First record of the Heterotoma planicornis (Hemiptera, Heteroptera, Miridae), an introduced species in the Korean Peninsula, with characterization of its complete mitochondrial genome

First record of the Heterotoma planicornis (Hemiptera, Heteroptera, Miridae), an introduced species in the Korean Peninsula, with characterization of its complete mitochondrial genome Minsuk Oh1,2, Soojeong Cho3, Seunghwan Lee1,2 1 Laboratory of Insect Biosystematics, Department of Agricultural Biotechnology, Seoul National University, Seoul, Republic of Korea 2 Research Institute of Agriculture and Life Sciences, Seoul National University, Republic of Korea 3 Beolbolil-itneun-saramdeul citizen science community, Idang-ro, Gunpo-si, Gyeonggi-do, Republic of Korea https://zoobank.org/0DC0D61B-F9C7-4C52-B1C1-FF46B94DEC7D Corresponding author: Seunghwan Lee ([email protected]) Academic editor: Dávid Rédei ♦ Received 14 March 2025 ♦ Accepted 9 September 2025 ♦ Published 14 October 2025 Abstract The West Palearctic genus Heterotoma Lepeletier & Serville, 1825, and its included species H. planicornis (Pallas, 1772) (Hemiptera, Heteroptera, Miridae, Orthotylinae) are reported from the Korean Peninsula. Heterotoma planicornis is diagnosed, and its dorsal habitus and the genitalia of both sexes are illustrated. The possible introduction of this European species into the Korean Peninsula is discussed. The complete mitochondrial genome of the species is provided. The circular mitogenome of H. planicornis is 16,179 bp long, comprising 13 protein-coding genes, two ribosomal RNA genes, and 22 transfer RNAs, and shows the same gene order as the closely allied species Mecomma ambulans (Fallén, 1807). The A–T content of the total sequence was 71.54%. An analysis using a cytochrome oxidase I (COI) neighbor-joining (NJ) tree revealed that the Korean H. planicornis population formed two distinctive groups, with a 1.8% sequence difference between them. Key Words species introduction, distribution, East Asia, mitogenome, phylogeny, plant bugs Introduction Orthotylinae is a subfamily of Miridae (Hemiptera: Heteroptera), comprising approximately six tribes, 260 genera, and over 2,100 described species (Schuh 2002–2013; Schuh and Weirauch 2020; Oh et al. 2023). A morphology-based analysis recovered this subfamily as a close relative of Phylinae (Schuh 1976), and this was further supported by subsequent molecular-based phylogenetic analyses (Menard et al. 2014; Oh et al. 2023). Orthotyline species are predominantly predators, and some of them are natural enemies of pests of various plants (Wheeler 2001; Oh et al. 2023). Heterotoma species are zoophytophagous and recognized as natural enemies of various pest arthropods; H. planicornis has been reported to prey on mites, aphids, psyllids, gall midges, eggs and larvae of Chrysomelidae, as well as eggs of heteropteran and lepidopteran insects (for review, see Kment and Bryja 2006). Currently, four Heterotoma species are known, all native to the Western Palearctic (Wheeler and Henry 1992; Kerzhner and Josifov 1999; Schuh 2002–2013). Among them, H. planicornis exhibits the widest distribution; in addition to its native Western Palearctic area, it has been introduced to North America (Canada, USA) (Knight 1917; Wheeler and Henry 1992). An observation from the Republic of South Africa (Western Cape) has been reported by iNaturalist (2025), but the correct identification of this record requires verification. In this paper, we present a new distribution record of Heterotoma planicornis from the Korean Peninsula. A morphological diagnosis is presented, and the habitus of the species and genitalia of both sexes are illustrated. The Dtsch. Entomol. Z. 72 (2) 2025, 225–235|DOI 10.3897/dez.72.152970 Copyright Minsuk Oh et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. dez.pensoft.net Minsuk Oh et al.: New record and mitogenome of Heterotoma 226 complete mitogenome sequence of H. planicornis is provided and compared to those of related taxa by a neighbor-joining (NJ) tree based on the cytochrome oxidase I (COI) region. Materials and methods Specimen examination and figure acquisition All voucher specimens are deposited in the collection of the Insect Biosystematics Laboratory, Research Institute for Agriculture and Life Science, Seoul National University, Korea (SNU). Digital images of external characteristics were taken with a DMC 5400 digital camera attached to a Leica Z16 APO motorized microscope. Genital structures were dissected and observed under a Leica DM 4000B microscope, and images were taken using a digital camera combined with the microscope (Lumenera Infinity 3). All measurements (mean and range) are provided in millimeters. Terminology for male and female genital structures follows Oh and Lee (2018). Mitogenome data construction Genomic DNA was extracted from the abdominal segments IV–VI and legs of specimens preserved in 95% to 99% alcohol using a DNeasy Blood and Tissue Kit (QIAGEN, Inc.) following the manufacturer’s protocols. Examined specimens were labeled and stored in alcohol depending on their condition. After DNA extraction, a sequencing library was prepared using the TruSeq Nano DNA Kit, based on the TruSeq Nano DNA Sample Preparation Guide, Part #15041110 Rev. D (Illumina, Inc.). NovaSeq X Plus was used as the sequencing platform. Read length was 151 bp (30 million reads per lane) according to the Macrogen Company (South Korea) sequencing facility. Sequences were aligned using Geneious Prime 2025.1.3 software (Kearse et al. 2012). A total of six reference sequences were used for alignment (Table 1), and the resulting data were annotated using MITOS2 (Bernt et al. 2013) and visualized as a circular map using the webserver Proksee (https://proksee.ca/, accessed on 26 July 2024) (Grant et al. 2023). Phylogenetic analysis A COI-based dataset, comprising 10 sequences from the NCBI database and four from the present study, was compiled. To evaluate interand intraspecific genetic distances, 563 bp of the mitochondrial protein-coding COI (cytochrome oxidase subunit I) gene was used (Folmer et al. 1994), with LCO1490 (5′-GGTCAACAAATCATAAAGATATTGG-3′) and HCO2198 (5′-TAAACTTCAGGGTGACCAAAAAATCA-3′) primers. During alignment, an L-INS-i strategy was used, without any manipulation except trimming the ends of reads. A neighbor-joining (NJ) tree (Saitou and Nei 1987) was generated, and sequence divergences were evaluated using the Kimura 2-parameter model (K2P) (Kimura 1980) in MEGA 7.0 (Kumar et al. 2016). Detailed information for the sequences used is provided in Table 2. Table 1. Sequences used for the annotation of the mitochondrial genome of Heterotoma planicornis. Subfamily Tribe Species Accession No. Locality bp Reference Mirinae Hyalopeplini Onomaus tenuis Zheng, 2004 MW619699.1 Unknown 14860 Ye et al. (2022) Mirinae Mecistoscelini Mystilus priamus Distant, 1904 MW619697.1 Unknown 14687 Ye et al. (2022) Mirinae Mirini Adelphocoris suturalis (Jakovlev, 1882) KJ020288.1 China 14327 Wang et al. (2014) Mirinae Stenodemini Trigonotylus caelestialium (Kirkaldy, 1902) KJ170899.1 China 15095 Wang et al. (2014) Orthotylinae Orthotylini Mecomma ambulans (Fallén, 1807) MW619692.1 Unknown 12613 Ye et al. (2022) Phylinae Pilophorini Pilophorus typicus (Distant, 1909) MW619694.1 Unknown 15335 Ye et al. (2022) Table 2. COI sequences used for phylogenetic analysis in this study. Subfamily Tribe Species Accession No. Locality bp Reference Orthotylinae Halticini Ectmetopterus comitans (Josifov & Kerzhner, 1972) GU194790.1 Unknown 527 Jung and Lee (2012) Orthotylinae Halticini Orthocephalus funestus Jakovlev, 1881 GU194802.1 Unknown 527 Jung and Lee (2012) Orthotylinae Halticini Strongylocoris leucocephalus (Linnaeus, 1758) KM022178.1 Germany 658 Raupach et al. (2014) Orthotylinae Orthotylini Blepharidopterus diaphanus (Kirschbaum, 1856) MZ407988 Korea 563 Oh et al. (2023) Orthotylinae Orthotylini Dryophilocoris miyamotoi Yasunaga, 1999 GU194785.1 Unknown 527 Jung and Lee (2012) Orthotylinae Orthotylini Heterotoma planicornis (Pallas, 1772) HQ105735.1 Canada 638 Park et al. (2011) Orthotylinae Orthotylini Heterotoma planicornis (Pallas, 1772) HQ105736.1 Canada 638 Park et al. (2011) Orthotylinae Orthotylini Heterotoma planicornis (Pallas, 1772) PV879916 Korea 16179 This work Orthotylinae Orthotylini Heterotoma planicornis (Pallas, 1772) PV789749 Korea 563 This work Orthotylinae Orthotylini Heterotoma planicornis (Pallas, 1772) PV850443 Korea 563 This work Orthotylinae Orthotylini Heterotoma planicornis (Pallas, 1772) PV850444 Korea 563 This work Orthotylinae Orthotylini Orthotylus pallens (Matsumura, 1911) GU194805.1 Unknown 527 Jung and Lee (2012) Orthotylinae Orthotylini Ulmica yasunagai Oh & Lee, 2018 MZ408099 Korea 563 Oh et al. (2023) Orthotylinae Orthotylini Zanchius tarasovi Kerzhner, 1988 MZ408103 Korea 563 Oh et al. (2023) Dtsch. Entomol. Z. 72 (2) 2025, 225–235 dez.pensoft.net 227 Taxonomy Genus Heterotoma Lepeletier & Serville, 1825 Pachycera Billberg, 1820: 68 (original description). Type species: Cimex spissicornis Fabricius, 1777 (= Cimex planicornis Pallas, 1772), by monotypy. Nomen oblitum. Synonymized by Kerzhner (1997: 245). Heterotoma Lepeletier & Serville, 1825: 326 (original description). Type species: Cimex planicornis Pallas, 1772, by monotypy. Acroderrhis Bergroth, 1914: 179 (original description). Type species: Heterotoma dentipennis Bergroth, 1914, by monotypy. Synonymized by Wagner (1968: 182). Heterotoma: Schuh (1995: 120) (catalog); Kerzhner and Josifov (1999: 248) (catalog); Schuh (2002–2013) (online catalog); Kment and Bryja (2006: 11) (diagnosis); Aukema (2018) (online catalog). Heterotoma planicornis (Pallas, 1772) Figs 1–6 Cimex planicornis Pallas, 1772: 23. (original description). Cimex spissicornis Fabricius, 1777: 300 (original description). Synonymized by Costa (1843: 55). Acanthia crassicornis Fabricius, 1794: 70 (original description). Synonymized by Reuter (1888: 667). Cimex crassipennis Turton, 1802: 609 (new name for Cimex crassicornis). Synonymized by Reuter (1888: 667). Heterotoma acinaciformis Costa, 1839: 20 (original description). Synonymized by Wagner (1968: 185), tentatively. Heterotoma planicornis: Tamanini (1962: 136) (diagnosis, description); Schuh (1995: 121) (catalog); Kerzhner and Josifov (1999: 248) (catalog); Schuh (2002–2013) (online catalog); Kment and Bryja (2006: 11) (redescription, figures, neotype designation, distribution, biolFigure 1. Heterotoma planicornis, live individuals. A, B, D. Male; C. Female; E. Last instar nymph; F. Collection site, with host plant Spiraea japonica ‘Goldflame’. dez.pensoft.net Minsuk Oh et al.: New record and mitogenome of Heterotoma 228 Figure 2. Dorsal and ventral habitus of Korean Heterotoma planicornis. A–C. Male; D, E. Female; F. Last instar nymph. Scale bar: 3.0 mm. ogy); Kment and Bryja (2012: 117) (correction); Aukema (2018) (online catalog). Diagnosis. This species can be distinguished by the combination of the following characters: dorsum covered with brown, suberect setae and pale, reclining sericeous setae (Figs 1A–E, 2A, D); antennal segment blackish brown, segments I and II densely covered with fuscous, suberect setae; segment II thick and elongated, somewhat flattened (Figs 1A–E, 2A–F); segments Dtsch. Entomol. Z. 72 (2) 2025, 225–235 dez.pensoft.net 229 Figure 3. Pygophore of Heterotoma planicornis. A. Dorsal view; B. Ventral view; C. Pygophore opening. Scale bar: 0.5 mm. III and IV short, thin, and linear, basally pale; hemelytra entirely dark brown; labium reaching mesocoxae; femora pale green, without dark or reddish tinge; tibiae yellowish green, subbasally tinged with green; tarsus yellowish green, widely darkened at segments I and III; pygophore with one elongated, pointed-end structure and short, bifurcate structure (Fig. 3A–C); male endosoma with two elongated, apically spinulate sclerites and thick and smooth, pointed-end basal sclerite (Fig. 4A–D); sensory lobe of left paramere with thick structure covered with stiff setae, hypophysis with minute, hook-like structure (Fig. 4E–G); sensory lobe of right paramere moderately curved and laterally with two dentation, hypophysis stout, apically blunt (Fig. 4H–J); female genitalia with thick sclerotized ring (Fig. 5A–C); interramal lobe spinulate, bifurcate proximally (Fig. 5D, E). For more diagnostic characters and figures, see Tamanini (1962, 1981) and Kment and Bryja (2006, 2012). Measurements. Male (n = 5) / Female (n = 2). Total body length 4.45–5.00 / 4.75–4.85; head width across eyes 0.79–0.84 / 0.83–0.85; vertex width 0.33–0.36 / 0.35–0.39; lengths of antennal segments I–IV 0.53– 0.59, 1.66–1.79, 0.49–0.53, 0.42–0.47 / 0.54–0.55, 1.64–1.70, 0.49–0.50, 0.44; labial length 1.57–1.63 / 1.53–1.65; mesal pronotal length including collar 0.63–0.71 / 0.61–0.62; basal pronotal width 1.01–1.12 / 1.04–1.10; width across hemelytron 1.28–1.39 / 1.39– 1.63; cuneal length 0.69–0.88 / 0.76–0.78; cuneal width 0.33–0.39 / 0.33–0.35; lengths of metafemur, tibia, and tarsus 1.49–1.66, 2.05–2.40, 0.47–0.55 / 1.56–1.65, 2.13–2.22, 0.53–0.54. Distribution. Europe: most of the territory. Asia: Türkiye, South Korea (introduced) (new record). North America: Canada, USA (Wheeler and Henry 1992; Kerzhner and Josifov 1999; Kment and Bryja 2006) (introduced). Biology. This species exhibits a wide host range, feeding on plants from 12 angiosperm families (Kment and Bryja 2006). Our observations revealed that the Korean population inhabits Spiraea japonica (Rosaceae) (Fig. 1F). The habitat of H. planicornis discovered in this study was limited to narrow artificial landscapes in urbanized areas, excluding larger available natural habitats such as parks. Material examined. South Korea, Seoul: 5♂, 1♀, near Hannam station, Dokseodang-ro 6, Yongsan-gu, on Spiraea japonica 3.vi.2024, Soojeong Cho (SNU); 1♀, ditto, 17.vi.2023, Soojeong Cho (SNU); 1♂, ditto, 5.vi.2024, Minsuk Oh (NIBR). dez.pensoft.net Minsuk Oh et al.: New record and mitogenome of Heterotoma 230 Figure 4. Male genitalia of Heterotoma planicornis. A–D. Endosoma; E–G. Left paramere; H–J. Right paramere. Scale bars: 0.5 mm. Result In this study, the mitogenome sequence of H. planicornis is documented for the first time (Fig. 6). Its full length is 16,179 bp, comprising 13 protein-coding genes (PCGs), two rRNA genes, 22 tRNA genes, and a control region. The gene composition is similar to that of other previously sequenced mitochondrial genomes of Miridae (Wang et al. 2014). The average GC content of the full sequence was 28.46%. Among PCGs, GC content was the lowest at ATP synthase subunit 8 (atp8, 22.01%) and the highest at cytochrome oxidase subunit I (COI, 36.39%). Unlike other mirid genome sequences, the mitogenome of H. planicornis did not have a positive GC skew except in the control region. Additional information about each gene sequence is provided in Table 3. In the NJ tree and pairwise distance analysis using COI sequences (Fig. 7, Table 4), the sequence extracted from the mitogenome of a male H. planicornis (PV879916) and one female (PV850444) showed a 1.90–1.98% genetic distance compared to the Canadian population (HQ105735.1, HQ105736.1), whereas sequences from a last instar nymph (PV850443) and a male (PV789749) were almost identical. Dtsch. Entomol. Z. 72 (2) 2025, 225–235 dez.pensoft.net 231 Figure 5. Female genitalia of Heterotoma planicornis. A. Bursa copulatrix (before dissecting posterior wall); B. Bursa copulatrix, dorsal view; C. Bursa copulatrix, ventral view; D, E. Posterior wall; F, G. Gonapophysis II. Scale bar: 0.6 mm. Table 3. Gene composition of the mitochondrial genome of Heterotoma planicornis. Sequences were annotated by MITOS2, powered by the Proksee webserver. GC contents of each gene were calculated using the VectorBuilder GC Content Calculator (Cyagen Biosciences Inc. 2024). Type Name Start Stop Strand GC contents (%) Anti-codon tRNA trnI (tRNA-Isoleucine) 1 66 1 21.21 GAT tRNA trnQ (tRNA-Glutamine) 64 132 -1 24.64 TTG tRNA trnM (tRNA-Methionine) 132 200 1 24.64 CAT PCG nad2 (NADH dehydrogenase subunits II) 258 1202 1 27.2 - tRNA trnW (tRNA-Tryptophan) 1201 1264 1 15.63 TCA tRNA trnC (tRNA-Cysteine) 1257 1319 -1 26.98 GCA tRNA trnY (tRNA-Tyrosine) 1319 1382 -1 26.56 GTA PCG cox1 (cytochrome oxidase subunits I) 1383 2921 1 36.39 - tRNA trnL2 (tRNA-Leucine2) 2917 2981 1 26.15 TAA PCG cox2 (cytochrome oxidase subunits II) 2983 3664 1 32.84 - tRNA trnK (tRNA-Lysine) 3662 3731 1 31.43 CTT tRNA trnD (tRNA-Aspartic Acid) 3731 3795 1 10.77 GTC PCG atp8 (ATP synthase subunits 8) 3796 3954 1 22.01 - PCG atp6 (ATP synthase subunits 6) 3948 4619 1 31.99 - PCG cox3 (cytochrome oxidase subunits III) 4619 5405 1 33.29 - tRNA trnG (tRNA-Glycine) 5403 5464 1 20.97 TCC PCG nad3 (NADH dehydrogenase subunits III) 5480 5830 1 29.34 - tRNA trnA (tRNA-Alanine) 5817 5878 1 25.81 TGC tRNA trnR (tRNA-Arginine) 5883 5946 1 34.38 TCG tRNA trnN (tRNA-Asparagine) 5946 6012 1 22.39 GTT tRNA trnS1 (tRNA-Serine1) 6012 6080 1 33.33 GCT tRNA trnE (tRNA-Glutamic Acid) 6080 6143 1 21.88 TTC tRNA trnF (tRNA-Phenylalanine) 6142 6205 -1 26.56 GAA PCG nad5 (NADH dehydrogenase subunits V) 6186 7898 -1 25.51 - tRNA trnH (tRNA-Histidine) 7899 7962 -1 21.88 GTG PCG nad4 (NADH dehydrogenase subunits IV) 7962 9287 -1 25.57 - PCG nad4l (NADH dehydrogenase subunits IV) 9281 9574 -1 23.47 - tRNA trnT (tRNA-Threonine) 9589 9654 1 21.21 TGT tRNA trnP (tRNA-Proline) 9655 9719 -1 27.69 TGG PCG nad6 (NADH dehydrogenase subunits VI) 9730 10212 1 25.05 - PCG cob (Cytochrome b) 10213 11349 1 34.39 - tRNA trnS2 (tRNA-Serine2) 11348 11413 1 22.73 TGA PCG nad1 (NADH dehydrogenase subunits I) 11421 12347 -1 29.34 - tRNA trnL1 (tRNA-Leucine1) 12348 12414 -1 19.4 TAG rRNA rrnL (rRNA-Large) 12413 13651 -1 23.08 - tRNA trnV (tRNA-Valine) 13650 13717 -1 25 TAC rRNA rrnS (rRNA-Small) 13717 14473 -1 22.85 - Control region 14474 16179 1 29.95 dez.pensoft.net Minsuk Oh et al.: New record and mitogenome of Heterotoma 232 Table 4. Pairwise distances between analyzed Heterotoma planicornis specimens. H. planicornis HQ105735.1 H. planicornis HQ105736.1 H. planicornis Korea-MalePV879916 H. planicornis Korea-MalePV789749 H. planicornis Korea-5th_instarPV850443 H. planicornis Korea-FemalePV850444 H. planicornis HQ105735.1 H. planicornis HQ105736.1 0.0000 H. planicornis KoreaMalePV879916 0.0190 0.0198 H. planicornis KoreaMale-PV789749 0.0000 0.0000 0.0180 H. planicornis Korea5th_instar-PV850443 0.0000 0.0000 0.0180 0.0000 H. planicornis KoreaFemale-PV850444 0.0190 0.0198 0.0000 0.0180 0.0180 Figure 6. Mitochondrial genome of Heterotoma planicornis. Abbreviations of each gene were documented in Table 3. GC contents were indicated as a black circular graph, and GC-skew was indicated as green(+) or purple(-) in a circular graph. GC contents and GC skew were plotted as the deviation from the mean GC value of the full sequence. The innermost side of the circular graph shows sequence length, numerically marked with a 2 kbp interval. The circular map of the mitochondrial genome was visualized by the web server Proksee and annotated by MITOS2. Dtsch. Entomol. Z. 72 (2) 2025, 225–235 dez.pensoft.net 233 Discussion Morphological features and anticipated introduction pathways of H. planicornis in Korea Heterotoma planicornis is morphologically similar to H. merioptera (Scopoli, 1763), leading to considerable taxonomic confusion reviewed by Kment and Bryja (2006, 2012). Current diagnostic characters primarily rely on differences in genital structures and the length ratio of antennal segments II and III. Heterotoma planicornis tends to have a shorter second segment and a longer third segment compared to H. merioptera (Wagner 1968, 1974; Goula 1990). Additionally, Kment and Bryja (2006) proposed the ratio of vertex width to compound eye width as an alternative diagnostic character, but they noted that it overlaps between the two species. The lengths of antennal segments III:II in the examined Korean specimens (0.289–0.303) were consistent with the value reported for H. planicornis by Kment and Bryja (2006). The presence of two apically sinuate, unfused endosomal sclerites and parameres in the examined specimens was consistent with the illustrations of H. planicornis provided by Tamanini (1962) and Goula (1990). However, a medially bifurcated sclerite, as in fig. 2 of Kment and Bryja (2012), was not observed in the Korean population. In the figures of the endosomal sclerites provided by Tamanini (1962), bifurcated sclerites were observed in both Heterotoma species. Considering that intraspecific variation in the branching of endosomal sclerites can occur in other species of Orthotylinae (Oh et al. 2022), the above differences are possibly due to variation. It is concluded that the Korean material examined pertains to H. planicornis. Our study confirmed that the Korean H. planicornis uses Spiraea japonica as a breeding host. This plant species is native to Japan but has been widely planted in Europe since its introduction in the mid-19th century (Trees and Shrubs Online 2025). In Korea, cultivated S. japonica plants are mostly imported from Japan and European countries such as Germany and the Netherlands. Although various cultivars of S. japonica are cultivated in Japan, no species of the genus Heterotoma have been reported from the country or from any other locality in East Asia. Considering that all species of the genus Heterotoma are native to Europe, it is highly likely that individuals detected in Korea were introduced from populations in Europe, presumably on their host plants. Since eggs of Heterotoma species can overwinter and be transported with host saplings (Kment and Bryja 2006), the population might have been imported as eggs. Molecular characteristics of Korean individuals of Heterotoma planicornis Individuals used in the analysis were collected from the same plant, and no significant differences were observed in their external characteristics. An analysis of COI sequences from H. planicornis collected in this study and from taxonomically related species revealed that the Korean population comprises two distinct populations, exhibiting a 1.80–1.98% sequence difference (Table 4). This was also reflected in the NJ tree, where one population clustered with the North American population, while another branched into a separate node. However, due to the unavailability of European specimens, the relationship between Korean and European populations could not be ascertained. With sufficient sequence data on H. planicornis from diverse geographic regions, it could be possible to trace the origin and introduction pathway of the Korean population. In species delimitation studies of Miridae using COI sequences, genetic distances less than 2% are largely considered conspecific (Jung and Lee 2012; Kim and Jung 2018). A recent DNA barcoding study by Kim and Jung (2018) on 273 individuals from 123 Miridae species revealed intraspecific variation ranging from 0 to 2.8%, with a mean variation of only 0.2%. Furthermore, in their analysis of 27 individuals from 14 Orthotylinae species, maximum intraspecific variation did not exceed 2%, while interspecific variation ranged from 1.6% to 27.5%. However, as currently only limited molecular data are available for populations of H. planicornis, further research is necessary. Acknowledgments We would like to express our sincere gratitude to Dr. Heungsik Lee (Animal and Plant Quarantine Agency, Gimcheon, Korea) for his invaluable assistance in the identification of the Korean Heterotoma population. Thanks are extended to Dr. Dávid Rédei (Department of Entomology, National Chung Hsing University, Taichung, Taiwan) and Dr. Petr Kment (Department of Entomology, National Museum, Prague, Czech Republic) Figure 7. Neighbor-joining tree based on 14 COI sequences, including molecular data of Korean Heterotoma planicornis.