Diogma glabrata (Meigen, 1818)

Kolcsar, Levente-Peter, Paramonov, Nikolai, Imada, Yume, Kato, Daichi, Gamboa, Maribet, Shinoka, Dai, Kato, Makoto & Watanabe, Kozo, 2022, Notes on the taxonomic status and distribution of some Cylindrotomidae (Diptera, Tipuloidea), with emphasis on Japanese species, ZooKeys 1083, pp. 13-88 : 13

Diogma glabrata (Meigen, 1818)

Figs 4C , 5C , 8B , 14 , 15 , 16

Phalacrocera megacauda in Alexander 1931: 349-350: original description.

Diogma glabrata megacauda in Alexander 1949: 196: comparison; Ishida 1955: 76: distribution; Takahashi 1960: 82: distribution, comparison, illustration; Alexander 1966: 122: distribution, faunistic records; Sidorenko 1999: 68-70: identification key, illustration, distribution.

Diogma glabrata ( Diogma glabrata Phalacrocera megacauda ) in Paramonov 2006: 888-889: identification key, illustration, distribution.

Diogma glabrata megacauda , D. glabrata in Gelhaus et al. 2007: synonymy, comparison, ecology, distribution, illustration.

Diogma glabrata in Nakamura 2014: 54: distribution; Imada 2020: biology and ecology of larvae.

Non-type material examined.

Belarus • 2 ♂, 1 ♀; Brest Oblast; Kamenets District, Belavezhskaya Pushcha National Park; 52.58807°N, 23.81746°E; alt. 160 m; 4 Aug. 1961; E.P. Narchuk leg.; ZIN. Denmark • 1 ♂;?Upilbo; 27 Jul. 1917; P. Nielsen leg.; USNM. Estonia • 1 ♀; Ida-Viru County, Narva-Jõesuu Town [Gungerburg]; 59.45°N, 28.03°E; 18 Jul. 1909; A.I. Chekini leg.; ZIN. Finland • 1 ♂, 2 ♀; Pihtipudas, Valkeispuro; 63.41082°N, 26.05336°E; alt. 170 m; 12 Jul. 2008 - 14 Aug. 2008; J. Salmela leg.; Malaise trap; LMM. • 1 ♂; Luvia, Niemenkyla; 61.39108°N, 21.56586°E; 12 m; 18 Jul. 2012; E. Viitanen leg.; CKLP. • 1 ♂; Mustasaari, Valassaaret; 63.43103°N, 21.07421°E; alt. 1 m; 2 Jul. 2019; E. Viitanen leg.; CKLP. • 1 ♀; Virolahti, Kurkela; 60.56858°N, 27.83847°E; alt. 8 m; 25 Jul. 2016; E. Viitanen leg.; CKLP. • 1 ♀; Fennia, Kb: 698:72, Ilomantsi, Tapionaho; 62.86016°N, 31.48371°E; alt. 190 m; 7 Jul. 1993 - 28 Jul. 1993; J.B. Jakovlev leg.; ZIN. • 1 ♀; Sotkamo, Iso-Matojarvi, Window trap № 3, Kn: 7086:590; 63.86638°N, 28.85971°E; alt. 210 m; 1 Jul. 1997 - 14 Jul 1997; Kuussaari leg.; ZIN. Japan • 2 ♂; Hokkaido, Higashikawa, Asahidake, River Yukomabetsu; 43.65226°N, 142.80229°E; alt. 1120 m; 23 Jul. 2019; L.-P. Kolcsár leg.; CKLP. • 4 ♂, 1 ♀; Hokkaido, Shari, Shiretoko Pass; 44.05331°N, 145.10166°E; alt. 716 m; 26 Jul. 2019; L.-P. Kolcsár leg.; CKLP. • 1 ♂; Iwate, Hachimantai, Toshiti Spa; 39.94253°N, 140.86804°E; alt. 1344 m; 3 Aug. 2013; • 2 ♀; same locality; 5 Aug. 2015; D. Kato leg.; BLKU. • 4 ♂; Nagano, Otakimura, Mt. Ontake; 35.86894°N, 137.51421°E; alt. 1990 m; 22 Jul. 2016; D. Kato leg.; BLKU. • 1 ♂; Toyama, Toyama, Arimine Jurodani; 36.46063°N, 137.42198°E; alt. 1130 m; 28 Aug. 2009 - 1 Sep. 2009; • 1 ♀; same locality; 1 Sep. 2009 - 8 Sep. 2009; M. Watanabe leg; Malaise trap; BLKU. Latvia • 1 ♀; Dolesmuiža, Doles sala; 56.866°N, 24.2014°E; alt. 4 m; 20 Jul. 2018; L.-P. Kolcsár leg.; CKLP. • 1 ♂, 1 ♀; Skaistkalne, small stream; 56.411°N, 24.637°E; alt. 12 m;; L.-P. Kolcsár leg.; birch-spruce forest; CKLP. Russia • 1 ♂; Altai Republic, Turochak District, near Artybash Settlement; 51.79299°N, 87.26535°E; alt. 430 m; 15 Jul. 2006; N.M. Paramonov leg.; ZIN. • 1 ♀; Amur Oblast, Shimanovsk District, Urochishche Samodon, 100 km W Svobodny City; 51.29°N, 126.83°E; alt. 320 m; 6 Aug. 1959; A.G. Zinovjev leg.; ZIN. • 1 ♂; Amur Oblast, Shimanovsk District, Simonovo Settlement, 75km W Svobodny City; 51.46°N, 126.98°E; alt. 305 m; 27 Jul. 1959; A.G. Zinovjev leg.; ZIN. • 1 ♂; Leningrad Oblast, Luga District, Jashhera Village; 58.89°N, 29.82°E; alt. 40 m; 23 Jul. 1963; A.A. Stackelberg leg.; ZIN. • 2 ♂, 1 ♀; Leningrad Oblast, Gobzhicy Village; 58.83°N, 30.13°E; 7 Jul. 1934 -16 Jul. 1934; A.A. Stackelberg leg.; ZIN. • 1 ♂, 2 ♀; Leningrad Oblast, Tolmachyovo Urban Locality; 58.86°N, 29.91°E, alt. 60 m; 16 Jul. 1935 - 26 Jul. 1935; A.A. Stackelberg leg.; ZIN. • 1 ♂; Leningrad Oblast, Kamenka River; 58.88°N, 29.76°E; alt. 65 m; 8 Jul. 1935; A.A. Stackelberg leg.; ZIN. • 2 ♂, 5 ♀, Leningrad Oblast, Vsevolozhsk District, Jukki Village; 60.11°N, 30.27°E; alt. 58 m; 13 Jul. 1931 - 22 Jul. 1933; A.A. Stackelberg leg.; ZIN. • 1 ♂; Leningrad Oblast, Vsevolozhsk District, Ostrovki Village; 59.81°N, 30.82°E; alt. 13 m; 21 Jun. 1906 - 22 Jun. 1906; G.G. Jakobson leg.; ZIN. • 1 ♂; Magadan Oblast, Magadan Urban Okrug, near Sokol Urban Settlement; 59.92°N, 150.71°E; alt. 177 m; 11 Jul. 2014 - 19 Jul. 2014; N.E. Vikhrev leg.; ZIN. • 2 ♀; Moscow Oblast, Naro-Fominsk District, Naro-Fominsk City, near Vostochnyy Community; 55.39094°N, 36.68878°E; alt. 195 m; 29 Jun. 2011; • 1 ♂; same locality, 29 Jun. 2014; D.I. Gavryushin leg.; ZIN. • 1 ♀; Moscow Oblast, Naro-Fominsky District, Vostochnyy [Oriental] settlement, within the settlement; 55.3741°N, 36.4984°E; alt. 205 m; 29 Jun. 2011; D.I. Gavryushin leg.; CKLP. • 1 ♂, Moscow Oblast, Naro-Fominsk, Nara River; 55.36075°N, 36.7404°E; alt. 174 m; 29. Jun. 2014; D.I. Gavryushin leg.; CKLP. • 1 ♀; Novgorod Oblast, Novgorod District, 1.5 km SE Glebovo Settlement; 58.54893°N, 31.83198°E; alt. 40 m; 2010; N.M. Paramonov leg.; ZIN. • 1 ♂; Primorsky Krai, Vladivostok City; 43.11553°N, 131.88548°E; alt. 20 m; 8 Aug. 2003; V.V. Sidorenko leg.; ZIN. • 1 ♂; Primorsky Krai, Chuguyevka District, Verchneussuri station; 44.067°N, 133.979°E; alt. 330 m; 30 Jul. 1979; A.G. Zinovjev leg.; ZIN. • 1 ♂; Primorsky Krai, Krasnoarmeysk District, Udegeyskaya Legenda National Park, apiary; 45.46052°N, 135.20451°E; alt. 700 m; 19 Jul. 2009; A.N. Ovtshinnikov leg.; ZIN. • 1 ♀; Primorsky Krai, Terney District, Terney Urban-type Settlement, Lower Serebryanka [Sanhobe] River; 45.09°N, 136.58°E; alt. 60 m; 6 Aug. 1941; K.J. Grunin leg.; ZIN. • 1 ♂; Primorsky Krai,Ussuriysk Urban Settlement, Gorno-Tajozhnoe Settlement, 25 km SE Ussuriysk; 43.69°N, 132.15°E; alt. 120 m; 3 Aug. 1963; E.P. Narchuk leg.; ZIN. • 1 ♀; Sakhalin Oblast, Severo-Kurilsky District, Kuril Islands, Paramushir Island, Rifovaya Bay; 50.4594°N, 156.0138°E; alt. 130 m; 30 Aug. 1999; A.S. Lelej, S.Y. Storozhenko leg.; ZIN. • 1 ♂; Sakhalin Oblast, Yuzhno-Kurilsk Urban Settlement, Kuril Islands, Kunashir Island, near Lagunnoe Lake; 44.062°N, 145.759°E; alt. 20 m; 25 Jul. 1955; N.A. Violovich leg.; ZIN. • 1 ♀; Sakhalin Oblast, Kuril Islands, Shikotan Island, near Malokurilskoye Village; 43.866°N, 146.827°E; alt. 30 m; 21 Aug. 1963; G.O. Krivoluckaja leg. ZIN. • 1 ♂; Sakhalin Oblast, Sakhalin Island, Yuzhno-Sakhalinsk City; 46.95°N, 142.73°E; alt. 50 m; 29 Jul. 1959; N.A. Violovich leg.; ZIN. • 1 ♀; Samara Oblast, Zhigulyovsk Urban Okrug, Zhiguli Nature Reserve, Bakhilova Polyana; 53.43543°N, 49.66252°E; alt. 45 m; 24 Jun. 2006; N.M. Paramonov leg.; ZIN. • 1 ♀; Tver Oblast, Udomlya District, 1,5 km NW Kaskovo Village; 57.98475°N, 35.03497°E; alt. 167 m; 19 Jul. 2017; A.G. Korobkov leg.; ZIN. • 1 ♂; Tver Oblast, Udomlya District, Moldino Settlement; 57.74807°N, 35.24965°E; alt. 155 m; 4 Jul. 2018; • 1 ♂; same locality; 5 Jul. 2018; A.G. Korobkov leg.; ZIN. • 1 ♂; Yaroslavl Oblast, Tutayev District, near former railway station Pustovo; 57.81438°N, 39.56016°E; alt. 122 m; 30 Jun. 2012; M.A. Klepikov leg.; pine forest, stream; ZIN.

Redescription.

Head. Dorsal part brown, ventral part yellowish brown (Fig. 14B, C ). Frons with white to yellowish-grey pubescence, visible only in dry specimens (Fig. 14C, F ). Rostrum yellowish brown, short without nasus; mouthparts pale brown to brown. Palpus pale brown to brown, 5 segmented; last segment slightly longer than penultimate segment (Fig. 14F ). Scape cylindrical 1.8-2 × longer than pedicel; pedicel ovate; pedicel and scape same colour or pedicel slightly darker (Figs 4C , 14E, F ); flagellum 14 segmented, gradually darkening from base to tip; flagellar segments simple in both sexes, not expanded ventrally; male flagellomere cylindrical, with short sparse whitish setae - sensilla, slightly denser in ventral and lateral sides; last segment 1.5-1.8 × longer than penultimate (Figs 4C , 14E ); female flagellomeres oval to cylindrical, first 4-6 flagellomeres oval, rest of segments cylindrical, sometimes all segment elongated, cylindrical as in male; only first 8-10 flagellomeres with sensilla; last flagellomere 1.8-2.8 × longer than penultimate (Figs 4C , 14F ); verticels black, shorter than length of flagellomere; generally one verticel in ventral surface and two or three in dorsal/dorsolateral sides of flagellomeres, first segment with 4-6 shorter verticels.

Thorax. General colouration yellowish with contrasting, shiny black markings. Pronotum yellow, middle part pale brown. Mesonotum yellow to pale brown with three separated black markings (Fig. 14C ) or one big black patch (Fig. 14D ). Scutellum yellow. Posterior part of mediotergite black (Fig. 14B ). Anepisternum and katepisternum well separated; ventral part of katepisternum black; ventral part of anepisternum yellowish (Japan, Fig. 14B ) or pale brown to brown (Finland, Russia). Ventral part of meron pale brown (Japan, Fig. 14B ) or brown to black (Finland, Russia). Laterotergite black at ventral corner. Coxa and trochanter yellow, femur pale brown; tibia gradually darkening from pale brown to dark brown/black; tarsus uniformly black; tarsomeres each with one spur. Wing hyaline; veins pale brown to brown; pterostigma pale; three branches of M reaching wing margin, M1 at same level as M1+2, cell a2 narrow,> 7 × longer than wide (Fig. 5C ); wing membrane with interference patterns, visible with dark background. Halter monochrome or knob darker, yellow to pale brown.

Abdomen. Tergites and sternites pale brown to brown, with paler longitudinal median line, poorly visible in dry specimens. Tergites and sternites 7 and 8 darker (Fig. 14A ). Pleural parts yellow to greenish yellow in living specimen.

Male terminalia. Large, black directed caudally (Fig. 14A ). Tergite 9 not fused with gonocoxite, partly cover gonocoxite (Fig. 15C ); medial part of tergite 9 rounded, with small tuft of hairs (Fig. 15A ); lateral lobe of tergite 9 greatly extended, complex, shorter than basal part of tergite 9; shape variable, rectangular to triangular in lateral view (Fig. 15C ); margin wavy, especially in caudal end (Fig. 15C , see also Gelhaus et al. 2007: figs 12-15); weakly curved inward in dorsal view (Fig. 15A ); ventral base of lateral lobe with small, black, heavily sclerotised lobe (named lamina by some authors) considerably variable in shape (Fig. 15E , see also, Gelhaus et al. 2007: figs 16-20, Takahashi 1960: figs 2, 3). Posterior margin of tergite 9 between median round part and lateral lobe with dense short, blunt ended setae (Fig. 15A ). Gonocoxite complex; apical and ventral lobe tips rounded with hairs; inner part of gonocoxite with less defined lobe, directed apically with hairs on tip (Fig. 15D ). Gonostylus simple, outer half wider (Fig. 15F ), with triangular, membranous lobe at inner side, poorly visible in dry specimens (Fig. 15G ). Sperm pump and ejaculatory apodeme small (Fig. 15H-J ), covered by parameres in lateral view (Fig. 15J ). Dorsal lobe between interbases complex, sclerotised (Fig. 15H, J ); interbase with ventral projection (Fig. 15J ). Aedeagus bifid, branches long, curved ventrally almost in right angle, then turned dorsally (Fig. 15J ).

Female terminalia. Brown, tip of cercus and hypopygial valve yellowish brown (Fig. 14G ). Tergite 8 separated at middle by membranous area (Fig. 16A ). Ventral corner of tergite 9 weakly rugged and with hairs (Fig. 16B ). Triangular sclerite separated from tip of tergite 10 (Fig. 16A ). Lateral lobe of tergite 10 relatively small, with long hairs (Fig. 16A, B ). Cercus and hypogynial valve simple, wide, blade-shaped, tips rounded (Fig. 16B ). Dorsal apical surface of cercus rough, formed by few blunt pyramid or round teeth (Fig. 16A, B ). Base of sternite 8 sclerotised (Fig. 16B ), lateral margins almost straight in ventral and inner dorsal view (Fig. 16C ), with transverse creases (Fig. 16B, C ). Two round spermathecae present, duct curved (Fig. 16D ). Lateral sclerite of genital fork triangular, two sperm ducts simple, without any clear markings (Fig. 16C ).

Distribution.

Austria, Belgium, Czech Rep., Denmark, Estonia, Finland, France, Germany, Great Britain, Ireland, Japan: Hokkaido I, Honshu I, Korea (North Korea or South Korea), Lithuania, Luxembourg, Netherlands, Norway, Poland, Romania, Slovakia, Sweden, Switzerland, and Russia (North European territory, Central European territory (Yaroslavl Oblast), Far East (Amur Oblast, Primorsky Krai, Sakhalin Oblast (Kuril Is: Kunashir I) ( Oosterbroek 2021).

First records from Belarus, Latvia, and Russia: Altai Republic, Amur Oblast, Novgorod Oblast, Magadan Oblast, Samara Oblast, and Kuril Islands (Shikotan I and Paramushir I). Occurrence data in Japan and surrounding areas are presented in Figure 8B .

Comments.

Diogma glabrata is a relatively common species in Europe, with a similar distribution to Cylindrotoma distinctissima . However, it is rarer or seemingly absent from southern Europe ( Kolcsár et al. 2018; Oosterbroek 2021). Alexander (1931) described the species Phalacrocera megacauda from Japan, based on the external morphological characters, without describing or illustrating the male terminalia. Later, Edwards (1938) designated a new genus, Diogma , for Cylindrotoma glabrata , which was previously included in Liogma by Osten Sacken (1859). Later, Alexander (1949) moved Phalacrocera megacauda to Diogma and mentioned it as a subspecies of Diogma glabrata , without detailing the difference or the reason for transferring it to subspecies rank. Takahashi (1960) illustrated the structural difference of the ventral lobe of tergite 9, called “lamina”, between Diogma glabrata megacauda , and Diogma glabrata glabrata . This lobe’s morphological variability was discussed and illustrated in detail by Gelhaus et al. (2007). They concluded that the two D. glabrata subspecies did not significantly differ in stable features and synonymised D. megacauda with D. glabrata . After morphological comparisons of the Japanese specimens with the West Palearctic specimens, our conclusion is the same. Only the body colouration shows slight differences between the two groups, however, colour variation is common among different populations of Cylindrotominae species. In this study, the European specimens are found to be genetically separated from the Japanese specimens, and the D. caudata sequences joined the Finnish D. glabrata clade. Additional sequences are needed for both Diogma species, from different areas of their distribution ranges, to resolve this genetic contradiction.

Figures

Figure 4. Antennae A Cylindrotoma distinctissima (Meigen, 1818) B Diogma caudata Takahashi, 1960 C Diogma glabrata (Meigen, 1818) D Liogma mikado (Alexander, 1919) E Liogma brevipecten Alexander, 1932 F Liogma serraticornis Alexander, 1919 G Triogma kuwanai (Alexander, 1913). Scale bar: 1 mm.

Figure 5. Wing A Cylindrotoma distinctissima (Meigen, 1818) B Diogma caudata Takahashi, 1960 C Diogma glabrata (Meigen, 1818) D Liogma mikado (Alexander, 1919) E Liogma brevipecten Alexander, 1932 F Liogma serraticornis Alexander, 1919 G Triogma kuwanai (Alexander, 1913) of " Triogma kuwanai " form H Triogma kuwanai (Alexander, 1913) of " Triogma limbinervis " form.

Figure 8. Occurrence data in Japan and surrounding areas of A Cylindrotoma distinctissima (Meigen, 1818) B Diogma glabrata (Meigen, 1818) and D. caudata Takahashi, 1960. Red dots indicate locations of investigated specimens, white dots indicate approximate locations of literature data. Green dot indicate approximate location of type locality of D. caudata Takahashi, 1960.

Figure 14. Diogma glabrata (Meigen, 1818) A habitus of male, lateral view (colouration of wings is artefact) B head and thorax, lateral view C head and thorax, dorsal view of pale form D thorax, dorsal view - dark form E antenna of male, dorsal view F head of female, lateral view G female terminalia lateral view.

Figure 15. Male genital structures of Diogma glabrata (Meigen, 1818) A terminalia, dorsal view B terminalia, ventral view C terminalia, lateral view D gonocoxite, inner lateral view E shape of heavily sclerotized lobe (lamina) of tergite 9 F shape of the gonostylus, caudal view G shape of the gonostylus, inner ventral view H aedeagus complex, dorsal view I aedeagus complex, ventral view J aedeagus complex, lateral view. Abbreviations: al - gonocoxite apical lobe; gsl - gonostylar lobe; vl - gonocoxite ventral lobe.

Figure 16. Female genital structures of Diogma glabrata (Meigen, 1818) A terminalia, dorsal view B terminalia, lateral view C sternite 8, hypogynial valve, genital fork, and sperm ducts, inner dorsal view D spermathecae.