Abstract
Background
The genus Tuponia Reuter, 1875 belongs to the subfamily Phylinae and comprises 91 species worldwide. Before this study, only T.koreana Kim & Jung had been recorded from the Korean Peninsula.
New information
Two species of Tuponia Reuter, 1910 are recognised from the Korean Peninsula including the first record of T.mongolica Drapolyuk, 1980. T.koreana Kim & Jung, 2021 is proposed as a junior synonym of T.chinensis Zheng & Li, 1992. The species is identified, based on the dorsal habitus and male and female genitalic structures. A brief discussion of the distribution of Korean Tuponia species also is presented.
Keywords: Exaeretini, Tuponia , Korean Peninsula, new record, synonymy
Introduction
The phyline plant bug genus Tuponia Reuter, 1875 comprises 91 species worldwide (Schuh 2002, Kim et al. 2021). This group is distributed in the Palaearctic, Oriental Asia and northern Afrotropical Region and a large number of species use diverse Tamarix plants as their breeding host (Kerzhner and Josifov 1999, Li and Liu 2016). In the United States, Tamarix has been recognised as an invasive species, producing negative effects on ecosystems. For example, they accelerate salinisation of the soil (Ladenburger et al. 2006), increase wildfire severity by replacing fire-resistant plants and deplete groundwater (Pattison et al. 2011, Drus et al. 2013). In Korea, Tamarix has been planted on a limited basis as an ornamental tree, with natural populations only observed in Ansan and Incheon, which are distributed throughout the western coast of the Peninsula (NIBR 2011, Lee et al. 2018). Since a natural population exists in Korea and the plants can survive in many environments, research on related insects is important to understand more about the Tamarix species and its effect on the local ecosystem.
After the genus Tuponia was first erected by Reuter (1875), taxonomic analysis for this genus was actively conducted. In East Asia, Drapolyuk revised the subgenera Chlorotuponia Wagner, 1964 and Tuponia in Russia and Mongolia with the addition of 18 new species (Drapolyuk 1980, Drapolyuk 1982) and Chinese Tuponia was recently revised by Li and Liu (2016) with movement of 16 species including three new species. Later, Konstantinov (2016) reassessed the validity of some Tuponia species and synonymised three endemic Chinese species, including two of the new species described by Li and Liu (2016). In Korea, Tuponiakoreana Kim & Jung was recently described by Kim et al. (2021). Herein we discuss two species of Tuponia in Korea, including a new distributional record of T.mongolica Drapolyuk and suggest a new synonym of Tuponiakoreana Kim & Jung syn. n. with T.chinensis Zheng & Li. Images of the dorsal habitus and genitalic structures of both sexes were presented for these two Korean Tuponia species.
Materials and methods
All examined specimens are deposited in the collection of Insect Biosystematics Laboratory, Research Institute for Agriculture and Life Science, Seoul National University, Korea (SNU) and National Institute of Biological Resources (NIBR), Incheon, Korea. External characteristics were observed under a Leica Z16 APO microscope and digital images were obtained with a Leica DMC 5400 camera. Genitalic 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 millimetres, unless otherwise noted.
Terminology used to describe the genitalic structures follows Menard et al. (2014), Li and Liu (2016) and Kim et al. (2021), with the following abbreviations: Male: HP: hypophysis; SG: secondary gonopore; SL: sensory lobe. Female: DP: dorsal labiate plate; IL: interramal lobe; IS: interramal sclerite; RM: ramus; SR: sclerotized ring; VP: ventral labiate plate.
Taxon treatments
Tuponia
Reuter, 1875
726A4055-615A-515D-B8D9-30C1C1F362AF
Tuponia Reuter, 1875 - Reuter 1875: 98 as a subgenus of Megalodactylus, upgraded by Reuter 1878: 16.
Tuponia Capsus tamarisci Perris, 1857 Synonyms: TuponiacunealisReuter OM (1902) Capsidae novae mediterraneae. IV. Öfversigt af Finska Vetenskapssocietetens Förhandlingar B 44: 51–70.: 65. TuponianotatusFieber FX (1858) Criterien zur generischen Theilung der Phytocoriden (Capsini auct.). Wiener entomologische Monatschrift 2: 289–327, 329–347, 388, 1 pl.: 338.
Diagnosis
Tuponia can be recognised by the following characters: body elongate oval; dorsum somewhat shining, without distinct punctures and covered with pale, sericeous setae and dark setae; basic colouration greenish or yellowish-brown with brown, dark brown or reddish spots; membrane dark grey, vein pale green to brown; endosoma elongated, C-, S- or J-shaped; sometimes with sclerotised apical structures and membranous lobes; secondary gonopore usually developed between subapical part of sclerotised lobes; sclerotised ring elongated oval, interramal sclerite elongate and smooth. For detailed diagnostic characters and figures, see Drapolyuk (1980), Drapolyuk (1982), Li and Liu (2016) and Kim et al. (2021).
Distribution
Afrotropical Region, Oriental Region, Palaearctic Region (Drapolyuk 1980, Drapolyuk 1982, Kerzhner and Josifov 1999, Li and Liu 2016).
SubgenusTuponia
The subgenus can be recognised by the following characters: body elongate oval (male) or suboval (female), comparatively moderate to large (2.1–4.1 mm, usually around 3.0 mm); dorsum somewhat shining, without distinct punctures and covered with pale, sericeous setae; apical 1/3 of clavus and median part of hemelytra usually with suberect, dark brown to reddish setae, forms transverse line; basic colouration pale green to yellowish-green, partly brownish or tinged with red; membrane dark grey, vein pale green to brown; endosoma C-, S- or J-shaped, usually with one or two sclerotised apical structures; endosomal membrane rather developed, situated along apical processes; secondary gonopore usually developed between the subapical part of sclerotised lobes; female sclerotised ring elongate oval, surrounded by wide and weakly sclerotised labiate plates; bursa copulatrix with a pair of distinct, round structures dorsally; posterior wall with elongated, distally round interramal sclerites and weakly sclerotised, rough interramal lobe.
SubgenusChlorotuponia
This subgenus was established by Wagner (1964) and can be recognised by the following characters: body elongate oval (male) or suboval (female), comparatively small (1.7–2.5 mm, usually around 2.0 mm); dorsum somewhat shining, without distinct punctures and covered with long, pale hair and short, suberect brownish setae; basic colouration green to yellowish-green, usually concolorous; membrane dark grey, vein pale green to brown; endosoma C-, S- or J-shaped, usually with one or two sclerotised apical structures; endosomal membrane weakly developed, indistinct; secondary gonopore usually developed between the subapical part of sclerotised lobes; female sclerotised ring elongate oval, surrounded by wide and weakly sclerotised labiate plates; inner margin of bursa copulatrix with thin, arch-shaped sclerotised structure; posterior wall with elongated, distally round interramal sclerites and weakly sclerotised, interramal lobe with rough surface
Tuponia (Chlorotuponia) chinensis
Zheng & Li, 1992
E6C1E63C-6EC6-5D76-B64D-737E1FE70C16
Tuponiachinensis Zheng & Li, 1992 - Zheng and Li 1992: 12.
Tuponiakoreana Kim & Jung, 2021 - Kim et al. 2021: 1268. New synonymy.
Materials
Type status: Other material. Occurrence: recordedBy: Jihwan Park; individualCount: 10; sex: 5♂, 5♀; lifeStage: adult; Taxon: scientificName: Tuponia (Chlorotuponia) chinensis Zheng & Li, 1992; Location: country: Republic of Korea; stateProvince: Incheon-si; locality: Sorae Wetlands Ecology Park, Nonhyeon-dong, Namdong-gu; Identification: identifiedBy: MinSuk Oh; Event: eventDate: 31.vii.2021; habitat: on Tamarixchinensis; Record Level: institutionCode: SNU
Type status: Other material. Occurrence: recordedBy: Jihwan Park; individualCount: 1; sex: 1♀; lifeStage: adult; Taxon: scientificName: Tuponia (Chlorotuponia) chinensis Zheng & Li, 1992; Location: country: Republic of Korea; stateProvince: Incheon-si; locality: Sorae Wetlands Ecology Park, Nonhyeon-dong, Namdong-gu; Identification: identifiedBy: MinSuk Oh; Event: eventDate: 31.vii.2021; habitat: on Tamarixchinensis; Record Level: institutionCode: NIBR
Diagnosis
Recognised by elongate oval body, 1.8–2.3 mm; basic colouration pale green to yellowish-green, weakly shining (Fig. 1A–C, Fig. 2A–D); dorsum covered with pale sericeous setae and dark brown setae; labium reaches metacoxa; hemelytra pale green, covered with long, sericeous setae and dark brown, simple setae; tibial spine blackish-brown, base without dark spot. Male genitalia (Fig. 4A–I): Endosoma S-shaped, with two elongated, twisted sclerites; secondary gonopore situated subapically between the two sclerites; left paramere with thick and round sensory lobe; sub-basal part of sensory lobe with elongated, pointed-end process; hypophysis sub-triangular, pointed-end; right paramere rather elongated and flat, slightly curved; hypophysis short. Female genitalia (Fig. 6A–C): Sclerotised ring ovoid, thin-rimmed; interramal sclerites slender and elongated. For detailed diagnostic characters and figures, see Zheng and Li (1992), Li and Liu (2016) and Kim et al. (2021).
Korean Tuponia species, live individuals.
Figure 1a.

A: Tuponiachinensis, male;
Figure 1b.
B: ditto, female;
Figure 1c.

C: ditto, last instar;
Figure 1d.
D: T.mongolica, male;
Figure 1e.
E: ditto, female;
Figure 1f.
F: ditto, last instar.
Figure 2.
Dorsal habitus of Korean Tuponiachinensis. A, B Male; C, D Female. Scale bar: 1.0 mm.
Figure 4.
Male genital structure of Korean Tuponiachinensis. A, E–H, endosoma; B, I phallotheca; C left paramere; D right paramere. Scale bar: 0.1 mm.
Figure 6.
Female genital structure of Korean Tuponia species. A–C, T.chinensis; D–G T.mongolica (A, D: bursa copulatrix (before dissected posterior wall); B, E bursa copulatrix (dorsal view); F bursa copulatrix (ventral view); C, G: posterior wall). Scale bar: 0.2 mm.
Measurements
Male (n = 5). Total body length 1.86–2.07; head width across eyes 0.53–0.57; vertex width 0.29–0.32; lengths of antennal segment I–IV 0.14–0.18, 0.54–0.62, 0.38–0.39, 0.20–0.21; labial length 0.68–0.74; mesal pronotal length including collar 0.33–0.38; basal pronotal width 0.68–0.82; width across hemelytron 0.76–0.93; cuneal length 0.32–0.36; cuneal width 0.19–0.23; lengths of metafemur, tibia and tarsus 0.74–0.76, 1.07–1.14, 0.35–0.39. Female (n = 5). Total body length 1.85–2.06; head width across eyes 0.55–0.60; vertex width 0.32–0.35; lengths of antennal segment I–IV 0.15–0.19, 0.55–0.63, 0.34–0.41, 0.22–0.25; labial length 0.71–0.74; mesal pronotal length including collar 0.36–0.39; basal pronotal width 0.71–0.81; width across hemelytron 0.83–1.01; cuneal length 0.33–0.37; cuneal width 0.20–0.26; lengths of metafemur, tibia and tarsus 0.79–0.82, 1.08–1.18, 0.32–0.37.
Distribution
Korea (Incheon), China (Tianjin, Hebei, Shandong, Ningxia, Shaanxi) (Zheng and Li 1992, Li and Liu 2016, Kim et al. 2021).
Biology
Known host plant is Tamarixchinensis (Tamaricaceae) (Zheng and Li 1992, Li and Liu 2016, Kim et al. 2021).
Notes
We examined specimens of Tuponiachinensis in SNU and propose T.koreana Kim & Jung as a junior synonym of T.chinensis Zheng & Li. The diagnostic characteristics of T.koreana nearly match those of T.chinensis and the genitalia of the two nominal species are identical. Kim et al. (2021) separated T.koreana from T.chinensis by the following characters: i) metatarsal segment II distinctly shorter than segment III, ii) left paramere with one long process pointing down and iii) endosoma without visible secondary gonopore. However, as in Fig. 7, they only measured the length of each segment at the ventral side of the metatarsus and the actual length of segment II is not distinctly shorter than that of segment III. Additionally, in the structure of the left paramere, our specimen shows a straight lateral process (Fig. 4C). However, we could not find other structural differences and concluded that these were minor intraspecific variations. In addition, Kim et al. (2021) stated that a secondary gonopore of T.koreana is ‘clearly invisible,’ but it can be seen upon displacement of the two apical sclerites of the endosoma, as shown in Fig. 4E–H. When compared with the description of Zheng and Li (1992), the secondary gonopore of the Korean specimen looks conspecific in its subapical location and rugged margin.
Figure 7.
Metatarsus of Tuponiachinensis. Red arrow indicates segment borders in the dorsal side of tarsus and blue arrow indicates segment borders in the ventral side. Ratio of tarsal segments visualised with red bar (dorsal view: a), blue bar (ventral view: b) and black bar (actual length: c). A male; B female; C holotype male, a traced picture of Kim et al. (2021): fig. 1C; D a traced picture of Kim et al. (2021): fig. 1D.
Tuponia (Tuponia) mongolica
Drapolyuk, 1980
1A125432-BF4C-5C1D-A052-FF25436C5EDA
Tuponiamongolica Drapolyuk, 1980 - Drapolyuk 1980: 63.
Tuponiatamaricicola Hsiao and Meng, 1963 - Hsiao and Meng 1963: 447, 449. (junior primary homonym of Tuponiatamaricicola Lindberg, 1939)
Tuponiahsiaoi Zheng and Li, 1992 - Zheng and Li 1992: 10.
Materials
Type status: Other material. Occurrence: recordedBy: WonGun Kim; individualCount: 7; sex: 2♂, 5♀; lifeStage: adult; Taxon: scientificName: Tuponia (Tuponia) mongolica Drapolyuk, 1980; Location: country: Republic of Korea; stateProvince: Incheon-si; locality: Sorae Wetlands Ecology Park, Nonhyeon-dong, Namdong-gu; Identification: identifiedBy: MinSuk Oh; Event: eventDate: 19.viii.2022; habitat: on Tamarixchinensis; Record Level: institutionCode: SNU
Type status: Other material. Occurrence: recordedBy: WonGun Kim; individualCount: 1; sex: 1♀; lifeStage: adult; Taxon: scientificName: Tuponia (Tuponia) mongolica Drapolyuk, 1980; Location: country: Republic of Korea; stateProvince: Incheon-si; locality: Sorae Wetlands Ecology Park, Nonhyeon-dong, Namdong-gu; Identification: identifiedBy: MinSuk Oh; Event: eventDate: 19.viii.2022; habitat: on Tamarixchinensis; Record Level: institutionCode: NIBR
Diagnosis
Recognised by elongate oval body, 2.8–3.5 mm; basic colouration pale yellowish-green, weakly shining (Fig. 1D–F, Fig. 3A–D); dorsum covered with pale sericeous setae and dark brown setae; labium reaches metacoxa; hemelytra pale yellowish-green, partly tinged with pale orange; posterior half of clavus and posterior 1/3 of corium densely covered with brown setae; tibial spine blackish-brown. Male genitalia (Fig. 5A–D): Endosoma S-shaped, with two elongated, pointed-end sclerites and laterally serrate, membranous lobe; secondary gonopore developed between two sclerites; left paramere laterally wide; hypophysis twisted, apically hooked, sensory lobe with small pointed-end protuberance laterally. Female genitalia (Fig. 6D–G): Sclerotised ring ovoid, thin-rimmed; interramal sclerites slender and elongated. For detailed diagnostic characters and figures, see Drapolyuk (1980), Zheng and Li (1992) and Li and Liu (2016).
Figure 3.
Dorsal habitus of Korean Tuponiamongolica. A, B Male; C, D Female. Scale bar: 2.0 mm.
Figure 5.
Male genital structure of Korean Tuponiamongolica. A endosoma; B phallotheca; C left paramere; D right paramere. Scale bar: 0.2 mm.
Measurements
Male (n = 2). Total body length 2.75–3.03; head width across eyes 0.76–0.79; vertex width 0.34–0.35; lengths of antennal segment I–IV 0.22, 0.90–0.99, 0.81, 0.36; labial length 1.15–1.16; mesal pronotal length including collar 0.54–0.55; basal pronotal width 0.99–1.07; width across hemelytron 1.09–1.16; cuneal length 0.47–0.54; cuneal width 0.28–0.29; lengths of metafemur, tibia and tarsus 1.02–1.09, 1.59–1.62, 0.50. Female (n = 5). Total body length 2.83–3.14; head width across eyes 0.75–0.80; vertex width 0.34–0.38; lengths of antennal segment I–IV 0.21–0.24, 0.97–1.00, 0.72–0.88, 0.33–0.38; labial length 1.08–1.23; mesal pronotal length including collar 0.50–0.55; basal pronotal width 1.07–1.14; width across hemelytron 1.23–1.29; cuneal length 0.53–0.57; cuneal width 0.29–0.32; lengths of metafemur, tibia and tarsus 1.04–1.16, 1.61–1.67, 0.50–0.56.
Distribution
Korea (Incheon), China (inner Mongolia, Shandong, Hebei, Ningxia), Mongolia (Drapolyuk 1980, Li and Liu 2016).
Biology
Known host plants are Tamarix sp. and Tamarixchinensis (Tamaricaceae) (Drapolyuk 1980).
Notes
This species can be confused with T.jaxartensis Drapolyuk and T.zhenyuanensis Li & Liu, from which it is easily distinguished by endosoma with laterally serrate and elongated apical sclerites, phallotheca with a fin-like protrusion at the inner margin and a structural difference of parameres.
Discussion
Distributions of Korean Tamarix population and Tuponia
In this work, T.chinensis Zheng & Li and T.mongolica Drapolyuk were found at a coastal wetland near Incheon. This area provides an adequate environment for Tamarix and is adjacent to an international Airport, which may have been its source of introduction. According to a recent study, it is assumed that Ansan and Incheon populations of Tamarix were introduced about 40 years ago from China (Beijing) and from another unknown origin (Lee et al. 2018). Kim et al. (2021) also mentioned this hypothesis and suggested the need for subsequent research on the distributions of T.koreana and the closely allied species T.chinensis. Since these two species are regarded as conspecific, we can assume that T.chinensis was introduced along with the Tamarix. To support this, a comparison of the relationships of Tamarix in naturalised and intentional ornamental populations is crucial. In addition, considering the Tuponia diversity of China (Li and Liu 2016), further investigations may yield additional records for Korean Tuponia.
Supplementary Material
Acknowledgements
We deeply thank to Dr. Tomohide Yasunaga (American Museum of Natural History, New York, USA), Dr. Nikolay Simov (Bulgarian Academy of Sciences, Sofia, Bulgaria) and Dr. Attilio Carapezza (University of Palermo, Palermo, Italy) for improving the manuscript with invaluable comments and suggestions. This research was supported by the Basic Science Research Program through the National Research Foundation of Korea (NRF), funded by the Ministry of Education (NRF2020R1I1A2069484) and by a grant from the National Institute of Biological Resources funded by the Ministry of Environment (MOE) of the Republic of Korea (NIBR 202304203, NIBR 202333201).
References
- Drapolyuk I. S. Review of the capsid bugs of the subgenusTuponia s. str. (Heteroptera, Miridae) from the USSR and Mongolia. Nasekomye Mongolii. 1980;7:43–68. Russian. [Google Scholar]
- Drapolyuk I. S. Review of capsid bugs of the subgenusChlorotuponia (Heteroptera, Miridae) from the USSR and Mongolia. Nasekomye Mongolii. 1982;8:148–183. Russian. [Google Scholar]
- Drus G. M., Dudley T. L., Brooks M. L., Matchett J. R. The effect of leaf beetle herbivory on the fire behavior of tamarisk (Tamarixramosissima Lebed.) International Journal of Wildland Fire. 2013;22:446–458. doi: 10.1071/WF10089. [DOI] [Google Scholar]
- Hsiao T., Meng H. The plant-bugs collected from cotton-fields in China (Hemiptera, Heteroptera, Miridae) Acta Zoologica Sinica. 1963;15:439–449. [Google Scholar]
- Kerzhner I. M., Josifov M. In: Catalogue of the Heteroptera of the Palaearctic Region. Vol. Aukema B., Rieger C., editors. Vol. 3. The Netherlands Entomological Society, Amsterdam; 1999. Cimicomorpha II, Miridae.577 [Google Scholar]
- Kim J., Roca-Cusachs M., Lee B. W., Kim I. K., Jung S. First record of the genus Tuponia Reuter (Hemiptera: Heteroptera: Miridae: Phylinae) from the Korean Peninsula, with description of a new species and notes on natural history with Tamarix. Journal of Natural History. 2021;55(19-20):1267–1275. doi: 10.1080/00222933.2021.1943555. [DOI] [Google Scholar]
- Konstantinov F. V. New synonymies in the plant bug family Miridae (Hemiptera: Heteroptera) from Northern China. Zootaxa. 2016;4205(5):496–500. doi: 10.11646/zootaxa.4205.5.10. [DOI] [PubMed] [Google Scholar]
- Ladenburger C. G., Hild A. L., Kazmer D. J., Munn L. C. Soil salinity patterns in Tamarix invasions in the Bighorn Basin, Wyoming, USA. Journal of Arid Environments. 2006;65:111–128. doi: 10.1016/j.jaridenv.2005.07.004. [DOI] [Google Scholar]
- Lee S. R., Gaskin J. F., Kim Y. D. Molecular diagnosis for a Tamarix species from two reclaimed lands along the Yellow Sea in Korea inferred from genome wide SNP markers. Journal of Systematics and Evolution. 2018;57:247–255. doi: 10.1111/jse.12432. [DOI] [Google Scholar]
- Li X. M., Liu G. Q. The genus Tuponia Reuter, 1875 of China (Hemiptera: Heteroptera: Miridae: Phylinae: Exaeretini) with descriptions of three new species. Zootaxa. 2016;4114(2):101–122. doi: 10.11646/zootaxa.4114.2.1. [DOI] [PubMed] [Google Scholar]
- Menard K. L., Schuh R. T., Woolley J. B. Total-evidence phylogenetic analysis and reclassification of the Phylinae (Insecta: Heteroptera: Miridae), with the recognition of new tribes and subtribes and a redefinition of Phylini. Cladistics. 2014;30:391–427. doi: 10.1111/cla.12052. [DOI] [PubMed] [Google Scholar]
- NIBR NIBR Biogeographic Information Service. https://species.nibr.go.kr/geo/html/index.do?ktsn=120000077653. [2023-01-02T00:00:00+02:00]. https://species.nibr.go.kr/geo/html/index.do?ktsn=120000077653
- Pattison R. R., D'Antonio C. M., Dudley T. L. Biological control reduces growth, and alters water relations of the saltcedar tree (Tamarix spp.) in western Nevada, USA. Journal of Arid Environments. 2011;75:346–352. doi: 10.1016/j.jaridenv.2010.11.006. [DOI] [Google Scholar]
- Reuter O. M. Revisio critica Capsinarum, praecipue Scandinaviae et Fenniae. Akademisk Afhandling, Helsingfors; 1875. 101pp.+190 [Google Scholar]
- Reuter O. M. Hemiptera Gymnocerata Europae. Hémiptères Gymnocérates d'Europe, du bassin de la Méditerranée et de l'Asie russe. I. Acta Societatis Scientiarum Fennicae. 1878;13:1–188. [Google Scholar]
- Schuh R. T. On-line systematic catalog of plant bugs (Insecta: Heteroptera: Miridae) http://research.amnh.org/pbi/catalog/ [2023-01-29T00:00:00+02:00]. http://research.amnh.org/pbi/catalog/
- Wagner E. Neuer Beitrag zur Systematik der Gattung Tuponia Reuter 1875 (Hem. Het. Miridae). Entomologische Berichten. 1964;24:192–200, 215, 220. [Google Scholar]
- Zheng L., Li H. Genus Tuponia Reuter from China (Insecta, Heteroptera: Miridae) Reichenbachia. 1992;29:9–13. [Google Scholar]
Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.










