A new species of the genus Donacia Fabricius (Coleoptera, Chrysomelidae, Donaciinae) from South Korea: morphology, ecology, and COI gene analysis

83 A new species of the genus Donacia Fabricius (Coleoptera, Chrysomelidae, Donaciinae) from South Korea: morphology, ecology, and COI gene analysis Joong Youb Kim1, Yeong-Deok Han1, Jong Eun Lee2 1 Research Center for Endangered Species, National Institute of Ecology, Yeongyang, Republic of Korea 2 Department of Biological Science, Gyeongkuk National University, Andong, Republic of Korea Corresponding author: Jong Eun Lee ([email protected]) Copyright: © Joong Youb Kim 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 Donacia koreana sp. nov. is described from South Korea, with detailed descriptions and illustrations of both the adult and larval stages. A key for species identification, distribution map, host plant associations, oviposition behaviors, feeding patterns, and other ecological information, along with partial sequences of mitochondrial cytochrome c oxidase subunit I, are also presented. Key words: Larva, new species, reed beetles, taxonomy Introduction The genus Donacia Fabricius, commonly known as reed beetles or pond lily leaf beetles, belongs to the family Chrysomelidae. In BugGuide.net and iNaturalist. com, Donaciinae are referred to as aquatic leaf beetles, while Donacia as a genus has not been assigned a more specific common name because the genus is too diverse. It is widely distributed across the world, with many species inhabiting freshwater environments, and plays a significant role in aquatic ecosystems. Most Donacia species occur in diverse freshwater habitats, such as ponds, wetlands, swamps, lakes, and streams. Their unique life cycles and ecological characteristics make them important environmental indicator species (Hayashi 2020; Geiser 2023; Askevold 2024). Adults feed on leaves and flowers above the water surface, whereas larvae live underwater and feed on sap from plant roots. The larvae breathe by tapping the aerenchyma of the plant using two hollow abdominal spiracle hooks connected to their tracheal system (Geiser 2023). Askevold (in press) summarized the known research on this subject, as how donaciines obtain oxygen is not definitively demonstrated to be just one mechanism. In South Korea, seven species belonging to the genus Donacia have been reported (Lee and An 2001; An 2019; Cho and An 2020; KSAE and ESK 2021; Jeong et al. 2024). Within this species list, D. simplex Fabricius was misidentified as D. bicoloricornis Chen, making their accurate differentiation crucial. In particular, Eastern Palearctic Donacia species are characterized by distinct differences in the presence or absence of hairs on the pronotum and elytra, Academic editor: Alexander Konstantinov Received: 18 August 2025 Accepted: 20 October 2025 Published: 21 November 2025 ZooBank: https://zoobank.org/ F7B1C36F-58E6-41EF-A82CDAC2DD6392BC Citation: Kim JY, Han Y-D, Lee JE (2025) A new species of the genus Donacia Fabricius (Coleoptera, Chrysomelidae, Donaciinae) from South Korea: morphology, ecology, and COI gene analysis. ZooKeys 1261: 83–99. https://doi.org/10.3897/ zookeys.1261.169034 ZooKeys 1261: 83–99 (2025) DOI: 10.3897/zookeys.1261.169034 84 ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034 Joong Youb Kim et al.: Donacia koreana sp. nov. from South Korea which has led in part to confusion in classification (Chen 1966; Askevold 1990). Kippenberg (2015) suggested a revised arrangement of Donacia species, proposing several new subgenera and clarifying some existing subgeneric names. Notably, the subgenus name Donacocia Gistel applies to the vast majority of members of Donacia sensu lato. Here, the new species would be placed in the D. (Donacocia) subgenus. The two other species of Donacia known to be in South Korea are both in the subgenus D. (Cyphogaster) Goecke. Adult D. (Donacocia) are typically characterized within the Donaciini tribe by the coarsely punctate-rugose pronotum, distinctly rugose to rugulose and punctulate elytra, typically entirely metallic color of elytra, pronotum and appendages. Some exception exist in which species are not metallic (e.g. D fennica), or with base of femora reddish (e.g. D. bicoloricornis and several others). Species of D. (Donacocia) are never entirely pubescent dorsally (D. pubescens of North America may not belong in this subgenus; Askevold in prep.) The main morphological characteristics of Donacia larvae include: a slightly convex body with a very small, sclerotized, prognathous head; distinct frons, clypeus, and labrum; mandible bearing two prominent teeth; short, slightly curved, and awl-shaped legs lacking pulvilli; five stemmata on each side of the head; a thorax that is unsclerotized and covered with numerous setae; abdominal segments lacking tubercles; and a large, sclerotized spiracle hook on the 8th abdominal segment (Lee 1991). In the present study, detailed descriptions, illustrations, and taxonomic keys were developed for both adult and larval stages of the new species. Furthermore, ecological data, distribution maps, host-plant associations, oviposition behaviors, and feeding behaviors of the genus Donacia were synthesized, and cytochrome c oxidase subunit I (COI) gene sequences were analyzed. Materials and methods Type specimens of the new species described have been deposited in the National Institute of Biological Resources, South Korea (NIBR) and J.Y. Kim’s private collection, South Korea (JYK). Biological observations were conducted from May to June of 2022–2023 at the type locality and under laboratory conditions. Adults were collected by sweeping and handpicking, whereas larvae were obtained from roots of host plants. Captured adults were placed in Ziplock bags (20 cm wide, 15 cm long, and 8 cm deep) with host plants and monitored daily, whereas larvae were preserved in 70% ethyl alcohol in the field. For morphological analysis of minute structures, some larvae were dissected, cleared in 10% KOH solution for 5 h, and rinsed in distilled water. Microscopic tissues were mounted on slides, and specimens were prepared using Neo-Shigaral solution (Shiga Konchu Fukyusha, Tokyo, Japan). To examine genitalia, specimens were boiled in hot water for approximately 3 min to soften tissues. Genitalia were then carefully dissected using entomological needles and subsequently immersed in 10% KOH solution for 1–2 h to facilitate clearing. Descriptions and illustrations were prepared using a stereoscopic microscope (LEICA M125, Olympus SZX-12) and an optical microscope (Olympus BX50) equipped with a drawing tube and Adobe Photoshop CS6 software (Adobe Systems, Inc., San Jose, CA, USA) using tablets (WACOM Intuos CLT-4100). Habitus images were captured using a a Canon EOS 5D Mark 5D IV digital camera attached to a 85 ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034 Joong Youb Kim et al.: Donacia koreana sp. nov. from South Korea Canon EF 100 mm F2.8 Macro USM. Morphological terminology for adults largely followed Gressitt and Kimoto (1961), Kimoto and Gressitt (1979), Mann and Crowson (1983), Askevold (1987, 1988, 1990, 1991), and Hayashi (2020), whereas terminology for larvae was based primarily on Anderson (1947) and LeSage (2012). Genomic DNA was extracted from the legs of two specimens using the LaboPas Tissue Genomic DNA Isolation Kit Mini (Cosmo Genetech Inc., Seoul, Korea) according to the manufacturer’s instructions. The COI barcode fragment was amplified using the universal primer set (LepF1 and HCO2198) under the following cycling conditions: initial denaturation at 94 °C for 2 min; followed by 35 cycles of 98 °C for 15 s, 53 °C for 30 s, and 68 °C for 60 s; and a final extension at 68 °C for 5 min. Amplified products were then sequenced using an ABI 3100 automated sequencer (Perkin-Elmer, CA, USA). Sequence assembly, alignment, and trimming were performed using the Geneious 8.1.9 software (Kearse et al. 2012). COI sequences amplified from the newly identified Donacia species were aligned with those of 19 other Donacia species retrieved from the National Center for Biotechnology Information database, using Geneious v. 10.2.6 (Table 1). Pairwise genetic distances were calculated using the Kimura 2-parameter substitution model in MEGA X v. 12.0.11 (Kumar et al. 2024). Phylogenetic relationships were inferred using maximum likelihood analysis implemented in PhyML v. 3.1, based on the GTR + I + G model selected using jModelTest v. 2.1.10, with 1000 bootstrap replicates. The COI sequence of Plateumaris braccata (Scopoli) was used as an outgroup for this analysis. Results Donacia koreana Kim & Lee sp. nov. https://zoobank.org/C52E8C52-E941-4A82-B007-2242FC6BB524 Figs 1–4 Type locality. South Korea, Kangwon State, Hyeoncheon-ri, Dunnae-myeon, Hoengseong-gun, swamp, 37°30'9.37"N, 128°9'53.57"E. Type materials. Holotype (NIBR no. NIBRIN0001069714): male, South Korea • Kangwon State, Hyeoncheon-ri, Dunnae-myeon, Hoengseong-gun, swamp, 37°30'9.37"N, 128°9'53.57"E, 520 m, 29.V.2022, J.Y. Kim. Paratypes (NIBR no. NIBRIN0001069715): • five males and seven females, same data as the holotype, except for 6.VI.2023, J.Y. Kim. Other materials. Twenty larvae were collected from the roots of Scirpus wichurae Boeckeler with the same data as for the paratype, except for 6.VI.2023. Diagnosis, adult (Figs 1, 2, 4). The new species closely resembles Donacia bicolora Zschach, but can be distinguished by the following diagnostic characteristics: the rows of punctures are still clearly visible even in the last third of the elytra (D. bicolora: these punctures are almost completely obscured by the silky, shiny, fine wrinkles in the last third of the elytra).; and it generally occurs in two distinct color morphs: cupreous and greenish (D. bicolora exhibits a variety of body colorations, commonly greenish, bluish, or blackish). The legs are entirely metallic. Description, adult (Figs 1, 2, 4). Male. Length 7.9–8.1 mm, width 2.7–2.8 mm. Body narrowly elongate; generally cupreous or greenish; head cupreous or greenish; antennomeres 1–4 coppery or greenish, 5 partially darkened, 6–11 dark; pronotum cupreous or greenish; legs cupreous or greenish. 86 ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034 Joong Youb Kim et al.: Donacia koreana sp. nov. from South Korea Head. Narrower than prothorax; eyes small and convex; supraocular furrow present but shallow; vertex slightly swollen, pubescent with a fine median line. Antenna filiform and pubescent; antennomere 5 longest amongst 2nd to 6th; antennomere 4 slightly longer than 3rd; antennomere 5 2.5 times longer than 2nd. Pronotum. Generally quadrate, width as long as length, posterior and anterior corners prominent; anterolateral calli present, callosal sulcus present but shallow; median line deep; disc coarsely punctate and transverse deep rugae; basal sulcus present but shallow, coarsely punctate. Scutellum. Subtriangular, closely covered with silvery pubescent. Elytron. Subparallel-sided, gradually narrowed apically; surface distinctly 10 longitudinal regular striate, striae regular and deeply punctate; with a shallow depression at the base near the suture (1st–4th sutural intervals), and a deep depression at the base of the median area (5th–8th sutural intervals); apex truncate, outer apical angle obtusely rounded, inner apical angle obtuse. Legs. Slender; metafemur robust in shape, tapering and not distinctly clavate, with a prominent large tooth. Pygidium. Trapezoidal and apex pubescent, emarginate in both sexes. Table 1. List of Donacia species used in molecular analysis and the respective references. Species Collection locality GenBank accession No. Reference Donacia koreana sp. nov. Korea This study Donacia aquatica Germany KU912172 Rulik and Ahrens 2017 Finland MZ659808 Roslin et al. 2022 Donacia aureocincta Finland MZ658052 Roslin et al. 2022 Donacia bicolora Finland MZ656234, MZ658244 Roslin et al. 2022 Donacia cincticornis Canada KM844344, HM411995 – Donacia cinerea Germany KU917042 Rulik et al. 2017 Finland MZ658078 Roslin et al. 2022 Donacia flemola China MK049870 Nie et al. 2020 Donacia hypoleuca Canada JF887697 – Donacia marginata Germany KU919476 Rulik et al. 2017 Finland KJ964703 Pentinsaari et al. 2014 Donacia obscura Italy MH323099 Magoga et al. 2018 Finland KJ962943 Pentinsaari et al. 2014 Donacia piscatrix Canada KM844563 – Donacia semicuprea Germany KM443995 Rulik et al. 2017 Sweden KJ965000 Pentinsaari et al. 2014 Donacia simplex Germany KU913219, KU906925 Rulik et al. 2017 Donacia sparganii Germany KU910558 Rulik et al. 2017 Donacia subtilis Canada KR483642, KR486904 Hebert et al. 2016 Donacia thalassina Germany KU912621 Rulik et al. 2017 Finland KJ967484 Pentinsaari et al. 2014 Donacia tomentosa France KM451348 Hendrich et al. 2015 Finland MZ631647 Roslin et al. 2022 Donacia versicolorea Germany KU917168, KU919222 Rulik et al. 2017 Donacia vulgaris Germany KU915372 Rulik et al. 2017 Finland MZ631910 Roslin et al. 2022 Donacia provostii Korea OL663187 – Plateumaris braccata Finland MZ633983 Roslin et al. 2022 87 ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034 Joong Youb Kim et al.: Donacia koreana sp. nov. from South Korea Male genitalia. Median lob of genitalia narrowed apically with median lip and both sides emarginate subapically; a cap of tegmen slender, apex with shallow depression; endophallus elongate, median process short and curving dorsal, BSB elongate. Sterna. Entirely greenish and pubescent; apical shape of last sternite truncate. Female. Length 9.0–9.3 mm, width 2.8–3.1 mm. Body larger than that of males; pronotum frontal outline quadrate, width longer than length; apical shape of last sternite pointed. Spermatheca. Short and broad, strongly C-shaped; apex tapered; collum and ramus distinct; cornu and nodulus fused into a globose basal swelling; horns widely separated. Description, first instar larva. Length 1.66–1.78 mm, width 0.40–0.41 mm, spiracle-hook length (0.213 mm), and spiracle-hook width (0.033 mm (n = 4). Creamy white, body slightly convex, thorax and abdomen with numerous setae, but not as setae as the last instar larva; egg buster absent. Diagnosis, last instar larva (Figs 3, 4C). The last instar larva is easily distinguished from the other Donacia larvae by the following characteristics: epicranium with six pairs of dorsal setae; two pairs of sensilla; labrum with three pairs of labral setae and one pair of sensilla; and prementum with one pair of setae and two pairs of sensilla. Description, last instar larva (Figs 3, 4C). Length 10.5–12.3 mm, width 3.7–4.3 mm, head length 0.78–0.84 mm, spiracle hook length 0.73–0.77 mm, spiracle hook width 0.42–0.44 mm (n = 5). Body yellowish, slightly convex, and C-shaped. Head, prothorax, and legs pale brown; setae and spiracles brown. Head. Prognathous, small, rounded, well sclerotized, strongly retractile into prothorax; hind corner of epicranium slightly posteriorly produced. Epicranial suture Y-shaped; frontal suture nearly straight. Stemmata pigmented, five in number. Epicranium with six pairs of dorsal setae, two pairs of sensilla, and three pairs of lateral setae. Frons with four pairs of frontal setae and one pair of sensilla. Antenna three-segmented: segment 1 with two sensilla, segment Figure 1. Donacia koreana sp. nov. A, B. Male (A. Holotype); C, D. Female (C. Paratype). Scale bars: 1.0 mm (A–D). 88 ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034 Joong Youb Kim et al.: Donacia koreana sp. nov. from South Korea 2 with conical sensory papilla and two sensilla basiconica, and segment 3 with two sensilla basiconica and one seta. Endocarina absent; epistomal sutures distinct. Clypeus with two pairs of clypeal setae and one pair of sensilla. Labrum slightly notched anterior margin with three pairs of labral setae and one pair of sensilla. Epipharynx with two pairs of spiniform setae. Mandible conical, strongly sclerotized, with two teeth, 2nd serrated, and one mandibular seta. Maxillar palp three-segmented; palpifer with two setae; stipes, Figure 2. Adult: Donacia koreana sp. nov. A. Leg (l.v.); B, C. Pygidium (d.v.); D, E. Abdomen (d.v.); F. Antenna (d.v.); G. Median lobe (l.v.); H. Apex of median lobe (d.v.); I. A cap of tegmen; J, K. Endophallus (d.v. and v.v.); L. Spermatheca. male: A, B, D, F–K; female: C, E, L. Scale bar: 1.0 mm (A, D, E, F); 0.5 mm (B, C, G, H); 0.2 mm (I–L). 89 ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034 Joong Youb Kim et al.: Donacia koreana sp. nov. from South Korea two setae and one sensillum; galea fused with lacinia, spear-like; cardo large, trapeziform, with one seta. Labial palp one-segmented with one sensillum; ligula with numerous setae; prementum with one pair of setae and two pairs of sensilla; postmentum with two pairs of setae and one pair of sensilla. Thorax. Pronotum pale brown, slightly sclerotized, with numerous setae. Meso-and metanotas not sclerotized. Thoracic spiracles annuliform, situated on the EPa; spiracular opening rounded. Legs short and stout; femur with ten setae and four sensilla; tibio-tarsus with three setae; claws slightly curved, awlshaped, based enlarged, with one seta; pulvillus absent. Figure 3. Larva: Donacia koreana sp. nov. A. Last instar larva (l.v.); B. Antenna (d.v.); C. Head (d.v.); D. Clypeus, labrum and epipharynx (d.v. and v.v.); E. Mandible (b.v.); F. Spiracle (d.v.); G. Maxilla and labium; H. Leg (v.v.); I. 8th abdominal spiracle. Scale bar: 1.0 mm (A); 0.5 mm (C, H, G, I); 0.1 mm (B, E, F). 90 ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034 Joong Youb Kim et al.: Donacia koreana sp. nov. from South Korea Abdomen. Nine segmented, unsclerotized. Typical abdominal segments with two folds. Abdominal spiracles present on segments 1–8 similar to mesothoracic spiracles except for 8th spiracle. Spiracle hook length about 2.3 times as long as width; projected, large, peg-like, strongly sclerotized. Distribution. South Korea. Host plants. Cyperaceae: Scirpus wichurae Boeckeler, Eleocharis ussuriensis G. Zinserl. Key to the adults of South Korean Donacia species 1 Entire dorsum of the beetle not really metallic at all; appendages predominantly bicolored to rufescent; pronotal disc with at most fine punctures and rugulae; elytra with at most few fine rugulae and minute punctulae in intervals ... 2 – Entire dorsum of beetle metallic; appendages predominantly dark; pronotal disc coarsely punctured and often transversely coarsely rugose; elytral disc generally coarsely rugose between striae ...........................................3 2 Third antennal segment slightly longer than, or as long as second ............ .......................................................................................... D. lenzi Schönfeldt – Third antennal segment much longer than second .......D. provostii Fairmaire 3 Femora and tibia entirely metallic-colored ..................................................4 – Femora and tibia partly metallic-colored (femoral base usually rufous) ... 5 Figure 4. Donacia koreana sp. nov. A, B. Adults; C. Larva; D. Pupal case; (C, D red circle). 91 ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034 Joong Youb Kim et al.: Donacia koreana sp. nov. from South Korea 4 Elytron with red longitudinal stripe ........................... D. aquatica (Linnaeus) – Elytron without red longitudinal stripe .........................................................6 5 Pronotal disc with distinct fine tum with short hairs (most specimens) ..... ...................................................................................D. clavareaui Jacobson – Pronotal disc without distinct short hairs .................D. bicoloricornis Chen 6 Body and legs entirely black, unicolored ......................... D. flemola Goecke – Body and legs entirely cupreous or green ......................D. koreana sp. nov. Key to the larvae of South Korean Donacia species 1 Head inside the prothorax .............................................................................2 – Head outside the prothorax ............................................. D. flemola Goecke 2 Hind corners of epicranium moderately produced .....................................3 – Hind corners of epicranium slightly produced ............................................4 3 Ephipharynx with three pairs of spiniform setae ....D. clavareaui Jacobson – Epipharynx with two pairs of spiniform setae ............... D. koreana sp. nov. 4 Head ovoid .....................................................................................................5 – Head rounded ................................................................................................6 5 Postmentum with two pairs of setae ............................. D. lenzi Schönfeldt – Postmentum with three pairs of setae..................... D. aquatica (Linnaeus) 6 Clypeus with two pairs of clypeal setae....................D. bicoloricornis Chen – Clypeus with three pairs of clypeal setae ..................D. provostii Fairmaire Biological note Feeding pattern To investigate the feeding behavior of the Korean Donaciinae, the classification system proposed by Bieńkowski (2015) was adopted. According to this framework, three distinct feeding patterns were identified. • Type I: where the insect inclines its head, turns it to the right or left, and inserts both mandibles into the leaf tissues. • Type II: characterized by the insertion of one mandible into the leaf, whereas the other remains on the leaf surface. • Type III: the head movement of the insect resembles that of Type II, but both mandibles capture the upper and lower surfaces of the leaf. Based on the observations following this classification, D. lenzi and D. provostii exhibited Type I feeding behavior. Donacia aquatica, D. bicoloricornis, and D. clavareaui displayed Type II feeding behavior, whereas D. flemola and D. koreana sp. nov. exhibited Type III behavior (Fig. 5). Notably, although D. koreana sp. nov. engaged in leaf feeding, its primary feeding sites were the flowers of Scirpus wichurae and Eleocharis ussuriensis rather than leaf tissues. Oviposition patterns Adult Donaciinae exhibit diverse oviposition behaviors that vary depending on the species and associated host plant (Bieńkowski 1996). In the present study, 98 ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034 Joong Youb Kim et al.: Donacia koreana sp. nov. from South Korea Fossil Insect Research Group for Nojiri-ko Excavation (1985) Atlas of the Japanese Donaciinae (Guide for identification of the fossil donaciinae beetles). Fossil Insect Research Group for Nojiri-ko Excavation, Osaka, 182 pp. Geiser E (2023) Revision of the Palaearctic species of the genus Plateumaris CG Thomson, 1859 (Coleoptera, Chrysomelidae, Donaciinae). ZooKeys 1177: 167–233. https:// doi.org/10.3897/zookeys.1177.103214 Gressitt JL, Kimoto S (1961) The Chrysomelidae (Coleopt.) of China and Korea, Part 1. Pacific Insects Monograph 1A: 1–299. Hayashi M (2020) Revision of extant and fossil Donaciinae (Coleoptera: Chrysomelidae) of Japan. Japanese Journal of Systematic Entomology, Supplementary Series (2): 61–160. Hebert PDN, Ratnasingham S, Zakharov EV, Telfer AC, Beaudin VL, Milton MA, Pedersen S, Jannetta P, deWaard JR (2016) Counting animal species with DNA barcodes: Canadian insects. Philosophical Transactions of the Royal Society B: Biological Sciences 371(1702): 20150333. https://doi.org/10.1098/rstb.2015.0333 Hendrich L, Morinière J, Haszprunar G, Hebert PD, Hausmann A, Köhler F, Balke M (2015) A comprehensive DNA barcode database for Central European beetles with a focus on Germany: Adding more than 3500 identified species to BOLD. Molecular Ecology Resources 15(4): 795–818. https://doi.org/10.1111/1755-0998.12354 Jeong YJ, Kim JY, Park JH, Cho GH, Lee JE, Hong KJ (2024) Revealing the Identity of Donacia aquatica (Linnaeus) and D. japana Chûjô & Goecke (Coleoptera: Chrysomelidae: Donaciinae) based on Literature Review, New Material, and DNA Barcodes. Korean Journal of Applied Entomology 63(3): 189–195. https://doi.org/10.5656/ KSAE.2024.07.0.018 Kearse M, Moir R, Wilson A, Havas SS, Cheung M, Sturrock S, Buxton S, Cooper A, Markowitz S, Duran C, Thierer T, Ashton B, Meintjes P, Drummond A (2012) Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics (Oxford, England) 28(12): 1647–1649. https://doi.org/10.1093/bioinformatics/bts199 Kimoto S, Gressitt JL (1979) Chrysomelidae (Coleoptera) of Thailand, Cambodia, Laos and Vietnam I. Pacific Insects 20(2–3): 191–256. Kippenberg H (2015) Strukturierung von artenreichen Chrysomelidae-Gattungen, am Beispiel der Gattung Donacia Fabricius (Coleoptera: Chrysomelidae) Koleopterologische Rundschau 85: 249–282. KSAE [Korean Society of Applied Entomology], ESK [The Entomological Society of Korea] (2021) Check List of Insects from Korea. Paper and Pencil, Daegu, 1055 pp. Kumar S, Stecher G, Suleski M, Sanderford M, Sharma S, Tamura K (2024) MEGA12: Molecular Evolutionary Genetic Analysis version 12 for adaptive and green computing. Molecular Biology and Evolution 41(12): msae263. https://doi.org/10.1093/molbev/msae263 Lee JE (1991) A taxonomic study on the larvae of the subfamily Donaciinae from Japan (Coleoptera: Chrysomelidae). Entomological Research Bulletin Korea 17: 33–46. Lee JE, An SL (2001) Family Chrysomelidae. Economic Insects of Korea 14. Insecta Koreana (Supplement 21): 1–231. https://kiss.kstudy.com/Detail/Ar?key=277086 LeSage L (2012) The eggs and larvae of Pachybrachis peccans and P. bivittatus, with a key to the known immature stages of the Nearctic genera of Cryptocephalinae (Coleoptera: Chrysomelidae). Canadian Entomologist 117(2): 203–220. https://doi. org/10.4039/Ent117203-2 99 ZooKeys 1261: 83–99 (2025), DOI: 10.3897/zookeys.1261.169034 Joong Youb Kim et al.: Donacia koreana sp. nov. from South Korea Magoga G, Sahin DC, Fontaneto D, Montagna M (2018) Barcoding of Chrysomelidae of Euro-Mediterranean area: Efficiency and problematic species. Scientific Reports 8(1): 13398. https://doi.org/10.1038/s41598-018-31545-9 Mann JS, Crowson RA (1983) On the internal anatomy and classification of Donaciinae (Col., Chrysomelidae). Entomologist’s Monthly Magazine 119: 17–27. Nie RE, Andújar C, Rodríguez CG, Bai M, Xue HJ, Tang M, Yang CT, Tang P, Yang XK, Vogler AP (2020) The phylogeny of leaf beetles (Chrysomelidae) inferred from mitochondrial genomes. Systematic Entomology 45(1): 188–204. https://doi.org/10.1111/syen.12387 Pentinsaari M, Hebert PD, Mutanen M (2014) Barcoding beetles: A regional survey of 1872 species reveals high identification success and unusually deep interspecific divergences. PLOS ONE 9(9): e108651. https://doi.org/10.1371/journal.pone.0108651 Roslin T, Somervuo P, Pentinsaari M, Hebert PDN, Agda J, Ahlroth P, Anttonen P, Aspi J, Blagoev G, Blanco S, Chan D, Clayhills T, deWaard J, deWaard S, Elliot T, Elo R, Haapala S, Helve E, Ilmonen J, Hirvonen P, Ho C, Itämies J, Ivanov V, Jakovlev J, Juslén A, Jussila R, Kahanpää J, Kaila L, Kaitila J-P, Kakko A, Kakko I, Karhu A, Karjalainen S, Kjaerandsen J, Koskinen J, Laasonen EM, Laasonen L, Laine E, Lampila P, LevesqueBeaudin V, Lu L, Lähteenaro M, Majuri P, Malmberg S, Manjunath R, Martikainen P, Mattila J, McKeown J, Metsälä P, Miklasevskaja M, Miller M, Miskie R, Muinonen A, Mukkala V-M, Naik S, Nikolova N, Nupponen K, Ovaskainen O, Österblad I, Paasivirta L, Pajunen T, Parkko P, Paukkunen J, Penttinen R, Perez K, Pohjoismäki J, Prosser S, Raekunnas M, Rahulan M, Rannisto M, Ratnasingham S, Raukko P, Rinne A, Rintala T, Romo SM, Salmela J, Salokannel J, Savolainen R, Schulman L, Sihvonen P, Soliman D, Sones J, Steinke C, Ståhls G, Tabell J, Tiusanen M, Várkonyi G, Vesterinen EJ, Viitanen E, Vikberg V, Viitasaari M, Vilen J, Warne C, Wei C, Winqvist K, Zakharov E, Mutanen M (2022) A molecular‐based identification resource for the arthropods of Finland. Molecular Ecology Resources 22(2): 803–822. https://doi.org/10.1111/1755-0998.13510 Rulik B, Ahrens D (2017) Using taxonomic consistency with semi-automatized data pre-processing for high quality Barcode data submissions: a case study on German beetles. https://doi.org/10.1111/2041-210X.12824 van der Valk A (2012) The Biology of Freshwater Wetlands. Oxford University Press, Oxford, 280 pp. https://doi.org/10.1093/acprof:oso/9780199608942.001.0001