Empis septentrionalis Shamshev & Sinclair sp. nov.

(Figs 6, 7)

urn:lsid:zoobank.org:act: 1E05618C-B300-4E56-B381-1A0865B7462E

Type material. HOLOTYPE ♂, labelled: [RUSSIA. Chukotka AO (Wrangel Island)] “[printed in Cyrillic, Russian] Chukotskiy AO [= Autonomous Okrug] o. [= ostrov, island] Wrangel/ Verch. tech. [= verkhnee techenie, upper flow] river Neizvest-/ naya 71°13′N 179°19′W / O.A. Khruleva 8.7.2015 / 2B”; “ Holotypus / Empis / septentrionalis / Shamshev, Sinclair sp. nov. [red label]” (INS_DIP_0000612, ZIN).

PARATYPES: RUSSIA. Chukotka AO (Wrangel Island): same locality as holotype, BT 7 A, 5.vii.2006, OAK (1 ♂, 1 ♀, CNC); same locality as holotype, BT 7 A, 10.vii.2015, OAK (1 ♂, ZIN); same locality as holotype, BT 7 B, 10.vii.2015, OAK (7 ♀, ZIN); same locality as holotype, BT C1, YPT, 4–15.vii.2015, OAK (1 ♂, ZIN); same locality as holotype, BT 1 A, 6–13.vii.2015, OAK (1 ♂, ZIN); Mineev Mtns, 71°00′N 179°31′W, BT 14 B, 14.vii.2006, OAK (1 ♂, ZIN); environs of Pervaya Mtn (71°10′N 179°27′W), BT A, hand collecting, 21.vii.2019, OAK (1 ♀, ZIN).

Diagnosis. Mid-sized (wing length 5–5.5 mm) robust flies with brownish pruinescent thorax and abdomen and almost entirely dark brown legs. Male dichoptic; occiput slightly convex with numerous long, flattened setae laterally behind eye; thorax black setose, scutum with 4 vittae; legs with only hind tarsomeres 1–3 brownish yellow, hind femur somewhat thickened, hind tibia very slender; apical part of phallus uniformly tubular. Female with brownish infuscate wing; abdomen densely brown pruinescent on tergites dorsally.

Description. Body length 5.2–5.9 (holotype 5.3); wing length 5–5.5 (holotype 5.1) mm. Male (Fig. 6). Head with dense brownish pruinescence on face, frons, postgena, ocellar triangle and occiput; entirely dark setose; clypeus shiny. Dichoptic; ommatidia equally small. Frons broad, below ocellar tubercle broader than distance between outer margins of posterior ocelli; with marginal setulae. Face broad, bare. Ocellar triangle with several long fine setae. Occiput slightly convex laterally behind eyes; with numerous long mostly flattened setae on convex part, fine long postoculars and similar setae on upper part; postgena with numerous hair-like setae. Antenna black; scape short, slightly longer than pedicel, both with short setulae; postpedicel conical, with slightly concave ventral margin on apical part, nearly 2X longer than wide; stylus rather long, slightly longer (1.3–1.4X) than postpedicel basal width. Palpus dark (sometimes yellowish, 2 ♂); with several black moderately long fine setae. Proboscis with labrum mostly brownish, reddish-brown apically, about 1.5X longer than head height.

Thorax dark in ground-colour, brownish pruinescent; with black setation; scutum with 4 indistinct narrow blackish brown vittae (dorsal view). Proepisternum with tuft of numerous mostly long and slightly flattened setae (with some additional shorter fine setae) on lower section; upper proepisternum in front of spiracle bare. Prosternum bare. Antepronotum with numerous short fine setae. Postpronotal lobe covered with very numerous subequally long setae. Mesonotum with hardly differentiated setae; several long fine intra-alars just behind postpronotal lobe; supra-alar space with several similar setae before suture and less numerous setae behind suture, notopleuron with numerous long setae, some posterior setae somewhat stronger; 2 long strong pal (sometimes with additional setulae), 8–10 sctl; acr long, fine, arranged in 2 irregular rows, lacking on prescutellar depression; presutural dc similar to acr, multiserial, postsutural dc less numerous, 3–4 prescutellars longest. Laterotergite with numerous long fine setae. Anterior and posterior spiracles brown.

Legs robust, almost entirely dark brown, only hind tarsomeres 1–3 (sometimes 1–4) brownish yellow; black setose; hind femur somewhat thickened, mid tibia slightly arcuate, hind tibia very slender. Coxae and trochanters with simple setae. Femora whitish pilose ventrally (less distinct on fore femur); fore femur with long fine setae anteroventrally, posteroventrally and posteriorly; mid femur with numerous spine-like setae of different lengths ventrally, posteroventrals longer, some moderately long anterodorsal setae; hind femur with numerous long flattened setae anterodorsally (except short subapical part), only pale pilose along lower half anteriorly (except several setae near extreme base), rows of long, strong anteroventral and posteroventral setae, short spine-like setae ventrally. Tibiae devoid of strong setae (including setae of subapical circlet); fore tibia with some short, fine setae posteriorly; mid tibia with erect setulae ventrally; hind tibia pale pilose ventrally, no seta in posteroapical comb. Hind tarsomeres with somewhat finer and sparser setulae.

Wing membrane very faintly infuscate, with brownish veins; CuA+CuP (anal vein) complete; cell dm with elongate apex. Pterostigma brownish yellow. Basal costal seta absent. Anal lobe well-developed; axillary incision acute. Squama dirty yellow, brownish fringed. Halter with yellow knob and brownish stem.

Abdomen dark, brownish pruinescent (somewhat denser on tergites laterally and sternites); covered with numerous long, dark fine setae (shorter on tergites dorsally). Segment 8 with tergite and sternite separated; tergite 8 mostly membranous, represented by two weakly sclerotized, narrow lateral sclerites each bearing 0–1 setae; sternite simple, nearly as long as sternite 7 (lateral view), with numerous long setae along posterior margin.

Terminalia (Fig. 7). Cerci broadly yellowish to brownish yellow, narrowly brownish along upper margin and anteriorly; separated from each other and from epandrium; cercus elongate oval (lateral view), with rounded apex, slightly extended beyond apex of epandrium, covered with short dark setae. Epandrium almost entirely brownish, narrowly yellowish brown along lower margin; subtriangular (lateral view), with dark setae more numerous and longer along lower margin. Hypandrium mostly membranous, rim-like sclerotized along upper margin, bare. Phallus almost uniformly broad, slightly narrowed near middle, S-like bend, sulcate on subapical portion, with pair of small projections near apex beneath. Ejaculatory apodeme extended far beyond basal curvature of phallus, with subequal lateral and vertical wings and somewhat narrower lower wing.

Female. Occiput not convex laterally, without flattened setae, ocellar tubercle with short setae; palpus somewhat paler, yellowish brown, sometimes with yellowish apex or yellowish. Thorax and legs with shorter setation; hind femur not thickened, hind tibia simple; hind femur without flattened setae and pale pilosity along lower half anteriorly, entirely covered with setulae anteriorly; fore tibia without fine setae posteriorly, mid and hind tibiae with usual setulae ventrally; hind tarsus brownish, with similar setation to fore and mid tarsi. Wing darker, brownish infuscate, pterostigma brownish. Abdomen densely brownish grey pruinescent on tergites laterally and sternites, brown pruinescent on tergites dorsally (except noted); tergites 7–8 extensively shiny (except posterior margin), also sternites 7–8 usually subshiny to shiny anteriorly; only tergites 1–2 with some short setae laterally, otherwise tergites covered with scattered setulae; cercus black, with dark setulae.

Distribution. Palaearctic, known only from Wrangel Island.

Etymology. The specific name is from the Latin septentrionalis (north, northern), in reference to the distribution of this species.

Remarks. Empis septentrionalis sp. nov. belongs to the group of species with uncertain subgeneric position within Empis (Shamshev 2001a). The group is known almost exclusively from the Asiatic part of the Palaearctic (except one species) and is especially diverse in Middle Asia. In addition, three undescribed species are known from Yukon in Canada. The new species is very similar to E. jacutiensis Shamshev occurring in Yakutia and Chukotka of Russia (Shamshev 2016). Empis septentrionalis sp. nov. can be readily distinguished from E. jacutiensis by uniformly tubular apical part of the phallus (cup-shaped apically in E. jacutiensis) (Fig. 8) and dark brown hind tibia (yellowish to brownish yellow in E. jacutiensis). In addition, the male of E. septentrionalis sp. nov. usually has dark palpi (yellowish in E. jacutiensis), but this character appears to be somewhat variable. The female of the new species differs from the female of E. jacutiensis by dark brown hind tibia and tarsus and by brown pruinescence on abdominal tergites dorsally (abdomen uniformly light grey pruinescent in the female of E. jacutiensis).

Habitat. Almost all specimens were collected in the warmer central region of Wrangel Island, mainly in dry habitats.

Genus Rhamphomyia Meigen

Key to Rhamphomyia of the Russia Arctic islands and Svalbard Archipelago

The key provided below follows generally the key compiled by Sinclair et al. (2019) to species of the Canadian Arctic Archipelago, Greenland and Iceland and we recommend that this publication be consulted for additional species that may be found in the Eurasian Arctic islands. Identification of Asiatic species of Rhamphomyia remains problematic due to very high diversity and weak species group definitions and classification. Barták & Kubík (2009) and Saigusa (2012) should be consulted for keys to species groups of Rhamphomyia and references to available revisions.

1 Male (unknown in R. nordqvistii, R. sp. 1).................................................................. 2

- Female (unknown in R. armipes, R. submacrura sp. nov., R. subfilicauda sp. nov.)................................ 28

2 Axillary lobe of wing very little developed, axillary excision extremely obtuse (Fig. 27); eyes dichoptic, frons shiny [Additional characters: occiput shiny on upper part, greyish pruinescent on middle.]....... R. (Lundstroemiella) hybotina (Zetterstedt)

- Axillary lobe of wing well developed, axillary excision at most 90°; eyes holoptic or frons pruinescent................. 3

3 Prosternum clothed with long white hair-like setae (Sinclair et al. 2019: fig. 3D); sternite 6 with cluster of strong golden setae; sternite 7 with pair of horn-like projections ventrally (Fig. 9)...................... R. (Ctenempis) albopilosa Coquillett

- Prosternum bare; sternite 6 without cluster of golden setae; sternite 7 unmodified or modified different than above........ 4

4 Cerci strongly prolonged anteriorly, extended at least to tergite 6 (Figs 10–13) (subgenus Dasyrhamphomyia)............ 5

- Cerci not extended anteriorly beyond tergite 8............................................................. 10

5 Halter yellow........................................................................................ 6

- Halter brown......................................................................................... 8

6 Cerci very long, extended nearly to middle of abdominal tergite 3 (Fig. 13); hind femur without setae on about apical 2/3 ventrally, only densely pale pilose.................................................... R. (D.) nigrita Zetterstedt

- Cerci much shorter, extended at most to abdominal tergite 5; hind femur with numerous short setae over entire length ventrally............................................................................................ 7

7 Hind tibia thickened; wing whitish (Fig. 14); epandrium brownish to reddish brown (Fig. 15)............................................................................... R. (D.) gorodkovi Shamshev, Sinclair & Saigusa sp. nov.

- Hind tibia slender; wing faintly infuscate (Fig. 20); epandrium yellowish (Fig. 21)......................................................................................... R. (D.) mallochi Shamshev, Sinclair & Saigusa sp. nov.

8 Frons very broad, in middle nearly as broad as distance between outer margins of posterior ocelli, almost parallel-sided; hind femur without setae on about apical 3/4 ventrally, only densely pale pilose (Fig. 11).......... R. (D.) erinacioides Malloch

- Frons very narrow on middle part, at most slightly broader than anterior ocellus or eyes touching; hind femur with numerous short setae over entire length ventrally..................................................................... 9

9 Mesonotal setae strong, bristle-like (Fig. 10); eyes separated by very narrow frons........... R. (D.) brusewitzii Holmgren

- Mesonotal setae almost uniformly fine, hair-like (Fig. 12); eyes usually touching............ R. (D.) hovgaardii Holmgren

10 Subepandrial sclerite prolonged into pair of medioposterior lobes beneath cerci (Fig. 31) (Sinclair et al. 2019: figs 25A, C) 11

- Cercus without pair of subepandrial lobes beneath.......................................................... 14

11 Hind femur with triangular posteroventral projection beyond mid-length; hind tibia with triangular posteroventral projection on basal third (Fig. 30)..................................................... R. (Pararhamphomyia) armipes Sack

- Hind femur and tibia without projections.................................................................. 12

12 Abdomen covered with black setae; fore coxa with thin setae anteriorly; hind tibia with 4–5 anterodorsal and several posterodorsal prominent setae, hind basitarsus with only short setae dorsally (Sinclair et al. 2019: fig. 34B); wing membrane whitish (Fig. 34).................................................................................... R. (P.) hoeli Frey

- Abdomen covered with pale setae; fore coxa with strong blunt-tipped setae anteriorly; hind tibia and basitarsus covered with long, dense, sometimes woolly, setae dorsally; wing membrane hyaline to slightly infuscate......................... 13

13 Phallus forming broad loops (Sinclair et al. 2019: fig. 36C); hind tibia clavate (Fig. 35)........ R. (P.) kjellmanii Holmgren

- Phallus not forming loops, only gently curved (Sinclair et al. 2019: fig. 25A, C); hind tibia more slender, evenly thickened towards apex (Fig. 33)........................................................... R. (P.) frigida Sinclair et al.

14 Abdominal segments 7 and 8 asymmetrical, strongly sclerotized (somewhat darker) than preceded segments, segment 7 enlarged; terminalia slightly rotated to right (Fig. 52). [Additional characters: hind tibia strongly curved inward near basal third (posterior view) (Fig. 49); phallus mostly hidden within epandrium, hook-like curved to left at apex (Fig. 53)].......................................................................... R. (P.) wrangeli Shamshev & Sinclair sp. nov.

- Abdominal segments symmetrical, normally sclerotized, segment 7 unmodified; terminalia unrotated................. 15

15 Upper proepisternum in front of spiracle with setae; basal costal seta usually present (often 2 setae in R. shewelli or sometimes indistinct in R. sublongiseta sp. nov.).................................................................... 16

- Upper proepisternum in front of spiracle without setae; basal costal seta usually absent (except R. taimyrensis).......... 21

16 Wing axillary incision very acute, at most 45°. Cercus without notch dorsally, at most tuberculate; phallus short, gently curved............................................................................................. 17

- Wing axillary incision more than 45°, usually close to 90°. Cercus with notch dorsally (Fig. 60); phallus long, sinuate.... 19

17 Halter yellow; abdomen pruinose dorsally and narrowly shiny laterally (Fig. 26) R. (Eorhamphomyia) shewelli Sinclair et al.

- Halter brown; abdomen entirely pruinose, although sometimes thin............................................. 18

18 Face with several fine setae; abdominal tergites faintly pruinescent, somewhat lustrous (Fig. 53)........................................................................................... R. (Rhamphomyia) morio (Zetterstedt)

- Face bare; abdominal tergites densely light grey pruinescent (Fig. 52)............... R. (R.) albosegmentata (Zetterstedt)

19 Hind basitarsus strongly expanded (Fig. 62); cercus with notch forming small projection slightly beyond middle dorsally (Fig. 64)..................................................... R. sublongiseta Shamshev, Sinclair & Saigusa sp. nov.

- Hind basitarsus slender; cercus with finger-like projection near base dorsally..................................... 20

20 Wing somewhat whitish (Fig. 57); mid tibia and basitarsus with numerous very long setae dorsally (2–2.5X longer than tibia width) (Fig. 58)........................................................................ R. kaninensis Frey

- Wing faintly infuscate; mid tibia and basitarsus with short setae (at most as long as tibia width) (Fig. 56)................................................................................................... R. hirtula Zetterstedt

21 Phallus with small, narrow U-shaped loop on subapical part (Fig. 67); cercus about as long as epandrium, with straight dorsal margin. [Additional character: hind tibia curved inward closer to base (posterior view)] (Fig. 66)....... R. taimyrensis Frey

- Phallus gradually curved; cercus distinctly shorter than epandrium, usually more or less concave dorsally.............. 22

22 Anal vein reaching wing margin, entirely sclerotized (Fig. 43). [Additional characters: halter brownish; mesonotum and abdomen with only black setae; scutum entirely pruinescent; hind tibia straight; hind basitarsus slightly expanded, cylindrical, covered with short setae; epandrium nearly 4X longer than cercus (Figs 43, 44).]............................................................................................... R. (P.) submacrura Shamshev & Sinclair sp. nov.

- Anal vein disappearing before wing margin, at most evanescent (or fold-like) apically.............................. 23

23 Halter with yellow knob; mesonotum (except stronger setae) and abdomen covered with pale thin setae; hind tibia curved (posterior view); hind basitarsus spindle-shaped, convex dorsally, longer than half length of tibia (Fig. 39). [Additional characters: scutum with 4 shiny vittae; hind basitarsus covered with numerous long setae dorsally; epandrium nearly 1.5X longer than cercus.]...................................................... R. (P.) septentrionalis Sinclair et al.

- Halter brown; mesonotum and abdomen with black setation; hind tibia straight; hind basitarsus cylindrical or clavate, shorter than half length of tibia................................................................................ 24

24 Anepisternum shiny.................................................................................. 25

- Anepisternum pruinose................................................................................ 26

25 Scutum faintly greyish pruinescent, with two shiny median vittae (sometimes additional less distinct vitta present on supra-alar space); hind tibia with uniformly long setae dorsally; hind basitarsus clavate (Fig. 48); cercus spatulate beyond median cavity (lateral view).................................................................... R. (P.) ursinella Melander

- Scutum shiny, at most very faintly pruinescent in front of scutellum; hind tibia with long setae only near apical half dorsally; hind basitarsus more or less cylindrical; cercus finger-like beyond median cavity (lateral view) (Figs 36, 37).............................................................................................. R. (P.) longestylata Frey

26 Acrostichals and presutural dorsocentrals very short, fine, the latter 1–2-serial; mesonotum and abdomen almost shiny. [Additional characters: fore and mid tibiae and tarsi with short setae, hind tibia with moderately long setae dorsally; hind basitarsus clavate, tarsomere 2 thickened, both with long, dense fine setae dorsally (Fig. 45).]................................................................................. R. (P.) subtenuiterfilata Shamshev & Sinclair sp. nov.

- Acrostichals and presutural dorsocentrals long, strong, the latter at least 3–4-serial; mesonotum and abdomen densely pruinose........................................................................................... 27

27 Smaller, wing 4.3 mm; hind tibia with long dense fine setae dorsally; wing somewhat whitish (Fig. 41)............................................................................. R. (P.) subfilicauda Shamshev & Sinclair sp. nov.

- Larger, wing 4.8–5.4 mm; hind tibia with bristle-like setae dorsally; wing faintly infuscate (Fig. 32)....................................................................................... R. (P.) filicauda Henriksen & Lundbeck

28 Axillary lobe of wing almost absent, axillary excision extremely obtuse; frons shiny. [Additional characters: occiput shiny on upper part, greyish pruinescent on middle.].......................................... R. (L.) hybotina (Zetterstedt)

- Axillary lobe of wing well-developed, axillary excision at most slightly more than 90°; frons pruinescent.............. 29

29 Prosternum clothed with long white hair-like setae..................................... R. (C.) albopilosa Coquillett

- Prosternum bare..................................................................................... 30

30 Anal vein (CuA+CuP) reaching wing margin, usually entirely sclerotized (sometimes slightly weaker about middle) (variable in R. erinacioides)................................................................................... 31

- Anal vein (CuA+CuP) not reaching wing margin or evanescent (fold-like) apically (variable in R. hoeli)............... 44

31 Basal costal seta present (in R. shewelli often 2 setae)........................................................ 32

- Basal costal seta absent............................................................................... 38

32 Upper proepisternum in front of spiracle without setae; halter with yellow knob; mid and hind femora with anterodorsal and posteroventral rows of pennate setae (except extreme base)..................................... R. taimyrensis Frey

- Upper proepisternum in front of spiracle with setae; halter brown; legs with simple setae........................... 33

33 Abdomen pruinose dorsally and narrowly shiny laterally............................... R. (E.) shewelli Sinclair et al.

- Abdominal tergites uniformly pruinose................................................................... 34

34 Face with several fine setae........................................................ R. (R.) morio (Zetterstedt)

- Face bare........................................................................................... 35

35 Acr and dc strong bristle-like; scape and pedicel slightly paler than postpedicel; thorax lighter, grey pruinose................................................................................... R. (R.) albosegmentata (Zetterstedt)

- Acr and dc fine; antenna uniformly dark; thorax darker, brownish pruinose....................................... 36

36 Anal lobe of wing darker than wing tip; hind tibia and basitarsus slightly thickened......................................................................................... R. sublongiseta Shamshev, Sinclair & Saigusa sp. nov.

- Wing membrane uniformly infuscate; at least hind basitarsus slender........................................... 37

37 Hind tibia slender, entirely short setose; abdomen lighter, densely greyish pruinose.................. R. kaninensis Frey

- Hind tibia somewhat dilated, almost bare anteriorly; abdomen darker, densely brownish pruinose..... R. hirtula Zetterstedt

38 Legs with pennate setae, at least mid and hind femora (somewhat variable in R. gorodkovi sp. nov.)................... 39

- Legs covered with simple setae, at most sometimes some setae on mid and hind femora slightly flattened.............. 41

39 Halter brown; hind tibia with pennate setae dorsally................................... R. (D.) hovgaardii Holmgren

- Halter with yellow knob; hind tibia with simple setae........................................................ 40

40 Hind tibia evenly broadened toward apex [see also couplet 42]... R. (D.) gorodkovi Shamshev, Sinclair & Saigusa sp. nov.

- Hind tibia slender........................................ R. (D.) mallochi Shamshev, Sinclair & Saigusa sp. nov.

41 Halter at least with yellow knob......................................................................... 42

- Halter brown........................................................................................ 43

42 Legs uniformly dark, hind tibia slender; wing membrane brownish.......................... R. (D.) nigrita Zetterstedt

- Legs yellowish to brownish yellow, hind tibia evenly broadened toward apex; wing membrane faintly infuscate................................................................ R. (D.) gorodkovi Shamshev, Sinclair & Saigusa sp. nov.

43 Smaller, wing length about 5 mm; mesonotum dark brown (dorsal view); presutural dorsocentrals multiserial.......................................................................................... R. (D.) erinacioides Malloch

- Larger, wing length 6–6.5 mm; mesonotum greyish (dorsal view); presutural dorsocentrals biserial................................................................................................. R. (D.) brusewitzii Holmgren

44 Abdomen covered with pale to yellowish setae............................................................. 45

- Abdomen covered with black setae...................................................................... 47

45 Mesoscutum with four shiny vittae; wing deeply brown, broadly rounded (Fig. 40); fore coxa with fine setae anteriorly........................................................................... R. (P.) septentrionalis Sinclair et al.

- Mesoscutum uniformly pruinose; wing faintly brownish infuscate, of normal shape; fore coxa with strong blunt-tipped setae anteriorly........................................................................................... 46

46 Hind tibia with dense dorsal and ventral fine setae at least as long as tibia width.............. R. (P.) kjellmanii Holmgren

- Hind tibia with ventral setae at most half-length of tibia width............................ R. (P.) frigida Sinclair et al.

47 Halter with yellowish knob................................................................ R. (P.) hoeli Frey

- Halter brownish..................................................................................... 48

48 Scutum almost shiny or with shiny vittae.................................................................. 49

- Scutum densely, more or less uniformly pruinescent......................................................... 51

49 Anepisternum pruinose..................................... R. (P.) subtenuiterfilata Shamshev & Sinclair sp. nov.

- Anepisternum shiny.................................................................................. 50

50 Scutum faintly greyish pruinose, with two shiny median vittae (sometimes additional less distinct vitta present on supra-alar space)......................................................................... R. (P.) ursinella Melander

- Scutum shiny, at most very faintly pruinose in front of scutellum............................. R. (P.) longestylata Frey

51 Anal lobe of wing darker than wing tip................................... R. (P.) filicauda Henriksen & Lundbeck

- Wing uniformly infuscate.............................................................................. 52

52 Scutellum with 4 setae [see also couplet 47]................................................... R. (P.) hoeli Frey

- Scutellum with 6–8 setae.............................................................................. 53

53 Upper proepisternum in front of spiracle with 3 setae; no distinct basal costal seta (Note: this species is known after the lectotype female only and the range of variability of these characters is unclear.)............ R. (P.) nordqvistii Holmgren

- Upper proepisternum in front of spiracle without setae; usually 1 more or less distinct short basal costal seta............ 54

54 Mesonotal setae stronger, bristle-like; presutural dorsocentrals biserial, separated from supra-alars by broad bare space................................................................... R. (P.) wrangeli Shamshev & Sinclair sp. nov.

- Mesonotal setae fine; presutural dorsocentrals 3–4-serial, barely separated from supra-alars (Fig. 68)........................................................................................................ Rhamphomyia sp. 1