Sphyracephala beccarii ( Rondani, 1873 )

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 beccarii ( Rondani, 1873)

Figs 2 , 4 , 33–36 , 37 , 38–42 , 43–47 , 48–53 , 105 , 106 , 107 , 108 , 109 , 110–112 , Tables 2 , 3 , 4

Diopsis beccarii Rondani, 1873: 289.

Hexechopsis beccarii (Rondani): Rondani 1875: 442. Osten Sacken 1882: 235; Brunetti 1907: 163; Bezzi 1922: 71; Eggers 1925: 488, 493; Séguy 1949: 74, 1955: 1123.

Sphyracephala beccarii (Rondani): Osten Sacken 1882: 235. Bezzi 1922: 69; Brunetti 1928: 273; Curran 1928: 274; Hennig 1941 a: 62, 1941 b: 6, figs 1, 5 d, 1965: figs 46, 49 c, 58 a; Séguy 1949: 75, figs 6, 7; 1950: 276; Vaillant 1953: 11, figs on p. 12; Collart 1954: 329; Lindner 1954: 28; 1962; 17; Séguy 1955: 1124; Guiglia 1957: 194; Descamps 1957: 17, figs 7 b – 7 f; van Bruggen 1961: 425, figs 12, 15; Steyskal 1972: 13; Feijen 1978: 19; 1983: fig. 5; 1987: 420; 1989: 67; Cogan and Shillito 1980: 584; Ferrar 1987: figs 28.34, 28.49, 28.62, 28.63, table 28.1; Rossi 1990: 3; Blackith and Guillet 1995: 66; Jakobs 1997: 13; McAlpine 1997: 172, figs 23, 24, 27; 2011: 150, figs 123, 125; Papp et al. 1997: 137; Hariri et al. 1998: 254, tabs 1–4; Baker 1999: tab 2-1, figs 2-2, 2-3, app. A; Lande and Wilkinson 1999: table 1; Wilkinson and Taper 1999: 1687, fig. 1, table 1; Simova-Tošić and Stojanović 2000: 149; Hilger 2000: 337, figs 7.3–7.7; Meier and Hilger 2000: 6, figs 9–11, table 2; Baker and Wilkinson 2001: figs 2, 3, table 1; Baker et al. 2001: figs 1, 2; Hurley et al. 2001: 408, 1 C-E, 2 D, 3 C (as beccarri); Hurley 2002: 2, figs 1.3, 2.1. C-E, 2.2. D. 2.3. C; Meier and Baker 2002: 333, figs 1 b, 2; Cotton 2004: 2, figs 4.1, 4.2, 4.3, table 4.1; Cotton et al. 2004: 1310, figs 1-3, table 1; Chapman et al. 2005: 534; Carr et al. 2005: 403, figs 4, 5, 2006 a: 5, figs 1 g, 2, 2006 b: fig. 22; Warren and Smith 2007: figs 1 a, 2, 3 b, 3 d; Carr 2008: 114, fig. 1, table 1; Dawah and Abdullah 2008: 89; Ribak et al. 2009; 861, figs 1, 3, 4, 5, 7, table 1; Hauser et al. 2011: 765, figs 1–3; O’Hara et al. 2011: 96; Baker et al. 2012: 2368; Marshall 2012: fig. 8 on p. 473; El-Hawagry et al. 2013: 60, 2016: 124, 2017: 17; Voje and Hansen 2013: figs 1, 2, tab 1; Baker et al. 2016: 898, figs 1, 2, table 1; Feijen et al. 2017: 76, figs 1, 5, 11, 17, 18, 20; Feijen et al. 2018: 142, figs 189, 194, 199; Mader 2017: 108; Nartshuk 2017: 129; Jackson 2019: suppl. fig. 1, 2; Picker et al. 2019: 344; Grace and Carr 2020: 18 / 24, table 1; Feijen and Feijen 2021: 1484, figs 4, 48; 2022: 1111, fig. 9.56 a; 2023: 84. [Given the frequent spelling errors in beccarii (1 or 2 c’s, 1 or 2 r’s, 1 or 2 i’s), this should be taken into account for digital searches.]

Sphyracephala africana Karsch, 1888: 380, pl. 4 fig. 11. Speiser 1910: 166, 1924: 100; Eggers 1925: 493; Brunetti 1926: 84, 1928: 273; Curran 1928: 274; Séguy 1933: 32, 1938: 237, 1949: 75, 1955: 1124; Collart 1954: 329; Lindner 1954: 28; Descamps 1957: 17; van Bruggen 1961: 426; Steyskal 1972: 13; Feijen 1978: 20; Cogan and Shillito 1980: 584; Feijen et al. 2017: 76.

Sphyracephala hearseiana (Westwood): Bezzi 1922: 69, Algeria (misidentification). Hennig 1941 b: 6; Séguy 1955: 1124; van Bruggen 1961: 426.

Type series.

Sphyracephala (Diopsis) beccarii . Eritrea: Sciotel, Bogos [1870, 15°35'N, 38°20'E, 780 m], 61 syntypes in MSNG ( Sforzi and Sommaggio 2021), more syntypes in various other museums (MLUH, NHMUK) . No lectotype has been nominated ( Sforzi and Sommaggio 2021), although Guiglia (1957) mentioned “ Tipo e numerosi cotipi ”.

Sphyracephala africana . Tanzania: holotype, ♀, Bondei , [~ 5°00'S, 39°00'E, 100 m, i.1886 / 87], ZMHB . The holotype is not listed in Rohlfien and Ewald (1970), but it is present in the ZMHB collection (Sven Marotzke, pers. comm. 2024).

Material examined.

It would go too far to list all the S. beccarii we have examined since 1971. Here we only list the totals examined per country or region: Eritrea, syntypes 3 ♀, 3 ♂, Sciotel, Bogos, 1870, O. Beccari ( MLUH, V. Röder collection); Algeria, 1? sex, Rhouffi, vii. 1949, Vaillant ( ZSM); Arabian Peninsula 75 ♀, 80 ♂ (see Feijen et al. 2017); Benin, 1 ♀ ( RMNH); Botswana, 1 ♀ ( RMNH); Burkina Faso, 1 ♂ ( RMNH); Cameroon, 1 ♀ ( BMSA); DR Congo, 3 ♀, 5 ♂ ( CSCA); Ethiopia, 2 ♀, 1 ♂ ( FBUB); Gambia 1 ♀ ( RMNH); Ghana, 19 ♀, 14 ♂ ( RMNH, CSCA); Kenya, 4 ♀, 4 ♂ ( RMNH); Madagascar, 528 ♀, 276 ♂ ( CAS); Malawi, 314 ♀, 322 ♂, 1971–1975 ( RMNH); Mozambique, 131 ♀, 135 ♂, 1976–1982 ( RMNH); Niger, 8 ♀ ( RMNH); Senegal, 2 ♀ ( RMNH); South Africa 6 ♀, 1 ♂ ( RMNH); Tanzania, 26 ♀, 35 ♂, 1982–1988 ( RMNH); Togo, 17 ♀, 32 ♂ ( FBUB, SMF, RMNH; Zambia, 1 ♀ ( RMNH); Zimbabwe 7 ♀, 2 ♂ ( AMGS, RMNH). In total 622 ♀ and 635 ♂ were examined for Continental Africa and the Arabian Peninsula, giving a balanced sex-ratio of 100 ♀: 102 ♂. However, a different picture emerged for Madagascar: 528 ♀ and 276 ♂ were found, based on 84 malaise trapping periods in 2002–2004, which gives a sex-ratio of 100 ♀: 52 ♂ (see also Table 4 ). This striking difference will be discussed in the section on sex-ratio.

Diagnosis.

Sphyracephala beccarii can be recognised by the following set of characters: head brown, thorax and abdomen blackish brown; sparsely covered with small setulae; frons with dark brown semicircular band; occiput yellowish brown; eye stalk stout (~ 0.75–0.80 × the widest sagittal eye diameter), moderately sized for a Sphyracephala ; very small eye span (~ 2.1 mm) in both ♀ and ♂ (respectively ~ 49 % and ~ 53 % of body length); monomorphic with rate of dimorphism D = 0.07; distinct precoxal bridge; apical seta / scutellar spine ratio: ~ 3.9; scutellar spine / scutellum ratio: 0.50; small, pale scutellar spines ~ 0.17 mm; transparent wings; fore femur brown with apical fifth dark brown, inner side with dark brown transverse stripe on central third, strongly incrassate, l / w ratio: 2.5–2.6, with two rows of black spinous setae, inner row with ~ 6.0 setae, outer row with ~ 1.2 setae; tergite 1 with vague transverse ridges, on the meson two parallel, longitudinal grooves; intersternite 1-2 very slender, laterally connected to main sternite 2; ♀ tergite 7 and sternite 7 divided in two small sclerites almost touching laterally; ♀ cerci broad, l / w ratio: ~ 1.9; ♀ sternite 8 represented by two small sclerites, almost touching on the meson; no sclerotised ring; surstyli articulate, almost touching on the meson, tapering apically towards an upturned apex, anterior side with microtrichia on basal third and ~ 25 setulae on apical half. Sphyracephala beccarii belongs to the S. hearseiana species group and can be considered the sister species of S. hearseiana .

Redescription.

The following redescription considers the original descriptions by Rondani (1873) and Karsch (1888), description and figures by Hennig (1941 b: figs 1, 5 d; 1965: figs 46, 49 c, 58 a), Séguy (1949: figs 6, 7), Vaillant (1953: figs on p. 12), the table of differences between S. beccarii and S. munroi by Collart (1954), descriptions and figures by Descamps (1957: fig. 7 b – f), description and illustrations by van Bruggen (1961: figs 13, 14, 16), Feijen (1983: fig. 5), description and figures of antenna by McAlpine (1997: 172, figs 23, 24, 27; 2011, figs 123, 125), figures by Hilger (2000: figs 7.3–7.7), egg description and figures by Meier and Hilger (2000: figs 9–11).

Measurements. Body length ♀ 4.26 mm ± SE 0.04 (range 3.54–4.64, n = 40), ♂ 3.91 mm ± 0.04 (range 3.32–4.27, n = 40), eye span ♀ 2.09 mm ± 0.02 (range 1.75–2.31, n = 40), ♂ 2.08 mm ± 0.02 (range 1.78–2.27, n = 40); wing length ♀ 3.46 mm ± 0.04 (range 3.17–3.60, n = 10), ♂ 3.09 mm ± 0.08 (range 2.75–3.54, n = 10); length of scutellar spine ♀ 0.176 ± 0.004 (range 0.169 –0.193, n = 10), ♂ 0.171 mm ± 0.004 (range 0.145 –0.193, n = 10). Baker and Wilkinson (2001) found ♀ mean body length 4.97 mm, ♂ 4.50 mm; ♀ mean eye span 2.10 mm, ♂ 2.05 mm.

Head. Central head (Figs 33 , 34 , 36 ) brown, arcuate groove dark brown; frons with dark brown semicircular band running from arcuate groove via base of inner vertical seta to ocellar tubercle; stalks dorsally and posteriorly largely blackish; occiput yellowish brown, slightly darker dorsally; head uniformly pruinose (Figs 34 , 36 ), head with a few small black setulae dorsally, ventrally more and longer whitish setulae; arcuate groove distinct blackish; frons with rectangular elevation below ocellar tubercle, grooves laterally of elevation; face flat, no facial teeth, lateroventral corners rounded, facial sulcus absent, but ventral facial edges slightly turned upward medially; eye stalk stout, ~ 0.75–0.80 × the widest sagittal eye diameter; eye span very small in both female (49.2 % ± SE 0.1 % of body length, n = 40) and male (53.3 % ± SE 0.1 % of body length, n = 40); a monomorphic species with rate of dimorphism D = 0.07 (Figs 37 , 105 , 106 , Table 2 ); inner vertical seta long,> 0.4 mm, 1.2 × diameter of eye stalk; outer vertical seta long,> 0.3 mm, 0.9 × diameter of eye stalk (Figs 34 , 36 ). A drawing of the antenna is provided by Feijen (1983: fig. 5). McAlpine (2011: figs 123, 125) provided a scanning electron microscope picture of the conus of the pedicel and a drawing of funiculus and basal segments of arista. McAlpine (1997: figs 23, 24) discussed the taxonomic importance of the ultrastructure of the face and provided electron micrographs of the lower part of face and parafacials and the microtrichose crazed cuticle of face. In a thesis, Jakobs (1997) studied the fine structure of the optical system. Jakobs measured 30 females and 30 males. She found for ratio eye span / body length in females 44.4 % and in males 45.8 %, whereas she found a rate of dimorphism D = 0.05, clearly indicating a monomorphic species. Baker and Wilkinson (2001) found for ratio eye span / body length in females 42.3 % and in males 45.6 %, whereas they found a rate of dimorphism D = 0.20 which still qualifies S. beccarii as a monomorphic species.

Thorax. Collar, scutum and scutellum blackish brown with few small setulae (Figs 35 , 36 ), scutum and scutellum with fine granulated structure; scutellar spines whitish with brown base; pleura blackish brown, uniformly pruinose; posterior notopleural seta quite long; infra-alar seta long, almost twice length of notopleural seta (Fig. 36 ); supra-alar carina indistinct; distinct precoxal bridge (Fig. 2 ); scutal length / scutal width ratio: 0.85; scutellum trapezoid, narrowing distally; scutellar spines small, straight, slightly turned upward, diverging at angle of ~ 60 °; scutellar spine / scutellum ratio: 0.50 ± 0.01 (n = 25, see Table 3 ); scutellar spine / length of body ratio: 0.044 ± 0.001 (n = 20); apical seta / scutellar spine ratio: 3.89 ± 0.07 (n = 25); scutellar length / scutellar width (at base) ratio: 0.69 ± 0.01 (n = 20). McAlpine (1997: fig. 27) illustrated the median ventral region of sternopleura, showing double series of pits.

Wing. Transparent with only the faintest brownish hue (Fig. 38 ); vein CuA + CuP from vein CuP onward extending under angle of 45 ° to two-thirds of wing margin in straight line; vein M 4 continuing distal of crossvein dm-m to one quarter of distance to wing margin; cell cua very narrow, width near base and apex equal (Fig. 38 ); crossvein h distinct; glabrous area only includes basal quarter of cell br. Wing pictures were provided by van Bruggen (1961: fig. 12). Hilger (2000: fig. 3), Feijen et al. (2017: fig. 11) and Feijen and Feijen (2018: fig. 189). In the drawing by Hilger, cell cua is clearly misrepresented.

Legs. Fore coxa and trochanter very pale, thinly pruinose, with some setulae; fore femur (Figs 39–41 ) brown, thinly pruinose, apical fifth dark brown on inner and outer side, inner side with dark brown transverse band, dorsally connected to dark apex, sparsely clothed in small setulae; fore tibia dark brown, thinly pruinose; basitarsus brown, other tarsomeres pale, thinly pruinose and with rows of blackish setulae (Fig. 41 ); mid and hind legs pale brown, femora with dark brown apical third, hind tibia with dark brown apex; fore femur strongly incrassate (Table 2 ), l / w ratio: 2.48 ± SE 0.02 in ♀ (n = 10) and 2.53 ± 0.03 in ♂ n = 15); fore femur with two rows of black spinous setae on distal half with in ♀ 7.5 ± SE 0.2 setae (n = 16) and in ♂ 6.8 ± 0.2 (n = 16) setae, inner row with 6.0 ± 0.1 (n = 32) setae and outer row with 1.2 ± 0.1 setae (n = 32), two rows of tubercles on distal three-quarters with in ♀ 49.8 ± 0.8 tubercles (n = 16) and in ♂ 46.9 ± 0.6 (n = 16), inner row with 22.3 ± 0.2 (n = 32) tubercles and outer row with 26.1 ± 0.3 (n = 32) tubercles. Curran’s (1928) key separating S. beccarii from S. munroi by the “ Tibiæ and tarsi largely or wholly yellowish ” for the former and the tibiae and tarsi brown for the latter should be disregarded. Already in Rondani’s (1873) description it was clearly stated “ antici .... tibiis, et metatarso nigricantibus ”. Collart (1954) in his table indicated as differential characters for S. beccarii that the fore femur was strongly incrassate, hind tibia only black at apex and for S. munroi that the fore femur was moderately incrassate, hind tibia completely black.

Preabdomen. Tergites (Fig. 42 ) blackish brown, thinly pruinose, small setulae laterally; tergite 1 with very vague transverse ridges, on the meson two parallel, longitudinal grooves (Fig. 42 ); suture between tergites 1 and 2 just visible; sternites 1–6 brown, all covering the width of the abdomen, sternite 1 and basal half of sternite 2 glossy, other sternites thinly pruinose, sparsely clothed in small white setulae; sternite 1 short, trapezoid; intersternite 1-2 absent, sternite 2 a uniform, slightly trapezoid plate (Fig. 46 ); sternites 3–6 rectangular.

Female postabdomen. Postabdomen short, broad (Fig. 43 ); tergite 7 represented by two small, strongly sclerotised, laterally located, sclerites; tergite 8 (Fig. 45 ) two square, thinly pruinose, sclerites, laterally located, broadly separated on the meson; tergum 10 short, extending posteriorly on the meson, thinly pruinose, one pair of apical setulae; cerci broad, l / w ratio: ~ 1.9, clothed in microtrichia and setulae; sternite 7 consisting of two small, strongly sclerotised, laterally located, angular sclerites, almost touching tergite 7; spiracle 7 located in membrane in between sternites 6 and 7 (Fig. 43 ); sternite 8 represented by two small sclerites, almost touching on the meson (Fig. 4 ), near the genital pore, clothed in microtrichia, 6 pairs of long setulae and some small setulae; subanal plate (Fig. 44 ) kidney-shaped with rounded apex and rounded anterolateral corners, apex with one pair of longer setulae, clothed in microtrichia and a few pairs of small setulae; spermathecae (Fig. 47 ) mushroom-shaped with large, bell-shaped, hollow, more sclerotised, striated, inner structure, no protuberances; sclerotised ring of ventral vagina absent. Hilger (2000: figs 4, 5) provides detailed drawings of ventral and lateral views of the postabdomen.

Male postabdomen. Syntergosternite 7 + 8 very slender, extending the width of the abdomen, slightly angular on the meson (Fig. 53 ); spiracles 7 in membrane; epandrium (Fig. 48 ) rounded, clothed in microtrichia and ~ 30 pairs of setulae; surstyli articulate, almost encircled by the epandrium, nearly touching on the meson, tapering apically towards an upturned apex (Figs 48–50 ), outer side with microtrichia on basal third, apical half with ~ 25 setulae (Fig. 49 ), inner side with only a few small setulae on apical half (Fig. 50 ); surstyli interconnected via broad processus longi; cerci tapering basally and apically, broadest at one-third from apex (Fig. 48 ), length / broadest width ratio: 2.8, clothed in microtrichia and on apical half with more than 30 setulae; phallapodeme (Fig. 52 ) with slender anterior arm, corners pointed, anterior arm slightly longer than posterior arm, lateral processes “ vane ” broad; phallus broad, short, male genital process hardly sticking out from apex; ejaculatory apodeme straight, very slender (Fig. 51 ), ejaculatory sac normal-sized.

Hennig (1941 b: figs 1, 5 d) illustrated the inner genitalia and epandrium with surstylus. Vaillant (1953: figs on p. 12) presents drawings of the postabdomen, but the surstylus is shown as fused to the epandrium, syntergosternite is lacking and the terms ventral and dorsal should be reversed.

Van Bruggen (1961) stated that male genitalia in Sphyracephala “ are too uniform to facilitate identification of the species ”. However, Hennig (1941 b) already stated that S. beccarii and S. hearseiana are extraordinarily similar, but can be distinguished by epandrium and surstyli (see also Figs 48 , 49 and Figs 132 , 133 ). Hilger (2000: figs 6, 7) gives drawings of the lateral views of male postabdomen and the phallic complex while Feijen and Feijen (2021: fig. 48) give a posterior view of the epandrium.

Egg, larva, and pupa. Descamps (1957) was the first to note the reticulation of the chorion of the egg as “ non strié longitudinalement ” and “ un fin réseau de petits polygones irréguliers. ” Feijen (1989) stated that the absence of longitudinal ridges might represent a apomorphic character for Sphyracephala . Meier and Hilger (2000) studying three Sphyracephala including S. beccarii , stated that the eggs are “ entirely covered with hexagonal reticulation, chorion never striated ”. They considered the egg ornamentation as a diagnostic character for the genus. Micrographs were provided of dorsal egg, micropyle, posterior pole ( Meier and Hilger 2000: figs 9, 10, 11). Descamps (1957) stated that eggs are laid on decomposing plant material. Descamps (1957: pl. 7, figs c – f) described larva and puparium and illustrated larval cephaloskeleton, larval posterior spiracles and puparium.

Biology.

Descamps (1957) reared S. beccarii in Cameroon. Eggs were laid on decomposing plant matter. Descamps described the saprophagous larvae and the time the various stages take. From egg to fly took approximately two weeks. Descamps indicated that S. beccarii often constitute large swarms in the dry season. The flies then disperse at the start of the rainy season. In Algeria, Vaillant (1953) detected among rocks in an oasis swarms of S. beccarii . Feijen (1984) found in the dry season in Malawi among rocks in a river bedding a dense mass of S. beccarii . After being disturbed they flew up. A single sweep of a net yielded more than 6,000 specimens. The size of the whole mass was estimated to be approximately 100,000 specimens, while the cluster took up an area of less than 0.2 m 2. Feijen et al. (2017) described and illustrated a cluster of more than 80,000 S. beccarii on a tree trunk near a river in Wadi Darbat, Oman. Picker et al. (2019) recorded for South Africa that S. beccarii forms groups in moist, rocky places near water. Their idea that these flies mimic small jumping spiders appears unlikely. Mader (2017) mentions for S. beccarii the antipodal position during copulation. However, all photographs for Sphyracephala and other diopsids show an epipodal position during copulation. Descamps (1957) noted that copulation takes several hours in S. beccarii .

Rossi (1990) described Stigmatomyces beccarii ( Laboulbeniales ) from S. beccarii , while Stigmatomyces elongatus was described from S. munroi . Both new fungi were described from flies from Malawi. It is interesting to note that these two sympatric Sphyracephala were parasitised by very different Stigmatomyces . Hariri et al. 1998 recorded the presence of the bacteria Type A Wolbachia in S. beccarii . Wolbachia can be associated with female-biased sex ratio distortion. As we report on a female-biased sex ratio in S. beccarii from Madagascar only, it would be interesting to compare the Wolbachia’s in flies from Madagascar and mainland Africa. Carr (2008) found no evidence for the presence of subfamilies of transposable elements in S. beccarii , though three independent lineages were found in S. babadjanidesi (as S. europaea ).

Some minor contradictions are found in the records for the rate of dimorphism for S. beccarii . Wilkinson and Taper (1999) considered S. beccarii a monomorphic species. They found an eye span / body length ratio of 0.44 for ♀ and 0.47 for ♂ (we found 0.49 and 0.53 respectively). As rate of dimorphism, they gave D 0.54–0.35 = 0.19 (we found D 0.56 - 0.49 = 0.07, see Fig. 37 ). The same data were presented in Baker and Wilkinson (2001), but due to a difference in rounding off a D of 0.20 is given, while S. beccarii was classified as a dimorphic species. Jakobs (1997) found an eye span / body length ratio of 44.4 % for ♀ and 45.8 % for ♂, and a rate of dimorphism D = 0.05, indicating a monomorphic species. Carr et al. (2005) stated that S. beccarii has identical mean eye span in males and females, but female body size is greater, leading to sexual dimorphism in relative eye span. However, that is not the way the rate of dimorphism is determined in Diopsidae , cf. Baker and Wilkinson (2001) and Feijen and Feijen (2009, 2021). Referring to the data of Baker and Wilkinson (2001), Cotton et al. (2004 a) considered S. beccarii a species with only slight (or weak) sexual dimorphism for eye span. Chapman et al. (2005) considered S. beccarii a sexually monomorphic species. Ribak et al. (2009) found for S. beccarii no significant differences in the mass-adjusted eye-span between the sexes. Voje and Hansen (2013) used for S. beccarii the data of Baker and Wilkinson (2001). Their calculations of allometric slope and intercept were based on the least-squares regression of log eye span as function of log body size. Accordingly, they found that slope and intercept are different between the sexes at the 95 % confidence level.

Distribution.

Sphyracephala beccarii is known to occur in almost all contiguous Sub-Saharan African countries and Madagascar. We have seen specimens or records from Benin, Botswana, Burkina Faso, Burundi, Cameroon, Chad, D. R. Congo, Eritrea, Eswatini, Djibouti, Ethiopia, Gambia, Ghana, Kenya, Malawi, Mali, Mozambique, Namibia, Niger, Nigeria, Rwanda, Senegal, South Africa, Sudan, Tanzania, Togo, Uganda, Zambia, and Zimbabwe.

Sphyracephala beccarii extends into the Palaearctic Region in Algeria ( Bezzi 1922, Hennig 1941 b, Séguy 1949, Vaillant 1953). Bezzi referred the Algerian flies to S. hearseiana , but Hennig considered that a likely misidentification for S. beccarii , while Séguy confirmed that the two flies studied by Bezzi belonged to S. beccarii . Vaillant illustrated male genitalia of his Algerian flies and provided reliable information on habitat and swarming. Only his assumptions on the predatory nature of these flies must be rejected. We examined a specimen collected in 1949 in Algeria by Vaillant ( ZSM). Papp et al. (1997) considered S. beccarii “ an Afrotropical species with one questionable record from Algeria ”. However, there is no reason to consider the Algerian records as doubtful. Moreover, Séguy (1950) recorded S. beccarii for Monts Bagzane in northern Niger, not far from Algeria. Sphyracephala beccarii also extends into the Palaearctic Region in the Arabian Peninsula ( Feijen et al. 2017). An extensive number of records were provided for Oman, Saudi Arabia, the United Arab Emirates and Yemen. They discussed the delimitation of the Afrotropical and Palaearctic Regions in the Arabian Peninsula.

Figures

Figure 37. Sphyracephala beccarii, eye span plotted against body length.

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 1, 2. Prosternum, ventral view 1 basiliform prosternum, ♂, Sphyracephala munroi, Arusha, Tanzania 2 precoxal bridge, ♂, Sphyracephala beccarii, Mboma, DR Congo. Scale bars: 0.1 mm.

Figures 3, 4. ♀, sternite 8, ventral view 3 Sphyracephala munroi, Arusha, Tanzania 4 Sphyracephala beccarii, Maputo, Mozambique. Scale bars: 0.1 mm.

Figures 33–36. Sphyracephala beccarii 33 live photograph, Mikumi, Tanzania (photograph © Stephen Marshall) 34 ♂, head, anterior view, Wadi Maharish, Saudi Arabia 35 ♀, thorax, lateral view, Mahavelo Forest, Madagascar 36 ♀, thorax, dorsal view, Niamey, Niger. Scale bars: 0.5 mm.

Figures 38–42. Sphyracephala beccarii 38 ♂, wing, Diampwe Malawi 39, 40 ♀, fore femur, Wadi Maharish, Saudi Arabia 39 inner view 40 outer view 41 ♂, Limbe, Malawi, fore leg, inner view 42 ♀, Dedza, Malawi, abdomen, dorsal view. Scale bars: 0.5 mm (38, 41, 42); 0.2 mm (39, 40).

Figures 43–47. Sphyracephala beccarii, ♀, 43, 44, 47 Maputo. Mozambique, 45, 46 Zomba, Malawi 43 postabdomen, ventral view 44 subanal plate, ventral view 45 tergite 8, 10 and cerci, dorsal view 46 sternites 1 and 2, ventral view 47 spermathecae. Scale bars: 0.2 mm (43, 46); 0.1 mm (44, 45, 47).

Figures 48–53. Sphyracephala beccarii, ♂, Diampwe River, Malawi 48 epandrium, cerci, surstyli, posterior view 49 surstylus, outer view 50 surstylus, inner view 51 ejaculatory apodeme and sac 52 phallapodeme 53 syntergosternite 7 + 8, anterodorsal view. Scale bars: 0.1 mm.

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.

Figures 132, 133. Sphyracephala hearseiana, Rajasthan, India 132 epandrium, cerci, surstyli, posterior view 133 surstylus, outer view. Scale bars: 0.1 mm.