Research Article
Print
Research Article
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
expand article infoVolker Mauss, Alexander V. Fateryga§, Erol Yildirim|, James M. Carpenter
‡ Stuttgart State Museum of Natural History, Entomology Department, Stuttgart, Germany
§ T.I. Vyazemsky Karadag Scientific Station, Nature Reserve of RAS, Feodosiya, Russia
| Atatürk University, Erzurum, Turkey
¶ Division of Invertebrate, Zoology, American Museum of Natural History, New York, United States of America
Open Access

Abstract

Celonites ivanovi sp. nov. is described as a new species from Dagestan where it has been recorded from dry habitats in a small area on the northern side of the Greater Caucasus. Celonites cagrii sp. nov. is described from Erzurum Province in east Turkey. As in other members of the C. cyprius-group, the females of both species were observed to visit flowers of Heliotropium (Boraginaceae). A morphological examination including the male genitalia of all species of the C. cyprius-group revealed that C. ivanovi sp. nov. and C. cagrii sp. nov. share the apomorphic characters of this group and are closely related to Celonites osseus Morawitz, 1888. Mean genetic distance between C. ivanovi sp. nov. and C. cagrii sp. nov. based on COI-5 sequences is 7.40%. The geographical distribution of all members of the C. cyprius-group is summarized and an illustrated key is provided for the identification of males and females of the species. A lectotype is designated for C. osseus.

Keywords

Boraginaceae, distribution, flower visiting behaviour, Heliotropium, key to species, male genitalia, Palaearctic region, taxonomy, trophic relationships

Introduction

The pollen wasp genus Celonites Latreille, 1802 forms a well defined monophylum (Carpenter 1993; Krenn et al. 2002) whose members share a number of distinctive characters such as the wide horizontal lamellae on the propodeum, the acute sides of the metasoma and the ability to roll up (Richards 1962; Gess and Gess 2010). The biogeographical distribution of Celonites is disjunct with 21 species occurring in the semi-arid to arid areas in the southwest of the Afrotropical region (Gess and Gess 2010) and approximately 42 species in the Palaearctic and adjacent dry areas in the north of the Afrotropical and Oriental region (according to the list of Carpenter 2001, combined with newly described or synonymized species by Gusenleitner 2002, 2007, 2012, 2018; Mauss 2013; Mauss et al. 2016; Mauss and Prosi 2018). Among the Palaearctic taxa of Celonites the subgenus Eucelonites Richards, 1962 is characterised by a small laterally directed process of the axilla which lies on the tegula or even fits into a slight emargination of the tegula opposite to the projection (Richards 1962). This unique apomorphy defines Eucelonites as a monophyletic group (Carpenter 2001; note that the nominate subgenus is evidently paraphyletic). Up till now 26 species and nine additional subspecies were assigned to this monophylum indicating high diversity. However, many of these taxa were described from small type series often containing only one sex and the last summarizing key provided by Richards (1962) left out several rarely collected species from the Middle East and Central Asia. Therefore, the identity and status of some described Eucelonites taxa is still uncertain and species identification can be difficult. Within Eucelonites, Celonites cyprius Saussure, 1854 and its closer relatives form a distinct group which is characterized by male genitalia with distally tapering more or less spatula-like harpides, the medial and lateral margin of which are partly raised ventrally in addition (Figs 12, 13). We suggest naming this group the Celonites cyprius-group.

The purpose of the present paper is to describe two new species of Celonites belonging to the C. cyprius-group from the Republic of Dagestan, Russia, and Erzurum Province in East Anatolia, Turkey, as well as to summarize taxonomic and chorological data on the related taxa and to provide a key for their identification.

Materials and methods

Field investigations of the hitherto unknown Celonites ivanovi sp. nov. were made by A. Fateryga on 23 June 2018 in the vicinity of Maydanskoye in the Untsukulskiy district of the Republic of Dagestan, Russia [locality 1: 42°37'36"N, 46°56'48"E], where 3 females were observed and 2 females were collected, and on 11 June 2019, 15 and 16 June 2021 at a second site in the vicinity of Maydanskoye, 3 km to the southeast of the first locality [locality 2: 42°36'07"N, 46°58'13"E], where numerous females and males were observed and 17 females and 10 males were collected. Wasp activity and flower visiting behaviour were observed with the naked eye for approximately one hour in the morning of 11 June 2019 and documented with a Canon PowerShot SX160 IS camera. The pictures were compared to single frames of video sequences of C. cyprius and C. rugiceps Bischoff, 1928 visiting Heliotropium flowers in Rhodes (taken with a Canon EOS 80D, 50 frames per second, scale up to 1:1, Mauss et al. in prep.). Flower preferences of the wasps were studied by counting the number of sightings (= first observations) of flower visiting individuals while walking randomly across the locality. A single female was recorded on 22 June 2021 in the vicinity of Turtsi in the Lakskiy district [locality 3: 42°11'34"N, 47°09'33"E]. Field observations of Celonites cagrii sp. nov. were made by E. Yildirim at Şenkaya-Akşar in the Erzurum province of Turkey on 12 July, 14 July and 1 August 2020. For identification of the collected Heliotropium plants the keys provided by Grossheim (1967) and Dönmez (2008) were used.

For the morphological and chorological investigations in total 479 dry specimens of the Celonites cyprius-group were examined from the following collections: Museum of Zoology Lausanne, Switzerland (MCZL), Museum of Natural History Mainz, Germany (NHM), Natural History Museum of Venice, Italy (MSNVE), Upper Austrian State Museum – Biology Centre, Linz, Austria (OLML), State Museum of Natural History Karlsruhe, Germany (SMNK), Stuttgart State Museum of Natural History, Germany (SMNS), Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia (ZISP), Zoological Museum of the M.V. Lomonosov Moscow State University, Russia (ZMMU) , as well as the private collections of A.V. Fateryga stored in the T.I. Vyazemsky Karadag Scientific Station – Nature Reserve of RAS – Branch of A.O. Kovalevsky Institute of Biology of the Southern Seas of RAS, Feodosia, Russia (AF), J. Gusenleitner, Linz, Austria (JG), V. Mauss, Michelfeld, Germany (VM), M. Schindler, Bonn, Germany (MS), C. Schmid-Egger, Berlin, Germany (CSE) and E. Yildirim, Erzurum, Turkey (EY) (Suppl. material 1: Table S1). Every specimen examined by V. Mauss was labelled with an individual, serial database number (dbM = database Mauss) printed on the determination label. The holotypes were deposited in OLML, paratypes in American Museum of Natural History (AMNH), OLML, ZISP, AF, VM and EY.

Specimens were investigated under a WILD M3 stereo microscope (maximum magnification 80 times). Numbers of terga and sterna apply to metasomal segments. Antennal articles are termed A1–A12 counted from the proximal to the distal end. Nomenclature of male genitalia follows that of Birket-Smith (1981). Measurements of the exoskeleton were made using an ocular micrometer (highest resolution 0.011 mm; microscope in monaxial position). Distances between the ocelli, the compound eyes and the mesonotum width were measured according to Eck (1978). Proportion of head width to head length in the key was calculated from measurements of the maximum distance between the lateral margins of the compound eyes and the distance in the median axis between the bottom of the ventral emargination of the clypeus and the dorsal end of the median ocellus, while the head was aligned with dorsal and median part of ventral margin of the clypeus in focus plane and the lateral margins of the compound eyes in symmetrical distance to the median axis. Drawings were made with a drawing tube (WILD Type 308700) in monaxial position. For drawings of the postero-lateral part of the propodeum the metasoma was removed from the specimens. The postero-lateral part of the propodeum was viewed from dorso-posterior with the dorso-anterior border of the pronotum becoming just visible anteriorly, so that the dorsal area of the pronotum was visible at a flat angle, while the apical ends of the postero-lateral processes were orientated exactly transversal. Male genitalia were removed after resoftening of the specimens and studied in 80% ethanol or glycerine. Macrophotos of whole specimens were taken with a Canon EOS 80D camera with a 100 mm macro-lens and extension tube (scale more than 1:1, resolution 24 mega pixel) and macro flash-lights. Microphotos were taken with a Keyence VHX-5000 Digital Microscope (combined stacking and stitching). Photographs of the lectotype of C. osseus Morawitz, 1888 were taken in ZISP with a Canon EOS 70D camera mounted on an Olympus SZX10 stereomicroscope. Multifocus-pictures were generated with Helicon Focus 6 Pro software. Final illustrations were post-processed for sharpness, contrast, and brightness using Adobe Photoshop CS2 software.

DNA barcoding was accomplished by AIM Advanced Identification Methods GmbH Leipzig following standard methods of DNA extraction from a single leg of dry specimens or specimens collected and stored in 96% pure ethanol, PCR for Cytochrome Oxidase subunit 1 (COI-5P), cycle sequencing of forward and reverse strand and sequence editing. Nucleotide sequences of ten individuals from four taxa of the C. cyprius-group were obtained uploaded and compared to the BOLD database (www.BOLDsystem.org). Another eight COI-5P sequences of three taxa of the C. cyprius-group were obtained from the BCHYM- and GBACU-project by courtesy of Christian Schmid-Egger and Stefan Schmidt and added to the data set. Finally genetic distances (Kimura 2 Parameter) and a neighbour joining BOLD TaxonID Tree were computed including 18 COI-5P sequences of morphologically identified specimens of C. cagrii sp. nov., C. cyprius, C. ivanovi sp. nov., C. rugiceps and C. yemenensis Giordani Soika, 1957 using the following parameters: distance model Kimura 2 Parameter; pairwise deletion of positions containing gaps and missing data; minimum complete overlap 100 bp; alignment with BOLD Aligner (Amino Acid based HMM); individual nucleotide sequence length 557–699 bp.

For the investigation of the geographic distribution label information of 479 specimens of the C. cyprius-group coming from 114 localities was entered into the database Mauss (Suppl. material 1: Table S1). Additionally, 55 published records were taken from Bischoff (1928), Blüthgen (1952), Bytinski-Salz and Gusenleitner (1971), Carpenter (2001), Ebrahimi and Carpenter (2008), Giordani Soika (1957), Gusenleitner (1966, 1973, 1997, 2005, 2010), Richards (1962, 1984) and Schmid-Egger (2017) (Suppl. material 1: Table S1). If not given on the label or in the publication, WGS 84 coordinates were reconstructed mainly with the aid of Google Maps (www.google.de/maps). The distribution map was created with Natural Earth (www.naturalearthdata.com) using the open-source geographic information system QGIS, version 3.4 (www.qgis.org).

Systematics

Celonites osseus Morawitz, 1888

Celonites osseus Morawitz, 1888: 268–269, ♀, type locality: “territorio transcaspico (Tschikischljar) [Turkmenistan]”, lectotype (designated here): ♀, <golden disc>, “Tschikischljar. Pom.[eranzev]”, “Celonites osseus ♀. F. Moraw.”, “к. Ф Моравица [coll. F Morawitz]”, “Lectotypus ♀ Celonites osseus Morawitz des. Fateryga, 2018 <red label>” ZISP (Fig. 3a–c); Kostylev, 1935: 114, Turkestan, Transcaspian sands, Armenia; Richards, 1962: 216 (key), 226–227, Turkmenistan (Repetek), misidentification of Celonites crenulatus Morawitz, 1888 not C. osseus; Carpenter, 2001: 11, Armenia, Turkmenistan.

Additional material studied

Armenia: Zanga [currently Hrazdan] River, near Yerevan, 19.07.1935 1♀ (dbM 5855) leg. G. Kostylev ZMMU; Yerevan, Nork [=Nor Nork], 23.06.1930 1♀ leg. A. Shelkovnikov ZISP. Turkmenistan, Balkan region: Krasnovodsk [currently Türkmenbaşy], 13.06.1928 1♀ leg. V. Gussakovskij ZISP, 16.06.1928 1♂ leg. V. Gussakovskij ZISP; Jebel, 25 km NW Balkanabat, 22.05.1993 1♂ (dbM 5521) leg. M. Halada OLML. Iran: Elburz, 30 km S of Ab Ali, 09.07.1965 1♀ (dbM 5788) leg. Giordani Soika & Mavromoustakis MSNVE; Elburz, Vana 50 km N di Ab Ali, 12–13.07.1965 10♀♀ (dbM 5676, 5677, 5678, 5780, 5781, 5782, 5783, 5784, 5785, 5786) leg. Giordani Soika & Mavromoustakis MSNVE; Elburz (Mazandaran), Pulour [=Polour] 22 km N of Ab Ali, 13–14.07.1965 1♀ (dbM 5787) leg. Giordani Soika & Mavromoustakis MSNVE; Fars, Daria Namak, steppe presso lago salato 27 km E Shiraz, 07.07.1965 3♀♀ (dbM 5679, 5778, 5779) leg. Giordani Soika & Mavromoustakis MSNVE; Fars prov., 20 km E Kazarun, 29°33.034'N, 51°49.416'E, 1256 m, 14.06.2019 1♂ leg. V.M. Gnezdilov ZISP; Golestan prov., 70 km E Minudasht, 1050 m, 12.06.2010 1♀ (dbM 4617) leg. M. Halada OLML; Teheran, Steppa presso Chitgar, 18 km W di Teheran, 15.07.1965, 1♀ (dbM 5675) leg. Giordani Soika & Mavromoustakis MSNVE.

Taxonomic remarks

The identity of C. osseus was probably not clear to O.W. Richards, since in his key and in his description of the species he emphasizes that the dorsal area of the mesoscutum has smooth shining interstices at least as wide as the punctures, which is clearly inconsistent with the original description of Morawitz (1888: 269), and that the metasomal terga II–V are not at all crenulated (Richards 1962: 216, 227). Moreover, the measurements given by him for the length of body and fore-wing seem to be too large to represent C. osseus. His description was based on a single female from Repetek collected on 7 June 1937 by G. Kostylev placed in ZMMU and determined by Kostylev as C. osseus (Richards 1962: 227). However, this particular specimen is lacking in the ZMMU, but there is another female (dbM 5854) from Repetek collected on 9 June 1937 labelled by Kostylev as C. osseus, which is clearly misidentified and belongs to Celonites crenulatus Morawitz, 1888 or an undescribed taxon close to it. The description of C. osseus given by Richards (1962: 226–227) is almost identical with the characters of this misidentified specimen. Moreover, there is a series of seven correctly identified males of C. crenulatus in the collection of the ZMMU collected on 1 and 9 June 1937 at Repetek by G. Kostylev, while genuine specimens of C. osseus are lacking from this locality. Obviously, Kostylev misinterpreted two females of his C. crenulatus series with weak crenulation on the metasomal terga as C. osseus. Almost certainly, Richards investigated one of these wrongly identified specimens (maybe it was given to him, because it was the only duplicate specimen identified as C. osseus in the collection of the ZMMU) and finally based his description on it. As a consequence, the key and the description provided by Richards (1962) cannot be used for the identification of C. osseus.

Distribution

Armenia, Turkmenistan, Iran (new record) (Fig. 17).

Bionomics

Unknown.

Figure 1. 

Imagines of the C. cyprius-group in lateral view, proboscis protruded (C. rugiceps female dbM 5447 male dbM 5450, C. cyprius female dbM 5428 male dbM 5434).

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).

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 3. 

Lectotype of Celonites osseus a lateral view b head in frontal view c labels.

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).

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).

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.

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).

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.

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).

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.

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).

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.

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).

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.

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).

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.

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).

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.

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.

Table 1.

Measurements of the exoskeleton of females and males of Celonites cagrii sp. nov., C. ivanovi sp. nov., C. osseus and C. clarus (x = median; min = minimum, max = maximum, n = sample size; maximum accuracy 0.011 mm, all distances in mm).

Genus female male
Species C. cagrii sp. nov. C. ivanovi sp. nov. C. osseus C. clarus C. cagrii sp. nov. C. ivanovi sp. nov. C. osseus
Parameter x min max n x min max n x min max n x min max n x min max n x min max n x n
lateral ocelli distance 0.39 0.36 0.42 13 0.41 0.37 0.43 11 0.39 0.36 0.43 10 0.41 0.36 0.42 3 0.34 0.34 0.35 3 0.39 0.32 0.43 5 0.39 1
med./lat. ocellus distance 0.13 0.12 0.14 13 0.14 0.13 0.17 11 0.14 0.12 0.15 10 0.14 0.12 0.14 3 0.11 0.11 0.12 3 0.13 0.10 0.14 5 0.13 1
compound eyes distance 1.17 1.11 1.22 13 1.21 1.16 1.24 11 1.17 1.12 1.24 10 1.21 1.16 1.22 3 0.90 0.86 0.94 3 0.86 0.72 1.09 5 1.09 1
A1 length 0.17 0.15 0.18 13 0.19 0.18 0.20 11 0.17 0.15 0.18 10 0.15 0.14 0.17 3 0.17 0.15 0.19 3 0.19 0.17 0.20 5 0.17 1
A3 length 0.23 0.22 0.24 13 0.24 0.23 0.25 11 0.23 0.21 0.24 10 0.18 0.18 0.19 3 0.23 0.23 0.26 3 0.24 0.23 0.26 5 0.22 1
A3 width 0.09 0.08 0.10 13 0.10 0.09 0.11 11 0.09 0.09 0.10 10 0.09 0.09 0.10 3 0.10 0.10 0.10 3 0.11 0.09 0.11 5 0.11 1
A4–A5 length 0.17 0.15 0.18 13 0.18 0.15 0.18 11 0.17 0.15 0.18 10 0.13 0.12 0.14 3 0.20 0.19 0.22 3 0.20 0.18 0.21 5 0.21 1
A8–A12 length 0.65 0.63 0.67 13 0.70 0.66 0.74 11 0.69 0.65 0.72 10 0.64 0.62 0.70 3 0.85 0.81 0.88 3 0.85 0.78 0.95 5 0.99 1
A8–A12 width 0.31 0.29 0.32 13 0.32 0.31 0.34 11 0.32 0.32 0.34 10 0.32 0.31 0.34 3 0.48 0.47 0.48 3 0.44 0.42 0.51 5 0.45 1
antennal sockets distance 0.56 0.54 0.59 13 0.58 0.54 0.61 11 0.58 0.54 0.61 10 0.57 0.54 0.63 3 0.41 0.40 0.42 3 0.42 0.40 0.48 5 0.47 1
clypeus max. width 0.87 0.85 0.90 13 0.94 0.87 0.97 11 0.90 0.88 0.95 10 0.88 0.86 0.92 3 0.70 0.68 0.73 3 0.70 0.67 0.84 5 0.77 1
clypeus apical width 0.46 0.43 0.51 13 0.54 0.44 0.56 11 0.46 0.40 0.48 10 0.41 0.31 0.44 3 0.34 0.33 0.37 3 0.45 0.33 0.50 5 0.39 1
clypeus length 0.62 0.59 0.65 13 0.68 0.65 0.70 11 0.65 0.62 0.69 10 0.58 0.56 0.61 3 0.53 0.52 0.55 3 0.56 0.50 0.64 5 0.58 1
mesonotum width 2.10 1.99 2.18 13 2.30 2.13 2.38 11 2.21 2.10 2.27 10 2.04 1.93 2.13 3 1.90 1.88 2.02 3 2.24 1.85 3.77 5 2.18 1
mesoscutum length 1.31 1.24 1.38 13 1.48 1.39 1.53 11 1.42 1.29 1.50 10 1.33 1.29 1.39 3 1.12 1.03 1.19 3 1.12 1.00 1.34 5 1.24 1
wing length 4.14 4.00 4.32 13 4.69 4.46 4.90 11 4.53 4.23 4.65 10 4.09 4.00 4.32 3 3.96 3.77 4.09 3 4.14 3.77 4.62 5 4.37 1
R+Sc length 2.21 2.18 2.32 13 2.49 2.41 2.58 11 2.44 2.24 2.55 10 2.16 2.13 2.35 3 2.13 2.04 2.21 3 2.16 2.04 2.60 5 2.32 1
number of hamuli 7 6 8 13 7 6 8 11 7 7 9 10 7 7 9 3 8 7 9 3 7 7 8 5 8 1
femur I length 1.01 0.97 1.06 13 1.11 1.08 1.14 10 1.04 0.94 1.10 10 0.97 0.95 0.99 3 0.90 0.84 0.95 3 1.00 0.87 1.07 4 0.95 1
tibia I length 0.67 0.62 0.69 13 0.76 0.74 0.77 11 0.69 0.66 0.74 10 0.64 0.63 0.65 3 0.62 0.62 0.65 3 0.67 0.62 0.75 5 0.64 1
tibia I width 0.36 0.34 0.40 13 0.42 0.40 0.45 11 0.20 0.19 0.20 10 0.36 0.36 0.40 3 0.17 0.17 0.17 3 0.33 0.29 0.36 5 0.34 1
metatarsus I length 0.45 0.40 0.48 13 0.51 0.46 0.54 11 0.42 0.39 0.43 10 0.45 0.43 0.51 3 0.28 0.28 0.29 3 0.45 0.42 0.48 5 0.51 1
metatarsus I width 0.19 0.18 0.20 13 0.20 0.19 0.21 11 0.13 0.13 0.14 10 0.18 0.18 0.20 3 0.10 0.10 0.10 3 0.17 0.14 0.19 5 0.19 1
tarsus I T2–T5 length 0.14 0.13 0.14 13 0.15 0.13 0.15 11 0.50 0.47 0.54 10 0.12 0.12 0.13 3 0.42 0.40 0.45 3 0.11 0.10 0.12 5 0.11 1
tergum I width 2.21 2.10 2.30 13 2.41 2.30 2.49 11 2.30 2.13 2.46 10 2.10 2.07 2.27 3 2.07 2.04 2.27 3 2.27 1.90 2.52 5 2.30 1
tergum I length 0.88 0.73 0.94 13 1.02 0.95 1.04 11 0.94 0.87 1.04 10 0.87 0.78 0.90 3 0.88 0.80 0.96 3 0.97 0.80 1.12 5 1.00 1
tergum II width 2.18 2.07 2.24 13 2.41 2.27 2.52 11 2.32 2.21 2.46 10 2.07 2.07 2.30 3 2.07 1.99 2.18 3 2.24 1.96 2.52 5 2.35 1
total length 5.7 5.3 6.6 13 6.0 5.6 6.5 11 6.1 5.4 6.2 10 5.4 5.3 6.5 3 5.4 5.3 5.4 2 6.2 5.0 7.0 4 6.5 1

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).

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).

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.

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).

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.

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).

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.

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).

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.

Figure 15. 

Female antennal articles A1–A6 in medial view (scaled to same size to show different proportions of A3) a C. clarus (dbM 5674) b C. osseus (dbM 5678).

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.

Figure 16. 

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

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).

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).

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.

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 Akşar, Turkey (aspect in autumn).

Key to the species of the Celonites cyprius-group

Male of Celonites clarus Gusenleitner, 1973 not known.

1 Female 2
Male 8
2 Larger, body length approximately 7–8 mm, more sturdily built (Fig. 1). Crenulation of terga II–IV considerably stronger (Fig. 6a). Emargination between lateral lamella and postero-lateral process of propodeum in outline comparatively small, its apical end nearly circular (Fig. 7d). Gena black. Celonites rugiceps Bischoff, 1928
Smaller, body length approximately 5–6.5 mm (Figs 1, 2). Crenulation of terga II–IV considerably weaker, sometimes almost obsolete (Fig. 6b). Emargination between lateral lamella and postero-lateral process of propodeum in outline comparatively larger, its apical end medially extended (Fig. 7a, b, e–g; except C. cagrii sp. nov. Fig. 7c). Gena black or with yellow marking behind compound eye 3
3 Postero-lateral process of propodeum with blunt apical end, outline of emargination between lateral lamella and postero-lateral process narrowly elongated (Fig. 7a). Posterior margin of sternum VI nearly transverse, with sharply bended postero-lateral corners (Fig. 8a). Mesoscutum strongly reticulate with sharply raised interstices, comb-like (Fig. 6e). Head very broad, in median 1.56 times as wide as long (1.52–1.58) (Fig. 4a) Celonites yemenensis Giordani Soika, 1957
Postero-lateral process of propodeum and outline of emargination between lateral lamella and postero-lateral process different (Fig. 7b, c, e–g). Posterior margin of sternum VI more continuously curved, without distinct postero-lateral corners (Fig. 8b; less so in C. cagrii sp. nov. Fig. 8c). Mesoscutum reticulate, with interstices often less sharply raised and more bluntly rounded partly forming lines (Fig. 6c, d) or with dense macropunctation with smooth shiny interstices (Fig. 6f). Head in some species less broad (Fig. 4c) 4
4 Head and especially clypeus appear less broad in frontal view, head in median 1.43 times as wide as long (1.38–1.48) (Fig. 4c). Clypeus on disk with smooth, shiny cuticula, moderately covered with fine punctures from which setae arise (Fig. 5a). Mandible on outside at base with distinct transverse depression with dull, shagreened cuticula, densely covered with pubescence of very tiny thin setae, well set off from adjacent apical smooth shiny area with few macropunctures (Fig. 5c). Fore-femur postero-ventrally only with weak carina, not produced into distinct anteriorly curved lobe in middle (Fig. 8d). Gena with extended yellow stripe behind compound eye Celonites cyprius Saussure, 1854
Head and especially clypeus appear distinctly broader in frontal view, head at least 1.45 times as wide as long (Fig. 4b, d–f). Clypeus on disk with some coarser depressions i.e. dorso-laterally, where punctures are in rows and interstices are slightly raised forming wrinkled lines; setae arise from larger punctures (Fig. 5b). Mandible on outside at base without distinct transverse depression, basal area with shagreened cuticula extending further apically, dorsal to condylar ridge becoming distinctly longitudinally striated, basally moderately covered with pubescence of tiny thin setae (longer than in C. cyprius); this area contrasts to smooth shiny cuticula on condylar ridge and ventro-apical outside (Fig. 5d). Fore-femur postero-ventrally produced in middle into distinct anteriorly curved lobe that distally changes into tapering carina along ventral margin of femur (Fig. 8e, f). Gena black or with yellow marking behind compound eye. 5
5 Mesoscutum with dense macropunctation with smooth shiny interstices (Fig. 6f). A3 shorter, in median 2.1 times as long as wide (Fig. 15a). Postero-lateral process of propodeum with blunter apical end, outline of emargination between lateral lamella and postero-lateral process with narrow elongated neck (Fig. 7b). Crenulation along posterior margin of metasomal terga weak and blunt, not exceeding the lower hyaline rim of the tergum (Fig. 2). Gena with yellow marking in middle behind outside bend of compound eye and sometimes additional yellow spot close to mandible. Head very broad, in median 1.56 times as wide as long (1.55–1.56) (Fig. 4b) Celonites clarus Gusenleitner, 1973
Mesoscutum reticulate, with distinctly raised narrow interstices partly forming lines (Fig. 6c, d). A3 longer, in median 2.5 times as long as wide (Fig. 15b). Postero-lateral process of propodeum with converging end (Fig. 7e–g). Gena black or with yellow marking 6
6 Outline of emargination between lateral lamella and postero-lateral process of propodeum with narrow short neck, its apical end small oval-shaped (Fig. 7c). Anterior pronotal carina short, forming anterior sharp edge along ventral half of crenate groove. Crenate groove straight, trough-like, its nearly vertical posterior wall separated from posterior margin of pronotum by a stripe of cuticula similarly covered with micropunctures and at same level as surface of surrounding antero-ventral area of pronotum. Bulged shiny rim of sternum VI postero-laterally strongly curved in medial direction, running obtuse to nearly transverse at posterior end resulting in blunter appearance of posterior end of sternum VI (Fig. 8c). Posterior section of bulged rim of sternum VI partly interrupted by large depressions of macropunctures (Fig. 8c). Dorso-posterior margin of sternum VI with distinct irregularly serrated crystalline horizontal lamella situated immediately above posteriorly directed stiff setae, dorso-medially fused with median spine and somewhat more protruded at postero-lateral edges (Fig. 8h, i). Light markings light yellowish-white. Head broad, in median 1.50 as wide as long (1.45–1.54) (Fig. 4f) Celonites cagrii Mauss & Yildirim, sp. nov.
Outline of emargination between lateral lamella and postero-lateral process of propodeum with broad short neck, its apical end medially enlarged (Fig. 7f, g). Anterior pronotal carina starts close to posterior margin of pronotum and runs in short parabolic curve antero-laterally and then parallel to posterior margin of pronotum forming anterior margin of crenate groove. Anterior pronotal carina fairly projecting in posterior direction over anterior border of groove, especially along its ventral parabolic curve. Crenate groove posteriorly open, posteriorly adjacent cuticula gradually rising to posterior margin of pronotum. Cuticula in this lowered area differs from surrounding surface of antero-ventral area of pronotum being obliquely striated with some folds that are ±continuing into bottom ribs of crenate groove. Bulged shiny rim of sternum VI forms regular curve postero-laterally resulting in more pointed appearance of posterior end of sternum VI (Fig. 8b). Posterior section of bulged rim of sternum VI not or barely interrupted by large depressions of macropunctures (Fig. 8b). Dorso-posterior margin of sternum VI at most with simple weak translucent cuticula crest (Fig. 8g). Light markings yellowish-white or yellow 7
7 Head in frontal view more triangular, outline of compound eye less strongly curved (Fig. 4e), in median 1.48 as wide as long (1.45–1.53). Light colouration yellowish-white, less extended, normally mesoscutum black (Fig. 6c) and outside of mid- and hind-tibia with dark ring-like markings. Dark colour of head and mesosoma deep black (Fig. 2) Celonites ivanovi Mauss & Fateryga, sp. nov.
Head in frontal view more elliptic, outline of compound eye nearly semi-circular (Fig. 4d), in median 1.57 as wide as long (1.50–1.62). Light colouration more yellowish to yellow, more extended, with yellow spot on mesoscutum (Fig. 6d) and outside of mid- and hind-tibia completely yellow. Dark colour of head and mesosoma black, partly with more or less brownish to reddish shade (Fig. 2) Celonites osseus Morawitz, 1888
8 Larger, body length approximately 7–8 mm, more sturdily built (Fig. 1). Sternum VIII semicircularly produced, not emarginated (Fig. 14a). Male genital as in Figs 10a, 11a. Harpide ventrally at base with unique ventro-medially extending sclerite (Figs 12a, 13a) Celonites rugiceps Bischoff, 1928
Smaller, body length approximately 5–6.5 mm (Figs 1, 2). Sternum VIII acutely produced, running into two tapering lobes at posterior end with deep median incision between them (Fig. 14b–f). Male genital different (Figs 10b–f, 11b–f). Harpide ventrally at base without ventro-medially extending sclerite (Figs 12b–f, 13b–f) 9
9 Posterior lobes of sternum VIII of medium length, wedge-shaped with acute-angled end (Fig. 14b). Male genital along posterior end of ventral plate of stipes with dense band of conspicuously long strong setae projecting well above volsella (Figs 12b, 13b). Ventral side of harpide densely covered with long setae (Figs 12b, 13b) Celonites cyprius Saussure, 1854
Posterior lobes of sternum VIII different (Figs 14c–f). Male genital on ventral plate of stipes at most with a sparse row of short setae (Figs 12c–f, 13c–f). Ventral side of harpide less densely covered with shorter setae (Figs 12c–f, 13c–f) 10
10 Posterior lobes of sternum VIII short with blunt ending (Fig. 14c). Male genital as in Figs 10c, 11c, 12c, 13c. Lateral margin of harpide strongly and continuously concave along most of its length Celonites yemenensis Giordani Soika, 1957
Posterior lobes of sternum VIII long, lanceolate with pointed tip (Fig. 14d–f). Male genital as in Figs 10d–f, 11d–f, 12d–f, 13d–f. Lateral margin of harpide with concavity at most along posterior half of its length 11
11 Male genital broader at base (Figs 10d, 11d), cupula wider, less curved (in ventral view; Figs 12d, 13d). Lateral margin of harpide without concavity near apical end, due to broad lamella along postero-lateral margin of harpide (Figs 12d, 13d). Median lobe of tergum VII broader at base between postero-lateral angles (Fig. 9c) Celonites osseus Morawitz, 1888
Male genital narrower at base (Figs 10e, f, 11e, f), cupula narrower, more strongly curved (in ventral view; Figs 12e, f, 13e, f). Lateral margin of harpide with shallow concavity near apical end. Median lobe of tergum VII narrower at base between postero-lateral angles (Fig. 10 d, e) 12
12 Head in frontal view more triangular (Fig. 9a). Antennal club (A8–12) in dorsal view about 1.8–1.9 times as long as broad, asymmetrical, with evenly rounded anterior margin and weakly concave distal margin that is running obliquely upwards. At postero-distal edge distal margin curved into posterior margin forming a blunt angle (Fig. 9f); posterior margin weakly convex. Median lobe of tergum VII markedly produced, its posterior margin and adjacent posterior translucent lamella entirely convex (Fig. 9d). Sternum VIII with longitudinal depression in centre largely oval, lancet-like tips and postero-median incision between them broader (Fig. 14e). Aedoeagus with narrowly rounded distal end, its lateral sides noticeably converging. Uncus thyrsos sharp (Fig. 12e). Apodema thyrsos robust, anteriorly curved laterad (Figs 10e, 11e). Outline of emargination between lateral lamella and postero-lateral process of propodeum with broad short neck, its apical end medially enlarged (Fig. 7g). Yellowish-white stripe antero-medial on antenna restricted to A3–A7, at most with light brownish shade on adjacent proximal margin of A8 Celonites ivanovi Mauss & Fateryga, sp. nov.
Head in frontal view strongly oval (Fig. 9b). Antennal club (A8–12) in dorsal view about 1.68 times as long as broad, asymmetrical, with evenly rounded anterior margin and straight distal margin, that is running transverse. At postero-distal edge distal margin bent into posterior margin at nearly right angle (Fig. 9g); posterior margin nearly straight. Median lobe of tergum VII moderately produced, its posterior margin weakly concave in middle, with adjacent posterior translucent lamella fairly emarginated (Fig. 10e). Sternum VIII with longitudinal depression in centre more elongated, lancet-like tips and postero-median incision between them narrower (Fig. 14f). Aedoeagus with broadly rounded distal end, its lateral sides only weakly converging. Uncus thyrsos comparatively broad and blunt (Fig. 12f). Apodema thyrsos delicate, anteriorly nearly straight (Figs 10f, 11f). Outline of emargination between lateral lamella and postero-lateral process of propodeum with narrow neck, its apical end small oval-shaped (Fig. 7c). Yellowish-white stripe antero-medial on antenna continues on proximal part of A8 Celonites cagrii Mauss & Yildirim, sp. nov.

Geographic distribution

The distribution of the members of the C. cyprius-group is shown in Fig. 17. All of the 20 studied specimens from collection sites in Iran and Armenia that previously had been determined as C. cyprius smyrnensis Richards, 1962 by J. Gusenleitner turned out to belong to C. osseus (dbM 4617, 5675–5679, 5778–5788) and C. cagrii sp. nov. (dbM 5789, 5790) respectively or even to C. clarus (dbM 4616). For that reason, the occurrence of C. cyprius is considered as doubtful in Iran and Armenia and all of the nine published records of C. cyprius from both countries by Gusenleitner (1973, 1997) and Ebrahimi and Carpenter (2008) were excluded from the data set. Celonites rugiceps seems to be absent from the whole Levant region (Fig. 17). Reinvestigation of a single specimen from Jordan (dbM 2523) previously reported by Mauss and Prosi (2013) to represent C. rugiceps revealed that it was misidentified and belongs to C. cyprius instead.

Discussion

Celonites cagrii sp. nov., C. ivanovi sp. nov. and C. osseus can be assigned to Eucelonites and to the Celonites cyprius-group without contradictions since the imagines share the potential apomorphic characters of these taxa, that is the axilla is produced into a curved tapering projection which fits into a slight emargination of the tegula (Richards 1962) and the harpide has a tapering spatula-like distal end with a vertical duplication ventrally along the medial margin. Within the Celonites cyprius-group C. cagrii sp. nov., C. ivanovi sp. nov. and C. osseus are especially similar in most morphological characters. Therefore they are probably closely related and we suggest naming this group the C. osseus-complex. The close relationship of the members of the C. osseus-complex is supported by the COI-5P sequence analyses that revealed that within the C. cyprius-group the lowest interspecific genetic distances exist between C. ivanovi sp. nov. and C. cagrii sp. nov., while they differ from the available sequences of the other species by at least 11% (Fig. 16). However, the COI-5P gene sequence of C. osseus is still unknown, so that the close relationship of C. osseus to C. cagrii sp. nov. and C. ivanovi sp. nov. deduced from morphological data cannot be proved genetically at the moment.

Celonites osseus differs from C. cagrii sp. nov. and C. ivanovi sp. nov. in the shape of the harpides and the base of the male genital which is associated with different proportions of tergum VII and sterna VII+VIII of the males. Differences between C. cagrii sp. nov. and C. ivanovi sp. nov. exist in the form of the male antennal club, the proportions of tergum VII and sternum VIII, as well as some structures of the aedoeagus. All of these characteristics can be assumed to be associated with mating behaviour in Celonites (cf. Mauss 2006; Mauss and Müller 2014). Thus it can be hypothesized that the dissimilarity between these traits functions as a reproductive isolation mechanism between the taxa so that they probably constitute different biospecies sensu Mayr (1967). This hypothesis is corroborated by the measured mean interspecific genetic distance between C. cagrii sp. nov. and C. ivanovi sp. nov. of more than 7%. In Central European Apoidea and Vespoidea a level of COI sequence divergence exceeding 2% commonly signals different species, although there are a few exceptions in which maximum intraspecific genetic distances of up to 13.1% are not correlated with equally pronounced morphological characters (Schmidt et al. 2015; Schmid-Egger and Schmidt 2021).

Besides other morphological characteristics many species of the C. cyprius-group vary in the shape of the postero-lateral process and the lateral lamella of the propodeum resulting in different outlines of the emargination between them. Species specific differences in the shape of the postero-lateral part of the propodeum were already observed between C. cyprius, C. rugiceps and C. clarus (Richards 1962; Gusenleitner 1973), and also exist among another four Palaearctic (Gusenleitner 1973) and four Afrotropical species of Celonites (Gess 2007). The function of this structure is unknown and therefore its contribution to reproductive isolation is unclear. Nevertheless, the repeated observation of species-specific differences in the postero-lateral part of the propodeum within Celonites suggests that the distinct structures are associated with an unknown mechanism of reproductive isolation or niche segregation. Hence, it is of note that C. cagrii sp. nov. differs distinctly in this character from C. ivanovi sp. nov. and C. osseus that, for their part, resemble C. cyprius.

The relation of Celonites clarus to the C. cyprius-group is still unresolved. It is only known from the type series collected near Teheran (Gusenleitner 1973; Rahmani et al. 2020) and the single female from Kerman province in south-east Iran assigned to this species in this study. In accordance with the description by Gusenleitner (1973) this female and two investigated paratype specimens differ distinctly from all other members of the C. cyprius-group in the structure of the cuticula on frons and mesoscutum and the unique postero-lateral part of the propodeum. On the other hand, C. clarus is otherwise quite similar to the members of the C. osseus-complex apart from having comparatively shorter antennae and a weaker or even absent crenulation along the posterior margins of the metasomal terga. Since the male of C. clarus is still unknown and genetic data are unavailable, the position of the taxon within Eucelonites remains uncertain and should be reinvestigated with care, when more material becomes available.

Celonites ivanovi sp. nov. and C. cagrii sp. nov. were exclusively recorded from flowers of different Heliotropium species. This corresponds to other members of the C. cyprius-group for which flower visiting records are available, that is C. cyprius and C. rugiceps, that were also found at flowers of various Heliotropium plants (Richards 1962; Mauss pers. obs.). The behaviour of C. ivanovi sp. nov. during pollen uptake from H. styligerum flowers is very similar to the behaviour of C. cyprius and C. rugiceps at flowers of other Heliotropium species (Mauss pers. obs.). Therefore, it is probably homologous.

The geographic range of Celonites osseus is similar to Iranian faunal elements sensu de Lattin (1967). The adjacent small and probably endemic distribution areas of the closely related C. ivanovi sp. nov. and C. cagrii sp. nov. on the northern side of the Greater Caucasus and in East-Anatolia and Armenia respectively can be hypothesized to be relicts of a previous interglacial range expansion of their last common ancestor shared with C. osseus. In the following glacial epoch, range regression probably resulted in geographic isolation of small relict populations that evolved into separate biospecies. Postglacially C. ivanovi sp. nov. and C. cagrii sp. nov. remained in their glacial refuges, while C. osseus expanded its range again. The other species of the C. cyprius-group can also be assigned to different glacial refuges sensu de Lattin (1967) with C. rugiceps and C. cyprius being of Ponto-Mediterranean origin, while C. yemenensis has a distinct Syroeremial distribution and C. clarus seems to represent the Iranoeremial type. This distribution pattern indicates that speciation within the C. cyprius-group was most likely influenced by geographic isolation during glacial periods. The sympatric occurrence of C. cyprius with C. rugiceps in the west and of C. osseus with C. clarus in the east implies that some kind of niche segregation evolved between these taxa.

Just three species of pollen wasps (all in the genus Celonites) were previously known from Russia (Antropov and Fateryga 2017; Fateryga 2020). The present contribution increases this number to four. Since the known distribution ranges of C. ivanovi sp. nov. and C. cagrii sp. nov. are very restricted, both species may be endangered. Therefore, a systematic search for further localities of C. ivanovi sp. nov. in the Caucasus or C. cagrii sp. nov. in East Anatolia and Armenia should be carried out in suitable habitats with occurrence of Heliotropium plants.

Acknowledgments

Sergey A. Belokobylskij and Yulia V. Astafurova supported the second author during his work in ZISP. Rainer Prosi kindly prepared the distribution map from our records. Christian Schmid-Egger and Stefan Schmidt provided COI Sequences of eight specimens of the C. cyprius-group for analysis. A. Rosenbauer identified the Heliotropium species from Turkey. C. Schmid-Egger, J. Gusenleitner, M. Uliana, E. Ockermüller kindly provided material from their private collections or public collections under their care. The useful and encouraging reviews by Jack Neff and Andreas Müller are sincerely acknowledged.

The work of A.V. Fateryga was a part of the State research project No. 121032300023-7.

References

  • Antropov AV, Fateryga AV (2017) Family Vespidae. In: Lelej AS, Proshchalykin MY, Loktionov VM (Eds) Annotated Catalogue of the Hymenoptera of Russia. Volume I. Symphyta and Apocrita: Aculeata. Proceedings of the Zoological Institute RAS, Supplement 6: 175–196. https://doi.org/10.31610/trudyzin/2017.supl.6.5
  • Birket-Smith SJR (1981) The male genitalia of Hymenoptera – a review based on morphology in Dorylidae (Formicidae). Entomologica Scandinavica Supplement 15: 377–397.
  • Bischoff H (1928) Hymenoptera (Vespidae, Scoliidae, Mutillidae, Chrysididae und Evaniidae). In: Roewer CF (Ed.) Zoologische Streifzüge in Attika, Morea und besonders auf der Insel Kreta. II. Abhandlungen des Naturwissenschaftlicher Verein zu Bremen 27: 85–90.
  • Blüthgen P (1952) Weitere neue oder bemerkenswerte paläarktische Faltenwespen aus der Zoologischen Staatssammlung in München (Hym., Eumenidae, Masaridae). Mitteilungen der Münchener Entomologischen Gesellschaft 42: 1–19.
  • Bytinski-Salz H, Gusenleitner J (1971) The Vespoidea of Israel (Hymenoptera). Israel Journal of Entomology 6: 239–299.
  • Carpenter JM (1993) Biogeographic patterns in the Vespidae (Hymenoptera): two views of Africa and South America. In: Goldblatt P (Ed.) Biological relationships between Africa and South America. Yale University Press, New Haven, 139–155. https://doi.org/10.2307/j.ctt22726mc.11
  • de Lattin G (1967) Grundriß der Zoogeographie. Gustav Fischer, Stuttgart, 602 pp.
  • Dönmez A (2008) Heliotropium samoliflorum subsp. erzurumicum (Boraginaceae), a new subspecies from Turkey. Annales Botanici Fennici 45: 396–399. https://doi.org/10.5735/085.045.0509
  • Eck R (1978) Biometrische Untersuchung zur Klärung der Artunterschiede bei sozialen Faltenwespen (Hymenoptera: Vespidae). Entomologische Abhandlungen des Staatlichen Museums für Tierkunde Dresden 42: 315–344.
  • Fateryga AV (2020) New records of Celonites kozlovi Kostylev, 1935 and C. sibiricus Gusenleitner, 2007 (Hymenoptera: Vespidae: Masarinae), with observations on their behavior at flowers. Far Eastern Entomologist 405: 20–32. https://doi.org/10.25221/fee.405.4
  • Gess FW (2007) Four new species of the wasp genus Celonites Latreille, 1802 (Hymenoptera: Vespidae: Masarinae) from south-western Africa, designation of neotype for C. michaelseni von Schulthess, 1923, species representation in Namibia, and key to species occuring in Namibia. Journal of Hymenoptera Research 16: 11–29.
  • Gess SK, Gess FW (2010) Pollen wasps and flowers in Southern Africa. South African National Biodiversity Institute, Pretoria, SANBI Biodiversity Series 18, 147 pp.
  • Giordani Soika A (1957) Hymenoptera: Vespidae. In: Expedition to South-West Arabia 1937–1938. Volume 1. British Museum (Natural History), London, 471–484. [6 figs]
  • Grossheim AA (1967) Flora Kavkaza [Flora of the Caucasus]. Second edition. Volume VII. Nauka, Leningrad, 894 pp. [In Russian]
  • Gusenleitner J (1966) Vespidae, Eumenidae und Masaridae aus der Türkei. Teil I. Polskie Pismo Entomologiczne 36: 343–363.
  • Gusenleitner J (1973) Über Masaridae aus dem nahen Osten (Vespoidea, Hymenoptera). Bolletino della Museo Civico di Storia Naturale di Venezia 24: 55–69.
  • Gusenleitner J (1997) Die europäischen Arten der Gattung Celonites Latreille 1802 (Hymenoptera, Masaridae). Linzer Biologische Beiträge 29: 109–115.
  • Gusenleitner J (2002) Neue oder bemerkenswerte Vespoidea aus dem Nahen Osten (Hymenoptera: Eumenidae, Masaridae). Linzer Biologische Beiträge 34: 335–343.
  • Gusenleitner J (2005) Über Faltenwespen aus dem Oman Teil 2 (Hymenoptera, Vespidae, Eumenidae, Masaridae). Linzer Biologische Beiträge 37: 1199–1201.
  • Gusenleitner J (2007) Eine neue Celonites-Art aus Sibirien (Hymenoptera: Vespidae, Masarinae). Linzer Biologische Beiträge 39: 133–135.
  • Gusenleitner J (2010) Order Hymenoptera, family Vespidae. Arthropod fauna of the UAE 3: 422–467.
  • Gusenleitner J (2012) Neue Masarinae aus der paläarktischen Region (Hymenoptera: Vespidae: Masarinae). Linzer Biologische Beiträge 44: 319–326.
  • Gusenleitner J (2018) Neue asiatische Faltenwespen (Hymenoptera, Vespidae: Eumeninae, Masarinae). Linzer Biologische Beiträge 50: 303–308.
  • Kostylev G (1935) Materialien zur Kenntnis der Masariden-Fauna der Paläarktis. Archives du Muzeé Zoologique de l’Université de Moscow 2: 85–116. [In German and Russian]
  • Mauss V (2006) Observations on flower association and mating behaviour of the pollen wasp species Celonites abbreviatus (Villers, 1789) in Greece (Hymenoptera: Vespidae, Masarinae). Journal of Hymenoptera Research 15: 266–269.
  • Mauss V (2013) Description of Celonites andreasmuelleri sp. n. (Hymenoptera, Vespidae, Masarinae) from the Middle East with a key to the Palaearctic species of the C. abbreviatus-complex of the subgenus Celonites s. str. Journal of Hymenoptera Research 31: 79–95. https://doi.org/10.3897/JHR.31.4235
  • Mauss V, Fateryga AV, Prosi R (2016) Taxonomy, distribution and bionomics of Celonites tauricus Kostylev, 1935, stat. n. (Hymenoptera, Vespidae, Masarinae). Journal of Hymenoptera Research 48: 33–66. https://doi.org/10.3897/JHR.48.6884
  • Mauss V, Müller M (2014) First contribution to the bionomics of the pollen wasp Celonites fischeri Spinola, 1838 (Hymenoptera, Vespidae, Masarinae) in Cyprus. Journal of Hymenoptera Research 39: 119–153. https://doi.org/10.3897/JHR.39.7841
  • Mauss V, Prosi R (2013) First record of the pollen wasp Celonites rugiceps Bischoff 1928 (Hymenoptera, Vespidae, Masarinae) from Central Europe. Linzer Biologische Beiträge 45: 697–701.
  • Mauss V, Prosi R (2018) Identity and distribution of Celonites hermon Gusenleitner, 2002 (Hymenoptera, Vespidae, Masarinae) from the Middle East with a description of the hitherto unknown male. Journal of Hymenoptera Research 66: 55–70. https://doi.org/10.3897/jhr.66.29795
  • Mayr E (1967) Artbegriff und Evolution. Paul Parey, Hamburg, 617 pp.
  • Morawitz F (1888) Hymenoptera aculeata nova. Horae Societatis Entomologicae Rossicae 22: 224–302.
  • Richards OW (1962) A revisional study of the masarid wasps (Hymenoptera, Vespoidea). British Museum of Natural History, London, 294 pp.
  • Richards OW (1984) Insects of Saudi Arabia. Hymenoptera: Fam. Masaridae (the Arabian species). Fauna of Saudi Arabia 6: 413–422.
  • Schmid-Egger C (2017) Order Hymenoptera, family Vespidae. Subfamily Masarinae. In: van Harten A (Ed.) Arthropod fauna of the UAE. Volume 6. Department of The President’s Affairs, Abu Dhabi, 341–349.
  • Schmid-Egger C, Schmidt S (2021) Unexpected diversity in Central European Vespoidea (Hymenoptera, Mutillidae, Myrmosidae, Sapygidae, Scoliidae, Tiphiidae, Thynnidae, Vespidae), with description of two species of Smicromyrme Thomson, 1870. ZooKeys 1062: 49–72. https://doi.org/10.3897/zookeys.1062.70763
  • Schmidt S, Schmid-Egger C, Morinière J, Haszprunar G, Hebert PDN (2015) DNA barcoding largely supports 250 years of classical taxonomy: identifications for Central European bees (Hymenoptera, Apoidea partim). Molecular Ecology Resources 15: 985–1000. https://doi.org/10.1111/1755-0998.12363
  • Snelling RR (1986) The taxonomy and nomenclature of some Australian paragiine wasps (Hymenoptera, Masaridae). Los Angeles County Museum of Natural History Contributions in Science 378: 1–19. https://doi.org/10.5962/p.208122

Supplementary material

Supplementary material 1 

Supplementary material

Volker Mauss, Alexander V. Fateryga, Erol Yildirim, James M. Carpenter

Data type: investigated specimens, published records, localities

Explanation note: List of all specimens, published records and localities included in the study.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (269.00 kb)