Research Article |
Corresponding author: Ding Yang ( dyangcau@126.com ) Academic editor: Torsten Dikow
© 2023 Jiale Zhou, Ding Yang.
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Citation:
Zhou J, Yang D (2023) A review of the subgenus Loxocera Meigen, 1803 (Diptera, Brachycera, Psilidae) in China. ZooKeys 1186: 71-96. https://doi.org/10.3897/zookeys.1186.108876
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The subgenus Loxocera (Loxocera Meigen, 1803) (Diptera, Brachycera, Psilidae) in China is reviewed. Six species are recognized, including four new species: L. (L.) chikuni sp. nov., L. (L.) lonsdalei sp. nov., L. (L.) maculithorax sp. nov., and L. (L.) obscura sp. nov. Two species originally placed in the subgenus Loxocera, L. anulata Wang & Yang, 1998 and L. tianmuensis Wang & Yang, 1998, are transferred to the subgenus Loxocera (Imantimyia Frey, 1925). A key to the species of the subgenus Loxocera occurring in China is provided.
Acalyptratae, Diopsoidea, Imantimyia, new species, Palaearctic Realm, Psilinae
Psilidae is a group of small to medium-sized, yellow to black acalyptrate flies which can be diagnosed externally by their peculiar wing venation and reduced setation (
The psiline genus Loxocera Meigen, 1803 is cosmopolitan and currently comprises about 50 species. This genus was traditionally divided into four subgenera: Loxocera s. str., Asiopsila Shatalkin, 1998, Platystyla Macquart, 1835, and Tropeopsila Shatalkin, 1983 (
In the present study, we review and key the Chinese fauna of the subgenus Loxocera and document six species, four of which are described here as new. The other two Chinese species previously placed in the subgenus Loxocera are here transferred to Imantimyia.
Outside of China, an additional eight species of the subgenus Loxocera are known, with these occurring exclusively within the Palaearctic and northern Oriental realms (
Specimens examined in this study are deposited in the
Entomological Museum of China Agricultural University, Beijing, China (
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 glycerol and stored in microvials pinned below the corresponding specimens.
Photographs were taken using a Canon 7D Mark II digital camera with a Canon macro lens EF 100 mm and MP-E 65 mm for habitus, and an Olympus BX51 microscope for terminalia. Figures were stacked using Helicon Focus v. 5.3 and assembled by Adobe Photoshop 2020. The distribution map was prepared using the online version of SimpleMappr (
Loxocera Meigen, 1803: 275. Type species: Musca aristata Panzer, 1801, by monotypy.
Platystyla
Macquart, 1835: 374. Type species: Loxocera hoffmannseggii Meigen, 1826, by monotypy. Synonymized by
Terarista Wang, 1999: 268. Nomen nudum.
Terarista
Yang & Wang in Wang & Yang, 2003: 563. Type species: Terarista fujiana Wang, 1999, by original designation. Synonymized by
The subgenus Loxocera can be recognized by the following combination of character states: frontal vitta desclerotized, dull, velvety; lunule sclerotized, broadly exposed between antennal base and anterior margin of frons; fore wing with alula glabrous except margin; hind femur with a subapical patch of microtomentum on ventral surface; male sternite 8 broadly exposed and setulose, fused to tergite 6 and epandrium; female tergite 10 and cerci separated. For details on the character states used to define the subgenus, see
1 | Arista blackish brown, laterally compressed and very high, arising at apex of antennal first flagellomere (Figs |
2 |
– | Arista whitish yellow, thin, arising near midpoint of antennal first flagellomere (Figs |
4 |
2 | Frontal vitta uniformly black | L. (L.) fujiana (Wang) |
– | Frontal vitta black with anterior part dark yellow or yellowish brown (Figs |
3 |
3 | Apex of antennal first flagellomere not produced beyond base of arista (Figs |
L. (L.) chikuni sp. nov. |
– | Apex of antennal first flagellomere clearly produced beyond base of arista (Fig. |
L. (L.) maculithorax sp. nov. |
4 | Antennal scape and pedicel subequal in length (Fig. |
L. (L.) omei Shatalkin |
– | Antennal scape and pedicel unequal in length (Figs |
5 |
5 | Antennal scape distinctly shorter than pedicel (Fig. |
L. (L.) lonsdalei sp. nov. |
– | Antennal scape distinctly longer than pedicel (Fig. |
L. (L.) obscura sp. nov. |
Holotype
(♂): China, Hubei, Shennongjia, Guanmenshan, 1560 m, 2019.viii.12, leg. Ding Yang (
Generally blackish brown; face blackish; antennal scape and pedicel subequal in length; apex of antennal first flagellomere not produced beyond base of arista; arista laterally compressed and very high, arising at apex of first flagellomere and 1.8 times as long as the latter; wing with broad, transverse, dark band at level of posterior crossvein; hypandrial lobe small, short, covered with long setae; pregonite stout, lobate, apically blunt with rounded processes; phallus tongue-like, relatively short, with rounded posterior margin.
Male. Body length 9.0 mm, wing length 7.2 mm, length of antenna 2.5 mm. Generally blackish brown, moderately shining (Fig.
Head (Figs
Thorax (Fig.
Abdomen elongate, with short, dense, whitish-yellow and black setae; syntergite 1+2 with several long, hair-like setae laterally.
Male genitalia
: Sternite 8 (Figs
Loxocera (Loxocera) chikuni sp. nov., male genitalia 5 sternite 8 to genitalia, caudal 6 same, lateral 7 pregonite and phallus, ventral 8 hypandrium and associated structures, ventral. Abbreviations: ce = cercus, ea = ejaculatory apodeme, ep = epandrium, ha = hypandrial arm, hb = hypandrial bridge, hl = hypandrial lobe, pg = pregonite, s8 = sternite 8, ph = phallus, pt = phallotrema.Scale bar: 0.25 mm.
Female. Unknown.
The specific epithet is dedicated to the Chinese entomologist Chikun Yang (1925–2006), in honor of his excellent contribution to Chinese insect taxonomy.
China – Hubei: Shennongjia (Fig.
This new species is similar to L. (L.) fujiana (Wang, 1999) and L. (L.) maculithorax sp. nov. by sharing the laterally compressed and high arista which arises at the apex of the antennal first flagellomere, and the broad, transverse, dark band on the wing. The new species differs from L. (L.) fujiana by the following character states: head blackish brown with anterior part of frontal vitta, parafacial, gena, and posterior eye margin yellowish brown [vs head blackish brown with gena slightly paler in L. (L.) fujiana]; arista 1.8 times as long as antennal first flagellomere [vs 2 times in L. (L.) fujiana]; hind tibia pale brown in basal half and blackish brown in apical half [vs uniformly dark brown in L. (L.) fujiana]; transverse dark band on wing interrupted along cell r2+3 [vs not interrupted in cell r2+3 in L. (L.) fujiana]. The new species can be separated from L. (L.) maculithorax sp. nov. by the following character states: apex of antennal first flagellomere not produced beyond base of arista [vs produced beyond base of arista in L. (L.) maculithorax sp. nov.]; arista 1.8 times as long as antennal first flagellomere [vs 2.1 times in L. (L.) maculithorax sp. nov.]; mesonotum uniformly blackish brown [vs largely blackish with irregular brownish margin in L. (L.) maculithorax sp. nov.]; abdomen uniformly blackish brown [vs reddish brown with both ends darkened in L. (L.) maculithorax sp. nov.]; male sternite 8 flattened [vs with a small blunt protrusion in L. (L.) maculithorax sp. nov.]; hypandrial lobe of male small and short [vs large and broad in L. (L.) maculithorax sp. nov.].
Terarista fujiana
Wang, 1999: 268 (protologue);
Loxocera (Loxocera) fujiana:
Lectotype
(♀): China, Fujian, Wuyishan, Guadun, 1991.x.7, leg. Hong Wu (
Generally blackish brown; face black; antennal scape and pedicel subequal in length; apex of antennal first flagellomere not produced beyond base of arista; arista laterally compressed and very high, arising at apex of antennal first flagellomere and 2 times as long as the latter; wing with broad, transverse, dark band at level of posterior crossvein; segment 8 of female with shallowly emarginated posterodorsal margin and deeply incised posteroventral margin.
China – Fujian: Wuyishan (Fig.
For redescription and photographs of this species, see
Holotype
(♂): China, Shaanxi, Xi’an, Huyi, Zhuque Forest Park, 2606 m, 2020.vii.10, leg. Bing Zhang (
Generally brown to blackish brown; face yellowish brown; antennal pedicel distinctly longer than scape; antennal first flagellomere about 3 times as long as pedicel, gradually narrowed towards apex; arista whitish yellow, thin, arising near midpoint of antennal first flagellomere and 1.3 times as long as the latter; mesonotum blackish, with irregular brownish margin; wing without transverse dark band; hypandrial lobe large, broad, covered with short setae; pregonite short, simple in shape, apically blunt; phallus shovel-like, relatively long, abruptly widened apically with rounded posterior margin.
Male and female. Body length 7.7–9.5 mm, wing length 6.0–6.5 mm, length of antenna 1.6–1.8 mm. Generally brown to blackish brown, moderately shining (Figs
Head (Figs
Thorax (Figs
Abdomen elongate, with short, dense, whitish-yellow setae; syntergite 1+2 with several long hair-like setae laterally.
Male genitalia
: Sternite 8 (Figs
Loxocera (Loxocera) lonsdalei sp. nov., male genitalia 14 sternite 8 to genitalia, caudal 15 same, lateral 16 pregonite and phallus, ventral 17 hypandrium and associated structures, ventral. Abbreviations: ce = cercus, ea = ejaculatory apodeme, ep = epandrium, ha = hypandrial arm, hb = hypandrial bridge, hl = hypandrial lobe, pg = pregonite, s8 = sternite 8, ph = phallus, pt = phallotrema. Scale bars: 0.25 mm (14, 15, 17); 0.1 mm (16).
Female terminalia : Segment 7 laterally compressed; sternite 7 keeled along midline. Segment 8 coriaceous and longitudinally striate, posterodorsal margin shallowly emarginate, posterolateral margin forming blunt angular protrusion, posteroventral margin with deep linear incision. Tergite 10 relatively long and broad, with tiny, scattered setulae. Cerci relatively broad, separate from tergite 10, with rows of long setulae on posterior margin.
The specific epithet is dedicated to the Canadian entomologist Owen Lonsdale (Agriculture and Agri-Food Canada, Ottawa, Canada), for his outstanding contribution to the systematics of Acalyptratae, and his encouragement to the first author.
China – Qinghai: Haibei, Haidong; Shaanxi: Xi’an (Fig.
This new species is most similar to L. (L.) omei Shatalkin, 1998, but it can be easily distinguished from the latter by the following character states: antennal pedicel distinctly longer than scape [vs subequal in length in L. (L.) omei]; antennal first flagellomere distinctly narrowed towards apex [vs weakly narrowed in L. (L.) omei]; mesonotum largely blackish with irregular brownish margin [vs uniformly blackish in L. (L.) omei]; abdomen yellowish brown with distinct ring-like markings [vs blackish brown with apical segments reddish in L. (L.) omei].
Antennal morphology similar to that of this new species also presents in L. (L.) hoffmannseggi Meigen, 1826 from Central and West Europe and L. (L.) matsumurai Iwasa, 1992 from Japan. However, the coloration of head, thorax, and abdomen of the new species is very different from that of the latter two species. Additionally, the head of the new species is nearly rounded in dorsal view with the frons strongly protruding anteriorly, while in the latter two species, the head is transverse oblong in dorsal view and the frons weakly protrudes anteriorly.
Holotype
(♂): China, Shaanxi, Baoji, Longxian, Guanshan Grassland, 2034 m, 2020.viii.13, leg. Bing Zhang (
Generally dark brown; face brown; antennal scape and pedicel subequal in length; apex of antennal first flagellomere produced beyond base of arista; arista laterally compressed and very high, arising at apex of antennal first flagellomere and 2.1 times as long as the latter; mesonotum blackish, with irregular, brownish margin; wing with broad, transverse, dark band at level of posterior crossvein; hypandrial lobe large, short, covered with long setae; pregonite stout, lobate, apically blunt, with some rounded processes; phallus tongue-like, relatively short, with rounded posterior margin.
Male and female. Body length 8.8–10.5 mm, wing length 6.2–7.0 mm, length of antenna 1.9–2.3 mm. Generally dark brown, moderately shining (Figs
Head (Figs
Thorax (Figs
Abdomen elongate, with short, dense, whitish-yellow setae; syntergite 1+2 with several long hair-like setae laterally.
Male genitalia
: Sternite 8 (Figs
Loxocera (Loxocera) maculithorax sp. nov., male genitalia 23 sternite 8 to genitalia, caudal 24 same, lateral 25 pregonite and phallus, ventral 26 hypandrium and associated structures, ventral. Abbreviations: ce = cercus, ea = ejaculatory apodeme, ep = epandrium, ha = hypandrial arm, hb = hypandrial bridge, hl = hypandrial lobe, pg = pregonite, s8 = sternite 8, ph = phallus, pt = phallotrema. Scale bars: 0.25 mm (23, 24, 26); 0.1 mm (25).
Female terminalia : Segment 7 laterally compressed; sternite 7 keeled along midline. Segment 8 coriaceous and longitudinally striate, posterodorsal margin shallowly emarginate, posterolateral margin forming acute angular protrusion, posteroventral margin with deep linear incision. Tergite 10 relatively short and narrow, with scattered tiny setulae. Cerci relatively broad, separate from tergite 10, with rows of long setulae on posterior margin.
The specific epithet is derived from Latin macula (meaning spotted) and Greek thorax (meaning thorax), referring to the mesonotum of the new species, which is blackish with an irregular brownish margin.
China – Shaanxi: Baoji (Fig.
This new species is similar to L. (L.) fujiana, but it can be readily separated from the latter by the different coloration of head, mesonotum, hind tibia, and abdomen, the apically curved and produced first flagellomere, and the shortened scutum. It also resembles L. (L.) chikuni sp. nov., and their differences are discussed above [see under L. (L.) chikuni sp. nov.]. The Japanese species L. (L.) monstrata Iwasa, 1992 shares a similar antennal morphology with the new species, but its arista is less than 2 times as long as the first flagellomere, and it has different coloration on thorax, legs and abdomen.
Holotype
(♂): China, Shaanxi, Xi’an, Zhouzhi, Houzhenzi, 2009.ix.29, leg. Maoling Sheng (
Generally dark brown; face blackish; antennal scape distinctly longer than pedicel; antennal first flagellomere stick-like, weakly narrowed towards apex of segment; arista whitish yellow, thin, arising before midpoint of antennal first flagellomere and 1.4 times as long as the latter; wing without transverse dark band; hypandrial lobe very large, broad, covered with short, sparse setae on inner surface; pregonite rather long, slender, curved apically with sharp apex; phallus droplet-like, elongate, abruptly narrowed apically.
Male. Body length 8.9–10.2 mm, wing length 6.6–7.2 mm, length of antenna 2.2–2.5 mm. Generally dark brown, moderately shining (Fig.
Head (Figs
Thorax (Figs
Abdomen elongate, with short, dense, white setae; syntergite 1+2 with several long, hair-like setae laterally.
Male genitalia
: Sternite 8 (Figs
Loxocera (Loxocera) obscura sp. nov., male genitalia 31 sternite 8 to genitalia, caudal 32 same, lateral 33 pregonite and phallus, ventral 34 hypandrium and associated structures, ventral. Abbreviations: ce = cercus, ea = ejaculatory apodeme, ep = epandrium, ha = hypandrial arm, hb = hypandrial bridge, hl = hypandrial lobe, pg = pregonite, s8 = sternite 8, ph = phallus, pt = phallotrema.Scale bars: 0.25 mm (31, 32, 34); 0.1 mm (33).
Female. Unknown.
The specific epithet is derived from Latin obscura (meaning dark, indistinct), referring to the dark-brown body color of the new species.
China – Shaanxi: Xi’an (Fig.
This new species is most similar to L. (L.) malaisei (Frey, 1955) (from Myanmar and Nepal) in having a relatively long antennal scape (longer than pedicel), a nearly parallel-sided and laterally compressed antennal first flagellomere, similar coloration of legs, and an enlarged and posteriorly inflated male sternite 8. It can be distinguished from the latter by the following character states: antennal scape about 1.5 times as long as pedicel [vs 2 times in L. (L.) malaisei]; antennal first flagellomere elongate, 4.6 times as long as pedicel [vs shorter, 4 times in L. (L.) malaisei]; arista slender [vs widened towards apex in L. (L.) malaisei]; phallus droplet-like with apex abruptly narrowed [vs elongate oval in L. (L.) malaisei].
Loxocera (Platystyla) omei
Shatalkin, 1998: 90, 97 (protologue). Holotype (♂): China, Sichuan, Emeishan,
Loxocera (Loxocera) omei:
Loxocera omei:
Holotype
(♂): China, Sichuan, Leshan, Emeishan, 1935.vii.21, leg. D.C. Graham (
Generally blackish brown; face yellowish brown; antennal scape and pedicel subequal in length; antennal first flagellomere laterally compressed, about 3.6 times as long as pedicel, weakly narrowed towards apex; arista whitish yellow, thin, arising near midpoint of antennal first flagellomere and 1.3 times as long as latter; wing without transverse dark band.
China – Sichuan: Leshan (Fig.
This species was described based on one male (the holotype) from Sichuan, China (
Loxocera (Loxocera) anulata
Loxocera annulata:
Loxocera anulata:
Holotype
(♂): China, Hubei, Shennongjia, Dajiuhu, 1977.vii.9, leg. Huanguang Zou (
China – Hubei: Shennongjia.
Loxocera anulata was originally described as a member of the subgenus Loxocera based on one male (the holotype) from Hubei, China (
The holotype of L. anulata was examined during the present study. It satisfied the diagnosis of Loxocera s. lat. provided by
Loxocera tianmuensis
Holotype
(♂): China, Zhejiang, Lin’an, Xitianmushan, 350 m, 1987.ix.3, leg. Qun Ma (
China – Zhejiang: Lin’an.
Loxocera tianmuensis was originally described based on one male (the holotype) from Zhejiang, China (
The holotype of L. tianmuensis was examined in the course of the present study. The date on the collection data label of the holotype does not match that in the original description. The authors (
The present study documents the Chinese fauna of the subgenus Loxocera, including two described species and four new species. The other two species originally assigned to the subgenus Loxocera are herein transferred to the subgenus Imantimyia based on examination of their holotypes. An identification key to the species of the subgenus Loxocera from China is also presented.
Antennal morphology of members of the subgenus Loxocera is diverse and useful in species identification (
Including the four species newly described in this study, the subgenus Loxocera currently comprises 14 species (Table
Described species of the subgenus Loxocera and their known distribution.
Species | Distribution | References |
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L. (L.) aristata (Panzer, 1801) | Europe, Iran, Israel |
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L. (L.) atriceps Bigot, 1886 | Europe |
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L. (L.) chikuni Zhou & Yang, sp. nov. | China | present study |
L. (L.) fujiana (Wang, 1999) | China |
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L. (L.) glandicula Iwasa, 1993 | Nepal |
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L. (L.) hoffmannseggi Meigen, 1826 | Europe |
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L. (L.) lonsdalei Zhou & Yang, sp. nov. | China | present study |
L. (L.) maculata Rondani, 1876 | Europe |
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L. (L.) maculithorax Zhou & Yang, sp. nov. | China | present study |
L. (L.) malaisei Frey, 1955 | Myanmar, Nepal |
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L. (L.) matsumurai Iwasa, 1992 | Japan, Russia |
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L. (L.) monstrata Iwasa, 1992 | Japan |
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L. (L.) obscura Zhou & Yang, sp. nov. | China | present study |
L. (L.) omei Shatalkin, 1998 | China |
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We sincerely appreciate Allen L. Norrbom, Alyssa Seemann and Torsten Dikow (
The authors have declared that no competing interests exist.
No ethical statement was reported.
This work was supported by grants from the National Natural Science Foundation of China (no. 31970444) and the China Scholarship Council (no. 202106350076).
Conceptualization, Jiale Zhou and Ding Yang; writing – original draft preperation, Jiale Zhou and Ding Yang; writing – review and editing, Jiale Zhou and Ding Yang; material, Jiale Zhou and Ding Yang; visualization, Jiale Zhou; funding, Jiale Zhou and Ding Yang.
Jiale Zhou https://orcid.org/0000-0002-8901-3187
Ding Yang https://orcid.org/0000-0002-7685-3478
All of the data that support the findings of this study are available in the main text.