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 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 Cerny 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. Table 3. A synopsis of morphological characteristics of Japanese bryophyte-associated Phytoliriomyza species.
Group Phytoliriomyza species Color of 1st flagellomere Color of maxillary pulpus Wing length (mm) No. pairs of acro-stichal setulae Color of scutum Color of scutellum Color of leg Medial, inter-alar and supra-alar stripes1 color of haltere Surstylus No. tubercle-like setae of comb on epan-drium2 No. hyper-trohied arm on epan-drium No. tubercle-like setae on surstylus Distal end of disti-phalus Length of disti-phalus Host plant genus
Phytoliriomyza phaeocerotis -G Phytoliriomyza megacerotis dark dark 1.2-1.4 2-3 dark dark dark 0-0-0 dark lobate 6 0 6 tapering out short Megaceros
- Phytoliriomyza foliocerotis yellow yellow 1.1-1.3 3-4 dark partly yellow yellow 0-0-0 yellow lobate - 0 0 tapering out long Folioceros
- Phytoliriomyza phaeocerotis dark yellow 1.2-1.5 0 dark dark yellow 0-0-0 dark lobate - 0 0 tapering out long Phaeoceros , Nothotylas , Anthoceros
Phytoliriomyza mesnili -G Phytoliriomyza caerulescens dark yellow 1.1-1.3 3-4 dark dark dark 0-0-0 dark elongated - 0 0 tapering out short Riccia
- Phytoliriomyza ricciae dark dark 1.0-1.3 0-2 dark dark yellow 1-1-1 dark elongated (2) 0 2 tapering out short Riccia
- Phytoliriomyza sexfasciata dark yellow 1.2-1.5 1-3 dark dark dark 0-0-0 dark elongated (40) 0 0 tapering out short Riccia
- Phytoliriomyza iriomotensis dark yellow 1.4-1.7 0 dark dark dark 0-0-0 dark elongated (3) 0 1 tapering out short Asterella
Phytoliriomyza alpicola -G Phytoliriomyza tsukuyomi dark dark 1.6-1.7 6-7 dark yellow dark 0-0-0 dark lobate 4 2 1 truncated short Marchantia
- Phytoliriomyza alpicola dark dark 1.7-1.8 5-6 dark yellow dark 0-0-0 dark lobate 6 0 1 truncated short Conocephalum
- Phytoliriomyza marchantiae dark dark 1.6-1.8 5-6 dark yellow dark 0-0-0 dark lobate 8 0 0 truncated short Marchantia
- Phytoliriomyza rebouliae dark dark 1.3-1.7 6-7 dark yellow dark 0-0-0 dark lobate 7 0 0 truncated short Reboulia
- Phytoliriomyza conocephali dark dark 1.3-1.7 5-6 dark yellow dark 0-0-0 dark lobate 5-6 0 2 truncated short Conocephalum
- Phytoliriomyza lanternaria dark dark 1.8-1.9 5-6 dark yellow dark 0-0-0 dark lobate 7 0 1 truncated short Conocephalum
- Phytoliriomyza suetsugui dark dark 1.3-1.5 5-6 dark dark dark 0-0-0 dark lobate 6 0 1 truncated short Conocephalum
- Phytoliriomyza dumortierae dark dark 1.9-2.2 7-8 dark yellow dark 0-0-0 yellow lobate 2 0 1 truncated short Dumortiera
- Phytoliriomyza wiesnerellae dark dark 2.0-2.3 7 dark yellow dark 0-0-0 yellow lobate (1) 0 2 truncated short Wiesnerella
- Phytoliriomyza nubatama dark yellow 1.3-1.6 5 dark yellow dark 0-0-0 yellow lobate 7 0 3 truncated short Marchantia
Phytoliriomyza dorsata -G Phytoliriomyza ugetsu dark yellow 2.1-2.7 10-12 dark dark yellow 0-0-0 dark lobate 6 0 1 truncated short Conocephalum
- Phytoliriomyza luteola dark yellow 1.9-2.0 8-9 yellow yellow yellow 0-0-0 yellow lobate 3-4 0 1 truncated short Conocephalum
- Phytoliriomyza helva yellow yellow 1.8-2.1 5-6 yellow yellow yellow 0-0-0 yellow lobate 3 0 1 truncated short Conocephalum
- Phytoliriomyza pallidofasciata dark yellow 1.9-2.0 8-9 yellow yellow yellow 0-1-1 yellow lobate 4-5 0 1 truncated short Conocephalum
- Phytoliriomyza argentifasciata yellow yellow 1.5-1.9 5 yellow yellow yellow 1-1-1 yellow lobate 4-5 0 1 truncated short Reboulia
- Phytoliriomyza longifurcae yellow yellow 1.5-1.6 5 yellow yellow yellow 1-1-1 yellow lobate 6 1 1 truncated short Reboulia
- Phytoliriomyza calcicola yellow yellow 1.6-1.7 7 yellow yellow yellow 1-1-1 yellow lobate 6 1 1 truncated short Plagiochasma
- Phytoliriomyza brunofasciata dark yellow 1.9-2.2 7-8 yellow yellow yellow 1-1-1 yellow lobate 5-7 0 1 truncated short Conocephalum
- Phytoliriomyza dorsata dark yellow 1.7-1.8 8-9 yellow yellow yellow 1-1-1 yellow lobate 6 0 2 truncated short Marchantia
- Phytoliriomyza nigroflava yellow/dark yellow 2.2-2.3 8-10 yellow yellow yellow 1-1-1 yellow lobate 5-7 0 1 truncated short Conocephalum
- Phytoliriomyza bifasciata dark yellow 2.2-2.3 22-26* yellow yellow yellow 1-1-0 yellow lobate 3-4 1 1 truncated short Conocephalum
- Phytoliriomyza cometiformis dark yellow 2.0-2.2 8 yellow yellow yellow 1-1/1 yellow lobate 3+3 2 1 truncated short Reboulia
- Phytoliriomyza luna dark yellow 2.7-2.9 8-10 yellow yellow yellow 1-1/1 yellow lobate 7-8 0 1 truncated short Conocephalum
- Phytoliriomyza izayoi dark yellow 2.4-2.5 7-9 yellow yellow yellow 1-1/1 yellow lobate 9-12 0 1 truncated short Conocephalum
- Phytoliriomyza chichibuensis dark yellow 2.2-2.9 8-9 yellow yellow yellow 1/1/1 yellow lobate 6 0 1 truncated short Conocephalum
- Phytoliriomyza caliginosa dark yellow 2.1-2.3 8-9 yellow yellow yellow 1/1/1 yellow lobate 8 0 1 truncated short Conocephalum
- Phytoliriomyza igniculus dark yellow 2.0-2.1 6-7 yellow yellow yellow 1/1/1 yellow lobate 5 0 1 truncated short Marchantia
- Phytoliriomyza crepusculum dark yellow 1.4-1.8 5 yellow yellow yellow 1/1/1 dark lobate 5 0 0 truncated short Dumortiera
- Phytoliriomyza arcus yellow yellow 1.3-1.6 6 yellow yellow yellow 1/1/1 yellow elongated (2) 1 1 truncated short Plagiochasma
- Phytoliriomyza plagiochasmatos yellow yellow 1.4-1.5 6 yellow yellow yellow 1/1/1 yellow elongated (2) 0 1 truncated short Plagiochasma , Asterella
- Phytoliriomyza falcata yellow yellow 1.6-2.0 6-8 yellow yellow yellow 1/1/1 yellow elongated (2) 1 1 truncated short Reboulia
- Phytoliriomyza aratriformis yellow yellow 1.9-2.3 7-8 yellow yellow yellow 1/1/1 yellow elongated (2) 1 1 truncated short Reboulia
1 1, present; 0, absent; 1-1, stripes unfused; 1/1, stripes fused.<br/> 2 Number of unfused tubercle-like setae forming a row is shown in parenthesis; -, lacking tubercle-like setae forming a row.<br/> * Number of setulae in four rows.
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 x 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