Phytoliriomyza Hendel, 1931

Kato, Makoto, Yamamori, Luna & Imada, Yume, 2022, Diversity underfoot of agromyzids (Agromyzidae, Diptera) mining thalli of liverworts and hornworts, ZooKeys 1133, pp. 1-164 : 1

Phytoliriomyza Hendel, 1931

Phytoliriomyza Hendel, 1931: 203 [as subgenus of Liriomyza ]. Type species: Agromyza perpusilla Meigen 1830: 181, by monotypy. Frick 1952: 410, 1959: 413; Spencer 1969: 201; Spencer and Steyskal 1986 b: 151; Papp and Černý 2017: 313; Londsale 2021: 376.

Xyraeomyia Frick, 1952: 412. Type species: Xyraeomyia conjuctimontis Frick 1952: 413, by original designation. Spencer 1965 [synonymy].

Pteridomyza Nowakowski, 1962: 97. Type species: Agromyza hilarella Zetterstedt 1848: 2776, by original designation. von Tschirnhaus 1971 [synonymy]. Manual of North American Agromyzidae 377.

Lemurimyza Spencer, 1965: 26. Type species: Liriomyza enormis Spencer 1963 c: 114, by original designation. von Tschirnhaus 1971 [synonymy].

Nesomyza Spencer in Spencer and Stegmaier 1973: 190. Type species: Nesomyza fusculoides Spencer in Spencer and Stegmaier 1973: 190, by original designation. Spencer 1973 [synonymy].

Diagnosis.

Head: Head yellow, frons often with pruinosity. Orbital plates more or less emerge from the plane of frons. Front orbitals three pairs; one ori directed inward and two ors directed up. 1-2(3) pairs of medio-clinate lower orbital setae, two pairs of reclinate upper orbital setae. Orbital setulae proclinate, upright or partly reclinate. Postocellar and ocellar setae well developed. First flagellomere round.

Thorax: Scutum with 1+3 dorsocentrals. Acrostichal setulae in two rows, but lacking in some species. Postpronotal, two propleural, presutural, and propleural setae normal or strong. Anepisternum with a long posterior seta and with two or three shorter posterior setae. Katepisternum with one or two long setae. Legs simple, only ventroapical seta on middle tibia present. Costal vein ends at apex of M1. The ratio of ultimate and penultimate sections of M4 various.

Abdomen: Male abdomen lacks stridulatory organs.

Male genitalia: Epandrium round apically, often with a comb of several fused tubercle-like setae and/or elongated tubercle-like setae on the inner surface; surstylus setigerous and sometimes with one or a few tubercle-like setae. Subepandrial sclerite short but very broad, connecting bases of surstyli through a special sclerite. Hypandrium very long but very thin. Hypophallus membranous often with a pair of parallel sclerites medially. Paraphallus present or absent. Mesophallus cylindrical and well sclerotized. Distiphallus with basal part formed by usually a paired sclerite and an apical part, which terminates in paired tubes or in extremely long less sclerotized tubules. Ejaculatory apodeme variable, but basal part usually broad and blade not too large.

Female postabdomen: Tergite 10 trifurcate or cruciform posteriorly. Cercus with two stout, apical, trichoid sensilla.

Classification of Phytoliriomyza species associated with bryophytes

The morphological characteristics of the species associated with bryophytes coincided with the characteristics unique to the genus Phytoliriomyza . In addition to these characteristics of adult males, we found characteristics of adult females unique to the bryophyte-associated species, in which tergite 10 was trifurcate or cruciform and each cercus bears two stout, apical, trichoid sensilla.

All the species associated with bryophytes are distinguished by color of antenna, color of maxillary palpus, color pattern of scutum and morphology of male genitalia. Colors of 1st flagellomere of antenna and maxillary palpus are dark or yellow, and unique to each species. Scutum was yellow or dark, and the yellow scutum often had three pairs of dark longitudinal stripes: medial stripes on anterior 2/3 (always merged together except in one species), intra-alar stripes (inner lateral stripes) and supra-alar stripes (outer lateral stripes) on anterior 4/5. In some species, the intra-alar stripe and the supra-alar stripe are merged together to shape a wide band, and often merged with the medial stripe. The color pattern of scutum is unique to each species at least among the species sharing the same host bryophyte genus. Although some species are very similar in external morphology, they could be clearly distinguished by the number of tubercle-like setae in a comb on the male epandrium and other male genital morphological characters.

The 39 recorded Phytoliriomyza species can be classified into four groups ( Phytoliriomyza phaeocerotis group, Phytoliriomyza mesnili group, Phytoliriomyza alpicola group, and Phytoliriomyza dorsata group) based on presumed synapomorphy of the following characteristics: distribution of acrostichal setulae on scutum, morphology of epandrium and distiphallus of male genitalia, and color of legs. By assessing the morphological characteristics of the Phytoliriomyza species (Table 3 ), we obtained a key to species as follows.

Key to the Japanese Phytoliriomyza

1	Acrostichal setulae vestigial, at most 4 pairs in 2 rows (Fig. 74D ); male epandrium with convex inner ventral margin (Fig. 74I ); distal end of distiphallus tapering out (Fig. 74G ); small species with wing length ranging from 1.1 to 1.8 mm	2
-	Acrostichal setulae at least 5 pairs in 2 (or rarely 4) rows (Fig. 1D ); male epandrium with concave inner ventral margin (Fig. 1J ); distal end of distiphallus truncated (Fig. 1H ); small to large species with wing length ranging from 1.3 and 2.9 mm	8
2	Surstylus small, lobate, not elongated, not setose apically (Fig. 74I )	3 [ Phytoliriomyza phaeocerotis -group: associated with hornworts]
-	Surstylus large, elongated, setose apically (Fig. 63K )	5 [ Phytoliriomyza mesnili -group: associated with Riccia and Asterella ]
3	Legs dark brown (Fig. 72A ); epandrium with a comb of tubercle-like setae (Fig. 72H ); distiphallus shorter than phallapodeme (Fig. 72G )	Phytoliriomyza megacerotis [host: Megaceros ]
-	Legs yellow or brownish yellow (Fig. 74A ); epandrium without a comb of tubercle-like setae (Fig. 74I ); distiphallus longer than phallapodeme (Fig. 74G )	4
4	1st flagellomere and haltere yellow (Fig. 70B ); scutellum dark brown, medially with a yellow patch (Fig. 70D )	P. foliocerotis [host: Folioceros ]
-	1st flagellomere and haltere dark (Fig. 74B ); scutellum wholly gray (Fig. 74D )	P. phaeocerotis [host: Phaeoceros , Notothylas , Anthoceros ]
5	Maxillary palpus dark-colored (Fig. 63C ); legs yellow (Fig. 63A ); scutum yellowish gray with dark intra-alar and supra-alar stripes (Fig. 63D )	P. ricciae [host: Riccia ]
-	Maxillary palpus yellow (Fig. 68F ); legs dark-colored (Fig. 68A ); scutum completely gray or gray with longitudinal dark stripes (Fig. 66I )	6
6	Scutum completely gray (Fig. 21D ); acrostichal setula absent (Fig. 21D )	P. iriomotensis [host: Asterella ]
-	Scutum bluish gray with longitudinal dark stripes (Fig. 66I ); acrostichal setulae 1-4 pairs in two rows (Fig. 66I )	7
7	Scutum with 3 pairs of longitudinal dark gray stripes (Fig. 66I ); epandrium with a dense cluster of short tubercle-like setae at inner margin (Fig. 66L )	P. sexfasciata [host: Riccia ]
-	Scutum with a medial stripe and a pair of lateral longitudinal dark gray stripes (Fig. 68I ); epandrium with 3 strong tubercle-like setae at inner margin (Fig. 68N )	P. caerulescens [host: Riccia ]
8	Legs dark-colored (Fig. 56B ); scutum uniformly dark-colored (Fig. 56 D )	9 [ Phytoliriomyza alpicola group]
-	Legs yellow (Fig. 1B ); scutum yellow with or without longitudinal lateral stripes (Fig. 1D ); if scutum dark-colored, arista yellow (Fig. 44D, F )	18 [ Phytoliriomyza dorsata group]
9	Haltere yellow (Fig. 10E )	10
-	Haltere dark-colored (Fig. 56B )	12
10	Maxillary palpus yellow (Fig. 8C ); wing length <2 mm	P. nubatama [host: Marchantia ]
-	Maxillary palpus dark-colored (Fig. 10B ); wing length> 2 mm	11
11	Pleuron with ventral half dark-colored (Fig. 10B ); surstylus with 1 tubercle-like seta (Fig. 10K )	P. dumortierae [host: Dumortiera ]
-	Pleuron wholly yellow (Fig. 35B ); surstylus with 2 tubercle-like setae (Fig. 35I )	P. wiesnerellae [host: Wiesnerella ]
12	Scutellum wholly gray (Fig. 62D )	P. suetsugui [host: Conocephalum ]
-	Scutellum wholly yellow or at least partly yellow (Fig. 56B )	13
13	Pedicel of antenna yellow (Fig. 5C ); scutum completely dark gray (Fig. 5D ); male surstylus basally with a hypertrophied columnar arm, bearing tubercle-like seta apically (Fig. 5I )	P. tsukuyomi [host: Marchantia ]
-	Pedicel of antenna brown (Fig. 56C ); scutum dark gray except for posterior margin or mid-posterior area (Figs 56D , 58D ); male surstylus lacking a hypertrophied columnar arm (Fig. 5I )	14
14	Legs dark brown (Fig. 56B ); Scutum dark gray with posterior margin yellow (Fig. 56D )	P. alpicola [host: Conocephalum ]
-	Legs yellowish brown (Fig. 58B ); scutum dark gray with a semicircular yellow patch at mid posterior area (Fig. 58D )	15
15	Male surstylus lacking tubercle-like seta (Fig. 6I ); epandrium lacking an anteriorly directed tubercle-like seta on inner-lateral surface (Fig. 6I )	16
-	Male surstylus with 1 or 2 tubercle-like setae (Fig. 59K ); epandrium with an anteriorly directed tubercle-like seta on inner-lateral surface (Fig. 59K )	17
16	Male epandrium with a comb comprising 8 basally fused, long, tubercle-like setae (Fig. 6I ); epandrium with one tubercle-like seta at inner-lateral margin (Fig. 6G )	P. marchantiae [host: Marchantia ]
-	Male epandrium with a comb comprising 7 basally fused, long, tubercle-like setae (Fig. 33J ); epandrium with 1 tubercle-like seta at inner-posterior margin (Fig. 33H )	P. rebouliae [host: Reboulia ]
17	Male epandrium with a comb comprising 5 or 6 basally fused, long, tubercle-like setae (Fig. 59I ); Surstylus with 2 tubercle-like setae (Fig. 59K )	P. conocephali [host: Conocephalum ]
-	Male epandrium with a comb comprising 7 basally fused long tubercle-like setae (Fig. 58I ); surstylus with 1 tubercle-like seta (Fig. 58I )	P. lanternaria [host: Conocephalum ]
18	Scutum and scutellum gray (Fig. 44D ); arista yellow (Fig. 44F )	P. ugetsu [host: Conocephalum ]
-	Scutum yellow at least in part, and scutellum yellow (Fig. 44D ); arista black (Fig. 44F )	19
19	Scutum wholly yellow, lacking dark stripes (Fig. 49D )	20
-	Scutum yellow, with 1 or 2 pairs of dark longitudinal stripes (Fig. 49D )	21
20	1st flagellomere black (Fig. 51B ); tubercle-like seta on inner surface of epandrium 1.5 × as long as the seta of a comb (Fig. 51I )	P. luteola [host: Conocephalum ]
-	1st flagellomere yellow (Fig. 52B ); tubercle-like seta on inner surface of epandrium as long as the seta of a comb (Fig. 52I )	P. helva [host: Conocephalum ]
21	Scutum lacking a medial dark stripe, but with brown unfused intra-alar and supra-alar stripes (Fig. 49D )	P. pallidofasciata [host: Conocephalum ]
-	Scutum with a medial dark stripe (Fig. 1D )	22
22	Intra-alar and supra-alar stripes of scutum separate and unfused (Fig. 1D )	23
-	Intra-alar and supra-alar stripes of scutum fused, forming a pair of wide bands (Fig. 3D )	29
23	Scutum subglossy with pruinosity; intra-alar and supra-alar stipes with similar thickness (Fig. 46B ); acrostichal setulae in 2 rows (Fig. 46B )	24
-	Scutum glossy; supra-alar stipe is pale and indistinct (Fig. 54D ); acrostichal setulae in 4 rows (Fig. 54D )	P. bifasciata [host: Conocephalum ]
24	Intra-alar stripes not confluent with a pair of lateral presutural dark ovoid spots (Fig. 46B ); 1st flagellomere yellow in male, black in female (Fig. 46A, D )	P. nigroflava [host: Conocephalum ]
-	Intra-alar stripes adjoining and confluent with 1 pair of lateral presutural dark ovoid spots (Fig. 1D ); 1st flagellomere yellow or black, same in both sexes	25
25	1st flagellomere yellow (Fig. 24B )	26
-	1st flagellomere black (Fig. 1B )	28
26	Intra-alar stripes and lateral presutural dark ovoid spots with silvery pruinosity (Fig. 24D ); male epandrium with a comb comprising 4 or 5 tubercle-like setae (Fig. 24H	P. argentifasciata [host: Reboulia ]
-	Intra-alar stripes and lateral presutural dark ovoid spots without silvery pruinosity (Fig. 27D ); male epandrium with a comb comprising 6 tubercle-like setae (Fig. 27I )	27
27	Male epandrium with an extremely elongated arm, bearing 2, dark, ventrally curved, tubercle-like setae borne at wide angle (Fig. 27H ); surstylus lobate, setose apically, with 1 short tubercle-like seta subapically (Fig. 27K )	P. longifurcae [host: Reboulia ]
-	Male epandrium with a basally enlarged, slightly flattened hypertrophied arm, which bears a dark laterally enlarged tubercle-like seta (Fig. 18G ); surstylus narrow, bare, with 1 long tubercle-like seta apically (Fig. 18G )	P. calcicola [host: Plagiochasma ]
28	Scutellum yellow with lateral margins dark-colored (Fig. 1D ); surstylus with 2 tubercle-like setae (Fig. 1J )	P. dorsata [host: Marchantia ]
-	Scutellum wholly yellow (Fig. 48D ); surstylus with 1 tubercle-like seta (Fig. 48I )	P. brunofasciata [host: Conocephalum ]
29	1st flagellomere black (Fig. 37C ); male epandrium with a comb of tubercle-like setae (Fig. 37I )	30
-	1st flagellomere yellow (Fig. 14B ); male epandrium lacking a comb of tubercle-like setae (Fig. 14K )	36
30	Scutellum brownish yellow, tergite of abdomen brown (Fig. 13D ); haltere brown (Fig. 13D ); wing length <1.8 mm	P. crepusculum [host: Dumortiera ]
-	Scutellum yellow, tergite of abdomen yellow (Fig. 47D ); haltere yellow (Fig. 47D ); wing length> 1.9 mm	31
31	Lateral fused band of scutum reaching just before scutellum (Fig. 22D ); male epandrium with a comb comprising 3 or 4 fused long tubercle-like setae, and with an enlarged arm bearing a strong tubercle-like seta (Fig. 22J )	P. cometiformis [host: Reboulia ]
-	Lateral fused band of scutum ending at anterior 7/8 length of scutum before scutellum (Fig. 37D ); male epandrium with a comb comprising 7-12 fused long tubercle-like setae (Fig. 37I )	32
32	Medial stripe of scutum not confluent with the lateral fused band, and the 2 bands are delimited by a yellow line (Fig. 37D ); yellow patch ranging from posterior scutum to scutellum well-defined (Fig. 37D ); male epandrium with 1 pair of tubercle setae at posterior end on inner margin (Fig. 37G )	33
-	Medial stripe of scutum confluent with the lateral fused band (Fig. 43D ); yellow patch ranging from posterior scutum to scutellum oblong and ill-defined (Fig. 43D ); male epandrium with 1 pair of tubercle setae on subposterior inner margin (Fig. 43G )	34
33	Scutellum wholly yellow (Fig. 37D ); male epandrium with a comb comprising 7 or 8 fused tubercle-like setae (Fig. 37I )	P. luna [host: Conocephalum ]
-	Scutellum yellow with lateral margins darkened (Fig. 40D ); male epandrium with a comb comprising 9-12 fused tubercle-like setae (Fig. 40J )	P. izayoi [host: Conocephalum ]
34	Abdomen dorsally with a pair of brown lateral semicircular patches on anterior half of the 2nd tergite (Fig. 3E ); male epandrium with a comb comprising 5 fused tubercle-like setae (Fig. 3I )	P. igniculus [host: Marchantia ]
-	Abdomen dorsally without dark patches on tergites (Fig. 42B ); male epandrium with a comb comprising 6-8 fused tubercle-like setae (Fig. 42J )	35
35	Male epandrium with a comb comprising 6 fused tubercle-like setae (Fig. 42J )	P. chichibuensis [host: Conocephalum ]
-	Male epandrium with a comb comprising 8 fused tubercle-like setae (Fig. 43I )	P. caliginosa [host: Conocephalum ]
36	Yellow patch of midposterior scutum darkened and ill-defined (Fig. 32D ); Tergite of abdomen brownish yellow (Fig. 32E )	P. aratriformis [host: Conocephalum ]
-	Yellow patch of midposterior scutum rectangular and well-defined (Fig. 14D ); Tergite of abdomen yellow (Fig. 14E )	37
37	Male epandrium with a pair of elongated, hypertrophied arms, each bearing a tubercle-like seta apically (Fig. 14K )	38
-	Male epandrium without a pair of elongated, hypertrophied arms (Fig. 17I )	P. plagiochasmatos [host: Plagiochasma ]
38	Hypertrophied arm on male epandrium strongly curved outward (Fig. 14K )	P. arcus [host: Plagiochasma ]
-	Hypertrophied arm on male epandrium protruded forward (Fig. 29I )	P. falcata [host: Reboulia ]

Taxonomy

We describe 39 Phytoliriomyza species in order of phylogenetic position of their host plants (as shown in Tables 1 and 2 ), focusing on morphological characteristics, geographical variation, host-plant records, and life history of each species.

Species associated with Marchantia

Figures

Figure 1. Phytoliriomyza dorsata A-E male at Yakumo A habitus B lateral body C frontal head D dorsal thorax E thorax in posterior view F, G female at Yakumo F dorsal thorax G lateral body H-J male genitalia H whole genitalia, ventral I phallic complex, ventral J epandrium, ventral K, L female postabdomen K oviscape and spermatheca L tergite 10 M-O mined thalli of Marchantia polymorpha M mined thallus dissected to show interior and puparium (arrow) N mined thallus with exit hole O the same thallus with the mine dissected to show a puparium (arrow).

Figure 3. Phytoliriomyza igniculus sp. nov. A-E holotype male A habitus B lateral body C frontal head and ventral body D dorsal thorax E thorax in posterior view F-H male genitalia F whole genitalia G phallic complex H postgonite I epandrium J a living female fly on a thallus of Marchantia polymorpha K, L female postabdomen K oviscape and spermatheca L tergite 10.

Figure 5. Phytoliriomyza tsukuyomi sp. nov. A-E holotype male A habitus, B lateral C frontal D dorsal E posterior F paratype female (MK-AG-a 442) G-J male genitalia G whole genitalia, ventral H phallus complex, lateral I epandrium J ejaculatory apodeme, lateral K, L female postabdomen K oviscape and spermatheca L tergite 10 M mined thallus of Marchantia polymorpha.

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-a 442), 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.

Figure 8. Phytoliriomyza nubatama sp. nov. A-D, G holotype male A habitus B lateral C frontal D dorsal E, F paratype female (MK-AG- 323), E dorsal F frontal G-L male genitalia (G-K type locality L Kume Island) G whole genitalia, ventral H phallic complex, lateral I epandrium, ventral J ejaculatory apodeme lateral K postgonite L phallic complex, ventral.

Figure 10. Phytoliriomyza dumortierae sp. nov. A-D holotype male A habitus B lateral C frontal D dorsal E paratype female (MK-AG- 241) F-L male genitalia (F at type locality G-I at Ichiu J-L at Tashiro) F whole genitalia, ventral G whole genitalia, lateral H phallic complex, lateral I ejaculatory apodeme, lateral J phallic complex, ventral K-L epandrium, posterior and anterior.

Figure 13. Phytoliriomyza crepusculum sp. nov. A-C holotype male A habitus B lateral C frontal D paratype male (MK-AG-a 309), dorsal E paratype female (MK-AG- 274), dorsal F-I male genitalia F ejaculatory apodeme, dorsal G whole genitalia, ventral H phallic complex, lateral I epandrium, ventral J puparium K-L female postabdomen K oviscape and spermatheca L tergite 10 M habitat of type locality N-P mined thalli of Dumortiera hirsuta (N type locality O Kiyosumi P Mt. Nabejiri). An arrow in N indicate an puparium.

Figure 14. Phytoliriomyza arcus sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female (MK-AG-a 16), dorsal G-K male genitalia (G at type locality H-K at Todai-shiraiwa) G, H whole genitalia, ventral I phallic complex, lateral J ejaculatory apodeme, lateral K epandrium, ventral.

Figure 17. Phytoliriomyza plagiochasmatos sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female (MK-AG-a 523), dorsal G-J male genitalia G whole genitalia, ventral H phallic complex, lateral I, J epandrium, ventral and anterior K ejaculatory apodeme, lateral L live male fly.

Figure 18. Phytoliriomyza calcicola sp. nov. A-D holotype male A habitus B lateral C frontal D dorsal E, F paratype female (MK-AG-a 222) E dorsal F posterior G, H male genitalia G epandrium H phallic complex in lateral view I live female fly.

Figure 21. Locality records of three Phytoliriomyza species associated with Asterella and Reboulia spp.: P. iriomotensis and P. cometiformis.

Figure 22. Phytoliriomyza cometiformis sp. nov. A-D holotype male A habitus B lateral C frontal D dorsal E paratype female, dorsal (MK-AG-a 428) F-J male genitalia (F-H at type locality I, J at Sagiura) F whole genitalia, ventral G phallic complex, lateral H, J epandrium, ventral I ejaculatory apodeme, lateral.

Figure 24. Phytoliriomyza argentifasciata sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female (MK-AG- 436) G-J male genitalia (G-J at type locality K at Kume Is.) G whole genitalia, ventral H epandrium, ventral I ejaculatory apodeme, lateral, J phallic complex, ventral K phallic complex, lateral.

Figure 27. Phytoliriomyza longifurcae sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F, G paratype female (MK-AG-a 497) F dorsal G lateral H-L male genitalia H epandrium, ventral I whole genitalia, ventral J, K phallic complex, ventral and lateral L ejaculatory apodeme, lateral.

Figure 29. Phytoliriomyza falcata sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female (MK-AG-a 18) dorsal G-L male genitalia (G-K, at type locality L at Tatsuyama) G whole genitalia, ventral H phallic complex, ventral I, J, L epandrium (I anterior J anterior-ventral L ventral) K ejaculatory apodeme, lateral.

Figure 32. Phytoliriomyza aratriformis sp. nov. A-D holotype male A habitus B lateral C frontal D dorsal E paratype female (MK-AG-a 346) F-I male genitalia (K-H type locality I, J Okuchihibu) F, I phallic complex, ventral (distiphallus lost) G phallic complex, lateral H epandrium, ventral J postgonite K, L female postabdomen K oviscape and spermatheca (one spermatheca split into two) L tergite 10 M live female N habitat at Nakatsugawa O mined thallus of Reboulia hemisphaerica orientalis at Nakatsugawa.

Figure 33. Phytoliriomyza rebouliae sp. nov. A-D holotype male A habitus B lateral C frontal D dorsal E, F paratype female (MK-AG-a 424) E posterior F lateral G live female fly H-K male genitalia H whole genitalia, ventral I phallic complex, lateral J epandrium, ventral K ejaculatory apodeme, lateral.

Figure 35. Phytoliriomyza wiesnerellae sp. nov. A-D holotype male A habitus B lateral C frontal D dorsal E paratype female (MK-AG-a 342), posterior F-J male genitalia F whole genitalia, ventral G epandrium, lateral H phallic complex, lateral I epandrium, ventral J ejaculatory apodeme, lateral.

Figure 37. Phytoliriomyza luna sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female (MK-AG- 512), dorsal G-K male genitalia G whole genitalia, ventral and lateral H phallic complex, lateral I epandrium, ventral J ejaculatory apodeme, lateral K postgonite L-N female postabdomen at Aizankei L oviscape and spermatheca M tergite 10 N spermatheca.

Figure 40.. Phytoliriomyza izayoi sp. nov. A-D holotype male A habitus B lateral C frontal D dorsal E, F paratype female (MK-AG-a 262) E posterior F lateral G-K male genitalia (G-I type locality J, K Renge-onsen) G whole genitalia, ventral H phallic complex, ventral I ejaculatory apodeme, dorsal J epandrium, ventral K phallic complex, lateral.

Figure 42. Phytoliriomyza chichibuensis sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F live male fly G-K male genitalia G whole genitalia, ventral H, I phallic complex, ventral and lateral J epandrium, ventral K ejaculatory apodeme, lateral, L, M female postabdomen L oviscape and spermatheca M tergite 10 N mined thallus of Conocephalum sp. 2.

Figure 43. Phytoliriomyza caliginosa sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female (MK-AG-a 224) G-J male genitalia G whole genitalia, ventral H phallic complex, ventral I epandrium, ventral J ejaculatory apodeme, lateral K, L female postabdomen K oviscape and spermatheca L tergite 10 M mined thallus of Conocephalum orientalis at Tsurara-kannon.

Figure 44. Phytoliriomyza ugetsu sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female (MK-AG- 554), frontal G-J male genitalia (G, H at type locality I Yaku Is. J Mikisato) G whole genitalia, ventral H phallic complex, lateral I epandrium, ventral J ejaculatory apodeme, lateral K a live female fly foraging on Conocephalum orientalis thallus at Tategasaki L, M female postabdomen L oviscape and spermatheca M tergite 10 N puparia O mined thallus of Conocephalum orientalis at type locality P habitat at Tategasaki.

Figure 46. Phytoliriomyza nigroflava sp. nov. A-C holotype male A habitus B dorsal C frontal D-G paratype female (MK-AG-a 22) D habitus E dorsal F frontal G posterior H-K male genitalia H whole genitalia, ventral I epandrium, ventral J phallic complex, lateral K ejaculatory apodeme, lateral B postgonite.

Figure 47. Female morphology and larval ecology of Phytoliriomyza nigroflava sp. nov. A, B female postabdomen A oviscape and spermatheca B tergite 10 C, D habitat (C type locality D at Shirabiso-toge) E-H mined thalli of Conocephalum salebrosum at type locality. Arrows in G and H shows internal puparia.

Figure 48. Phytoliriomyza brunofasciata sp. nov. A-E holotype male A habitus B dorsal C frontal D dorsal E posterior F paratype female (MK-AG-a 42), dorsal G-J male genitalia G whole genitalia, ventral H phallic complex, ventral I epandrium, ventral J ejaculatory apodeme, dorsal L, M female postabdomen L oviscape and spermatheca M tergite 10 N habitat at Nishiyama-onsen O-R mined thalli of Conocephalum salebrosum. Arrows in Q and R indicate puparia.

Figure 49. Phytoliriomyza pallidofasciata sp. nov. A-E holotype male A habitus B dorsal C frontal D dorsal E posterior F paratype female (MK-AG-a 520), dorsal G-J male genitalia G whole genitalia H ejaculatory apodeme, lateral I phallic complex, lateral J epandrium, ventral K live fly L, M female postabdomen L oviscape and spermatheca M tergite 10 N-P mined thalli of Conocephalum orientalis. An arrow in O indicates an internal puparium.

Figure 51. Phytoliriomyza luteola sp. nov. A-E holotype male A habitus B dorsal C frontal D dorsal E posterior F paratype female (MK-AG-a 1), dorsal G-K male genitalia (G at type locality K-M at Mt. Horoiwa K at Sounkyo) G, H whole genitalia, ventral I epandrium, ventral J phallic complex, lateral K ejaculatory apodeme, lateral L, M female postabdomen L oviscape and spermatheca M tergite 10 N habitat at type locality O mined thallus of Conocephalum salebrosum P live female fly at Eniwa.

Figure 52. Phytoliriomyza helva sp. nov. A-E holotype male A habitus B dorsal C frontal D dorsal E posterior F paratype female (MK-AG-a 541), frontal G-J male genitalia G whole genitalia, ventral H phallic complex, lateral I epandrium, ventral J ejaculatory apodeme, lateral.

Figure 54. Phytoliriomyza bifasciata sp. nov. A-E holotype male A habitus B dorsal C frontal D dorsal E posterior F paratype female (MK-AG-a 272), dorsal G-J male genitalia G whole genitalia H phallic complex I epandrium J ejaculatory apodeme, dorsal K, L female postabdomen K oviscape and spermatheca L tergite 10 M live female fly at Hozaka N habitat at Hachimori O-Q mined thalli (O Conocephalum orientalis at Hachimori P C. salebrosum at Haccho-toge Q C. salebrosum at Akka).

Figure 56. Phytoliriomyza alpicola A-E male at Ikawa-toge A habitus B dorsal C frontal D dorsal E posterior F live female fly at Mitsumine-Jinja G-K male genitalia (G-J at Ikawa-toge K at Samani) G whole genitalia H phallic complex I epandrium J, K ejaculatory apodeme, ventral and lateral.

Figure 58. Phytoliriomyza lanternaria sp. nov. A-D holotype male A habitus B dorsal C frontal D dorsal E paratype female (MK-AG- 430), dorsal F-I male genitalia (F-H at type locality I Mt. Horoiwa) F whole genitalia, ventral G phallic complex, lateral H, I epandrium, ventral, J, K female postabdomen J oviscape and spermatheca K tergite 10 L habitat at type locality M, N mined thalli of Conocephalum orientalis. An arrow in N indicates an internal puparium O live female fly at Yakumo.

Figure 59. Phytoliriomyza conocephali sp. nov. A-E holotype male A habitus B dorsal C frontal D dorsal E posterior F paratype female (MK-AG-a 444), dorsal G-K male genitalia (G on Conocephalum orientalis H-K on C. japonicum) G, H whole genitalia, ventral I phallic complex, lateral J ejaculatory apodeme, dorsal K epandrium, ventral.

Figure 62. Phytoliriomyza suetsugui sp. nov. A-D holotype male A habitus B lateral C frontal D dorsal E-H paratype female (MK-AG-a 434) E dorsal F frontal G lateral H posterior I-L male genitalia I whole genitalia, ventral J phallic complex, lateral K epandrium, ventral L ejaculatory apodeme M, N female postabdomen M oviscape and spermatheca N tergite 10 O, P habitat at type locality Q mined thallus of Conocephalum orientalis.

Figure 63. Phytoliriomyza ricciae sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female, frontal (MK-AG- 201) G-K male genitalia (G-J on Riccia nipponica K on R. huebeneriana) G whole genitalia, ventral H phallic complex, lateral I phallic complex, ventral L ejaculatory apodeme, lateral M epandrium, ventral.

Figure 66. Phytoliriomyza sexfasciata sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F-J paratype female (MK-AG-a 573) F habitus F lateral G frontal H dorsal I posterior K-O male genitalia K whole genitalia, ventral L epandrium, ventral M phallic complex, lateral N genitalia, lateral O ejaculatory apodeme, lateral P live female fly walking on soil.

Figure 68. Phytoliriomyza caerulescens sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F-J paratype female (MK-AG-a 558) F frontal G habitus H lateral I dorsal J posterior K female fly ovipositing to thallus of Riccia billardieri L-O male genitalia L whole genitalia, ventral M phallic complex, lateral N epandrium, ventral O ejaculatory apodeme, ventral.

Figure 70. Phytoliriomyza foliocerotis sp. nov. A-D holotype male A habitus B lateral C frontal D dorsal E, F paratype female (MK-AG-a 450) E dorsal F posterior G-J male genitalia G whole genitalia, ventral H ejaculatory apodeme, lateral I phallic complex, lateral J epandrium, posterior-ventral K, L female postabdomen K oviscape and spermatheca with spermatheca L tergite 10 M habitat at type locality N mined thallus of Folioceros fuciformis

Figure 72. Phytoliriomyza megacerotis sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female (MK-AG-a 14), dorsal G-J male genitalia (G at type locality H, I at Tashiro J at Yasukawa-keikoku) G whole genitalia, ventral H epandrium, ventral I ejaculatory apodeme, lateral J phallic complex, lateral K, L female postabdomen K oviscape and spermatheca L tergite 10.

Figure 74. Phytoliriomyza phaeocerotis sp. nov. A-E holotype male A habitus B lateral C frontal D dorsal E posterior F paratype female (MK-AG-a 10), dorsal G-K male genitalia (G, H emerged from Notothylas temperata at type locality I emerged from Phaeoceros carolinianus at Kakeroma Is. J emerged from Notothylas temperata at Kibi-kogen K emerged from Anthoceros punctatus at Inago) G whole genitalia, ventral H, J phallic complex, lateral and ventral I epandrium, ventral K ejaculatory apodeme, lateral L phallic complex, lateral.