#ZooKeys ZooKeys 1215: 151-183 (2024) DOI: 10.3897/zookeys.1215.134871 Research Article The genus Argopistes Motschulsky from Japan and Taiwan, with descriptions of three new species from Taiwan (Coleoptera, Chrysomelidae, Galerucinae, Alticini) Chi-Feng Lee’™®, Ming-Yao Chiang'™®, Haruki Suenaga? 1 Applied Zoology Division, Taiwan Agricultural Research Institute, Taichung 413, Taiwan 2 Nakashima, 108-11, Kurashiki-shi, Okayama, 710-0803 Japan Corresponding author: Ming-Yao Chiang (mingyaw@tari.gov.tw) OPEN Qaccess Academic editor: Ron Beenen Received: 16 August 2024 Accepted: 15 September 2024 Published: 15 October 2024 ZooBank: https://zoobank.org/ C57CB315-F1 5F-4D98-868A- EFEA22BC64A3 Citation: Lee C-F, Chiang M-Y, Suenaga H (2024) The genus Argopistes Motschulsky from Japan and Taiwan, with descriptions of three new species from Taiwan (Coleoptera, Chrysomelidae, Galerucinae, Alticini). Zookeys 1215: 151-183. https://doi. org/10.3897/zookeys.1215.134871 Copyright: © Chi-Feng Lee 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). Abstract Four previously described species of Argopistes are recognized and redescribed from Japan and Taiwan: A. biplagiatus Motschulsky, 1860, A. rufus Chen, 1934, A. tsekooni Chen, 1934, and A. unicolor Jacoby, 1885. Three new species from Taiwan, A. jungchani sp. nov., A. tsoui sp. nov., and A. yuae sp. nov., are described. Descriptions of species include illustrations of aedeagi, antennae, gonocoxae, abdominal ventrite VIII, and sper- mathecae. Argopistes rufus Chen, 1934, stat. nov. is raised to species status from a variety of A. biplagiatus Motschulsky, 1860. Argopistes coccinelliformis Csiki, 1940, syn. nov. and A. ryukyuensis Shigetoh & Suenaga, 2022, syn. nov. are proposed as junior synonyms of A. rufus Chen, 1934 Lectotypes are designated for A. undecimmaculata Jacoby, 1885, A. unicolor Jacoby, 1885, and A. biplagiatus var. rufus Chen, 1934. Key words: Chionanthus, Fraxinus, Jasminum, Ligustrum, Olea, Oleaceae, Osmanthus, Syringa Introduction The flea beetle genus Argopistes Motschulsky, 1860 contains 44 species re- corded from Afrotropical, Australian, Neotropical, Oriental, and Palearctic re- gions (Blanco and Konstantinov 2013; Biondi et al. 2024). Four species were known from Japan and reviewed by Kimoto (1965) with emphasis on male aedeagi. A new species was also described from Ryukyu Islands and Dait6 Is- lands (Shigetoh and Suenaga 2022). Chdj6é (1936) was the first to record the genus from Taiwan as A. biplagiatus Mostschulsky, 1860, although Gressitt and Kimoto (1963) indicated that it was a misidentification of A. coccinelliformis Csiki, 1940. No other records have been reported from Taiwan since then. Adults and larvae of Argopistes are oligophagous on Oleaceae (Jolivet and Hawkeswood 1995). A number of species of Oleaceae are ornamental trees popu- lar in Japan, including Osmanthus x fortunei Carriére, O. heterophyllus (G. Don) P. S. Green, and Ligustrum japonicum Thunb. Although few insect pests are reported for these ornamental trees, A. rufus Chen, 1934 and A. biplagiatus Motschulsky, 1860 are major pests. Ecology of both species have been studied in this context (Inoue 151 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan and Shinkaji 1989a-c, 1990; Inoue 1990a, b, 1991a, b, 1992, 1993, 1994, 1996, 1998, 2001, 2014). In contrast, Chinese privet, Ligustrum sinense Lour., is one of the worst invasive plants in the U.S. Argopistes tsekooni Chen, 1934 was evaluated as a promising biological control agent of Chinese privet (Zhang et al. 2008a, b, 2009). In Taiwan, Chionanthus retusus Lindley & Paxton (ijt f«) (Fig. 1A—D, G), Chi- nese fringetree, and Osmanthus fragrans (Thunb.) Lour. (#:74¢), sweet osman- thus, are popular ornamental plants. They have been attacked by Argopistes species during recent years. This phenomenon also occurs on small islands, including Kinmen Island (Fig. 1E, F), Nangan Island (Fig. 1C, D), and Beigan Island (Fig. 1G, H). Taxonomic studies on Argopistes in Taiwan and Japan are needed to describe diagnostic characters in addition to male aedeagi. Figure 1. Field photographs of Argopistes rufus Chen A adults feeding on leaves of Chionanthus retusus surrounding Hsinchu City Government on April 23, 2021 B mature larvae mining leaves of the same tree C blooming C. retusus at Qingshui Village (7K 4), Nangan Island (fF 53), on April 21, 2024 D larvae mining leaves near the ground of the same tree E larvae mining leaves of Osmanthus fragrans at Yingshan Temple (% 11/23), Kinmen Island (42F4 &), on April 11, 2023 F larvae mining leaves of Osmanthus fragrans at the guesthouse, Jinhu Township (4=ii//A), Kinmen Island, on May 20, 2024 G C. retusus (red arrow) and Ligustrum japonicum (blue arrow) planting surrounding Chinbe Village (7 #), Beigan Island (4t4F &) on 22 April 22 2024 H feeding marks caused by adults on leaves of L. japonicum. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 152 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Materials and methods For taxonomic study, abdomens of adults were separated from the forebodies and boiled in 10% KOH solution, followed by washing in distilled water to prepare genitalia for illustrations. The genitalia were then dissected from the abdomens, mounted on slides in glycerin, and studied and drawn using a Leica M165 stereomi- croscope. For detailed examinations, a Nikon ECLIPSE 50i microscope was used. At least three males and females from each species were examined to de- limit variability of diagnostic characters. For species collected from more than one locality or with color variations, at least one pair of each sex from each locality and color morph was examined. Length was measured from the ante- rior margin of the eye to the elytral apex, and width at the greatest width of the elytra. Nomenclature for morphological structures of adults follows Duckett and Daza (2004). Names of plant species follows the Taiwan Encyclopedia of Life (2024; TaiEOL). Specimens studied herein are deposited at the following institutes and col- lections: HAPC Private Collection of Haruki Suenaga, Okayama, Japan; HIPC Private Collection of Hiroaki Shigetoh, Sapporo, Japan; IZAS Institute of Zoology, Chinese Academy of Sciences, Beijing, China [Yongying Ruan]; NHMUK _ The Natural History Museum, London, UK [Michael F. Geiser, Max- well V. L. Barclay]; SEHU The Laboratory for Systematic Entomology, Hokkaido University, Sapporo, Japan [Takuya Takemoto]; TAFI Forest Arthropod Collection of Taiwan, Taiwan Forestry Research Institute, Taipei City, Taiwan [Sheng-Shan Lu]; TARI Applied Zoology Division, Taiwan Agricultural Research Institute, Taichung, Taiwan [Chi-Feng Lee]; ZMMU Zoological Museum of Moscow State University, Moscow, Russia [Vladimir Savitsky]. Exact label data are cited for all type specimens of described species; a dou- ble slash (//) divides the data on different labels and a single slash (/) divides the data in different rows. Other comments and remarks are in square brackets: [p] — preceding data are printed, [h] — preceding data are handwritten, [w] - white label, [y] — yellow label, [g] - green label, [b] — blue label, and [r] — red label. Traditional Chinese fonts are added to the names of localities. Taxonomic account Argopistes biplagiatus Motschulsky, 1860 Figs 2A-F, 3, 4 Argopistes biplagiatus Motschulsky, 1860: 236 (Amur: Russian Far East and northeastern China); Csiki 1940: 523 (catalogue); Chdjé and Kimoto 1961: 174 (catalogue); Kimoto 1965: 436 (redescription); Lee and An 2001: 182 (South Korea); Lee and Cho 2006: 91 (host plants); Takizawa 2012: 38 (faunistics). ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 153 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Argopistes flavitarsis Motschulsky, 1860: 137 (chromatic variation). Argopistes limbatus Motschulsky, 1860: 137 (chromatic variation). Argopistes suturalis Motschulsky, 1860: 137 (chromatic variation). Argopistes undecimmaculata Jacoby, 1885: 738 (Japan: Sapporo); Chdjé 1936: 109 (catalogue); Csiki 1940: 524 (catalogue). Type material examined. Argopistes biplagiatus. * 11 syntypes glued on the same card (ZMMU) (Fig. 2A-D): “type [h, w] // Amur [h, r] // Argopistes / bipla- giatus / Amur. m. Motsch [h, w, with black border] // Syntypus [p, r] // 3oomy3en Mry (Mockba, POCCNR) / No ZMMU Col 03056 / Zool. Mus. Mosq. Univ. / (Mosquae, RUSSIA) / ex coll. V. |. Motschulsky [p, pink label]”. Argopistes undecimmaculata. Lectotype « (here designated, sex undetermined, NHMUK) (Fig. 2E, F): “Type / H.T. [p, w, circle label with red border] // SYN- / TYPE [p, w, circle label with blue border] // Sapporo / 5.VIII-16.VIII.80. [p, w] // Japan / G. Lewis. / 1910-320 [p, w] // Sap [h, w]”. Paralectotypes + 1 (sex undetermined, NHMUk): “SYN- / TYPE [p, circle label with blue border] // Sapporo / 5.VIII-16. VIII.80. [p, w] // Japan/ G. Lewis. / 1910-320 [p, w] // Argopistes / 11maculata Jac [h, b]”; * 12 (TARI): “Sapporo [h] / JAPAN [p] / 10.VIII.1880 [h] / Col. G. LEWIS [p, w] // Argopistes / undecimmaculata / Jacoby [h] / DET. M. CHUJO [p, w] // CO / Type [p, w, circle label with yellow letters and border] / 1526 [p, w]”. | | Pi: eer | Ld Pop, BMNH(E) #1024843 H Figure 2. Type specimens and labels A Argopistes biplagiatus Motschulsky, 1860, syntypes B one syntype with typical color form C one syntype with enlarged red spots on elytra D labels pinned with syntypes E A. undecimmaculata Jacoby, 1885, lectotype F labels pinned with lectotype G A. coccinelloides Baly, 1874, holotype H labels pinned with holotype. = pe ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 154 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan G Figure 3. Argopistes biplagiatus Motschulsky A antenna, male B antenna, female C aedeagus, dorsal view D aedeagus, lateral view E abdominal ventrite VIII, female F spoermatheca G gonocoxae. Additional material examined. JAPAN. Hokkaido: - 12 (HAPC), Sapporo-shi, Hokkaido University, 15.X.2011, leg. H. Suenaga; Honshu. Aichi: » 14 (SEHU), Toyohashi-shi, Imou-shitsugen, 8.IV.1989, leg. Y. Komiya; Ibaraki: * 12 (HIPC), Daigo, Uenomiya, Mt. Yamizo-san, 28.V.2917, leg. H. Yoshitake; * 1’ (SEHU), Sakura-mura, Sakura-gawa Riv., 1.V1.1986, leg. Y. Komiya; Ishikawa: * 19 (HAPC), ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 155 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Figure 4. Habitus of Argopistes biplagiatus Motschulsky A typical color form, female, dorsal view B ditto, ventral view C ditto, lateral view D yellowish brown color form, female, dorsal view E ditto, ventral view F ditto, lateral view. = Mt. Haku-san, Betsuzan-d6, 21.V.2016, leg. H. Kawase; Shizuoka: + 14, 2° (SEHU), Izu-peninsula, Mt. Manzaburo-dake, 19.V.1980, leg. J. Okuma; * 19 (SEHU), Tagata-gum, Tohi, 4.V.1985, leg. Y. Komiya; Tokyo: * 1< (NHMUK), Kat- sushika-ku, Mizumoto K6en Park, 8.V.2005, leg. Y. Komiya; Shikoku. Ehime: + 13) (HAPC), Kumakdégen-ché, Mt. Saragamine, 7.VI.2009, leg. H. Suenaga; + 14', 29 (HAPC), Matsuyama-shi, Mt. Takanawa-san, 12.V.2007, leg. S, Sejima; Kyushu. 33,19 (TARI), Mt. Hiko-san, 14.VIII.1941, leg. M. Chajé; Fukuoka: 24 (HAPC), Soeda-machi, Mt. Hiko-san, 8.VIII.2009, leg. S. Sejima; RUSSIAN FAR EAST. Pri- morsky Krai: * 2 (NHMUK), Lazovski Zapovednik, 170 m E Vladivostok, Korpad, 28.V.-6.VI.2001, leg. M. Quest; * 14’ (NHMUk), Odarkovskij, Zavod, 25.IV.1911, leg. A. Tsherskij; * 14’ (NHMUK), Wladiwostok, leg. Herman Frieb.; SOUTH KOREA. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 156 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan * 19 (TARI), Sulgen, 15.VII.1932, leg. D. Okamoto; TAIWAN. Taipei: * 1, 19 (TARI), Kueitzukeng (4f45t), 4.XII.2006, leg. H.-T. Cheng; * 19 (TARI), same but with “leg. H. Lee”; * 14 (TARI), same locality, 9.1X.2007, leg. M.-H. Tsou; + 12 (TARI), same but with “18.X1.2007”; + 29 (TARI), Tienmu (7c4#), 8.XI1.2006, leg. S.-F. Yu. Diagnosis. Adults of Argopistes biplagiatus are similar to those of A. rufus with similar color pattern but differing from A. rufus possessing line of punc- tures that are less coarse than those between the lines, sometimes confused (lines of punctures much coarser than those between lines in A. rufus) and a wider interspace between eyes. Genitalic characters are more diagnostic for both species. Those of A. biplagiatus possess pointed apices (Fig. 3C) and are wider in lateral view (Fig. 3D) (widely rounded apex (Fig. 5C) and narrow aedea- gus in lateral view (Fig. 6D) in A. rufus); females have narrow, parallel-sided bas- es of gonocoxae (Fig. 3G) (medially widened gonocoxae (Fig. 5G) in A. rufus), and ventrite VIII evenly rounded and with dense setae on apical margin (Fig. 4E) (medially depressed and without setae on median area of apical margin of ab- dominal ventrite VIII (Fig. 5E) in A. rufus). Figure 5. Distribution map of Argopistes species in Taiwan, solid line: 1000 m, broken line: 2000 m. Red dots A. rufus Chen; blue dots A. biplagiatus Motschulsky; green dots A. tsoui sp. nov.; orange dots A. yuae sp. nov.; purple dot A. jung- chani sp. nov. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 157 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan In addition, adults of A. biplagiatus in Taiwan are larger (4.7-4.9 mm) than those of A. rufus (3.8-4.3 mm). Moreover, distinct color patterns occur in both species respectively (black elytra with reddish brown at middle in A. biplagiatus; yellowish brown elytra with distinct arrangement of black spots in A. rufus). Redescription. Length 4.4—4.9 mm, width 3.5-3.8 mm. Color variable (see below). Pronotum broad, convex, lateral margin narrowly explanate; 2.0-2.2 x wider than long, disc with dense coarse punctures; lateral margin rounded, an- terior margin strongly concave, posterior margin moderately convex. Intercoxal prosternal process flattened and with coarse punctures, delimited by narrow ridge on apical and lateral margins, truncate or slightly rounded at apex. Elytra broadly oval, 1.1 x longer than wide, disc with dense, confused, coarse punc- tures. Abdominal ventrite | with intercoxal area 2.0 x as long as wide, widest at basal 1/5, disc glabrous, rounded by reversed U-shaped ridge, provided with a row of coarse punctures inside subparallel lateral ridges. Male. Antenna filiform (Fig. 3A), antennomere | much longer than others, ap- proximate ratios of length of antennomeres I-XI 1.0: 0.3: 0.2: 0.4: 0.4: 0.3: 0.4: 0.4: 0.4: 0.4: 0.6; approximate ratios of length to width of antennomeres |I-XI 4.4: 1.9: 1.7: 2.4: 2.0: 1.6: 1.6: 1.8: 1.7: 1.7: 2.9. Aedeagus (Fig. 3C, D) apically and strongly narrowed from apical 1/5, slightly narrowed from apical 2/10-3/10, then slightly and basally widened towards basal 1/6, apex pointed; anterior opening very small, from apex to apical 3/10; tectum composed of one pair of sclerotized processes with apices twisted; extremely wide and straight in lateral view; paired processes straight in lateral view; endophallic sclerite laterally flat- tened, with small process near apex, and with basal processes membranous. Female. Antenna (Fig. 3B) similar to males, ratios of length of antennomeres I-XI 1.0: 0.3: 0.2: 0.3: 0.3: 0.3: 0.3: 0.3: 0.3: 0.3: 0.6; ratios of length to width of antennomeres I-XI 4.5: 2.0: 1.7: 2.0: 1.9: 1.7: 1.5: 1.6: 1.3: 1.5: 2.4. Ventrite VIII (Fig. 3E) membranous, only apical margin sclerotized, T-shaped, with dense long setae along apical margin, apical margin widely rounded, spiculum long. Sper- mathecal receptaculum (Fig. 3F) much longer than pump, moderately swollen, curved in lateral view; pump slightly emarginated at inner side of base; sperma- thecal duct with long basal part, ramus rounded. Gonocoxae (Fig. 3G) wide and separated, base membranous, each gonocoxa longitudinal and asymmetric, apically narrowed from middle, with dense long setae along apical areas. Color variation. In Japan, two distinct color patters of adults, typical color form (Fig. 4A—C): general color black, each elytron with one large red spot, lateral margin sometimes yellowish brown, legs dark brown but tarsi yellowish brown, head entirely black, or with one yellowish brown spot on vertex, or entirely yel- lowish brown except above eyes, abdominal ventrites yellowish brown but medi- ally black; yellowish brown color form: general color yellowish brown (Fig. 4D-F; undecimmaculata form), pronotum with one pair of small lateral black spots, elytra with 11 black spots, two pairs arranged into transverse lines near base and middle, one transverse pair near suture at middle, others longitudinal, one additional transverse pair near apex, one spot along suture from basal 1/3 to apical 1/3, medially widened, head yellowish brown but black below eyes except mouthparts, thoracic and abdominal ventrites black but abdominal ventrites lat- erally yellowish brown, legs black but tarsi, pro- and mesotibiae yellowish brown. At the type locality (Russian Far East and northeastern China), some individu- als represent the typical form (Fig. 2B) but with yellowish margins of pronotum ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 158 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan and elytra, some with enlarged red spots on the elytra connected with each other, some with entirely yellowish-brown bodies (Fig. 2C). In Taiwan, some specimens represent the typical form, but some have en- larged red spots on elytra that extend into the basal margin and connect with each other, and have reddish brown thoracic and abdominal ventrites. Host plants. Inoue (1990a) recorded the following species as host plants: Osmanthus x fortunei, O. heterophyllus, O. fragrans (#£7¢), O. fragrans var. auran- tiacus Makino, Ligustrum japonicum (H A #3), L. ovalifolium Hassk., L. licid- um W. T. Aiton, Syringa vulgaris L., and S. reticulata (Blume) H. Hara. Chaj6é and Kimoto (1961) recorded one additional host, Fraxinus mandshurica Rupr. var. japonica Maxim. Lee and Cho (2006) recorded Ligustrum obtusifolium Siebold & Zucc for Korean populations. Biology. Various aspects of biology of A. biplagiatus were studied in Japan, including feeding habits, seasonal development, habitat selection, host plant preference, and adult diapause (Inoue 1990a, b, 1991b, 1992, 1993, 1994). Gen- erally, the species has a univoltine life cycle. Eggs and/or larvae of this species are observed in the spring. Mature larvae fall from the host trees rather than crawling (Inoue 2014). Remarks. Syntypes of A. biplagiatus Motschulsky display great color varia- tion. Several names (A. flavitarsis, A. limbatus, and A. suturalis) have been pro- posed for different color patterns. Distribution. China, Japan (Hokkaido, Honshu, Shikoku, Kyushu), Russian Far East, South Korea, and new to Taiwan (Fig. 5). Argopistes rufus Chen, 1934, stat. nov. Figs 1, 2G, H, 6, 7 Argopistes coccinelloides Baly, 1874 (nec Suffrian, 1868): 202 (Japan); Chajé 1935a: 87 (Japan: Okinawa); Chijé 1935b: 211 (catalogue). Argopistes biplagiatus: Schénfeldt 1890: 175 (Japan: Loochoo); Chujé 1936: 110 (Taiwan), misidentification (Gressitt and Kimoto 1963). Argopistes biplagiatus var. rufus Chen, 1934a: 72 (China). Argopistes coccinelliformis Csiki, 1940: 524 (new replacement name for A. coc- cinelloides Baly, 1874); Chdj6 and Kimoto 1961: 174 (catalogue); Gressitt and Kimoto 1963: 812 (South China); Kimoto 1965: 436 (redescription); Takizwa, 2012: 38 (faunistics). Argopistes ryukyuensis Shigetoh & Suenaga, 2022: 4 (Japan: Okinawa). syn. nov. Type material examined. Argopistes coccinelloides. Holotype + (sex undeter- mined, NHMUK) (Fig. 2G, H): “Argopistes / coccinelloides / Baly / Japan [h, b] // Type / H.T. [p, circle label with red border] // Baly Coll. [h, w] // BMMH(E) / #1024843 [p, w]”. Argopistes biplagiatus var. rufus. Lectotype * 2° (here designated, NHMUKk): “China [p] // Bowring / 63°37* [p] // Argopistes coccinelliformis / Csiki, 1940 / det C.-F. Lee, 2023 [p] 2 [h, w] // NHMUK 015998267 [with OR Code, p, w]”. Paralec- toypes * 392 (NHMUKk), same as lectotype but with “0155998268-0155998270”. Argopistes ryukyuensis. Paratypes. JAPAN: Kitadaité-jima Island (JEAIR SS): - 14, 39 (HAPC), Kitadait6-jima, 21.IV.2018, leg. H. Kawase; Okinawa-jima ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 159 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Island: » 14’ (HAPC), Tomigusuku-shi, Tomigusuku, 10.V.2020, leg. H. Shige- toh; 1 3, 19 (HAPC), same but with “11.1I1.2021”; Ou-jima Island: + 19 (HIPC), Nanjo-shi, Tamashiro-ou, 6.V.2019, leg. H. Shigetoh; * 34’, 19 (14: HAPC; 2, 19: HIPC), same but with “2.1I1.2021”; Tonaki-jima Island: + 1¢', 12 (HIPC), SM | ‘| | V4 \ \ ZY, S tis Figure 6. Argopistes rufus Chen A antenna, male B antenna, female C aedeagus, dorsal view D aedeagus, lateral view 0.1mm Ze E abdominal ventrite VIII, female F spermatheca G gonocoxae. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Figure 7. Habitus of Argopistes rufus Chen A typical color form, male, dorsal view B ditto, ventral view C ditto, lateral view D yellowish brown color form, female, dorsal view E ditto, ventral view F ditto, lateral view G reddish brown color form, male, dorsal view H ditto, ventral view I ditto, lateral view. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 161 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Tonaki-son, Uaki, 1.1X.2018, leg. H. Shigetoh; Tsuken-jima Island: * 19 (HAPC), Uruma-shi, Katsurentsuken, 14-16.VII.2020, leg. H. Shigetoh; Yonaguni-jima Is- land: * 14, 12 (HAPC), Kita-Bokujé, 28.1II.2001, leg. S. Tsuyuki. Additional material examined. CHINA. Guandong: « 7, 62 (TARI), Yangtais- han (fH Gil), 23.1V.2022, leg. Y.-Y. Ruan; * 134, 112 (TARI), Wutongshan (#24 iL), 5.IV.2023, leg. Y.-Y. Ruan; Hong Kong: * 2, 12 (NHMUK), 56 / 157, 894 / 7/8/63; * 13 (NHMUK), Walker Coll., 93—58; * 14 (NHMUK), Tailung National Park, 12.111.1963, leg. P. Y. So; JAPAN. Honshu. Gumma: * 12 (NHMUK), Mae- bashi-shi, lwakami-ché, 18.IV.2003, leg. Y. Komiya; Okayama: * 19 (HAPC), Mi- masaka-shi, Yono, 10.IV.2016, leg. H. Suenaga; * 14 (HAPC), Okayama-shi, Ki- ta-ku, Kibi service area, 1.VII.2012, leg. O. Yamaji; * 2¢', 22 (HAPC), Tsuyama-shi, Yamakita, 3.IV.2014, leg. H. Suenaga; Tokyo: + 3¢', 39 (HIPC), Hachidji-shi, Kinu- gaoka, 18.VI.2016, leg. H. Shigetoh; Hachij6-jima Island: * 2<', 29 (SEHU), Okag6, 3.VIII.1963, leg. Y. Kamiya; Ogasawara Haha-jima Island: * 14 (SEHU), Funamid- ai, 24.V1.1987, leg. H. Akiyama; Kyushu. Fukuoka: * 12 (HAPC), Fukuoka-shi, Higashi-ku, Hakozaki Kyushu Univ., 7.VI.2008, leg. Y. Matsumura; + 12 (HAPC), Fukuoka-shi, Hakozaki, 16.VIII.2011, leg. H. Suenaga; Kagoshima: Koshiki-jima Island: * 19 (SEHU), Teuchi, 16.V.1965, leg. Y. Komiya; the Ryukyus. Okinawa: Ki- ta-dait6-jima Island: * 2¢, 29 (HIPC), Kita-dait6-jima, 21.IV.2018, leg. H. Kawase; TAIWAN. Hsinchu: * 3, 69 (TARI), Hsinchu (#177 TH), 23.IV.2021, leg. C.-Y. Tsai; Kinmen: Kinmen Island (4=/"4 &): * 19 (TARI), Botanic Park (444 fl), 12.V1I.2023, leg. C.-F. Lee; * 34, 149 (TARI), Jinsha (47), 12.1V.2023, leg. C.-F. Lee; Matsu Islands: + 104, 119 (TARI), Beigan Island (Jif 5), 12.IV.2024, leg. C.-F. Lee; * 63, 109 (TARI), Nangan Island (fF), 12.IV.2024, leg. C.-F. Lee; Nantou: * 4¢', 59 (TARI), Chichi (4242), 26.V.2023, leg. T.-W. Hsu; * 24 (TARI), Mingchien (4 fi), 14.VII.2022, leg. Y.-J. Tung; Taipei: - 1,19 (TARI), Kuantu (7), 8.1V.2020, leg. M.-H. Tsou; * 44’, 19 (TARI), same locality, 18.X.2010, leg. S.-F. Yu; = 12 (TARI), same but with “20.11.2011”; + 53, 69 (TARI), Kuanyinshan (iL), 21.11.2016, leg. H.-T. Cheng; * 2<' (TARI), same locality, 20.V.2011, leg. H. Lee. Diagnosis. Adults of A. rufus look similar to those of A. biplagiatus with a similar color pattern, but differ from A. biplagiatus in having lines of punctures much coarser than those between the lines (lined punctures slightly coarser than those between lines, sometime confused in A. biplagiatus) and a narrower interspace between eyes. Genitalic characters are diagnostic for both species. Those of A. rufus possess widely rounded apices (Fig. 6C) and the aedeagus is narrow in lateral view (Fig. 6D) (pointed apex (Fig. 3C) and wider aedeagus in lateral view (Fig. 3D) in A. biplagiatus); females have medially widened gono- coxae (Fig. 6G) (narrow and parallel-sided base of gonocoxae (Fig. 3G) in A. biplagiatus), and abdominal ventrite VIII medially depressed and without setae on median area (Fig. 6E) (evenly rounded and with dense setae on apical mar- gin of abdominal ventrite VIII (Fig. 3E) in A. biplagiatus). In addition, adults of A. rufus in Taiwan are smaller (3.8-4.3 mm) than those of A. biplagiatus (4.7-4.9 mm). Moreover, distinct color patterns occur to both species respectively (yellowish brown elytra with distinct arrangement of black spots in A. rufus; black elytra with reddish brown central area in A. biplagiatus). Redescription. Length 3.6—4.3 mm, width 2.9-3.4 mm. Color variable (see below). Pronotum broad, convex, lateral margin narrowly explanate; 2.1-2.2 x wider than long, disc with dense fine punctures; lateral margin rounded, anterior margin strongly concave, posterior margin moderately convex. Elytra broadly ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 162 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan oval, 1.1 x longer than wide, disc with coarse punctures arranged into longitudi- nal striae and with dense fine punctures between striae. Male. Antenna filiform (Fig. 6A), antennomere | much longer than others, ap- proximate ratios of length of antennomeres I-XI 1.0: 0.4: 0.3: 0.4: 0.4: 0.4: 0.4: 0.4: 0.4: 0.4: 0.5; approximate ratios of length to width of antennomeres |-XI 4.3: 2.0: 2.0: 2.0: 2.0: 1.8: 1.5: 1.6: 1.7: 1.7: 2.7. Aedeagus (Fig. 6C, D) apically and strongly narrowed from apical 1/3, apex truncate; anterior opening large, ~ 0.52 as long as aedeagus, from apical 1/8-3/5; tectum composed of one pair of sclerotized processes with bifurcate apices, outer apex hooked, small, ~ 0.36 as long as anterior opening; narrow and slightly curved in lateral view; paired processes apically curved in lateral view; endophallic sclerite laterally flattened, with small process near apex, and with basal processes. Female. Antenna (Fig. 6B) similar to males, ratios of length of antennomeres I-XI 1.0: 0.4: 0.3: 0.4: 0.4: 0.3: 0.3: 0.4: 0.4: 0.4: 0.5; ratios of length to width of antennomeres III-XI 4.4: 1.9: 2.0: 2.2: 1.8: 1.6: 1.4: 1.7: 1.6: 1.6: 2.3. Ventrite VIII (Fig. 5E) weakly sclerotized, T-shaped, with several setae along apical margin, apical margin medially depressed, spiculum long. Spermathecal receptaculum (Fig. 6F) longer than pump, moderately swollen, curved in lateral view; pump emarginate at inner side of base; spermathecal duct with long basal part, ra- mus rounded. Gonocoxae (Fig. 6G) wide and separated, base membranous, each gonocoxa asymmetric, apically narrowed from basal 1/3, with sparse long setae along apical areas. Color variation. In Japanese populations, antennae yellowish brown; pro- notum and elytra black, each elytron with one large red spot, sometimes wid- ened and spots connected to each other, red spot reduced in some individu- als; mesoventrite and abdominal ventrites reddish brown but medially black; femora blackish brown, tibiae dark brown, tarsi yellowish brown; few individ- uals have entirely reddish-brown bodies. In the Ryukyus, adults usually have larger red spots on the elytra and reddish-brown elytral margins (described as A. ryukyuensis Shigetoh & Suenaga, 2022). On Taiwan Island, adults separate into two color forms. Typical form (Fig. 7A—-C): black elytron with one large red spot, same as Japanese popu- lations; yellowish brown color form (Fig. 7D—F): elytra with wide black stripe along suture, starting from base, apically narrowed and abbreviated at basal 1/3, with two pairs of black spots halfway between suture and lateral margin, anterior pair at base, posterior pair at apical 1/3, one wide black stripe along lateral margin, starting from base, apically narrowed, abbreviated at basal 1/3 or 1/4; abdominal ventrites medially darker. This color form is also found in Nangan Island. In China and Kinmen Island, almost all adults belong to the typical form. A few specimens have entirely reddish-brown bodies (Fig. 7G-l). Three speci- mens collected from Hong Kong also have reddish bodies. Host plants. Inoue (2014) recorded Osmanthus x fortunei, O. heterophyllus, O. fragrans (#£7¢), O. fragrans var. aurantiacus, O. insularis Koidz, Ligustrum ja- ponicum (H Ax), L. licidum, L. ovalifolium, L. obtusifolium, Syringa vulgaris, S. reticulata, Jasminum nudiflorum Lindl., and Olea europaea L. as host plants in Japan. In Taiwan, larvae mine the leaves of the following plants: Chionanthus retusus (in Taiwan Island, Nangan, and Beigan islands), Ligustrum japonicum (in Beigan island), and Osmanthus fragrans (in Kinmen Island). ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 163 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Biology. Various aspects of the biology of A. rufus were studied in Japan, includ- ing feeding habits, habitat selection, seasonal development, and developmental biology on various host trees, developmental success of larvae on two different host trees, seasonal trends of feeding and oviposition activities of adults, effects of food condition on oviposition, overwintering and oviposition ability of adults that emerge late in the season, effects of photoperiod and temperature on induc- tion of reproductive diapause in newly emergence adults, and occurrence on olive trees (Inoue and Shinkaji 1989a-—c, 1990; Inoue 1990b, 1991a, 1998, 2001, 2014). The seasonal development of this species was studied in the field in southern Kanto, Central Japan (Inoue 1996). Overwintered adults appeared on host trees beginning mid-March, with a peak in mid-April to early May. Females began to deposit eggs from mid- to late April. The eggs were laid singly, embedded in young leaves, and coated with excrement. Leaf-mining larvae only developed in new leaves. Larvae underwent three larval instars and mature larvae crawled down to pupate in the upper layers of soil. Adults eclosed in mid-June, with a peak in later June-early July. They mainly fed on mature leaves. Adults passed the winter near the ground, mainly under fallen leaves. The egg, larval, prepupal, and pupal period took ~ 10, 20-30, 10-15, and 10-15 days respectively during spring to early summer. In Taiwan, larvae and adults can be found during April. Remarks. Argopistes biplagiatus var. rufus was described by Chen (1934a) based on four reddish brown individuals (Fig. 7G—1) deposited in the NMHUK. We found the determination label: “Argopistes / biplagiatus / var. rufa”, hand- written by Chen pinned with one typical form (Fig. 7A-C). Four adjacent fe- males fit the original description (reddish brown body form) and bore two labels “China / Bowring” although no determination labels were found. Thus, those specimens were designated as lectotypes and paralectotypes. Bezdek and Konstantinov (2024) placed this name as a junior synonym of A. coccinel- liformis Csiki, 1940. Actually, it is a distinct species and attributed to the oldest available name. Thus, the valid name is Argopistes rufus Chen, 1934, stat. nov. Adults of A. rufus and A. ryukyuensis are not separable when Taiwanese and Chinese specimens are included. Aedeagi of both areas are intermediate be- tween A. rufus and A. ryukyuensis. Moreover, one distinct color pattern (yel- lowish-brown elytra with black spots) occurs in Taiwanese populations. Thus, color patterns may not be considered as diagnostic characters. Other diagnos- tic characters provided by Shigetoh and Suenaga (2022) are not diagnostic for species delimitation. Thus A. ryukyuensis Shigetoh & Suenaga, 2022 is regard- ed as junior synonym of A. rufus Chen, 1934. Distribution. China, Japan (Honshu, the Izu Isls., Ogasawara Isls., Shikoku, Kyushu, Okinoshima Is., Kashiwa-jima Is., the Koshiki-jima Isls., Yakushima Is., the Ryukyu Isls.), Taiwan including Kinmen Island and Matsu Islands (Beigan and Nangan Islands) (Fig. 5). Argopistes tsekooni Chen, 1934 Figs 8A-C, 9 Argopistes tsekooni Chen, 1934b: 316 (China: Shanghai, Hangchow); Csiki 1940: 525 (catalogue); ChOj6 and Kimoto 1961: 174 (China, Japan); Gressitt and Kimoto 1963: 813 (China: Jiangsu); Kimoto 1965: 437 (redescription); ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 164 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Lee and An 2001: 183 (South Korea); Lee and Cho 2006: 91 (host plant); Takizwa, 2012: 38 (faunistics); Cho and An 2020: 15 (North Korea); Won et al. 2023: 9 (South Korea: Ulleungdo). Argopistes biplagiatus: Baly 1874: 202 (misidentification). Type material examined. One syntype + (sex undetermined, IZAS): “YZ. (= Zhe- jiang): tit (= Hangchow) / 1934. [h] / A EI#}S2 [se (= Chinese Academy of Sci- ences) [h, p] // 27k is##t (attacking Ligustrum obtusifolium) [h, w] // Argopistes / tsekooni / Chen [h, w]”. Although this specimen does not bear any type label, it should be regarded as type specimen since it fit the original description. Additional material examined. JAPAN. * 12 (NHMUk): “Argopistes / biplagia- tus / Motsch / Japan [h, w] // Baly Coll. [p, w]”; Honshu. Shizuoka: * 14 (SEHU), Tagata-gun, Tohi, 4.V.1985, leg. Y. Komiya; Tokyo: + 1< (HAPC), Komae-shi, Ko- mai-machi, 10.VI.2021, leg. R. Seki; Yamaguchi: * 14 (NHMUK); Kyushu. Fu- kuoka: * 19 (HAPC), Fukuoka-shi, Higashi-ku, Shimobaru (alt. 100-360 m), 27.V.2009, leg. S. Sejima; * 19 (NHMUK), Mt. Mikazuki, 2.V.1954, leg. K. Morim- oto; Nagasaki: * 14, 29 (SEHU), Sasebo-shi, Mt. Yahirodake, 14.1V.1981, leg. J. Okuma; * 14’ (SEHU), same locality but with “21.1V.1981”; Oita: * 2’, 32 (HAPC), Hita-shi, Miwa, Chikura, 11.IV.2016, leg. S. Sasaki. Diagnosis. Adults of A. tsekooni are recognized easily by their small body sizes (< 3.5 mm; > 3.5 mm in others except A. unicolor), elongate ovate body shapes (elytra 1.2 x longer than wide; but 1.1 x longer than wide in others), and the combined red spots on elytra (usually separate red spots on the elytra in others); additionally, most genitalic characters are unique, such as the tube-like apex of the aedeagus (Fig. 9C); few setae on apical margin of abdominal ven- trite VIII in females (Fig. 9E); and transverse gonocoxae with dense, long setae on the widely rounded apical margin (Fig. 9G). Redescription. Length 2.8-3.2 mm, width 2.1-2.4 mm. Color (Fig. 8A—-C) blackish brown, elytron with one transverse orange area at basal 1/3, and nar- rowed towards suture; tarsi and front tibiae yellow; antennae dark brown but seven basal antennomeres yellow. Pronotum broad, convex, lateral margin nar- rowly explanate; 2.0-2.1 x wider than long, disc with dense coarse punctures; lateral margin rounded, anterior margin strongly concave, posterior margin moderately convex. Elytra elongate oval, 1.2 x longer than wide, disc with con- fused, dense, coarse punctures. Male. Antenna filiform (Fig. 9A), antennomere | much longer than others, ap- proximate ratios of length of antennomeres I-XI 1.0: 0.4: 0.3: 0.5: 0.5: 0.5: 0.5: 0.5: 0.5: 0.5: 0.7; approximate ratios of length to width of antennomeres |-XI 3.1: 1.8: 2.0: 2.5: 2.3: 1.9: 1.8: 2.1: 2.1: 2.2: 2.9. Aedeagus (Fig. 9C, D) gradually widened from basal 1/9-1/2, then gradually narrowed to basal 1/2, strongly widened posterior—basal 1/2; anterior opening large, ~ 0.39 as long as aedea- gus, from apex to apical 2/5; tectum composed of one pair of sclerotized pro- cesses, long, ~ 0.85 as long as anterior opening, wide and slightly curved from basal 2/4 to apex in lateral view, recurved near apex; endophallic sclerite later- ally flattened, with base twisted. Female. Antenna (Fig. 9B) similar to males, but antennomeres VII—X wider, ra- tios of length of antennomeres I—XI 1.0: 0.4: 0.3: 0.4: 0.4: 0.4: 0.4: 0.4: 0.4: 0.4: 0.6; ratios of length to width of antennomeres I-XI 3.8: 1.8: 2.0: 1.8: 1.9: 1.9: 1.6: 1.5: 1.5: 1.5: 2.4. Ventrite VIII (Fig. 9E) weakly sclerotized, only part of apical margin ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 165 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Figure 8. Habitus of Argopistes species A A. tsekooni Chen, male, dorsal view B ditto, ventral view C ditto, lateral view D A. unicolor Jacoby, female, dorsal view E ditto, ventral view F ditto, lateral view. well sclerotized, with several setae along apical margin, spiculum long and base wider. Spermathecal receptaculum (Fig. 9F) longer than pump, moderately swol- len, curved in lateral view; pump emarginated at inner side of base; spermathecal duct with long basal part, ramus truncate. Gonocoxae (Fig. 9G) wide and separat- ed, base membranous, each gonocoxa asymmetric, apically narrowed from near base, with sparse setae along apical areas, setae longer at apical 1/2. Color variation. One male has a black body and lacks transparent spots on elytra. Another male has an entire yellowish-brown body. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 166 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Figure 9. Argopistes tsekooni Chen A antenna, male B antenna, female C aedeagus, dorsal view D aedeagus, lateral view E abdominal ventrite VIII, female F spermatheca G gonocoxae. Host plants. Oleaceae: Ligustrum obtusifolium (Chij6 & Kimoto, 1961); Sy- ringa oblata Lindl., L. japonicum, L. licidum, and L. sinense (Zhang et al. 2008b). Biology. The biology and life history of A. tsekooni were studied under labora- tory and outdoor conditions in Huangshan City of Anhui Province, China (Zhang ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 167 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan et al. 2009). Argopistes tsekooni overwintered as adults and had three overlap- ping generations in Anhui Province. Distribution. China, Japan (Honshu, Kyushu, the Goto Isls., Hirado-jima Is. Tsushima Is.), North Korea, South Korea. Argopistes unicolor Jacoby, 1885 Figs 8D-F, 10 Argopistes unicolor Jacoby, 1885: 738 (Japan: Yuyama); Chuj6 1936: 109 (cata- logue); Csiki 1940: 524 (catalogue); ChOj6 and Kimoto 1961: 174 (catalogue); Kimoto 1965: 438 (redescription); Takizawa, 2012: 38 (faunistics). Type material examined. Lectotype « 3 (here designated, NHMUk): “(aedeagus glued on the transparent card) // Yuyama / 10.V.-14.V.81. [p, w] // Japan / G. Lew- is. / 1910-320 [p, w] // Type / H.T. [p, w, circle label with red border] // Argopistes / unicolor. Jac. [h, b] // Argopistes unicolor JACOBY, / LECTOTYPUS 1885 / J. Kral m. dit 1969! [h, w] // lecto- / typus [p, r]”. Paralectotype. + 14 (TARI): “Yuyama [h] / Japan [p] / 10.V.1881 [h] / Col. G. LEWIS [p, w] // Argopistes / unicolor Jac. [h] / Det. T. Shiraki [p, w] // Co / Type [p, w, circle label with yellow letters and border] // Argopistes / unicolor Jacoby [h] // DET. M. CHUJO [p, w] // 1527 [p, w]”. Additional material examined. JAPAN. Kyushu. Nagasaki: * 3, 39 (SEHU), Shimbara-shi, Senbuki, 8.V.1984, leg. S. Imsaka; * 23’, 29 (HAPC), same but with “Senfujiki”, collected on Ligustrum japonicum. Diagnosis. Adults of A. unico/or are recognized easily by their small body siz- es (< 3.5 mm; > 3.5 mm in others except A. tsekooni), black antenna with three basal antennomeres paler (entirely yellowish-brown antennae in others except A. tsekooni with five dark apical antennomeres), and the entirely black elytra. Additionally, most genitalic characters are unique, including strongly curved ae- deagus in lateral view (Fig. 10E), and anterior opening from apex to middle (Fig. 10C); straight apical margin of abdominal ventrite VIII in females (Fig. 10F) with setae reduced at medial area (other species with straight margin of abdominal ventrite VIII in female always with setae on median area); and longitudinally square gonocoxae (Fig. 10H). Redescription. Length 3.2-3.4 mm, width 2.3-2.5 mm. Color (Fig. 8D-F) black; legs and mouthparts dark brown; antenna black but three basal antenno- meres dark brown; abdominal ventrites yellowish brown but medially darkened. Pronotum broad, convex, lateral margins narrowly explanate; 1.9-2.0 x wider than long, disc with dense coarse punctures; lateral margin rounded, anterior margin strongly concave, posterior margin moderately convex. Elytra broadly oval, 1.1 x longer than wide, disc with coarse punctures arranged into longitudi- nal striae, and with fine punctures between striae. Male. Antenna filiform (Fig. 10A), antennomere | much longer than others, approximate ratios of length of antennomeres I-XI 1.0: 0.4: 0.3: 0.4: 0.4: 0.4: 0.4: 0.4: 0.4: 0.5: 0.7; approximate ratio of length to width of antennomeres I- XI 3.8: 1.8: 1.8: 2.0: 1.8: 1.6: 1.6: 1.7: 1.5: 1.6: 2.7. Aedeagus (Fig. 10C-E) widest at apical 1/4, slightly narrowed at middle, apically narrowed from apical 1/4, apex broadly rounded; anterior opening large, ~ 0.45 as long as aedeagus, from apex to middle; tectum composed of one pair of sclerotized processes, long, ~ 0.78 as ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 168 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan 0.2 mm E F HY/ Figure 10. Argopistes unicolor Jacoby A antenna, male B antenna, female C apex of aedeagus, front view D base of ae- deagus, dorsal view E aedeagus, lateral view F abdominal ventrite VIII, female G spermatheca H gonocoxae. 0.1 mm long as anterior opening, paired processes with apices recurved in lateral view; endophallic sclerite laterally flattened, with one pair of long apical processes. Female. Antenna (Fig. 10B) similar to males, ratios of length of antenno- meres I-XI 1.0: 0.4: 0.3: 0.3: 0.4: 0.3: 0.4: 0.4: 0.4: 0.4: 0.6; ratios of length to width of antennomeres III—XI 3.4: 1.9: 1.9: 1.9: 1.7: 1.5: 1.6: 1.7: 1.5: 1.6: 2.4. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 169 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Ventrite VIII (Fig. 10F) weakly sclerotized, T-shaped, with several pairs of setae along apical margin, spiculum long. Spermathecal receptaculum (Fig. 10G) lon- ger and wider than pump, moderately swollen; pump emarginate at inner side of base; spermathecal duct with long basal part, ramus rounded. Gonocoxae (Fig. 10H) wide and separated, base membranous, each gonocoxa subquad- rate, with sparse setae along apical areas. Color variation. One male has a black body and lacks transparent spots on the elytra. Another male has an entire yellowish-brown body. Host plants. Oleaceae: Osmanthus heterophyllus (= Olea ilicifolia Hassk.) (Chdjé6 and Kimoto 1961), Ligustrum japonicum (based on collecting data). Biology. Unknown. Distribution. Japan (Honshu, Kyushu, Hirado-jima Is.). Argopistes jungchani sp. nov. https://zoobank.org/313C271 1-2603-4D4D-A690-8E060F85480C Figs 11A, B, 12 Type material examined. Holotype - ¢ (TARI). TAIWAN. Pingtung: Jinshuiying (7K), 14.IV.2021, leg. J.-C. Chen. Paratype - 12 (TARI), same but with “27. VI.2012”. Diagnosis. Adults of this new species are easily recognized by their color pattern: black bodies with yellowish-brown lateral margins of pronotum and elytra; also, genitalic characters are diagnostic: tube-like apex of aedeagus sim- ilar to that of A. tsekooni but parallel-sided from near apex to middle (Fig. 12C) (apically narrowed from near apex to middle (Fig. 9C) in A. tsekooni), paired elongate tectum small, 0.76 as long as anterior opening (Fig. 12C) (paired elon- gate tectum long, 0.85 as long as anterior opening (Fig. 9C) in A. tsekooni), and anterior opening from apical 1/13-2/5 (Fig. 12C) (anterior opening from apex to apical 2/5 (Fig. 9C) in A. tsekooni); only two pair of long setae on apical margin of abdominal ventrite VIII (Fig. 12E) in females (more than two pair of setae on apical margin of abdominal ventrite VIII in females of other species), and cylindrical gonocoxae (Fig. 12G). Description. Length 3.5-3.6 mm, width 2.7-2.9 mm. Color (Fig. 11A, B) blackish brown, sides of pronotum and elytra paled, tarsi, front femur and tibi- ae, and antennae yellowish brown. Pronotum broad, convex, lateral margin nar- rowly explanate; 2.3 x wider than long, disc with dense coarse punctures; lateral margin rounded, anterior margin strongly concave, posterior margin moderate- ly convex. Elytra broadly oval, 1.1 x longer than wide, disc with fine punctures arranged into longitudinal lines, confused, dense, fine punctures present be- tween longitudinal punctures. Male. Antenna filiform (Fig. 12A), antennomere | much longer than others, approximate ratios of length of antennomeres |-XI 1.0: 0.5: 0.3: 0.5: 0.5: 0.5: 0.5: 0.5: 0.5: 0.5: 0.6; approximate ratios of length to width of antennomeres I-X1 4.4: 2.2: 1.9: 2.6: 2.3: 2.2: 1.9: 1.8: 1.7: 1.7: 2.6. Aedeagus (Fig. 12C, D) parallel-sided, strongly and subapically narrowed, apex tube-like and extremely small; anterior opening small, ~ 0.30 as long as aedeagus, from apical 1/13- 2/5; tectum composed of one pair of sclerotized processes, large, ~ 0.76 as long as anterior opening; wide and slightly curved in lateral view; paired pro- ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 170 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Figure 11. Habitus of Argopistes species A A. jungchani sp. nov., female, dorsal view B ditto, lateral view C A. tsoui sp. nov., male, dorsal view D ditto, ventral view E ditto, lateral view. = —<— cesses apically curved in lateral view; endophallic sclerite laterally flattened, with basal processes slightly sclerotized. Female. Antenna (Fig. 12B) much smaller than in males, ratios of length of antennomeres I-XI 1.0: 0.4: 0.3: 0.3: 0.3: 0.3: 0.3: 0.4: 0.4: 0.4: 0.6; ratios of length to width of antennomeres I-XI 4.2: 1.8: 1.8: 1.6: 1.7: 1.5: 1.5: 1.7: 1.6: 1.4: 2.3. Ventrite VIII (Fig. 12E) membranous, only apical margin sclerotized, T-shaped, with two pairs of short setae at sides of apical margin, apical margin truncate, spiculum long. Spermathecal receptaculum (Fig. 12F) much longer than pump, moderately swollen; pump slightly emarginate at inner side of base; spermathecal duct with long basal part, ramus rounded. Gonocoxae (Fig. 12G) cylindrical and separated, base membranous, each gonocoxa symmetric, with dense long setae along apical and outer margin. Host plant. Unknown Biology. Unknown. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 171 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Etymology. This new species is named for Jung-Chan Chen (28 =), the first person to collect specimens. Distribution. Only known from the type locality (Fig. 5). 0.1 mm Figure 12. Argopistes jungchani sp. nov. A antenna, male B antenna, female C aedeagus, dorsal view D aedeagus, lateral view E abdominal ventrite VIII, female F spermatheca G gonocoxae. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Argopistes tsoui sp. nov. https://zoobank.org/7FA87DFC-55AB-45B2-AD05-A2EA64530D90 Figs 11C, D, 13, 14A, B Type material examined. Holotype - 3 (TARI). TAIWAN. Hsinchu: Tahunshan (Ai), 24.11.2009, leg. S.-F. Yu. * Paratypes * 14, 19 (TARI), same as holo- type; * 12 (TARI), same but with “8.1X.2009”; Ilan: * 3, 39 (TARI), Fushan (48 i) Chihwuyan (#44 ] = Botanic Park), 15.11.2009, leg. M.-H. Tsou; * 44, 69 (TARI), same locality, 8.VI.2023, leg. S.-S. Lu; * 14’ (TAFI), same but with “27. X1.2023”; + 19 (TAFI), same but with “28.X1.2023”; * 14, 12 (TAFI), same but with “29.X1.2023”; © 14, 19 (TARI), same but with “5.XII.2023”; - 19 (TARI), same but with “7.XII.2023”; * 19 (TARI), same but with “18.XII.2023”; « 13 (TARI), same but with “11.1.2024”; + 14 (TARI), same but with “8.111.2024”; - 44, 39 (TARI), same but with “7.V.2024”: Keelung: * 14, 12 (TARI), Hungtanshan (AL 1), 10.V.2008, leg. M.-H. Tsou; Pingtung: * 12 (TARI), Lilungshan (£7 LL), 5.X1.2009, leg. M.-H. Tsou; * 19 (TARI), Tahanshan (AY LL), 22.1.2009, leg. S.-F. Yu; > 12 (TARI), same locality, 20.VIII.2022, leg. Y.-T. Chung; Taoyuan: * 2 (TARI), Tungyanshan (3H iL), 8.VII.2007, leg. M.-H. Tsou; * 19 (TARI), Yongfu (7k #4), 24.11.2014, leg. H. Lee. Diagnosis. Adults of A. tsoui sp. nov. are similar to those of A. biplagiatus with reddish-brown elytra with wide black lateral margins, but differ from A. bi- plagiatus by the reddish-brown pronotum with wide black lateral margins (en- tirely black pronotum in A. biplagiatus). Diagnostic genitalic characters include pointed apex of aedeagus similar (Fig. 13C) to that of A. biplagiatus (Fig. 3C) but relatively narrower in lateral view (Fig. 13D) (relatively wider (Fig. 3D) in A. biplagiatus), longer, longitudinal paired sclerites of tectum (Fig. 13C) (short, curved paired sclerites of tectum (Fig. 3C) in A. biplagiatus), anterior opening from apical 1/10 to middle (Fig. 13C) (anterior opening from apex to apical 3/10 (Fig. 3C) in A. biplagiatus); triangular gonocoxae similar to those of A. rufus but expanding inwardly at basal 1/3 (Fig. 13G) (expanding outward at basal 1/3 (Fig. 6G) in A. rufus); dense setae along apical margin of abdominal ventrite VIII similar to those of A. biplagiatus but much denser and shorter (Fig. 13C) (less denser and longer setae on apical margin of abdominal ventrite VIII (Fig. 3E) in A. biplagiatus). Description. Length 3.9-4.3 mm, width 3.2-3.5 mm. Color (Fig. 11C-E) red- dish brown, sides of pronotum and elytra darker, tarsi and antennae yellow. Pronotum broad, convex, lateral margin narrowly explanate; 2.2 x wider than long, disc with dense coarse punctures; lateral margin rounded, anterior mar- gin strongly concave, posterior margin moderately convex. Elytra broadly oval, 1.0-1.1 x longer than wide, disc with dense, confused, coarse punctures. Male. Antenna filiform (Fig. 13A), antennomere | much longer than others, approximate ratios of length of antennomeres I-XI 1.0: 0.4: 0.3: 0.4: 0.4: 0.4: 0.4: 0.4: 0.4: 0.4: 0.6; approximate ratios of length to width of antennomeres I- XI 4.7: 2.1: 2.2: 2.4: 2.0: 2.0: 1.9: 1.9: 1.9: 1.7: 2.7. Aedeagus (Fig. 13B, C) strong- ly narrowed from apical 1/5 to apex, apex pointed; anterior opening small, ~ 0.35 as long as aedeagus, from apical 1/10 to middle; tectum composed of one pair of sclerotized processes with bifurcate apices, outer apex hooked, large, ~ 1.1 as long as anterior opening; narrow and slightly curved in lateral view; endophallic sclerite laterally flattened, with small process near apex, and with basal processes membranous. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 173 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan ALU Yl Y) 0.2 mm 0.2 mm G Figure 13. Argopistes tsoui sp. nov. A antenna, male B antenna, female C aedeagus, dorsal view D aedeagus, lateral view E abdominal ventrite VIII, female F spermatheca G gonocoxae. Female. Antenna (Fig. 13B) similar to males, ratios of length of antenno- meres I-XI 1.0: 0.4: 0.3: 0.4: 0.4: 0.3: 0.4: 0.4: 0.4: 0.4: 0.5; ratios of length to width of antennomeres III-—XI 4.1: 2.2: 2.4: 2.4: 2.2: 1.9: 2.1: 2.0: 1.9: 1.8: 2.9. Ventrite VIII (Fig. 13E) membranous, only apical margin sclerotized, T-shaped, with dense short setae along apical margin, spiculum long. Spermathecal re- ceptaculum (Fig. 13F) longer than pump, moderately swollen, curved in lateral ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 174 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Figure 14. Field photographs of Argopistes species A adults of A. tsoui sp. nov. feeding on leaves of Osmanthus hetero- phyllus B close-up shot of adults of A. tsoui sp. nov. C adult of A. yuae sp. nov. resting on underside of leaf of Chionanthus ramiflorus D larva mining new leaf of C. ramiflorus. view; pump emarginate at inner side of base; spermathecal duct with elongate basal part, ramus rounded. Gonocoxae (Fig. 13G) wide and separated, base membranous, each gonocoxa asymmetric, apically narrowed from apical 1/3, with dense long setae along apical areas. Host plant. Oleaceae: Osmanthus heterophyllus (Fig. 13A, B), O. kaoi (T. S. Liu & J. C. Liao) S. Y. Lu, O. enervius Masam. & T. Mori, O. fragrans. Biology. This species seems to be univoltine. The larvae were found only during late March. Etymology. This new species is named for Mei-Hua Tsou (#/ 5 #£), the first person to collect specimens. Distribution. This new species is widespread in lowlands of Taiwan (Fig. 5). Argopistes yuae sp. nov. https://zoobank.org/D4D59AB2-7496-44F6-AD3B-C05251B4E355 Figs 14C, D, 15, 16 Type material examined. Holotype : ¢ (TARI). TAIWAN. Taitung: Lanyu (fH), 16.1V.2023, leg. Y.-F. Hsu. Paratypes + 116, 72 (TARI), same data as holotype; * 104, 59 (TARI), same but with “20.1I1.2023”; * 84’, 59 (TARI), same but with “17. V1.2023”; + 23’, 32 (TARI), same island, 14.1II.2023, leg. Y.-Y. Liu & Y.-F. Hsu; * 12 (TARI), same island, 28.IV.2022, leg. S-F. Yu; * 14 (TARI), same island, 4.IV.2016, ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 15 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan D E Figure 15. Habitus of Argopistes yuae sp. nov. A typical color form, male, dorsal view B ditto, ventral view C ditto, lateral view D Black color form, female, dorsal view E ditto, ventral view F ditto, lateral view. leg. Y.-T. Chung; * 14 (TARI), same island, 14.1V.2013, leg. B.-X. Guo; * 14' (TARI), same island, 26.IV.2009, leg. U. Ong; + 34, 39. (TARI), same island, 18.1I1.2024, leg. Y.-F. Hsu; * 4, 39 (TARI), same island, 24.1V.2024, leg. J.-C. Chen; + 60, 39 (NHMUK), same island, Lanyu Weather Station (flil4a 3244), 22°02.238'N, 121°33.287'E, 26.VII.2008, hand collecting, leg. M. V. L. Barclay & H. Mendel; - 23,192 (TARI), same island, Tataienchih (A Ath), 19.11.2024, leg. Y.-F. Hsu. Diagnosis. Adults of this new species are not separable from those of A. rufus except by genitalic characters, including parallel-sided apex of aedeagus with anterior opening very close to apex of aedeagus, from apical 1/12-3/5 (Fig. 16C) (apically narrowed aedeagus with anterior opening not so close to apex of aedeagus, from apical 1/8-3/5 (Fig. 6C) in A. rufus); deeply notched apical margin of abdominal ventrite VIII (Fig. 16E) in females (shallowly notched apical margin of abdominal ventrite VIII (Fig. 6E) in females of A. rufus). In addition, this new species is restricted to Lanyu Island, and thus is isolated from other ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 176 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Figure 16. Argopistes yuae sp. nov. A antenna, male B antenna, female C aedeagus, dorsal view D aedeagus, lateral view E abdominal ventrite VIII, female F spermatheca G gonocoxae. species geographically. Moreover, larvae and adults of this new species feed on leaves of Chionanthus ramiflorus Roxb. (Fig. 14C, D) but not those of Osmanthus fragrans based on laboratory rearing tests. Thus, both species are allopatric eco- logically since Osmanthus fragrans is one of the host plants for A. rufus. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 177 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Description. Length 4.2—4.3 mm, width 3.5 mm. Color (Fig. 15A—-C) blackish brown, elytron with one large transparent area at basal 1/3, near or connected with suture; tarsi and antennae yellowish brown. Pronotum broad, convex, lat- eral margin narrowly explanate; 2.3 x wider than long, disc with dense, coarse punctures; lateral margin rounded, anterior margin strongly concave, posterior margin moderately convex. Elytra broadly oval, 1.1 x longer than wide, disc with confused, dense, fine punctures. Male. Antenna filiform (Fig. 16A), antennomere | much longer than others, approximate ratios of length of antennomeres |-XI 1.0: 0.4: 0.3: 0.5: 0.4: 0.4: 0.4: 0.5: 0.5: 0.4: 0.6; approximate ratios of length to width of antennomeres I-X1 3.8: 2.0: 2.0: 2.7: 2.0: 1.9: 1.9: 1.9: 1.8: 1.9: 2.8. Aedeagus (Fig. 16C, D) parallel-sided from basal 1/3-2/3, apically narrowed from apical 1/3-1/6, apex tube-like; anterior opening large, ~ 0.53 as long as aedeagus, from api- cal 1/12-3/5; tectum composed of one pair of sclerotized processes, small, ~ 0.43 as long as anterior opening; wide and slightly curved in lateral view; paired processes curved at apical 1/3 in lateral view; endophallic sclerite later- ally flattened, with basal processes slightly sclerotized, and one pair of small processes near apex. Female. Antenna (Fig. 16B) similar to males, ratios of length of antenno- meres I-XI 1.0: 0.4: 0.3: 0.4: 0.4: 0.4: 0.4: 0.4: 0.4: 0.4: 0.5; ratios of length to width of antennomeres III—X1 4.9: 1.9: 2.3: 2.5: 2.0: 1.8: 1.9: 1.8: 1.8: 1.9: 2.5. Ventrite VIII (Fig. 15E) weakly sclerotized, T-shaped, with several pairs of setae along apical margin, setae smaller at sides, apical margin medially depressed, spiculum long. Spermathecal receptaculum (Fig. 16F) longer than pump, mod- erately swollen, curved in lateral view; pump emarginate at inner side of base; spermathecal duct with long basal part, ramus rounded. Gonocoxae (Fig. 16G) wide and separated, base membranous, each gonocoxa asymmetric, apically narrowed from near base, with sparse setae along apical areas, setae longer at apical 1/2. Variation. A few specimens have black bodies and lack red spots on elytra (Fig. 14D-F). Host plant. Oleaceae: Chionanthus ramiflorus Roxb. Biology. This species seems to be univoltine. The larvae (Fig. 14D) were found only during late March. Etymology. This new species is named for Su-Fang Yu (42477), the first per- son to collect specimens. Distribution. Endemic to Lanyu Island (Fig. 5). Key to Taiwanese species of Argopistes Kimoto (1965) provided a key to Japanese species of Argopistes. We think that the key is appropriate for Japan but not other countries due to color variations or/and color patterns for species elsewhere. A key to Taiwanese species of Argopistes is provide as follows: 1 General color yellowish brown, elytra with black spots; aedeagus (Fig. 7C, D) apically narrowed, and apex widely rounded, apical margin of anterior opening far from (apical 1/8) apex of male aedeagus.......... A. rufus Chen —- General color black or reddish Drown ...............ccccccccccessssececeessssceeeeeeesseeeeees 2 ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 178 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan 2 General color reddish brown, with black, wide margins along pronotum and elytra; aedeagus (Fig. 13B, C) strongly narrowed from apical 1/5 to apex, apex pointed; anterior opening from apical 1/10 to middle; narrow and slightly curved in lateral VieW ................cccccccceessseeeeeeees A. tsoui sp. nov. - General color black, with or without yellowish brown margins along prono- LUI PATI SLY deter Me ae. ees ree, rear a, Rese pe A adh vets hie auth tee Rede Dee 3 3 Elytra with one pair of red spots, but lacking yellowish brown margins ALONG PrFOMOLUM ANG ely thes. wie 8. ccceest cease deta Seam ee ovens ran ee ook eee ergo 4 — Elytra without red spots, and with yellowish-brown margins on pronotum and elytra; aedeagus (Fig. 12C, D) parallel-sided, strongly subapically nar- rowed, apex tube-like and extremely small; anterior opening from apical 1/13-2/5; wide and slightly curved in lateral view.....A. jungchani sp. nov. 4 Only coarse punctures on elytra, wider interspace between eyes; aedeagus (Fig. 3C, D) apically and strongly narrowed from apical 1/5, apex pointed; anterior opening very small, from apex to apical 3/10; extremely wide in lateral MGW. casessorcemverrncsedorstnecitenemmmeee recurs Tuas A. biplagiatus Motschulsky - Coarse and fine punctures confused on elytra, narrower interspace be- PWVICCNIEOV CS hitter nis | tascam usahaten he tuat ident ata csiuea te uaa beth setae Reon 5 5 Aedeagus (Fig. 7C, D) apically narrowed, apex widely rounded, apical mar- gin of anterior opening far from apex of male aedeagus (apical 1/8)......... Se eee cst oN acet nna see ahi te URE Sica vee a cee na toe a eee A. rufus Chen - Aedeagus (Fig. 16C, D) apically narrowed, and apex parallel-sided, apex widely rounded, apical margin of anterior opening close to apex of male aedeagus (apioalhly lA}... «ccc get bane mets as Hea east A. yuae sp. nov. Discussion Some Taiwanese species can be identified by their characteristic color patterns, including A. tsoui sp. nov., A. jungchani sp. nov., and the yellowish-brown form of A. rufus Chen. Male aedeagi are more diagnostic. Genital characters in fe- males such as abdominal ventrite VIII, soermatheca, and gonocoxae are more or less diagnostic, but combinations of these morphological characters and biological information can form the basis for a sound taxonomy of the genus. Although members of Argopistes are oligophagous or monophagous on Oleaceae, only A. rufus has established populations in small islands surround- ing Japan, China, and Taiwan. Five islands (Beigan Island 1b“f &, Nangan Is- land “#55, Dongju Island # £5, Xiju Island fH 2, Dongyin Island #4] &) of the Matsu Islands were investigated during the spring 2024 field season. Populations of A. rufus were found on only Beigan Island and Nangan Island feeding on Chionanthus retusus and Ligustrum japonicum. Chionanthus retusus and Ligustrum japonicum were transported to both islands, the largest of the Matsu Islands, as ornamental trees. This supports the idea that A. rufus can be an invasive insect pest, invading islands as a result of exportation of ornamen- tal trees of Oleaceae. In addition, populations of A. rufus expanded dramatically in Kinmen Island (42F'J&). One population was found at one locality of Jin- sha township (4748) in the middle of April 2023 and one adult was collected at another locality (Botanical Park) at Jinhu township (<=34%H) in the summer (July) of 2023. We have now found larvae attacking new leaves of Osmanthus fragrans at a guesthouse in Jinhu township (47 i44$8), in late May 2024. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 179 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Acknowledgements We are grateful to the Taiwan Chrysomelid Research Team (TCRT) and citizen scientists, including Jung-Chan Chen (fk 28 =), Hsing-Tzung Cheng (2) 85), Yi- Ting Chung (#222), Yu-Feng Hsu (4/4), Hsueh Lee (224), Sheng-Shan Lu (4 1), Mei-Hua Tsou (#32), and Su-Fang Yu (497) for assistance in col- lecting material. We thank for Yi-Chia Chiu (582£3%) and Chi-Lung Lee (42 #242) for photographs of specimens, Mei-Hua Tsou (#/5¢2#) and Yu-Feng Hsu (#14 ll) for field photography, and Chih-Kai Yang for identification of host plants. We would like to thank all curators for giving us the opportunity to study the speci- mens from their collections. We thank Messrs. Osaku Yamaji, Hiroaki Shigetoh, Rydta Seki, and Shoma Sejima for providing valuable specimens, and Chang- Chin Chen ([S #98) for assisting our study in various ways. We thank Chris Carlton for reading the draft and editing for American English style. Finally, we thank Maurizio Biondi for reviewing the manuscript, and Ron Beenen for editing the manuscript. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Funding This study was supported by the National Science and Technology Council NSTC 112-2313-B-055-001-MY3 and the Animal and Plant Health Inspection Agency, Ministry of Agriculture 113AS-5.5.4-VP-01. Author contributions Investigation: HS. Supervision: MYC. Writing - original draft: CFL. Author ORCIDs Chi-Feng Lee © https://orcid.org/0000-0003-1996-0557 Ming-Yao Chiang ® https://orcid.org/0000-0002-1514-7086 Data availability All of the data that support the findings of this study are available in the main text. References Baly JS (1874) VI. Catalogue of the phytophagous Coleoptera of Japan, with descrip- tions of the species new to science. Transactions of the Entomological Society of London 1874: 161-217. https://doi.org/10.1111/j.1365-2311.1874.tb00164.x Bezdék J, Konstantinov AS (2024) Supertribe Alticitae. In: Bezdék J, Sekerka L (Eds) Cat- alogue of Palaearctic Coleoptera. Vol. 6/2. Updated and Revised 2™ edn. Chrysom- eloidea (Orsodacnidae, Megalopodidae, Chrysomelidae). Brill, Leiden/Boston, 468- 580. https://doi.org/10.1163/9789004707283 ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 180 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Biondi M, lannella M, D’Alessandro P (2024) Argopistes Motschulsky from Madagascar with descriptions of six new species (Coleoptera, Chrysomelidae, Galerucinae, Altici- ni). ZooKeys 1202: 303-327. https://doi.org/10.3897/zookeys.1202.122977 Blanco J, Konstantinov A (2013) Review of the New World species of the genus Ar- gopistes Motschulsky (Coleoptera: Chrysomelidae: Galerucinae: Alticini). Zootaxa 3626: 249-267. https://doi.org/10.11646/zootaxa.3626.2.3 Chen SH (1934a) On some species of Chrysomelidae (Col.) in the British Museum. Sty- lops 3: 66-72. https://doi.org/10.1111/j.1365-3113.1934.tb01547.x Chen SH (1934b) Revision of the Halticinae (Col. Chrysomelidae) of Yunnan and Tonkin. Sinensia 5: 225-416. Cho HW, An SL (2020) An annotated list of leaf beetles (Coleoptera: Chrysomelidae) of Korea, with comments and new records. Far Eastern Entomologist 404: 1-36. https://doi.org/10.25221/fee.404.1 Chdj6 M (1935a) Chrysomelidae of Loo-choo Archipelago (I). Transactions of the Natu- ral History Society of Formosa 25: 69-89. Chdj6 M (1935b) Chrysomelidae of Loo-choo Archipelago (Il). Transactions of the Natu- ral History Society of Formosa 25: 203-211. Chdjd M (1936) Studies on the Chrysomelidae in the Japanese Empire (VIII). Subfamily Halticinae (6). Transactions of the Natural History Society of Formosa 26: 108-114. Chajo M, Kimoto S (1961) Systematic catalog of Japanese Chrysomelidae. Pacific In- sects 3: 117-202. Csiki E (1940) [new name. In: Heikertinger F, Csiki E: Pars 169.] Chrysomelidae: Halticinae. In: Schenkling S (Ed.) Coleopterorum Catalogus. W. Junk, s-Gravenhage, 337-635. Duckett CN, Daza JD (2004) A new species of flea beetle in the genus Alagoasa Bechyné (Coleoptera: Chrysomelidae: Galerucinae) from highland Costa Rica. Coleopterists Bulletin 58(1): 84-96. https://doi.org/10.1649/608 Gressitt JL, Kimoto S (1963) The Chrysomelidae of China and Korea, Part 2. Pacific Insects Monograph 1B: 301-1026. Inoue T (1990a) Feeding habits and seasonal development of the flea beetle Argopistes biplagiatus Motschulsky (Coleoptera: Chrysomelidae) in Chiba Prefecture, Central Japan. Japanese Journal of Applied Entomology and Zoology 34(2): 153-160. [in Japanese with English summary] https://doi.org/10.1303/jjaez.34.153 Inoue T (1990b) Habitat selection of two flea beetle species of the genus Argopistes (Coleoptera: Chrysomelidae) on their host trees. Japanese Journal of Applied Ento- mology and Zoology 34(3): 217-226. [in Japanese with English summary] https:// doi.org/10.1303/jjaez.34.217 Inoue T (1991a) Studies on the life history of the flea beetle, Argopistes coccinelliform- is Csiki (Coleoptera: Chrysomelidae). V. Effects of Food Conditions on Oviposition. Japanese Journal of Applied Entomology and Zoology 35(3): 213-220. [in Japanese with English summary] https://doi.org/10.1303/jjaez.35.213 Inoue T (1991b) Host plant preference of flea beetle Argopistes biplagiatus Motschulsky adults (Coleoptera: Chrysomelidae). Applied Entomology and Zoology 26(3): 403- 406. https://doi.org/10.1303/aez.26.403 Inoue T (1992) Adult diapause of the flea beetle, Argopistes biplagiatus Motschulsky (Coleoptera: Chrysomelidae). |. Seasonal trends in feeding and oviposition activities. Japanese Journal of Applied Entomology and Zoology 36(3): 159-168. [in Japanese with English summary] https://doi.org/10.1303/jjaez.36.159 Inoue T (1993) Adult diapause of the flea beetle, Argopistes biplagiatus Motschulsky (Coleoptera: Chrysomelidae). Il. Effects of photoperiodic condition on feeding and ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 181 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan oviposition activities. Applied Entomology and Zoology 28(3): 313-318. https://doi. org/10.1303/aez.28.313 Inoue T (1994) Adult diapause of the flea beetle, Argopistes biplagiatus Motschulsky (Coleoptera: Chrysomelidae). Ill. Effects of low temperatures on termination of winter diapause. Applied Entomology and Zoology 29(3): 433-436. https://doi. org/10.1303/aez.29.433 Inoue T (1996) Biology of two Argopistes (Chrysomelidae: Alticinae). In: Jolivet PHA, Cox ML (Eds) Chrysomelidae Biology, Vol. 3: General Studies, Academic Publishing, Amsterdam, 327-355. Inoue T (1998) Studies on the life history of the flea beetle, Argopistes coccinelliform- is Csiki (Coleoptera: Chrysomelidae). VI. Overwintering and oviposition abilities of adults which emerged late in the season. Applied Entomology and Zoology 33(1): 21-27. https://doi.org/10.1303/aez.33.21 Inoue T (2001) Studies on the life history of the flea beetle, Argopistes coccinelliformis Csiki (Coleoptera: Chrysomelidae). VII. Effects of photoperiod and temperature on in- duction of reproductive diapause in newly emerged adults. Applied Entomology and Zoology 36(1): 53-58. https://doi.org/10.1303/aez.2001.53 Inoue T (2014) Life history of the flea beetle, Argopistes coccinelliformis Csiki (Coleop- tera: Chrysomelidae). VIll. Occurrence on olive tree. Japanese Journal of Environ- mental Entomology and Zoology 25(2): 71-73. https://doi.org/10.11257/jjeez.25.71 Inoue T, Shinkaji N (1989a) Studies on the life history of the flea beetle, Argopistes coccinelliformis Csiki (Coleoptera: Chrysomelidae). |. Feeding habits and seasonal development on various trees of Oleaceae. Japanese Journal of Applied Entomol- ogy and Zoology 33(4): 217-222. [in Japanese with English summary] https://doi. org/10.1303/jjaez.33.217 Inoue T, Shinkaji N (1989b) Studies on the life history of the flea beetle, Argopistes coc- cinelliformis Csiki (Coleoptera: Chrysomelidae). Il. Developmental ability on various host trees. Japanese Journal of Applied Entomology and Zoology 33(4): 223-230. [in Japanese with English summary] https://doi.org/10.1303/jjaez.33.223 Inoue T, Shinkaji N (1989c) Studies on the life history of the flea beetle, Argopistes cocci- nelliformis Csiki (Coleoptera: Chrysomelidae). Ill. Developmental success of larvae on two different host plants. Japanese Journal of Applied Entomology and Zoology 33 (4): 262-263. [in Japanese with English summary] https://doi.org/10.1303/jjaez.33.262 Inoue T, Shinkaji N (1990) Studies on the life history of the flea beetle, Argopistes cocci- nelliformis Csiki (Coleoptera: Chrysomelidae). IV. Seasonal trends of feeding and ovi- position activities of adults. Japanese Journal of Applied Entomology and Zoology 34: 55-62. [in Japanese with English summary] https://doi.org/10.1303/jjaez.34.55 Jacoby M (1885) Descriptions of the phytophagous Coleoptera of Japan obtained by Mr. George Lewis during his second journey, from February 1880 to September 1881. — Part ll. Halticinae and Galerucinae. Proceedings of the Scientific Meetings of the Zoological Society of London 1885: 719-755. https://doi.org/10.1111/j.1096-3642.1885.tb02918.x Jolivet P Hawkeswood TJ (1995) Host-plants of Chrysomelidae of the World. Backhuys Publishers, Leiden, 281 pp. Kimoto S (1965) The Chrysomelidae of Japan and the Ryukyu Islands. IX. Subfamily Aliticinae II. Journal of the Faculty of Agriculture, Kyushu University 13: 431-459. https://doi.org/10.5109/22730 Lee JE, An SL (2001) Family Chrysomelidae. Economic Insects of Kore 14. Insecta Ko- reana Supplement 21. National Institute of Agricultural Science and Technology, Su- won, 231 pp. ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 182 Chi-Feng Lee et al.: Argopistes from Japan and Taiwan Lee JE, Cho HW (2006) Leaf beetles in the crops (Coleoptera: Chrysomelidae). Econom- ic Insects of Korea 27. Insecta Koreana Supplement 34. National Institute of Agricul- tural Science and Technology, Suwon, 130 pp. Motschulsky V de (1860) Coléoptéres rapportés de la Sibérie orientale et notamment des pays situés sur le bords du fleuve Amour par MM. Schrenck, Maack, Ditmar, Voz- nessenski ect. In: Reisen und forschungen Petersburg ausgefuhrt und in Verbind- ung mit mehreren Gelehrten herausgegeben. Band II. Zweite Lieferung. Coleopteren. Eggers & Comp., St. Petersburg, 79-257. Schonfeldt H von (1890) Ein Beitrag zur Coleopterenfauna der Liu-Kiu-Inseln. Entomolo- gische Nachrichten 16: 168-175. Shigetoh H, Suenaga H (2022) Description of a new species of the genus Argopistes Motschulsky (Coleoptera, Chrysomelidae, Galerucinae, Alticini) from the Ryukyu Is- lands and Daito Islands, southwestern Japan, and redescription of A. coccinelliformis Csiki. Elytra, New Series, 12: 1-13. Suffrian E (1868) Verzeichniss der von Dr. Gundlach auf der Insel Cuba gesammelten Chrysomelinen. Archiv fur Naturgeschichte 34(1): 163-252. Taiwan Encyclopedia of Life (2024) TaiEOL. https://taieol.tw/ [accessed 3 July 2024] Takizawa H (2012) Notes on biology of Chrysomelidae in Japan (6). Kanagawa-chtho 177: 33-51. [in Japanese] Won MH, Choi JW, Bang WJ, Lee DY, Moon MK, Kim YK, Kim DG, Kim DY, Suh SJ, Choi KS (2023) Insect fauna including unrecorded species in Ulleungdo, South Korea. Biodi- versity Data Journal 11: e100783. https://doi.org/10.3897/BDJ.11.e100783 Zhang Y, Hanula J, Sun J (2008a) Survey for potential insect biological control agents of Ligustrum sinense (Scrophulariales: Oleaceae) in China. Florida Entomologist 91(3): 372-382. https://doi.org/10.1653/001 5-4040(2008)91[372:SFPIBC]2.0.C0;2 Zhang Y, Hanula J, Sun J (2008b) Host specificity of Argopistes tsekooni (Co- leoptera, Chrysomelidae), a potential biological control agent of Chinese privet. Journal of Economic Entomology 101(4): 1146-1151. https://doi. org/10.1603/0022-0493(2008)101[1146:HSOATC]2.0.CO;2 Zhang Y, Sun J, Hanula J (2009) Biology and life history of Argopistes tsekooni (Co- leoptera: Chrysomelidae) in China, a promising biological control agent of Chinese privet. Annals of the Entomological Society of America 102: 508-516. https://doi. org/10.1603/008.102.0322 ZooKeys 1215: 151-183 (2024), DOI: 10.3897/zookeys.1215.134871 183