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Mycophagous Gall Midges (Diptera: Cecidomyiidae) in Korea: Newly Recorded Species with Discussion on Four Years of Taxonomic Inventory PDF

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Preview Mycophagous Gall Midges (Diptera: Cecidomyiidae) in Korea: Newly Recorded Species with Discussion on Four Years of Taxonomic Inventory

Anim. Syst. Evol. Divers. Vol. 36, No. 1: 60-77, January 2020 https://doi.org/10.5635/ASED.2020.36.1.055 Review article Mycophagous Gall Midges (Diptera: Cecidomyiidae) in Korea: Newly Recorded Species with Discussion on Four Years of Taxonomic Inventory Daseul Ham1, Mathias Jaschhof 2, Yeon Jae Bae3,* 1Korean Entomological Institute, Korea University, Seoul 02841, Korea 2Station Linné, Ölands Skogsby 161, SE-38693 Färjestaden, Sweden 3Division of Environmental Science and Ecological Engineering, College of Life Sciences, Korea University, Seoul 02841, Korea ABSTRACT Cecidomyiidae (Diptera) consists of six subfamilies, which are divided into three groups according to larval ecological habits (phytophagous, mycophagous, and zoophagous). The five basal subfamilies of Cecidomyiidae consist entirely of mycophagous species, with approximately 1500 species described worldwide and 29 previously known to occur in Korea. In this study, 37 named species (1 Lestremiinae, 29 Micromyinae, 4 Winnertziinae, and 3 Porricondylinae species) are newly reported from South Korea. We excluded Lestremia yasukunii Shinji from the list of Korean mycophagous cecidomyiids as it is a nomen nudum. Therefore, we herein officially recognize 65 species, 30 genera, and four subfamilies for the Korean mycophagous cecidomyiid fauna. We also provide diagnoses and photographs to aid species identification and discussion on the four years of gall midge taxonomic inventory in South Korea. Keywords: C ecidomyiidae species new to Korea, taxonomic diagnosis, gall-midge taxonomy, insect identification, Korean gall-midge fauna, Korean indigenous species survey INTRODUCTION Sweden (Jaschhof, unpublished data). The Swedish fauna of fungus-feeding gall midges has been, for nearly 15 years, the Cecidomyiidae (gall midges) is well-known as a family of subject of intensive taxonomic inventory work (e.g., Jaschhof plant-feeders; however, it also contains many predatory and and Jaschhof, 2009, 2013), and, as a result, it may be regard- mycophagous species. Unlike many other insect groups, in- ed as the best-studied national fauna worldwide. At a higher cluding gall-inducing insects and other plant-feeding gall level, the family Cecidomyiidae is possibly currently the most midges, mycophagous cecidomyiids are exclusively studied species-rich family of Diptera, with an estimated 2,000,000 by professional researchers. No amateur entomologist is in- species (Hebert et al., 2016). More than 800 species of Ce- terested in these tiny, elusive midges, nor in their faunistic cidomyiidae were recorded from one site at a Neotropical aspects. It is fair to say that mycophagous gall midges are cloud forest, rendering this family the most speciose of all the probably the most poorly researched, large subgroup of the dipteran families recorded there (Borkent et al., 2018; Brown order Diptera. In contrast, it is known that mycophagous cec- et al., 2018). ids occur worldwide - especially in woodlands - and that they The mycophagous Cecidomyiidae are classified into five are so speciose that the number of species in specific regions subfamilies, namely Catotrichinae, Lestremiinae, Micromy- or worldwide is impossible to estimate. There are currently inae, Winnertziinae, and Porricondylinae (the sixth subfamily, ~1,500 named species of mycophagous cecidomyiids in the Cecidomyiinae, contains all the herbivorous and predatory world, comprising approximately one-fourth of all Cecido- species). Micromyinae, with 38 extant genera and 579 extant myiidae (Gagné and Jaschhof, 2017). Of these, 560 species species, and Porricondylinae, with 84 extant genera and 501 have been recorded in only one country in northern Europe: extant species, are the most diverse fungus-feeding subfam- This is an Open Access article distributed under the terms of the Creative *To whom correspondence should be addressed Commons Attribution Non-Commercial License (http://creativecommons.org/ Tel: 82-2-3290-3408, Fax: 82-2-3290-3623 licenses/by-nc/3.0/) which permits unrestricted non-commercial use, distribution, E-mail: [email protected] and reproduction in any medium, provided the original work is properly cited. eISSN 2234-8190 Copyright The Korean Society of Systematic Zoology Mycophagous Gall Midges in Korea ilies (Gagné and Jaschhof, 2017). Most of these genera and lished report) are documented in publications (e.g., Jaschhof, species were discovered and named from the Palearctic re- 2000, 2001). In summary, it can be stated that no long-term, gion, as Europe has traditionally been the center of taxonomic concerted effort has ever been made to investigate the Far studies and a home base for the research of faunas from oth- Eastern hotspot systematically. Consequently, we have only er continents (cf. Jaschhof and Jaschhof, 2009: 18ff.; 2013: vague ideas of the faunal size and composition in that region. 22ff.). Moreover, tools for species identification, such as specimen The biology of the midges in question is even more insuffi- collections, published descriptions and keys, and DNA bar- ciently researched than their taxonomy. Their larvae, which is code libraries, are of limited use because of incomplete spe- the ontogenetic stage occupied with feeding (adults usually do cies coverage and unsatisfactory quality (see Discussion). not assimilate food), are described for only a small proportion Clearly, the Far Eastern Palearctic is the place to perform pio- of the named species. Details regarding host (fungus) special- neer work for scholars focused on mycophagous cecids. ization, habitat requirements, and interactions with other spe- Korea was a blank spot on the world map of mycophagous cies are poorly understood. Larvae are usually found in asso- cecid diversity until 2003, when a species of Mycophila Felt ciation with plant litter, with soil and rotting wood being the (in Micromyinae) was reported as a pest infesting mushroom most common larval habitats. Furthermore, larvae are gener- cultivations (Lee and Kim, 2003). Two similar cases, but re- ally considered to pierce and suck fungal hyphae penetrating garding the genus Camptomyia Kieffer (in Porricondylinae), plant detritus (Mamaev and Krivosheina, 1965). Their eco- were reported by Shin et al. (2011). Eventually, a taxonomic logical role is, therefore, best described as mycophagous de- inventory focused on the mycophagous subfamilies of Ce- tritobiont, or in more general terms, detritivores. Krivosheina cidomyiidae was started in 2016 as part of the framework (2006) reviewed the ecology of bark- and wood-dwelling program “Korean Indigenous Species Survey-Insects,” ini- (xylobiont) species in an evolutionary context. However, tiated by the National Institute of Biological Resources in much remains unknown regarding the biology of adults, Incheon, South Korea (https://www.nibr.go.kr/cmn/sym/mnu/ which is the ontogenetic stage commonly used for species mpm/512010000/enHtmlMenuView.do). Since then, a total identification. Moreover, the classification of mycophagous of 29 species have been recorded as occurring in the country Cecidomyiidae is largely based on interpretation from adult (Ham and Bae, 2017; Ham et al., 2018, 2019; Jaschhof et morphology, with males having an advantage over females al., 2018), Lestremia yasukunii Shinji, a dubious name that in providing data for taxonomic and phylogenetic interpreta- was herein deleted from the Korean checklist (cf. Gagné and tion (Jaschhof and Jaschhof, 2009, 2013). In recent years, an Jaschhof, 2017), not included. integrated approach using both morphological and molecular In the present paper, we document findings of 37 additional data has been increasingly applied to species delimitation (e.g., species, namely 1 Lestremiinae, 29 Micromyinae, 4 Winnert- Jaschhof, 2016b) and phylogeny reconstruction (Sikora et al., ziinae, and 3 Porricondylinae. This more than doubles the 2019). number of species recorded from Korea, which now stands The realization that the Far East of the Palearctic region is a at 66. Besides these species, we are aware of the existence hotspot of mycophagous cecid diversity has developed grad- of twice as many species among the material that we cannot ually since the 1970s, initiated by studies by Boris M. Ma- name at this stage, either because the available literature is maev in Far Eastern Russia (many different articles, see list in not sufficient for a positive identification or because they are Gagné and Jaschhof, 2017) and Junichi Yukawa in Japan (e.g., unnamed. Yukawa, 1971). Additional data have been published since the 1990s by Wenjun Bu and subsequent authors in China. At the beginning of the 2000s, the Russian researcher Zoya MATERIALS AND METHODS A. Fedotova started publishing on mycophagous cecids from Far Eastern Russia (cf. Gagné and Jaschhof, 2017, for a list of The number of male specimens and species as examined ma- her articles) and authored the Cecidomyiidae chapter in the terials used here are 80 and 37. All samples were collected famed Keys to the Insects of the Russian Far East (Fedotova, using malaise traps in 2019, except for two, which were col- 2004). The latter publication is still the most comprehensive lected in Cheongsong-gun, 2017 by sweep net and Sangju- review available on mycophagous cecid diversity in the Far si, 2018 by malaise trap. Gangwon-do (Mt. Odaesan, Mt. Eastern Palearctic, even though it is of limited suitability for Gariwangsan) and Gyeongsangbuk-do (Cheongsong-gun, species identification (see Discussion). One of us, MJ, studied Sangju-si) were investigated and primarily the virgin forests for a limited time (1998-2000) the taxonomy of Japanese Ca- were examined (Fig. 1). The survey point is briefly marked totrichinae, Lestremiinae, and Micromyinae. However, only with the area code (Table 1). All samples were cleared in Cre- part of the then found 325 different species (Jaschhof, unpub- osote reagent and made slide preparations were made under Anim. Syst. Evol. Divers. 36(1), 60-77 61 Daseul Ham, Mathias Jaschhof, Yeon Jae Bae A B Fig. 1. Survey areas. A, Odaesan National Park, April 2019; B, Gariwangsan Recreational Forest, August 2019. Table 1. Collecting sites in South Korea Locality GPS Method Collector Regional codea Odaesan National Park, Odaesan-ro, 37°47ʹ34.58ʺN, 128°33ʹ37.97ʺE, Malaise trap Daseul Ham, ONP1 Jinbu-myeon, Pyeongchang-gun, alt. 929 m Sunghwan Park Gangwon-do 37°47ʹ55.33ʺN, 128°34ʹ8.24ʺE, Malaise trap Daseul Ham, ONP2 alt. 1,226 m Sunghwan Park 37°47ʹ5.67ʺN, 128°34ʹ16.97ʺE, Malaise trap Daseul Ham, ONP3 alt. 830 m Sunghwan Park Gariwangsan Recreational Forest, 37°25ʹ23.72ʺN, 128°33ʹ23.19ʺE, Malaise trap Daseul Ham, GRF1 Hoehdong-ri, Jeongseon-eup, alt. 462 m Sunghwan Park Jeongseon-gun, Gangwon-do 37°24ʹ35.51ʺN, 128°32ʹ3.10ʺE, Malaise trap Daseul Ham, GRF2 alt. 750 m Sunghwan Park Research center for endangered species 36°38ʹ21.32ʺN, 129°9ʹ9.70ʺE, Malaise trap Honggeun Kim, NIE1 of National Institute of Ecology, Gowol-gil, alt. 300 m Yejin Choi Yeongyang-eup, Yeongyang-gun, 36°38ʹ53.38ʺN, 129°9ʹ18.84ʺE, Malaise trap Honggeun Kim, NIE2 Gyeongsangbuk-do alt. 241 m Yejin Choi 36°38ʹ52.15ʺN, 129°9ʹ13.34ʺE, Malaise trap Honggeun Kim, NIE3 alt. 256 m Yejin Choi Neogu Village, Cheongsong-eup, 36°26ʹ10.89ʺN, 129°9ʹ22.51ʺE, Sweeping net Daseul Ham NV Cheongsong-gun, Gyeongsangbuk-do alt. 454 m Ipseok-ri, Hwabuk-myeon, 36°39ʹ1.79ʺN, 127°52ʹ51.54ʺ, Malaise trap Wanggyu Kim SJ Sangju-si, Gyeongsangbuk-do alt. 249 m aIn the taxonomic accounts section, the investigation areas where samples were collected are abbreviated for brevity. a stereomicroscope (Olympus SZ51, Tokyo, Japan) using referred to here are following Jaschhof (1998b), Jaschhof and Canada balsam (Jaschhof and Jaschhof, 2009; Ham et al., Jaschhof (2009, 2013), and Gagné and Jaschhof (2017) except 2019). Specimens were examined with bright-field and opti- for synonyms. Subfamilies are listed according to the taxo- cal microscopy (Olympus BX50). Microscopic images of the nomic hierarchy (Sikora et al., 2019), and genera and species specimens were taken using a optical microscopy (Olympus are listed alphabetically. Generic diagnoses can be found in BX50), with the aid of a microscope-attached camera (Nikon the books by Jaschhof and Jaschhof (2009, 2013). Species di- Z6, Tokyo, Japan). The terminology of adult morphological agnoses given here are primarily compiled from these books features used in this study follows Jaschhof and Jaschhof (Jaschhof and Jaschhof, 2009, 2013) and original papers, (2009, 2013). The world distribution and history of species which were listed type specimens. Arrows of photographs 62 Anim. Syst. Evol. Divers. 36(1), 60-77 Mycophagous Gall Midges in Korea indicate the diagnostic character. All specimens are deposited Material examined. 1♂, ONP1, 1 Jun-14 Jul 2019, slide in NIBR (National Institute of Biological Resources, Incheon, No. 19C-93 (in NIBR). Korea) or KU (Korean Entomological Institute of Korea Uni- Diagnosis. A typical Aprionus is referred to Jaschhof and versity, Seoul, Korea). Jaschhof (2009: 219). Characters specific to A. heothinos are as follows (Jaschhof and Jaschhof, 2009: 279): Gonostylus without claw, slightly convex basally and tapered. Tegmen SYSTEMATIC ACCOUNTS with 4-5 pairs of thin overlapping fingers. Subanal plate with dark markings along the longitudinal axis and sublaterally. Order Diptera Linnaeus, 1804 Distribution. Finland, Germany, Russia (Western Siberia), Family Cecidomyiidae Newman, 1834 new to South Korea. Subfamily Lestremiinae Rondani, 1840 Genus Anaretella Enderlein, 1911 4. 5*Aprionus laevis Mohrig, 1967 (Fig. 2F) Aprionus laevis Mohrig 1967: 453. 1. 1*Anaretella iola Pritchard, 1951 (Fig. 2A) Anaretella iola Pritchard 1951: 250. Material examined. 2♂♂, ONP1, 1 Jun-14 Jul 2019, slides No. 19C-115 (in NIBR), 19C-13 (in KU). Material examined. 3♂♂, ONP1, May 11-26 2019, slides Diagnosis. A typical Aprionus is referred to Jaschhof and No. 19-2 (in NIBR), 19-7, 10 (in KU). Jaschhof (2009: 219). Characters specific to A. laevis are as Diagnosis. A typical Anaretella is referred to Jaschhof and follows (Jaschhof and Jaschhof, 2009: 268): First segment of Jaschhof (2009: 64). Characters specific to A. iola are as fol- palpus enlarged. Gonostylus convex with sharp spine apical- lows (Jaschhof and Jaschhof, 2009: 66): Apex of gonostylus ly. Tegmen constricted below and beyond the midlength with protruded. Tegmen with two large, horn-shaped sclerotized several delicate folds. Subanal plate absent. Crenulation of processes. male poorly developed. Remarks. This species could be split into several species. Distribution. Northern Europe, Germany, Austria, new to Distribution. USA, Europe, Russia (Western Siberia), New South Korea. to South Korea. 5. 6*Aprionus pseudispar Jaschhof, 1997 (Fig. 2G, H) Subfamily Micromyinae Rondani, 1856 Aprionus pseudispar Jaschhof 1997b: 118. Genus 2*Aprionus Kieffer, 1894 Material examined. 1♂, ONP1, 1 Jun-14 Jul 2019, slide 2. 3*Aprionus adventitius Jaschhof, 2009 (Fig. 2B, C) No. 19C-48 (in NIBR). Aprionus adventitius Jaschhof 2009: 275. Diagnosis. A typical Aprionus is referred to Jaschhof and Jaschhof (2009: 219). According to Jaschhof and Jaschhof Material examined. 2♂♂, ONP1, 1 Jun-14 Jul 2019, slides (2009: 268), A. pseudispar is similar to A. dispar, differing No. 19C-31 (in NIBR), 19C-21 (in KU). from it as follows: Gonogtylus thicker. Tegmen slenderer. An- Diagnosis. A typical Aprionus is referred to Jaschhof and tennal translucent sensilla 2-3 branched. Second, third seg- Jaschhof (2009: 219). Characters specific to A. adventitius are ments of palpus elongated. as follows (Jaschhof and Jaschhof, 2009: 275): Gonostylus Remarks. Korean A. pseudispar has a wing in which apicR1 convex basally, tapered with claw apically. Tegmen has a cap is the same as the length of Rs. However, European A. pseud- apically with 5-6 pairs overlapping fingers. Subanal plate ispar has a wing in which apicR1 is 2-2.5 times the length of curtain-shaped, upper portion dark, lower portion membra- Rs. nous, spread distolaterally. Distribution. Sweden, Germany, new to South Korea. Distribution. Sweden, Finland, Russia (Europe, Far East), New to South Korea. 6. ‌7*‌Aprionus separatus Mamaev & Jaschhof, 1997 ‌ (Fig. 2I, J) 3. 4*Aprionus heothinos Jaschhof, 2009 (Fig. 2D, E) Aprionus separatus Mamaev & Jaschhof 1997: 457. Aprionus heothinos Jaschhof 2009: 279. Material examined. 1♂, ONP1, 1 Jun-14 Jul 2019, slide No. Korean name: 1*쌍뿔둥근날개혹파리 (신칭), 2*고랑애혹파리속 (신칭), 3*덧고랑애혹파리 (신칭), 4*동쪽고랑애혹파리 (신칭), 5*저라고랑애혹파리 (신칭), 6*나도고랑애혹파리 (신칭), 7*가시고랑애혹파리 (신칭) Anim. Syst. Evol. Divers. 36(1), 60-77 63 Daseul Ham, Mathias Jaschhof, Yeon Jae Bae A B C D E F G I J H K Fig. 2. Genitalia (A-G, I-K) and flagellomere (H) of male mycophagous cecidomyiids (Lestremiinae, Micromyinae). A, Anaretella iola; B, C, Aprionus adventitious; D, E, Aprionus heothinos; F, Aprionus laevis; G, H, Aprionus pseudispar; I, J, Aprionus separatus; K, Aprionus spiniger. c, claw of gonostylus; ca, cap; f, fingers in tegmen; g, gonostylus; sp, subanal plate; t, tegmen; ts, translucent sensilla. Scale bars: A-K 0.05 mm. = 64 Anim. Syst. Evol. Divers. 36(1), 60-77 Mycophagous Gall Midges in Korea A B C D Fig. 3. Genitalia (A-D) of male mycophagous cecidomyiids (Micromyinae). A, Bryomyia apsectra; B, Bryomyia bergrothi; C, D, Bryo- myia gibbosa. dmp, dorsomesal projection; ve, ventral emargination; tg9, ninth tergite. Scale bars: A-D 0.05 mm. = 19C-20 (in NIBR). fingers and numerous hair-like fingers below. Subanal plate Diagnosis. A typical Aprionus is referred to Jaschhof and spread laterally with dark longitudinal axis. Jaschhof (2009: 219). Characters specific to A. separatus are Distribution. USA (Maine), widespread Europe, Far East, as follows (Jaschhof and Jaschhof, 2009: 254): Gonocoxites new to South Korea. pointed ventroapically. Gonostylus slightly convex basally with a small spine apically. Tegmen papered with five pairs of Genus 2*Bryomyia Kieffer, 1895 large, overlapping fingers. Antennal sensilla simple. Distribution. Sweden, Finland, Latvia, Germany, Russia (Far 8. 3*Bryomyia apsectra Edwards, 1938 (Fig. 3A) East), new to South Korea. Bryomyia apsectra Edwards 1938a: 210. 7. 1*Aprionus spiniger (Kieffer), 1894 (Fig. 2K) Material examined. 3♂♂, ONP1, 1 Jun-14 Jul 2019, slides Apriona spinigera Kieffer 1894: clxxvi. No. 19C-1 (in NIBR), 19C-7, 61 (in KU). Aprionus spiniger Edwards 1938b: 231. Diagnosis. A typical Bryomyia is referred to Jaschhof and Jaschhof (2009: 281). Characters specific to B. apsectra are as Material examined. 1♂, ONP1, 1 Jun 2019-14 Jul 2019, follows (Jaschhof and Jaschhof, 2009: 182): Gonostylus with- slide No. 19C-25 (in NIBR). out distinctive dorsomesal projection. Apical margin of Ninth Diagnosis. A typical Aprionus is referred to Jaschhof and tergite sinuous with two rounded lobes. Jaschhof (2009: 219). Characters specific to A. spiniger are Distribution. Widespread Palearctic, new to South Korea. as follows (Jaschhof and Jaschhof, 2009: 282): Gonotylus flat, broad, tapered with apical spine, and several subapical 9. 4*Bryomyia bergrothi Kieffer, 1895 (Fig. 3B) bristles. Tegmen has a flat apical cap with 8-9 pairs of large Bryomyia bergrothi Kieffer 1895: 78. Korean name: 1*톱니고랑애혹파리 (신칭), 2*이끼애혹파리속 (신칭), 3*새새이끼애혹파리 (신칭), 4*오대이끼애혹파리 (신칭) Anim. Syst. Evol. Divers. 36(1), 60-77 65 Daseul Ham, Mathias Jaschhof, Yeon Jae Bae Material examined. 1♂, ONP1, 1 Jun-14 Jul 2019, slide No. Material examined. 3♂♂, NIE3, 10-17 Apr 2019, slides 19C-24 (in NIBR). No. 19AY-22, 24, 19Aya-1 (in KU). Diagnosis. A typical Bryomyia is referred to Jaschhof and Diagnosis. A typical Campylomyza is referred to Jaschhof Jaschhof (2009: 281). Characters specific to B. bergrothi are and Jaschhof (2009: 89). Characters specific to C. flavipes as follows (Jaschhof and Jaschhof, 2009: 183): Ninth tergite are as follows (Jaschhof, 1998b: 168): Tegmen with serrate with subrectangular lobes apicolaterally. Gonocoxites with processes on apex. Mesal gonocoxal bridge with two narrow deeply U-shaped ventral emargination. Gonostylus stout with processes. indistinct dorsomesal border. Distribution. Russia (Far East), new to South Korea. Distribution. Widespread Europe, Russia (Western Siberia, Near East), China, new to South Korea. Genus 4*Catocha Haliday, 1833 10. 1*Bryomyia gibbosa (Felt), 1907 (Fig. 3C, D) 13. 5*Catocha latipes Haliday, 1833 (Fig. 4C) Campylomyza gibbosa Felt 1907: 3. Catocha latipes Haliday 1833: 156. Bryomyia gibbosa Pritchard 1947: 69-70, 86-87. Material examined. 1♂, ONP1, 11-26 May 2019, slide No. Material examined. 1♂, ONP1, 1 Jun-14 Jul 2019, slides 19B-18 (in KU). No. 19C-14 (in NIBR), 19C-39 (in KU), 1♂, GRF1, 13 Apr- Diagnosis. A typical Catocha is referred to Jaschhof and 12 May 2019, slide No. 19I-1 (in KU). Jaschhof (2009: 78). Characters specific to C. latipes are as Diagnosis. A typical Bryomyia is referred to Jaschhof and follows (Jaschhof, 1998b: 80): Gonostylus large, thick with Jaschhof (2009: 284). Characters specific to B. gibbosa are as smaller claw than other species belonging to Catocha. Teg- follows (Jaschhof and Jaschhof, 2009: 184): Ninth tergite sep- men narrowly rounded apically. Aedeagus membranous with arated by deep cleft with subrectangular apicolateral lobes. dense microtrichia apically and 10-11 small and large sclero- Gogocoxites shallowly emarginated by rounded, setulose pro- tized spines laterally. jections ventrally. Gonostylus with bare dorsomesal border. Distribution. Widespread Holarctic, new to South Korea. Loops of copulatory organ distinctive. Distribution. Widespread Holarctic, new to South Korea. Genus 6*Heterogenella Mamaev, 1963 Genus Campylomyza Meigen, 1919 14. 7*Heterogenella cambrica (Edwards), 1938 (Fig. 4D) Bryomyia cambrica Edwards 1938a: 210. 11. 2*Campylomyza flavipes Meigen, 1818 (Fig. 4A) Heterogenella cambrica Jaschhof 1998b: 223. Campylomyza flavipes Meigen 1818: 102. Material examined. 1♂, ONP1, 11-26 May 2019, slide No. Material examined. 2♂♂, ONP2, 11-26 May 2019, slides 19B-6 (in NIBR). No. 19A-3,7 (in KU), 1♂♂, NIE2, 13-20 Mar 2019, slide Diagnosis. A typical Heterogenella is referred to Jaschhof No. 19AU-3 (in KU), 6♂♂, NIE3, 10-17 Apr 2019, slides and Jaschhof (2009: 187). Characters specific to H. cambrica No. 19AY-3, 16, 18-20, 25, 19Aya-4 (in KU), 5♂♂, NERC 3, are as follows (Jaschhof and Jaschhof, 2009: 188): Palpus 4. 3-10 Apr 2019, slides No. 19AZ-1-5 (in KU). Projections on the ventral emargination of gonocoxites ab- Diagnosis. A typical Campylomyza is referred to Jaschhof and sent. Gonostylus straightened, elongate, slightly excavated Jaschhof (2009: 89). Characters specific to C. flavipes are as dorsomesally. Antennal translucent sensilla 2-3 furcate. follows (Jaschhof and Jaschhof, 2009: 106): Tegmen points Distribution. Widespread Europe, China (Sichuan), new to lamellate; dorsal processes leaf-shaped directed to bottom lat- South Korea. erally. Distribution. Widespread Holarctic, New Zealand, new to Genus 8*Monardia Kieffer, 1895 South Korea. 15. 9*Monardia (Xylopriona) atra (Meigen), 1804 (Fig. 12. 3*Campylomyza spinata Jaschhof, 1998 (Fig. 4B) 4E, G) Campylomyza spinata Jaschhof 1998a: 264. Cecidomyia atra Meigen 1804: 40. Korean name: 1*이끼애혹파리 (신칭), 2*옷깃애혹파리 (신칭), 3*거스러미애혹파리 (신칭), 4*다발애혹파리속 (신칭), 5*다발애혹파리 (신칭), 6*앙증애혹파리속 (신칭), 7*앙증애혹파리 (신칭), 8*왕애혹파리속 (신칭), 9*큰왕애혹파리 (신칭) 66 Anim. Syst. Evol. Divers. 36(1), 60-77 Mycophagous Gall Midges in Korea A B C D E Fig. 4. Genitalia (A-F) and flagellomere (G) of male mycophagous cecidomyiids (Micromyinae). A, Campylomyza fla- vipes; B, Campylomyza spinata; C, Catocha latipes; D, Heterogenella cambrica; E, G, Monardia (Xylopriona) atra; F, Monardia (Xylopriona) furcifera. ad, aedeagus; c, claw of gonostylus; dp, dorsal processes of tegmen; ea, ejaculatory apodeme; g, gonostylus; p, processes of mesal gonocoxal bridge; t, F G tegmen; ts, translucent sensilla. Scale bars: A-G 0.05 mm. = Anim. Syst. Evol. Divers. 36(1), 60-77 67 Daseul Ham, Mathias Jaschhof, Yeon Jae Bae Monardia (Xylopriona) atra Jaschhof 1998b: 301. Latvia, Germany, Kirghizstan, new to South Korea. Material examined. 1♂, ONP2, 11-26 May 2019, slide No. 18. 3*Peromyia boreophila Jaschhof, 2001 (Fig. 5B) 19A-8 (in NIBR), 2♂♂, ONP1, 11-26 May 2019, slides No. Peromyia boreophila Jaschhof 2001: 107. 19B-8, 20 (in KU). Diagnosis. A typical Monardia is referred to Jaschhof and Material examined. 6♂♂, ONP1, 1 Jun-14 Jul 2019, slides Jaschhof (2009: 195). Characters specific to M. atra are as No. 19C-12 (in NIBR), 19C-18, 30, 43, 58, 65 (in KU). follows (Jaschhof and Jaschhof, 2009: 210): Robust, black Diagnosis. A typical Peromyia is referred to Jaschhof and color. Body size 2-3 mm long in male. Palpus 4. Flagel- Jaschhof (2009: 140). Characters specific to P. boreophila lomere nodes shorter than necks with leaf-shaped translucent are as follows (Jaschhof and Jaschhof, 2009: 161): Postcular sensilla. Gonotylus large with small apical spine. Tegmen bristle one long row irregular posterior row. Flagellomeres narrowly rounded apically, weakly sclerotized. with one mesal and one distal whorl of hair-shaped sensilla. Distribution. USA (California), widespread Palearctic, new Gonostylus long, tapered. Tegmen parallel-sided medially to South Korea. and rounded apically; Ventral plate clear, horseshoe-shaped. Distribution. Sweden, Finland, Russia (Europe), Japan (Hok- 16. 1*Monardia (Xylopriona) furcifera Mamaev, 1963 kaido, Honshu), new to South Korea. (Fig. 4F) Monardia furcifera Mamaev 1963a: 444. 19. 4*Peromyia brevispina Yukawa, 1967 (Fig. 5C, D) Monardia (Xylopriona) furcifera Jaschhof 1998b: 305. Peromyia brevispina Yukawa 1967: 188. Peromyia fungicola Yukawa 1971: 26. Material examined. 1♂, ONP1, 11-26 May 2019, slide No. Peromyia brevispina Jaschhof 2016b: 87. 19B-4 (in NIBR). Diagnosis. A typical Monardia is referred to Jaschhof and Material examined. 1♂, ONP1, 1 Jun-14 Jul 2019, slides Jaschhof (2009: 195). Characters specific to M. furcifera are No. 19C-88 (in NIBR). as follows (Jaschhof and Jaschhof, 2009: 211): Palpus 4. Diagnosis. A typical Peromyia is referred to Jaschhof and Neck of flagellomeres slightly shorter than nodes. Gonogtylus Jaschhof (2009: 140). According to Jaschhof (2016b: 88), P. slightly tapered with a tiny dorsosubapical spine. Ejaculatory brevispina is mostly similar to P. fungicola and distinguished apodeme large, broad, and furcated apically. by the characters: Gonostylus slenderer with a shorter, spine- Distribution. Widespread Europe, Russia (Far East), new to shaped claw apically. On gonocoxites, ventroposterior lobe South Korea. sma ll er; membranous, asetose area of ventral gonocoxal bridge more extensive. Tegmen smaller. Ninth tergite with Genus Peromyia Kieffer, 1894 gentle angles. Distribution. Sweden, Finland, Germany, Russia (Karelia, 17. 2*Peromyia borealis (Felt), 1920 (Fig. 5A) Far East), Japan, new to South Korea. Joannisia borealis Felt, 1920: 280. Peromyia borealis Jaschhof 1998b: 444. 20. 5*Peromyia curta Jaschhof, 1997 (Fig. 5E) Peromyia curta Jaschhof 1997a: 53. Material examined. 1♂, NIE3, 24 Apr-2 May 2019, slide No. 19BD-3 (in NIBR). Material examined. 1♂, GRF2, 12 May-21 Jun 2019, slide Diagnosis. A typical Peromyia is referred to Jaschhof and No. 19K-6. Jaschhof (2009: 140). Characters specific to P. borealis are Diagnosis. A typical Peromyia is referred to Jaschhof and as follows (Jaschhof and Jaschhof, 2009: 149): Gonocoxites Jaschhof (2009: 140). Characters specific to P. curta are as short; Ventrobasal portion asetose; Ventral gonocoxal bride follows (Jaschhof and Jaschhof, 2009: 157): Flagellomeres thick. Gonostylus stout rounded apically with microtrichia with appendiculate sensilla. Gonostylus flat, stout, tapered densely. Tegmen large, rounded apically with membranous toward apex. Tegmen parallel-sided with narrowly rounded margin apically. Flagellomeres with one distal and one mesal apical point; ventral plate visible only apical margin. whorl of hair-shaped sensilla, both irregular. Distribution. Widespread Europe, Russia (Near East), Japan Distribution. USA (Minnesota, New York), UK, Sweden, (widespread), new to South Korea. Korean name: 1*작은왕애혹파리 (신칭), 2*영양어리애혹파리 (신칭), 3*북어리애혹파리 (신칭), 4*난장이애혹파리 (신칭), 5*작은어리애혹파리 (신칭) 68 Anim. Syst. Evol. Divers. 36(1), 60-77 Mycophagous Gall Midges in Korea A B C D E F G H I Fig. 5. Genitalia of male mycophagous cecidomyiids (Micromyinae). A, Peromyia borealis; B, Peromyia boreophila; C, D, Peromyia brevispina; E, Peromyia curta; F, Peromyia fagiphila; G, Peromyia gemella; H, I, Peromyia imperatoria. c, claw of gonostylus; g, gonostylus; t, tegmen; tg9, ninth tergite; vgb, ventral gonocoxal bridge; vp, ventral plate. Scale bars: A-I 0.05 mm. = 21. 1*Peromyia fagiphila Jaschhof, 1997 (Fig. 5F) Jaschhof (2009: 140). Characters specific to P. fagiphila are Peromyia fagiphila Jaschhof 1997a: 36. as follows (Jaschhof and Jaschhof, 2009: 152): Flagellomeres with one mesal and one distal whorl of hair-shaped sensil- Material examined. 4♂♂, ONP1, 1 Jun-14 Jul 2019, slides la. Gonocoxites long. Gonostylus long, cylindrical. Tegmen No. 19C-4, 46, 54, 68, 1♂, GRF2, 12 May-21 Jun 2019, large, tapered, ventral plate long. slide No. 19K-14. Remarks. This species could be split into several species. Diagnosis. A typical Peromyia is referred to Jaschhof and Distribution. Northern Europe, Germany, Austria, Russia Korean name: 1*곰팡이애혹파리 (신칭) Anim. Syst. Evol. Divers. 36(1), 60-77 69

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