Sphyracephala babadjanidesi Zaitzev, 1919

Feijen, Hans R., Feijen, Frida A. A. & Feijen, Cobi, 2025, A revision of the four Afrotropical and Palaearctic Sphyracephala Say (Diptera, Diopsidae) with an illustrated overview of the other five Sphyracephala, ZooKeys 1241, pp. 1-81 : 1-81

Sphyracephala babadjanidesi Zaitzev, 1919

Figs 5 , 6, 7 , 8–11 , 12, 13 , 14–17 , 18–23 , 24–28 , 29–31 , 32 , 105 , 106 , 107 , 108 , 109 , 110–112 , Tables 1 , 2 , 3 , 4

Sphyracephala babadjanidesi Zaitzev, 1919: 3 (in Russian), 5 (in English), fig. 1. Hennig 1941 a: 60, 1941 b: 6, fig. 6 (repetition of Zaitzev’s description); Steyskal 1972: 13; Feijen 1989: 67; Papp et al. 1997: 137; Hilger 2000: 340; Simova-Tošić and Stojanović 2000: 149; Nartshuk 2003: 179, pl. 44, fig. 13, 2017: 128.

Sphyracephala europaea Papp & Földvári, 1997: 138, figs 1–13. Simova-Tošić and Stojanović 2000: 149, figs 1–5, table 1 (as S. europea [sic]); Hilger 2000: 338, figs 7.1, 7.2; Földvári and Meier 2002: 71; Carr et al. 2006 a: 5, figs 1 h, 2; Oosterbroek 2006: 130, fig. 496; Carr 2008: 114, fig. 1, table 1; Kotrba 2014: 98, fig. on p. 99; Nartshuk 2017: 129; KMNP 2018: 10 th p. (unpaginated); Kutsarov and Hubenov 2019: 145, figs 1–3); Jackson 2019: 61, suppl. fig. 1, 2; Turista Magazin 2023: unpag., fig. 4 / 6. Syn. nov.

Link.

https://www.flickr.com/search/?text=sphyracephala%20europaea.

Type series.

Sphyracephala babadjanidesi . Azerbaijan: 6 syntypes (♂ and ♀), Elizavetpol [later Ganja, then Kirovabad, now Ganja], vi.1916, vi.1917 ; type series lost according to Nartshuk (2017), who designated a neotype, ♂, Azerbaijan, окр. Ганжи, р. Качкарка [okr. Ganzhi, r. Kachkarka , 40°40'12"N, 46°16'33"E], 2.vii.1933, Lukyanovich ( ZIN) . Type location and the nearby neotype location are well into the Asian part of Azerbaijan.

Sphyracephala europaea . Hungary: holotype, ♂, Szeged, Maros-torok, magaspart [~ 46°14'24"N, 20°14'14"E ~ 100 m], 26. iv. 1997. Paratypes, 10 ♂, 7 ♀, same locality and date; 1 ♀, same locality, 16. x. 1996 (all in HNHM).

Material examined.

Azerbaijan: 1 ♀, 1 ♂, Болчалы ЮЗ Гянджи, Азербайджан, Лукъянович 17.vii.1933, Sphyracephala babadjanides [sic], det. Nartshuk ( RMNH) [Bolchaly (= Balchili ), sw Ganja, Lukyanovich, 40°40'12"N, 46°16'33"E, 17.vii.1933, ~ 500 m] . Azerbaijan is mainly located in West Asia, but a small part (5 ½ districts) in the North is part of Europe as the Caucasus form the division between Eastern Europe and Western Asia. From 1920 to 1991, Azerbaijan was part of the Soviet Union ( USSR). Fyodor K. Lukyanovich (1904–1942) was a Russian entomologist. Hungary: paratypes of S. europaea , 2 ♀, 2 ♂, Szeged, Maros-Torok , magaspart, 26.iv.1997, Paulovics and Földvári ( HNHM) ; Georgia: 1?, Kakheti, Signagi, Vakiri , 41°38'43.368"N, 45°55'27.3"E [390 m], 17.vii.2024, S. Kiladze (photographs, see https://www.inaturalist.org/observations/229960704) .

Diagnosis.

Sphyracephala babadjanidesi can be recognised by the following set of characters: head mainly blackish brown, face and anterior edge of frons brown; thorax and abdomen blackish; clothed in sparse, small white setulae; eye stalk very stout (~ 0.94–1.02 × the widest sagittal eye diameter); very small eye span (~ 1.7–2.2 mm) in ♀ and ♂ (respectively ~ 52 % and ~ 60 % of body length); very low rate of dimorphism D = 0.39; rectangular basiliform prosternum; apical seta / scutellar spine ratio: 5.4–5.5; scutellar spine / scutellum ratio: 0.43 in ♀ and 0.41 in ♂; very small, scutellar spines whitish but darker basally ~ 0.13 mm; wing almost transparent with brown central and apical spots; fore femur brown with apical third blackish brown, inner side centrally with dark brown diagonal transverse band, strongly incrassate (l / w ratio: 2.7–2.8), two rows of pale slender spinous setae, inner row with ~ 4.5 setae and outer row with ~ 4.0 hardly spinous setae; tergite 1 with distinct subcircular groove; intersternite 1-2 a solid, straight, rod-like sclerite, laterally linked to sternite 2; ♀ tergite 7 consisting of two anteriorly located, triangular sclerites; ♀ sternite 7 with anteriorly two subtriangular plates, posteriorly connected to two subrectangular plates; ♀ sternite 8 two large elongate sclerites; ♀ cerci rather elongate, l / w ratio: ~ 3.2, sharply tapering apically, remarkably curled upward; small sclerotised ring present; surstyli articulate, medially directed, subrectangular with slightly concave apical side, without microtrichia, clothed in setulae, diagonal ridge on basal half of inner side. Sphyracephala babadjanidesi belongs to the S. brevicornis species group and comes closest to S. munroi .

Redescription.

The following redescription considers the original descriptions by Zaitzev (1919) and Papp et al. (1997), and especially also the description and illustrations by Simova-Tošić and Stojanović (2000). Philipp Adamovich Zaitzev (1877–1957) was the founder of research on the insect fauna of the Caucasus.

Measurements. Zaitzev (1919) studied six specimens and gave as length of body 3.7–4.2 mm, eye span 2.2–2.5 mm and wingspan 7–8 mm. Papp et al. (1997) gave as length of body 3.48 mm (holotype ♂), 3.10–3.50 mm (paratype ♂♂), 3.38–4.05 mm (paratype ♀♀); as wing length 3.13 mm (holotype), 2.75–3.20 mm (paratype ♂♂), 3.05–3.65 mm (paratype ♀♀); and as eye span 2.20 mm holotype), 1.90–2.20 mm (paratype ♂♂) and 1.70–2.15 mm (paratype ♀♀). The best series of measurements were given in Simova-Tošić and Stojanović (2000: table 1). However, measurements were given in μm and not, as stated, in mm, so values given must be divided by 1000 to get mm). Their most important measurements are body length ♀ 3.92 mm ± SE 0.05 (range 3.33–4.38, n = 35), ♂ 3.69 mm ± 0.04 (range 3.15–4.13, n = 38), eye span ♀ 2.05 mm ± 0.03 (range 1.68–2.28, n = 35), ♂ 2.21 mm ± 0.04 (range 1.75–2.83, n = 38). We measured the two flies from Azerbaijan and four paratypes from Hungary. For comparison, relevant measurements are summarised in Table 1 . All data points for body length and eye span are plotted in Fig. 5 . The actual Azerbaijan measurements for the ♀ fit well with the measurements of Zaitzev, but the ♂ is clearly at the lower end of the size range (Fig. 5 ). In general, measurements for the three countries are well in agreement with each other (Table 1 ).

Head. Face and anterior edge of frons brown (Figs 6 , 7 , showing head of flies from Azerbaijan and Hungary); arcuate groove blackish brown; remainder of frons and stalks blackish brown, occiput brown but blackish medially (Figs 8 , 9 ); head uniformly pruinose (Figs 6 – 9 ), clothed in sparse white setulae; frons flat; face flat, no facial teeth, lateroventral corners rounded, facial sulcus absent, but ventral facial edges slightly turned upward medially; eye stalk very stout, ~ 0.94–1.02 × the widest sagittal eye diameter; eye span very small in both female (52.3 % ± SE 0.2 % of body length, n = 38) and male (59.7 % ± SE 0.5 % of body length, n = 41); a dimorphic species, rate of dimorphism very low D = 0.39 (Fig. 5 , 105 , 106 , Table 2 ) [ Carr et al. 2006 a also indicated this species as mildly dimorphic]; inner vertical seta long, ~ 0.50 mm, 1.3 × diameter of eye stalk; outer vertical seta long, ~ 0.35 mm, 0.9 × diameter of eye stalk (Figs 6 , 7 ). For additional figures of the head refer to Zaitzev (1919: fig. 1 a), Papp et al. (1997: figs 1–3) and Simova-Tošić and Stojanović (2000: figs 1, 2 b, 2 c (antennae), 4 a, 4 b, 4 c).

Thorax. Collar and scutum uniformly black, pruinose (Figs 6 , 7 , 31 ), scutellum slightly more blackish brown, pruinose; scutellar spines whitish but darker basally, densely pruinose (Figs 8 , 9 ); scutum and scutellum with sparse setulae; pleura dark black, largely pollinose, only katepisternum and ventral section of anepimeron glossy (Fig. 9 ); posterior notopleural seta long, infra-alar seta very long (Fig. 8 ), infra-alar seta more than twice longer than posterior notopleural seta ( Simova-Tošić and Stojanović (2000) give for the notopleural seta a length of ~ 0.29 mm and for the infra-alar seta a length of ~ 0.67 mm); supra-alar carina just visible; basiliform prosternum large, rectangular, with deep medial groove, prosternum laterally close to propleuron but distinct; scutal length / scutal width ratio: ~ 0.9; scutellum trapezoid; scutellar spines very small, straight, slightly turned upward, diverging at angle of ~ 50 °; scutellar spine / scutellum ratio: 0.43 in ♀ and 0.41 in ♂ (Table 3 ); scutellar spine / length of body ratio: 0.055 in ♀ and 0.051 in ♂; apical seta / scutellar spine ratio: 5.4 in ♀ and 5.5 in ♂; scutellar length / scutellar width (at base) ratio: 0.57 in ♀ and in ♂. For additional figures of the thorax can be referred to Simova-Tošić and Stojanović (2000: figs 3 e, 3 f, scutellum and postscutellum).

Wing. Almost transparent with brown central and apical spots (Figs 12 , 13 showing wing of flies from Azerbaijan and Hungary); apex with rounded spot in cells r 2 + 3 and r 4 + 5 just extending in cells r 1 and m 1; central irregular spot running from posterior end of crossvein dm-m to almost vein R 2 + 3, section in cell r 2 + 3 rounded, section in cell r 4 + 5 running from crossvein r-m to crossvein dm-m, section in cell bm + dm in anterodistal corner and along crossvein dm-m; vein CuA + CuP from vein CuP onward extending under angle of 45 ° to just past halfway wing margin in straight line; vein M 4 continuing distal of crossvein dm-m to almost three-quarters the wing margin; cell cua slightly broadening distally, apically rounded (Figs 12 , 13 ); crossvein h distinct; glabrous area only includes tiny basal spot in cell br.

Legs. Fore coxa and trochanter brown, especially anteriorly pruinose, clothed in some white setulae; fore femur (Figs 14 , 15 showing femora of flies from Azerbaijan and Hungary) brown, apical third blackish brown on inner and outer side, inner side centrally with dark brown diagonal transverse band, thinly pruinose, sparsely clothed in small setulae; fore tibia dark brown, thinly pruinose; basitarsus dark brown, other tarsomeres brown (Figs 16 , 17 ), thinly pruinose and with rows of blackish setulae; mid and hind legs brown, femora with darker brown apices, hind tibia with darker brown apex; fore femur strongly incrassate (Table 2 ), l / w ratio in Azerbaijan ♀ 2.69 and in Azerbaijan ♂ 2.82, l / w ratio in two S. europaea paratype ♀ from Hungary 2.77 ± 0.03 (range 2.73–2.80) and in two S. europaea paratype ♂ 2.82 ± 0.03 (range 2.79–2.85) [ Papp et al. (1997) stated that the fore femur in S. europaea is definitely thicker than in S. babadjanidesi and a major differential character. Basing themselves on Zaitzev’s (1919) drawing, they gave a ratio width / length for S. babadjanidesi of 27 / 75 (i. e., l / w ratio: 2.78). According to Papp et al., the ratio length / width in the holotype of S. europaea came to 2.41, while in the paratype females it was 2.44. However, the data at our disposal clearly show no difference in the ratio l / w between S. europaea and S. babadjanidesi .]; fore femur with two rows of pale spinous setae on distal two-thirds (Figs 14 , 15 ), especially the setae on the outer site are very slender and hardly qualify as “ spinous ”, in total 8.5 ± SE 0.2 setae (n = 10, range 8–9, ♀ and ♂ combined), inner row with 4.5 ± 0.2 (n = 10, range 4–5) setae and outer row with 4.0 ± 0.0 setae (n = 11, range 4); two rows of tubercles on distal three-quarters with in total 50.2 ± 0.4 tubercles (n = 10, range 48–52, ♀ and ♂ combined), inner row with 24.4 ± 0.3 (n = 10, range 23–26) tubercles and outer row with 25.5 ± 0.4 (n = 11, range 23–27) tubercles. Simova-Tošić and Stojanović (2000: fig. 3 a) illustrated setae and tubercles on the fore femur.

Preabdomen. Tergites (Fig. 8 ) blackish brown, thinly pruinose, small white setulae especially laterally; tergite 1 with very distinct subcircular groove and vague transverse ridges (Fig. 8 , groove also shown in Zaitzev (1919: fig. 1), Papp et al. (1997: fig. 1), and Simova-Tošić and Stojanović (2000: fig. 4 d, e); suture between tergites 1 and 2 very distinct; sternites 1–6 dark brown; sternites 1 and 2 glossy, clothed in small setulae (Fig. 18 ); other sternites thinly pruinose, sparsely clothed in small white setulae; sternite 1 short, trapezoid; intersternite 1-2 a solid straight rod-like sclerite, laterally linked to sternite 2 (Fig. 18 ); sternite 2 a slightly trapezoid plate; sternites 3–5 rectangular plates; sternite 6 (Fig. 19 ) somewhat curved, strongly sclerotised, anteromesally a non-sclerotised section.

Female postabdomen. Postabdomen (Fig. 19 ) with “ normal ” shape, not long and narrow like S. munroi ; tergite 7 represented by two glossy, well sclerotised, anteriorly located, triangular sclerites, just separated mesally (Fig. 20 ); tergite 8 two subrectangular, thinly pruinose, sclerites, separated on the meson; tergum 10 short, on the meson broader and posteriorly rounded, thinly pruinose, one pair of apical setulae; cerci curled upward, a striking feature clearly visible in both females from Hungary and Azerbaijan (Figs 9 , 10 ; see also Simova-Tošić and Stojanović 2000: fig. 4 i), rather elongate, l / w ratio: ~ 3.2, sharply tapering apically, clothed in microtrichia and setulae, apically 3 tiny spine-like setulae (Fig. 20 ); sternite 7 (Fig. 19 ; see also Papp et al. 1997: fig. 10) with anteriorly two subtriangular, almost bare, plates well separated on the meson, posteriorly these plates are connected to two posterior subrectangular plates just separated on the meson, these latter plates pruinose and clothed in some setulae; spiracle 7 in membrane; sternite 8 represented by two elongate sclerites, well separated on the meson (Fig. 19 ), pruinose and posteriorly with some setulae; subanal plate (Fig. 21 ) triangular to mitre-shaped with apically a tiny extension, pruinose, apex with one pair of longer setulae, ~ 9 pairs of setulae posteriorly; spermathecae (Fig. 23 ) mushroom-shaped with medium-sized, bell-shaped, hollow, more sclerotised, striated, inner structure, no protuberances, spermathecal ducts very short and broadening distally; sclerotised ring of ventral vagina (Fig. 22 ) small, anteriorly straight and posteriorly semi-circular, arms very slender.

Male postabdomen. Syntergosternite 7 + 8 very slender and wide, weakly sclerotised; spiracles 7 in membrane just anteriorly of syntergosternite; epandrium (Fig. 24 ) rounded, with a large mesal gap, clothed in microtrichia and ~ 10 pairs of setulae; surstyli articulate, l / w ratio: ~ 1.5–1.7, almost touching on the meson, (Figs 24–26 ), simple, subrectangular with slightly concave apical side, outer side (Fig. 25 ) without microtrichia, clothed in ~ 50 setulae, inner side (Fig. 26 ) with only 20 setulae and an almost diagonal ridge on basal half; surstyli interconnected via slender processus longi, processus broadening medially (Fig. 24 ); cerci slender, rather elongate, l / w ratio: 3.9, clothed in microtrichia and on apical half with ~ 10 setulae; phallapodeme (Fig. 28 ) with slender anterior arm, corners rounded, anterior arm 1.5 × longer than posterior arm, lateral processes broad; ejaculatory apodeme straight, slender but apically widening to twice its width (Fig. 27 ), ejaculatory sac normal-sized. Papp et al. (1997: figs 4, 5, 8) give lateral views of the male genital complex, also illustrating the phallic complex with short male genital process. Simova-Tošić and Stojanović (2000: fig. 5 a, b) likewise illustrate the lateral view of the male genital complex.

Biology.

Zaitzev (1919) reported that S. babadjanidesi was attracted by electric light in the evening. Papp et al. (1997) reported on hundreds of S. europaea in October on an overwintering site. Returning in April, the flies were there again, though in smaller numbers. Paulovics (1998) presented data on summer occurrences and distribution along the Hungarian Maros river as well as observations on its ethology. The fly preferred plain and sandy parts of the riverbank and avoided the zone covered by mud. Flies were found 0.5–1.5 m from the river edge. On or near a dead frog, seven or eight Sphyracephala were found. A single fly was also found feeding on a dead ant. In a two-week period in October, the number of assembling flies increased from several to ~ 500. Simova-Tošić and Stojanović (2000) found a cluster of a few thousand flies of S. europaea in early November. Males and females occurred in approximately equal numbers. Kutsarov and Hubenov (2019) reported on clusters of thousands of flies in Bulgaria (see Figs 29 , 30 ). Nartshuk (2017) noted the gregarious behaviour of S. babadjanidesi in Azerbaijan. She also noted that there were only small differences in the descriptions for S. babadjanidesi and S. europaea and that the study of type material would be necessary to determine their synonymy. Papp et al. (1997) and Simova-Tošić and Stojanović (2000) reported on 44 ♀♀ and 55 ♂♂ which would give a sex ratio of 1 ♀: 1.25 ♂ (Table 4 ). However, the latter authors stated that ♂♂ and ♀♀ occurred in equal numbers.

Laboulbeniales (Ascomycota) have never been found on S. babadjanidesi , S. nigrimana , and the two Nearctic Sphyracephala . The long hibernation period might form the reason for this. Rossi and Feijen (2018) noted that Laboulbeniales are common on African Diopsidae , but considerably less common on Oriental Diopsidae . In Afrotropical Sphyracephala Laboulbeniales are very common, but in Oriental Sphyracephala they are rare. It should be noted that the first fossil record of the order Laboulbeniales was found on a fossil diopsid Prosphyracephala in Baltic Amber ( Rossi et al. 2005). Grace and Carr (2020) recorded for S. babadjanidesi (as europaea ) 4 subfamilies of mariner transposons against none in S. beccarii .

Distribution.

The original type series for S. babadjanidesi ( Zaitzev 1919) and its neotype ( Nartshuk 2017) all originate from the Ganja Region in the Asian part of Azerbaijan. The neotype forms part of a large collection ( ZIN) of more than 500 specimens originating from almost the same place as the type series ( Nartshuk 2017). In 2024, S. babadjanidesi was photographed in Georgia (https://www.inaturalist.org/observations/229960704). The type series for S. europaea came from the Maros River in Hungary. This river is a tributary of the Tisza River which in its turn is the main tributary of the Danube. According to Paulovics (1998), the species exists all along the Hungarian part of the Maros (Fig. 31 ). The species was later ( KMNP 2018) recorded from the Körös-Maros Nemzeti Park in Hungary, near the border with Romania (~ 46°41'23"N, 21°10'28"E, ~ 80 m). Rivers in this park are tributaries of the Tisza. Rahmé published pictures (https://www.flickr.com/photos/eurythyrea/5126015816) taken in Makó, Csongrád, Hungary (46°12'11"N, 20°27'11"E, 83 m). Simova-Tošić and Stojanović (2000) extensively reported on the presence of S. europaea in Serbia along the Danube, ca 2 km from the mouth of the Tisza at Stari Slankamen (45°9'5"N, 20°14'44"E, ~ 100 m) Stojanović (pers. comm.) observed the flies again on 14. x. 2006 in the same locality. Early in the morning (8: 00 am) ~ 40 specimens could be observed in the same hollow on the loess profile. Later, at around 4: 00 p. m., more than 200 specimens were gathered in the same place, spread out over an area of ca 3 m 2. Kutsarov and Hubenov (2019) recorded S. europaea in Bulgaria, east of the town of Nikopol, next to the rocky monastery St. Stefan (43°42'36"N, 24°54'51"E, 60 m) on the limestone rocks along the Danube River on the border with Romania. On the internet references for S. europaea in Romania can be found (https://www.flickr.com/photos/eurythyrea/5126015816). All locations for S. europaea are along the Danube River and its tributaries. Papp et al. (1997) stated “ that Hennig (1941 b) hypothesized the occurrence of Sphyracephala in South Europe including Hungary ”. However, that view, also repeated in Földvári and Meier (2002), cannot be deduced from Hennig’s paper. The various collecting localities and the two type localities are shown on the map (Fig. 32 ). In Hungary, this fly has a nature conservation status: collecting it carries a 10,000 HUF fine ( Turista Magazin 2023).

Remarks on synonymy.

Papp et al. (1997) described S. europaea and declared it to be the first known species of the family Diopsidae in Europe. However, fossil species are well known from Europe. Papp et al. (1997) considered S. babadjanidesi the most closely related species to S. europaea . By mistake, they reported Armenia as type locality of S. babadjanidesi . Papp and Földvári (in Papp et al. (1997)), while describing S. europaea , had no access to specimens of S. babadjanidesi . They had to rely on the description and illustrations by Zaitzev (1919), which for its time were certainly of a good standard. However, Zaitzev did not study the genitalia of S. babadjanidesi . Describing a closely related species, without access to flies from the type locality of S. babadjanidesi and without knowledge of genitalia morphology, is not a procedure to be recommended.

Papp et al. (1997) list three “ features ” that are different between S. babadjanidesi and S. europaea . The first one concerns the colour of the fore tarsi. According to Papp et al. “ both description and figure [of Zaitzev] say that fore basitarsus and tarsomeres are yellow in Sphyracephala babadjanidesi , contrasting those of S. europaea . ” In the description, Papp et al. (1997) state “ Fore basitarsus all black, dorsal surface of other fore tarsomeres dark grey, at most 5 th tarsomere light ”. Comparison of photographs (Figs 16 , 17 ) clearly shows that the colour of basitarsus and tarsomeres are similar for flies from Azerbaijan and Hungary. The second difference they listed is “ No dark hue in r 1 cell of Sphyracephala europaea , contrary to S. babadjanidesi . ” This feature is certainly a bit overrepresented in Zaitzev (1919: fig. 1), but is not repeated in the text. Now, comparison of the wings shows no difference in this regard (Figs 12 , 13 ). In wings from both Azerbaijan and Hungary, the very apex of cell r 1 can be slightly darker. The third difference given by Papp et al. (1997) concerns the l / w ratio of the fore femur. In the section on the legs of S. babadjanidesi , their statement that fore femora of S. europaea are more incrassate, has already been rejected (see above, and Table 1 ). The measurements given by Papp et al. (1997) are somewhat haphazard and not well presented. Fortunately, Simova-Tošić and Stojanović (2000) later presented high quality measurements for many characters and based on large series of males and females. The graphs for the ratio’s eye span / body length for flies from Azerbaijan, Hungary, and Serbia (Fig. 5 ) also clearly show no differences in this regard. Likewise, study of the wing morphometrics (Figs 107 – 109 ) supports the conspecificity of S. babadjanidesi and S. europaea .

We present comparative colour photographs for flies from Azerbaijan and Hungary for anterior head (Figs 6 , 7 ), wings (Figs 12 , 13 ) and inner side of fore femur and whole fore legs (Figs 14–17 , 31 ). These already give a strong indication that the same species is involved. Comparison with the large sets of genitalia drawings by Papp et al. (1997) and Simova-Tošić and Stojanović (2000) confirms the view that flies from Azerbaijan, Hungary, and Serbia are conspecific.

Figures

Figure 5. Sphyracephala babadjanidesi, eye span plotted against body length. Most data points were supplied by A. Stojanović (NHMBEO). These Serbian data were also used by Simova-Tošić and Stojanović (2000). Data points for two Azerbaijan flies are marked in blue, while data points of four Hungarian flies are marked in black and white.

Figure 32. Distribution map for records of Sphyracephala babadjanidesi (red dots). The yellow dot indicates the type locality of S. babadjanidesi, while the blue dot indicates the one for S. europaea.

Figure 105. Eye span plotted against body length for males of eight Sphyracephala species. A single data point for S. nigrimana is also shown.

Figure 106. Eye span plotted against body length for females of eight Sphyracephala species. A single data point for S. nigrimana is also shown.

Figure 107. Geometric morphometric PCA analysis of Sphyracephala wing venation patterns. Grey dots represent landmarks that were placed with the aid of straight (dotted) lines through other landmarks (white dots). Asterisks indicate wings from paratypes of S. europaea (junior synonym of S. babadjanidesi).

Figure 108. Hierarchical clustering analysis of principal component scores from the PCA in Fig. 107, using the average clustering method. Branch labels indicate the Approximately Unbiased p-value (AU) and Bootstrap Probability (BP). Asterisks indicate wings from paratypes of S. europaea (junior synonym of S. babadjanidesi).

Figure 109. Hierarchical clustering analysis of principal component scores from the PCA in Fig. 107, using the complete clustering method. Branch labels indicate the Approximately Unbiased p-value (AU) and Bootstrap Probability (BP). Asterisks indicate wings from paratypes of S. europaea (junior synonym of S. babadjanidesi).

Figures 6, 7. Sphyracephala babadjanidesi, head, anterior view 6 ♀, sw Ganja, Azerbaijan 7 ♀, paratype S. europaea, Szeged, Hungary. Scale bars: 0.2 mm.

Figures 8–11. Sphyracephala babadjanidesi 8, 9, 11 ♀, sw Ganja, Azerbaijan 8 habitus, dorsal view 9 habitus, lateral view 10 ♀, paratype S. europaea, abdominal apex with curved cerci, ventral view 11 basiliform prosternum. Scale bars: 1.0 mm (8, 9); 0.2 mm (10); 0.1 mm (11).

Figures 12, 13. Sphyracephala babadjanidesi, wing 12 ♂, sw Ganja, Azerbaijan 13 ♀, paratype S. europaea, Szeged, Hungary. Scale bars: 0.5 mm.

Figures 14–17. Sphyracephala babadjanidesi 14, 15 fore femur, inner side 16, 17 fore leg 14 ♀, sw Ganja, Azerbaijan 15 ♀, paratype S. europaea, Szeged, Hungary 16 ♀, outer side, sw Ganja, Azerbaijan 17 ♂, inner side, paratype S. europaea, Szeged, Hungary. Scale bars: 0.2 mm.

Figures 18–23. Sphyracephala babadjanidesi, ♀, sw Ganja, Azerbaijan 18 sternite 1, intersternite 1-2 and sternite 2, ventral view 19 postabdomen, ventral view 20 postabdomen, dorsal view 21 subanal plate, ventral view 22 sclerotised ring 23 spermathecae. Scale bars: 0.5 mm (18); 0.2 mm (19, 20); 0.05 mm (21–23).

Figures 24–28. Sphyracephala babadjanidesi, ♂, sw Ganja, Azerbaijan 24 epandrium, cerci, surstyli, posterior view 25 surstylus, outer view 26 surstylus, inner view 27 ejaculatory apodeme + sac 28 phallapodeme. Scale bars: 0.1 mm (24, 27, 28); 0.05 mm (25, 26).

Figures 29–31. Sphyracephala babadjanidesi, live 29 cluster of flies in cleft, Nikopol, Danube River, Bulgaria (photograph © Yordan Kutsarov) 30 three flies, same data as for 29 31 fly showing fore femur, Szeged. Maros River, Hungary (photograph © Walter Pflieger).

Figures 110–112. 110 Cladogram of Sphyracephala species groups according to Feijen (1989) 111 preferred maximum likelihood phylogeny of the Sphyracephalinae, based on molecular analysis (Jackson 2019, supplemental figure 1, in slightly adapted form). Numbers at nodes represent Ultrafast Bootstrap supports 112 revised cladogram of Sphyracephala species groups.