Celonites ivanovi Mauss & Fateryga, 2022

Mauss, Volker, Fateryga, Alexander V., Yildirim, Erol & Carpenter, James M., 2022, Contribution to the taxonomy, bionomics and distribution of the Palaearctic Celonites cyprius - group (Hymenoptera, Vespidae, Masarinae) with the description of two new species from the North Caucasus and East Anatolia, Journal of Hymenoptera Research 89, pp. 109-155 : 109

Celonites ivanovi Mauss & Fateryga sp. nov.

Holotype.

♀ (dbM 5492), "[Russia] Dagestan, Maydanskoye 42°36'16"N 46°58'10"E [corrected to 42°36'07"N, 46°58'13"E in 2021] 11.VI.2019 on Heliotropium styligerum leg. [A.V.] Fateryga" OLML (Figs 5e , 6c ).

Paratypes.

"[Russia] Dagestan, Untsukulskiy distr. vicinity of Maydanskoye [42°37'36"N 46°56'48"E] on Heliotropium styligerum 23.06.2018 leg. [A.V.] Fateryga", 1♀ AF, 1♀ (dbM 5287) VM; "[Russia] Dagestan Maydanskoye 42°36'16"N 46°58'10"E [corrected to 42°36'07"N, 46°58'13"E in 2021] 11.VI.2019 leg. [A.V.] Fateryga", 1♂ (dbM 5497) AMNH, 1♂ (dbM 5498) OLML, 1♂ ZISP, 3♂ AF, 2♂ (dbM 5496, 5499) VM; "[Russia] Dagestan, Maydanskoye 42°36'16"N 46°58'10"E [corrected to 42°36'07"N, 46°58'13"E in 2021] 11.VI.2019 on Heliotropium styligerum leg. [A.V.] Fateryga", 1♀ (dbM 5494) AMNH, 1♀ ZISP, 5♀ AF, 3♀ (dbM 5490, 5491, 5493) 1♂ (dbM 5495) VM; "[Russia] Dagestan, Maydanskoye 42°36'07"N, 46°58'13"E 15.VI.2021 on Heliotropium styligerum leg. [A.V.] Fateryga", 4♀ (dbM 5856, 5857, 5858, 5995) VM; "[Russia] Dagestan, Maydanskoye 42°36'07"N, 46°58'13"E 16.VI.2021 on Heliotropium styligerum leg. [A.V.] Fateryga", 2♀ (dbM 5859, 5860) VM; "[Russia] Dagestan, Maydanskoye 42°36'07"N, 46°58'13"E 16.VI.2021 leg. [A.V.] Fateryga", 1♂ (dbM 5996) VM; "[Russia] Dagestan, vicinity of Turtsi 42°11'34"N, 47°09'33"E on Heliotropium styligerum 22.VI.2021 leg. [A.V.] Fateryga", 1♀ (dbM 5997) VM.

Diagnosis.

See key.

Description.

Female. Colour (Figs 2 , 4e ): Black. The following are yellowish-white: large rectangular spot dorso-medial on clypeus; two small spots adjoining ventral margin of clypeus to both sides of median emargination; small spot on each ocular sinus and two spots on frons; medium-sized spot on antero-dorsal angle of pronotum (humeral spot); broad stripe along dorso-medial (inner) margin of pronotum, anteriorly somewhat angularly enlarged with little median dent; large spot on dorsal mesepisternum; laterally directed process of axilla; medium-sized spot postero-medially on scutellum; dorsal and ventral side of propodeal lamella; antero-lateral one-fifth and posterior two-fifth of tegula, interrupted by brownish translucent area on bulge changing into black towards antero-medial margin of tegula; posterior band on tergum I occupying whole of sides but less than half of middle part, somewhat widened anteriorly in median axis, anteriorly with small brownish tinge towards adjacent black area; laterally and medially widened posterior bands on terga II-V, anteriorly with small brownish tinge towards adjacent black area, interrupted on each side of middle by brownish area (Fig. 6b ); little longitudinal spot antero-medial on tergum VI; small spots on postero-lateral edges of sterna II-III; outside of distal tips of fore-, mid- and hind-femora; outside of proximal end of fore-tibia; outside of mid-tibia except small diagonal blackish-brown interruption in middle; outside of hind-tibia except blackish ring covering one-third below middle. Brown are: distal half of mandible; maxillary and labial palpi, protrudeable parts of proboscis; ventral margin of labrum; translucent ventro-medial margin of clypeus; postero-lateral margin of scutellum; median third of metanotum; postero-lateral process of propodeum; humeral plate (at base of wing underneath tegula); sides and posterior translucent margin of tergum VI; tarsi; sternum I; sterna II-V, lighter along posterior translucent margin. Antenna black except: weak yellowish markings medial on A4-A6; brown-suffused area ventral on A9-A11 and proximal part of A12. Wings moderately infuscate, pterostigma black, veins black becoming somewhat lighter at base.

Variation (number of specimens in brackets): Yellowish-white markings: clypeus with two additional lateral spots (1), only with two small separate spots dorso-medial on clypeus (1) or clypeus completely black (1); spot on ocular sinus and frons narrowly fused (1); spot on ocular sinus or frons asymmetrically reduced (4) or completely reduced (1); short narrow spot on gena along postocular carina at dorso-lateral corner of head (3); humeral spot small (4); little spot antero-ventrally on mesepimeron (1); little spot postero-medially on mesoscutum (5); bands on terga II-V interrupted on each side of middle by blackish-brown area (3); antero-medial spot on tergum VI absent (2); continuous stripe on outside of mid-tibia (5); continuous stripe on outside of hind-tibia (2) or marking on outside of mid- and hind-tibia reduced to small area at distal and proximal end (3); little spot disto-medially on A3 (1), weak yellowish markings medial on A7 (3).

Structure: Head in front view 1.48 times as wide as long in median (min 1.45, max 1.53, n=5) (Fig. 4e ). Mandible with two large blunt incisivi at distal end separated by acute-angled cleft and two smaller more acute subapical incisivi on antero-medial margin. External side of mandible (Fig. 5d ) distally bearing longitudinal rows of long stiff setae; at base without distinct transverse depression; basal area with shagreened cuticula moderately covered with pubescence of tiny thin setae (longer than in C. cyprius ); anterior to condylar ridge cuticula of basal area extends further apically becoming distinctly striated in longitudinal direction; this area contrasts to smooth shiny cuticula on condylar ridge and postero-apically adjoining surface; condylar ridge distinct at basal two-third of mandible continuing in more angled curve into apical side (strongest bend approximately after basal third of mandible). Labrum matt shining, finely shagreened and longitudinally wrinkled; densely covered with pale stiff setae directing obliquely downwards; setae as long as A7 maximum wide, with distal end curved ventro-medially, laterally at apex of labrum thicker with larger diameter at base. Clypeus 1.4 times wider than long (Fig. 5b ); translucent ventro-medial margin becoming much narrower medially resulting in distinct median emargination; cuticula shiny, ventro-medial above emargination with sparse micropunctation becoming moderately spaced dorsally and laterally on disc with larger irregular flat depressions and wrinkles in addition; dorso-lateral vertical parts of clypeus smooth with moderately spaced micropunctation partly striated at base; covered with pale thin stiff setae arising from micropunctures; setae on disc about as long as A4, vertically erected with distal ends strongly curved in ventro-medial direction, on sides shorter lying more flatly. Frons very coarsely punctured, interstices shining, raised to ± transversal little rounded ridges; protruding central part of supra-antennal area smooth without punctures in middle; semi-circular depression of antennal groove wrinkly shagreened; slight median depression dorsal to supra-antennal area, frontal line weak; sparsely covered with pale short setae arising from coarse punctures, setae on supra-antennal area as on clypeus. Vertex with close macropunctation, becoming more closely reticulate behind ocelli with smaller punctures and interstices more strongly raised forming short sharp-edged ± transversal ridges; sparsely covered with pale short setae arising from punctures; cuticula of interstices shiny, weakly shagreened (Fig. 5e ). Median ocellus 1.2 times larger in diameter than lateral ocelli; median ocellus somewhat bilateral symmetric with anterior sector less strongly curved than posterior sector; lateral ocelli ± circular (in dorso-lateral view). Compound eyes sparsely covered with tiny setae. Preoccipital carina (sensu Snelling 1986) sharp; medially straight, nearly transversal; laterally behind dorso-lateral end of each compound eye curved downwards for short distance becoming obsolete posterior to dorsal end of postocular carina (sensu Snelling 1986). Gena narrow, less than half as wide as basal width of A3. Postocular carina sharp; extends dorsad from posterior mandibular articulation along posterior margin of gena; ends level with dorsal end of compound eye anterior to preoccipital carina that runs parallel for short distance. Antennal articles A8-A12 forming ventrally flattened club about 2.1-2.3 times as long as broad (in dorsal view).

Pronotum with anterior margin raised to carina; anterior pronotal carina (sensu Carpenter 1988) in antero-ventral area of pronotum starting close to posterior margin, running in a short parabolic curve antero-laterally and then parallel to posterior margin, preceding distinct crenate groove; fairly projecting in posterior direction over anterior border of crenate groove, especially along ventral parabolic curve; crenate groove with anterior wall nearly vertical, bottom with sulcature of transverse ribs, posteriorly gradually inclined up to posterior margin of pronotum, cuticula in this area obliquely striated with some folds ± continuing into bottom ribs; on lateral quarter distinct posterior pronotal carina sharply separating semicircular antero-ventral area from dorsal area of pronotum; antero-medial front behind head nearly vertical; slight depression along dorso-medial margin especially anteriorly; posterior margin raised to short translucent carina dorsally in front of upper half of tegula; cuticula of antero-ventral area shiny, shagreened, with moderately spaced small shallow punctures; cuticula of dorso-lateral area shiny, with close coarsely reticulate macropunctation, smooth interstices raised to sharp edges postero-laterally forming lines; cuticula of pronotal lobe and dorsally continuing concavely curved depression in front of tegula smooth with a few distinct punctures but without reticulation, distinctly set off from adjacent parts of pronotum. Mesoscutum with distinct median notal suture on anterior third; cuticula shiny, coarsely reticulate with close deep macropunctation and narrow distinctly raised interstices (Fig. 6c ). Mesoscutellum with distinct transverse sulcature of longitudinal cuticula-ribs separated by intercostal spaces along antero-medial margin; laterally with distinct smooth carina along posterior margin, carina medially increasingly reduced so that cuticula of medial lobe continues evenly into crenulate margin; cuticula more coarsely reticulate than on mesoscutum. Metanotum laterally with distinct sulcature of longitudinal cuticula-ribs separated by intercostal spaces; carina along posterior margin medially with small irregular indentations continuing in vertical median keel. Axilla produced into curved tapering projection which fits into slight emargination of tegula. Tegula shiny, closely covered by macropunctures except completely smooth central convex area. Antero-ventral parts of pronotum, ventral corner of dorsal mesepisternum (= prepectus sensu Richards 1962) and ventral mesepisternum form continuous antero-ventral cavity delimited from lateral parts of mesosoma by posterior pronotal carina, carina along ventral margin of dorsal mesepisternum and epicnemial carina. Dorsal mesepisternum separated from ventral mesepisternum by weak mesepisternal groove; with distinct carina along ventral margin, which is in one line with epicnemial carina though separated from it by little notch. Ventral mesepisternum with pronounced epicnemial carina, posteriorly deflexed backwards running medially in a curve to front of mid-coxa. Mesepimeron feebly separated by weak scrobal groove; postero-ventrally bearing mesopleural process (= process at or below mesepisternal scrobe sensu Richards 1962) of moderate size, distally rounded, its posterior side shagreened matt shiny without punctures. Cuticula laterally on mesopleurum and dorsal metapleurum shiny, with closely reticulate macropunctation; longitudinally striated by raised interstices in parts; ventral mesepisternum coarsely punctured with some interstices strongly raised to knife-like edges forming coarse rugose sculpture. Propodeum with horizontal propodeal triangles and dorso-lateral margins of posterior face of propodeum reduced to two pointed protuberances dorsally on each side of middle; posteriorly with narrow medial cuticula-fold running from dorsal margin to postero-medial flange of propodeum; posterior surface ventrally striated by strong vertical cuticula-folds arising below anterior transversal carina of postero-medial flange of propodeum, weakly coriaceous, with shallow macropunctures between folds, moderately covered with fine pale setae arising from macropunctures, laterally and dorsally continuing into coarsely reticulate macropunctation with shorter setae. Cuticula below lateral lamella shiny, on metepisternum densely horizontally wrinkled, on side of propodeum shagreened with moderately spaced small shallow punctures. Lateral lamella moderate, slightly curved laterally downwards, its outer margin gently curved, its apex truncate, outer and posterior margins somewhat crenate; inner margin of lateral lamella and postero-lateral process of postero-medial flange of propodeum separated by moderate gap; anterior margin of postero-lateral process straight transverse, while posterior margin converges in a gentle curve towards lateral apex; outline of emargination being broad at its base with short moderately wide neck between lateral apex of postero-lateral process and inner margin of lateral lamella and deep medially extended apical part (Fig. 7g ); dorsal cuticula of lateral lamella and adjacent dorso-lateral part of propodeum shiny, with reticulate macropunctation. On whole exoskeleton single thin seta arises from bottom of each macropuncture, seta short if not stated otherwise.

Fore-femur postero-ventrally produced in middle forming anteriorly curved lobe (Fig. 8f ) distally changing into tapering carina along ventral margin of femur; end of tibia when folded against femur coinciding with produced region; tarsomeres I-IV broad and flattened; underside of tibia and tarsomere I with strong obliquely distally directing setae forming stiff brush; underside of tarsomere I and II with comb-like row of particularly strong setae along distal margin. Claws ventrally with small tooth.

Metasomal terga with postero-lateral corners slightly produced; posterior margin of tergum I weakly crenulated, crenulation not produced into spines and not projecting over smooth translucent lower posterior margin of tergum; posterior margin of terga II-V weakly to moderately crenulated, crenulation in middle of terga II-IV produced into little slightly raised teeth projecting approximately to end of translucent lower posterior margin of terga (Fig. 6b ); cuticula moderately shining, densely covered with reticulate macropunctation, punctures distinct, smaller and more regular than on mesoscutum; interstices finely shagreened. Tergum VI with lateral margins converging in weakly convex curve, at transition to posterior median lobe strongly bend inwards forming distinct postero-lateral angle on each side; posterior margin of posterior median lobe running in convex oval curve formed by distinct translucent lamella; posterior median lobe set off from more strongly sloping median area of tergum VI by slight concave curvature at its base; cuticula covered with fine pubescence of thin pale setae arising from micropunctures on interstices of reticulate macropunctation, slightly projecting beyond postero-median translucent lamella and lateral margins; on ventral side (viewed from ventral) posterior translucent lamella of median lobe continues on both sides into distinct carina running anteriorly along medial margin adjoining sternum VI, thereby slightly but continuously diverging from lateral margin of tergum VI.

Metasomal sternum I shiny, finely shagreened, with tiny setae but without punctures. Sterna II-V posteriorly with broad stripe of asetose, translucent cuticula adjacent to posterior margin of more strongly sclerotized cuticula; small sparse row of setae along posterior sclerotized margin somewhat projecting over anterior part of translucent stripe of cuticula; outer area of postero-lateral corners distinctly depressed, densely covered with macropunctures; rest of sclerotized cuticula shiny, finely shagreened on sterna II-III, shagreening weaker or missing on sternum IV and absent on sternum V; sternum II antero-laterally with moderately spaced shallow macropunctation and a few micropunctures becoming barely punctured towards posterior margin, whole medial area densely covered with small macropunctures from which short pale setae arise; sterna III-V anteriorly with moderate to dense shallow macropunctation, posteriorly changing into nearly unpunctured area along posterior margin. Posterior margin of sterna I-IV straight, posterior margin of sternum V medially concave running in a gentle curve. Sternum VI tapering towards distal end; with outer margin forming bulged rim, anteriorly raised to inwardly bent carina, posteriorly running in regular curve postero-medially protruded into little median spine (Fig. 8b ); cuticula with smooth median area tapering posteriorly, slightly raised to weak median keel at posterior end that continues into median spine, laterally with moderately spaced deep macropunctures becoming densely spaced and partly fused along lateral rim; stiff setae of moderate length arising obliquely backwards from macropunctures; posterior along distal end of rim densely covered with posteriorly directed stiff setae medially of same length as median spine becoming shorter anteriorly; at dorso-posterior margin dorsal (inner) cuticula weakly protruded into horizontal lamella, situated immediately above the posteriorly directed stiff setae, dorso-medially slightly raised and fused with median spine, becoming continuously smaller postero-laterally (Fig. 8g ).

Male. Colour (Fig. 2 ): Resembles female, except as follows. Yellowish-white are: large M-shaped band on frons, laterally nearly filling each ocular sinus except small area along upper inner margin of eye, narrowly interrupted medially on supra-antennal area; clypeus except dorso-lateral vertical sides and brownish translucent ventro-medial margin; complete longitudinal stripe on outside of fore-tibia; whole outside of mid- and hind-tibia; outside of hind-metatarsus. Labrum translucent brown with two yellowish-white antero-lateral spots. Tergum VI as in terga II-V with laterally and medially widened posterior band, anteriorly with small brownish tinge towards adjacent black area, interrupted on each side of middle by brownish area. Tergum VII blackish-brown posteriorly changing into brown. Antenna black, with yellowish-white stripe antero-medial on A3-A7.

Variation (number of specimens in brackets): yellowish-white M-shaped band on frons filling ocular sinus completely (1), medially not completely interrupted (1); short narrow interrupted yellowish-white streak on gena along occipital carina at dorso-lateral corner of head (2); one or two little yellowish-white spots postero-medial on mesoscutum (2); humeral plate with yellowish-white marking (4); outside of mid-metatarsus yellowish-white (1).

Structure: Resembles female, except as follows. Head in front view 1.4-1.6 times as wide as long (Fig. 9a, h ). Mandible with single pointed tooth at distal end and two smaller acute teeth distally on antero-medial margin. Labrum with flat ventro-median area, shiny with few thin short pale setae; dorsal and lateral area set off by tiny edge, strongly convex, weakly shagreened with rows of micropunctures from which short thin pale setae arise. Clypeus 1.3 times wider than long, strongly convex; shiny, ventro-medial area above emargination smooth, sparsely covered with micropunctures, dorsally and laterally changing into moderately spaced micro- and dense uneven shallow macropunctation; covered with fine pale erected setae, only very few with distally curved ends. Frons with distinct depression dorsal to protruding centre of supra-antennal area. Antennal club formed by A8-A12 about 1.8-1.9 times as long as broad (in dorsal view); asymmetrical (Fig. 9f ), with anterior margin evenly rounded, strongly curved at distal end into weakly concave distal margin that is running obliquely upwards, and posterior margin weakly convex curved into distal margin at postero-distal edge forming blunt angle; with distinct longitudinal depression on posterior half of ventral side bearing three somewhat oval shaped tyloids, situated within A9, A10 and A11, tyloid of A9 smaller than others. Mid-coxa without small spine at distal end on anterior side close to anterior-medial angle.

Tergum VII at posterior end with characteristically narrow median lobe and well set off postero-lateral angle on each side (Fig. 9d ); median lobe with translucent lamella, that continues on ventral side (in ventral view) at its base on both sides into distinct carina running anteriorly along medial margin adjoining sternum VII+VIII (fused); medial margin of postero-lateral angles running in semi-circular curve medially continuing into cuticula of median lobe slightly dorsal to base of translucent lamella; posterior median lobe and postero-lateral angles nearly horizontal distinctly set off at their base by sharp bend from anteriorly adjacent rising part of tergum; posteriorly with increasingly close and deep macropunctation, strongest medially above sharp bend; interstices anteriorly distinctly shagreened, posteriorly smooth and more shiny, postero-medially moderately covered with tiny pale setae.

Sternum VIII acutely produced running into two pointed lancet-like tips at posterior end with deep median incision between them (Fig. 14e ); convex with large longitudinal oval depression in centre, lateral margins in proximal two-third bent horizontally; cuticula shiny, postero-medially with shallow macropunctures, becoming denser towards apical end; pale postero-medially directed setae arising from macropunctures, posteriorly increasing in length, forming little tuft projecting over posterior median incision. Sparse transverse fringe of tiny setae along distal end of fused sternum VII projecting over base of sternum VIII.

Male genital as in Figs 10e , 11e , 12e , 13e . Genital comparatively narrow and elongated; in lateral view broadest at base of stipites tapering into flat distal ends of harpides, in dorsal view basal opening narrow with stipites curved towards cupula without substantial lateral enlargement. Dorsal part of stipes distally continuing into harpide, with dorsal outline of harpide nearly straight in lateral view. Harpide in ventral view with tapering spatula-like distal end with distinctly concave latero-distal margin; medial margin strongly bent in ventral direction resulting in longitudinal vertical duplication, upper margin of which curved towards longitudinal axis of harpide in addition; ventro-lateral margin continues proximally into curved sides of stipes; distally moderately covered with thin setae, short on ventral side, dorsally longer with longest setae along apical margin. Volsella continues ventro-proximally into ventral plate of stipes; medially set off from ventral plate of stipes by deep emargination of medial margin; ventrally moderately covered with strong setae that are longer apically; apically on dorsal side with strongly sclerotized large, dark tubercles. Aedoeagus with narrowly rounded distal end; thyrsoi not distinctly separated from surrounding transparent soft cuticula, though clearly stronger sclerotized laterally along basal two-third of aedoeagus, converging towards distal end; each thyrsos ventrally with distinct ventro-anteriorly directed sharp process (uncus thyrsos); apodema thyrsos robust, anteriorly curved laterad. Basal region with cupula and basal sclerite; cupula fused with base of each stipes connecting both stipites dorsally, while ventro-medial ends of cupula are separated by wide gap from each other; basal sclerite forming half ring on ventral side basal to cupula; medially slightly convex in ventral direction, laterally strongly curved upwards forming vertical sides, rounded at dorsal end.

Measurements.

Measurements of the exoskeleton are listed in Table 1 .

DNA barcoding.

COI-5P gene sequences were obtained from three specimens and entered in BOLD database (AIMEJ036-20, AIMEJ037-20, AIMEJ038-20). The intraspecific sequence divergence of C. ivanovi sp. nov. is low, reaching at most 0.18%. The clade is distinctly separated from the other investigated Celonites taxa (Fig. 16 ). The lowest interspecific genetic distance exists towards C. cagrii sp. nov. with a minimum of 6.86% (mean 7.40%).

Etymology.

The species is named after Prof. Sergey P. Ivanov, a Crimean entomologist and the scientific advisor of A. Fateryga.

Distribution.

Russia (Dagestan) (Fig. 17 ).

Bionomics.

Habitat. Imagines were observed at roadsides with richly flowering ruderal herbaceous vegetation (Fig. 18f ). The road was running along a water reservoir located at the bottom of a valley. The localities 1 and 2 were situated at 575 m and 550 m a.s.l. respectively, while the mountains surrounding them were significantly higher (up to approximately 1000 m a.s.l.). The slopes of these mountains were very dry and just sparsely covered with shrubs of Paliurus spina-christi Mill. ( Rhamnaceae ). Mean annual temperature is approximately 9.1 °C, mean precipitation 592 mm (calculated for the nearby village Untsukul by https://de.climate-data.org). At both localities, the most abundant flowering plants were Heliotropium styligerum Trautv. ( Boraginaceae ), as well as Xanthium sp. and X. spinosum L. ( Asteraceae ). In 2019 and 2021 neither H. styligerum nor the wasps were found at locality 1. The third locality was situated at 1345 m a.s.l. on a slope next to a road with very sparse vegetation. The most abundant plant species was H. styligerum followed by Vicia alpestris Steven ( Fabaceae ).

Flower association.

Adults of Celonites ivanovi sp. nov. visited exclusively flowers of Heliotropium styligerum . A total of 36 females and 1 male were recorded on flowers of this plant ("first observations"). A female visiting a flower, stood on the corolla holding on to the margins or distal parts of the petals of the same or adjacent flowers of the inflorescence with her mid- and hind-legs, while her head was situated above the corolla opening. The fore-legs were on the level surface of the corolla postero-laterally to the sides of her head. Then she rapidly protruded the proboscis thereby inserting it into the corolla tube (Fig. 18a ). The proboscis was partially pro- and retracted in a high frequency, accompanied by up and down movements of the head. Immediately thereafter, pollen was transferred from the proboscis to the fore-tarsi by moving the fore-legs characteristically parallel downwards from the lower part of the head towards the corolla opening along the simultaneously retracting proboscis, while mid- and hind-legs were still used to hold on to the flower (Fig. 18b, c ). Then she placed her fore-legs back on the surface of the petals, postero-laterally to the sides of her head, protruded her proboscis again into the corolla tube, and the whole sequence started anew (Fig. 18d ). After a few cycles of pollen removal and transfer to the fore-tarsi, she consumed the accumulated pollen from the fore-tarsi with her mouthparts. In the process the fore-legs were alternately drawn backwards through the opened mandibles from the proximal towards the distal end of the fore-tarsi (Fig. 18e ). Nectar may have been collected simultaneously with pollen, since a discrete nectar collecting behaviour was not observed. A visit to a single flower usually took just some seconds. The females walked from flower to flower of a particular inflorescence visiting several flowers one after another. Females flew between inflorescences.

Flower visits were periodically interrupted by alighting on the ground. Standing on a stone a female repeatedly regurgitated and withdrew again a mass of pollen and nectar that became visible as a droplet of liquid between her mouthparts (Fig. 18g ). This behaviour may have served to thicken the pollen and nectar mass. Occasionally, females that stood on the ground were observed to brush over their heads with their fore-legs (Fig. 18h ).

Male behaviour.

Males performed patrol flights across the area covered with H. styligerum in a low constant flight. Patrolling was regularly interrupted by perching on the ground. Courtship and copulation were not observed.

Figures

Figure 2. Imagines of the C. cyprius - group in lateral view, proboscis retracted (C. yemenensis female dbM 4944 male dbM 4945, C. osseus female dbM 4617 male dbM 5521, C. ivanovi sp. nov. female dbM 5493 male dbM 5498, C. cagrii sp. nov. female dbM 5607 male dbM 5619, C. clarus female dbM 5674).

Figure 4. Female head in frontal view a C. yemenensis (dbM 5528) b C. clarus (dbM 4616) c C. cyprius (dbM 5431) d C. osseus (dbM 5678) e C. ivanovi sp. nov. (dbM 5490) f C. cagrii sp. nov. (dbM 5609).

Figure 5. a, b Female clypeus in frontal view a C. cyprius (dbM 5431) b C. ivanovi sp. nov. (dbM 5490) c, d female mandible outside c C. cyprius (dbM 5431) d C. ivanovi sp. nov. (dbM 5490) e, f female ocelli and vertex in dorsal view e C. ivanovi sp. nov. (dbM 5492) f C. cagrii sp. nov. (dbM 5609).

Figure 6. a, b Metasomal tergum II and III of female in dorsal view a C. rugiceps (dbM 5446) b C. ivanovi sp. nov. (dbM 5481) c-f mesonotum of female in dorsal view c C. ivanovi sp. nov. (dbM 5492) d C. osseus (dbM 4617) e C. yemenensis (dbM 5528) f C. clarus (dbM 4616).

Figure 7. Right side of postero-lateral part of the propodeum of females in dorso-posterior view a C. yemenensis (dbM 4759) b C. clarus (dbM 4616) c C. cagrii sp. nov. (dbM 5611) d C. rugiceps (dbM 5446) e C. cyprius (dbM 5433) f C. osseus (dbM 5677) g C. ivanovi sp. nov. (dbM 5491) (ps = posterior surface of propodeum fp = postero-medial flange of propodeum pp = postero-lateral process ll = lateral lamella).

Figure 8. a-c Female sternum VI in ventral view a C. yemenensis (dbM 4759) b C. ivanovi sp. nov. (dbM 5491) c C. cagrii sp. nov. (dbM 5611) d-f female fore femur in posterior view d C. cyprius (dbM 5431) e C. osseus (dbM 5678) f C. ivanovi sp. nov. (dbM 5493) g-i female sternum VI posterior margin in dorso-posterior view g C. ivanovi sp. nov. (dbM 5493) h C. cagrii sp. nov. (dbM 5607) i C. cagrii sp. nov. (dbM 5618).

Figure 9. a, b Male head in frontal view a C. ivanovi sp. nov. (dbM 5498) b C. cagrii sp. nov. (dbM 5619) c-e male tergum VII in dorso-posterior view c C. osseus (dbM 5521) d C. ivanovi sp. nov. (dbM 5498) e C. cagrii sp. nov. (dbM 5619) f, g male antennal club in dorsal view f C. ivanovi sp. nov. (dbM 5499) g C. cagrii sp. nov. (dbM 5619) h, i male head in dorsal view h C. ivanovi sp. nov. (dbM 5498) i C. cagrii sp. nov. (dbM 5619).

Figure 10. Male genitalia in dorsal view a C. rugiceps (dbM 2834) b C. cyprius (dbM 5452) c C. yemenensis (dbM 5531) d C. osseus (dbM 5521) e C. ivanovi sp. nov. (dbM 5497) f C. cagrii sp. nov. (dbM 5619) (ad = aedoeagus at = apodema thyrsos bs = basal sclerite cu = cupula ha = harpide sp = stipes).

Figure 11. Microphotos of male genitalia in dorsal view a C. rugiceps (dbM 5450) b C. cyprius (dbM 5439) c C. yemenensis (dbM 5532) d C. osseus (dbM 5521) e C. ivanovi sp. nov. (dbM 5498) f C. cagrii sp. nov. (dbM 5619) (ad = aedoeagus at = apodema thyrsos bs = basal sclerite cu = cupula ha = harpide sp = stipes).

Figure 12. Male genitalia in ventral view a C. rugiceps (dbM 2834) b C. cyprius (dbM 5452) c C. yemenensis (dbM 5531) d C. osseus (dbM 5521) e C. ivanovi sp. nov. (dbM 5497) f C. cagrii sp. nov. (dbM 5619) (ad = aedoeagus bs = basal sclerite cu = cupula ha = harpide sp = stipes vo = volsella ut = uncus thyrsos).

Figure 13. Microphotos of male genitalia in ventral view a C. rugiceps (dbM 5450) b C. cyprius (dbM 5439) c C. yemenensis (dbM 5532) d C. osseus (dbM 5521) e C. ivanovi sp. nov. (dbM 5498) f C. cagrii sp. nov. (dbM 5619) (ad = aedoeagus bs = basal sclerite cu = cupula ha = harpide sp = stipes vo = volsella ut = uncus thyrsos).

Figure 14. Male fused sterna VII + VIII in ventral view a C. rugiceps (dbM 5289) b C. cyprius (dbM 5439) c C. yemenensis (dbM 5532) d C. osseus (dbM 5521) e C. ivanovi sp. nov. (dbM 5498) f C. cagrii sp. nov. (dbM 5619).

Figure 16. Neighbour joining BOLD taxon ID tree of 18 specimens of the C. cyprius - group based on COI- 5 P gene sequences (individually marked by BOLD process ID and assigned to species a priori by morphological characters; COI- 5 P gene sequence of C. osseus and C. clarus unknown).

Figure 17. Geographic distribution of members of the C. cyprius - group (full circles specimens investigated, open circles records from literature; previously published records of C. cyprius from Iran and Armenia excluded; made with Natural Earth, www. naturalearthdata. com).

Figure 18. a-e, g, h Flower visiting behaviour of Celonites ivanovi sp. nov. females at flowers of Heliotropium styligerum and behaviours associated with it (details see text) f habitat of C. ivanovi sp. nov. at locality 2 in the vicinity of Maydanskoye, Dagestan i habitat of C. cagrii sp. nov. near Aksar, Turkey (aspect in autumn).