The pentatomine bug genus PlautiaStål, 1865, comprises 28 species and subspecies reported worldwide (Aukema, 2018;Ishikawa and Moriya, 2019), primarily across the Palearctic region (mostly East and Southeast Asia) and Australia (Linnavuori, 1975;Rider, 2006, 2015;Aukema, 2018;Ishikawa and Moriya, 2019). Among them, several species, such as P. staliScott, 1874, P. affinis (Dallas, 1851) and P. crossota (Dallas, 1851) are significant agricultural pests (Yasunaga et al., 1993;Jang et al., 2010;Sriram, 2020). These species cause frequent damage to various fruit trees and crops throughout East, Southeast and South Asia (Figs. 1A–I) (Kwon et al., 2001;Rider, 2015).
In East Asia, a total of seven species have been recorded from China (Rider et al., 2002), and a recent taxonomic study raised Japanese Plautia diversity from three to five species with the description of three new species and one synonym (Ishikawa and Moriya, 2019). The Korean Plautia were first recorded by Okamoto (1924) with P. stali from Jeju (also known as Quelpart Is.), and Lee (1971) added P. splendens Distant, 1900 based on material from Daegu and Ulleung Is. However, Kim et al. (2025) recently excluded the Korean distribution record of P. splendens, though they suggested that this exclusion requires future verification based on additional specimens.
Natural enemies of the genus Plautia were gradually reported from the mid-1950s onward (Rider, 2015). Almost all parasitoid records are concentrated on major pest species, P. affinis, P. crossota, P. stali, and P. splendens (Table 1). Among these records, the natural enemy record for P. splendens likely pertains to P. himechabaneIshikawa and Moriya, 2019, because Watanabe (1954) conducted his work on a population collected from Zetsuji (Shikoku) (Matsuo et al., 2014). Hymenopteran parasitoids pose a potential threat to Plautia eggs and after emergence, nymphs and adults can also be affected by oviposition by tachinid flies and the parasitic wasp Aridelus. Egg parasitoids fully utilize the energy from their host, and nymph and adult parasitoids also cause destructive damage to host fitness and fecundity through consumption of internal muscles, intestines and reproductive organs. While diverse parasitoids have been highlighted and studied, little has been reported about non-insect parasites observed on Plautia.
Acarine parasitoids affecting stink bugs include several reported observations of Leptus sp., and with parasitic Leptus impacting mortality and reproduction rates and may even causing the death of adult mirid bugs (González-Moraga et al., 2015;Vanitha and Saroj, 2015).
Our recent survey recognized two Plautia species, P. stali and P. himechabane. Plautia splendens was excluded from the Korean list based on the review of previous references and examination of the Korean population. Images of dorsal habitus and genitalic structures of both sexes are presented for two Korean Plautia species, with figures for observable variation in coloration and female spermatheca. We also describe updated two new associated mites of Plautia, Lobogynium sp. and Leptus andongensisHakimitabar et al., 2020, with detailed figures of both specimens.
Materials and Methods
All examined specimens 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. For the specimens of Plautia, 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 millimeters, or otherwise noted. For the mites, digital images of phoretic status were taken with a Leica DMA 5400 camera with a Leica Z16 APO microscope. Detached mites were cleared in lactic acid solution and mounted in PVA medium (Downs, 1943), and digital images of the mounted specimen were obtained with an Olympus DP27 camera attached to Olympus BX53 compound microscope equipped with differential interference contrast optical systems.
The terminology used to describe the genitalic structures of Plautia follows Redei (2017) and Ishikawa and Moriya (2019), and nomenclature for the general external morphologies of parasitic mites follows the Lindquist and Evans (1965) and Southcott (1992).
Results
Systematic accounts
Clade Mandibulata
Class Insecta
Order Hemiptera
Family Pentatomidae
Genus PlautiaStål, 1865 꼬마갈색노린재속(Figs. 2–10, 11A–G)
PlautiaStål, 1865: 191 (new genus). Type species: Cimex fimbriatus Fabricius, 1787, subsequent designation by Distant, 1902: 180 (= Pentatoma crossota Dallas, 1851).
Diagnosis.Plautia can be recognized by the following characters: Body ovoid, lateral margin sub-parallel (Figs. 2A– D, 3A–D); dorsum rather glabrous, with distinct, brown to dark brown punctures; basic coloration yellowish green to green; head subtriangular, apically blunt; pronotum glabrous and punctate, with weakly protruded humeral angle (Figs. 4A –F); scutellum wide and triangular, apically blunt; hemelytra usually with brown to reddish brown corium and greenish embolium (Figs. 5A–I); membrane pale grey; paramere wide and flat, with variable protrusion; phallus elongated oval, with well sclerotized vesical and a pair of membranous expansion; valvifer VIII sector-like, inner margin covered with stiff setae. For detailed diagnostic characters and figures, see Ishikawa and Moriya (2019).
Distribution. Australasia region, Palaearctic region (East Asia, Southeast Asia, South Asia, North Africa) (Linnavuori, 1975;Rider, 2006, 2015;Aukema, 2018;Ishikawa and Moriya, 2019).
Plautia staliScott, 1874 갈색날개노린재(Figs. 1A–I, 2A– D, 4D–F, 5A–H, 6A–F, 8A–H, 9A, C;10A;11A–F)
Plautia staliScott, 1874: 299 (new species).
Nezara amurensis Reuter in Autran and Reuter, 1888: 200 (new species, syn. by Kiritshenko, 1961: 443).
Diagnosis. Recognized by ovoid body, 9.0–11.0mm; dorsum rather glabrous; basic coloration yellowish green (Figs. 2A–D), brown to reddish brown before hibernation (Fig. 5G); apex 1/3 of antennal segment III, apex 1/2 of antennal segment IV and V blackish; lateral margin of pronotum usually with distinct blackish line; labium reaches or extending over metacoxa; hemelytra brown to reddish brown; Male genitalia (Figs. 6A–F, 8A–H): Ventral rim of genital capsule curved laterally, medially flat; dorsal sclerite ear-shaped, proximal margin comparatively thick; parameres flat and wide, lateral margin with two blunt protrusions, hypophysis straight, subtriangular; vesica well sclerotized and tapering apically, widened at apex and process of vesica developed laterally. Female genitalia (Figs. 9A, C;10A): Valvifer VIII subtriangular, with distinctly angled inner margin; apical part of spermatheca with one or two horn-shaped process. For detailed diagnostic characters and figures, see Ishikawa and Moriya (2019).
Measurements. Male (n = 5)/Female (n = 5). Total body length 8.92–10.97/ 9.39–11.16; head length 1.70–1.93/ 1.77–2.25; head width across eyes 2.36–2.74/ 2.40–2.80; vertex width 1.46–1.75/ 1.52–1.68; lengths of antennal segment I–V 0.45–0.56, 0.90–1.03, 0.94–1.07, 1.25–1.41, 1.41–1.54/ 0.43–0.58, 0.88–1.05, 1.08–1.26, 1.27–1.54, 1.42–1.71; labial length 4.54–5.34/ 4.75–5.18; mesal pronotal length 2.22–2.83/ 2.22–2.85; basal pronotal width 5.44–6.84/ 5.86–7.05; scutellum length 3.64–4.62/ 3.77– 4.44; scutellum width 3.47–4.36/ 3.69–4.38; width of abdomen across segment III 5.35–6.67/ 5.71–6.85; lengths of metafemur, tibia and tarsus 3.09–4.06, 2.74–3.29, 0.96– 1.24/ 3.44–4.35, 2.88–3.58, 1.15–1.33.
Material Examined. In total, 1,139 specimens (NIBR, SNU) were examined from the following localities: South Korea: Busan-si, Chungcheongbuk-do (Jecheon, Yeongdong), Chungcheongnam- do (Boryeong, Geumsan), Daegu-si, Daejeon-si, Gyeongsangbuk-do (Bonghwa, Pohang, Sangju, Ulleung Is.), Gyeonggi-do (Ansan, Anyang, Bucheon, Gapyeong, Gunpo, Gimpo, Gwacheon, Gwangju, Hwaseong, Icheon, Namyangju, Osan, Paju, Pocheon, Seongnam, Suwon, Uijeongbu, Yangpyeong, Yeoncheon, Yongin), Gyeongsangnam-do (Haman, Hamyang, Hapcheon, Jinju, Sancheong, Tongyeong), Gangwon- do (Chuncheon, Donghae, Gangneung, Inje, Yeongwol), Incheon-si, Jeju (Jeju, Seogwipo), Jeollabuk-do (Wanju), Jeollanam- do (Gurye, Gwangyang, Haenam, Jangseong, Suncheon, Wando).
Distribution. Korea (Whole), China, Japan, Russia (Far East), North America (Hawaii) (introduced) (Beardsley, 1979;Kwon et al., 2001;Ishikawa and Moriya, 2019).
Remarks. This species frequently occurs on the Korean Peninsula. According to Kwon et al. (2001), P. stali is regarded as an important economic pest for various orchards and crops, including more than 54 species in 11 families in Korea.
Normally, the basic coloration of P. stali is yellowish green (Figs. 2A–D), but it sometimes appears bluish green (Fig. 5C), brownish (Figs. 5H) or dark yellowish green (Figs. 5D– F). In the hibernation season, near fall to winter, their basic coloration turns brownish, or even reddish brown without any greenish tinge in some individuals (Fig. 5G). This color change returns to green in spring when the appropriate temperature change occurs (Kotaki, 1998), perhaps to better align with the coloration of their deciduous hosts. Ishikawa and Moriya (2019) described a blackish lateral pronotal margin as a diagnostic character of P. stali. Most of the Korean population shows a similar color pattern to the Japanese population (Fig. 4D), but some individuals have a somewhat brownish stripe (Fig. 4E) or widely obsolete stripe (Fig. 4F). This variation makes it difficult to distinguish this species from P. himechabane (Fig. 4A–C), so consideration of other characteristics like genitalic structure is essential for the Korean population.
When Lee (1971) first reported P. splendens from the Korean Peninsula, they distinguished this species from P. stali based on the following differences: i) smaller body size (less than 8mm); ii) brighter body coloration (light green to bluish green); and iii) lateral margin of pronotum without a dark line. However, these characteristics largely overlap with the diverse color variation of P. stali (Figs. 5A–H), leading to misidentification. As an example, we found specimens labeled as P. splendens from NIBR and SNU, and most were eventually identified as P. stali with a comparatively small size and obsolete dark stripe, not P. splendens nor P. himechabane.
Plautia himechabaneIshikawa and Moriya, 2019 꼬마갈색 노린재(Figs. 3A–D, 4A–C, 5I, 7A–F, 8I–P, 9B, D;10B– D;11G)
Plautia himechabaneIshikawa and Moriya, 2019: 475.
Diagnosis. Recognized by ovoid body, 8.0–10.0mm; dorsum rather glabrous; basic coloration yellowish green (Figs. 3A–D); apex 1/3 of antennal segment III, apex 1/2 of antennal segment IV and V brown to dark brown; lateral margin of pronotum without any dark stripe; labium extending over metacoxa; hemelytra brown to reddish brown; Male genitalia (Figs. 7A–F, 8I–P): Ventral rim of genital capsule curved laterally, medially flat; dorsal sclerite well sclerotized, with a minute protrusion; parameres flat and beak-shaped, darkened along lateral margin; vesica well sclerotized and cylindrical, situated with cordiform lateral process. Female genitalia (Figs. 9B, D;10B–D): Valvifer VIII sector-like, with brown spot near inner margin; apical part of spermatheca with elongated and medially branched horn-shaped process. For detailed diagnostic characters and figures, see Ishikawa and Moriya (2019).
Measurements. Male (n = 5)/Female (n = 5). Total body length 7.98–9.49/ 9.43–10.11; head length 1.56–1.96/ 1.89 –2.16; head width across eyes 2.15–2.48/ 2.40–2.61; vertex width 1.35–1.56/ 1.49–1.64; lengths of antennal segment I– V 0.45–0.52, 0.70–0.78, 0.87–1.07, 1.08–1.19, 1.22– 1.34/ 0.43–0.55, 0.76–0.92, 0.93–1.05, 1.08–1.34, 1.34– 1.47; labial length 4.87–5.10/ 4.77–5.46; mesal pronotal length 1.75–2.25/ 2.23–2.55; basal pronotal width 4.48– 5.61/ 5.52–6.08; scutellum length 2.91–3.60/ 3.68–3.98; scutellum width 2.82–3.51/ 3.48–3.88; width of abdomen across segment III 4.51–5.55/ 5.59–6.15; lengths of metafemur, tibia and tarsus 2.55–3.26, 2.30–2.89, 1.04– 1.06/ 2.95–3.43, 2.66–2.94, 1.05–1.17.
Material Examined. South Korea: Chungcheongnam-do: 1 ♀, Mt. Gyaeryeong, Banpo-myeon, Gongju-si, 27.v.2015, Hwaseop Song (NIBR). Gyeonggi-do: 1♂, SNU Suwon campus, Gwonseon-gu, Suwon-si, 18.ix.1999, Dong-Youl Lee (SNU). Gyeongsangnam-do: 1♀, Forest trail, Yuho-ri, Jangmok- myeon, Geoje-si, alt. 98m, 35°1'13''N 128°42'2''E, 22.v. 2009, Cheong Jiwoong (NIBR); 1♂, Mt. Sanseong, Yongtaeri, Hail-myeon, Goseong-gun, from light trap, 29-30.vi.1998, collector unknown (NIBR); 1♂, Geunho, Deokho-ri, Hi-myeon, Goseong-gun, from light trap, 21-22.vi.2001, Junseong Shin (NIBR); 1♀, Hyangro peak, Dongsan-ri, Sangri-myeon, Goseong-gun, from light trap, 14-15.viii.1996, Hyunggyu Joo (NIBR); 1♀, Mt. Imaeng, Jikjeon-ri, Bukcheon-myeon, Hadong- gun, from light trap, 28-29.viii.2000, Gyeongryun Han (NIBR); 2♂, Daesong-ri, Geumnam-myeon, Hadong-gun, alt. 279m, 34°58'35''N 127°52'11''E, 7.vii.2010, Park Joongsuk (NIBR); 1♀, Mt. Geum-o, Goryeong-ri, Jingyo-myeon, Hadonggun, from light trap, 28-29.viii.2000, Jaeshik Jeon (NIBR); 1 ♀, Guyang-ri, Macheon-myeon, Hamyang-gun, 19.vi.2015, Park Joongsuk (NIBR). Jeollanam-do: 1♂, Mountain trail, Mt. Gaya, Gwangyeong-dong, Gwangyang-si, 15.v.1999, Juhwan Son (NIBR); 1♀, Hanseok Tourist Farm, Mt. Gaya, Joongundong, Gwangyang-si, from light trap, 28-29.vii.1998, Taeho Ahn (NIBR); 1♀, same locality, from light trap, 30-31.viii.2000, Taeho Ahn (NIBR); 1♀, Chusan-ri, Okryeong-myeon, Gwangyang- si, 27.vii.1998, Hikyeong Choi (SNU); 1♀, Mt. Baekun, Chusan-ri, Okryeong-myeon, Gwangyang-si, 17.vi.2009, Soohyun Yang (NIBR).
Distribution. Korea (Central, South), China, Japan (Ishikawa and Moriya, 2019).
Remarks.Ishikawa and Moriya (2019) recently suggested that previous records of P. splendens were misidentified except for the endemic population in the Ogasawara islands, Japan. In addition, their work reclassified that ‘misidentified’ population as a new species, P. himechabane. However, they were unaware of the existence of Korean P. splendens, which also requires reexamination. In Korea, the first record of P. splendens can be found in the Lee (1971)’s illustrated encyclopedia, which includes a review of Korean Hemiptera. In this book, they described P. splendens with darkened antennal segment III– V, which correlates more closely to P. himechabane, and this record is followed by catalogues of Korean fauna without detailed revision of this taxon (Lee and Kwon, 1994;Kwon et al., 2001;Lee et al., 2013;Cho & Kwon, 2017). Recently, a monograph on Korean Pentatomidae reviewed the genus Plautia (Roca-Cusachs and Jung, 2020), but they also addressed P. himechabane as P. splendens. In conclusion, we have not been able to verify the existence of P. splendens on the Korean peninsula, so we also exclude P. splendens from the species list of Korean Plautia following Kim et al. (2025), and reclassify it as P. himechabane. Based on our examination of available specimens, Korean P. himechabane is far rarer than P. stali, with a population ratio > 1:70.
Class Chelicerata
Subclass Acari
Order Mesostigmata
Suborder Trigynaspida
Family Diplogyniidae
Lobogynium sp.(Figs. 11I, 12A)
Diagnosis. Body circular-shaped, brownish color, slightly longer than width. Most of dorsal setae short (Z5 longer), four pairs of thick and extended setae (apical tip barbed) bears along with lateral side of dorsal shield. Posterior-median part of body with deep incision from dorsum to ventral. Ventri-anal shield triangular-shaped, not abutted the posterior margin of body (Fig 12A). Peritremes long, extended at level of coxa I. Leg I slender and longest, without claws; barbed seta on genu III and IV. See more detailed characteristics of this genus in Trägårdh (1950), Plumari and Kazemi (2012).
Material Examined. Gyeonggi-do: 1♂, Anyang-si, on the left compound eye of Plautia stali, 27.xi.1997, ByungWoo Lee (SNU).
Remarks. This species has not previously been collected with Hemipteran insects, but has been observed on various other insects or beneath tree bark (Trägårdh, 1950;Ishikawa, 1968;Plumari and Kazemi, 2012;Trach, 2013;Saito, 2022). In addition, based on the phoretic behavior of Diplogyniid mites, we can presume that they climb onto stink bugs during overwintering, rather than acting as natural enemy of their host (Fig 11I). In this study, species-level identification of the specimen was not possible due to the lack of sufficient material. The suborder Trigynaspida was recently recorded in Korea for the first time by the second author (Oh et al., 2025) and was associated with passalid beetles (Coleoptera: Passalidae).
Order Trombidiformes
Family Erythraeidae
Leptus andongensisHakimitabar et al., 2020.(Figs. 11F–H, 12B–D)
Diagnosis. Body oval-shaped, yellowish color in mounted condition. Dorsal idiosoma covered strongly barbed setae (Fig 12B). Palp femur and genu with a single barbed seta (Fig 12C). Shape of scutum wider than longer, anterior margin of scutum distinctly concaved (Fig 12D). See more detailed characteristics in Hakimitarbar et al. (2020).
Material Examined. Jeollanam-do: 1 larva, Chusan-ri, Okryeong-myeon, Gwangyang-si, on the left side of mesopleuron of Plautia stali, 27.vii.1998, Dongsik Jung (SNU); 1 larva, Mt. Baekun, Chusan-ri, Okryeong-myeon, Gwangyangsi, on the left side of abdominal venter VII of Plautia himechabane, 17.vi.2009, Soohyun Yang (SNU).
Remarks.Leptus andongensis was reported from the body of Meimuna sp. (Auchenorrhyncha: Cicadidae) as an ectoparasite (Hakimitabal et al., 2020). In this study, we report L. andongensis from two other hemipteran hosts. This suggests that this species possesses a potentially broad host range across the order Hemiptera (Figs 11F–H).
Notes on observed genitalic structure variation on Korean Plautia species
In our observation, Plautia shows somewhat diverse female spermatheca structure between conspecific populations. Similar to the Japanese population, Korean P. himechabane has a subapically placed sclerotized zone, a round and distinct apical receptacle and an elongated horn-shaped process. However, the Korean population has a short (Fig. 10B) or bifurcate (Fig. 10C), sometimes elongated and protruded structure (Fig. 10D) situated at the median part of horn-shaped process. Despite this prominent difference, highly sclerotized structures like the valvifer and laterotergite maintain a certain structure, and variability of the spermatheca can be observed in Plautia (Staddon et al., 1994;Ishikawa and Moriya, 2019). In the case of Palomena (Pentatomidae), 15 different types of spermatheca have been reported in a single species, P. viridissima (Belousova, 2018). As in P. himechabane, P. stali also presents morphological variation of the spermatheca, which possesses one (Fig. 10A) or two horn-shaped processes, as Ishikawa and Moriya (2019) described. In conclusion, to conduct a more thorough classification of this highly variable taxon, overall consideration of their diverse coloration, size, and particular structures is necessary.
Distribution records of Plautia species from Korea, focusing on P. himechabane
Among two Korean Plautia, P. stali is widely distributed across South Korea, and its distribution in North Korea was also confirmed by Josifov and Kerzhner (1978). However, the distribution of P. himechabane remains problematic. Ishikawa and Moriya (2019) state that most of the Japanese P. himechabane population inhabits regions within a few kilometers of the coast. In our examination, this species is mainly distributed in the southern part of the Korean peninsula, supporting the observations of Ishikawa and Moriya (2019), but some specimens were also found at the central region (GG, CN) (red dots in Fig. 13). Distribution records of Lee (1971) and subsequent studies (Kwon et al., 2001) shown as highly sporadic (blue dots in Fig. 13), showing clearly different distribution tendencies compared with our observation. As mentioned above, a large number of misidentifications of Korean P. himechabane were found in some collections, and previous records may include misidentification. To construct a proper distribution map for this tentative pest, reconfirmation of previous records is necessary.





















KSAE