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: ) 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: Cecidomyiidae‌ species new to Korea, taxonomic diagnosis, gall-midge , 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INTRODUCTION Sweden (Jaschhof, unpublished data).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 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 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 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* 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, tegmen; ts, translucent sensilla. Scale F G 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. small­ 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 Daseul Ham, Mathias Jaschhof, Yeon Jae Bae

(Western Siberia, Far East), Japan, new to South Korea. as follows (Jaschhof, 2001: 113): Flagellomere nodes with long and short setae basally, 1 row of long sensory whorl 22. 1*Peromyia gemella Jaschhof, 2001 (Fig. 5G) medially, several sensory peaks distally. Gonocoxites with Peromyia gemella Jaschhof 2001: 84. U-shaped ventral emargination. Gonostylus cylindrical, ta- pered with long setae distally, claw absent. Tegmen tapered, Material examined. 2♂♂, ONP1, 1 Jun-14 Jul 2019, slides ventral plates distinct. No. 19C-94 (in NIBR), 19C-119 (in KU). Distribution. Japan, new to South Korea. Diagnosis. A typical Peromyia is referred to Jaschhof and Jaschhof (2009: 140). Characters specific to P. gemella are as 26. 5*Peromyia neomexicana (Felt), 1913 (Fig. 6C) follows (Jaschhof, 2001: 84): Flagellomeres with appendicu- Joannisia neomexicana Felt 1913: 160. late sensory whorl. Gonostylus long, slightly cylindrical with Peromyia neomexicana Pritchard 1947: 79-80. brush of long dense hairs apically. Tegmen elongated, slender, weakly sclerotized with distinct ventral plate. Material examined. 1♂, NIE1, 24 Apr-5 May 2019, slide Distribution. Japan, new to South Korea. No. 19BA-3 (in KU). Diagnosis. A typical Peromyia is referred to Jaschhof and 23. 2*Peromyia imperatoria Jaschhof, 2001 (Fig. 5H, I) Jaschhof (2009: 140). Characters specific to P. neomexicana Peromyia imperatoria Jaschhof 2001: 51. are as follows (Jaschhof, 2001: 53; Jaschhof, 1998b: 441): Outline of terminalia circular ventrally. Gonotylus tapered Material examined. 1♂, NIE1, 24 Apr-5 May, slide No. with long brush-shaped claw apically. Tegmen peaked, round- 19BA-2 (in KU). ed apically. Ventral plate weakly sclerotized. Diagnosis. A typical Peromyia is referred to Jaschhof and Distribution. USA (New Mexico), Germany, Ukraine, Russia Jaschhof (2009: 140). Characters specific to P. imperatoria (Europe, Far East), Japan, new to South Korea. are as follows (Jaschhof, 2001: 84): Gonocoxites long with U-shaped ventral emargination. Ventral gonocoxal bridge 27. 6*Peromyia photophila (Felt), 1907 (Fig. 6D) thick, heavy. Tegmen heavy, blunt apically with short, scle- Campylomyza photophila Felt 1907: 3. rotized ventral plate. Gonostylus cylindrical, rounded with Peromyia photophila Pritchard 1947: 76-78. brush hair apically. Distribution. Japan, new to South Korea. Material examined. 1♂, ONP1, 11-26 2019, slide No. 19B- 5 (in KU). 24. 3*Peromyia maetoi Jaschhof, 2001 (Fig. 6A) Diagnosis. A typical Peromyia is referred to Jaschhof and Peromyia maetoi Jaschhof 2001: 58. Jaschhof (2009: 140). Characters specific to P. photophila are as follows (Jaschhof, 2001: 129; Jaschhof, 1998b: 481): Material examined. 2♂♂, ONP1, 1 Jun-14 Jul 2019, slides Gonocoxites subrectangular basally with narrowly shallowed No. 19C-16 (in NIBR), 19C-112 (in KU). ventral emargination. Gonostylus slender, narrow with sharp Diagnosis. A typical Peromyia is referred to Jaschhof and claw apically. Tegmen brunt, ventral plate darker, smaller Jaschhof (2009: 140). Characters specific to P. maetoi are as than tegmen, rounded apically. follows (Jaschhof, 2001: 58): Gonocoxites with slightly shal- Distribution. USA (widespread eastern), widespread Europe, low ventral emargination. Gonostylus long, slender. Tegmen Russia, Japan, new to South Korea. elongated, parallel-sided with darkened bar inside. Distribution. Japan, new to South Korea. 28. 7*Peromyia trifida Jaschhof, 2001 (Fig. 6E-G) Peromyia trifida Jaschhof 2001: 102. 25. 4*Peromyia montivaga Jaschhof, 2001 (Fig. 6B) Peromyia montivaga Jaschhof 2001: 113. Material examined. 1♂, NIE3, 10-17 Apr 2019, slide No. 19AY-1, (in NIBR), 1♂, NIE3, 7 24 Apr-May 2019, slide Material examined. 1♂, ONP1, 1 Jun-14 Jul 2019, slide No. No. 19BD-2 (in KU). 19C-9 (in NIBR). Diagnosis. A typical Peromyia is referred to Jaschhof and Diagnosis. A typical Peromyia is referred to Jaschhof and Jaschhof (2009: 140). Characters specific to P. trifida are as Jaschhof (2009: 140). Characters specific to P. montivaga are follows (Jaschhof, 2001: 102): Gonocoxites with U-shaped

Korean name: ‌1*쌍둥이애혹파리 (신칭), 2*고월어리애혹파리 (신칭), 3*얇은어리애혹파리 (신칭), 4*앙증어리애혹파리 (신칭), 5*영양애혹파리 (신칭), 6*뾰족어리애혹파리 (신칭), 7*장갑어리애혹파리 (신칭)

70 Anim. Syst. Evol. Divers. 36(1), 60-77 Mycophagous Gall Midges in Korea

A B C

D E F

G H I

Fig. 6. Genitalia of male mycophagous cecidomyiids (Micromyinae). A, Peromyia maetoi, B, Peromyia montivaga; C, Peromyia neomexicana; D, Peromyia photophila; E-G, Peromyia trifida; H, Peromyia valens; I, Polyardis bispinosa. c, claw of gonostylus; g, gonostylus; gr, genital rod; gx, gonocoxites; t, tegmen; ve, ventral emargination; vp, ventral plate. Scale bars: A-I=0.05 mm. ventral emargination, membranous. Gonostylus with three Peromyia valens Jaschhof 2001: 145. lobes; medial lobe directed upward, cylindrical without claw; inner lobe short with dense hairs; dorsal lobe small with long Material examined. 1♂, ONP1, 1 Jun-14 Jul 2019, slide setae. Tegmen as long as length of gonocoxies, slightly peaked No. 19C-19 (in NIBR). apically; ventral plate smaller than tegmen, rounded apically. Diagnosis. A typical Peromyia is referred to Jaschhof and Distribution. Japan, new to South Korea. Jaschhof (2009: 140). Characters specific to P. valens are as follows (Jaschhof, 2001: 145): Gonocoxites shallow emargi- 29. 1*Peromyia valens Jaschhof, 2001 (Fig. 6H) nated ventrally with long gonocoxal ventral bridge. Gonosty-

Korean name: 1*센어리애혹파리 (신칭)

Anim. Syst. Evol. Divers. 36(1), 60-77 71 Daseul Ham, Mathias Jaschhof, Yeon Jae Bae

A B C

D E F

G H I

Fig. 7. Genitalia of male mycophagous cecidomyiids (Winnertziinae, Porricondylinae). A-C, Japoplusia honshuensis; D, Winnertzia discretella; E, Winnertzia globifera; F, Winnertzia pravdini; G, Cassidoides fulvus; H, Spungisomyia aberrans; I, Spungisomyia me- dia. ad, aedeagus; c, claw of gonostylus; ea, ejaculatory apodeme; g, gonostylus; gp, gonocoxal processes; p, parameres; t, teg- men; ve, ventral emargination; vgb, ventral gonocoxal bridge. Scale bars: A-I=0.05 mm. lus slender, tapered, covered by microtrichia. Tegmen large, Monardia bispinosa Mamaev 1963a: 443. widest in distal half. Polyardis bispinosa Jaschhof 1998b: 314. Distribution. Japan, new to South Korea. Material examined. 1♂, ONP1, 11-26 May 2019, slide No. Genus 1*Polyardis Pritchard, 1947 19B-19 (in KU). Diagnosis. A typical Polyardis is referred to Jaschhof and 30. 2*Polyardis bispinosa (Mamaev), 1963 (Fig. 6I) Jaschhof (2009: 215). Characters specific to P. bispinosa are

Korean name: 1*수풀애혹파리속 (신칭), 2*뾰족수풀애혹파리 (신칭)

72 Anim. Syst. Evol. Divers. 36(1), 60-77 Mycophagous Gall Midges in Korea

as follows Jaschhof and Jaschhof (2009: 217): Flagellomeres NIBRIN0000857608 (HDS-694) (in NIBR). 12. Gonocoxites with deep V-shaped ventral emargination. Diagnosis. A typical Winnertzia is referred to Jaschhof and Gonostylus slender, tapered, slightly curved inwardly with Jaschhof (2013: 74). Characters specific to W. discretella are thin spine apically. Tegmen peaked apically. Genital rod as follows (Jaschhof and Jaschhof, 2013: 77): Gonocoxites straightened. with extended posterior portion dorsally. Ninth tergite slightly Distribution. Widespread Europe, Russia (Near East, Far sinuous emargination. Gonostylus with brushed claw inward- East), new to South Korea. ly in distal half. Distribution. Widespread Europe, Russia (Far East), new to Subfamily Winnertziinae Panelius, 1965 South Korea. Genus 1*Japoplusia Jaschhof, 2016 34. 5*Winnertzia pravdini Mamaeva & Mamaev, 1971 ‌ 31. 2*Japoplusia honshuensis Jaschhof, 2016 (Fig. 7F) (Fig. 7A-C) Winnertzia pravdini Mamaeva & Mamaev 1971: 9. Japoplusia honshuensis Jaschhof 2016a: 225. Material examined. 1♂, ONP1, 1 Jun-14 Jul 14 2019, slide Material examined. 1♂, SJ, 26 Jul 2018, slide No. NIBRIN No. 19C-109 (in NIBR). 0000857602 (HDS-631) (in NIBR). Diagnosis. A typical Winnertzia is referred to Jaschhof and Diagnosis. Characters specific to J. honshuensis are as fol- Jaschhof (2013: 74). Characters specific to W. pravdini are as lows (Jaschhof, 2016a: 224): Flagellomeres 12; node with 1 follows (Jaschhof and Jaschhof, 2013: 102): Anepimeron se- row crenulate whorls medially, dense hair-like sensilla ba- tose. AntGA long, thin. Gonostylus with long pectinated claw sally, and short hair-like sensilla distally. Gonocoxal ventral subapically. Tegmen slightly sclerotized laterally. emargination V-shaped deeply with angled lobe ventroposte- Distribution. Sweden, Estonia, Latvia, Ukraine, Near East rially. Gonostylus concaved medially with pectinate tooth on (Caucasus Mts.), new to South Korea. protruded boundary of apical margin. Parameres long, thin, separated each, directed dorsally. Subfamily Porricondylinae Kieffer, 1913 Remarks. Genus Japoplusia is a monotypic taxon. Genus 6*Cassidoides Mamaev, 1960 Distribution. Japan, new to South Korea. 35. 7*Cassidoides fulvus (Kieffer), 1896 (Fig. 7G) Genus Winnertzia Rondani, 1860 Holoneurus fulvus Kieffer 1896: 18. Cassidoides fulvus Jaschhof & Jaschhof 2013: 300. 32. 3*Winnertzia discretella Spungis, 1992 (Fig. 7D) Winnertzia discretella Spungis 1992: 19. Material examined. 1♂, ONP1, 1 Jun-14 Jul 2019, slide No. 19C-64 (in NIBR). Material examined. 1♂, ONP1, 11-26 May 2019, slide Diagnosis. A typical Cassidoides is referred to Jaschhof and No. 19B-17 (in NIBR). Jaschhof (2013: 297). Characters specific to C. fulvus are Diagnosis. A typical Winnertzia is referred to Jaschhof and as follows (Jaschhof and Jaschhof, 2013: 300): Ejaculatory Jaschhof (2013: 74). Characters specific to W. discretella are apodeme with sclerotized cap apically. Gonocoxal processes as follows (Jaschhof and Jaschhof, 2013: 80): Gonocoxites curved ectad. Gonostylus compact with pectinated claw. with U-shaped ventral emargination. AntGA long beyond the Distribution. North Carolina, France, Russia, new to South gonocoxal ventral bridge. Ninth tergite sinuous. Gonostylus Korea. rounded apically with dense, broad microtrichia row on tip. Distribution. Sweden, Latvia, Russia (Europe, Far East), Genus 8*Spungisomyia Mamaev & Zaitzev, 1996 new to South Korea. 36. 9*Spungisomyia aberrans Jaschhof, 2013 (Fig. 7H) 33. 4*Winnertzia globifera Mamaev, 1963 (Fig. 7E) Spungisomyia aberrans Jaschhof 2013: 259. Winnertzia globifera Mamaev 1963b: 566. Material examined. 1♂, ONP1, 1 Jun-14 Jul 2019, slide Material examined. 1♂, NV, 2017. 17 May 2017, slide No. No. 19C-82 (in NIBR).

Korean name: ‌1*한일타슬혹파리속 (신칭), 2*한일타슬혹파리 (신칭), 3*앙증타슬혹파리 (신칭), 4*콧수염타슬혹파리 (신칭), 5*수염타슬혹파리 (신칭), 6*동글곧은혹파리속 (신칭), 7*동글곧은혹파리 (신칭), 8*수염곧은혹파리속 (신칭), 9*둥근수염곧은혹파리 (신칭)

Anim. Syst. Evol. Divers. 36(1), 60-77 73 Daseul Ham, Mathias Jaschhof, Yeon Jae Bae

Diagnosis. A typical Spungisomyia is referred to Jaschhof including exchange of experiences and specimens, and dia- and Jaschhof (2013: 258). Characters specific to S. aberrans logue about results and problems, is hardly possible (because are as follows (Jaschhof and Jaschhof, 2013: 260): Prescute- of the lack of fellow students). As a third reason, the sheer llar window present. Gonocoxites with U-shaped ventral diversity of genera and species can both encourage and dis- emargination. Gonostylus subglobular. Tegmen narrowed to courage any scholar, especially a beginner, which is why it is apex. Aedeagus parallel-sided, rounded apically, longer than not advisable to try dealing with all five subfamilies at once. tegmen. Moreover, the amount of unnamed taxa, which is usually Distribution. Sweden, new to South Korea. underestimated by the non-specialist, can evoke excitement as well as frustration. Regional faunas that have never been 37. 1*Spungisomyia media (Spungis), 1981 (Fig. 7I) studied before, especially those outside the Palearctic region, Porricondyla media Spungis 1981: 54. usually comprise a large part of unnamed species (including Spungisomyia media Mamaev & Zaitzev 1996: 209. many that cannot be placed in existing genera); this raises the question of where to obtain the resources to describe all Material examined. 1♂, ONP2, 11-26 May 2019, slide No. those novelties. Not all institutions that fund taxonomic re- 19-61 (in NIBR). search are excited about such challenges, and some might Diagnosis. A typical Spungisomyia is referred to Jaschhof and tend to regard groups such as mycophagous Cecidomyiidae Jaschhof (2013: 258). Characters specific to S. media are as as bottomless pits. follows (Jaschhof and Jaschhof, 2013: 263): Scutal windows Considering all the information available on the world’s absent. Parameres slightly split apically. Aedeagus membra- mycophagous cecids, we conservatively estimate the exis- nous apically. Gonocoxal processes small. Gonostylus sub- tence of approximately 1,000 species in Korea, which means globular. we have discovered 7% of the expected total, or 20% if we Distribution. Sweden, Finland, Latvia, Russia (Far East), include the unnamed species identified to date. The Swedish new to South Korea. fauna, on the other hand, is currently known to contain slight- ly over 700 species (including 140 unnamed species), whose discovery took 11 years of full-time employment of one expe- DISCUSSION rienced research taxonomist and one skilled technician. Addi- tions to the Swedish fauna are continuously discovered with At any time in the history of entomo-taxonomy, there were ongoing study, with species new to science by far outnumber- only few scholars dedicated to the study of mycophagous ing new faunistic records, and most of the species being rare Cecidomyiidae. There are many reasons for mycophagous or rarely collected. cecidomyiids being shunned by most professionals, even So, what are the essential prerequisites for a comprehensive those that are primarily dipterists, taxonomists, or ecolo- taxonomic inventory of the mycophagous Cecidomyiidae of gists. The first reason is the unattractiveness of the study ob- Korea? First of all, one must be aware that such an enterprise jects: both larvae and adults are very small in size, with no is a long-term effort that must be carefully designed regarding eye-catching macroscopic feature, and inconspicuous in their the costs involved and the results generated; a source of fund- habits. The fact is that many dipterists are not even aware ing that is both predictable and flexible enough to adjust to that the midges in question exist at all. Another reason is that the magnitude of the task is thus required. Furthermore, dedi- students attempting to learn more about mycophagous cecids cated personnel prepared to tackle a long-term challenge and are inevitably faced with considerable practical problems, that is either experienced with the taxa in question or is given such as: skills and equipment (including the high cost of the chance to develop such experience during the course of Malaise traps) are needed to collect specimens for studying the project is required. One of us, MJ, estimates that he need- them effectively; specimens have to be slide-mounted before ed more than five years to feel prepared to appropriately solve their morphology can be examined for species identification all types of taxonomic problems occurring in Micromyinae (which requires preparation skills plus laboratory facilities (which is only one of five mycophagous subfamilies). Dipte- prepared for work with dangerous chemicals); the positive ro-taxonomists agree that the midges in question are excep- identification of specimens requires access to a large body tionally demanding when it comes to species identification of literature (written in several different languages, including and classification based on morphology. Russian and Chinese) and a well-stocked reference collec- What should motivate us to tackle the described research? tion (only very few of which exist in the world); teamwork, Firstly, there are only few subgroups in Diptera with such a

Korean name: 1*수염곧은혹파리 (신칭)

74 Anim. Syst. Evol. Divers. 36(1), 60-77 Mycophagous Gall Midges in Korea potential of new discovery, even in the Palearctic region. Of for providing the samples of Malaise traps. 560 species discovered in Sweden, a country with an excep- tionally long taxonomic tradition, nearly 200 species were REFERENCES described as being new to science (with other 140 new spe- cies awaiting description). Secondly, there is an economic Borkent A, Brown BV, Adler PH, Amorim DS, Barber K, Bickel demand for thorough taxonomic knowledge of mycophagous D, Boucher S, Brooks SE, Burger J, Burington ZL, Capellari Cecidomyiidae in the region. Issues with mushroom pests, RS, Costa DNR, Cumming JM, Curler G, Dick CW, Epler such as species of Mycophila and Camptomyia, as well as JH, Fisher E, Gaimari SD, Gelhaus J, Grimaldi DA, Hash J, nuisance species, such as Asynapta groverae Jiang & Bu (cf. Hauser M, Hippa H, Ibáñez-Bernal S, Jaschhof M, Kamene- Ham et al., 2018), can only be solved in collaboration with va EP, Kerr PH, Korneyev V, Korytkowski CA, Kung GA, skilled taxonomists. Thirdly, taxonomists specialized in the Kvifte GM, Lonsdale O, Marshall SA, Mathis W, Michelsen field of mycophagous cecids are scarce, so the development V, Naglis S, Norrbom AL, Paiero S, Pape T, Pereira-Colavite of young talents capable of carrying our collective knowl- A, Pollet M, Rochefort S, Rung A, Runyon JB, Savage J, Sil- edge further into the future is crucial. The Swedish Taxon- va VC, Sinclair BJ, Skevington JH, Stireman JO III, Swann omy Initiative funds researches on mycophagous cecids in J, Vilkamaa P, Wheeler T, Whitworth T, Wong M, Wood Sweden and is explicitly concerned about safeguarding the DM, Woodley N, Yau T, Zavortink TJ, Zumbado MA, 2018. Remarkable (Diptera) diversity in a patch of Costa Ri- threatened taxonomic expertise, this being even stated in can cloud forest: why inventory is a vital science. Zootaxa, their funding criteria. Fourthly and finally, thorough knowl- 4402:53-90. https://doi.org/10.11646/zootaxa.4402.1.3 edge of the species in Korea will help to solve taxonomic Brown BV, Borkent A, Adler PH, Amorim DS, Barber K, Bickel problems pertaining to the entire region. Many species de- D, Boucher S, Brooks SE, Burger J, Burington ZL, Capellari scribed from the Russian Far East and China are so poorly RS, Costa DNR, Cumming JM, Curler G, Dick CW, Epler documented in the literature that fresh specimens are re- JH, Fisher E, Gaimari SD, Gelhaus J, Grimaldi DA, Hash J, quired to clarify their identity, and these species are likely to Hauser M, Hippa H, Ibáñez-Bernal S, Jaschhof M, Kamene- be obtained in Korea. Altogether, Korea is now in a position va EP, Kerr PH, Korneyev V, Korytkowski CA, Kung GA, to become a significant player in the taxonomy of mycopha- Kvifte GM, Lonsdale O, Marshall SA, Mathis W, Michelsen gous Cecidomyiidae. V, Naglis S, Norrbom AL, Paiero S, Pape T, Pereira-Colavite A, Pollet M, Rochefort S, Rung A, Runyon JB, Savage J, Sil- va VC, Sinclair BJ, Skevington JH, Stireman JO III, Swann J, ORCID Thompson FC, Vilkamaa P, Wheeler T, Whitworth T, Wong M, Wood DM, Woodley N, Yau T, Zavortink TJ, Zumbado

Daseul Ham: https://orcid.org/0000-0002-1900-3637 MA, 2018. Comprehensive inventory of true (Diptera) Yeon Jae Bae: https://orcid.org/0000-0001-7810-5409 at a tropical site. Communications Biology, 1:21. https://doi. org/10.1038/s42003-018-0022-x Edwards FW, 1938a. On the British Lestremiinae, with notes CONFLICTS OF INTEREST on exotic species. 5. (Diptera, Cecidomyiidae). Proceedings of the Royal Entomological Society of London, 7:199-210. No potential conflict of interest relevant to this article was https://doi.org/10.1111/j.1365-3113.1938.tb01229.x reported. Edwards FW, 1938b. On the British Lestremiinae, with notes on exotic species. 6. (Diptera, Cecidomyiidae). Proceedings of the Royal Entomological Society of London, 7:229-243. ACKNOWLEDGMENTS https://doi.org/10.1111/j.1365-3113.1938.tb01232.x Fedotova ZA, 2004. Supplement. 22. Fam. Cecidomyiidae - gall This study was supported by a grant from the National In- midges. In: Keys to the insects of the Russian Far East in six

volumes. VI. Diptera and Siphonaptera. Part 3 (Ed., Ler PA). stitute of Biological Resources (NIBR), with funds from the Dal’nauka, Vladivostok, pp. 565-629 (in Russian). Ministry of Environment of the Republic of Korea (grant no. Felt EP, 1907. New species of Cecidomyiidae. New York State NIBR201902205). We sincerely thank Sunghwan Park (Korea Education Department, Albany, NY, pp. 1-53. University), Wanggyu Kim (Korea University), for helping Felt EP, 1913. Appendix: a study of gall midges. In: His 28th re- to collect specimens and Catrin Jaschhof (Station Linné) for port of the State Entomologist on injurious and other insects assisting in arrange the samples. The Korea National Park of the State of New York 1912. Bulletin of the New York

Service (KNPS) is thanked for permitting to collect insects at State Museum, 165:5-265. Odaesan National Park, and the National Institute of Ecolo- Felt EP, 1920. New gall midges or Itonididae from the Adiron- gy_ Research center for endangered species (NIE) is thanked dacks. Journal of the New York Entomological Society,

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