Diversity underfoot of agromyzids (Agromyzidae, Diptera) mining thalli of liverworts and hornworts Author Kato, Makoto kato@zoo.zool.kyoto-u.ac.jp Author Yamamori, Luna https://orcid.org/0000-0002-5342-1277 Author Imada, Yume https://orcid.org/0000-0003-2173-7389 text ZooKeys 2022 2022-11-30 1133 1 164 http://dx.doi.org/10.3897/zookeys.1133.94530 journal article http://dx.doi.org/10.3897/zookeys.1133.94530 1313-2970-1133-1 D7A37FE0DC2A4ECCA6A10E873C7C7A5A 4C6A7FCDB0B55088AED4D861AF5BE503 4. Phytoliriomyza marchantiae Kato sp. nov. Figs 6 , 7 Material examined. Holotype : Japan: 1♂ (MK-AG-a320), Namari-kawa, Yakumo, Futami, Hokkaido ( 42.187765°N , 140.122182°E , 190 m asl), 2-VI-2021 (as larva on Marchantia paleacea paleacea ), emerged on 17-VI-2021, NSMT-I-Dip 31903. Paratypes : Japan: 1♂2♀ (MK-AG-a484, a485, a442), same data as holotype, emerged on 18-VI-3-VII-2021, NSMT-I-Dip 31904-31906; 1♂ (MK-AG-a322), Renge-onsen, Itoigawa, Niigata Pref., 11-VII-2021 (as larva on M. p. paleacea ), emerged on 5-VIII-2021, NSMT-I-Dip 31907; 1♂ (MK-AG-a321), Nippara, Okutama, Tokyo Pref., 27-III-2021 (as larva on M. p. paleacea ), emerged on 27-V-2021, NSMT-I-Dip 31908; 1♀ (MK-AG-302), Umegashima, Aoi-ku, Shizuoka Pref., 5-I-2014 (as larva on M. p. diptera ), emerged on?-V-2014, NSMT-I-Dip 31909; 1♂ (MK-AG-a356), Inago, Shibakawa, Fujinomiya, Shizuoka Pref., 20-III-2000 (as larva on M. p. diptera ), emerged on 17-IV-2000, NSMT-I-Dip 31910; 1♂ (MK-AG-a355), Muramatsu, Iwakura, Sakyo-ku, Kyoto Pref., 11-VI-2021 (as larva on M. polymorpha ), emerged on 21-VII-2021, NSMT-I-Dip 31911; 1♂ (MK-AG-a358), Naiku, Oe, Fukuchiyama, Kyoto Pref., 19-V-2021 (as larva on M. papillata grossibarba ), emerged on 3-VII-2021, NSMT-I-Dip 31912. Other material. Japan: On Marchantia paleacea paleacea : 22♂16♀, Namari-kawa, Yakumo, Futami, Hokkaido, 2-VI-2021 (as larva), emerged on 9-VI-7-VII-2021; 38♂44♀, Renge-onsen, Itoigawa, Niigata Pref., I-VII-2021 (as larva), emerged on 29-VII-9-VIII-2021; 2♂1♀, Narahara, Ueno, Tano, Gunnma Pref., 18-IV-2021 (as larva), emerged on 30-31-V-2021; 2♂4♀, Kanna-gawa, Nakatsugawa, Chichibu, Saitama Pref., 14-XI-2010 (as larva), emerged on 26-IV-2010; 2♂1♀, Nippara, Okutama, Tokyo Pref., 27-III-2021 (as larva), emerged on 27-IV-21-V-2021; 5♂5♀, Yashajin-toge, Minami-arupusu, Yamanashi Pref., 25-III-2021 (as larva), emerged on 2-V-1-VI-2021. On Marchantia paleacea diptera : 2♂3♀, Inago, Shibakawa, Fujinomiya, Shizuoka Pref., 26-V-2021 (as larva), emerged on 17-27-VI-2021; 1♂5♀, Abe-toge, Aoi-ku, Shizuoka Pref., 30-XI-2014 (as larva), emerged on 30-IV-8-V-2014; 3♂5♀, Kuchisakamoto, Aoi-ku, Shizuoka Pref., 26-V-2021 (as larva), emerged on 1-26-VI-2021; 6♂8♀, Tsudono, Aoi-ku, Shizuoka Pref., 30-XI-2014 (as larva), emerged on 30-IV-6-V-2014; 2♂4♀, Tosayama, Kochi, Kochi Pref., 27-II-2011 (as larva), emerged on 23-28-IV-2011; 1♂3♀, Mt. Nabejiri, Taga, Shiga Pref., 4-V-2021 (as larva), emerged on 27-V-15-VI-2021; 6♂3♀, Naiku, Oe, Fukuchiyama, Kyoto Pref., 19-V-2021 (as larva), emerged on 2-10-VII-2021; 2♂3♀, Iya-kei, Ikeda, Miyoshi, Tokushima Pref., 1-II-2014 (as larva), emerged on 4-4-V-2014; 2♂5♀, Okujisso, Isa, Kagoshima Pref., 17-XII-2017 (as larva), emerged on 22-30-III-2012. On Marchantia polymorpha : 1♀, Matsubara-ko, Koumi, Nagano Pref., 18-IV-2021 (as larva), emerged on 18-VI-2021; 7♂2♀, Odarumi, Makioka, Yamanashi Pref., 30-VI-2021 (as larva), emerged on 28-VII-4-VII-2021; 5♂15♀, Muramatsu, Iwakura, Sakyo-ku, Kyoto Pref., 31-XII-2013 (as larva), emerged on?-V-2013. On Marchantia papillata grossibarba : 2♂1♀, Inago, Shibakawa, Fujinomiya, Shizuoka Pref., 26-V-2021 (as larva), emerged on 18-VII-2021; 3♂5♀, Naiku, Oe, Fukuchiyama, Kyoto Pref., 19-V-2021 (as larva), emerged on 2-10-VII-2021; 2♂1♀, Seikandoro, Kumanogawa, Shingu, Wakayama Pref., 7-VII-2021 (as larva), emerged on 27-VII-4-VIII-2021; 1♀, Nagabuchi, Ume, Saeki, Oita Pref., 29-XI-2011 (as larva), emerged on 3-V-2011. Diagnosis. A medium-sized species (wing length 1.6-1.8 mm) having dark gray scutum, dark-cornered yellow scutellum, black 1st flagellomere, black maxillary palpus, gray halteres, and brown legs. Male epandrium with a comb comprising seven or eight fused long tubercle-like setae. Larva mines the thallus of Marchantia spp. Description. Adult male (Fig. 6A-E ). Head : Head light yellow, with ocellar tubercle brown, frons light yellow, back of head dark brown (Fig. 6C ). Antenna porrect, first flagellomere black, pedicel and scape brown. Arista subbasal, black, pubescent. Clypeus, face, gena, parafacial and postgena yellow. Proboscis normal, yellow; palpus brown, cylindrical. Chaetotaxy : Front orbitals three pairs; one ori directed inward; two ors directed upward (Fig. 6B ). Orbital setulae minute and erect, in a single row. Figure 6. Phytoliriomyza marchantiae sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female (MK-AG-a442), dorsal G-I male genitalia G-I emerged from Marchantia paleacea paleacea at type locality J-M emerged from M. papillata grossibarba at Mt. Oe. G whole genitalia, ventral H phallus complex, lateral I, L epandrium, ventral J, K phallic complex, ventral and lateral M ejaculatory apodeme, lateral. Thorax : Thorax pruinose. Scutum pruinose gray, with a dark gray medial stripe on anterior 2/3, one pair of dark gray lateral stripes, and with narrow yellow patch along posterior margin (Fig. 6D ). Scutellum light yellow with lateral corners widely brownish. Subscutellum brown with anterior margin light yellow. Mediotergite, anatergite and katatergite dark brown. Pleuron largely yellow; postpronotal lobe with anterior brown spot; notopleuron with brown narrow spot along anterior lower margin; anepisternum and anepimeron with brown venters; katepisternum and meron with dark brown patches on venter (Fig. 6B ). Haltere yellowish gray, with yellow stalk. Calypter margin and hairs gray. Leg segments brown; tibia and tarsus darker (Fig. 6A ). Chaetotaxy : Scutum with 1+3 dorsocentrals, shortened anteriorly (Fig. 6D ). Acrostichal setulae five or six pairs largely in two rows. Wing : Wing length 1.6-1.7 mm, costa reaching M1 (Fig. 6A ). Length of ultimate section of vein M4 divided by penultimate section 1.6-1.8. Abdomen : Abdomen dorsally subshiny brown (Fig. 6E ). Genitalia : (Fig. 6G-M ) Epandrium dark brown, rounded apically, inner-lateral surface medially with one tubercle-like setae; inner-anterior surface ventrally with a comb comprising eight fused long tubercle-like setae; ventral margin with a row of several minute tubercle-like setae (Fig. 6I, L ). Surstylus subrectangular, setose apically. Cercus narrow, setose. Subepandrial sclerite with one pair of spines and one pair of plate-like ventral processes; tips hook-like (Fig. 6I ). Hypandrium slightly sclerotized along outer margin. Postgonite bare and goose barnacle-shaped (Fig. 6H ). Phallophorus with shallow incision below (Fig. 6J ), articulated with phallapodeme, fused to epiphallus (Fig. 6H, K ). Basiphallus with subtriangular dorsal sclerite; 2/3 x as long as mesophallus (Fig. 6H, K ). Hypophallus subtriangular membranous, lined with numerous microtrichia ventrally, medially with a pair of fused linear pale sclerites (Fig. 6H, K ). Paraphalli consisting of a pair of incurved narrow pigmented sclerites and a pair of triangular plate-like pigmented sclerites (Fig. 6H, K ). Mesophallus dark, cylindrical, widest subbasally, slightly longer than distiphallus (Fig. 6H ). Distiphallus comprising one pair of stout tubules; basal half composed of ventral dark scalpel-shaped sclerite and weaker medial region; distal half cylindrical, dorsally and laterally pigmented, with truncated, unpigmented apex (Fig. 6H, K ). Ejaculatory apodeme fan-shaped, blade pale with apical half clear; sclerite of sperm pump with lateral extension; sperm pump clear (Fig. 6M ). Female (Fig. 6I-K ). Similar to male, but scutellum darker than male (Fig. 6F ). Wing length 1.6 mm. Postabdomen : (Fig. 7A, B ) Oviscape dark brown, setigerous (Fig. 7A ). Tergite 10 cruciform, laterally uniting narrow pleural sclerites (Fig. 7B ). Cercus with two stout, apical, trichoid sensilla, 1/2 length of cercus. Spermathecae semi-orbicular (Fig. 7A ). Figure 7. Female morphology and larval ecology of Phytoliriomyza marchantiae sp. nov. A, B female postabdomen (at Renge-onsen) A oviscape and spermatheca B tergite 10 C-E habitat and mined thalli on Marchantia paleacea paleacea C habitat at type locality D, E mined thalli at Yashajin-toge and Renge-onsen F-H habitat and mined thalli on M. p. diptera F habitat at Kuchisakamoto G, H mined thalli at Iwaya-kei and Tosayama. Arrows indicate internal puparia. Variation. The yellow patch on posterior scutum slightly varied from distinct to obscure ones among localities but not among host liverwort species/subspecies (Fig, 6F-K). Etymology. The specific name refers to host plant genus, Marchantia . Japanese name. Uzumibi-zenigoke-hamoguribae. Host plants. The main host plants are Marchantia paleacea paleacea and M. p. diptera ( Marchantiaceae ), with M. polymorpha and M. papillata grossibarba also recorded as host in some localities. Mine. Larvae construct linear mines in the thallus, particularly in the midrib, and pupate in the mined thalli (Fig. 7D, E, G, H ). Biological notes. Marchantia paleacea paleacea grows on rocky substrates of cliffs, slopes and riverbanks in cool temperate deciduous forests (Fig. 7C ) and Marchantia p. diptera in warm temperate evergreen forests (Fig. 7F ). While these subspecies are frequently utilized by P. marchantiae , they are never utilized by the other three Marchantia -associated species, P. igniculus , P. tsukuyomi , and P. nubatama . P. marchantiae is also recorded from M. polymorpha in cool temperate forests and M. papillata grossibarba in warm temperate evergreen forests in some localities. Our rearing records suggest that this species is multivoltine. Distribution. Japan: Hokkaido, Honshu, Shikoku, Kyushu (Fig. 2 ). Remarks. This species resembles P. rebouliae , P. lanternaria , and P. conocephali in the narrow yellow posterior margin of the scutum and medial yellow stripe of the scutellum; it is distinguished from P. rebouliae by number of tubercle-like setae in a comb of the male epandrium (8 in P. marchantiae ; 7 in P. rebouliae ), from P. lanternaria and P. conocephali by the number of tubercle-like setae on the surstylus of the male epandrium (0 in P. marchantiae ; 1-2 in P. lanternaria and P. conocephali ). This species also resembles Phytoliriomyza miki (Strobl, 1898) in color pattern of the scutum and morphology of the male genitalia; it is distinguished from the latter by the number of tubercle-like setae in a comb on the male epandrium (8 in P. marchantiae ; 5 in P. miki ).