A comparative study of the external male genitalia in the subfamily Zeugophorinae, and a new species of genus Zeugophora (Coleoptera, Megalopodidae) from China

277 A comparative study of the external male genitalia in the subfamily Zeugophorinae, and a new species of genus Zeugophora (Coleoptera, Megalopodidae) from China Kai-Qin Li1,2 , Yuan Xu3, Hong-Bin Liang4 1 Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650223, China 2 Yunnan Key Laboratory of Biodiversity Information, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650201, China 3 College of Forestry, Southwest Forestry University, Kunming 650224, China 4 Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China Corresponding author: Hong-Bin Liang ([email protected]) Copyright: © Kai-Qin Li et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract The external male genitalia of 21 Chinese species of the genus Zeugophora (subfamily Zeugophorinae) are compared. A key is compiled to Chinese Zeugophora species based on external male genitalia with aid of other external morphological traits. One new species, Zeugophora (Pedrillia) liupanshanensis Li & Liang, sp. nov., is described from Ningxia Hui Autonomous Region, China. Key words: Key, median lobe, median struts, Pedrillia, spiculum, tegmen Introduction The external male genitalia of Megalopodidae consist of a median lobe, a pair of median struts, a tegmen, an internal sac and a spiculum (belonging to the 9th abdominal segment) (Snodgrass 1935; Chen 1940; Verma 1996). They are among the most important identification characteristics in the Megalopodidae and receive much attention from taxonomists. Chûjô (1952, 1953) studied the morphology of external male genitalia of the Megalopodinae and Zeugophorinae extensively and found similarities between these two subfamilies. Subsequently, taxonomists began to pay attention to the male genitalia of the Megalopodidae when describing new species and constructing phylogenetic relationships within the Chrysomeloidea (Reid 1989, 1998; Kuschel and May 1990; Mann and Crowson 1996; Verma 1996; Medvedev 1997; Li et al. 2013; Li and Liang 2018, 2020; Rodríguez-Mirón et al. 2021). The current classification of the genus Zeugophora is unstable. Some authors treated Pedrillia Westwood, 1864 as a distinct genus of the Zeugophorinae (Baly 1873; Jacoby 1908; Chûjô 1935, 1937) or as a subgenus of Zeugophora (Monrós 1959; Gressitt and Kimoto 1961; Chen and Pu 1962; Kimoto and Gressitt 1979; Medvedev 1985, 1997; Reid 1989, 1992, 1995; Schöller 2009; Silfverberg 2010; Warchałowshi 2010; Li and Liang 2018, 2020), while other authors treated Pedrillia as a synonym of the genus ZeuAcademic editor: Ron Beenen Received: 19 August 2025 Accepted: 6 November 2025 Published: 2 December 2025 ZooBank: https://zoobank. org/08B2CC39-AB8A-4100-B77A5907862AB3C4 Citation: Li K-Q, Xu Y, Liang H-B (2025) A comparative study of the external male genitalia in the subfamily Zeugophorinae, and a new species of genus Zeugophora (Coleoptera, Megalopodidae) from China. ZooKeys 1261: 277–306. https://doi. org/10.3897/zookeys.1261.169189 ZooKeys 1261: 277–306 (2025) DOI: 10.3897/zookeys.1261.169189 278 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae gophora (Sekerka and Vives 2013; Rodríguez-Mirón, 2018; Takemoto, 2019; Rodríguez-Mirón et al. 2021). We accept that Pedrillia is a subgenus of the genus Zeugophora because Pedrillia can be separated from Zeugophora on external morphology, external male and female genitalia, and host plants, at least for Chinese species. A total of 34 species of subfamily Zeugophorinae has been recorded in China (Lee and Cheng 2007; Li and Liang 2018, 2020). In Pedrillia, it is difficult to determine certain species based only on external morphological characteristics; therefore, we attempted to find characteristics of external male genitalia to aid its classification. The comparison of female genitalia was not included in this study owing to insufficient specimen material in several species. In this study, we describe and compare 21 available Chinese species of Zeugophorinae to summarize key characteristics of external male genitalia for distinguishing subgenera and species within the Chinese fauna. With accumulating information about the host plants of Zeugophorinae during expeditions in China, we found that plants of the family Celastraceae are hosted by a few species, such as Zeugophora (Pedrillia) annulata (Baly, 1873), Z. (P. ) bicolor (Kraatz, 1879), Z. (P. ) nigricollis (Jacoby, 1885), Z. (P. ) ruficollis Chûjô, 1932, Z. (P. ) decorata Chûjô, 1937, Z. (P. ) formosana (Gressitt, 1945), Z. (P. ) tricolor Chen & Pu, 1962, Z. (P. ) euonymorum Li & Liang, 2020 (Lee and Cheng 2007; Li and Liang 2018, 2020). In recent years, we focused on trees of Celastraceae when surveying Zeugophorinae. In 2018, two specimens were collected by beating plants of the family Celastraceae in Ningxia Hui Autonomous Region. We compared them with all known species, and found they represented a new species, described in this paper. Materials and methods Preparation of specimens Dry specimens were soaked in water for 1–2 h. The lateral margin of the abdomen was opened and the genitalia were pulled out of the abdomen with fine forceps. The genitalia were soaked in a warm solution of 10% KOH for 10–20 min. After treatment, these organs were washed with water. The genitalia were then detached and transferred to glycerin for observation, photography, and preservation. All measurements were made using a Nikon SMZ1500 or Nikon SMZ18 stereoscopic microscope with the aid of an ocular micrometer. Body length (BL) = the linear distance along the midline from the anterior margin of the labrum to the apex of the elytra; body width (BW) = elytra width (EW) = the maximum linear distance across the elytra; pronotum length (PL) = the linear distance along the median line of the pronotum; pronotum width (PW) = the linear distance across the widest part of the pronotum; elytra length (EL) = the linear distance from the base of the elytra to the apex of the sutural angle; median lobe length = the linear distance from the base to the apex; median struts length = the linear distance from the base to the apex. Ratios cited in descriptions are based on these measurements. Photographs of male genitalia were taken using a Nikon SMZ-1500 stereoscopic dissecting microscope fitted with a Cannon 450D digital camera, or a 279 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Nikon SMZ18 stereoscopic dissecting microscope fitted with a Nikon D610 digital camera. For each final image, several photographs were taken using different focal planes, combined with Helicon Focus software to obtain one synthesized photograph, and finally edited with Adobe Photoshop software. Terminology Morphological terminology for male genitalia follows Snodgrass (1935), Verma (1996), Chûjô (1952, 1953), Kasap and Crowson (1985), Li and Liang (2018, 2020), and Li et al. (2013). Abbreviations and acronyms The abbreviation TL means type locality. The abbreviation TD means type depository. The abbreviations used in the descriptions are as follows: BFU Beijing Forestry University, Beijing, China. CAS California Academy of Sciences, San Francisco, California, USA. IZCAS National Zoological Museum of China, Institute of Zoology, Chinese Academy of Sciences, Beijing, China. KIZ Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China. MZPW Polish Academy of Science, Museum of the Institute of Zoology, Warsaw, Poland. NHMUK The Natural History Museum, London, UK. NSMK National Science Museum, Korea. SCBG Shanghai Chenshan Botanical Garden, Shanghai, China. SDEI Deutsches Entomologisches Institut, Eberswalde, Germany. ZIN Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia. Species studied A total of 21 available Zeugophora (subgenera Zeugophora and Pedrillia) species belonging subfamily Zeugophorinae were examined (Table 1). Results Key to species of the genus Zeugophora 1 Median strut short, ratio of median strut / median lobe ≤ 1.5 (subgenus Zeugophora) ...................................................................................................2 – Median strut long, ratio of median strut / median lobe ≥ 2.0 (subgenus Pedrillia) .........................................................................................................7 2 Median lobe with apical portion gradually narrowed towards apex, apex blunt or sharp (Figs 2, 18, 23); host plant genus Populus ........................... 3 – Median lobe with apical portion abruptly narrowed towards apex, apex sharp (Figs 7, 10, 15); host plant genus Salix ..............................................5 280 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae 3 Median lobe with apical portion broad, apex blunt (Fig. 2) .......................... ......................................................................Zeugophora (Z.) ancora Reitter – Median lobe with apical portion narrow, apex sharp (Figs 18, 23).............4 4 Dorso-central portion of median lobe without a sclerite (Fig. 18) ............... ..............................................................Zeugophora (Z.) scutellaris Suffrian – Dorso-central portion of median lobe with a sclerite (Fig. 23) ..................... .......................................................................Zeugophora (Z.) turneri Power 5 Apical portion of median lobe slightly broad (Fig. 7) .................................... ....................................................................... Zeugophora (Z.) cribrata Chen – Apical portion of median lobe slightly narrow (Figs 10, 15) .......................6 6 Apex of median lobe slightly long (Fig. 15) ................................................... ................................................................ Zeugophora (Z.) nigroarea Lopatin – Apex of median lobe slightly short (Fig. 10) .................................................. ........................................................................ Zeugophora (Z.) cyanea Chen 7 Sides of the median lobe narrow in basal and apical portion (Figs 39, 65) ........................................................................................................................8 – Sides of median lobe parallel or sub-parallel ..............................................9 8 Median basal portion lobe broader, apex strongly narrowed and bent downward;median strut short; tegmen strongly sclerotized (Figs 38–40) . .................................................................Zeugophora (P. ) emeica Li & Liang – Median lobe elongate, apex slightly swollen laterally; median strut long; tegmen slightly sclerotized (Figs 64–66) ...................................................... ...........................................................Zeugophora (P. ) nigroapica Li & Liang 9 Sides of median lobe parallel; dorso-central portion of median lobe with a sclerite (Figs 34, 73) ....................................................................................10 – Sides of median lobe parallel, sub-parallel or not parallel; dorso-central portion of median lobe without a sclerite ..................................................11 10 Sclerite of median lobe in dorso-central area weak; sides of spiculum longer than central portion (Figs 34, 36) ............................................................. ................................................................ Zeugophora (P. ) dimorpha Gressitt – Sclerite of median lobe in dorso-central area strong; sides of spiculum shorter than central portion (Figs 72, 75) ...................................................... ............................................................Zeugophora (P. ) trifasciata Li & Liang 11 Median lobe with dorso-central portion membranous, basal 1/3 not tubular; the sides of median lobe thin (Figs 47, 58, 82); host plant Symplocaceae ......................................................................................12 – Median lobe with dorso-central portion membranous, basal 1/3 tubular; sides of median lobe thick; host plant Lamiaceae or Celastraceae ........14 12 Ratio of median strut / median lobe ~3.0; apex of median lobe sharp (Figs 81, 82); apical portion of spiculum trifid ............................................... ...........................................................Zeugophora (P. ) yunnanica Chen & Pu – Ratio of median strut / median lobe 1.5–2.0; apex of median lobe blunt or prominent; apical portion of spiculum not trifid (Figs 46, 47, 57, 58) ......13 13 Median lobe apical portion slightly broad, apical margin projecting and flattened (Fig. 47) ............................. Zeugophora (P. ) flavithorax Li & Liang – Median lobe apical portion slightly narrow, apical margin blunt and bent downward medially (Fig. 58) .................... Zeugophora (P. ) maculata Chûjô 281 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae 14 Ratio of median strut / median lobe > 3.0; host plant Lamiaceae ...........15 – Ratio of median strut / median lobe < 3.0; host plant Celastraceae ........17 15 Median lobe strongly curved in lateral view, apical portion strongly upward curved, apex sharp (Figs 54, 55) ....Zeugophora (P. ) longicornis (Westwood) – Median lobe curved in lateral view, apical portion flattened, apex blunt ..... ......................................................................................................................16 16 Median lobe elongate, sides slightly narrowed in middle, apex slightly short (Figs 76, 77) ......................................Zeugophora (P. ) yuae Li & Liang – Median lobe slightly broad and short, sides paralleled, apex slightly long (Figs 49, 50) ..................................................Zeugophora (P. ) indica Jacoby 17 Median lobe basal portion curved in lateral view (Figs 41, 60, 92, 97) ....18 – Median lobe basal and apical portion curved in lateral view (Figs 25, 30, 69) ...........................................................................................20 18 Media lobe short, apex broad (Figs 41, 42) ................................................... .......................................................Zeugophora (P. ) euonymorum Li & Liang – Median lobe elongate, apex narrow ...........................................................19 19 Apical portion of median lobe broad (Fig. 61) ............................................... ..............................................................Zeugophora (P. ) nigricollis (Jacoby) – Apical portion of median lobe narrow (Figs 93, 98) ...................................... ..................................................... Zeugophora (P. ) liupanshanensis sp. nov. 20 Apex of median lobe slightly curved Leftward (Fig. 31), basal piece of tegmen long (Fig. 32) ..................................... Zeugophora (P. ) bicolor (Kraatz) – Apex of median lobe not curved Leftward (Figs 26, 70), basal piece of tegmen short (Figs 27, 71) .........................................................................21 21 Apex of the median lobe long and sharp (Fig. 26) ........................................ .....................................................................Zeugophora (P. ) annulata (Baly) – Apex of median lobe short and blunt (Fig. 70) .............................................. ................................................................Zeugophora (P. ) tricolor Chen & Pu Zeugophorinae Boving & Craighead, 1931 Genus Zeugophora Kunze, 1818 Subgenus Zeugophora Kunze, 1818 External male genitalia. (Figs 1–24) Median lobe generally strongly sclerotized, slender, dorso-central portion membranous with or without a long narrow sclerite, curved in lateral view, sides thin, sub-parallel, apex narrower than base, apical portion tongue shaped, blunt, or triangular and sharp; median struts rodshaped, widely separated from each other, ~1.1–1.4× as long as median lobe; base of tegmen slightly Y-shaped, tegminal ring with base broad and gradually narrow to apical portion, apex sub-square, paramere sub-square, apical margin with dense setae; endophallus membranous, with a paired granulated area, sclerotized or weakly sclerotized. Spiculum long, Y-shaped, strongly sclerotized, apical portion trifid, slightly sclerotized. The apical portion of median lobe with a long narrow sclerite in Zeugophora (Zeugophora) ancora, Z. (Z.) cribrata, Z. (Z.) cyanea, Z. (Z.) nigroaerea, and Z. (Z.) turneri (Figs 2, 7, 10, 15, 23) but not in Z. (Z.) scutellaris (Fig. 18). 282 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Table 1. Species list of Zeugophorinae studied in this paper. Species Locality Specimen status Host plant Zeugophora (Zeugophora) ancora Reitter, 1900 Ningxia common Salicaceae: Populus nigra var. italica (Moench) Koehne, Populus alba L. Zeugophora (Z.) cribrata Chen, 1974 Qinghai paratype Salicaceae: Salix L. Zeugophora (Z.) cyanea Chen, 1974 Sichuan, Qinghai specimen compared with holotype Salicaceae: Salix L. Zeugophora (Z.) nigroaerea Lopatin, 2008 Gansu, Shaanxi specimen compared with holotype Salicaceae: Salix L. Zeugophora (Z.) scutellaris Suffrian, 1840 Inner Mongolia common Salicaceae: Populus acuminata Rydberg, P. alba L., P. deltoides Marshall, P. grandidentata Michaux, P. nigra L., P. tremula L., Salix sp. Zeugophora (Z.) turneri Power, 1863 Beijing common Salicaceae: Populus sp. Zeugophora (Pedrillia) annulata (Baly, 1873) Ningxia common Celastraceae: Euonymus alatus (Thunb.) Siebold, Euonymus przewalskii Maxim., Euonymus phellomanus Loes. ex Diels. Euonymus maacki Rupr. and E. sacrosancta Koidz. Zeugophora (P. ) bicolor (Kraatz, 1879) Liaoning specimen compared with holotype Celastraceae: Celastrus orbiculatus Thunb. Zeugophora (P. ) dimorpha Gressitt, 1945 Hunan specimen compared with paratype unknown Zeugophora (P. ) emeica Li & Liang, 2018 Sichuan holotype unknown Zeugophora (P. ) euonymorum Li & Liang, 2020 Yunnan holotype Celastraceae: Euonymus alatus (Thunb.) Siebold, Euonymus hamiltonianus Wall. Zeugophora (P. ) flavithorax Li & Liang, 2020 Yunnan holotype Symplocaceae Zeugophora (P. ) indica Jacoby, 1903 Yunnan specimen compared with holotype possible Lamiaceae: Vitex quinata (Lour.) Will. Zeugophora (P. ) longicornis (Westwood, 1864) Yunnan specimen compared with type possible Lamiaceae: Vitex quinata (Lour.) Will. Zeugophora (P. ) maculata Chûjô, 1941 Guizhou common Symplocaceae: Symplocos sumuntia Buch.-Ham. ex D. Don Zeugophora (P. ) nigricollis (Jacoby, 1885) Ningxia common Celastraceae: Euonymus alatus (Thunb.) Siebold Zeugophora (P. ) nigroapica Li & Liang, 2018 Yunnan holotype unknown Zeugophora (P. ) tricolor Chen & Pu, 1962 Ningxia specimen compared with holotype Celastraceae: Euonymus alatus (Thunb.) Siebold Zeugophora (P. ) trifasciata Li & Liang, 2020 Yunnan holotype unknown Zeugophora (P. ) yuae Li & Liang, 2020 Yunnan holotype possible Lamiaceae: Vitex quinata (Lour.) Will. Zeugophora (P. ) yunnanica Chen & Pu, 1962 Yunnan specimen compared with holotype Symplocaceae: Symplocos sp. 283 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Zeugophora (Zeugophora) ancora Reitter, 1900 Figs 1–4 Zeugophora ancora Reitter, 1900: 164, pl. 1, fig. 9. TL: C. Asia (Donkyr). TD: unknown. Zeugophora ancora var. pseudancora Reitter, 1900: 165. Zeugophora (Pedrillia) ancora: Jolivet 1957: 12. Host plants. Salicaceae: Populus nigra var. italica (Moench) Koehne, Populus alba L. (Wu et al. 1982). Remarks. The median lobe of this species is similar to that of Zeugophora (Zeugophora) cyanea, Z. (Z.). cribrata, Z. (Z.) nigroaerea, and Z. (Z.) turneri in having the dorso-central portion of the median lobe with a sclerite (Figs 7, 10, 15, 23), but differs in having the apex of median lobe blunt. Zeugophora (Zeugophora) cribrata Chen, 1974 Figs 5–8 Zeugophora cribrata Chen, 1974: 43, 44, 47, fig. 1A. TL: China, Qinghai. TD: IZCAS. Host plant. Salicaceae: Salix L. (Chen 1974; pers. obs.). Remarks. The external male genitalia of this species are similar to those of Zeugophora (Zeugophora) cyanea, but differ in having the median lobe slightly broader (median lobe slightly narrower in Z. (Z.) cyanea, Fig. 10). Figures 1–8. Male genitalia of Zeugophora (Zeugophora) species. 1–4. Z. (Z.) ancora Reitter, 1900; 1. Median lobe and median struts, lateral view; 2. Median lobe, dorsal view; 3. Tegmen, dorsal view; 4. Spiculum, dorsal view; 5–8. Z. (Z.) cribrata Chen, 1974; 5. Spiculum, dorsal view; 6. Median lobe and median struts, lateral view; 7. Median lobe, dorsal view; 8. Tegmen, dorsal view. Scale bar: 2 mm. 284 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Zeugophora (Zeugophora) cyanea Chen, 1974 Figs 9–12 Zeugophora cyanea Chen, 1974: 44, 45, 47. TL: China, Qinghai. TD: IZCAS. Host plant. Salicaceae: Salix L. (Chen 1974; pers. obs.). Remarks. The similarities and differences of external male genitalia between this species and Zeugophora (Zeugophora) cribrata are provided in the remarks under Z. (Z.) cribrata. Zeugophora (Zeugophora) nigroaerea Lopatin, 2008 Figs 13–16 Zeugophora nigroaerea Lopatin, 2008: 918–927. TL: China (Gansu). TD: ZIN. Host plant. Salicaceae: Salix L. (pers. obs.). Remarks. The external male genitalia of this species are most similar to those of Zeugophora (Zeugophora) cyanea (Figs 9–12), but differ in having the median lobe slightly longer (median lobe slightly shorter in Z. (Z.) cyanea, Fig. 10). This species is also similar to Z. (Z.) cribrata (Figs 5–8), but differs from it in having the apex of the median lobe slightly narrower (apex of median lobe slightly broader in Z. (Z.) cribrata, Fig. 7). Zeugophora (Zeugophora) scutellaris Suffrian, 1840 Figs 17–20 Zeugophora scutellaris Suffrian, 1840: 99. TL: Germany (Aschersleben, Magdeburg). TD: MZPW. Host plants. Salicaceae: Populus acuminata Rydberg, P. alba L., P. deltoides Marshall, P. grandidentata Michaux, P. nigra L., P. tremula L., Salix sp. (Medvedev and Rogynskaya 1988; Bukejs 2009; Sergeev and Legalov 2022). Remarks. The external male genitalia of this species differ from those of other species of the subgenus Zeugophora (Zeugophora) in having the dorso-central portion without a sclerite (Fig. 18). Zeugophora (Zeugophora) turneri Power, 1863 Figs 21–24 Zeugophora turneri Power, 1863, 21: 8735. TL unknown. TD unknown. Zeugophora rufotestacea Kraatz, 1871, 15: 162. Host plant. Salicaceae: Populus sp. (Bukejs 2009). Remarks. The external male genitalia of this species are similar to those of Zeugophora (Zeugophora) cribrata and Z. (Z.) cyanea in the dorso-central 285 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae portion of median lobe with a sclerite, but differ in having apex of median lobe gradually narrowed (apex of median lobe abruptly narrowed in Z. (Z.) cribrata and Z. (Z.) cyanea). Subgenus Pedrillia Westwood, 1864 External male genitalia. (Figs 25–83, 92–99) Median lobe sclerotized, slender or short and broad, curved in lateral view, sides generally thickened, dorso-central portion entirely membranous, sides parallel or diamond-shaped, or dorso-central portion membranous with basal 1/3 tubular, sides parallel, apex generally strongly protruding, blunt or sharp; median struts rodshaped, widely separated from each other, ~1.5–3.3× as long as median lobe; base of tegmen Yor V-shaped, basal piece generally short, tegminal ring with base generally narrower than apical portion, paramere generally tongue-shaped or sub-trapezoid, apical margin with setae; endophallus membranous, with a paired granulated area. Spiculum long, basal part long, apex trifid or not, sclerotized. Zeugophora (Pedrillia) annulata (Baly, 1873) Figs 25–28 Pedrillia annulata Baly, 1873: 79. TL: Japan. TD: NHMUK. Pedrillia annulata var. disconotata Pic, 1906: 27. Synonymized by Ohno 1961: 37. Pedrillia annulata ab. theresae Pic, 1945: 13. Zeugophora annulata: Gressitt, 1945: 137. Zeugophora (Pedrillia) annulata: Jolivet, 1957: 12. Pedrillia annulata f. inannulata Chûjô, 1959: 1. Pedrillia annulata f. melanaria Chûjô, 1959: 1. Pedrillia biguttata Kraatz, 1879a: 119. Synonymized by Ohno, 1961: 37. Pedrillia annulata f. biguttata: Ohno, 1961: 37. Host plants. Celastraceae: Euonymus alatus (Thunb.) Siebold, Euonymus przewalskii Maxim., Euonymus phellomanus Loes. ex Diels., Euonymus maacki Rupr., and Euonymus sacrosancta Koidz. (Medvedev and Roginskaya 1988; Takemoto 2019; pers. obs.). Remarks. The external male genitalia of this species are similar to those of Zeugophora (Pedrillia) bicolor and Z. (P. ) tricolor in having the median lobe long and arched in lateral view, and the shapes of median lobe, tegmen, and spiculum. This species differs from Z. (P. ) bicolor in having the apex of the median lobe slightly narrower, not bending / curving leftwards, basal piece of tegmen shorter (apex of median lobe slightly broader, tending slightly leftwards, basal piece of tegmen longer in Z. (P. ) bicolor, Figs 31, 32). This species differs from Z. (P. ) tricolor in having the apex of the median lobe slightly longer and sharper (apex of median lobe slightly shorter and blunt in Z. (P. ) tricolor, Fig. 70). 292 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Zeugophora (Pedrillia) nigricollis (Jacoby, 1885) Figs 60–63 Pedrillia nigricollis Jacoby, 1885: 195. TL: Japan, Wada-tôge. TD: NHMUK. Synonymized as Pedrillia bicolor Kraatz, 1879a: 120 by Kimoto 1986: 309. Zeugophora (Pedrillia) nigricollis: Crowson 1946: 95; Gressitt & Kimoto 1961: 24, 27; Chûjô 1954: 51; Chûjô & Kimoto 1961: 119; Kimoto 1964: 108. Zeugophora (Pedrillia) bicolor: Kimoto 1986: 309; Jolivet 1957: 12; Kimoto & Takizawa 1994: 6, 99, 267; An & Kwon 2002: 272; Silfverberg 2010: 334; Li & Liang 2018: 133. Zeugophora bicolor: Gressitt 1945: 139; An & Kwon 1995: 91–92; Takizawa 2006: 2; Rodríguez-Mirón 2018: 291. Zeugophora nigricollis (Jacoby, 1885). Restored as a valid species by Takemoto 2019: 15–19. Host plants. Celastraceae: Euonymus sieboldiana Blum., Euonymus alatus (Thunb.) Siebold (Takemoto 2019; Li and Liang 2020). Remarks. The external male genitalia of this species are similar to those of Zeugophora (Pedrillia) annulata, Z. (P. ) bicolor, Z. (P. ) tricolor, and Z. (P. ) euonymorum, but differ from Z. (P. ) annulata, Z. (P. ) bicolor, and Z. (P. ) tricolor in having the median lobe less curved in the lateral view and slender, with the apex of the median lobe triangular and blunt; genitalia differ from that of Z. (P. ) euonymorum in having the median lobe slender and the apex slightly narrower. Zeugophora (Pedrillia) nigroapica Li & Liang, 2018 Figs 64–67 Zeugophora (Pedrillia) nigroapica Li & Liang, 2018: 143. TL: China (Yunnan). TD: IZCAS. Host plant. Unknown. Remarks. The similarities and differences of external male genitalia between this species and Zeugophora (Pedrillia) emeica Li & Liang are presented in the remarks of Z. (P. ) emeica. The external male genitalia of this species differ from other species of the subgenus Pedrillia except that of Z. (P. ) emeica in having a sub-oval median lobe. Zeugophora (Pedrillia) tricolor Chen & Pu, 1962 Figs 68–71 Zeugophora (Pedrillia) tricolor Chen & Pu, 1962: 116, fig. 2. TL: China (Gansu). TD: IZCAS. Host plant. Celastraceae: Euonymus alatus (Thunb.) Siebold (pers. obs.). Remarks. The similarities and differences of external male genitalia of this species, Zeugophora (Pedrillia) annulata, and Z. (P. ) bicolor are described in the remarks under Z. (P. ) annulata and Z. (P. ) bicolor. 293 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Zeugophora (Pedrillia) trifasciata Li & Liang, 2020 Figs 72–75 Zeugophora (Pedrillia) trifasciata Li & Liang, 2020, ZooKeys, 975: 69–72, figs 58–66. TL: China: Yunnan. TD: KIZ. Host plant. Unknown. Remarks. The similarities and differences of external male genitalia between this species and Zeugophora (Pedrillia) dimorpha are described in the remarks under Z. (P. ) dimorpha. The external male genitalia of this species are similar to that of Zeugophora (Pedrillia) yunnanica Chen & Pu in shape, but differ in having the median lobe dorso-central with a strongly sclerotized sclerite (without a strongly sclerotized sclerite in Z. (P. ) yunnanica, Fig. 82). Figures 57–63. Male genitalia of Zeugophora (Pedrillia) species. 57–59. Z. (P. ) maculata Chûjô, 1941; 57. Median lobe and median struts, lateral view; 58. Median lobe, dorsal view; 59. Tegmen, dorsal view; 60–63. Z. (P. ) nigricollis (Jacoby, 1885); 60. Median lobe and median struts, lateral view; 61. Median lobe, dorsal view; 62. Tegmen, dorsal view; 63. Spiculum, dorsal view. Scale bar: 0.2 mm. 294 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Zeugophora (Pedrillia) yuae Li & Liang, 2020 Figs 76–79 Zeugophora (Pedrillia) yuae Li & Liang, 2020, ZooKeys, 975: 72–76, figs 67–79. TL China: Yunnan. TD: KIZ, IZCAS. Host plant. Lamiaceae: Vitex quinata (Lour.) Will. (Li and Liang 2020). Remarks. The external male genitalia of this species are similar to that of Zeugophora (Pedrillia) tricolor in shape, but differ in having the median lobe less curved in the lateral view, the apex of the median lobe slightly broader and blunt, and the ratio of median struts / median lobe ~3.5 (the median lobe is more curved in the lateral view, the apex of the median lobe slightly narrower and sharper, and the ratio of median struts / median lobe is ~2.2 in Z. (P. ) tricolor). Figure 64–71. Male genitalia of Zeugophora (Pedrillia) species. 64–67. Z. (P. ) nigroapica Li & Liang, 2018; 64. Aedeagus, lateral view; 65. Median lobe, dorsal view; 66. Tegmen, dorsal view; 67. Spiculum, dorsal view; 68–71. Z. (P. ) tricolor Chen & Pu, 1962; 68. Spiculum, dorsal view; 69. Median lobe and median struts, lateral view; 70. Median lobe, dorsal view; 71. Tegmen, dorsal view. Scale bar: 0.2 mm. 295 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Zeugophora (Pedrillia) yunnanica Chen & Pu, 1962 Figs 80–83 Zeugophora (Pedrillia) yunnanica Chen & Pu, 1962: 116, fig. 3. TL: China (Yunnan). TD: IZCAS. Host plant. Symplocaceae: Symplocos sp. (Chen and Pu 1962). Remarks. The similarities and differences of external male genitalia of this species, Zeugophora (Pedrillia) dimorpha, and Z. (P. ) trifasciata are presented in the remarks under Z. (P. ) dimorpha and Z. (P. ) trifasciata. Figures 72–75. Male genitalia of Zeugophora (Pedrillia) trifasciata Li & Liang, 2020. 72. Median lobe and median struts, lateral view; 73. Median lobe, dorsal view; 74. Tegmen, dorsal; 75. Spiculum, dorsal view. Scale bar: 0.2 mm. 296 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Zeugophora (Pedrillia) liupanshanensis sp. nov. https://zoobank.org/1751E637-1A2B-4A46-8740-B2AC4DCD64E8 Suggested Chinese name 六盘山小距甲 Figs 84–106 Specimens examined. Holotype (KIZ): male, China, Ningxia, Jingyuan, Liupan Shan National Nature Reserve, Danangou, 35.48659°N, 106.27045°E / 2130 m, 2018.vii.28, Kaiqin Li coll., Kunming Institute of Zoology, Chinese Acad. Sci. / Holotype, Zeugophora (Pedrillia) liupanshanensis sp. nov., des. by K.Q. Li & H.B. Liang, 2024 [red label]. Paratype (IZCAS): 1 male, China, Ningxia, Jingyuan, Liupan Shan National Nature Reserve, Longtan Forest Farm, 35.39160°N, 106.35299°E / 1918 m, 2018.vii.26, Kaiqin Li & Likun Zhang coll., Kunming Institute of Zoology, Chinese Acad. Sci. / Paratype, Zeugophora (Pedrillia) liupanshanensis sp. nov., des. by K.Q. Li & H.B. Liang, 2024 [yellow label]. Figures 76–83. Male genitalia of Zeugophora (Pedrillia) species. 76–79. Z. (P. ) yuae Li & Liang, 2020; 76. Median lobe and median struts, lateral view; 77. Median lobe, dorsal view; 78. Tegmen, dorsal view; 79. Spiculum, dorsal view; 80–83. Z. (P. ) yunnanica Chen & Pu, 1962; 80. Spiculum, dorsal view; 81. Median lobe and median struts, lateral view; 82. Median lobe, dorsal view; 83. Tegmen, dorsal view. Scale bar: 0.2 mm. 297 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Diagnosis. Head brown, pronotum, scutellum, and underside brown; elytra brown, posterior portion with two black spots; antennae black; legs brown with posterior portion black; antennomeres 3 and 4 long, 5–11 short; apex of median lobe triangular and narrow. Description. BL = 3.8 mm, BW = 1.7 mm. Head brown, except posterior portion of mandible black; pronotum, scutellum, and underside brown; elytra brown, but with two black spots posteriorly; antennae black; legs brown but distal femora, tibiae, and tarsi black. Head: eye prominent, inner margin with distinct canthus; vertex sparsely punctate and pubescent centrally, densely punctate and pubescent laterally; Figures 84–91. Holotype and paratype of Zeugophora (Pedrillia) liupanshanensis sp. nov. 84–87. Holotype, male; 84. Dorsal view; 85. Ventral view; 86. Head, anterior view; 87. Pronotum, dorsal view; 88–91. Paratype, male; 88. Dorsal view; 89. Ventral view; 90. Head, anterior view; 91. Pronotum, dorsal view. Scale bars: 1.0 mm (84, 85, 88, 89); 0.5 mm (86, 87, 90, 91). 298 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae occiput strongly constricted; frons sparsely punctate and pubescent, center with a shallow concave; clypeus rectangular, width 2× length, punctate and pubescent laterally and anteriorly, separated from frons by deep clypeal suture; labrum rectangular, narrower than clypeus, anterior margin with punctures and pubescence; antennae slender, exceeding humeri in length, antennomere 1 long and swollen, antennomere 2 short, 2/3 as long as antennomere 1, antennomere 3 as long as 4 and 1, antennomere 5 shorter than antennomere 4, antennomeres 6–10 as long as antennomere 2, antennomere 11 acute at apex, as long as antennomere 5; antennomeres 1–4 sparsely punctate and pubescent, antennomeres 5–11 densely punctate and pubescent. Thorax: PW/PL = 1.3; anterior margin slightly flattened; posterior margin extended posteromedially ; length of posterior margin slightly longer than anterior margin; anterior groove distinct laterally, obsolete medially; posterior groove deep laterally, shallow medially; anterior portion of lateral margin subparallel, then gradually expanding from anterior portion to middle, strongly constricted behind middle; lateral tubercle rounded; behind the lateral tubercle, an oblique groove extending to basal portion; disc convex with dense and coarse punctures and pubescence, basal portion slightly depressed; basal portion of each side slightly prominent. Scutellum triangular, slightly emarginate at apex, densely punctate and pubescent. Figures 92–99. Holotype and paratype of Zeugophora (Pedrillia) liupanshanensis sp. nov. 92–95. Male genitalia of holotype; 92. Median lobe and median struts, lateral view; 93. Median lobe, dorsal view; 94. Tegmen, dorsal view; 95. Spiculum, dorsal view; 96–99. Male genitalia of paratype; 96. Spiculum, dorsal view; 97. Median lobe and median struts, lateral view; 98. Median lobe, dorsal view; 99. Tegmen, dorsal view. Scale bar: 0.2 mm. 299 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Elytra: EL/EW = 1.5–1.6; elytral humeri projecting antero-laterally, humeral groove shallow, lateral part of humeri densely punctate and pubescent; lateral margin slightly expanding from the base to the middle, widest behind the middle, apex slightly rounded; disc slightly flattened, coarsely punctate and pubescent; suture with one row of punctures and pubescence; epipleura narrow, two rows of punctures and pubescence at base and one row at apex. Abdomen and legs: underside sparsely punctate and pubescent. Legs moderately long, femora robust, midand hind-tibiae slightly curved. Pygidium moderately long, apical portion exposed. Apical margin of last abdominal ventrite slightly prominent. Median lobe sclerotized, short and broad, curved in lateral view, dorso-central portion membranous, basal 1/3 tubular, sides thickened and parallel, apex triangular, strongly constricted and slightly sharp; median struts rod-shaped, widely separated from each other, ~2.2–2.4× as long as median lobe; basal portion of tegmen Y-shaped, tegminal ring subdiamond-shaped, paramere tongue-shaped, apical margin of paramere with setae; endophallus membranous, with weakly sclerotized area. Spiculum spoon-shaped, apical margin prominent. Distribution. China (Ningxia). Host plants. Euonymus przewalskii, E. giraldii (Celastraceae). We collected this species by beating the Celastraceae trees in the forest. One specimen was collected from E. giraldii, the other was from E. przewalskii. Several leaves were observed to have damage from bite marks (Fig. 106). Figures 100–102. Habitat of Zeugophora (Pedrillia) liupanshanensis sp. nov. (Ningxia, Liupan Shan, Danangou). 100. Habitat; 101, 102. Host plants Euonymus przewalskii (Celastraceae) of Z. (P. ) liupanshanensis sp. nov. 300 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae Etymology. The specific name liupanshanensis refers to the type locality, Liupan Shan. Remarks. The pronotum shape of the new species is similar to that of Zeugophora (Pedrillia) eumonymorum Li & Liang, 2020 which is also found in Liupan Shan, but differs from the latter in having body mostly brown elytra with black spots, shorter antennae, the apex of the median lobe narrower (vs body dark blue or green, underside black, longer antennae, apex of median broader in Z. (P. ) eumonymorum). The external male genitalia of this species are similar to those of Zeugophora (Pedrillia) annulata, Z. (P. ) bicolor, and Z. (P. ) tricolor (Figs 25, 26, 30, 31, 69, 70), but differ from them in having a less curved median lobe in lateral view, and a shorter and less sclerotized apex of the median lobe, (more curved Figures 103–106. Habitat and host plants of Zeugophora (Pedrillia) liupanshanensis sp. nov. (Ningxia, Liupan Shan, Longtan forest farm). 103. Habitat; 104–106. Host plant Euonymus giraldii (Celastraceae) of Z. (P. ) liupanshanensis sp. nov.; 106. Feeding traces on the abaxial leaf surface caused by adults. 301 ZooKeys 1261: 277–306 (2025), DOI: 10.3897/zookeys.1261.169189 Kai-Qin Li et al.: Study of male genitalia and description one new species of Zeugophorinae median lobe in lateral view, apex of median lobe longer and more sclerotized in Z. (P. ) annulata, Z. (P. ) bicolor, and Z. (P. ) tricolor). This species is similar to Zeugophora (Zeugophora) tetraspilota Medvedev, 1998 (Fig. 110) in having posterior spots on elytra, but differs in having brown elytra with two black spots, a brown pronotum, black antennae, brown legs with black coloration, and a broader lateral tubercle on the pronotum (elytra yellowish brown with four black spots; pronotum, antennae, and legs yellowish brown, narrower lateral tubercle on the pronotum in Z. tetraspilota). This species is also similar to Zeugophora (Pedrillia) trisignata An & Kwon, 2002 and Z. (P. ) bimaculata Kratz, 1879 in having posterior spots on the elytra and in body shape, but differs from the latter in having a brown head, black antennae, brown legs with the posterior portion black, and antennomeres 3 and 4 longer (vs black head, antennomeres 1–4 yellowish brown, yellowish brown legs, antennomeres 3–4 shorter in Z. trisignata and Z. bimaculata, Figs 108, 109, respectively). The external male genitalia of this species differ from that of Z. (P. ) bimaculata in having a narrower apex of the median lobe (apex of median lobe broader in Z. (P. ) bimaculata). Moreover, we found the elytra colorations of Z. (P. ) bimaculata and Z. (P. ) trisignata to be variable. The species Z. (P. ) bimaculata has three kinds of coloration (Sergeev and Legalov 2022): one of them is the same as the type specimen of Z. (P. ) trisignata (type locality Odaesan, Korea). The species Z. (P. ) trisignata has two kinds of coloration (An and Kim 2020), of which one is the same as the type of Z. (P. ) bimaculata (type locality Amur, Russia). Therefore, we compared descriptions and illustrations of the types of Z. (P. ) bimaculata (Kraatz, 1879b; Fig. 108) and Z. (P. ) trisignata (An and Kwon 2002; Fig. 109) as no specimens were available, and no significant difference was found in external morphology. We speculate Z. (P. ) trisignata is probably conspecific with Z. (P. ) bimaculata, but their genitalia should be compared to confirm this synonymy. Figures 107–110. Types of Zeugophora. 107, 108. Type of Z. (Pedrillia) bimaculata Kraatz, 1879; 109. Holotype of Z. (Pedrillia) trisignata An & Kwon, 2002; 110. Type of Z. (Zeugophora) tetraspilota Medvedev, 1998.