Revision of Japanese species of Nipponomyia Alexander, 1924 (Diptera, Pediciidae) Kolcsar, Levente-Peter Kato, Daichi Gamboa, Maribet Watanabe, Kozo ZooKeys 2020 1000 71 105 87JGG 2728202 D5C92A25-F65D-5116-B1B1-3813B178B06F Alexander, 1924 Alexander 1924 Insecta Pediciidae Nipponomyia CoL Animalia Nipponomyia Diptera 0 71 Arthropoda genus  Type species.   Tricyphona kuwanaiAlexander, 1913 by original designation in Alexander (1924): pages 158-159.  Descriptive notes on  NipponomyiaAlexander, 1924 based on Japanese species. 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. 2A, B). Eyes widely separated. Antenna short in both sexes, only little longer than head. Scape 1.2-1.4 xlonger and wider than pedicel. Pedicel 1.8-2.2 xwider than first flagellomere. Flagellum 11-13-segmented, evenly narrow toward apical segment. Flagellomeres oval to cylindrical, first 9 or 10 flagellomeres with 1 or 2 long, erected verticils dorsally (Fig. 2C, D). Last 3 or 4 flagellar segments with 3 or 4 verticils arranged irregularly. Last flagellar segment with 3 or 4 dark apical verticils, slightly curved upward, differing in shape to other verticils. Ventral part of flagellomeres densely covered with whitish sensilla, shorter than diameter of basal segment (Fig. 2E). Additional microtrichia on flagellomere (Fig. 2E). Palpi 5-segmented, length varying among species.   Figure 2.Characters of head parts of  Nipponomyia trispinosa(Alexander), SEM Acompound eye, 300 x(magnification) Bcompound eye, 1500 x Cantenna, 100 x Dflagellomeres 1 to 5, 300 x Eflagellomere 9, 700 x. 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. 3A, C). Cervical sclerite elongated fusiform. Pronotum well developed, medial part of antepronotum with hump and long setae; antepronotal lobe well developed, dorsal part slipping under medial part of antepronotum; postpronotum relative narrow. Prescutum with anterior part rounded, greatly protruding anteriorly, above to the pronotum in lateral view. Scutum usually with conspicuous spots. Presutural area of scutum without longitudinal suture, just with solid line of some long hairs (Fig. 3D); area under line of hairs before transverse suture bare in SEM photo (  N. trispinosa) (Fig. 3D); not evident under stereomicroscopes. Transverse suture deep, V-shaped, generally with dark patch in middle. Mediotergite elongated, dorsal margin almost straight in lateral view (Fig. 3A, C). Episternum, epimeron, and laterotergite each virtually not divided. Pit between episternum and epimeron deep (Fig. 3C). Meron relatively small, narrow in middle, forming two triangular parts, ventral one bigger. Metepisternum angular, additional divisions indistinct.   Figure 3.Characters of anterior body parts of  Nipponomyia trispinosa(Alexander) Adrawing, lateral view Bfirst two sternites, ventral view CSEM, lateral view Ddorsolateral view of presutural area of scutum, 150 x(magnification) Etarsal claves 500 x. 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. 3E). Average relative lengths of each segment (in percentage %) to the total length of corresponding leg (100%) listed in Table 1for both sexes.   Table 1.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. pentacanthaAlexander, 1958 (n = 4), and  N. yakushimensis Kolcsar& 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 Kolcsar& 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. 4A. Longitudinal veins with setae; crossveins bare. Sc long, ending beyond fork of Rs. Crossvein sc-r before origin of Rs and before or on same level as A2. Usually Rs forking into R2+3+4 and R5 (Fig. 4A-D, F) or rarely into R2+3 and R4+5 (Fig. 4E, G); highly variable within species. Crossvein r-m before fork of Rs, except in  N. khasianaAlexander, 1936. R1 and R3 approaching each other at position of R2. Cell r4 wider at middle. Usually cell d closed, longer than cell m2. Direction of crossvein m-m variable, usually almost perpendicular (Fig. 4A-E, G) or oblique (Fig. 4F). Anterior margin of wing with yellow band, bordered with different sized and shaped brown-black patches. Additional transverse markings (dashes, dots) in costal cell present in some species (Fig. 4B-E). Additional brown markings along veins, from fork of Rs to m-m and to m-cu (Fig. 4F, G).   Figure 4.  Nipponomyiawings Awing venation of  N. trispinosa(Alexander) B  N. kuwanai(Alexander) from Aomori Prefecture, Honshu C  N. kuwanaifrom Ishikari Mts, Hokkaido D  N. pentacanthaAlexander E  N. okinawensis Kolcsar& Kato, sp. nov. F  N. trispinosa(Alexander) G  N. yakushimensis Kolcsar& Kato, sp. nov.   Abdomen:Covered with relative long and dense hairs. Membranous area of second sternite well developed, shaped as in Fig. 3A, B. Usually tergites and sternites each with longitudinal dark line on lateral side (Figs 8B, 10B, 14B, 15B, 18B) and/or with spots and transverse lines (non-Japanese species).   Male terminalia:Relatively simple. Tergite 9 (epandrium) and sternite 9 (hypandrium) fused; border indistinct, forming wide ring, bulging in ventral side (Figs 5E, F, 11E, F, 16E, F, 19E, F). Tergite 9 simple without any lateral projections/arms. Gonocoxite well developed, stout, membranous on inner side. Basal lobe on ventral side of gonocoxite variable in size among species. Apical lobe of gonocoxite (sometimes referred to as outer gonostylus) partly separated from gonocoxite, elongated and directed dorso-ventrally, covered with short dark spines (Fig. 5A, B, G, H). Interbase long, well developed, fused with gonocoxite (Fig. 5G, H), with a few pale setae on ventral side. Gonostylus with two parts (Fig. 5B); inner (anterior) part of gonostylus always elongated, directed inwards; outer (posterior) part of gonostylus always shorter, wide (Figs 5G, H, 11G, H) or slender (Figs 16G, H, 19G, H) bearing 2-14 black spines. Aedeagus complex simple in shape as in most species of Pediciidae; difference among species more distinct in lateral view (Figs 5I, J, 11I, J, 16I, J, 19I, J). Aedeagus complex fused with sternite 9; relatively hard to separate from it; fused part referred in this article as aedeagal guide. Shape and length of aedeagus variable among species.   Figure 5.Male terminalia of  Nipponomyia pentacanthaAlexander A, Bdorsal view C, Dventral view E, Flateral view G, Hgonocoxite and gonostylus inner lateral view I, Jaedeagus 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. 6A, B). Pair of small pits situated between tergites 8 and 9. Tergite 8 at least twice as wide as tergite 9 in lateral view. Cercus longer than combined length of tergites 8-10. Cercus almost straight (Fig. 12C, E) or curving dorsally (Figs 6B, 12A). Hypogynal valve dorsally with 5-7 strong setae pointing caudally, terminal seta well separated from penultimate one and situated laterally to anterior setae (Fig. 6B, C). Genital fork well-developed, spoon-like or cruciform. Pair of membranous invaginations ("interbase sheath") present on ventral side of genital fork, holding interbases during copulation (Fig. 7B). Sternite 9/genital plate with two sclerites lateral of genital fork, variable in shape and development among species and even within species (Figs 7A, B, 12B, D, F). Pair of sclerotized (darker) area between genital fork and genital opening present in some species. Area around genital opening sclerotized, T- or Y-shaped; Three small, light brown spermathecae closely situated to genital opening (Figs 6D, 7C). Sternite 10 rounded apically, with 5-10 longer hairs (Figs 6D, 7B, 12B, D, F).   Figure 6.Female terminalia of  Nipponomyia pentacanthaAlexander Adorsal view Blateral view Cinner dorsal view of sternite 8 and hypogynial valve Dinner 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   Figure 7.Female terminalia of  Nipponomyia pentacanthaAlexander Alateral view Bsternite 9/genital plate and sternite 10, ventral view Cgenital 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.  Distribution. Eastern Palearctic and Oriental (Fig. 1).  Biology. 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 kuwanaiand  N. trispinosamales walk fast on the vegetation and fly short distances to find females.  Nipponomyia kuwanaifemales were observed ovipositing in muddy, wet soil, near mosses on a mountain lakeshore. A  N. trispinosafemale 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. pentacanthainhabit the same habitat.