The species name, achyranthesae, is based on the generic name the host plant, Achyranthes bidentata (Amaranthaceae).

Holotype: 1♂ (KUEC): reared by A. K. Elsayed from a larva obtained from a fruit gall on A. bidentata, collected from Mount Tara, Saga Prefecture, Japan, on 7.x.2015, M. Tokuda leg., the larva departed from gall between 10–19.x.2015 and the adult male emerged on 3.ix.2016. Paratypes: All paratypes were reared from fruit galls on A. bidentata in Japan. 3 larvae: collected from Tokushima City, Tokushima Prefecture on 6.x.2001, M. Yukinari leg., larvae departed from galls on 12.x.2001; 5 larvae: collected from Tokushima City, Tokushima Prefecture on 30.x.2012, J. Yukawa et al. leg., larvae departed from galls on 30.x.2012; 8♀, 4 Pupal exuviae: collected from Takeo City, Saga Prefecture on 10.x.2015, A. Kita leg., larvae departed from galls between 13–19.x.2015, adults emerged on 1.ix.2016; 3♀: collected from Mount Hinokuma, Saga Prefecture on 16.x.2014, A. K. Elsayed & M. Tokuda leg., larvae departed from galls on 22.x.2014, adults emerged in summer 2015; 4 pupal exuviae: collected from Takeo City, Saga Prefecture on 10.x.2015, larvae departed from galls between 13–19.x.2015, adults emerged on 29.viii.2016; 10♂, 6♀: same data as holotype.

Head (Fig. 5): Compound eyes separated on vertex by a diameter of 0.0–0.75 facets, eye bridge 7–8 facets long, facets hexagonal. Fronto-clypeus with 15–21 setae (n = 6). Labrum and labella setose. Palpus 4-segmented: first segment ca 34 μm, second 1.3 times as long as the first, third 1.5 as long as the second, fourth 1.3 as long as the third. Antenna: scape slightly wider than long, pedicel rounded, flagellomeres I and II partially fused; female flagellomeres I–IX cylindrical with 2 connected rings of circumfila (Fig. 6); distal flagellomeres successively shorter; flagellomere X about as long as wide; flagellomere XI rounded, slightly wider than long; flagellomere XII rudimentary, partially fused with XI; male flagellomeres cylindrical with sinuous circumfila (Fig. 7).

Figures 5–11. 

Schizomyia achyranthesae. 5 Head 6 Ventral view of female antennal flagellomere V 7 Ventral view of male flagellomere V 8 Wing 9 Tarsomere V and acromere 10 Terminal part of female abdomen 11 Ovipositor apex. Scale bars: 50 µm (6, 7, 9–11), 200 µm (5, 8).

Thorax: Wing (Fig. 8) length 1.60–2.02 mm (n = 6) in female, 1.43–1.73 mm (n = 5) in male: R₁ join C before wing midlength, R₅ join C just after wing apex, C broken after the conjunction with R₅; wing fold present; M₃₊₄ forked with Cu. Tarsal claws untoothed, bent after midlength; empodia shorter than claws, with long setulae apically; pulvilli not discernable (Fig. 9). Anepimeral setae 19–23 (n = 8); mesanepisternum scales 22–38 (n = 6); lateral scutal setae 21–28 (n = 8). Lengths of leg segments as in Suppl. material 1: Table S1.

Female abdomen (Figs 10, 11): Tergites with anterior pair of trichoid sensilla; tergites I–VII rectangular and evenly covered with scales, tergites I–VI with a posterior row of setae and some scattered setae laterally at midlength; tergite VII with 1–2 posterior rows of setae and some scattered setae laterally at midlength; tergite VIII bare, notched laterally, posterior margin with a pair of well-developed dorsal lobes. Sternites II–VI rectangular, bare and less pigmented medially, with the lateral pair of trichoid sensilla situated anterior to the sclerotized sternite, several scattered setae on the anterior half, and 1–2 rows of setae posteriorly; sternite VII about 2.6 times as long as preceding sternites, with anterior pair of trichoid sensilla laterally situated on the sternite and setae covering posterior two-thirds. Ovipositor: protrusible needlelike portion about 4 times as long as sternite VII (Fig. 10); cerci fused, each lobe with few setae (Fig. 11).

Male abdomen: Tergites I–VII as in female; tergite VIII weakly sclerotized medially, with anterior pair of trichoid sensilla. Sternites II–VI as in female; sternite VII with lateral pair of trichoid sensilla situated anterior to the sclerotized sternite, several setae scattered anteriorly, median membranous bare area and 1–2 posterior row of setae; sternite VIII setose, with lateral pair of trichoid sensilla situated intersegmentally between sterna VII and VIII. Terminalia (Fig. 12): gonocoxite extending ventrally as convex lobe beyond of gonostylus; gonostylus stout, with unfused and compressed denticles, dorsally with setae on distal third, ventrally with a cluster of setae on the basal half. Cerci notched, each cercus with 4 strong setae and a few fine setae. Hypoproct shorter than cerci and aedeagus, bilobed, with a seta on each lobe. Parameres about half length of hypoproct. Aedeagus gradually tapering, acute apex, longer than cerci.

Figures 12–17. 

Schizomyia achyranthesae. 12 Male terminalia 13 Larval spatula 14 Larval anus 15 Terminal larval segment dorsally 16 Ventral view of pupal head 17 Pupal prothoracic spiracle. Scale bars: 50 µm (12–15), 100 µm (16, 17).

Mature larva (Figs 13–15): Cylindrical, yellow. Number and position of thoracic and abdominal spiracles as in other Asphondyliini (see Möhn 1961); 6 dorsal papillae present on all thoracic segments and abdominal segments I–VII, each with a seta; abdominal segment VIII with 2 dorsal lobes, each with a setose dorsal papilla. Sternal spatula bilobed (Fig. 13) with posterior portion about 3.6 times as wide as the base of the anterior free portion. Two groups of lateral papillae on all thoracic segments, the inner group of 2 setose papillae and the outer group of 2 setose and 1 asetose papillae. Two sternal papillae on each thoracic segment and 4 sternal papillae on abdominal segments I–VIII, all without setae and situated on slight swellings. Ventral papillae each with a seta on meso- and metathoracic segments and on abdominal segments I–VII. Anus situated ventrally, with simple opening and 4 asetose anal papillae (Fig. 14). Two pairs of pleural papillae present on all thoracic segments and abdominal segment VIII, and 3 pairs on abdominal segments I–VII. Terminal segment (Fig. 15) with 2 setose and 2 corniform terminal papillae.

Pupa (Figs 16, 17): Exuviae not pigmented except prothoracic spiracles and dorsal spines of abdomen. Antennal horns short; 2 pairs of cephalic papillae present, a pair with short seta; 2 pairs of lower facial papillae, a pair with seta; 3 lateral facial papillae present on each side, 1 with short seta, 2 without setae. Prothoracic spiracle, slightly curved, 23–29 μm long (n = 6), connected with trachea to the tip. Spiracles present on abdominal segments II–VI. Abdominal terga I–VIII each with anterior pair of trichoid sensilla and 2 pleural papillae; terga I–VII each with 3 pairs of dorsal papillae, only outermost pair with a seta; tergum VIII with a pair of dorsal papillae, each with a seta. Abdominal terga II–VIII each with 3–4 rows of spines on median third.

Japan: Honshu, Shikoku, Kyushu (Yukawa and Masuda 1996) and Tanegashima Island (Yukawa et al. 2013).

Schizomyia achyranthesae induces subglobular fruit galls on A. bidentata, 5.07–5.17 mm in diameter (n = 5) (Fig. 1) [Gall No. C-246 in Yukawa and Masuda (1996)]. Based on Yukawa and Masuda (1996) and present observations, each gall contains 1–13 chambers and each chamber contains a single larva. The galls appear in September. The mature larvae leave the galls between October and November and overwinter in the soil. The adults of S. achyranthesae emerge during the flowering season of the host plant in August and September.

Schizomyia achyranthesae is distinguishable from the known Schizomyia species, except S. asteris and S. solidaginis, by its shallowly constricted male flagellomeres, lateral position of anterior pair of trichoid sensilla and presence of four larval terminal papillae, as well as two setose papillae in inner group of lateral papillae. S. achyranthesae can be separated from S. solidaginis based on the larval characters as follows: S. achyranthesae possesses a more elongated sternal spatula than S. solidaginis; the inner group of lateral papillae consists of two setose papillae in S. achyranthesae, but one setose and one asetose papillae in S. solidaginis; the anal opening is simple in S. achyranthesae, while branched in S. solidaginis. Then, S. achyranthesae can be separated from S. asteris by the following features: female cerci is less divided in S. achyranthesae; dorsal setae are present on the gonostylus in S. achyranthesae, but absent in A. asteris; and the gonocoxite is only slightly extends ventrally beyond the gonostylus in S. achyranthesae, and the larval anal opening is simple in S. achyranthesae while branched in S. asteris.

Characters given in S. achyranthesae except for the following:

The species name, diplocyclosae, is based on the generic name of the host plant, Diplocyclos palmatus (Cucurbitaceae).

Holotype: 1♂ (KUEC): reared from a larva obtained from a flower bud gall on D. palmatus, collected from Hantagawa, Naha City, Okinawa Prefecture, Japan on 10.ii.2016, T. Ganaha-Kikumura leg., emerged on 14.iii.2016. Paratypes: All paratypes were reared from flower bud galls on D. palmatus in Japan. 4 larvae: collected from Gogayama, Nakijin Village, Okinawa Prefecture on 4.iii.2002, M. Tokuda leg., departed from galls on 9.iii.2002; 2 larvae: collected from Kinjo cho, Naha City, Okinawa Prefecture on 13.i.1977, S. Yamauchi leg.; 4 pupal exuviae: collected from Shuri, Naha City, Okinawa Prefecture, emerged in February 1977, S. Yamauchi leg.; 2 pupal exuviae, 3♂, 5♀: same data as holotype; 5 pupal exuviae, 3♂, 2♀: collected from Hantagawa, Naha City, Okinawa Prefecture on 10.ii.2016, T. Ganaha-Kikumura leg., emerged on 15.iii.2016; 1 pupal exuviae: collected from Hantagawa, Naha City, Okinawa Prefecture on 10.ii.2016, T. Ganaha-Kikumura leg., emerged on 16.iii.2016; 1 pupal exuviae: collected from Hantagawa, Naha City, Okinawa Prefecture on 10.ii.2016, T. Ganaha-Kikumura leg., emerged on 21.iii.2016.

Head (Fig. 18): Fronto-clypeus with 17–24 setae (n = 6). Palpus: first segment ca 38.3 μm, second 1.4 times as long as the first, third 1.3 as long as the second, fourth 1.5 as long as the third.

Figures 18–24. 

Schizomyia diplocyclosae. 18 Head 19 Ventral view of female flagellomere V 20 Dorsal view of male flagellomere V 21 Wing 22 Tarsomere V and acromere 23 Terminal part of female abdomen 24 Ovipositor apex Scale bars: 100 µm (18, 19), 50 µm (20–24).

Thorax: Wing (Fig. 21) length 2.15–2.26 mm (n = 5) in female, 1.70–2.12 (n = 5) in male. Empodia slightly longer than tarsal claws (Fig. 22). Anepimeral setae 11–18 (n = 8); mesanepisternum scales 20–38 (n = 8); lateral scutum setae 26–48 (n = 8). Lengths of leg segments as in Suppl. material 1: Table S1.

Female abdomen (Figs 23, 24): Posterior margin of tergite VIII with a pair of slightly developed dorsal lobes. Sternite VII about 2.5 times as long as preceding sternites. Ovipositor: distal protrusible needle-like portion about 3 times as long as sternite VII.

Male abdomen: Terminalia (Fig. 25): Gonostylus dorsally with several setae on distal half.

Figures 25–29. 

Schizomyia diplocyclosae. 25 Male terminalia 26 Larval spatula 27 Terminal larval segments dorsally 28 Ventral view of pupal head 29 Prothoracic spiracle in pupa. Scale bars: 50 µm (25–27), 100 µm (28, 29).

Mature larva: Sternal spatula with posterior portion about 2.8 times as wide as the base of the anterior free portion (Fig. 26). Larval anus with 2 asetose anal papillae.

Pupa (Figs 28, 29): Prothoracic spiracle 280–310 μm long (n = 6).

Japan: Kikaijima Island and Okinawa-honto Island (Yukawa and Masuda 1996).

Schizomyia diplocyclosae induces subglobular and pale green flower bud galls on D. palmatus, about 6–10 mm in diameter. Each gall consists of 10–45 chambers and each chamber contains a single larva [Gall No. C-409 in Yukawa and Masuda (1996)]. Galls become mature between December and March and larvae depart from galls to drop to the ground. The adults of S. diplocyclosae emerge in February and March when the larvae were reared under laboratory temperature (Yukawa and Masuda 1996, present data). Similar flower bud galls were found on Melothria liukiuensis Nakai (Cucurbitaceae) and considered to be induced by this species or a closely related one (Yamauchi et al. 1982, Yukawa and Masuda 1996).

Schizomyia diplocyclosae is morphologically very similar to S. achyranthesae but differs from it by the following characters: S. diplocyclosae has a shorter ovipositor (protrusible needle-like-portion three times as long as sternite VII while four times as long in S. achyranthesae), less developed dorsal lobes on the posterior margin of female tergite VIII, gonocoxite more pointed posteroapically, empodia longer than claws and larva with only two anal papillae (four in S. achyranthesae).

Characters as in S. achyranthesae except for the following:

The species name, castanopsisae, is based on the generic name of the host plant, Castanopsis sieboldii (Fagaceae).

Holotype: 1♂ (KUEC): reared from a larva obtained from an inflorescence gall on C. sieboldii by A. K. Elsayed, collected from Hachijojima Island on 6.xii.2014, T. Kikuchi leg., emerged on 24.ii.2015. Paratypes: All paratypes were reared by A. K. Elsayed from inflorescence galls on C. sieboldii in Japan. 2 larvae: collected from Hachijojima Island on 6.xii.2014, T. Kikuchi leg., departed from galls on 22.xii.2014; 5 larvae: collected from Hachijojima Island on 6.xii.2014, T. Kikuchi leg., departed from galls on 25.xii.2014; 6 pupal exuviae, 2♀, 3♂: collected from Shikinejima Island on 10.xii.2014, M. Tokuda leg., emerged between 24.i–20.ii.2015; 3 pupal exuviae, 4♀, 2♂: collected from Hachijojima Island on 6.xii.2014, T. Kikuchi leg., emerged between 20.ii–5.iii.2015.

Head (Fig. 30): Fronto-clypeus with 10–16 setae (n = 6). Palpus: first segment ca 43 μm, second 1.4 times as long as the first, third 1.5 as long as the second, fourth 1.2 as long as the third.

Figures 30–36. 

Schizomyia castanopsisae. 30 Head 31 Ventral view of female flagellomere V 32 Ventral view of male flagellomere V 33 Wing 34 Tarsomere V and acromere 35 Terminal part of female abdomen 36 Ovipositor apex. Scale bars: 50 µm (31, 32, 34–36), 100 µm (30, 33).

Thorax: Wing (Fig. 33) length 2.04–2.74 mm (n = 6) in female, 2.04–2.56 mm (n = 4) in male. Anepimeral setae 8–15 (n = 8); mesanepisternum scales 15–26 (n = 7); lateral scutum setae 15–27 (n = 8). Lengths of leg segments as in Suppl. material 1: Table S2.

Female abdomen (Figs 35, 36): Sternite VII about 3 times as long as preceding. Ovipositor: protrusible needle-like portion about 3.3 as long as sternite VII.

Male abdomen: Terminalia (Figs 37, 38): Gonocoxite length about 3.3 times as long as gonostylus.

Figures 37–42. 

Schizomyia castanopsisae. 37 Male terminalia 38 Male cerci 39 Larval spatula 40 Larval anus 41 Terminal larval segments dorsally 42 Ventral view of pupal head and thorax. Scale bars: 50 µm (37–41), 100 µm (42).

Mature larva: Sternal spatula (Fig. 39) with posterior portion about 2.8 times as wide as the base of the anterior free portion. Anus opening branched (Fig. 40).

Pupa (Fig. 42): Prothoracic spiracle 280–330 μm long (n = 6).

Japan: The Izu Islands (from Niijima to Aogashima) (Tokuda et al. 2012a, b, 2013, 2015, Tokuda and Kawauchi 2013a), Kyushu (Kagoshima and Miyazaki Prefectures) (Nagai 2010, Tokuda and Kawauchi 2013b), Tanegashima Island (Yukawa et al. 2013) and Okinawa-honto Island (Yamauchi et al. 1982).

Castanopsis sieboldii inflorescences galled by S. castanopsisae are rather irregularly swollen, 5.7–15.7 mm in diameter and 6.2–30.9 mm in length (Fig. 2) [Gall No. C-163 in Yukawa and Masuda (1996)]. Each gall consists of up to 30 chambers and each chamber contains a single larva. Mature larvae of S. castanopsisae left the galls collected in the Izu Islands in December within few days after the collection. Larvae were kept with soil in the laboratory and adults emerged the following January, February and March.

Schizomyia castanopsisae is morphologically close to S. asteris, S. achyranthesae and S. diplocyclosae. Schizomyia castanopsisae can be separated from S. asteris by a shorter ovipositor (protrusible needle-like portion 3.3 times as long as sternite VII, while 5.7 times in S. asteris), the presence of dorsal setae on the gonostyli, and the tooth of gonostylus, which extends more dorsally than in that of S. asteris; from S. achyranthesae by a shorter ovipositor (four times as long as sternite VII in S. achyranthesae), more posteroapically pointed gonocoxite, and branched anal opening of larva; and from S. diplocyclosae by shorter empodia than tarsal claws (empodia are as long as claws in S. diplocyclosae) and the number of larval anal papillae (four in S. castanopsisae while two in S. diplocyclosae).

Characters as in S. achyranthesae except for the following:

The species name, usubai, honors the late Mr Shigeshi Usuba who reared adults of this species for the first time.

Holotype: 1♂ (KUEC): reared by A. K. Elsayed from a larva obtained from a fruit gall on T. asiaticum, collected from Torinosu, Tanabe City, Wakayama Prefecture, Japan, I. Matoba leg., emerged on 22.v.2017. Paratypes: All paratypes were reared from fruit galls on T. asiaticum in Japan. 4 larvae: collected from Mount Takakuma, Kagoshima Prefecture in 1969, J. Yukawa leg.; 4 larvae: galls collected from Imuta Lake-side, Kedouin, Satsuma-sendai City, Kagoshima Prefecture on 2.xi.1978, S. Sako leg.; 4 pupal exuviae, 2♂, 2♀: collected from Torinosu, Tanabe City, Wakayama Prefecture, I. Matoba leg., reared by A. K. Elsayed, emerged on 18.v.2017; 2 pupal exuviae, 1♀, 2♂: same data as holotype.

Head (Fig. 43): Compound eyes separated on vertex by a diameter of 0.0–0.5 facets. Fronto-clypeal setae 15–16 setae (n = 4). Palpus: first segment ca 53.5 μm, second about as long as the first, third 1.6 as long as the second, fourth 1.4 as long as the third.

Figures 43–49. 

Schizomyia usubai. 43 Head 44 Ventral view of female flagellomere V 45 Ventral view of male flagellomere V 46 Wing 47 Tarsomere V and acromere 48 Terminal part of female abdomen 49 Ovipositor apex. Scale bars: 50 µm (44, 45, 47, 49), 100 µm (43, 46, 48).

Thorax: Wing (Fig. 46) length 2.03–2.34 mm (n = 4) in female, 1.80–1.95 mm (n = 3) in male. Empodia as long as claws (Fig. 47). Anepimeral setae 10–20 (n = 4); mesanepisternum scales 17–40 (n = 4); lateral scutum setae 19–27 (n = 4). Lengths of leg segments as in Suppl. material 1: Table S2.

Female abdomen (Figs 48, 49): Sternite VII about 2.6 times as long as preceding. Ovipositor: protrusible needle-like portion about 4.5 as long as sternite VII.

Male abdomen: Terminalia (Fig. 50): Gonocoxite with developed, pointed apical lobe extending beyond gonostylus.

Figure 50–55. 

Schizomyia usubai. 50 Male terminalia 51 Larval spatula 52 Larval anus 53 Terminal larval segments dorsally 54 Ventral view of pupal head 55 Pupal prothoracic spiracle. Scale bars: 50 µm (50–53), 100 µm (54, 55).

Mature larva: Sternal spatula (Fig. 51) with posterior portion about 3.5 times as wide as the base of the anterior free portion. Anus with branched opening (Fig. 52).

Pupa (Figs 54, 55): Prothoracic spiracle 250–350 μm long (n = 6).

Japan: The Izu Islands (Tokuda et al. 2012b, 2013, Tokuda and Kawauchi 2013) Honshu, and Kyushu (Yukawa and Masuda 1996).

The normal fruit of Trachelospermum asiaticum (Apocynaceae) is V-shaped, consisting of a pair of very long and thin seed pods. When the fruits are galled by S. usubai, the apical parts of the fruit become fused and swollen, more or less cat-bell shaped (Fig. 3), about 12–18 mm in diameter and 27 mm in length [Gall No. D-033 in Yukawa and Masuda (1996)]. Each gall consists of 10–25 chambers and each chamber contains 10–25 larvae. Galls mature between late September and October and the larvae depart from galls to overwinter in soil. The adults of S. usubai emerge between late April and July (Yukawa 1978; Yukawa and Masuda 1996; present study). Similar galls probably induced by this species were found on Trachelospermum gracilipes Hook. f. var. kiukiuense (Hatus.) Kitam. on Tanegashima Island (Yukawa et al. 2013).

Schizomyia usubai is close to S. asteris, S. achyranthesae, S. diplocyclosae and S. castanopsisae. Schizomyia usubai can be distinguished from S. asteris by a shorter ovipositor (protrusible needle-like portion about 4.5 times as long as sternite VII, while 5.5 times in S. asteris), longer empodia, and the presence of dorsal setae on gonostyli; from S. achyranthesae and S. diplocyclosae by a longer ovipositor (four and three times as long as sternite VII in S. achyranthesae and S. diplocyclosae, respectively), longer empodia, and branched opening of the larval anus. In addition, larva of S. usubai has four anal papillae, but two in S. diplocyclosae. Schizomyia castanopsisae is very similar to S. usubai, but can be separated by a shorter ovipositor (protrusible needle-like portion about 3.3 times as long as sternite VII, while 4.5 times in S. usubai), longer empodia, and less compressed circumfila of female flagellomeres.

Characters as in S. achyranthesae except for the following:

The species name, paederiae, is based on the generic name of the host plant, Paederia foetida (Rubiaceae).

Holotype: 1♂ (KUEC): reared from a larva obtained from a flower bud gall on P. foetida, collected from Misawa, Ogori City, Fukuoka Prefecture, Japan, K. Matsunaga leg., emerged between 11–15.viii.2017. Paratypes: All paratypes were reared from flower bud galls on P. foetida in Japan. 11 larvae: collected from Nishinoomote, Nishinoomote City, Kagoshima Prefecture, on 24.ix.2014, K. Ogata leg.; 4 pupal exuviae, 2♂, 7♀: same data as holotype.

Head (Fig. 56): Compound eyes separated on vertex by a diameter of 0.0–1.5 facets, eye bridge consist of 6–7 facets long. Fronto-clypeus with 11–13 setae (n = 4). Palpus: first segment ca 28.6 μm, second 1.3 times as long as the first, third 1.4 as long as the second, fourth 1.2 as long as the third.Male flagellomeres with deep basal constriction and elongated necks (Fig. 58).

Figures 56–62. 

Schizomyia paederiae. 56 Head 57 Dorsal view of female flagellomere V 58 Dorsal view of female flagellomere V 59 Wing 60 Tarsomere V and acromere 61 Terminal part of female abdomen 62 Ovipositor apex. Scale bars: 50 µm (57, 58, 60, 62), 100 µm (56, 59, 61).

Thorax: Wing (Fig. 59) length 1.16–1.57 mm (n = 5) in female, 1.04–1.36 mm (n = 3) in male. Anepimeral setae 9 or 10 (n = 5); mesanepisternum scales 5–10 (n = 6); lateral scutum setae 23–28 (n = 5). Empodia as long as tarsal claws (Fig. 60). Lengths of leg segments as in Suppl. material 1: Table S3.

Female abdomen (Figs 61, 62): Anterior pair of trichoid sensilla situated medially on abdominal sternites II–VI; sternite VII about 3.4 times as long as preceding sternites. Ovipositor: protrusible needle-like portion about 4.8 times as long as sternite VII.

Male abdomen: Anterior pair of trichoid sensilla situated medially on sternites II–VI and laterally on sternite VIII, sternite VIII with scattered setae. Terminalia (Fig. 63): Gonostylus dorsally with several setae on distal half, with unfused and compressed denticles.

Figures 63–67. 

Schizomyia paederiae. 63 Male terminalia 64 Larval spatula 65 Terminal larval segments dorsally 66 Ventral view of pupal head 67 Pupal prothoracic spiracle. Scale bars: 50 µm (63–65), 100 µm (66, 67).

Mature larva: Abdominal segment VIII with 2 setose dorsal papillae. Posterior portion of sternal spatula about 3.3 times as wide as the base of the anterior free portion (Fig. 64); 2 groups of lateral papillae present on all thoracic segments, each consisting of 2 setose and 1 asetose papillae. Terminal segment with 8 terminal papillae, consisting of 4 setose, 2 asetose and 2 corniform ones (Fig. 65).

Pupa (Figs 66, 67): Prothoracic spiracle 230–290 μm long (n = 4).

Japan: Honshu, Shikoku, Kyushu, and Yakushima Island (Yukawa and Masuda 1996).

Schizomyia paederiae induces flower bud galls on P. foetida. Basal parts of the galled flower buds are swollen, 3.0–5.6 mm in diameter and 4.0–6.1 mm in length (Fig. 4) [Gall No. D-037 in Yukawa and Masuda (1996)]. Galls are single-chambered and each gall contains 1–10 larvae. The larvae depart from mature galls from late August to September and overwinter in the soil. The adults of S. paederiae emerge in early August when the flower buds are available on the host plant (Yukawa and Masuda 1996).

Schizomyia paederiae is distinguishable from other Schizomyia species, except four Russian species, i.e. S. calathidiphaga, S. clematidis, S. spiraeae, and S. veronicastrum, by its deeply constricted male flagellomeres (Kovalev 1964; Fedotova 2002). Firstly, the adults of S. paederiae differs from S. calathidiphaga by a slightly longer ovipositor (protrusible needle-like portion about 4.8 times as long as sternite VII, while 4.5 times in S. calathidiphaga), longer empodia (empodia are as long as claws in S. paederiae, but shorter in S. calathidiphaga), the position of gonostylus tooth (mostly covering only the apical margin in S. paederiae, but on the posteroapical margin in S. calathidiphaga), and the arrangement of papillae on the larval terminal segment (the two asetose terminal papillae are situated more posteriorly in S. paederiae, while more anteriorly in S. clathidiphaga). Then, the adults of S. paederiae can be separated from S. clematidis, S. spiraeae and S. veronicastrum by a longer neck of male flagellomere III, which is about 0.25 as long as node in S. paederiae but about 0.15 as long as node in other species, the position of gonostylus tooth (mostly covering the apical margin in S. paederiae, but on the posteroapical margin in the other species), and a much narrower hypoproct than S. clematidis.

Characters as in S. achyranthesae except for the following:

2♂, 3♀ (Mamaev collection: slide no. B1-251369): collected from Rybatskij, Lithuania on 19.vii.1969; 1♀, 1 pupal exuviae (J. J. Kieffer’s specimen in Felt collection).

Head: Compound eyes with rounded facets; facets on vertex and eye bridge unobservable because the specimens mounted laterally. Palpus: first segment ca 23.4 μm, second 1.6 times as long as the first, third 1.4 as long as the second, fourth 1.4 as long as the third.

Figures 68–73. 

Schizomyia galiorum. 68 Lateral view of female flagellomere V 69 Lateral view of male flagellomere V 70 Wing 71 Tarsomere V and acromere 72 Terminal part of female abdomen showing the ovipositor 73 Ovipositor apex. Scale bars: 50 µm (68, 69, 71, 73), 100 µm (70, 72).

Thorax: Wing (Fig. 70) length 1.33–1.55 mm (n = 2) in male, 1.93 mm (n = 1) in female; R₅ joining C just before wing apex. Empodia as long as claws (Fig. 71). Anepimeral setae 9–15 (n = 5); mesanepisternum scales 14–18 (n = 4); lateral scutum setae 17–37 (n = 5). Lengths of leg segments as in Suppl. material 1: Table S3.

Female abdomen: Posterior margin of tergite VIII without dorsal lobes. Sternites with median pair of trichoid sensilla. Sternite VII about twice as long as VI. Ovipositor (Figs 72, 73): protrusible needle-like portion short, about 1.9 times as long as sternite VII; cerci fused, with few fine setae (Fig. 73).

Male abdomen: Sternites with median pair of trichoid sensilla. Sternite VII with two posterior rows of setae. Terminalia (Fig. 74): Gonocoxite with slightly developed apical lobe. Gonostylus with several setae on distal half dorsally, with group of setae on the basal half ventrally, and with distinctive unfused denticles. Hypoproct slightly longer than cerci.

Figure 74–76. 

Schizomyia galiorum. 74 Male terminalia 75 Ventral view of pupal head 76 Pupal prothoracic spiracle. Scale bars: 50 µm (74), 100 µm (75, 76).

Mature larva: Sternal spatula bilobed, the anterior free portion slightly wider than the posterior portion (Kieffer 1889). Two groups of lateral papillae present on each side of the spatula, each group of 2 setose and 1 asetose papillae. Terminal segment with 6 setose and 2 corniform terminal papillae (Möhn 1955).

Pupa (Figs 75, 76): Antennal horns slightly developed. Prothoracic spiracle 240 μm long (n = 1).

Widespread Europe, Algeria and Kazakhstan (Gagné and Jaschhof 2017).

This species is distinguished from eastern Holarctic congeners by the distinctly short ovipositor, the absence of dorsal lobes on the posterior margin of female tergite VIII, and the conjunction of wing vein C with R₅ before wing apex.

Characters as in S. achyranthesae except for the following:

All obtained from flower bud galls on Patrinia villosa (Valerianaceae) in Japan. 6 larvae: collected from Maruyama, Ojiya City, Niigata Prefecture on 12.x.1981, K. Yamagishi leg. 1♂: Iozan, Kanazawa City, Ishikawa Prefecture on 17.x.1978, emerged on 22.iv.1979, J. Yukawa leg. 2♂, 4♀, 2 pupal exuviae: collected from Kyuragi, Karatsu City, Saga Prefecture on 12.x.2015, M. Tokuda leg., emerged on 23.viii.2016, reared by A. K. Elsayed. 2♂, 3♀, 2 pupal exuviae: collected from Kyuragi, Karatsu City, Saga Prefecture on 12.x.2015, M. Tokuda leg., emerged on 28.viii.2016, reared by A. K. Elsayed. 2♂: collected from Kyuragi, Karatsu City, Saga Prefecture on 12.x.2015, M. Tokuda leg., emerged on 29.viii.2016, reared by A. K. Elsayed.

Head (Fig. 77): Compound eyes separated on vertex by a diameter of 0.25–1 facets, eye bridge consist of 5–6 facets long. Fronto-clypeus with 15–20 setae (n = 9). Palpus: first segment ca 34.1 μm, second 1.4 times as long as the first, third 1.4 as long as the second, fourth segment 1.5 as long as the third.

Figures 77–83. 

Schizomyia patriniae. 77 Head 78 Dorsal view of female flagellomere V 79 Ventral view of female flagellomere V 80 Wing 81 Tarsomere V and acromere 82 Terminal part of female abdomen 83 Ovipositor apex. Scale bars: 50 µm (78, 79, 81, 83), 200 µm (77, 80, 82).

Thorax: Wing (Fig. 80) length 1.83–2.09 mm (n = 5) in female, 1.50–1.83 mm (n = 5) in male. Anepimeral setae 17–23 (n = 8); mesanepisternum scales 16–20 (n = 7); lateral scutum setae 32–46 (n = 8). Lengths of leg segments as Suppl. material 1: Table S4.

Female abdomen (Figs 82, 83): Median pair of trichoid sensilla present on sternites II–VII. Sternite VII about 3 times as long as preceding. Ovipositor: protrusible needle-like portion about 4.8 as long as sternite VII.

Male abdomen: Trichoid sensilla present on sternites II–VIII in median position, except on VIII in lateral position. Terminalia (Fig. 84): Gonocoxite length about 3.3 times as long as gonostylus. Gonostylus with slim tooth in the dorsal and ventral views.

Figures 84–85. 

Schizomyia paederiae. 84 Male terminalia 85 Ventral view of pupal head and prothorax. Scale bars: 50 µm (84), 200 µm (85).

Mature larva: Abdominal segment VIII without dorsal papillae. Two groups of lateral papillae on all thoracic segments, each consisting of 2 setose and 1 asetose papillae. The terminal segment with 1 setose and 6 asetose terminal papillae (Yukawa 1983).

Pupa (Fig. 85): Prothoracic spiracle 260–290 μm long (n = 5).

Japan: Hokkaido, Honshu and Shikoku (Yukawa and Masuda 1996).

This species had been described by Shinji (1938) under the genus Schizomyia. Then, Yukawa (1983) combined the species with Asteralobia because of its shallowly constricted male flagellomeres. Because Asteralobia is synonymized under Schizomyia in this paper, S. patriniae is combined again with Schizomyia.

Schizomyia patriniae is distinguishable from known Schizomyia species, except three species that were previously treated as Asteralobia and newly combined here under Schizomyia, i.e. S. sasakii, S. soyogo, and S. humuli, by the presence of shallow constrictions on male flagellomeres and the absence of corniform papillae on the terminal larval segment. S. patriniae can be easily separated from S. sasakii, S. soyogo and S. humuli based on the number of papillae on the larval terminal segment: S. patriniae possesses two setose and six asetose terminal papillae, but S. sasakii and S. soyogo have only six setose terminal papillae, and S. humuli has four setose terminal papillae.

Characters as in S. achyranthesae except for the following:

2♂, 2♀: (Mamaev collection: slide no. B1-251363 & 251364), galls collected from Aster sp. in Kedrovaja Pad reserve, Russian Far East on 28.viii.1964. 4 larvae: galls collected from A. tataricus in Smolyaninovo, Primorsky Territory, Russian Far East on 13.ix.2002, M. Tokuda leg. 6 larvae: galls collected from A. scaber in Smolyaninovo, Primorsky Territory, Russian Far East on 13.ix.2002, M. Tokuda leg.

Head: Compound eyes with rounded facets; facets on the vertex and eye bridge unobservable. Palpus: first segment ca 34.5 μm, second 1.8 times as long as the first, third 1.2 as long as the second, fourth 1.1 as long as the third.

Figures 86–93. 

Schizomyia asteris. 86 Lateral view of female flagellomere V 87 Lateral view of male flagellomere V 88 Tarsomere V and acromere 89 Ovipositor apex 90 Male terminalia 91 Larval spatula 92 Terminal larval segments dorsally 93 Larval anus. Scale bar: 50 µm.

Thorax: Wing length 1.93 mm (n = 1) in female, 1.55–2.02 mm (n = 2) in male. Anepimeral setae 14–15 (n = 3); mesanepisternum scales 17–23 (n = 7); lateral scutum setae 23–29 (n = 4). Lengths of leg segments as in Suppl. material 1: Table S4.

Female abdomen: Ovipositor: protrusible needle-like portion about 5.7 times as long as sternite VII; cerci divided medially, with sclerotized margins and few setae (Fig. 89).

Male abdomen: Terminalia (Fig. 90): Gonocoxite with pointed apical lobe extending beyond gonostylus. Gonocoxite length about 3.5 times as long as gonostylus. Gonostylus without setae ventrally and dorsally.

Mature larva: Sternal spatula with posterior portion about 3.3 times as wide as the base of the anterior free portion (Fig. 91). Abdominal segment VIII with three dorsal lobes, the outer two each with 1 setose dorsal papilla (Fig. 92). Anus with branched opening, and 4 asetose anal papillae (Fig. 93).

Pupa: Prothoracic spiracle about 220 μm long (n = 1).

Russian Far East (Kovalev 1964; Tokuda et al. 2003).

Although Kovalev (1964) mentioned that S. asteris is closest to S. doellingeriae, Tokuda et al. (2003) showed that they are distinguishable by the number of lateral papillae, terminal papillae, and the shape of anal opening.

Figures 94–95. 

Dorsal view of pupal abdominal segments. 94 Schizomyia soyogo 95 Schizomyia sasaki. Scale bar: 100 µm.

Figure 96. 

A phylogenetic reconstruction based on partial sequences of cytochrome oxidase subunit I (COI) and 12S small ribosomal subunit genes. The topology and branch length were produced by the maximum likelihood method (note the scale bar). Bootstrap values are indicated at branches gaining more than 50% support (10³ replications).