Ceriana vespiformis ( Latreille, 1809 )

Steenis, Jeroen Van, Ricarte, Antonio, Vujić, Ante, Birtele, Daniele & Speight, Martin C. D., 2016, Revision of the West-Palaearctic species of the tribe Cerioidini (Diptera, Syrphidae), Zootaxa 4196 (2), pp. 151-209 : 183-186

Ceriana vespiformis ( Latreille, 1809)

Figs 11, 12 , 29, 30 , 47, 48 , 50 , 93 , 109–113 , 170

Ceria vespiformis Latreille, 1809: 328 . Type locality: Barbaria , [North Africa] (LT ♂?, possibly lost).

Ceria scutellata Macquart, 1842: 70 . Type locality Algeria (HT ♂ MNHN).

Ceria intricata Saunders, 1845:64 . Type locality: Albania (ST 2 ♂, 1 ♀ NHM).

Ceria conopsoides: Kassebeer (1999) : 145. Rectification of Lucas (1849).

Ceria scutellata: Kassebeer (1999) : 145. Rectification of Lucas (1849).

Ceria vespiformis: Rondani (1857) , Palma (1865), Rondani (1869), Costa (1882, 1883), Minà Palumbo (1887), Strobl (1893b), Bezzi & De Stefani-Perez (1897), Bezzi (1900), Tuccimei (1908).

Cerioides vespiformis: Efflatoun (1922) comb. nov., Bezzi (1925), Gil Collado (1930), Sack (1932), Venturi (1960), Glumac (1972), Gomes (1978).

Ceriana vespiformis: Becker et al. (1910) comb. nov., Becker (1921), Shaumar & Kamal (1978), Marcos-García (1985), Claussen & Lucas (1988), Peck (1988), Claussen (1989), Brădescu (1989, 1991), Claussen & Hauser (1990), Dirickx (1994) in part, Speight (1994), Belcari et al. (1995), Lucas (1996), Reemer & van Aartsen (2000), Marcos-García & Louis (2001), van Veen (2004), Mason et al. (2006), Riddiford & Ebejer (2006), Rotheray et al. (2006), Reemer & Smit (2007), Standfuss & Claussen (2007), Reemer et al. (2009), Birtele (2011), van Eck (2011), Rotheray & Gilbert (2011), Ricarte et al. (2012), Djellab et al. (2013), Radenković et al. (2013), Sarıbıyık (2014), Burgio et al. (2015), van Steenis et al. (2015).

Ceriana vespiformis Latreille, 1804 , wrong year: Reemer & van Aartsen (2000), Speight et al. (2005), Riddiford & Ebejer (2006), Speight (2013), Sarıbıyık (2014), Ssymank & Weitzel (2014).

Redescription. MALE ( Figs 11 , 29 ). Body length: 8.8–10.7 mm; wing length: 6.3–7.3 mm. Head ( Fig. 47 ). Face slightly protruding antero-ventrally with weakly demarcated tubercle and slightly concave below antennae. Head 2.2–2.3 times as wide as face just below the antennae; width of facial vitta 0.14–0.22 times as wide as width of face; length of eye contiguity 0.53–0.63 times as long as length of frons; angle of eyes at eye contiguity 115–125o. Amount of yellow varies; medial black vitta broad to narrow, sometimes not reaching the mouth edge; a broad black fascia at frontal prominence black to only a black triangular macula around frontal prominence; hypostomal bridge black to entirely yellow; genae with small yellow macula to predominantly yellow with only a broad black vitta from eye margin towards mouth edge. Frons yellow with broad posterior black triangle and broad medial vitta to yellow with small posterior black triangle. Vertical and ocellar triangle black. Dorsal surface of head capsule entirely yellow, evenly broad anteriorly at ocellar triangle. Frontal prominence 3.7–5.0 times longer than wide; relative length of pedicel is as 1.0–1.1: 1: 0.60–0.65. Antennae and frontal prominence brown-yellow to black coloured; arista white pilose. Thorax. Scutum black with yellow; a yellow macula on postpronotum; notopleuron yellow to black, in yellow specimens with additional dark-yellow to yellow posterolateral vittae near scutellum; with white pollinose maculae medially from the transverse suture and two vague white-grey vittae antero-medially. Pleuron black with two or three yellow maculae, one each on posterior 1/2–3/5 of posterior anepisternum, on dorsal 1/4–1/3 of katepisternum and a small one on anepimeron, sometimes this last one is missing. Metasternum with short pile and a low, rounded elevation medially on antero-ventral part. Scutellum entirely yellow.

Legs. Coxae and trochanter black; pro- and mesofemur predominantly black to predominantly yellow; metafemur with apical black to entirely yellow; with apico-medial 1/10–1/3 of tibiae black; pro- and mesotarsi dark-brown, dark-yellow to entirely yellow; metatarsus dark-brown to black. Pile on legs predominantly short and white; pile on apico-posterior part of mesofemur longer, very seldom with 1 or 2 black pili; metatrochanter without setulae. Procoxa broad, about as wide as long, with clear demarcated groove on entire length on dorsal part ( Fig. 50 ); mesotrochanter dorsally normal, straight; mesofemur without flattened area antero-basally; metatrochanter with weak sulcus and narrow rim laterally. Metatibia without appendix on apico-ventral part; metafemur elongate and slightly curved ( Fig. 93 ). Wing. Hyaline except for anterior 1/2 and along spurious vein and along vein M and cross-vein bm-cu. Loop in vein R4+5 weak with short appendix into cell r4+5. Cross-vein r-m slightly curved. Membrane entirely covered in microtrichia. Alula relatively broad, 2.8–3.4 times longer than wide. Abdomen. Black and yellow coloured. Length of tergite I: II: III: IV is as 1: 1.3–1.5: 1.8–2.1: 1.7–1.9. Width of yellow fascia medially: length of tergite of respectively tergite II, III and IV as 1: 3.1–3.3, 1: 4.5–4.8 and 1: 3.1–3.5. Tergite II wider than long, anterior part most narrow; length of tergite II: width of tergite II at respectively anterior: posterior as 1: 1.4–1.5: 1.5–1.7. Tergite I with large triangular yellow maculae on antero-lateral corner, very broadly separated to nearly connected medially; fascia on tergite II straight, hardly narrowed towards lateral margin; fascia on tergite III straight, gradually narrowed towards lateral margin; fascia on tergite IV straight, gradually narrowed towards lateral margin and clearly separated from the lateral margin; tergite II in some specimens with yellow posterolateral maculae. Tergite IV weakly emarginated. Tergite I with low medial elevation; tergites III and IV with low longitudinal elevation; with vague grey-white pollinosity medio-laterally. Sternites I–III with posterior straight yellow fascia, the one on sternite III sometimes missing. Genitalia. Epandrium with narrow ventral rim ( Figs 109– 112 ); in dorsal view, cerci broadly triangular shaped ( Fig. 111 ), pile about 1/4 as long as width of cerci; surstylus bi-lobed, dorsal lobe short broadly elongate with slightly narrow base, ventral lobe rather narrowly circular with membranous basal part ( Figs 109, 110, 112 ); surstylar apodeme narrow circular shaped; hypandrium ( Fig. 113 ) with broad base and narrower apex; with short, broad ventral lobe; superior lobe irregular circular shaped, articulating with hypandrium; aedeagus with baso-ventral elongated and pointed projection, apex with three projections, dorsally short and sharp pointed, medially a short rectangle and ventrally short slightly broad and pointed. FEMALE ( Figs 12 , 30 ). Body length: 9.3–11.8 mm; wing length: 7.2–9.2 mm. Similar to male, except for normal sexual dimorphism and the following characters. Head ( Fig. 48 ). Head 2.0–2.1 times wider than face just below the antennae; width of facial vitta 0.11–0.20 times as wide as width of face. Frontal prominence 5.0–5.7 times longer than wide; relative length of pedicel is as 1.0–1.2: 1: 0.67–0.85. Vertex black. Yellow fascia on dorsal surface of head capsule sometimes with narrow black posterior margin posterior to ocellar triangle. Wing. Alula relatively broad, 2.6–3.2 times longer than wide. Abdomen. Length of tergite I: II: III: IV: V is as 1: 1.6–1.7: 2.1– 2.3: 1.8–2.1: 0.63–0.73. Width of yellow fascia medially: length of tergite of respectively tergite II, III and IV as 1: 3.1–3.9, 1: 3.9–4.9 and 1: 2.5–3.1. Tergite II wider than long, anterior part most narrow; length of tergite II: width of tergite II at respectively anterior: posterior as 1: 1.2–1.4: 1.6–1.8.

Variation. This species is variable in body colour, in such a way that two different phenotypes can be distinguished, a black phenotype and a yellow phenotype. The black phenotype has the following characters, with characters of yellow phenotype between parentheses: medial facial vitta reaching to mouth edge (not reaching mouth edge), fascia at frontal prominence entire (only triangular macula present), hypostomal bridge black (predominantly yellow), frons with broad medial vitta (without black vitta), notopleuron black (yellow), pleuron with two yellow maculae (three yellow maculae), legs extensively black (extensively yellow); alula somewhat narrower (somewhat broader). Several intermediate specimens have been studied and a detailed examination of the male genitalia of different phenotypes, as well as preliminary DNA analyses have shown that these phenotypes do not correspond to separate species. The black phenotype occurs mostly in the northern part of the distributional range and in the spring in the southern part, the yellow phenotype mostly occurs in the southern part in summer and autumn, but no differences have been found in habitat preferences.

Material examined. Holotype of Ceria scutellata : 1 Ƌ, " MNHN, Paris / ED 5760", " Holotype " [red label, "126 / 88" [round white label], "No 1076 / Ceria / scutellata ", " Holotype Ƌ / Ceria scutellata / Macquart , 1842 / design. J. van Steenis, 2016" [red label] (MNHN) . Syntypes of Ceria intricata : 1 Ƌ, "Syn- / type " [round white label with blue margin], " Ceria / Ƌ / Type / intricata / Saund. [ers]" [round white label with orange margin], "73 / 34"[round white label], " Albania / 25 ♀ ", " intricata W. W. S.", " Albania / S.S. Saunders / Ex. coll. / W.W. Saunders / 73.34", " Lectotype Ƌ / Ceria intricata / Saunder, 1845 / design van Steenis, 2016" [red label]; 1 Ƌ, and 1 ♀ with corresponding labels and a red paralectotype label (all NHM) . Additional material: 396 Ƌ, 285 ♀ ( CCF, CEUA, CNBF, DBV, FSUNS, GPA, HMAG, IRSNB, JSA, MNHN, MRL, MSD, MSNF, MTD, MZLCH, NBC, NHM, NHRS, WRL, ZIN) .

Distribution ( Fig. 170 ). Albania, Algeria, Bosnia and Herzegovina, Croatia, Egypt, France, Germany, Greece, Israel, Italy, Malta, Montenegro, Morocco, the Netherlands, Portugal, Serbia, Spain, Tunisia, Turkey.

Biology. The preferred environments are Mediterranean evergreen oak forests ( Quercus ilex and Q. ruber ), wet Liquidamber orientalis forests, alluvial Fraxinus forests and even ruderal places close to some of the before mentioned forest types. Fast flying species, visiting open places at up to 2 m above the ground, settles on vegetation or bare ground. Females have been seen investigating rot holes in live Quercus pubescens . Adults mimic small Hymenoptera both in flight technique and sound produced ( Reemer & Smit 2007; Rotheray & Gilbert 2011; Speight 2013; van Steenis et al. 2015). The larvae are found in decaying roots of Fraxinus angustifolius ( Rotheray et al. 2006) and in sap runs of dead Morbus alba ( Shaumar & Kamal 1978) . The flowers visited, besides those listed by Speight (2013) are: Daucus carota , Elaeoselinum asclepium , Euphorbia luteola , Ferula communis , Ferulago galbanifera , Foeniculum vulgare , Helichrysum stoechas , Paliurus spina-christi , and large Apiaceae . Adults fly from mid April to late September, with a peak in June.

Discussion. The type of Ceria vespiformis was not studied. It could not be found in MNHN nor MHNL and it is possibly lost. The description leaves no doubt about the identity of this species: frontal prominence and brownish pedicel, yellow "caput", genae with yellow macula, entirely yellow scutellum and yellow legs it is our concept of the yellow phenotype of C. vespiformis and we see no need in designating a neotype at the moment.

The holotype of Ceria scutellata has been studied. The description, figure and labels on the type specimen leave no doubt about the type status of the specimen. It is clearly Ceriana vespiformis and belongs to the black phenotype. The synonymy of C. scutellata with C. vespiformis is herewith confirmed.

The syntypes of Ceria intricata have been studied. The description and labels on the specimens studied leave no doubt about the type status of the specimens. All three specimens studied are similar and belong to the black phenotype, hence the synonymy of C. intricata with C. vespiformis is herewith confirmed.

Figures

FIGURES 7 – 12. Ceriana adults, overall appearance, dorsal view. 7. Ceriana naja male, Tajikistan. 8. Ceriana naja female, Turkmenistan. 9. Ceriana sartorum male, Turkmenistan. 10. Ceriana sartorum female, Tajikistan. 11. Ceriana vespiformis male, Greece. 12. Ceriana vespiformis female, Montenegro. Scale line = 5 mm.

FIGURES 25 – 30. Ceriana adults, overall appearance, lateral view. 25 Ceriana naja male, Tajikistan 26 Ceriana naja female, Turkmenistan 27 Ceriana sartorum male, Turkmenistan 28 Ceriana sartorum female, Tajikistan 29 Ceriana vespiformis male, Greece 30 Ceriana vespiformis female, Rhodos. Scale line = 5 mm.

FIGURES 37 – 48. Head, frontal view. 37 Ceriana brunettii male, Tajikistan 38 Ceriana caesarea male, Turkmenistan 39 Ceriana conopsoides male, Greece 40 Ceriana conopsoides female, Russia 41 Ceriana glaebosa male paratype 42 Ceriana media male paratype, Afghanistan 43 Ceriana naja male, Tajikistan 44 Ceriana sartorum male, Turkmenistan 45 Ceriana sartorum female, Tajikistan 46 Ceriana skevingtoni male paratype, Azerbaijan Talish 47 Ceriana vespiformis male, Greece 48 Ceriana vespiformis female, Rhodos. Scale line = 2 mm.

FIGURES 49 – 55. 49 – 52 Procoxa of Cerioidini species, posterior view: 49 Ceriana conopsoides, male, Russia 50 Ceriana vespiformis, male, Montenegro 51 Primocerioides regale, female, Cyprus 52 Sphiximorpha subsessilis, male, The Netherlands. 53 Basisternum of Ceriana conopsoides, frontal view, male, Russia 54 Mesotrochanter of Ceriana caesarea, ventral view, male, Turkmenistan 55 Basisternum of Sphiximorpha subsessilis, frontal view, male, Greece. Scale line = 0.5 mm (a: 49, 50; b: 51, 52; c: 53 – 55).

FIGURES 90 – 93. Metafemur of Ceriana species, male 90 Ceriana caesarea (with tibia), Uzbekistan 91 Ceriana naja, Tajikistan 92 Ceriana sartorum, Kyrgyzstan 93 Ceriana vespiformis, Montenegro. Scale line = 1 mm.

FIGURES 109 – 113. Male genitalia of Ceriana vespiformis 109 Epandrium, both-sides view, Smyrna 110 Epandrium, bothsides view, Rhodos 111 Epandrium, dorsal view, Morocco 112 Epandrium, both-sides view, Morocco 113 Hypandrium, lateral view, Morocco. Scale line = 0.5 mm.

FIGURES 169 – 172. Distribution maps of the West-Palaearctic Cerioidini. 169 Ceriana glaebosa (red square), Sphiximorpha euprosopa (green circle) and Sphiximorpha petronillae (blue triangle) 170 Ceriana vespiformis (red circle) 171 Primocerioides regale (red circle) and Sphiximorpha garibaldii (blue triangle) 172 Sphiximorpha subsessilis (yellow circle).