Celonites cagrii Mauss & Yildirim, 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 cagrii Mauss & Yildirim sp. nov.

Holotype.

♀ (dbM 5610) "[Turkey] TR-Erzurum Şenkaya-Akşar [40.648262°N, 42.342351°E] 14.VII.2020-1275 m Leg. E. Yildirim [on Heliotropium ellipticum Ledeb.]" OLML, BOLD process ID CECYP006-20.

Paratypes.

"[Turkey] TR-Erzurum Şenkaya-Akşar [40.648262°N, 42.342351°E] 20.VIII.2011-1275 m Leg. E. Yildirim" 1♀ (dbM 4574) EY; "[Turkey] TR-Erzurum Şenkaya-Akşar [40.648262°N, 42.342351°E] 12.VII.2020-1275 m Leg. E. Yildirim [on Heliotropium ellipticum Ledeb.]" 3♀ (dbM 5607, 5608 (BOLD process ID CECYP005-20), 5609) VM; "[Turkey] TR-Erzurum Şenkaya-Akşar [40.648262°N, 42.342351°E] 01.VIII.2020-1275 m Leg. E. Yildirim [on Heliotropium ellipticum Ledeb.]" 1♀ (dbM 5616) AMNH, 1♂ (dbM 5619) 3♀ (dbM 5611, 5612, 5615) VM, 4♀ (dbM 5613, 5614, 5617, 5618) EY; "ARMENIA Erevan Monti desertici Aighepat 40 Km. SE 23-VII-63 [leg. Giordani Soika]" 2♂♂ (dbM 5789, 5790) MSNVE.

Diagnosis.

See key.

Description.

Female. Colour (Figs 2 , 4f ): Black. The following are light yellowish-white: rectangular spot dorso-medial on clypeus; large spot on each ocular sinus dorso-medially extending over lateral part of 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; small irregular postero-medial spot on mesoscutum; large spot on dorsal mesepisternum; laterally directed process of axilla; medium-sized narrow transversal spot postero-medial 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; 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 blackish area; two weak minute little spots medial on tergum VI; outside of distal tips of fore-, mid- and hind-femora; outside of proximal third to half of fore-, mid- and hind-tibia and little marking on outside at distal end of mid-, and hind-tibia. Brown are: distal half of mandible; maxillary and labial palpi, protrudeable parts of proboscis; ventral margin of labrum; postero-lateral margin of scutellum; median third of metanotum; postero-lateral process of propodeum; humeral plate; posterior translucent margin of tergum VI; tarsi; sterna I-V, posterior margin of sterna II-V translucent. Antenna black except: yellowish white markings medial on A4-A7 and proximal part of A8; brown-suffused area ventral on A10-A11. Wings moderately infuscate, pterostigma blackish-brown, veins blackish-brown becoming somewhat lighter at base.

Variation (number of differing specimens in brackets): Light yellowish-white markings: spot on each ocular sinus dorso-medially separated from spot on lateral part of frons (1); frons with two isolated little (4) or medium-sized (2) spots above antennal sockets, or both spots above antennal sockets large and dorso-laterally fused with large marking extending from ocular sinus over lateral part of frons (1); continuous narrow stripe on gena and postgena along occipital carina from dorso-lateral corner of head to postero-ventral corner of compound eye (1), this stripe can be shortly interrupted (1) or reduced to short narrow spot on gena and postgena along occipital carina at dorso-lateral corner of head (3) and/or little spot on postgena adjacent to occipital carina level with postero-ventral corner of compound eye (7); minute spot on malar area above condylus (7); humeral spot small (2); little spot antero-dorsally on ventral mesepisternum (2); mesoscutum completely black (1) or with rectangular spot (2) or triangular spot with top directing posteriorly (1); transversal spot postero-medial on scutellum large (2); narrow transversal stripe on median third of metanotum (10); minute spot on pointed protuberance on posterior face of propodeum dorsally on each side of middle (1); tergum I with little (1) or medium sized (1) separated longitudinal antero-medial spot that can be posteriorly fused with posterior band in median axis in addition (1); posterior band on tergum II (4) or on terga II-III (2) not completely interrupted along posterior margin on each side of middle; tergum VI with large medial spot and little spot on each postero-lateral angle (2) or only with large (5) or small (2) medial spot that can be reduced to four weak minute little dots (1); continuous stripe on outside of mid-tibia only (1) or on outside of mid- and hind-tibia (7). Other markings: light yellowish-white marking on A7 darkened (1); medial on proximal part of A8 only light brownish suffused (3) or with same colour than distal part (1).

Structure: Head in frontal view 1.50 times as wide as long in median (min 1.45, max 1.54, n=5) (Fig. 4f ). 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 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 but somewhat longitudinally striated cuticula on condylar ridge and postero-apically adjoining surface; condylar ridge distinct at basal two-third of mandible continuing in more gentle 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; translucent ventro-medial margin becoming much narrower medially resulting in distinct median emargination; cuticula shiny, ventro-medially above emargination smooth 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, with moderately to sparsely spaced macropunctures and few micropunctures; 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. 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 not shagreened (Fig. 5f ). 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 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. 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 2.0-2.2 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 weak, forming anterior sharp edge along ventral half of crenate groove dorsally continuing into sharply bent but rounded anterior border of crenate groove; crenate groove straight small trough-like, with nearly vertical anterior and posterior wall running parallel, bottom with sulcature of transverse ribs; running at very acute angle along posterior margin of pronotum slightly diverging from the posterior margin dorsally; cuticula between posterior margin of crenate groove and posterior margin of pronotum at same level as surrounding surface, shiny, with some micropunctures; 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 few small shallow punctures; cuticula of dorso-lateral area shiny, with close coarsely reticulate macropunctation, smooth interstices raised to narrow 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, ventro-laterally more distinctly set off from adjacent parts of pronotum than dorsally. Mesoscutum with distinct median notal suture on anterior third; cuticula shiny, coarsely reticulate with close deep macropunctation and narrow, distinctly raised interstices. 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 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 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, cuticula on protuberances rugose with interstices raised to knife-like edges; 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, 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 lateral apex of postero-lateral process of postero-medial flange of propodeum separated by small gap; anterior margin of postero-lateral process straight transverse, while posterior margin converges in weak curve towards lateral apex; outline of emargination being broad at its base with short narrow neck between lateral apex of postero-lateral process and inner margin of lateral lamella and small oval-rounded apical part (Fig. 7c ); 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 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; 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, nearly straight or slightly concave 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 well-definded 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; small outer area of postero-lateral corners distinctly depressed with some deep macropunctures; rest of sclerotized cuticula shiny, at least on anterior half of each sternum finely shagreened becoming weaker towards its posterior end; sternum II with median area moderately covered with small shallow macropunctures from which short pale setae arise, laterally sparsely covered with shallow macropunctures and micropunctures, micropunctation becoming fairly denser along posterior margin; sterna III-V anteriorly with moderate to dense shallow macropunctation, posteriorly changing into nearly unpunctured cuticula becoming moderately to densely covered by micropunctures further postero-laterally and along posterior margin. Posterior margin of sterna I-IV straight, posterior margin of sternum V medially concave running in a gentle curve. Sternum VI (Fig. 8c ) tapering towards distal end; with outer margin forming bulged shiny rim, anteriorly raised to inwardly bent carina, postero-laterally strongly curved medially resulting in blunter appearance of distal end of sternum VI; rim at posterior end obtuse to nearly transverse, partly interrupted by depressions of macropunctures, postero-medially protruded into little median spine; 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 protruded into irregularly serrated crystalline horizontal lamella situated immediately above the posteriorly directed stiff setae (Fig. 8h-i ), dorso-medially fused with median spine, at postero-lateral edges somewhat more protruded.

Male. Colour (Fig. 2 ): Resembles female, except as follows. Light yellowish-white are: large M-shaped band on frons, laterally filling each ocular sinus (Fig. 9b ); clypeus except dorso-lateral vertical sides and brownish translucent ventro-medial margin; little spot proximally on outside of mandible; 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 medially fused. Terga II-VI with laterally and medially widened posterior band, anteriorly with small brownish tinge towards adjacent blackish-brown area, on terga III-VI interrupted on each side of middle by blackish-brown area. Tergum VII blackish-brown. Antenna black, with light yellowish-white stripe antero-medial on A3-A7 and proximal part of A8.

Structure: Resembles female, except as follows. Head in front view 1.49 times as wide as long (Fig. 9b, i ). 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 1.36 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 short fine pale erected setae with distal end not curved. Frons with distinct depression dorsal to protruding centre of supra-antennal area. Antennal club formed by A8-A12 in dorsal view about 1.68 times as long as broad; asymmetrical (Fig. 9g ), with anterior margin evenly rounded, strongly curved at distal end into straight-line transverse distal margin, and posterior margin nearly straight, bent into distal margin at postero-distal edge forming nearly right angle; with distinct longitudinal depression on posterior two-fifth 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 narrow median lobe and well set off postero-lateral angle on each side (Fig. 9e ); median lobe moderately produced, its posterior margin weakly concave in middle, with adjacent posterior translucent lamella fairly emarginated; translucent lamella 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 (Fig. 14f ) acutely produced running into two pointed lancet-like tips at posterior end with deep median incision between them, lancet-like tips and median incision between them narrower than in C. ivanovi sp. nov.; convex with large longitudinal depression in centre, lateral margins in proximal half bent horizontally; cuticula shiny, with shallow macropunctures; 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 sternum VII projecting over base of fused sternum VIII.

Male genital as in Figs 10f , 11f , 12f , 13f . 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 slightly curved towards longitudinal axis of harpide in addition; ventro-lateral margin continues proximally into curved sides of stipes; distally moderately covered with thin setae, 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 broadly rounded distal end; thyrsoi not distinctly separated from surrounding transparent soft cuticula, though clearly stronger sclerotized laterally along basal two-third of aedoeagus, only weakly converging towards distal end; each thyrsos ventrally with distinct ventro-anteriorly directed comparatively broad and blunt process (uncus thyrsos); apodema thyrsos delicate, anteriorly running nearly straight. 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 two specimens and entered in BOLD database (CECYP005-20, CECYP006-20). The intraspecific sequence divergence of C. cagrii sp. nov. is 0.37%. The clade is clearly separated from the other investigated Celonites taxa (Fig. 16 ). The lowest interspecific genetic distance exists between C. cagrii sp. nov. and C. ivanovi sp. nov. with a minimum of 6.86% (mean 7.40%).

Etymology.

The species is named after M. Çağrı Yildirim, the son of E. Yildirim.

Distribution.

Turkey, Armenia (Fig. 17 ).

Bionomics.

Habitat. The locus typicus of C. cagrii sp. nov. is situated in an arid mountainous area at an altitude of 1275 m a.s.l. in a valley near Akşar, which is a village in the Şenkaya district of Erzurum. Mean annual temperature is approximately 7.2 °C, annual precipitation is 456 mm (calculated by https://de.climate-data.org). The sides of the valley are formed by the steep slopes of adjacent dry mountains and a stream is running at its bottom. The upper part of the valley is covered with rocks, while the lower part is characterized by stony ruderal sites mainly used for grazing, and a few dry pastures and fields (Fig. 18i ). General plant diversity is low, but plants of Heliotropium ellipticum Ledeb. ( Boraginaceae ) were growing solitarily or in patches on the sides of a little dirt road in an area that extended over 300 m, where the imagines of C. cagrii sp. nov. were observed.

Flower association.

Twelve females and a single male of C. cagrii sp. nov. were collected while they were visiting flowers of Heliotropium ellipticum .

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 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).