Orthonevra brevicornis ( Loew, 1843 )

Orthonevra brevicornis ( Loew, 1843) View in CoL

( Figs 1C–D View Fig , 2C–D View Fig , 3B View Fig , 4B View Fig , 5D–F View Fig , 6A–C View Fig , 7F–J View Fig , 9B, 9C View Fig )

Chrysogaster brevicornis Loew, 1843: 269 View in CoL (description of a male, drawings of the postpedicel, face profile and wing). Type locality: Poland, Posen [= Poznań].

Chrysogaster brevicornis View in CoL : BFCKFR (1921): 14 (in part: Schlesien /Sobten ( Poland) and Berlin ( Germany) are O. brevicornis View in CoL ; but Gastein ( Austria) are O. tristis View in CoL and O. montana View in CoL ; Schlesien /Wölfelsfall ( Poland) is O. montana View in CoL ; Schweiz (= Switzerland) are O. tristis View in CoL ); BAŇKowsKA (1963): 147–148 (short description in the key, drawings of wing and postpedicel).

Orthoneura brevicornis ( Loew, 1843) : LoFw (1857): 7 (description of a female, in part: Meseritz [= Międzyrzecz], Poland); SACK (1930): 23 (in part, short description in the key); STACKFLBFRG (1970): 54–55 (short description in the key and drawing of male terminalia).

Type material. NFoTYPF: J ( Figs 1C View Fig , 2C–D View Fig , 3B View Fig , 4B View Fig , 6A View Fig , 7F View Fig ), labelled: “ Poland XT 19 // Wielkopolski Nat. Park // 52°18’25’’N 16°40’14’’E // “Trzcielińskie Bagno” // 10 / V / 2006 // P. Trzciński leg.” and with our red neotype designation label “ NEOTYPE J // Chrystogaster brevicornis Loew, 1843 // design. R. Żóralski & F. Van de Meutter, 2024 ”. Matches type locality and all characters of original description by LoFw (1843), deposited in ZMHB.

Additional material studied. Voucher specimens. GERMANY: Frankfurt am Main [“Frankf. a/M. // Coll. H. Loew // 12930 // Typus (red label) // Orthoneura brevicornis m // NOT A TYPE locality mism. R. Żóralski, 2023 ”], 1 J ( Figs 7H View Fig , 9B, 9C View Fig ) ( ZMHB, ex coll. H. Loew). POLAND: 1 ♀ ( Figs 1D View Fig , 7G View Fig ), labelled “Poland XT 19 // Wielkopolski Nat. Park // 52°18’25’’N 16°40’14’’E // “Trzcielińskie Bagno” // 10 / V / 2006 // P. Trzciński leg.” and “VOUCHER SPECIMEN ♀ // Orthoneura brevicornis ( Loew, 1843) // det. R. Żóralski // & F. Van de Meutter, 2024 ”, intentionally designated voucher specimen to unambiguously reflect characters of opposite sex of the neotype, matches all characters of original description of female (LoFw 1857), caught together with neotype, deposited in ZMHB; 2 JJ, bearing our label “VOUCHER SPECIMEN J // Orthonevra brevicornis ( Loew, 1843) // det. R. Żóralski // & F. Van de Meutter, 2024 ”: Jeleń, 4.v.1986, 1 J ( Figs 5D, 5E, 5F View Fig , 6B View Fig , 7I View Fig ), leg. B. Soszyński ( USMB); Mrzezino, near Reda river [CF35], 29.iv.2019, 1 J ( Fig.6C View Fig ) (GenBank acc. no. OR859720), leg. RZ ( MZPW). GEORGIA: Dviri, 12.v.2022, 1 J ( Fig. 6J View Fig ), leg. FM ( RBINS).

Other specimens. 229 JJ 96 ♀♀. CZECH REPUBLIC: Veltrusy- Obora, deciduous forest, 7.v.1985, 1 J, leg. Barták ( ZFMK). GERMANY: Berlin [= Berlin; Berlin 55949.], 15.v.1906, 1 ♀ ( ZMHB, ex. coll. T. Becker); Berlin [= Berlin; Orth. brevicornis Lw. ; det. Oldenberg; Berlin 55949.], 25.v.1906, 1 ♀ ( ZMHB, ex. coll. T. Becker). Berlin, Schildhorn, vi.1948, 1 ♀ ( ZMHB, ex coll. A. Riedel). GREECE: Grevena, 505 m, 40.015N 21.395E, 8.v.2022, 1 J (GenBank acc. no. OR859727), leg. FM ( FMTB). HUNGARY: Hild, date unknown, 1 J, leg. Thalhammer ( ZFMK). POLAND: Ślęża Mtn.[= Zobten 12310], 3.v, ( ZMHB, ex coll. T. Becker).Niemcza [= Nimpsch Schles.Duda; O. plumbago Lw. ], 14.v.1908, 1 J 1 ♀, 17.v.1908, 1 ♀, 23.v.1908, 3 ♀♀, 26.v.1908, 1 ♀, 13.vi.1910, 1 ♀, 14.v.1912, 1 J 1 ♀, ( ZMHB, ex coll. O. Duda); Jędrzejów, oddz. 210, 3.v.1954, 1 ♀, on Padus avium , leg. J. Karczewski ( MZPW) (TRoIANo- wA- BAŇKowsKA 1959, BAŇKowsKA 1961, as O. frontalis ); Zdziar-Łopatki ad. Staroźreby, alder carr near Płonka river, DD33, 17.v.1971, 1 J 2 ♀♀, leg. J. K. Kowalczyk ( RZRP); Aleksandrów Łódzki, 1.v.1973, 1 ♀, leg. B. Soszyński ( RZRP); Rogów ad Łódź, 27.v.1975, 1 J, leg. B. Soszyński ( RZRP); Świętokrzyski National Park: Święty Krzyż, 2.vi.1979, 1 ♀, leg. J. K. Kowalczyk ( RZRP) (ŻÓRALsKl et al. 2017); Teofilów, DC40, 1.v.1983, 1 ♀, leg. B. Soszyński ( RZRP); Spalski Landscape Park: Konewka, DC41, 19.v.1985, 1 ♀, leg. B. Soszyński ( RZRP), 13.v.2017, 1 ♀, leg. M. Soszyński ( RZRP); Łagiewnicki Forest: Marianka, 4.v.1995, 1 ♀, leg. B. Soszyński ( RZRP); Biebrza National Park: Sztabin, 17.v.2003, 1 J, leg. Wanat M. ( RZRP); Wiry, XT19, 15.v.2004, 1 J, leg. P. Trzciński ( PTPP); Biebrza National Park: Grobla Honczarowska, 8.vi.2006, 1 ♀, Malaise trap, leg. J. Sawoniewicz ( RZRP); Trzcielińskie Bagno [XT19], 1.–10.v.2008, 192 JJ (GenBank acc. no. OR859703, OR859667, OR859676, OR859690, OR859714, OR859726, OR859728, OR859700, OR859693), 54 ♀♀, Moericke traps (part of it: 125 JJ and 13 ♀♀ continued to be stored in alcohol),leg.P.Trzciński ( PTPP, RZRP), 1.–10.v.2008, 2 JJ 1 ♀, Moericke traps, leg. P. Trzciński ( ZFMK), 1.–10.v.2008, 1 J 1 ♀, Moericke traps, leg. P. Trzciński ( SBHN), 14.v.2008, 1 J, leg. P. Trzciński ( PTPP), 23.v.2008, 1 J, leg. P.Trzciński ( PTPP), 24.iv.2010, 5 JJ 4 ♀♀, leg. P. Trzciński ( PTPP); Oleśnickie Jodły, CC 64, 24.iv.2009, 1 ♀, leg. B. Soszyński ( RZRP); Kuzki ad Włoszczowa, 3.v.2012, 1 J, leg. Ł. Mielczarek ( LMMP);Spalski Landscape Park:Spała, DC41, 24.iv.2015, 1 J, leg.RZ ( RZRP) (WlTFK et al. 2015).Spalski Landscape Park: Ceteń, DC50, 4.iv.2014, 1 J, leg.M. Soszyński ( RZRP), 22.iv.2014, 1 J, leg. Ł. Mielczarek ( LMMP), 27.iv.2015, 1 ♀, leg.B.Soszyński ( RZRP) (WlTFK et al.2015), 27.iv.2015, 1 J, leg.RZ ( RZRP) (WlTFK et al.2015), 27.iv.2015, 1 J 1 ♀, leg. P. Trzciński ( PTPP) (WlTFK et al. 2015), 9.v.2015, 1 ♀, leg. M. Soszyński ( RZRP), 25.v.2015, 1 J, leg. Ł. Mielczarek ( LMMP); Suwalski Landscape Park: Sidorówka, 22.v.2015, 1 ♀, leg.Ł. Mielczarek ( LMMP); Karniowce ad Trzebinia, 19.vi.2015, 1 J, leg. Ł. Mielczarek ( LMMP); Krynica Morska, 7.v.2016, 1 J, leg. Ł. Mielczarek ( LMMP); Nowa Wieś near Maniówka river, 24.v.2016, 1 ♀, leg. RZ ( RZRP) (ŻÓRALsKl et al. 2016); Wiatrołuża III, peatbog, 24.v.2016, 1 J, leg. Ł. Mielczarek ( LMMP) (ŻÓRALsKl et al. 2016); Gdańsk-Oliwa: Kwietna Street near the Oliva stream, 1.vi.2016, 1 ♀, leg. RZ ( RZRP); Gibała near Drwęca river, 13.v.2017, 1 J, leg. RZ ( RZRP); Zaskalskie-Bodnarówka, 24.vi.2017, 1 ♀, leg.Ł. Mielczarek ( LMMP); Lisówki ad Stęszew, XT19, 22.v.2018, 1 J, leg.P.Trzciński ( PTPP); Szumny Zdrój ad Górzno, DD19, 27.iv.2019, 1 J, leg. Ł. Mielczarek ( LMMP); Gdynia-Leszczynki near Chylonka stream, 14.v.2019, 1 ♀, leg. RZ ( RZRP);Włoszczowa “klekot”, vi.2019, 2 ♀♀, Malaise trap, leg. Ł. Mielczarek ( LMMP); Kartoszyno, CF16, 30.v.2021, 1 J (GenBank acc. no. OR859651), on Crataegus sp. , leg. RZ ( RZRP), 24.vi.2023, 2 ♀♀ (GenBank acc. no. OR859686), leg. RZ ( RZRP); Zakrzewska Osada, XV61, 20.vi.2022, 1 ♀, leg. & coll. D. Tarnawski. GEORGIA: Dviri, 10.v.2022, 1 J, leg. FM ( FMTB); Dviri, 12.v.2023, 2 JJ 1 ♀, leg. W. Opdekamp ( WORB). SWITZERLAND: Chevroux, 8.–15.v.2020 (Malaise trap), 1 J, leg.Association de la Grande Cariçaie, Nina Perret-Gentil ( MZLS).

Diagnosis. See notes on O. brevicornis within diagnosis of O. atlantica .

Redescription. Male. Body length 5.0–6.4 mm (n = 45; average = 5.9 mm; median = 5.9 mm; neotype = 5.9 mm), overall black to grey ( Fig. 1C View Fig ).

Head. Eyes bare. Antennae ( Fig. 3B View Fig ): scape black; pedicel dark-brown with a few white hairs ventrally that are as long as segment width and with a few shorter black hairs dorsally; postpedicel slightly longer than wide (length to width ratio 1.1–1.3×), not angular at tip, bicoloured: basoventral orange area extending over more than half of postpedicel and rest dark-brown. Sensory pit present on outer side. Arista brown, twice as long as postpedicel. Face ( Fig. 2C View Fig ) in most cases relatively narrow (neotype: upper face width 0.75 mm, lower face width 1 mm), less than half width of head, dull grey with diverging eye margins. Face with wrinkled texture, central part above mouth protrusion often with small, bare surface. Central symmetrical trapezoid area of dense white pilosity just below antennal sockets connected to triangular areas of white pilosity near eyes.All facial hairs white to yellow, covering part of face, largely missing in central area. Facial hairs shorter near mouth. In side view ( Fig. 2D View Fig ), facial profile slightly convex in middle, but varies between individuals between slightly convex to straight. Mouth edge protruding only slightly beyond antennal sockets. Frons not prominent, dull grey, punctuated, covered with white to yellowish hairs. Vertex dull grey, punctuated, with long yellowish hairs slightly bent forward. Ocelli forming almost equilateral triangle. Occiput with band of white pilosity along eyes.

Thorax. Scutum dull grey with metallic, densely punctuated cover. Covered with short erect hairs of equal length. All hairs on scutum white, exceptionally (two specimens from Georgia) with a few black hairs. Four darker longitudinal vittae where this cover is less dense or punctation is absent, best visible from anterodorsal side. Scutellum dull grey and slightly lighter than scutum, covered with white hairs, with rim on its posterior margin and with row of hairs along its hind rim, some slightly bent towards midline, shorter than half length of scutellum. Anterior anepisternum shiny, without hairs, except for very short pilosity on its posterior dorsal part. Posterior anepisternum shiny, covered with long hairs, except for its bare anteroventral part. Hair patches on upper and lower katepisternum widely separated.

Wings transparent and with microtrichia all over; rarely with brown cloud in centre. Venation between dark brown and black, but not pitch black. Vein M 1 with its anterior end almost perpendicular to R 4+5, non-recessive because of location of junction M 1 /R 4+ 5 in wing topology as illustrated on Fig. 7F View Fig ; much varying between individuals as on Figs 7H–J View Fig . In almost all cases, if line is drawn between vein junctions M 1 /M 2 and C/R 2+3, then junction M 1 /R 4+5 is located very close to it, usually a bit to wing tip ( Figs 7F, 7G View Fig ), but never much to wing base as it is the case in O. atlantica sp. nov. ( Figs 7A, 7B View Fig ). Rarely crossvein M 1 more deeply curved ( Fig. 7I View Fig ) or even wavy, according to the observed rule that the narrower wing cell r 4+5 is, the more bent vein M 1 is. Very rarely specimens have extra drop swelling on vein M 1 on one or two wings, or extra denticles on vein M 2 on two wings. Pterostigma light brown with brown anterior margin. Halteres yellow-grey.

Legs. All legs black, usually with shade of grey. Femora slightly swollen, two times maximum tibia width. Tibia covered with short adpressed white hairs, fore and mid femora covered with upstanding longer whitish hairs on the posterior side, longest hairs on posteroventral side (some more than 3/4 th of the width of the femora), hind femora with short hairs (less than half femur width). Ventral side of hind femora and ventral side of all tarsomeres of middle legs covered with short adpressed black bristles. First tarsomeres of middle legs with four longitudinal rows of these bristles, inner rows located close to each other. Claws orange at base, black on top.

Abdomen oval, grey. Dull and lightly pollinose in central areas, shiny grey metallic on sides of all tergites. Shiny parts of tergites covered with white erect or adpressed hairs, mostly directed towards posterior edge of tergites. Dull central area covered with shorter and much sparser hairs. Sternites covered with white hairs: sternite I shiny with sparse hairs, but lateral parts hairless; sternite II with erect and long hairs; sternites III and IV with shorter and inclined hairs, pointing backwards or to centre of sternite.

Terminalia. Phallus, in anterior view ( Fig. 5F View Fig ) laterally flattened. In lateral view ( Figs 5D View Fig , 6A–C View Fig ), with relatively short and broad anterodorsal hook-shaped appendix and prominent flat anteroventral appendix. Some individual variation in shape of flat anteroventral appendix of phallus was observed, as illustrated on Fig. 6 View Fig . Delicate, long and bristle-like process of phallus present. Postgonites not extending beyond tip of phallus, with visible hook-shaped extension anteroventrally ( Fig. 5E View Fig ), and without small denticle on anterior edge found in O. atlantica sp. nov. Posteroventral process of surstyli present ( Fig. 5E View Fig ).

Female. Body length 5.2–6.8 mm (n = 23; average = 6.2 mm; median = 6.3 mm). Resembling male ( Fig. 1D View Fig ), except for the following: eyes dichoptic; frons with 5 lateral furrows, abdomen larger, face in lateral view more concave. Tergite V with shallowly curved posterior margin and without keel or incision in middle. The same variation in wing topology and postpedicel length/width ratio as reported for males is observed in females.

Biology and behaviour. Adults have been found sitting on vegetation near marshland or feeding on flowers with a preference for shrubs and trees (maples Acer spp. , willows Salix spp. , rowans Sorbus spp. ) but also on low flowers. The area where this species occurs in very large numbers near Poznań (locus typicus) is the “Trzcielińskie Bagno” [= Trzcielińskie Marsh], a protected area within the Wielkopolska National Park, where many species of Stratiomyidae were also found (TRZClŇsKl 2007), including Actina chalybea Meigen, 1804 in its north edge of the range, and endangered Stratiomys cenisia Meigen, 1822 , Oxycera nigricornis Olivier, 1822 and O. trilineata (Linnaeus, 1767) . The swamp is a remnant of a former post-glacial lake fed by the waters of the small river Samica Stęszewska, surrounded by a thicket of willows and reeds, and the floodplain of the river has a variable water level, probably creating favourable conditions for the development of the larvae. Over 250 adults of both sexes of O. brevicornis were collected there using Moericke traps in hard-to-reach thickets of flowering willows in the first days of May 2008, and this species was dominant in Syrphidae material. Other findings, documented in the material section, are generally sporadic and associated with the immediate proximity of wetlands.

Distribution. Mainly distributed over Central and Eastern Europe ( Fig. 8 View Fig ) and further east into Russia and Transcaucasia. Most records come from Poland (locus typicus) but the species is also confirmed from eastern Germany, Ukraine (Fig. 21 in PRoKHoRov et al. 2023 shows the typical male terminalia), Russia (Fig. 12 in STACKFLBFRG 1953 shows the typical male terminalia; present in European parts of Russia and western Siberia), the Czech Republic, Hungary, northern Greece, the Republic of Georgia and few, isolated locations in Western Europe: in the Netherlands and Switzerland. Unverified records of O. brevicornis sensu lato that would fit into the known distribution come from Romania (BRǍDFsCU 1991), Turkey ( TÓTH 2013) and Austria (HFlMBURG et al. 2022). Reports of O. brevicornis from Iran (SHAKFRYARl et al. 2012) and Morocco (SAHlB et al. 2020) belong to different species, and the presence of O. brevicornis in these countries needs reassessment. Previous reports of the species from Japan were extensively commented on by ICHlGF (2006) as referring to another species; the slender surstyli and highly asymmetric postgonites in male, as well as the deep incision in the 5 th sternite in female, as described and shown for O. brevicornis in OHlsHl (2011), are clearly not conspecific with O. brevicornis , but close to O. incisa .

Molecular data. We were successful in sequencing 14 specimens of O. brevicornis from Poland, Georgia and Greece. In our NJ tree all DNA barcodes of O. brevicornis cluster with sequences of O. atlantica sp. nov. with high support (BS = 100%), but there is no support for grouping sequences of O. brevicornis alone ( Fig. 10 View Fig ). The COI sequences of O. brevicornis have an uncorrected intraspecific pairwise distance between 0.0 and 0.0035 (or a difference between 0.0 and 0.35%).

Notes on types. LoFw (1843) described the species Chrysogaster brevicornis based on two male specimens from Posen [= Poznań] in Poland (LoFw 1843, PFCK 1988). The original type locality was specified based on two independent statements in the Loew’s publication:

“ Professor Dr. H. Loew in Posen. (...) Ich besitze von dieser Art zwei ganz übereinstimmende hier gefangene Männchen [= I have two identical males of this species caught here.]” Further in the paper, after expanding on the species Chrysogaster nobilis Fall. , Chrysogaster plumbago Loew and Chrysogaster brevicornis nov. sp., and before describing Chrysogaster frontalis nov. sp. and Chrysogaster fumipennis nov. sp., he wrote: “Zu den bereits beschriebenen drei schwarzbeinigen Arten der zweiten Abtheilung von Chrysogaster , die in der Umgegend Posens vorkommen, treten nur noch zwei, dem südlichen Europa und benachbarten Kleinasien angehörige Arten hinzu, (...) [= To the three black-legged species of the second chapter of Chrysogaster already described, that occur in the vicinity of Poznań, two other species are added now, occurring in southern Europe and neighbouring Asia Minor, (...).]”

Only one specimen of O. brevicornis of the collection of Hermann Loew was logged into the ZMHB register book of accessions ( Fig. 9D View Fig ) to the collection from 1858–1884 (under registration no. 12930). This specimen is still present ( Figs 9B, 9C View Fig ). Loew’s first handwritten label under the specimen reads “Frank. a/M” [= Frankfurt am Main], a German city located 630 km west from the type locality. Loew’s second handwritten label reads “ Orthoneura brevicornis m.”, with “m.” meaning “mihi” or “mein”, to explicitly indicate the species was described by himself, as in LoFw (1857). Note that Loew uses “ Orthoneura ” instead of the original “ Chrysogaster ” as the genus name. The name Orthoneura (wrong spelling of Orthonevra Macquart ) became widely accepted by authors only after the publications of WALKFR (1851) and RoNDANl (1857) and became finally also accepted by LoFw (1857) but was not used by LoFw (1843) when describing Chrysogaster brevicornis . In conclusion, both the location and deduced timing of this specimen do not match with the type series. The specimen clearly comes from Loew’s collection, but it is not one of the two original syntypes. Below the two above-mentioned labels is a red label with “Type” typed on it. ZlFGLFR et al. (2020) clarify that specimens of the Loew collection only have two original handwritten labels and that all other labels were added later by museum workers. In this case, the specimen was clearly misinterpreted as a syntype and the type indication is false. Article 72.4 of the Code explicitly allows to take any evidence into account, published or unpublished, when determining which specimens belong to the type series. In this case the documented doubts, especially the mismatch in locality, do not allow designation of the mentioned specimen as the lectotype according to the Article 74.5 of the ICZN (1999). Type material of O. brevicornis was actively searched for by authors. Careful analysis of the collections of ZMHB (main collection and all supplementary drawers) and MZPW did not result in finding Loew’s specimens that could be considered syntypes. Thus, the type series is considered lost or destroyed, and a neotype male specimen from a locality close to the original type locality ( Poland, close vicinity of Posen) is hereby designated for the purpose of stabilisation of the taxonomic nomenclature and for stabilisation of the type locality of a nominal taxon. It must be noted, that the terminalia of Loew’s specimen (from Frankfurt am Main, the one rejected to be a syntype) were extracted and examined by the first author in 2023 and is the same as the terminalia of the neotype being here designated. The shape of the terminalia is also consistent with the drawing of the terminalia of O. brevicornis published by STACKFLBFRG (1953, 1970) and further reproduced by other authors: VAN DFR GooT (1981), BRǍDFsCU (1991), and ICHlGF (2006).

A female was described by LoFw (1857) under the name Orthoneura brevicornis based on an unspecified number of specimens from Meseritz [= Międzyrzecz] ( Poland) with Loew making a note that he also had specimens collected in Florence ( Italy) in his collection. None of these female specimens were donated by H. Loew to the ZMHB, nor are present elsewhere to our knowledge. Female specimen(s) used for the description are considered lost or destroyed. A voucher female specimen caught together with the neotype and matching the original description will be deposited in a well-curated accessible collection (ZMHB) together with the neotype.