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Research Article
A review of the Strongylophthalmyia coarctata subgroup (Diptera, Brachycera, Strongylophthalmyiidae) from China, with the descriptions of three new species
expand article infoJiale Zhou, Neal L. Evenhuis§, Ding Yang
‡ China Agricultural University, Beijing, China
§ Department of Natural Sciences, Bernice Pauahi Bishop Museum, Honolulu, United States of America
Open Access

Abstract

The species belonging to the Strongylophthalmyia coarctata subgroup of the S. punctata group (Diptera: Brachycera: Strongylophthalmyiidae) from China are reviewed. Six species are recognized, including three new species: S. corniculata sp. nov., S. flagellicornis sp. nov., and S. tangwangana sp. nov. Strongylophthalmyia narwhal Evenhuis, 2020 and S. raricornis Shatalkin, 1981 are recorded from China for the first time, and S. raricornis is also recorded from South Korea for the first time. An identification key to the Asian species of the S. coarctata subgroup is provided.

Key words

Acalyptratae, Diopsoidea, East Asia, Oriental Realm, Palearctic Realm, taxonomy

Introduction

The acalyptrate family Strongylophthalmyiidae currently contains 90 species divided into two genera: the monotypic Nartshukia Shatalkin, 1993 from Vietnam, and the speciose Strongylophthalmyia Heller, 1902 from the Australasian, Nearctic, Oriental, and Palearctic realms (Evenhuis 2016, 2020; Galinskaya and Shatalkin 2018). Members of Strongylophthalmyia are mostly assigned into four species-groups (S. crinita group, S. fascipennis group, S. punctata group, and S. ustulata group), with nearly 40% of species belonging to the S. punctata group, while the other 12 species are unplaced and two are treated as nomen dubia (Shatalkin 1996; Evenhuis 2016). The S. punctata group is characterized by the highly modified male antennal first flagellomere having a variable process, and is further divided into the S. coarctata subgroup (10 spp.) and S. punctata subgroup (25 spp.) (Shatalkin 1996; Evenhuis 2016).

The Chinese species belonging to the S. coarctata subgroup are reviewed in the present study. Prior to this study, only one species of this subgroup, S. coarctata Hendel, 1913, had been recorded from China (Yang and Wang 1998; Qilemoge et al. 2021). Here we describe three new species, and record two further species for the Chinese fauna and one new to the South Korean fauna. A key to the species of the S. coarctata subgroup from Asia is provided.

Materials and methods

Specimens examined or cited in this study are deposited in the following institutions:

NHMUK Natural History Museum, London, UK;

CAU Entomological Museum of China Agricultural University, Beijing, China;

CSCA California State Collection of Arthropods, California Department of Food & Agriculture, Sacramento, USA;

QSBG Queen Sirikit Botanic Garden, Chiang Mai, Thailand;

SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany;

ZMUM Zoological Museum of Moscow State University, Moscow, Russia.

Male terminalia were prepared by macerating the apical portion of the abdomen in heated 10% KOH solution for approximately 10 min, and then rinsing in distilled water. External structure and terminalia were examined using a Nikon SMZ745 stereoscopic microscope. After examination, the terminalia were transferred to fresh glycerin and stored in microvials pinned below the corresponding specimens.

Photographs were taken using a Canon 7D Mark II digital camera with a Canon micro lens MP-E 65 mm for habitus, and an Olympus BX51 microscope for terminalia. Figures were stacked with Helicon Focus v. 5.3 and assembled by Adobe Photoshop 2020. The distribution map was prepared using the online version of SimpleMappr (Shorthouse 2010). Terminology follows Evenhuis (2016) and Lonsdale (2020); we use the term “phallic plate” following Lonsdale (2020) to refer the structure posteroventral to the basiphallus, although the homology of this structure is arguable. Measurements were obtained using a calibrated micrometer. Distributional data are given from specimens examined in this study and from literature records; new distributional records are marked by asterisk (*); data from literature records (without specimen examination) are given the source.

Taxonomy

Strongylophthalmyia Heller, 1902

Strongylophthalmus Hendel, 1902: 179. Type species: Chyliza ustulata Zetterstedt, 1847, by original designation. Preoccupied by Strongylophthalmus Mannheim, 1853 (Insecta: Coleoptera).

Strongylophthalmyia Heller, 1902: 226. Replacement name for Strongylophthalmus Hendel, 1902.

Labropsila Meijere, 1914: 24. Type species: Labropsila polita Meijere, 1914, by subsequent designation (Hennig 1941: 36). Synonymized by Meijere (1916: 87).

Notes

For an extensive treatment of the family, systematic considerations, definitions, a key to species-groups, and a checklist of global species, see Evenhuis (2016) and Lonsdale (2020).

The Strongylophthalmyia coarctata subgroup

Diagnosis. Recognized within the genus by the following characters: antennal first flagellomere of male modified, having variable antennal process; arista bare; fore femur of male simple, lacking thorn-like spicules dorsally.

Diversity and distribution. This subgroup currently contains ten described species, most of which are distributed in the Oriental Realm, while one species occurs in the northeastern Palearctic and three species are endemic to Papua New Guinea (Evenhuis 2016, 2020). Strongylophthalmyia coarctata Hendel was previously known from China (Taiwan). Five species, including 3 described herein, are now added to the Chinese fauna.

Key to species of Strongylophthalmyia coarctata subgroup from Asia (based on males)

1 Antennal first flagellomere bifid, with dorsal and ventral processes (Fig. 38) 2
Antennal first flagellomere with a single process below insertion of arista (Figs 1, 6, 17, 27, 49) 3
2 Antennal processes short and stout, distinctly shorter than arista (Fig. 38); mesonotum with 7 dorsocentral setae (Fig. 39); femora blackish brown (Fig. 34) Strongylophthalmyia raricornis Shatalkin
Antennal processes extremely long and thin, distinctly longer than arista; mesonotum with 1 dorsocentral seta; femora yellow, hind femur with a dark brown subapical ring Strongylophthalmyia mekistocera Evenhuis
3 Antennal process short, distinctly shorter than arista (Figs 6, 49) 4
Antennal process long, distinctly longer than arista (Figs 1, 17, 27) 6
4 Frons yellowish brown in anterior half (Fig. 4); antennal first flagellomere with a short, conical process (Fig. 6) Strongylophthalmyia corniculata sp. nov.
Frons entirely black; first antennal flagellomere with a small, blunt process (Fig. 49) 5
5 Antennal first flagellomere yellowish brown with dorsal half dark brown; hind tibia darkened in middle Strongylophthalmyia gibbifera Shatalkin
Antennal first flagellomere yellowish brown (Fig. 49); hind tibia largely dark brown (Fig. 45) Strongylophthalmyia tangwangana sp. nov.
6 Antennal process slender, wipe-like, placed dorsally on first flagellomere in close proximity to arista (Figs 1, 17) 7
Antennal process thick, sword-like, placed anteriorly on first flagellomere (Fig. 27) Strongylophthalmyia narwhal Evenhuis
7 Frons partly to entirely yellow; antennal process covered with dense black setulae (Fig. 17); wing with dark suffusion at apex (Fig. 19) 8
Frons entirely black; antennal process covered with dense whitish setulae (Fig. 1); wing fully hyaline 9
8 Frons black with anterior part yellow; antennal first flagellomere bicolorous, ovate; wing with median transverse band at level of dm-m Strongylophthalmyia freidbergi Shatalkin
Frons entirely yellow (Fig. 15); antennal first flagellomere yellow, subrhombic (Fig. 17); wing without median transverse band at level of dm-m (Fig. 19) Strongylophthalmyia flagellicornis sp. nov.
9 Antennal first flagellomere ovate (Fig. 1); postpronotum and propleuron partly yellowish brown (Figs 1, 2) Strongylophthalmyia coarctata Hendel
First antennal flagellomere subrhombic; thorax entirely black Strongylophthalmyia stylocera Shatalkin

Strongylophthalmyia coarctata Hendel, 1913

Figs 1, 2

Strongylophthalmyia coarctata Hendel, 1913: 87 (protologue); Hennig (1940: 309) (in key, figure); Steyskal (1971: 142) (in key); Rohlfien and Ewald (1972: 413) (type material, record); Steyskal (1977: 21) (catalogue, distribution); Yang and Wang (1998: 458) (in key, distribution, record); Lin and Chen (1999: 68) (listed); Papp (2005: 195) (record); Shatalkin (2007: 560) (type material, redescription); Iwasa and Evenhuis (2014: 103) (listed, distribution); Evenhuis (2016: 207) (listed, distribution); Qilemoge et al. (2021: 236) (catalogue, distribution). Syntypes (2♂4♀): China, Taiwan, Hengchun, SDEI.

Material examined

China. Taiwan: Pingtung, Manzhou, Kankau [= Gangkou], 1912.iv, leg. H. Sauter (1♂, NHMUK).

Diagnosis

Generally shiny black (Figs 1, 2); antennal first flagellomere of male yellow, ovate (Fig. 1), with a long slender process covered with dense white setulae (Figs 1, 2); mesonotum blackish, postpronotum and propleuron partly yellowish brown (Figs 1, 2); wing hyaline (Fig. 2); mid and hind femora yellow, with apex dark brown (Figs 1, 2); distiphallus with small apical “glans” (see Hennig 1940: fig. 11).

Figures 1, 2. 

Strongylophthalmyia coarctata Hendel, 1913, non-type male, habitus: 1 dorsal view 2 lateral view. Scale bar: 1 mm.

Distribution

China – Taiwan: Chiayi (Papp 2005), Kaohsiung (Rohlfien and Ewald 1972), Pingtung (Fig. 56).

Remarks

This species was originally described based on “Mehrere ♀, 2♂” (syntype) from “Kankau (Koshun)” (now Hengchun, Pingtung, Taiwan, China) (Hendel 1913). According to Rohlfien and Ewald (1972), the type material of this species was preserved in the collection of SDEI and included two males and four females. Shatalkin (2007) redescribed this species based on a male and a female which he referred as “holotype” and “paratype”, respectively, and mentioned that these specimens were deposited in the Museum für Naturkunde, Berlin, Germany (ZMHB). However, no such specimens could be found in the collection of ZMHB (J. Pohl, pers. comm.), and the collection of SDEI was unavailable to us during the present study. We therefore presume that the type material of this species is still kept in SDEI, and Shatalkin (2007) made an incorrect interpretation on its depository. The uses of the term “holotype” and “paratype” by Shatalkin (2007) did not constitute valid lectotype designations (ICZN 1999, Art. 74.7), therefore the status of the type specimens of this species should remain as syntypes.

During the present study, we have examined a male from “Kankau” (now Gangkou, Pingtung, Taiwan, China) in the collection of NHMUK. Images of the dorsal and lateral habitus of this specimen (Figs 1, 2) are provided herein for facilitating the identification of this species.

Strongylophthalmyia corniculata sp. nov.

Figs 3, 4–8, 9–12

Type material

Holotype (♂): China, Yunnan, Honghe, Lvchun, Huanglianshan, 2018.v.19, by Malaise trap (CAU).

Diagnosis

Generally shiny black (Fig. 3); anterior half of frons yellowish brown (Fig. 4); antennal first flagellomere of male yellowish brown, ovate, with a short conical process (Figs 5, 6); wing hyaline (Fig. 8); mid and hind femora yellow, with narrow dark brown ring subapically (Fig. 3); hind femur of male lacking inner basal process; distiphallus less than half as long as phallapodeme, lacking apical “glans” (Fig. 12).

Figure 3. 

Strongylophthalmyia corniculata sp. nov., male, holotype, habitus, lateral view. Scale bar: 1 mm.

Figures 4–8. 

Strongylophthalmyia corniculata sp. nov., male, holotype: 4 head, dorsal view 5 same, lateral view 6 left antenna, lateral view 7 thorax, dorsal view 8 wing. Scale bars: 0.25 mm.

Figures 9–12. 

Strongylophthalmyia corniculata sp. nov., male genitalia: 9 external genitalia, posterior view 10 same, lateral view 11 internal genitalia, ventral view 12 same, lateral view. Abbreviations: cerc = cercus; dph = distiphallus; ea = ejaculatory apodeme; ep = epandrium; hyp = hypandrium; pgt = pregonite; phap = phallapodeme; pp = phallic plate; sur = surstylus. Scale bar: 0.1 mm.

Description

Body length 2.2 mm, wing length 2.2 mm.

Male. Generally shiny black (Fig. 3). Face, parafacial and anterior half of frons yellowish brown (Figs 4, 5). Antenna yellowish brown with arista dark brown (Fig. 5). Proboscis and palpus brown. Wing hyaline (Fig. 8); wing veins brown to dark brown. Halter white with base slightly darkened. Legs yellow with mid and hind femora narrowly dark brown subapically (Fig. 3); tarsomeres 4 and 5 dark brown.

Head (Figs 4, 5) with frons finely inflated; parafacial with dense silvery tomentose stripe; gena with silvery tomentose stripe along eye margin; postgena bulging. Head chaetotaxy: 1 inner vertical seta, 1 outer vertical seta, 3 fronto-orbital setae, 1 ocellar seta, 1 postocellar seta. Clypeus band-like; palpus elongate, with long sparse golden setulae. Antennal scape with scattered marginal setae and 1 dominant dorsal seta; pedicel with single strong seta dorsally; first flagellomere (Figs 5, 6) ovate, wider than long, densely covered with long white setulae, with a short conical process dorsally; antennal process (Fig. 6) with short dense white setulae, 0.7× as long as first flagellomere, sharp at apex; arista longer than antennal process.

Thorax with mesonotum (Fig. 7) densely covered with short scattered golden setulae, in dorsal view with distinct transverse suture. Anepisternum with short setulae along notopleural suture. Scutellum (Fig. 7) subtriangular, broad, slightly inflated. Thoracic chaetotaxy: 1 postpronotal seta, 1 anepisternal seta, 2 notopleural setae, 1 dorsocentral seta, 1 posterior supra-alar seta, 1 scutellar seta. Wing (Fig. 8) with R4+5 and M1+2 almost parallel apically; apical section of M1+2 finely arched; M4 and CuA+CuP not reaching but very closely approaching wing margin; r-m located near basal one-third (0.36) of cell dm; apical section of M4 nearly as long as dm-m; alula small; anal lobe slightly narrowed. Legs with dense whitish yellow setulae; hind femur lacking inner basal process.

Abdomen covered with long dense setae. Tergite 1 weakly sclerotized. Pregenital sclerites relatively weakly sclerotized.

Male genitalia: Epandrium (Figs 9, 10) short and broad, with long dense setae. Surstylus (Figs 9, 10) with short stout setae on inner distal surface. Cerci (Figs 9, 10) narrow, elongate, finger-like, with short dense setae. Hypandrium (Figs 11, 12) broadly rounded anteriorly, strongly arched medially, with one pair of bifid, long anterior lobes. Phallapodeme (Figs 11, 12) long, rod-like. Pregonite (Fig. 11) long, narrow, band-like, basally fused to inner surface of hypandrium. Phallic plate (Fig. 12) thickened, divided into two articulating sclerites. Distiphallus (Fig. 12) extremely short, less than half as long as phallapodeme, lacking apical “glans”, membrane microtrichose. Ejaculatory apodeme (Fig. 12) small, slightly curved.

Female unknown.

Etymology

The specific epithet is derived from Latin corniculata, referring to the short conical antennal process of this new species.

Distribution

China – Yunnan: Lvchun (Fig. 56).

Comparative notes

This new species is similar to S. gibbifera Shatalkin, 1993 from Vietnam in that both have an ovate first flagellomere with a small, short antennal process, and a fully hyaline wing. The new species differs from S. gibbifera in the following characters: anterior half of frons yellowish brown (frons entirely black in S. gibbifera); first flagellomere yellowish brown (yellowish brown with dorsal half dark brown in S. gibbifera); antennal process conical, with sharp apex (short and blunt in S. gibbifera); mid and hind femora narrowly dark brown subapically, hind tibia largely darkened (femora yellow with hind femur weakly darkened at apex, hind tibia darkened in middle in S. gibbifera).

Strongylophthalmyia flagellicornis sp. nov.

Figs 13–14, 15–19, 20–23

Type material

Holotype (♂): China, Yunnan, Xishuangbanna, Mengla, Yaoqu, 840 m, 2020.xi.22, leg. Liang Wang (CAU). Paratypes: Same collection data as for holotype (9♂, CAU); China, Yunnan, Honghe, Gejiu, Lvshuihe, 505 m, 2019.iii.30, leg. Xin Li & Liang Wang (5♂1♀, CAU); China, Yunnan, Honghe, Lvchun, Huanglianshan, 1300 m, 2018.vi.16, leg. Liang Wang (2♂, CAU).

Diagnosis

Generally shiny black (Figs 13, 14); frons yellow (Fig. 15); antennal first flagellomere of male yellow, subrhombic, with a long slender process covered with dense black setulae (Figs 16, 17); wing infumate, with large dark suffusion at apex (Fig. 19); mid and hind femora yellow with narrow dark brown ring subapically (indistinct on mid femur) (Figs 13, 14); hind femur of male with one thorn-like inner basal process; distiphallus nearly as long as phallapodeme, with small apical “glans” (Figs 22, 23).

Figures 13, 14. 

Strongylophthalmyia flagellicornis sp. nov., lateral habitus, lateral view: 13 male, holotype 14 female, paratype. Scale bars: 1 mm.

Figures 15–19. 

Strongylophthalmyia flagellicornis sp. nov., male, holotype: 15 head, dorsal view 16 same, lateral view 17 left antenna, lateral view 18 thorax, dorsal view 19 wing. Scale bars: 0.25 mm.

Figures 20–23. 

Strongylophthalmyia flagellicornis sp. nov., male genitalia: 20 external genitalia, posterior view 21 same, lateral view 22 internal genitalia, ventral view 23 same, lateral view. Abbreviations: cerc = cercus; dph = distiphallus; ea = ejaculatory apodeme; ep = epandrium; hyp = hypandrium; pgt = pregonite; phap = phallapodeme; pp = phallic plate; sur = surstylus. Scale bar: 0.1 mm.

Description

Body length 4.2–4.7 mm, wing length 2.4–3.0 mm.

Male. Generally shiny black (Fig. 13). Frons, face, parafacial, and gena yellow (Figs 15, 16). Antenna yellow; antennal process and arista blackish brown (Figs 16, 17). Clypeus yellow to yellowish brown; proboscis brown; palpus yellow. Wing infumate, with large dark suffusion at apex (Fig. 19); wing veins brown to dark brown. Halter white with base slightly darkened. Legs yellow; mid and hind femora with narrow dark brown ring subapically (indistinct on mid femur) (Fig. 13); mid and hind tarsomeres 4 and 5 dark brown. Abdominal syntergite 1+2 with large median yellow patch in posterior half.

Head (Figs 15, 16) with frons with minute shallow striations; parafacial with dense silvery tomentose stripe; gena with silvery tomentose stripe along eye margin; postgena bulging, with several white long setulae. Head chaetotaxy: 1 inner vertical seta, 1 outer vertical seta, 3 fronto-orbital setae, 1 ocellar seta, 1 postocellar seta. Clypeus band-like; palpus elongate, with short dense golden setulae. Antennal scape with scattered marginal setae and 1 dominant dorsal seta; pedicel with single strong seta dorsally; first flagellomere (Figs 16, 17) subrhombic, wider than long, densely covered with whitish yellow setulae, with a long slender process dorsally; antennal process (Fig. 17) with dense erect black setulae, 4.8× as long as first flagellomere; arista shorter than antennal process.

Thorax with mesonotum (Fig. 18) densely covered with short scattered golden setulae, in dorsal view with distinct transverse suture. Anepisternum with short dense golden setulae posteriorly. Scutellum (Fig. 18) subtriangular, broad, slightly inflated. Thoracic chaetotaxy: 1 anepisternal seta, 2 notopleural setae, 2 dorsocentral setae, 2 posterior supra-alar setae, 1 scutellar seta. Wing (Fig. 19) with R4+5 and M1+2 slightly convergent apically; apical section of M1+2 clearly arched; M4 and CuA+CuP not reaching but very closely approaching wing margin; r-m located at basal two-fifth (0.4) of cell dm; apical section of M4 shorter than dm-m; alula small; anal lobe well developed. Legs with dense dark setulae; hind femur with one thorn-like inner basal process.

Abdomen covered with short to long dense setae. Tergite 1 normally sclerotized. Pregenital sclerites weakly sclerotized.

Male genitalia: Epandrium (Figs 20, 21) long and narrow, subovate in lateral view, with long dense setae. Surstylus (Figs 20, 21) with short stout setae on inner distal surface. Cerci (Figs 20, 21) narrow, elongate, finger-like, fused along length, with 1 long subapical seta and several short setae. Hypandrium (Figs 22, 23) narrowly rounded anteriorly, arched medially, with one pair of bifid anterior lobes. Phallapodeme (Figs 22, 23) long, straight, rod-like, nearly as long as distiphallus. Pregonite (Fig. 22) very long, narrow, band-like, basally fused to inner surface of hypandrium. Phallic plate (Fig. 23) divided into two articulating sclerites. Distiphallus (Figs 22, 23) extremely long, nearly as long as phallapodeme, unsegmented, with sclerotized bands and apical “glans”, membrane microtrichose. Ejaculatory apodeme (Fig. 23) relatively long, straight, narrow.

Female. Antennal first flagellomere yellow with anterior margin darkened, ovate, lacking process (Fig. 14); clypeus thick, bulbous; hind femur lacking process; abdomen without yellow patch on syntergite 1+2. Other characters same as those of male.

Etymology

The specific epithet is derived from Latin flagell- and -cornis, referring to the long, whip-like antennal process of this new species.

Distribution

China – Yunnan: Gejiu, Lvchun, Mengla (Fig. 56).

Comparative notes

This new species resembles S. shatalkini Iwasa & Evenhuis, 2014 from Papua New Guinea and S. stylocera Shatalkin, 1996 from Philippines by having the long and slender antennal process. The new species differs from S. shatalkini in the following characters: frons yellow (entirely black in S. shatalkini); antennal first flagellomere of male subrhombic, with antennal process 4.8× as long as first flagellomere (ovate, with antennal process 4× as long as first flagellomere in S. shatalkini); femora yellow with mid and hind femora narrowly dark brown subapically (femora uniformly dark brown in S. shatalkini); wing with distinct large dark suffusion at apex, and R4+5 and M1+2 slightly convergent apically (wing with broad median transverse band and faint small apical suffusion, and R4+5 and M1+2 almost parallel in S. shatalkini). The new species can be separated from S. stylocera in the following characters: frons yellow (entirely black in S. stylocera); antennal process of male with dense dark setulae (with white setulae in S. stylocera); wing with distinct large dark suffusion at apex (wing hyaline in S. stylocera); abdomen with a large median yellow patch on syntergite 1+2 in male (uniformly black in S. stylocera).

Strongylophthalmyia narwhal Evenhuis, 2020

Figs 24, 25–29, 30–33

Strongylophthalmyia narwhal Evenhuis, 2020: 5 (protologue). Holotype (♂): Thailand, Chiang Mai, Chiang Mai, QSBG.

Type material examined

Holotype (♂): Thailand, Chiang Mai, Queen Sirikit Botanical Garden, 2013.vii.11–25, leg. M. Hauser, by Malaise trap (QSBG).

Other materials examined

China. Yunnan: Honghe, Lvchun, Huanglianshan, 380 m, 2018.iv.15, leg. Liang Wang (1♂, CAU); same locality and collector as preceding, 1300 m, 2018.iv.16 (1♂, CAU); Xishuangbanna, Jinghong, Naban River Watershed National Nature Reserve, Mandian Waterfall, 700 m, 2020.xi.25, leg. Liang Wang (1♂, CAU).

Diagnosis

Generally shiny blackish brown (Fig. 24); antennal first flagellomere of male yellowish brown with apical half darkened, round, with a long, thick, sword-like process (Figs 26, 27); wing infumate, with large dark suffusion at apex and narrow median transverse band at level of dm-m (Fig. 29); fore femur yellow with narrow brown ring subapically (Fig. 24); mid and hind femora yellowish brown, with apical half of mid femur and broad subbasal and subapical rings of hind femur dark brown (Fig. 24); hind femur of male with one wart-like inner basal process; distiphallus approx. 1.3× as long as phallapodeme, with small apical “glans” (Figs 32, 33).

Figure 24. 

Strongylophthalmyia narwhal Evenhuis, 2020, non-type male, habitus, lateral view. Scale bar: 1 mm.

Figures 25–29. 

Strongylophthalmyia narwhal Evenhuis, 2020, non-type male: 25 head, dorsal view 26 same, lateral view 27 left antenna, lateral view 28 thorax, dorsal view 29 wing. Scale bars: 0.25 mm.

Figures 30–33. 

Strongylophthalmyia narwhal Evenhuis, 2020, male genitalia: 30 external genitalia, posterior view 31 same, lateral view 32 internal genitalia, ventral view 33 same, lateral view. Abbreviations: cerc = cercus; dph = distiphallus; ea = ejaculatory apodeme; ep = epandrium; hyp = hypandrium; pgt = pregonite; phap = phallapodeme; pp = phallic plate; sur = surstylus. Scale bar: 0.1 mm.

Description of male genitalia

Epandrium (Figs 30, 31) short and narrow, with long, dense setae. Surstylus (Figs 30, 31) with short stout setae on inner distal surface. Cerci (Figs 30, 31) narrow, elongate, finger-like, fused along length, with 1 long subapical seta and several short setae. Hypandrium (Figs 32, 33) narrowly rounded anteriorly, strongly arched medially, with one pair of bifid anterior lobes. Phallapodeme (Figs 32, 33) short, slightly curved, distinctly shorter than distiphallus. Pregonite (Fig. 32) very long, narrow, band-like, basally fused to inner surface of hypandrium. Phallic plate (Fig. 33) divided into two articulating sclerites. Distiphallus (Figs 32, 33) extremely long, nearly 1.3× as long as phallapodeme, unsegmented, with sclerotized bands and apical “glans”, membrane microtrichose. Ejaculatory apodeme (Fig. 33) relatively large, slightly curved.

Distribution

China – Yunnan: Jinghong*, Lvchun* (Fig. 56). Thailand – Chiang Mai: Chiang Mai (Evenhuis 2020).

Remarks

This species was described based on a male holotype from Chiang Mai, Thailand (Evenhuis 2020). Newly available specimens from Yunnan, China are identical in most of the features described by Evenhuis (2020), but differs in the following aspects: (i) gena blackish brown as general body color; (ii) both wings with narrow median transverse band at level of dm-m; (iii) legs yellow to yellowish brown, with basal half of fore coxa, narrow subapical ring of fore femur, apical half of mid femur (except extreme apex), broad subbasal and subapical rings of hind femur, mid and hind tibiae (except bases and apices) and tarsomeres 4 and 5 brown to dark brown.

The chaetotaxy, which was not mentioned in the original description, can now be added as: head with 1 inner vertical seta, 1 outer vertical seta, 3 fronto-orbital setae, 1 ocellar seta and 1 postocellar seta; thorax with 1 anepisternal seta, 1 postpronotal seta, 2 notopleural setae, 2 dorsocentral setae, 2 posterior supra-alar setae and 1 scutellar seta; mid femur with row of long erect black setae. The male genitalia are described as above.

Strongylophthalmyia raricornis Shatalkin, 1981

Figs 34–35, 36–40, 41–44

Strongylophthalmyia raricornis Shatalkin, 1981: 792 (protologue); Krivosheina (1984: 27) (catalogue, distribution); Shatalkin (1993: 124, 126) (in key, bionomics, figure); Shatalkin (1996: 156) (diagnosis, redescription, record, figure); Krivosheina (1999: 508) (in key, distribution, figure); Ozerov (2010: 156) (type material); Iwasa and Evenhuis (2014: 103) (listed, distribution); Evenhuis (2016: 207) (listed, distribution, figure). Holotype (♂): Russia, Khabarovsk Krai, Maly Khingan, ZMUM.

Material examined

China. Beijing: Haidian, Jiufeng, 101 m, 2018.vi.20, leg. Jiale Zhou & Yike Cao (2♂1♀, CAU). Shaanxi: Mei County, Haopingsi, 1120 m, 2020.vi.29, leg. Bing Zhang (4♂, CAU); Tongchuan, Miaowan, 1233 m, 2019.vii.28, leg. Qicheng Yang & Jiaojie Wang (1♂10♀, CAU); Tongchuan, Yuhuagong, 1385 m, 2019.vii.30, leg. Jiaojie Wang (3♂, CAU); Tongchuan, Liulin, 1020 m, 2019.vii.27, leg. Qicheng Yang & Weijian Huang (4♂2♀, CAU); Xunyi, Shimenshan, 1577 m, 2019.vii.26, leg. Qicheng Yang (4♂, CAU); same locality, 1605m, 2019.vii.25, leg. Jiaojie Wang (1♂1♀, CAU). Shanghai: Baoshan, Gucun Park, 2021.v.9, photographed by Deyao Zhou (1♂, photo voucher only). Tianjin: Jizhou, Baxianshan, 221–706 m, 2019.vi.28, leg. Ding Yang (1♂4♀, CAU). SOUTH KOREA. Chungbuk: Okcheon, Dongi, Soesan, 150 m, leg. P. Tripton, by Malaise trap (1♂1♀, CSCA).

Diagnosis

Generally shiny black (Figs 34, 35); antennal first flagellomere of male yellowish brown, bifid, C-shaped (Figs 37, 38); wing hyaline (Fig. 40); legs blackish brown with fore and mid coxae (except base), trochanters, extreme apex of fore and mid femora, fore tibia, apex of mid and hind tibiae, and tarsomeres 1–3 yellow (Figs 34, 35); hind trochanter of male with a tiny thorn-like process on internal surface; hind femur of male with three small warty inner basal processes, middle one bearing setae, other two bare; distiphallus slightly shorter than phallapodeme, with large apical “glans” (Figs 43, 44).

Figures 34, 35. 

Strongylophthalmyia raricornis Shatalkin, 1981, habitus, lateral view: 34 non-type male 35 non-type female. Scale bars: 1 mm.

Figures 36–40. 

Strongylophthalmyia raricornis Shatalkin, 1981, non-type male: 36 head, dorsal view 37 same, lateral view 38 left antenna, lateral view; 39 thorax, dorsal view 40 wing. Scale bars: 0.25 mm.

Figure 41–44. 

Strongylophthalmyia raricornis Shatalkin, 1981, male genitalia: 41 external genitalia, posterior view 42 same, lateral view 43 internal genitalia, ventral view 44 same, lateral view. Abbreviations: cerc = cercus; dph = distiphallus; ea = ejaculatory apodeme; ep = epandrium; hyp = hypandrium; pgt = pregonite; phap = phallapodeme; pp = phallic plate; sur = surstylus. Scale bar: 0.1 mm.

Redescription

Body length 3.5–4.6 mm, wing length 2.5–2.9 mm.

Male. Generally shiny black (Fig. 34). Antenna yellowish brown with arista dark brown (Figs 37, 38). Proboscis and palpus brown. Wing hyaline (Fig. 40); wing veins brown to dark brown. Halter white with base slightly darkened. Legs blackish brown; fore and mid coxae (except base), trochanters, extreme apex of fore and mid femora, fore tibia, apex of mid and hind tibiae, and tarsomeres 1–3 yellow (Fig. 34).

Head (Figs 36, 37) with frons flattened; parafacial with dense silvery tomentose stripes; gena with silvery tomentose stripe along eye margin; postgena bulging. Head chaetotaxy: 1 inner vertical seta, 1 outer vertical seta, 3 fronto-orbital setae, 1 ocellar seta, 1 postocellar seta. Clypeus subquadrate; palpus elongate, with short dense black setulae. Antennal scape with scattered marginal setae and 1 dominant dorsal seta; pedicel with elongate dorsal seta; first flagellomere (Figs 37, 38) bifid, C-shaped, with short dense white setulae; arista longer than first flagellomere.

Thorax with mesonotum (Fig. 39) densely covered with long setulae located in rows, in dorsal view with distinct transverse suture. Anepisternum with short dense setulae. Katepisternum with short setulae ventrolaterally near mid coxa. Scutellum (Fig. 39) subtriangular, slightly inflated. Thoracic chaetotaxy: 1 anepisternal seta, 1 presutural intra-alar seta, 2 notopleural setae, 7 dorsocentral setae, 2 posterior supra-alar setae, 1 scutellar seta. Wing (Fig. 40) with R4+5 and M1+2 almost parallel apically; apical section of M1+2 clearly arched; M4 and CuA+CuP not reaching but very closely approaching wing margin; r-m located near basal one-third (0.34) of cell dm; apical section of M4 shorter than dm-m; alula relatively large; anal lobe well developed. Legs with dense dark setulae; fore coxa with several long white setulae antero-apically; hind trochanter with a tiny thorn-like process on internal surface; hind femur with three small warty inner basal processes, middle one bearing setae, other two bare.

Abdomen covered with short to long dense setae. Tergite 1 weakly sclerotized. Pregenital sclerites relatively weakly sclerotized.

Male genitalia: Epandrium (Figs 41, 42) long and narrow, subovate in lateral view, with long dense setae. Surstylus (Figs 41, 42) with several long dense setae at apex and short stout setae on inner distal surface. Cerci (Figs 41, 42) relatively short and broad, fused along length, with one rather long subapical seta and several short setae. Hypandrium (Figs 43, 44) narrowly rounded anteriorly, with one pair of bifid, long anterior lobes. Phallapodeme (Figs 43, 44) long, curved, slightly longer than distiphallus. Pregonite (Fig. 43) very long, narrow, band-like, basally fused to inner surface of hypandrium. Phallic plate (Fig. 44) divided into two articulating sclerites. Distiphallus (Figs 43, 44) extremely long, slightly shorter than phallapodeme, unsegmented, with sclerotized bands and large apical “glans”, membrane microtrichose. Ejaculatory apodeme (Figs 43, 44) large, slightly swollen at apex.

Female. Antennal first flagellomere ovate (Fig. 35); clypeus thick, bulbous; hind trochanter and femur lacking process. Other characters same as those of male.

Distribution

China – Beijing: Haidian*, Shaanxi: Mei County*, Tongchuan*, Xunyi*, Shanghai: Baoshan*, Tianjin: Jizhou* (Fig. 56). Russia – Khabarovsk Krai: Maly Khingan (Shatalkin 1981), Primorsky Krai: Kamenushka, Lazovskiy Nature Reserve (Shatalkin 1996). South Korea – Chungbuk: Okcheon* (Fig. 56).

Remarks

The male genitalia of this species were illustrated by Shatalkin (1996) and Krivosheina (1999) and are described here for the first time. This species was previously reported from the Russian Far East (Shatalkin 1996; Krivosheina 1999). Considering the new distributional records from China and South Korea, this species seems to be widely distributed in the eastern Palaearctic Realm.

Strongylophthalmyia tangwangana sp. nov.

Figs 45–46, 47–51, 52–55

Type material

Holotype (♂): China, Shaanxi, Tongchuan, Tang Wang Hunting Ground, 1429 m, 2019.vii.30, leg. Qicheng Yang (CAU). Paratypes. Same collection data as for holotype (1♂2♀, CAU).

Diagnosis

Generally shiny black (Figs 45, 46); antennal first flagellomere of male yellowish brown, ovate, with a small, bump-like process (Figs 48, 49); wing hyaline (Fig. 51); mid and hind femora yellow, with narrow dark brown ring subapically (indistinct on mid femur) (Figs 45, 46); hind femur of male lacking inner basal process; distiphallus nearly half as long as phallapodeme, lacking apical “glans” (Figs 54, 55).

Figures 45, 46. 

Strongylophthalmyia tangwangana sp. nov., habitus, lateral view: 45 male, holotype 46 female, paratype. Scale bars: 1 mm.

Figures 47–51. 

Strongylophthalmyia tangwangana sp. nov., male, holotype: 47 head, dorsal view 48 same, lateral view 49 left antenna, lateral view 50 thorax, dorsal view 51 wing. Scale bars: 0.25 mm.

Figures 52–55. 

Strongylophthalmyia tangwangana sp. nov., male genitalia: 52 external genitalia, posterior view 53 same, lateral view 54 internal genitalia, ventral view 55 same, lateral view. Abbreviations: cerc = cercus; dph = distiphallus; ea = ejaculatory apodeme; ep = epandrium; hyp = hypandrium; pgt = pregonite; phap = phallapodeme; pp = phallic plate; sur = surstylus. Scale bar: 0.1 mm.

Description

Body length 2.5–3.2 mm, wing length 2.1–2.4 mm.

Male. Generally shiny black (Fig. 45). Anterior margin of frons slightly paler; face and parafacial yellowish brown. Antenna yellowish brown with arista dark brown (Figs 48, 49). Proboscis and palpus yellowish brown. Wing hyaline (Fig. 51); wing veins brown to dark brown. Halter white with base darkened. Legs yellow; mid and hind femora with narrow dark brown ring subapically (indistinct on mid femur) (Figs 45, 46); hind tibia largely dark brown (Fig. 45); tarsomeres 4 and 5 dark brown.

Head (Figs 47, 48) with frons finely inflated; parafacial with dense silvery tomentose stripe; gena with silvery tomentose stripe along eye margin; postgena bulging. Head chaetotaxy: 1 inner vertical seta, 1 outer vertical seta, 3 fronto-orbital setae, 1 ocellar seta, 1 postocellar seta. Clypeus subquadrate; palpus elongate, with short dense golden setulae. Antennal scape with scattered marginal setae and 1 dominant dorsal seta; pedicel with single strong seta dorsally; first flagellomere ovate, wider than long, densely covered with short white setulae, with a small, bump-like process dorsally (Figs 48, 49); arista longer than first flagellomere.

Thorax with mesonotum (Fig. 50) densely covered with short scattered golden setulae, in dorsal view with distinct transverse suture. Anepisternum with short setulae posteriorly. Scutellum (Fig. 50) subtriangular, broad, slightly inflated. Thoracic chaetotaxy: 1 anepisternal seta, 2 notopleural setae, 1 dorsocentral seta, 2 posterior supra-alar setae, 1 scutellar seta. Wing (Fig. 51) with R4+5 and M1+2 slightly convergent apically; apical section of M1+2 straight; M4 and CuA+CuP not reaching but very closely approaching wing margin; r-m located near basal one-third (0.34) of cell dm; apical section of M4 shorter than dm-m; alula small; anal lobe slightly narrowed. Legs with dense whitish yellow setulae; fore coxa with several long white setulae antero-apically; hind femur lacking inner basal process.

Abdomen covered with long dense setae. Tergite 1 weakly sclerotized. Pregenital sclerites normally sclerotized.

Male genitalia: Epandrium (Figs 52, 53) short and broad, with long dense setae. Surstylus (Figs 52, 53) with short stout setae on inner distal surface. Cerci (Figs 52, 53) relatively broad, shorter than surstylus, with short dense setae. Hypandrium (Figs 54, 55) broadly rounded anteriorly, strongly arched medially, with one pair of bifid, long anterior lobes. Phallapodeme (Figs 54, 55) extremely long, slightly curved. Pregonite (Fig. 54) long, narrow, band-like. Phallic plate (Fig. 55) strongly thickened, divided into two articulating sclerites. Distiphallus (Figs 54, 55) extremely short, nearly half as long as phallapodeme, lacking apical “glans”, membrane microtrichose. Ejaculatory apodeme (Fig. 55) small.

Female. Antennal first flagellomere lacking process (Fig. 46); clypeus thick, bulbous. Other characters same as those of male.

Etymology

This species is named after its type locality.

Distribution

China – Shaanxi: Tongchuan (Fig. 56).

Figure 56. 

Known distribution of species of the Strongylophthalmyia coarctata subgroup in China and South Korea.

Comparative notes

This new species is similar to S. corniculata sp. nov., but can be distinguished from it in the following characters: frons black, at most anterior margin slightly paler (anterior half of frons yellowish brown in S. corniculata sp. nov.); first flagellomere of male with a small, bump-like process (with a small, conical, apically acute process in S. corniculata sp. nov.); thorax with two posterior supra-alar setae (one in S. corniculata sp. nov.); apical section of M1+2 straight (slightly arched in S. corniculata sp. nov.); distiphallus nearly half as long as phallapodeme (distinctly shorter than half of length of phallapodeme in S. corniculata sp. nov.).

This new species also resembles S. gibbifera Shatalkin, 1993, but differs in having a different color pattern on the frons, first flagellomere, and the mid and hind femora and hind tibia, and in the different shape of the antennal process.

Discussion

The present study is part of our ongoing taxonomic study of the Chinese Strongylophthalmyiidae, documenting the Strongylophthalmyia coarctata subgroup from China, including three known species (two newly recorded in China) and three new species. All species are keyed.

The main characters currently used to distinguish species in the S. coarctata subgroup include body color (including color patterns on antenna and wing), male antennal morphology, and the inner basal process on male hind femur. In this study, we newly described the male genitalia of most Chinese species, and found that the morphology of the external genitalia, phallapodeme, and distiphallus differed among species in this subgroup: the external genitalia are short and broad in some species, while in others they are long and narrow; the phallapodeme is straight or clearly curved, and the relative length of the distiphallus varies among species; the distiphallus of different species has different forms of tricha on the membrane and apical “glans”, and the apical “glans” is absent in S. corniculata sp. nov. and S. tangwangana sp. nov. These male genital characters are useful for species-level identification. In addition, the thoracic chaetotaxy varies between species, which may also be helpful in identifying female specimens that lack male-specific features.

The S. coarctata subgroup has high diversity in the Oriental Realm. However, most of the specimens currently available to us were collected from several scattered localities in China. There is no doubt that a considerable number of other species could be discovered in China (especially in southern China) following more thorough field investigations.

Acknowledgements

We sincerely thank Owen Lonsdale (Agriculture and Agri-Food Canada, Ottawa, Canada), Stephen D. Gaimari (CSCA) and Stephen A. Marshall (University of Guelph, Guelph, Canada) for their critical reading and helpful comments on the manuscript, and Vera C. Silva (São Paulo State University, Jaboticabal, Brazil) for her editorial work. We are very grateful to Duncan Sivell (NHMUK) for his kind help during our examination of dipteran specimens under his curation, to Deyao Zhou (Jiading Agriculture Technology Extension Service Center, Shanghai, China) for providing the photo voucher of S. raricornis from Shanghai, China, and to Jenny Pohl and Sven Marotzk (ZMHB) for checking and sharing information of specimens in ZMHB.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by grants from the National Natural Science Foundation of China (No. 31970444) and the China Scholarship Council (No. 202106350076).

Author contributions

Conceptualization, Jiale Zhou and Ding Yang; writing – original draft preperation, Jiale Zhou, Neal L. Evenhuis and Ding Yang; writing – review and editing, Jiale Zhou, Neal L. Evenhuis and Ding Yang; material, Jiale Zhou, Neal L. Evenhuis and Ding Yang; visualization, Jiale Zhou; funding, Jiale Zhou and Ding Yang.

Author ORCIDs

Jiale Zhou https://orcid.org/0000-0002-8901-3187

Neal L. Evenhuis https://orcid.org/0000-0002-1314-755X

Ding Yang https://orcid.org/0000-0002-7685-3478

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

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