wide as mesofemur length Etymology. The name of the new species in Latin means narrow and (eew/mf) (Fig. 6f). refers to flattened lateral margins of the elytra which are compara- Abdomen. Hind margin of male ventrite 5 truncate to weakly tively narrow throughout of their length. emarginate. Hind margin of male ventrite 6 weakly emarginate. Male tergite VIII scarcely emarginate. Tergite  transverse truncate Differential Diagnosis. Epiverta angusta is most similar to E. albopilosa at apex. In females apical margin of ventrite 5 truncate; ventrite 6 in its dorsal coloration with the background and maculae more irregular subtruncate apically; tergite VIII rounded to subtruncate apically. (as compared with E. chelonia and E. supinata) with background color Male genitalia. Aedeagus stout. Tegmen (Fig. 7b and c)with of the elytra hard to distinguish (Fig. 2d and j). Both species are also cov- parameres slightly shorter than penis guide; tegminal strut ered with white or greyish elytral pubescence (yellowish in E. chelonia rounded apically; anterior margin of tegminal plate weakly pro- and E. supinata). E.angusta, however, can be distinguished from E. albo- duced, simply rounded medially or scarcely emarginate medially. pilosa by: (1) lateral elytral flattened margins comparatively narrow Penis guide with pointed apex distinctly upturned backwards (vis- throughout of their length, (2) antennomere 3 about 1.25 times longer ible in lateral view) (Fig. 7b). Penis with remnant of inner arm of than4(1.8times longer in E. albopilosa), (3) antennomeres 3 and 5 sub- reduced basal T-shaped capsule comparatively large, irregularly equal in length (antennomere 3 about 1.38 times longer than 5 in E. albo- triangular (Fig. 7a). pilosa), (4) different shape of penis base and spermatheca. Female genitalia (Fig. 7d) with proctiger (TX) scarcely excised at apex; coxites with styli well developed; spermatheca variable, shorter weakly curved near base or longer curved near mid length; Description. Length 5.62–7.30 mm; TL/EW ¼ 1.11–1.27; PL/ accessory gland about 3 times longer than spermatheca. PW ¼ 0.39–0.42; EL/EW ¼ 1.00–1.10; TL/BH ¼ 2.27–2.74. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 8 Journal of Insect Science, 2017, Vol. 17, No. 2 Fig. 8. Epiverta angusta sp. nov.: (a) pronotum and base of elytra, (b) antero-lateral margin of elytron, (c) mouthparts, (d) pro- and mesothorax, ventral view, (e) habitus, dorsal view, (f) habitus, ventral view. Body oval in males and elongate oval in females (Figs. 2g and j (eew/mp) and 0.83–1.00 as wide as mesofemur length (eew/mf) and 8e). Dorsum bicoloured: brownish black and orange-yellow (Fig. 8f). butwithbackgroundof elytrahardtodistinguish;maculae irreg- Abdomen. Hind margin of male ventrite 5 distinctly emarginate. ular as in Fig. 2g and j. Pubescence moderately dense and long, Hind margin of male ventrite 6, tergite VIII and tergite X with greyish. scarce emargination/excision at apex. In females apical margin of Head (Fig. 8a) with interocular distance 0.70 times as wide as ventrite 5 and tergite VIII subtruncate apically; ventrite 6 subtrun- head across eyes. Antenna (Fig. 8d) with antennomere 3 about 1.25 cate to weakly triangularly angulate (with hind margin weakly proj- times as long as antennomere 4 and about 1.04 times as long as 5; ected backwards medially). antennomere 4 about 0.83 times as long as antennomere 5. Male genitalia. Aedeagus stout. Tegmen (Fig. 9b and c) with par- Terminal maxillary palpomere about as wide as long (Fig. 8c). ameres slightly shorter than penis guide; tegminal strut rounded api- Pronotum with lateral margins evenly arcuate (Fig. 8a). cally; anterior margin of tegminal plate not produced, simply Prosternal process 0.62–0.73 times as wide as mesoventral pro- rounded medially. Penis guide with pointed apex weakly upturned cess (pp/mp) (Fig. 8d); 0.86–1.04 times as wide as length of pros- backwards (visible in lateral view) (Fig. 9b). Penis with remnant of ternum in front of procoxa (pp/pl). Elytra with lateral flattened inner arm of reduced basal T-shaped capsule comparatively small, margins comparatively narrow, scarcely narrowing toward elytral irregularly triangular (Fig. 9a). apices (Fig. 8e); humeral margins simple; elytral total width 1.27– Female genitalia (Fig. 9d) with proctiger (TX) subtruncate or 1.35 times of elytral disc width (EW/ED); elytral epipleuron in sometimes weakly emarginated at apex; coxites with styli well devel- widest part 3.15–3.77 times as wide as epipleuron in narrowest oped; spermatheca curved in mid length; accessory gland 2.5–2.8 part (eew/een), 2.90–3.73 times as wide as mesoventral process times as long as than spermatheca. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 Journal of Insect Science, 2017, Vol. 17, No. 2 9 Fig. 9. Epiverta angusta sp. nov.: (a) penis, (b) tegmen, lateral, (c) tegmen, ventral, (d) female genitalia. Type Material. Holotype, male, SICHUAN PROVINCE, “Ta-Tsien- recognize these specimens as belonging to the new species, described Lu, 30.V.93, Potanin (translated and transcripted here from here as E. angusta. Russian, hand-writting)/E. chelonia (Mader), det. R. Bielawski 1959)/Inst. Zool. P.A.N, Warszawa, 86/60” (MIZ). Paratypes, “Fu- Distribution. Southwest China: SW Sichuan (Fig. 3). dian-xo (translated and transcripted here from Russian, hand writ- ing)/K. A. Semenowa/E. chelonia (Mader), det. R. Bielawski 1958)/ Epiverta supinata sp. nov Inst. Zool. P.A.N, Warszawa, 95/59” (one male: MIZ); “Thibet, (Figs. 2a–f, 3, 10a–g, 11a–d) Chasseurs de, T a-tsien-lou, ^ 1895” (two males, two females and 13: (urn:lsid:zoobank.org:act:0BC7054D-3368-422D-9D97- MNHN; 2: MIZ); “Chasseurs Indige ` nes, des Missionnaires, de Ta- EBAC94345B88) tsien-Lou,  1906” (one male, one female and nine: MNHN); “Sichuan Province, Kangding County, 2,400 m, 1963.VII.13, Mao Jinlong leg.” (one male: IOZ); “Thibet, Tatsienlou, mgr F. Birt/ Etymology. The name of the new species is derived from Latin Museum Paris, ex Coll., R. Oberthu ¨ r” (two: MNHN; 1: MIZ); “Su- “supinatas”, which means upturned and refers to humeral margins Tchuen, Mo-Sy-Mien, 1897” (two males, four females and 12: of the elytra upturned backwards. MNHN; 3: MIZ); “Su-Tchuen, Chass. Thibetains, 1897” (two: MNHN); same but 1899 (one: MNHN; one: MIZ); “CHINA: Differential Diagnosis. Epiverta supinata can be easily separated Sichuan Prov., 1999, Zeng Tao leg.” (two males, two females: from its congeners by having humeral margins of the elytra thick- SCAU; one male and one female: MIZ). ened and upturned backwards (Fig. 10c and f), lateral margins of the pronotum subparallel along basal 2/3 length, antennomere 4 and Comment. Bielawski (1960) studied Epiverta material from “der 5 subequal in length, female genitalia with comparatively short styli Sammlung des Zoologischen Institut der Akademie der and anterior margin of tegminal plate regularly, triangularly pro- Wissenschafte der UdSRR in Leningrad (now Saint Petersburg). He duced medially. studied 14 specimens from Sichuan Province (Ta-Tsien-Lu and Mao-nju-ku-Tal) and treated them as E. chelonia. He also provided Description. Length 6.10–6.30 mm; TL/EW ¼ 1.03–1.17; PL/ drawings of the male genitalia of this species unknown until then. PW ¼ 0.38–0.43; EL/EW ¼ 0.90–0.98; TL/BH ¼ 2.11–2.60. After study of two specimens from that series (retained in the MIZ Body oval (Figs. 2a and d and 10f). Background of elytra brown- collection), his pictures of the male genitalia (Bielawski 1960) (with ish black with regular yellow maculae arranged as in Fig. 2a and d, tegmen with parameres slightly shorter than penis guide, tegminal most of them with inclusions of blackish spots. Pubescence moder- strut rounded apically, anterior margin of tegminal plate not pro- ately dense and moderately long, yellowish. duced and simply rounded medially, and penis guide with pointed Head (Fig. 10b) with interocular distance 0.71 times as wide as apex weakly upturned backwards in lateral view) and the drawings head across eyes. Antenna (Fig. 10e) with antennomere 3 about 1.10 of elytral pattern commented by him, that “Die Flecken auf den times as long as antennomere 4 and about 1.15 times as long as 5; Flu ¨ geldecken verbindensich miteinender auf sehr verschidene Weise antennomere 4 about as long as antennomere 5. Terminal maxillary und bilden eine Mosaik Schwarz und braungrau gef€ arbt”, we palpomere about as wide as long (Fig. 10e). Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 10 Journal of Insect Science, 2017, Vol. 17, No. 2 Fig. 10. Epiverta supinata sp. nov.: (a) pronotum and base of elytra, (b) pronotum, (c) antero-lateral margin of elytron, (d) prosternum and mesoventrite, (e) mouthparts, (f) habitus, dorsal view, (g) habitus, ventral view. Female genitalia (Fig. 11d) with proctiger (TX) scarcely to Pronotum with lateral margins subparallel along basal 2/3 length then arcuate anteriorly (Fig. 10a and b). Prosternal process 0.65– weakly excised medially at apex; coxites with styli small to well de- 0.80 times as wide as mesoventral process (pp/mp) (Fig. 10d); 0.83– veloped; spermatheca curved in mid length; accessory gland 2.5–3.0 1.15 times as wide as length of prosternum in front of procoxa (pp/ times longer than spermatheca. pl). Elytra with lateral flattened margins scarcely narrowing towards elytral apices (Fig. 10f); humeral margins thickened and upturned Type Material. Holotype, male, SICHUAN PROVINCE, “Thibet, backwards (Fig. 10c and f); elytral total width 1.49–1.60 times of Yare ´ gong [Sichuan], P. Soulie ´ , 1900” (MNHN). Paratypes, same elytral disc width (EW/ED); elytral epipleuron in widest part 2.70– data as holotype (one male, two females and two: MNHN; two: 3.70 times as wide as epipleuron in narrowest part (eew/een), 3.90– MIZ); “Sichuan Province, Xiangcheng County, 2900 m, 4.80 times as wide as mesoventral process (eew/mp) and 1.10–1.30 17.VI.1982, Wang Shuyong leg./IOZ(E) 626295” (*one: IOZ); as wide as mesofemur length (eew/mf) (Fig. 10g). same but IOZ(E) 626296 and IOZ(E) 626297” (*one and male, re- Abdomen. Hind margin of male ventrite 5 emarginate. Hind spectively: IOZ); “Sichuan Province, Xiangcheng County, 3200 m, margin of male ventrite 6 weakly excised; tergite VIII and tergite X 26.VI.1982, Chai Huaicheng leg./IOZ(E) 626293” (*one: IOZ); with weakly emarginate at apex. In females apical margin of ventrite same but IOZ(E) 626294 (*one female: MIZ). TIBET, “Tibet, 5 truncate; ventrite 6 subtruncate to weakly rounded apically; tergite Mangkang County, Haitong, 3250 m, 12.VIII.1982, Wang Shuyong VIII subtruncate to scarcely emarginate apically. leg./IOZ(E) 626298” (male: *IOZ). “Tibet, Mangkang County, Male genitalia. Aedeagus stout. Tegmen (Fig. 11b and c) with Haitong 3200 m, 9.VIII.1982, Wang Shuyong leg./IOZ(E) 626301” parameres slightly shorter than penis guide; tegminal strut rounded (*one: MIZ); same data but IOZ(E) 626302” (*one female: MIZ); apically; anterior margin of tegminal plate triangularly produced “Tibet, Mangkang County, Haitong 3250 m, 11.VIII.1982, Wang medially. Penis guide with pointed apex weakly upturned back- Shuyong leg./IOZ(E) 626300” (*one male: IOZ); same data but wards (visible in lateral view) (Fig. 11b). Penis with remnant of inner IOZ(E) 626303” (*one female: IOZ); “Tibet, Mangkang County, arm of reduced basal T-shaped capsule most often trigger like Haitong 3250 m, 12.VIII.1982, Wang Shuyong leg./IOZ(E) (Fig. 11a). 626259” (*one: IOZ); same data but IOZ(E) 626299” (*one: IOZ). Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 Journal of Insect Science, 2017, Vol. 17, No. 2 11 Fig. 11. Epiverta supinata sp. nov.: (a) penis, (b) tegmen, lateral, (c) tegmen, ventral, (d) female genitalia. YUNNAN PROVINCE, “Yunnan Province, Deqin County, Adong – Body covered with greyish pubescence (Fig. 2g–l); elytra 1.00– Village, 2,700–3,000 m, 6.IX.1981, Wang Shuyong leg./IOZ(E) 1.10 times as long as wide; total width of elytra 1.27–1.35 times as 626255” (*one male: IOZ); same but IOZ(E) 626256 and IOZ(E) wide as elytral disc; antennomere 3 about as long as antennomere 5; 626257 (*two: IOZ). penis guide slightly longer parameres (Fig. 9b); tegminal strut rounded apically (Fig. 9c) ................................. . E. angusta sp. nov. Distribution. Southwest China: E Tibet, NW Yunnan, SW Sichuan (Fig. 3). Acknowledgments We thank Roger Booth (BMNH), Antoine Mantilleri (MNHN), Harald Schillhammer and Rudolf Schuh (NHM) and Kuiyan Zhang (IOZ) for the loan of specimens used in this study. Adam Slipinski  (CSIRO) read an early Key to Species of Epiverta draft of this manuscript and provided helpful suggestions. Magdalena 1. Lateral margins of pronotum subparallel along basal 2/3 Kowalewska-Groszkowska (MIZ) helped with SEM images. length (Fig. 10a and b); humeral margins of elytra thickened and up- turned backwards (Fig. 10c, f); total width of elytra 1.49-1.60 times References Cited as wide as elytral disc; elytral epipleuron in widest part 1.10-1.30 Bielawski, R. 1960. Materialien zur Kenntnis der Coccinellidae (Coleoptera). times of mesofemur length ............................... E. supinata sp. nov. Ann. Zool. 18: 435–458. – Lateral margins of pronotum regularly at least weakly arcuate; Bocak, L., C. Barton, A. Crampton-Platt, D. Chesters, D. Ahrens, and A. humeral margins of elytra simple and not upturned backwards; total Vogler. 2014. Building the Coleoptera tree-of-life for >8000 species: com- width of elytra 1.22–1.48 times as wide as elytral disc; elytral epi- position of public DNA data and fit with Linnaean classification. Syst. pleuron in widest part 0.70–1.07 times of mesofemur length ........ 2 Entomol. 39: 97–110. 2. Elytra covered with dense and comparatively long, white pu- Crowson, R. A. 1955. The natural classification of the families of Coleoptera, bescence (Fig. 2s and v); antennomere 3 about 1.8 times as long as 187 pp. Nathaniel Lloyd, London. antennomere 4 (Fig. 6c); elytral epipleuron in widest part 0.70-0.82 Dieke, G. H. 1947. Ladybeetles of the genus Epilachna (sens. lat.) in Asia, Europe and Australia. Smithsonian Miscellaneous Collect. 106: 1–183. times of mesofemur length (Fig. 6f); .............. E. albopilosa sp. nov. Jadwiszczak, A., and P. We˛grzynowicz. 2003. World Catalogue of – Elytra covered with sparser and shorter, yellowish or greyish Coccinellidae. Part I—Epilachninae, 264 pp. Mantis, Olsztyn. pubescence; antennomere 3 about 1.25 times as long as antenno- Kovar, I. 2007. Family Coccinellidae Latreille, 1807, pp. 568–631. In I., Lobl mere 4; elytral epipleuron in widest part 0.83–1.07 times of mesofe- & A., Smetana (Ed.), Catalogue of Palaearctic Coleoptera, Vol. 4. mur length .................................................................................... 3 Stenstrup, Apollo Books, 935 pp. 3. Body covered with yellowish pubescence (Fig. 2m–r); elytra at Lord, N., C. S. Hartley, J. F. Lawrence, J. V. McHugh, M. F. Whiting, and K. most as long as wide; total width of elytra 1.35–1.48 times as wide as B. Miller. 2010. Phylogenetic analysis of the minute brown scavenger bee- elytral disc; antennomere 3 about 1.15 times as long as antennomere 5; tles (Coleoptera: Latridiidae), and recognition of a new beetle family, penis guide at most as long as parameres (Fig. 5b); tegminal strut emar- Akalyptoischiidae fam. n. (Coleoptera: Cucujoidea). Syst. Entomol. 35: ginate or truncate apically (Fig. 5c) ....................... E. chelonia (Mader) 753–763. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018 12 Journal of Insect Science, 2017, Vol. 17, No. 2 Mader, L. 1933. Uber bekannte und neue Coccinelliden. Entomologischer Szawaryn, K., and W. Tomaszewska. 2013. Two new genera of Epilachnini Anzeiger, Wien. 13: 79–84. Mulsant from New Guinea and Aru Islands (Coleoptera: Coccinellidae). Pang, X. F., and J. L. Mao. 1979. Economic Insect Fauna of China (XIV) , 170 J. Nat. Hist. doi:10.1080/00222933.2012.763067. pp. Coleoptera: Coccinellidae II. Science Press, Beijing Szawaryn, K., L. Bocak, A. Slipinski,  H. E. Escalona, and W. Tomaszewska. Robertson, J. A., F. Whiting, and J. V. McHugh. 2008. Searching for natural 2015. Phylogeny and evolution of phytophagous ladybird beetles lineages within the Cerylonid Series (Coleoptera: Cucujoidea). Mol. (Coleoptera: Coccinellidae: Epilachnini), with recognition of new genera. Phylogenet. Evol. 46: 193–205. Syst. Entomol. 40: 547–569. Robertson, J., A. Slipinsk  i, M. Moulton, F. W. Shockley, A. Giorgi, N. P. Slipinski,  S. A., and W. Tomaszewska. 2010. Coccinellidae Latreille, 1802, Lord, D. D. McKenna, W. Tomaszewska, J. Forrester, K. B. Miller, et al. pp. 454–72. In: R.A.B., Leschen, R.G, Beutel and J.F. Lawrence (ed.): 2015. Phylogeny and classification of Cucujoidea and the recognition of a Handbook of Zoology, Vol. 2, Coleoptera. Berlin/New York: Walter de new superfamily Coccinelloidea (Coleoptera: Cucujiformia). Syst. Entomol. Gruyter GmbH & Co. KG. XIII þ 786 pp. 40: 745–778. Tomaszewska, W., and K. Szawaryn. 2013. Revision of the Asian species of Seago, A. E., A. Giorgi, J. Li, and A. Slipinski.  2011. Phylogeny, classification Afidentula Kapur, 1958 (Coleoptera: Coccinellidae: Epilachnini). Zootaxa. and evolution of ladybird beetles (Coleoptera: Coccinellidae) based on si- 3608: 26–50. multaneous analysis of molecular and morphological data. Mol. Tomaszewska, W., and K. Szawaryn. 2016. Epilachnini (Coleoptera: Phylogenet. Evol. 60: 137–151. Coccinellidae)—a revision of the world genera. J. Insect Sci. 16: 101. 1–91. Downloaded from https://academic.oup.com/jinsectscience/article-abstract/17/2/60/3739027 by Ed 'DeepDyve' Gillespie user on 17 July 2018

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