Celonites ivanovi Mauss & Fateryga, 2022

Fateryga, Alexander V., Mauss, Volker, Proshchalykin, Maxim Yu. & Maharramov, Mahir M., 2025, Taxonomic review of the pollen wasps (Hymenoptera: Vespidae: Masarinae) from the Caucasus, with the description of two new species, new nest and flower-visiting records, and a key to the species, Zootaxa 5716 (4), pp. 514-540 : 520-522

Celonites ivanovi Mauss & Fateryga, 2022

( Figs 4 , 10B, K )

Celonites ivanovi Mauss & Fateryga in Mauss et al. 2022a: 130, figs 2, 4e, 5b, d, e, 6b, c, 7g, 8b, f, g, 9a, d, f, h, 10e, 11e, 12e, 13e, 14e, 18a–e, g, h, ♀ ♂ (type locality: “[ Russia] Dagestan, Maydanskoye 42°36'16"N 46°58'10"E [corrected to 42°36'07"N, 46°58'13"E in 2021] … on Heliotropium styligerum ”), holotype, ♀, OLML (examined).

Material examined. HOLOTYPE: ♀, labeled “ Dagestan, Maydanskoye / 42°36'16"N 46°58'10"E / on Heliotropium styligerum / 11.VI.2019 leg. Fateryga // ♀ Celonites sp. aff. cyprius / de Saussure, 1854 / det. Fateryga, 2019 // HOLOTYPUS / Celonites ivanovi MAUSS / & FATERYGA, 2022 ♀ / designated by V. Mauss 2021 / db Mauss Nr.: 5492 [red label]” [ OLML]. PARATYPES: RUSSIA. Dagestan: Untsukulskiy District, vicinity of Maydanskoye, on Heliotropium styligerum , 23.VI.2018, 2 ♀ (dbM 5287), leg. A. Fateryga [ CAFK, CVMM]; Maydanskoye, 42°36'16"N, 46°58'10"E [actually 42°36'07"N, 46°58'13"E], 11.VI.2019, 8 ♂ (dbM 5497, 5498, 5496, 5499), leg. A. Fateryga [ 1 ♂, AMNH; 3 ♂, CAFK; 2 ♂, CVMM; 1 ♂, OLML; 1 ♂, ZISP]; ibid., on Heliotropium styligerum , 11.VI.2019, 10 ♀ (dbM 5494, 5490, 5491, 5493), 1 ♂ (dbM 5495), leg. A. Fateryga [ 1 ♀, AMNH; 5 ♀, CAFK; 3 ♀, 1 ♂, CVMM; 1 ♀, ZISP]; Maydanskoye, 42°36'07"N, 46°58'13"E, on Heliotropium styligerum , 15.VI.2021, 4 ♀ (dbM 5856, 5857, 5858, 5995), leg. A. Fateryga [ CVMM]; ibid., 16.VI.2021, 1 ♂ (dbM 5996), leg. A. Fateryga [ CVMM]; ibid., on Heliotropium styligerum , 16.VI.2021, 2 ♀ (dbM 5859, 5860), leg. A. Fateryga [ CVMM]; Vicinity of Turtsi, 42°11'34"N, 47°09'33"E, on Heliotropium styligerum , 22.VI.2021, 1 ♀ (dbM 5997), leg. A. Fateryga [ CVMM]. ADDITIONAL MATERIAL: RUSSIA. Dagestan: Maydanskoye, 42°36'07"N, 46°58'13"E, on Heliotropium styligerum , 28.V.2022, 1 ♀, leg. A. Fateryga [ CAFK]; Vicinity of Maydanskoye, 42°35'39"N, 46°58'17"E, 19.VI.2023, 1 ♂, leg. A. Fateryga [ CAFK]; Khotoch, 42°24'52"N, 46°57'10"E, 17.VI.2023, 1 ♂, leg. M. Proshchalykin [ CAFK].

Distribution. Russia ( Dagestan) ( Fig. 11 ).

Biology. Four old nests of this species were found in the vicinity of Maydanskoye in Dagestan ( 42°35'39"N,

46°58'17"E) on 19.VI.2023. The nesting site was a steep rocky slope facing south; at the bottom of the slope, there was a shale scree covered with sparse herbs predominated by Heliotropium styligerum Trautv. ( Boraginaceae ) ( Fig. 3A ). Three females ( Fig. 4A ) and one male of C. ivanovi were observed at this locality during 2 h. All four nests were attached to subvertical surfaces of medium-sized stones ( Fig. 3B ). The nests were made of fine clayey soil with a small proportion of sand grains and consisted of one to three cells. In the case of more than one cell, they were arranged in a longitudinal row in which each subsequent cell abutted with its basal end onto the apical end of the preceding cell. An additional nest covering was not present. Nest no. 1 consisted of one cell oriented vertically, with the entrance opening downwards ( Fig. 3C ); nest no. 2 consisted of one cell oriented diagonally, with the entrance opening rather upwards ( Fig. 3E ); nest no. 3 consisted of three cells oriented vertically, with the entrance opening downwards ( Fig. 3F ); nest no. 4 consisted of two cells oriented diagonally, with the entrance opening rather downwards ( Fig. 3G ). The lengths of the nests were 9.5, 7.5, 22, and 17 mm, respectively. The outer diameter of the cells was 3.5– 4 mm, with the narrowest part at the plug and the broadest one in 1–2 mm from the basal end. The thickness of the cell walls was about 0.2 mm. Each cell was sealed with a flat plug, also about 0.2 mm thick. The inner length of the cells was 6.0–6.5 mm. Outside around the cell plug, the cell walls continued and covered either the rounded bottom (basal end) of the next cell or the closing plug of the last cell, which had the same structure as the bottom of the cells. The bottom of the next cell or the closing plug were partially fused with the plug of the previous cell at the center and detached from it laterally, resulting in a small hollow space ( Fig. 3D ). In the case of the last (or a single) cell of the nest, the cell walls continued beyond the cell plug for 1–4 mm, forming a small rim, which had one or two deep notches and was slightly bent outwards ( Fig. 3C ).

Nest no. 1 contained an empty cocoon of Celonites , the first cell of nest no. 3 was broken and empty, while five other cells (nest no. 2, the second and the third cells in nest no. 3, and both cells in nest no. 4) contained empty cocoons of a cuckoo wasp, presumably of the genus Spintharina Semenov ( Hymenoptera : Chrysididae ). The cocoon of Celonites was whitish and soft, covering the whole inner volume of the cell and hardly detachable from the cell walls, with fecal pellets in the basal third ( Fig. 3D ). Cocoons of Spintharina were only 3.5– 4 mm long; they covered somewhat more than a basal half of the cell walls and then terminated on the head end by a transverse cap, which was often separated from the remaining part of the cocoon by the emerged imago. The cocoons were also whitish but denser than that of Celonites , parchment-like and semitransparent ( Fig. 3G ), easy detachable from the cell walls. Separated caps of the cocoons were sometimes visible through the holes that had been made by the emerging cuckoo wasps in lateral walls of the cells ( Fig. 3F ). After emergence of the imagines, nest no. 1 and the first cell in nest no. 4 were reused by spiders that placed their egg cocoons inside of them.

Nests of five Palaearctic species of the genus Celonites were described previously: C. abbreviatus (Villers) , C. mayeti Richards , C. fischeri Spinola , C. tauricus Kostylev , and C. jousseaumeii du Buysson ( Lichtenstein 1869, 1875; Bellmann 1984; Mauss & Müller 2014; Mauss et al. 2016, 2022b). In all cases, the nests consist of usually several rather cylindrical mud cells with thin walls that are attached to either stones or twigs. In most species, at least some cells are attached longitudinally to each other. Linearly arranged brood cells were observed in a few nests of C. abbreviatus ( Lichtenstein 1869) and C. fischeri (Mauss & Müller 2014) , but in these nests at least a single cell was also attached longitudinally to the others in addition. An exclusively linear arrangement of the brood cells was previously only described in C. jousseaumei ( Mauss et al. 2022b) . Taking into account the phylogenetic relationships of C. ivanovi and C. jousseaumei ( Mauss et al. 2022 a, 2024; Fateryga et al. 2023), we can assume that this type of linear cell arrangement belongs to the ground pattern of the clade containing C. jousseaumei and the members of the C. cyprius -group (sensu Mauss et al. 2022a). Nests of C. ivanovi are very similar to those of C. jousseaumei , except that the outer cell surface shows a less distinct “fish scale” pattern than in the latter species ( Mauss et al. 2022b). However, the slight difference in surface structure could be the result of an aging and weathering process, since the investigated cells of C. jousseaumei were freshly build, while the examined empty cells of C. ivanovi were at least one year old.

Cuckoo wasps of the genus Spintharina are known to be associated with the pollen wasp genus Celonites . Spintharina versicolor (Spinola) was recorded as a brood parasite of C. abbreviatus ( Blüthgen 1961; Erlandsson 1972), while S. arnoldi (Brauns) and S. bispinosa (Mocsáry) were reared from brood cells of two different Afrotropical species of Celonites ( Brauns 1913; Gess 1996). A cocoon with a larva of Spintharina innesi (du Buysson) was recently recorded in a brood cell of C. jousseaumei ( Mauss et al. 2022b) . There are at least three species of Spintharina in the Caucasus, which are expected to be distributed in its Russian part ( Rosa et al. 2019). It is worthy to mention that in the nests of C. ivanovi the cocoons of Spintharina sp. were not located inside of a hostcocoon, which is similar to the position of a cocoon of S. innesi in a nest of C. jousseaumei ( Mauss et al. 2022b) . Pauli et al. (2018) found a close phylogenetic relationship between Spintharina and the Nearctic genus Chrysurissa Bohart , which also appears to parasitize predominately pollen wasps ( Hicks 1927, 1929; Hungerford 1937; Parker 1967). This phylogenetic placement indicates that both genera are descendants of a last common ancestor which may already have exploited pollen wasps as hosts, suggesting an old and exclusive association with pollen wasps in this clade ( Pauli et al. 2018; Mauss et al. 2022b). However, the cocoons of Chrysurissa densa (Cresson) were found inside of the cocoons of their host, Pseudomasaris vespoides (Cresson) ( Hicks 1927) . This may indicate a difference between larval behaviors of the two cuckoo-wasp genera: the larva of Spintharina probably consumes a full-grown host larva before it starts cocooning, while the larva of Chrysurissa consumes the host as a prepupa or even pupa that has already finished spinning its cocoon. Cocoons of cuckoo wasps inside host cocoons are common in some other groups of the family Chrysididae , particularly in species associated with bees ( Martynova 2020), and Ch. densa is also reported to parasitize bees besides the genus Pseudomasaris Ashmead ( Hicks 1927, 1929).

Imagines of C. ivanovi exclusively visit flowers of Heliotropium styligerum ( Mauss et al. 2022 a, 2024) ( Fig. 4B ). Females of this species demonstrate a special case of narrow oligolecty (sensu Müller & Kuhlmann 2008): pollen collection on only one plant species in the absence of coflowering congenerics. Morphological and behavioral adaptations of C. ivanovi for pollen uptake from flowers of H. styligerum were described in detail by Mauss et al. (2024).

Figures

FIGURE 3. Nests of Celonites ivanovi Mauss & Fateryga, 2022 in Dagestan. A, nesting site (arrow indicates the position of nest no. 1); B, position of nest no. 1 (indicated with an arrow); C, nest no. 1, close up; D, the same nest longitudinally dissected; E, nest no. 2; F, nest. no. 3; G, nest no. 4. Scale bars 1 cm.

FIGURE 4. Celonites ivanovi Mauss & Fateryga, 2022. A, female standing on a small stone; B, female at flower of Heliotropium styligerum Trautv.

FIGURE 10. Characters of pollen wasps. A, I, Celonites tauricus Kostylev, 1935, ♀, Azerbaijan; B, C. ivanovi Mauss & Fateryga, 2022, ♀, Dagestan; C, J, C. tudesculptus Kostylev, 1935, ♀, Azerbaijan; D, H, C. laetus Panfilov, 1968, ♀, holotype; E, C. osseus Morawitz, 1888, ♀, Armenia; F, C. tauricus, ♂, Dagestan; G, C. osseus, ♂, Iran; K, C. ivanovi, ♂, paratype; L, Jugurtia eurycara Kostylev, 1935, ♀, Azerbaijan; M, J. irana Kostylev, 1935, ♀, Azerbaijan. A–C, emargination between lateral lamella and postero-lateral process of propodeum; D, E, tegula; F, G, apex of metasoma in dorsal view; H, metasomal terga II–VI; I–K, metasoma in dorsal view; L, M, metasomal terga II–IV. Scale bars 1 mm.

FIGURE 11. Distribution of pollen wasps in the Caucasus. A, Celonites cagrii Mauss & Yildirim, 2022; B, C. carpenteri Fateryga & Mauss, sp. nov.; C, C. ivanovi Mauss & Fateryga, 2022; D, C. laetus Panfilov, 1968; E, C. osseus Morawitz, 1888; F, C. tauricus Kostylev, 1935; G, C. tudesculptus Kostylev, 1935; H, Ceramius caucasicus André, 1884; I, Jugurtia eurycara Kostylev, 1935; J, J. irana Kostylev, 1935; K, Quartinia aliyevi Fateryga, Mauss, Proshchalykin & Maharramov, sp. nov.; L, Q. araxana Giordani Soika, 1960.