Research Article |
Corresponding author: Levente-Péter Kolcsár ( kolcsar.peter@gmail.com ) Academic editor: Gunnar Kvifte
© 2020 Levente-Péter Kolcsár, Daichi Kato, Maribet Gamboa, Kozo Watanabe.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Kolcsár L-P, Kato D, Gamboa M, Watanabe K (2020) Revision of Japanese species of Nipponomyia Alexander, 1924 (Diptera, Pediciidae). ZooKeys 1000: 71-105. https://doi.org/10.3897/zookeys.1000.55021
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Japanese species of the genus Nipponomyia Alexander, 1924 are revised. Two new species, Nipponomyia okinawensis Kolcsár & Kato, sp. nov. and N. yakushimensis Kolcsár & Kato, sp. nov. are described from the Ryukyu Islands. Images of habitus and wings, illustrations of male and female terminalia, and distribution maps are provided for the Japanese species. A key to the world species of Nipponomyia is added. DNA barcodes of three Japanese Nipponomyia are provided, representing the first barcodes from the genus.
COI, crane flies, distribution, genitalia, new species, ovipositor, taxonomy, Tipuloidea
Nipponomyia Alexander, 1924 is a small crane fly genus belonging to the Pediciidae (Diptera: Tipuloidea). The genus was established based on the Japanese species Tricyphona kuwanai Alexander, 1913 and named after Japan (Nippon in Japanese). Another three species were included in the genus in the original designation, Tricyphona novempunctata (Senior-White, 1922) (originally described as Amalopis) from India, T. symphyletes Alexander, 1923 from Taiwan, and T. trispinosa Alexander, 1920 from Japan. Nipponomyia is morphologically characterized by the combination of the following characters: compound eye appearing bare, wing with a conspicuous yellow longitudinal stripe along the posterior margin, and gonostylus bearing 2–14 black chitinized spines (
The genus includes 15 species known from the Eastern Palearctic and Oriental regions so far (Fig.
In the present paper we review the genus and describe two new species, N. okinawensis Kolcsár & Kato, sp. nov. and N. yakushimensis Kolcsár & Kato, sp. nov. from Ryukyu Islands, Japan. A descriptive note of the genus, images of wings and habitus, and illustrations of male and female terminalia are presented. Additional faunistic records and distribution maps are reported for Japanese species. A key to the world species of the genus is provided based on information from the literature. Finally, we present DNA barcodes for N. kuwanai, N. trispinosa, and N. pentacantha Alexander, 1958 with GenBank accession numbers.
Fresh materials were collected using sweep nets and stored in 90% ethanol or were dry pinned. In total, 76 specimens of Nipponomyia belonging to five species were examined. Male and female terminalia were described and illustrated from observations in glycerol, after maceration in 10–15% KOH and neutralization with 3% acetic acid, both at room temperature. The cleared terminalia were preserved in terminalia tubes with glycerol. Illustrations were made in Adobe Photoshop CC 2019. Photographs of wing and body were taken using a Zeiss Stemi 508 stereomicroscope equipped with a Canon Kiss M digital camera. Those of terminalia were taken using a Leica M165C stereomicroscope equipped with a Leica MC170HD camera. Stack photos were combined using Zerene Stacker software. Scanning electron microscope photos were taken with a Topcon Electron Microscope SM-200. Morphological terminology in this study follows Cumming and Wood (2017), in the case of the wing venation we follow the venation system, based on
A specimen level dataset was made available as a Darwin Core Archive (http://ipt.pensoft.net/resource?r=nipponomyia) and is deposited in GBIF (https://doi.org/10.15468/tr5595).
Genomic DNA was individually extracted using DNease blood tissue kits (Qiagen GmbH, Hilden, Germany) following the manufacturer’s instructions. Extracted DNA was amplified using primers LCO-1490 and HCO-2198 (
Depositories
CKLP Private Collection of L.-P. Kolcsár.
Tricyphona kuwanai Alexander, 1913 by original designation in
General coloration yellow to black, with or without conspicuous marking on thorax. Markings of body not differing significantly between sexes.
Head: Rostrum short. Eye appearing bare; however, a few small setae present between ommatidia, near to border of compound eye (Fig.
Characters of head parts of Nipponomyia trispinosa (Alexander), SEM A compound eye, 300 × (magnification) B compound eye, 1500 × C antenna, 100 × D flagellomeres 1 to 5, 300 × E flagellomere 9, 700 ×. Abbreviations: mt – microtrichia, pe – pedicel, sc – scape, sen – sensilla, ve – verticel. Scale bars: 33.3 μm (A, D), 6.67 μm (B), 100 μm (C), 14.3 μm (E).
Thorax: Elongated in dorso-ventral direction (Fig.
Characters of anterior body parts of Nipponomyia trispinosa (Alexander) A drawing, lateral view B first two sternites, ventral view C SEM, lateral view D dorsolateral view of presutural area of scutum, 150× (magnification) E tarsal claves 500×. Abbreviations: aprn – antepronotum, bw – base of the wing, bh – base of the halter, cerv scl – cervical sclerite, cx – coxa, epm – epimeron, epst – episternum, lbl – labellum, ltg – laterotergite, mr – meron, ma – membranous area of sternite 2, mtg – mediotergite, mtast – metepisternum, p1–p5 – palpomeres, pprn – postpronotum, prsa – presutural area of scutum, posa – postsutural area of scutum, psct –prescutum, scp – scape, sct – scutum, sctl – scutellum, st – sternite, tg – tergite, tr – trochanter. Scale bars: 500 μm (C), 150 μm (D), 40 μm (E).
Legs: Longer in male than in female. Fore coxa elongated, extending ventrally beyond episternum. Tibia longest segment in both sexes. Male fore tarsomere 1 as long as fore femur or slightly longer. Tibial spur formula: 1, 2, 2, spurs just half length of width of tibia. Tarsomeres with 2 spurs. Male tarsomere 5 shorter than tarsomere 4. Female tarsomere 5 longer than tarsomere 4. Tarsal claw simple, without teeth, covered with small hairs on base, arolium present (Fig.
The average relative lengths of each segment (as a percentage) to the total length of the corresponding leg (100%). Male data are based on Nipponomyia kuwanai (Alexander, 1913) (n = 8), N. trispinosa (Alexander, 1920) (n = 7), N. pentacantha Alexander, 1958 (n = 4), and N. yakushimensis Kolcsár & Kato, sp. nov. (n = 2), Female data are based on Nipponomyia kuwanai (n = 4), N. trispinosa (n = 6), N. pentacantha (n = 2), and N. okinawensis Kolcsár & Kato, sp. nov. (n = 1).
Male | Female | |||||
---|---|---|---|---|---|---|
Fore | Mid | Hind | Fore | Mid | Hind | |
femur | 26.7 | 29.9 | 29.7 | 28.2 | 31.1 | 31.2 |
tibia | 30.4 | 31.4 | 32.5 | 32.0 | 32.9 | 34.4 |
tarsomere 1 | 28.6 | 24.1 | 23.2 | 26.2 | 22.2 | 20.8 |
tarsomere 2 | 7.9 | 7.9 | 7.7 | 7.0 | 6.9 | 6.7 |
tarsomere 3 | 3.6 | 3.8 | 3.9 | 3.5 | 3.3 | 3.5 |
tarsomere 4 | 1.5 | 1.6 | 1.6 | 1.5 | 1.6 | 1.6 |
tarsomere 5 | 1.3 | 1.4 | 1.4 | 1.7 | 2.0 | 1.9 |
Wing: General wing venation as in Fig.
Nipponomyia wings A wing venation of N. trispinosa (Alexander) B N. kuwanai (Alexander) from Aomori Prefecture, Honshu C N. kuwanai from Ishikari Mts, Hokkaido D N. pentacantha Alexander E N. okinawensis Kolcsár & Kato, sp. nov. F N. trispinosa (Alexander) G N. yakushimensis Kolcsár & Kato, sp. nov.
Abdomen: Covered with relative long and dense hairs. Membranous area of second sternite well developed, shaped as in Fig.
Male terminalia: Relatively simple. Tergite 9 (epandrium) and sternite 9 (hypandrium) fused; border indistinct, forming wide ring, bulging in ventral side (Figs
Male terminalia of Nipponomyia pentacantha Alexander A, B dorsal view C, D ventral view E, F lateral view G, H gonocoxite and gonostylus inner lateral view I, J aedeagus complex lateral view. Abbreviations: ae – aedeagus, aeg – aedeagal guide, algc – apical lobe of gonocoxite, gc – gonocoxite, gs – gonostylus, ib – interbase, igs – inner part of gonostylus, ogs – outer part of gonostylus, st9 – sternite 9, tg9 – tergite 9, vlgc – ventrobasal lobe of gonocoxite. Scale bars: 0.5 mm (A–F), 0.2 mm (G, H), 0.2 mm (I, J).
Female terminalia, ovipositor: Elongated, tergites 8–10 fused (Fig.
Female terminalia of Nipponomyia pentacantha Alexander A dorsal view B lateral view C inner dorsal view of sternite 8 and hypogynial valve D inner ventral view of sternites 9 and 10 and cerci. Abbreviations: cr – cercus, hyp vlv – hypogynial valve, st9/gp – sternite 9/genital plate, st10 – sternite 10, tg – tergite
Female terminalia of Nipponomyia pentacantha Alexander A lateral view B sternite 9/genital plate and sternite 10, ventral view C genital opening and spermathecae. Abbreviations: gf – genital fork, go – genital opening, ibs – interbase sheath, ls – lateral sclerite, sa – sclerotized area, spt – spermatheca.
Larva: Unknown.
Pupa: Unknown.
Eastern Palearctic and Oriental (Fig.
Adults swarm in the air close to the ground or above the vegetation, in shadow and windless conditions. They rest on ventral surfaces of substrates like leaves, spreading their wings horizontally, even during copulation. Nipponomyia kuwanai and N. trispinosa males walk fast on the vegetation and fly short distances to find females. Nipponomyia kuwanai females were observed ovipositing in muddy, wet soil, near mosses on a mountain lakeshore. A N. trispinosa female was observed searching for oviposition sites around wet soil, rich of organic matter next to a waterfall, but the oviposition has not yet been observed. Sometimes N. kuwanai, N. trispinosa, and N. pentacantha inhabit the same habitat.
Species groups
Japanese species of the genus can be classified into two morphological species groups. The kuwanai species group is characterized by the presence of black transverse lines (dashes) in costal cell (Fig.
Pairwise distances between species
We successfully extracted and amplified COI barcode sequence from the three previously described species, Nipponomyia kuwanai (GenBank: MT874511), N. trispinosa (MT874512, MT874513), and N. pentacantha (MT874514). However, attempts to extract DNA from the type specimens of the two new species were unsuccessful. The pairwise genetic distance between species using Kimura 2-parameter ranged between 13.1% and 15.3%, the overall genetic distance is 14.2% (Table
Nipponomyia pentacantha: Alexander, 1958: 293–294, plate 3, figs 14, 17 – original description, wing and male terminalia illustration;
Holotype male: Japan, Nagano, Echigo, Mount Amakazari; alt. 300–600 m; 25–26 Jun. 1955; Baba leg. Paratype male: same location; alt. 300 m; 26 Jun. 1955; Baba leg. Type specimens deposited in National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA; not studied.
Non-types: Japan: [Honshu] • 2♂; Aomori, Nishimeya, Shirakami Nature observation garden, Kawaratai; alt. 255 m; 40°31.13'N, 140°12.89'E; 4 Jul. 2013; leg. D. Kato (pinned,
Yellowish species with 11 dark spots on thorax (7 dark spots in N. okinawensis Kolcsár & Kato, sp. nov., 11–13 dark spots in N. kuwanai). Wing with transverse dark lines in costal cell. Brown marking extending from R2+3 to crossvein m-m (brown marking extending from R2+3 to maximum to base of M1 in N. kuwanai and N. okinawensis Kolcsár & Kato, sp. nov.). Sternite 2 without black marking at corner of membranous area (with black marking at corner of membranous area in N. kuwanai), a diffuse line positioned same level as line on sternite 3 (N. kuwanai without this line, N. okinawensis Kolcsár & Kato, sp. nov. with any line and dark marking on sternite 2). Gonostylus with 4 or 5 spines (11–14 spines in N. kuwanai). Aedeagus short, twice as long as wide, rounded (as long as wide in N. kuwanai). Cercus curved upward (straight in N. okinawensis Kolcsár & Kato, sp. nov.). Female genital opening Y-shaped (T-shaped in N. okinawensis Kolcsár & Kato, sp. nov.), lateral sclerite 1/3 of length of genital fork (less than 1/5 of length of genital fork in N. kuwanai and less than 1/6–1/7 of length of genital fork in N. okinawensis Kolcsár & Kato, sp. nov.), genital fork cross-shaped (spoon-shaped in N. kuwanai, cross-shaped in N. okinawensis Kolcsár & Kato, sp. nov. but lateral branch curved caudally).
Body length: male 9.5–11 mm, female: 12–13 mm.
Wing length: male 9.5–10.5 mm, female 10–11 mm.
Head: Light brown to brown (Fig.
Thorax: In dry specimens general coloration yellow (Fig.
Legs: General coloration yellow, covered with yellowish setae (Fig.
Wing: As in Fig.
Abdomen: Abdomen covered with comparatively long pale setae. Tergites 2–6 in both sexes, each with longitudinal narrow black line on lateral side, situated on basal 1/3–1/2 of each tergite in male (Fig.
Male terminalia: dark yellow to brown, always darkest part of abdomen (Fig.
Female terminalia, ovipositor: General coloration dark yellow. Cercus curved upward (Fig.
Larva: Unknown.
Pupa: Unknown.
Japan: Honshu Island (
Distributions data of Nipponomyia species: for N. pentacantha Alexander white circles designate literature data while red circles are new data obtained in this study; N. gracilis Savchenko (yellow pentagon); N. yakushimensis Kolcsár & Kato, sp. nov. green triangle; N. okinawensis Kolcsár & Kato, sp. nov. blue square.
Middle of June to middle of September.
Tricyphona kuwanai:
Nipponomyia kuwanai:
Holotype female: Japan, Tokyo; 7 May 1912; S.I. Kuwana leg. Type specimens deposited in National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA; not examined.
Allotype male: Japan, Tokyo, Meguro; 15 Apr. 1919; R. Takahashi leg. Type specimens deposited in National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA; not examined.
Non-types: Japan: [Honshu] • 1♂; Aomori, Hirosaki, Ichinowatari-washinosu; alt. 205 m; 40°31.15'N, 140°26.33'E; 17 Jun. 2013; D. Kato leg. (pinned,
Yellowish species with 11–13 dark spots on thorax (7 dark spots in N. okinawensis Kolcsár & Kato, sp. nov., 11 dark spots in N. pentacantha). Wing with transverse dark lines in costal cell. Brown marking extending from base of R2+3 to base of M1, often not reaching M1 (brown marking extending from R2+3 to base of M1 in N. okinawensis Kolcsár & Kato, sp. nov. and to m-m in N. pentacantha). Second sternite with black marking at corner of membranous area (without this marking in N. pentacantha and N. okinawensis Kolcsár & Kato, sp. nov.), and without other line (a diffuse line positioned same level as line on sternite 3 in N. pentacantha). Gonostylus with 11–14 spines (4 or 5 spines in N. pentacantha). Aedeagus short, as long as wide in lateral view, tip rounded (twice as long as wide in N. pentacantha). Cercus curved upward (straight in N. okinawensis Kolcsár & Kato, sp. nov.). Genital opening Y-shaped (T-shaped in N. okinawensis Kolcsár & Kato, sp. nov.), lateral sclerite less than 1/5 of length of genital fork (1/3 of length of genital fork in N. pentacantha and less than 1/6–1/7 of length of genital fork in N. okinawensis Kolcsár & Kato, sp. nov.), genital fork spoon-shaped (cross-shaped in N. pentacantha and N. okinawensis Kolcsár & Kato, sp. nov.).
Body length: male 9.5–11 mm, female: 12–14 mm.
Wing length: male 9–12 mm, female 9.5–11.5 mm.
Head: Brown with grayish pruinosity (Fig.
Thorax: General coloration yellow for specimens in alcohol, dark yellow, with reddish shade in dry specimens, dorsal parts light brown (Fig.
Legs: General coloration yellow, covered with yellowish setae. Femora without apical darkened area, apical part of tibia slightly brownish, with darker setae. Apical ends of tarsomeres 1–3 each with narrow brown to dark brown ring, tarsomeres 4 and 5 light brown to brown (Fig.
Wing: As in Fig.
Abdomen: Abdomen covered with relatively long pale setae, dorsal setae darker than ventral ones. Tergites 2–6 (male) and 2–7 (female) each with a longitudinal narrow black line on lateral side, its length 1/2 of tergite in male (Fig.
Male terminalia: Dark yellow to brown (Fig.
Female terminalia, ovipositor: General coloration dark yellow. Cercus curved upward (Fig.
Larva: Unknown.
Pupa: Unknown.
The species flies from April to early August.
Holotype
♀, pinned. Original label: “Japan, Okinawa Island, Okinawa, Kunigami, Mt Fuenchiji-dake, Yona; alt. 250 m; 26°44.93'N, 128°14.54'E; 21 May 2016; D. Kato leg.” “Holotype Nipponomyia okinawensis Kolcsár & Kato, sp. nov. [red label]” (
Anterior part of thorax dark brown to black, posterior part yellowish brown (yellow in N. pentacantha and N. kuwanai), abdomen yellow. Thorax with 7 darker patches (11 in N. pentacantha and 11–13 in N. kuwanai), 2 in presutural area of scutum. Wing with transverse dark lines in costal cell. Brown marking extending from R2+3 to base of M1 (brown marking usually not extending to base of M1 in N. kuwanai and extending to crossvein m-m in N. pentacantha). Second sternite without dark line (with black marking at corner of membranous area in N. kuwanai and with a diffuse line positioned same level as line on sternite 3 in N. pentacantha). Cercus straight (curved upward in N. kuwanai and N. pentacantha). Genital opening T-shaped (Y-shaped in N. kuwanai and N. pentacantha), lateral sclerite very small, less than 1/6–1/7 of length of genital fork (1/3 of length of genital fork in N. pentacantha and less than 1/5 of length of genital fork in N. kuwanai). Genital fork cross-shaped, lateral branch curved caudally (spoon-shaped in N. kuwanai and cross-shaped in N. pentacantha but lateral branch almost straight).
Body length: female: 12 mm.
Wing length: female 10 mm.
Head: General coloration brown (Fig.
Thorax: Apical half of thorax dark brown, almost black, partly due to decay inside, posterior part yellowish brown (Fig.
Legs: General coloration yellow, covered with yellowish setae. Femora without apical dark area, tip of tibiae with a narrow darker ring. Apical ends of tarsomeres 1 to 4 each with narrow dark yellow to light brown ring. Tarsomeres 4 and 5 yellowish (Fig.
Wings: As in Fig.
Abdomen: Yellow to light brown, relatively short setae dark on tergites and pale on sternites. Tergites 2–6, each with longitudinal narrow black line on lateral side, 1/4–1/3 length of tergite length, less prominent compared to other species. Sternite 2 without dark mark. Sternites 3–5 each with narrow brown line, not continuous in sternite 3 (Fig.
Female terminalia, ovipositor: General coloration dark yellow (Fig.
Male: Unknown.
Larva: Unknown.
Pupa: Unknown.
Japan: Ryukyu Islands: Okinawa Island (Fig.
Type specimen collected at the end of May.
Okinawa Island is the largest island of the Ryukyu Archipelago, located roughly midway between Kyushu and Taiwan. The island was formed by complex process of Paleogene volcanic activities and Neogene-Quaternary sedimentations and reef deposits (
Tricyphona trispinosa:
Nipponomyia trispinosa:
Holotype male: Japan, Honshu, leg. Akio Nohiro. – without further data, probably Kyoto (see
Non-types: Japan: [Hokkaido] • 2♂, (GenBank # MT874512); Hokkaido, Sobetsu, River Benkei; alt. 238 m; 42°33.52'N, 140°59.29'E; 29 Jul. 2019; L.-P. Kolcsár leg. (pinned or in ethanol, CKLP). [Honshu] • 3♂, 1♀; Aomori, Hirosaki, Ichinowatari-washinosu; alt. 205 m; 40°31.15'N, 140°26.33'E; 5 Sep. 2013; D. Kato leg. (pinned,
Yellowish species with 11 darker spots on thorax (N. yakushimensis Kolcsár & Kato, sp. nov. dark yellow species with 11 large dark spots, N. gracilis without dark spots on thorax). Wing without transverse dark line in costal cell. Brown band running from base of R2+3 to tip of M4 and to m-cu (brown band not reaching wing margin in N. yakushimensis Kolcsár & Kato, sp. nov.). Dark band along crossveins r-m and m-cu conspicuous. Second sternite with black marking at corner of membranous area, but without other line. Gonostylus with 3 spines (2 spines in N. yakushimensis Kolcsár & Kato, sp. nov. and N. gracilis), aedeagus short, triangular, and acute at tip in lateral view (aedeagus long, rod-shaped in N. yakushimensis Kolcsár & Kato, sp. nov. and N. gracilis). Cercus long and straight, just gently curved upward. Genital opening T-shaped, genital fork cross-shaped, lateral sclerite large, half as long as genital fork.
Body length: male 8.5–12 mm, female: 13–15 mm.
Wing length: male 8.5–12 mm, female 11–12 mm.
Head: Yellowish brown (Fig.
Thorax: Specimens stored in ethanol whitish yellow. General coloration yellow, dorsal parts somewhat darker yellow in pinned specimens (Fig.
Legs: General coloration yellow, covered with yellowish setae. Femora without apical dark area, apical part of tibiae brownish, with a few darker setae. Apical ends of tarsomeres light brown. Apical half of tarsomere 4 brown, tarsomere 5 slightly lighter than tarsomere 4 (Fig.
Wing: As in Fig.
Abdomen: Abdomen covered with relatively long pale setae. Tergites 2–6 in male and 2–7 in female each with a longitudinal narrow black line on lateral side, its length ranging from 1/3–1/2 of tergite length (Fig.
Male terminalia: Dark yellow to brown (Fig.
Female terminalia, ovipositor: General coloration dark yellow. Cercus almost straight, only weakly curved upward (Fig.
Larva: Unknown.
Pupa: Unknown.
Usually flying between the end of July and the middle of November, but also collected in May.
Holotype
♂, pinned. Original label: “Japan, Kagoshima, Yakushima Island, Yakushima, near Shirataniunsui-kyo Valley, Yakushima-cho, alt. 600 m, 30°23.04'N, 130°34.37'E, 25 Apr. 2018, D. Kato leg.” “Holotype Nipponomyia yakushimensis Kolcsár & Kato, sp. nov. [red label]” (pinned,
Paratype
♂, same data as holotype (pinned,
Dark yellow species with 11 large darker spots on thorax (N. trispinosa light yellowish species with 11 smaller dark spots, N. gracilis brownish species without any dark spots on thorax). Wing without transverse dark line on costal cell. Brown band running from base of R2+3 to crossvein m-m, but not reaching wing margin (reaching the wing margin in N. trispinosa). Brown band along crossveins r-m and m-cu conspicuous. Second sternite with black marking at corner of membranous area, but without other line. Gonostylus with 2 spines (3 spines in N. trispinosa), aedeagus long, rod-shaped and acute at tip (aedeagus short, triangular in N. trispinosa).
Body length: male 8–8.5 mm.
Wing length: male 8–8.5 mm.
Head: Light brown to brown with grayish pruinosity (Fig.
Thorax: General coloration dark yellow, dorsal parts somewhat darker (Fig.
Legs: General coloration yellow, covered with yellowish setae. Femora without clear apical dark area, but with some darker setae. Apical part of tibiae light brown, with a few darker setae. Apical ends of tarsomeres narrowly dark yellow to light brown (Fig.
Wing: As in Fig.
Abdomen: Abdomen covered with relative long dark setae. Tergites 2–6 each with a longitudinal narrow black line on lateral side, its length ranging from 1/2–3/4 of tergite length. Sternite 2 with a short black line at corner of membranous fold. Sternites 3–7 each with a broad brown patch, covering anterior half of segment (Fig.
Male terminalia: Dark yellow to light brown (Fig.
Female: Unknown.
Larva: Unknown.
Pupa: Unknown.
Japan: Ryukyu Islands: Yakushima Island (Fig.
Type specimens were collected at the end of April.
Yakushima Island is one of the northmost members of Ryukyu Islands, and also the largest island of the Osumi Archipelago. Yakushima is located approximately 70 km south of Kyushu and formed by a combination of sedimentary and orogenic volcanism processes (
1 | Costal cell of wing without black transverse lines or dark points (similar to Fig. |
2 |
– | A series of black transverse lines or points in costal cell of wing (Fig. |
9 |
2 | Presutural area of scutum with brown-black spots (Figs |
3 |
– | Presutural area of scutum without any markings | 7 |
3 | Presutural area of scutum with 2 large black spots, separated by a thin yellow line, post sutural area of scutum almost black; wing with cloudy area around end of vein A2 | N. sumatrana (de Meijere, 1924) |
– | Presutural area of scutum with 4 spots (similar to Figs |
4 |
4 | Tip of the wing yellow, without darker bordering patches; only a small patch around crossvein m-cu | N. khasiana Alexander, 1936 |
– | Tip of wing darker, yellowish pattern bordered with darker patches; distinct pattern on wing as in Fig. |
5 |
5 | In addition to lateral black line on tergite, tergites with 4 brown spots on basal half; sternite with 2 lateral longitudinal lines, base of gonostylus relatively long before forking, outer part with 3 or 4 spines | N. mannheimsiana Alexander, 1969 |
– | No additional spot on tergite; only 1 dark line on lateral side of sternite; gonostylus with basal part relatively short | 6 |
6 | Lateral black line on tergite shorter than half length of corresponding tergite; brown line on sternite small (Fig. |
N. trispinosa (Alexander, 1920) |
– | Lateral black line on tergite longer than half length of corresponding tergite; line on sternite forming a broad patch (Fig. |
N. yakushimensis Kolcsár & Kato, sp. nov. |
7 | Femora and tibiae uniformly yellow; gonostylus with 3 or 4 spines | N. szechwanensis Alexander, 1935 |
– | Femora and tibiae each with dark ring at tips; gonostylus with 2 spines | 8 |
8 | Presutural area of scutum brownish, wing with typical pattern of genus, similar to Fig. |
N. gracilis Savchenko, 1983 |
– | Presutural area of scutum light yellow; wing with 5 or 6 darker and 5 or 6 paler spots, darker spots around base of wing, at sc-r, origin of Rs, tip of Sc, and m-cu | N. flavicollis Edwards, 1933 |
9 | Thorax, abdomen, and legs uniformly black | N. nigrocorporis Alexander, 1944 |
– | Yellowish species, with dark markings on thorax and abdomen | 10 |
10 | Dark markings on costal cell spot-shaped, not forming clear transverse lines | 11 |
– | Costal cell with clear transverse lines as in Fig. |
12 |
11 | Scutellum brown, posterior half of mediotergite brown, tip of femora darkened | N. kamengensis Alexander, 1967 |
– | Scutellum and mediotergite yellow, tip of femora not darkened | N. kulingensis Alexander, 1937 |
12 | Central dark spot on scutal suture absent | N. joshii Alexander, 1957 |
– | Central dark spot on scutal suture present (Figs |
13 |
13 | Tip of femora darkened, a dark cloud at tip of vein A2, extending to vein A1; pattern on tip of wing brown | N. novempunctata (Senior-White, 1922) |
– | Femora yellow; tip of A2 without dark patch; tip of wing yellowish with brown border | 14 |
14 | Legs uniformly light yellow, segments not distinctly darkened at tip | N. symphyletes (Alexander, 1923) |
– | Legs light yellow, segments except femora clearly darkened at tips (Figs |
15 |
15 | Presutural area of scutum with a large spot on each lateral side (Fig. |
N . okinawensis Kolcsár & Kato, sp. nov. |
– | Presutural area of scutum with 4 dark spots (Figs |
16 |
16 | Second sternite with a relatively dark, submarginal line, no line at corner of membranous area (Fig. |
N. pentacantha Alexander, 1958 |
– | Second sternite without submarginal line, but with line at corner of membranous area (Fig. |
N. kuwanai (Alexander, 1913) |
We would like to thank Moritz Fahldieck and Jukka Salmela for reviewing the manuscript and the subject editor of ZooKeys, Gunnar Mikalsen Kvifte, for providing constructive comments. L.-P. Kolcsár was supported by the Japan Society for the Promotion of Science (JSPS) Postdoctoral Fellowships for Research in Japan (Short-term Program) (Grant Number: PE18038). This research was financially supported by the JSPS Grant-in-Aid for Scientific Research (Grant Numbers: 19K21996, 19H02276).