JHR 89: 109-155 (2022) ger, JOURNAL OF eeertennscnn nn

doi: 10.3897/jhr.89.79832 RESEARCH ARTICLE () Hymenopter a
4

https://jhr.pensoft.net Thelnternaonl Society of Hymenopexriss, RESEARCH

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

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

I Stuttgart State Museum of Natural History, Entomology Department, Rosenstein 1, D-70191 Stuttgart,
Germany 2. 1:1. Vyazemsky Karadag Scientific Station, Nature Reserve of RAS, Branch of A.O. Kovalevsky Institute
of Biology of the Southern Seas of RAS, Nauki Str. 24, Kurortnoye, 298188 Feodosiya, Russia 3 Atatiirk University,
Faculty of Agriculture, Department of Plant Protection, TK 25240 Erzurum, Turkey 4 Division of Invertebrate,
Zoology, American Museum of Natural History, Central Park West at 79” street, New York, NY 10024, USA

Corresponding author: Volker Mauss (volker.mauss@gmx.de)

Academic editor: Michael Ohl | Received 27 December 2021 | Accepted 24 January 2022 | Published 28 February 2022
http://zoobank.org/6FA46014-BC04-4A27-9F49-B4437925DE4E

Citation: Mauss V, Fateryga AV, Yildirim E, Carpenter JM (2022) Contribution to the taxonomy, bionomics and
distribution of the Palaearctic Celonites cyprius-group (Hymenoptera, Vespidae, Masarinae) with the description of two
new species from the North Caucasus and East Anatolia. Journal of Hymenoptera Research 89: 109-155. https://doi.
org/10.3897/jhr.89.79832

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

Copyright Volker Mauss et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY
4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

110 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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 Ce/onites 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 Pal-
aearctic 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 identifica-
tion 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 be-
longing 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

Taxonomy, bionomics and distribution of the Celonites cyprius-group 111

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 Senkaya-Aksar 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 Dénmez (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, AK 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 Al—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

112 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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 $1). Additionally, 55 published records
were taken from Bischoff (1928), Bliithgen (1952), Bytinsky-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).

Taxonomy, bionomics and distribution of the Celonites cyprius-group 113

Systematics

Celonites osseus Morawitz, 1888

Celonites osseus Morawitz, 1888: 268-269, &, type locality: “territorio transcaspico
(Tschikischljar) [Turkmenistan]”, lectotype (designated here): 9, <golden disc>,
“Tschikischljar. Pom.[eranzev]”, “Celonites osseus 9. F. Moraw.”, “k. 0 Mopasnia
[coll. F Morawitz]”, “Lectotypus 9 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), misiden-
tification of Celonites crenulatus Morawitz, 1888 not C. osseus; Carpenter, 2001:
11, Armenia, Turkmenistan.

Additional material studied. Armenia: Zanga [currently Hrazdan] River, near Ye-
revan, 19.07.1935 192. (dbM 5855) leg. G. Kostyleyv ZMMU; Yerevan, Nork [=Nor
Nork], 23.06.1930 19 leg. A. Shelkovnikov ZISP. Turkmenistan, Balkan region:
Krasnovodsk [currently Tirkmenbasy], 13.06.1928 19 leg. V. Gussakovskij ZISP,
16.06.1928 14 leg. V. Gussakovskij ZISP; Jebel, 25 km NW Balkanabat, 22.05.1993
13 (dbM 5521) leg. M. Halada OLML. Iran: Elburz, 30 km S of Ab Ali, 09.07.1965
12 (dbM 5788) leg. Giordani Soika & Mavromoustakis MSNVE; Elburz, Vana 50 km
N di Ab Ali, 12—13.07.1965 109292 (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 192 (dbM 5787)
leg. Giordani Soika & Mavromoustakis MSNVE; Fars, Daria Namak, steppe presso
lago salato 27 km E Shiraz, 07.07.1965 32.2. (dbM 5679, 5778, 5779) leg. Giorda-
ni Soika & Mavromoustakis MSNVE; Fars prov., 20 km E Kazarun, 29°33.034'N,
51°49.416'E, 1256 m, 14.06.2019 14 leg. V.M. Gnezdilov ZISP; Golestan prov., 70
km E Minudasht, 1050 m, 12.06.2010 19 (dbM 4617) leg. M. Halada OLML; Te-
heran, Steppa presso Chitgar, 18 km W di Teheran, 15.07.1965, 12 (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 H—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

114 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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 |. 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).

Taxonomy, bionomics and distribution of the Celonites cyprius-group 115

Celonites cagrii Mauss & Yildirim, sp. nov.
http://zoobank.org/36232674-C53A-462F-8FFA-B47D84ACC4DF

Holotype. 2 (dbM 5610) “[Turkey] TR-Erzurum Senkaya-Aksar [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 Senkaya-Aksar [40.648262°N, 42.342351°E]
20.VIII.2011-1275 m Leg. E. Yildirim’ 19 (dbM 4574) EY; “[Turkey] TR-Erzurum
Senkaya-Aksar [40.648262°N, 42.342351°E] 12.VII.2020-1275 m Leg. E. Yildi-
rim [on Heliotropium ellipticum Ledeb.]” 32. (dbM 5607, 5608 (BOLD process ID
CECYP005-20), 5609) VM; “[Turkey] TR-Erzurum Senkaya-Aksar [40.648262°N,
42.342351°E] 01.VUI.2020-1275 m Leg. E. Yildirim [on Heliotropium ellipticum
Ledeb.]” 12 (dbM 5616) AMNH, 13 (dbM 5619) 39 (dbM 5611, 5612, 5615) VM,
49 (dbM 5613, 5614, 5617, 5618) EY; “ARMENIA Erevan Monti desertici Aighepat
40 Km. SE 23-VII-63 [leg. Giordani Soika]” 24.3 (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 dor-
so-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; lat-
erally 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; poste-
rior band on tergum | occupying whole of sides but less than half of middle part, some-
what widened anteriorly in median axis, anteriorly with small brownish tinge towards
adjacent black area; laterally and medially widened posterior bands on terga II—V, ante-
riorly 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 trans-
lucent margin of tergum VI; tarsi; sterna I—-V, posterior margin of sterna H—V translu-
cent. Antenna black except: yellowish white markings medial on A4—A7 and proximal
part of A8; brown-suffused area ventral on A10—Al1. 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 anten-
nal sockets, or both spots above antennal sockets large and dorso-laterally fused with

Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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).

Taxonomy, bionomics and distribution of the Celonites cyprius-group 117

hed Colouctas

Td Chey ty'tth - gid oe a
(an Fok: F, Ao tas

Figure 3. Lectotype of Celonites osseus a lateral view b head in frontal view € labels.

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 inter-
rupted (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 in-
terrupted 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 mark-
ings: light yellowish-white marking on A7 darkened (1); medial on proximal part of A8
only light brownish suffused (3) or with same colour than distal part (1).

Structure: Head in frontal view 1.50 times as wide as long in median (min 1.45,
max 1.54, n=5) (Fig. 4f). Mandible with two large blunt incisivi at distal end separated

by acute-angled cleft and two smaller more acute subapical incisivi on antero-medial

Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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

Taxonomy, bionomics and distribution of the Celonites cyprius-group 119

margin. External side of mandible distally bearing longitudinal rows of long stiff setae;
at base without distinct transverse depression; basal area with shagreened cuticula
moderately covered with pubescence of tiny thin setae (longer than in C. cyprius); anterior
to condylar ridge cuticula of basal area extends further apically becoming distinctly
striated in longitudinal direction; this area contrasts to smooth shiny but somewhat
longitudinally striated cuticula on condylar ridge and postero-apically adjoining surface;
condylar ridge distinct at basal two-third of mandible continuing in more gentle curve
into apical side (strongest bend approximately after basal third of mandible). Labrum
matt shining, finely shagreened and longitudinally wrinkled; densely covered with pale
stiff setae directing obliquely downwards; setae as long as A7 maximum wide, with distal
end curved ventro-medially, laterally at apex of labrum thicker with larger diameter at
base. Clypeus 1.4 times wider than long; translucent ventro-medial margin becoming
much narrower medially resulting in distinct median emargination; cuticula shiny,
ventro-medially above emargination smooth with sparse micropunctation becoming
moderately spaced dorsally and laterally on disc with larger irregular flat depressions
and wrinkles in addition; dorso-lateral vertical parts of clypeus smooth with moderately
spaced micropunctation partly striated at base; covered with pale thin stiff setae arising
from micropunctures; setae on disc about as long as A4, vertically erected with distal
ends strongly curved in ventro-medial direction, on sides shorter lying more flatly. Frons
very coarsely punctured, interstices shining, raised to transversal little rounded ridges;
protruding central part of supra-antennal area smooth, with moderately to sparsely
spaced macropunctures and few micropunctures; semi-circular depression of antennal
groove wrinkly shagreened; slight median depression dorsal to supra-antennal area,
frontal line weak; sparsely covered with pale short setae arising from coarse punctures.
Vertex with close macropunctation, becoming more closely reticulate behind ocelli
with smaller punctures and interstices more strongly raised forming short sharp-edged
+transversal ridges; sparsely covered with pale short setae arising from punctures; cuticula
of interstices shiny not shagreened (Fig. 5f). Median ocellus 1.2 times larger in diameter
than lateral ocelli; median ocellus somewhat bilateral symmetric with anterior sector
less strongly curved than posterior sector; lateral ocelli circular (in dorso-lateral view).
Compound eyes sparsely covered with tiny setae. Preoccipital carina sharp; medially
straight, nearly transversal; laterally behind dorso-lateral end of each compound eye
curved downwards for short distance becoming obsolete posterior to dorsal end of
postocular carina. Gena narrow, less than half as wide as basal width of A3. Postocular
carina sharp; extends dorsad from posterior mandibular articulation along posterior
margin of gena; ends level with dorsal end of compound eye anterior to preoccipital
carina that runs parallel for short distance. Antennal articles A8—A12 forming ventrally
flattened club about 2.0—2.2 times as long as broad (in dorsal view).

Pronotum with anterior margin raised to carina; anterior pronotal carina (sensu
Carpenter 1988) in antero-ventral area of pronotum weak, forming anterior sharp
edge along ventral half of crenate groove dorsally continuing into sharply bent but

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. 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).

Taxonomy, bionomics and distribution of the Celonites cyprius-group 124

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 mar-
gin of crenate groove and posterior margin of pronotum at same level as surrounding
surface, shiny, with some micropunctures; on lateral quarter distinct posterior pro-
notal 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 translu-
cent 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 adja-
cent 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 sulca-
ture 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 ex-
cept 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 sha-
greened 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. Pro-
podeum with horizontal propodeal triangles and dorso-lateral margins of posterior
face of propodeum reduced to two pointed protuberances dorsally on each side of

122 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

; ing: i 3 ft Sires 200 um

Figure 6. a,b Metasomal tergum II and HI 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. ivanovi sp. nov. (dbM 5492) d C. osseus
(dbM 4617) e C. yemenensis (dbM 5528) f C. clarus (dbM 4616).

Taxonomy, bionomics and distribution of the Celonites cyprius-group 123

middle, cuticula on protuberances rugose with interstices raised to knife-like edges;
posteriorly with narrow medial cuticula-fold running from dorsal margin to poste-
ro-medial flange of propodeum; posterior surface ventrally striated by strong vertical
cuticula-folds arising below anterior transversal carina of postero-medial flange of pro-
podeum, with shallow macropunctures between folds, moderately covered with fine
pale setae arising from macropunctures, laterally and dorsally continuing into coarsely
reticulate macropunctation with shorter setae. Cuticula below lateral lamella shiny,
on metepisternum densely horizontally wrinkled, on side of propodeum shagreened
with moderately spaced small shallow punctures. Lateral lamella moderate, slightly
curved laterally downwards, its outer margin gently curved, its apex truncate, outer
and posterior margins somewhat crenate; inner margin of lateral lamella and lateral
apex of postero-lateral process of postero-medial flange of propodeum separated by
small gap; anterior margin of postero-lateral process straight transverse, while posterior
margin converges in weak curve towards lateral apex; outline of emargination being
broad at its base with short narrow neck between lateral apex of postero-lateral process
and inner margin of lateral lamella and small oval-rounded apical part (Fig. 7c); dorsal
cuticula of lateral lamella and adjacent dorso-lateral part of propodeum shiny, with
reticulate macropunctation. On whole exoskeleton single thin seta arises from bottom
of each macropuncture, seta short if not stated otherwise.

Fore-femur postero-ventrally produced in middle forming anteriorly curved lobe
distally changing into tapering carina along ventral margin of femur; end of tibia when
folded against femur coinciding with produced region; tarsomeres I-IV broad and
flattened; underside of tibia and tarsomere I with strong obliquely distally directing
setae forming stiff brush; underside of tarsomere I and I] 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

124 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

a e

Awol

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. 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 sur-
face of propodeum fp = postero-medial flange of propodeum pp = postero-lateral process ll = lateral lamella).

along medial margin adjoining sternum VI, thereby slightly but continuously diverging
from lateral margin of tergum VI.

Metasomal sternum | 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

Taxonomy, bionomics and distribution of the Celonites cyprius-group 125

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 II[—V anteriorly with moderate to dense shallow macropunctation, posteriorly
changing into nearly unpunctured cuticula becoming moderately to densely covered by
micropunctures further postero-laterally and along posterior margin. Posterior margin
of sterna I-IV straight, posterior margin of sternum V medially concave running in
a gentle curve. Sternum VI (Fig. 8c) tapering towards distal end; with outer margin
forming bulged shiny rim, anteriorly raised to inwardly bent carina, postero-laterally
strongly curved medially resulting in blunter appearance of distal end of sternum VI;
rim at posterior end obtuse to nearly transverse, partly interrupted by depressions
of macropunctures, postero-medially protruded into little median spine; cuticula
with smooth median area tapering posteriorly, slightly raised to weak median keel
at posterior end that continues into median spine, laterally with moderately spaced
deep macropunctures becoming densely spaced and partly fused along lateral rim; stiff
setae of moderate length arising obliquely backwards from macropunctures; posterior
along distal end of rim densely covered with posteriorly directed stiff setae medially of
same length as median spine becoming shorter anteriorly; at dorso-posterior margin
dorsal (inner) cuticula protruded into irregularly serrated crystalline horizontal lamella
situated immediately above the posteriorly directed stiff setae (Fig. 8h-i), dorso-
medially fused with median spine, at postero-lateral edges somewhat more protruded.
Male. Colour (Fig. 2): Resembles female, except as follows. Light yellowish-white
are: large M-shaped band on frons, laterally filling each ocular sinus (Fig. 9b); clypeus
except dorso-lateral vertical sides and brownish translucent ventro-medial margin; lit-
tle 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 brown-
ish tinge towards adjacent blackish-brown area, on terga II-VI interrupted on each
side of middle by blackish-brown area. Tergum VII blackish-brown. Antenna black,
with light yellowish-white stripe antero-medial on A3—A7 and proximal part of A8.
Structure: Resembles female, except as follows. Head in front view 1.49 times
as wide as long (Fig. 9b, i). Mandible with single pointed tooth at distal end and
two smaller acute teeth distally on antero-medial margin. Labrum with flat ventro-
median area, shiny with few thin short pale setae; dorsal and lateral area set off by tiny
edge, strongly convex, weakly shagreened with rows of micropunctures from which
short thin pale setae arise. Clypeus 1.36 times wider than long, strongly convex; shiny,
ventro-medial area above emargination smooth, sparsely covered with micropunctures,
dorsally and laterally changing into moderately spaced micro- and dense uneven
shallow macropunctation; covered with short fine pale erected setae with distal end not

126 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

Nin tl”

Figure 8. a—c Female sternum VI in ventral view a C. yemenensis (dbM 4759) b C. ivanovi sp. nov.
(dbM 5491) ¢ 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).

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

Taxonomy, bionomics and distribution of the Celonites cyprius-group 127

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, Al10 and A11, tyloid of A9
smaller than others. Mid-coxa without small spine at distal end on anterior side close
to anterior-medial angle.

Tergum VII at posterior end with narrow median lobe and well set off postero-
lateral angle on each side (Fig. 9e); median lobe moderately produced, its posterior
margin weakly concave in middle, with adjacent posterior translucent lamella fairly
emarginated; translucent lamella continues on ventral side (in ventral view) at its base
on both sides into distinct carina running anteriorly along medial margin adjoining
sternum VII+VIII (fused); medial margin of postero-lateral angles running in semi-cir-
cular 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 ter-
gum; posteriorly with increasingly close and deep macropunctation, strongest medially
above sharp bend; interstices anteriorly distinctly shagreened, posteriorly smooth and
more shiny, postero-medially moderately covered with tiny pale setae.

Sternum VIII (Fig. 14f) acutely produced running into two pointed lancet-like
tips at posterior end with deep median incision between them, lancet-like tips and
median incision between them narrower than in C. ivanovi sp. nov.; convex with large
longitudinal depression in centre, lateral margins in proximal half bent horizontally;
cuticula shiny, with shallow macropunctures; pale postero-medially directed setae aris-
ing from macropunctures, posteriorly increasing in length, forming little tuft project-
ing over posterior median incision. Sparse transverse fringe of tiny setae along distal
end of sternum VII projecting over base of fused sternum VUI.

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

128 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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. osseus (dbM 5521) d C. ivanovi sp. nov. (dbM 5498) e C. ca-
grii 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) iC. cagrii sp. nov. (dbM 5619).

Taxonomy, bionomics and distribution of the Celonites cyprius-group 129

0.5mm

Figure 10. Male genitalia in dorsal view a C. rugiceps (dbM 2834) b C. cyprius (dbM 5452) € 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).

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

130 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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 Ce/onites 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. Cagri 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 maz.s.l. in a valley near Aksar, which is a vil-
lage in the Senkaya 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.

http://zoobank.org/1AB3C8D3-A445-409F-8443-C3BC920350B8

Holotype. 2 (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’,
12 AE 19 (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’, 13
(dbM 5497) AMNH, 14 (dbM 5498) OLML, 13' ZISPB 33 AE 24 (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.V1.2019 on Heliotropium styligerum leg. [A.V.]
Fateryga’, 12 (dbM 5494) AMNH, 19 ZISP, 59 AK 39 (dbM 5490, 5491, 5493)
13 (dbM 5495) VM; “[Russia] Dagestan, Maydanskoye 42°36'07"N, 46°58'13"E
15.V1.2021 on Heliotropium styligerum leg. [A.V.] Fateryga’, 492. (dbM 5856, 5857, 5858,
5995) VM; “[Russia] Dagestan, Maydanskoye 42°36'07"N, 46°58'13"E 16.VI.2021 on

131

Taxonomy, bionomics and distribution of the Celonites cyprius-group

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132 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

Heliotropium styligerum leg, [A.V.] Fateryga’, 22 (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.V1.2021 leg. [A.V.] Fateryga’, 12 (dbM 5997) VM.

Diagnosis. See key.

Description. Female. Colour (Figs 2, 4e): Black. The following are yellowish-
white: large rectangular spot dorso-medial on clypeus; two small spots adjoining ventral
margin of clypeus to both sides of median emargination; small spot on each ocular
sinus and two spots on frons; medium-sized spot on antero-dorsal angle of pronotum
(humeral spot); broad stripe along dorso-medial (inner) margin of pronotum,
anteriorly somewhat angularly enlarged with little median dent; large spot on dorsal
mesepisternum; laterally directed process of axilla; medium-sized spot postero-
medially on scutellum; dorsal and ventral side of propodeal lamella; antero-lateral one-
fifth and posterior two-fifth of tegula, interrupted by brownish translucent area on
bulge changing into black towards antero-medial margin of tegula; posterior band on
tergum I occupying whole of sides but less than half of middle part, somewhat widened
anteriorly in median axis, anteriorly with small brownish tinge towards adjacent black
area; laterally and medially widened posterior bands on terga II-V, anteriorly with
small brownish tinge towards adjacent black area, interrupted on each side of middle
by brownish area (Fig. 6b); little longitudinal spot antero-medial on tergum VI; small
spots on postero-lateral edges of sterna II-III; outside of distal tips of fore-, mid- and
hind-femora; outside of proximal end of fore-tibia; outside of mid-tibia except small
diagonal blackish-brown interruption in middle; outside of hind-tibia except blackish
ring covering one-third below middle. Brown are: distal half of mandible; maxillary
and labial palpi, protrudeable parts of proboscis; ventral margin of labrum; translucent
ventro-medial margin of clypeus; postero-lateral margin of scutellum; median third
of metanotum; postero-lateral process of propodeum; humeral plate (at base of wing
underneath tegula); sides and posterior translucent margin of tergum V]; 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—Al1 and
proximal part of Al2. 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); hu-
meral 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).

Taxonomy, bionomics and distribution of the Celonites cyprius-group 133

b Cc

0.5 mm

Figure | 1. Microphotos of male genitalia in dorsal view a C. rugiceps (dbM 5450) b C. cyprius (dbM 5439)
CC. 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

134 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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 Ftransversal 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

Taxonomy, bionomics and distribution of the Celonites cyprius-group 135

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 tcontinuing into bottom ribs; on lateral quarter
distinct posterior pronotal carina sharply separating semicircular antero-ventral area

b Cc

NAY
ANY \\

\

—— =

Nt

IRN
cide
ii

——

—

aa ih
\ QO

Nii
HEL,

0.5mm

Figure | 2. Male genitalia in ventral viewa C. rugiceps (dbM 2834) b C. cyprius (dbM 5452) € 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).

136 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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

Taxonomy, bionomics and distribution of the Celonites cyprius-group 137

0.5mm

Figure 13. Microphotos of male genitalia in ventral view a C. rugiceps (dbM 5450) b C. cyprius (dbM
5439) ¢ 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).

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

138 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

converges in a gentle curve towards lateral apex; outline of emargination being broad at
its base with short moderately wide neck between lateral apex of postero-lateral process
and inner margin of lateral lamella and deep medially extended apical part (Fig. 7g);
dorsal cuticula of lateral lamella and adjacent dorso-lateral part of propodeum shiny,
with reticulate macropunctation. On whole exoskeleton single thin seta arises from
bottom of each macropuncture, seta short if not stated otherwise.

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

Metasomal terga with postero-lateral corners slightly produced; posterior margin
of tergum I weakly crenulated, crenulation not produced into spines and not pro-
jecting 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. Gb); 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 me-
dian lobe running in convex oval curve formed by distinct translucent lamella; poste-
rior 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 pro-
jecting beyond postero-median translucent lamella and lateral margins; on ventral side
(viewed from ventral) posterior translucent lamella of median lobe continues on both
sides into distinct carina running anteriorly along medial margin adjoining sternum
VI, thereby slightly but continuously diverging from lateral margin of tergum VI.

Metasomal sternum I shiny, finely shagreened, with tiny setae but without punc-
tures. 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 translu-
cent 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 se-
tae arise; sterna HI—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 run-
ning in a gentle curve. Sternum VI tapering towards distal end; with outer margin

Taxonomy, bionomics and distribution of the Celonites cyprius-group 139

wv

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

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-anten-
nal area; clypeus except dorso-lateral vertical sides and brownish translucent ventro-
medial margin; complete longitudinal stripe on outside of fore-tibia; whole outside

140 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

Figure 15. Female antennal articles Al—A6 in medial view (scaled to same size to show different propor-

tions of A3) a C. clarus (dbM 5674) b C. osseus (dbM 5678).

of mid- and hind-tibia; outside of hind-metatarsus. Labrum translucent brown with
two yellowish-white antero-lateral spots. Tergum VI as in terga H—V with laterally
and medially widened posterior band, anteriorly with small brownish tinge towards
adjacent black area, interrupted on each side of middle by brownish area. Tergum VII
blackish-brown posteriorly changing into brown. Antenna black, with yellowish-white
stripe antero-medial on A3—A7.

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

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

Tergum VII at posterior end with characteristically narrow median lobe and
well set off postero-lateral angle on each side (Fig. 9d); median lobe with translucent
lamella, that continues on ventral side (in ventral view) at its base on both sides into
distinct carina running anteriorly along medial margin adjoining sternum VH+VUI

Taxonomy, bionomics and distribution of the Celonites cyprius-group 141
Celonites cagrii CECYP006-20
-Celonites cagrii CECY P005-20
Celonites ivanovi AIMEJ038-20
Celonites ivanovi AIMEJ037-20
Celonites ivanovi AIMEJ036-20
Celonites cyprius BCHYM3010-14
Celonites cyprius BCHYM10881-15
Celonites cyprius ATMEJ002-20
‘elonites cyprius AIMEJ003-20
Celonites cyprius AIMEJ001-20
Celonites yemenensis GBACU2693-13
Celonites pemenensis GBACU2692-13
Celonites rugiceps BCHYM10873-15
Celonites rugiceps BCHYM10869-15
Celonites rugiceps BCHYM10857-15
Celonites rugiceps AIMEJ004-20
Celonites rugiceps AIMEJ005-20

1%
: Celonites rugiceps BCHYM10816-15

Figure 16. Neighbour joining BOLD taxon ID tree of 18 specimens of the C. cyprius-group based on
COL5P 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).

(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

142 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

with increasingly close and deep macropunctation, strongest medially above sharp
bend; interstices anteriorly distinctly shagreened, posteriorly smooth and more shiny,
postero-medially moderately covered with tiny pale setae.

Sternum VIII acutely produced running into two pointed lancet-like tips at poste-
rior end with deep median incision between them (Fig. 14e); convex with large longi-
tudinal oval depression in centre, lateral margins in proximal two-third bent horizon-
tally; 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, lle, 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 mini-
mum 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.

Taxonomy, bionomics and distribution of the Celonites cyprius-group 143

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 as.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 (cal-
culated for the nearby village Untsukul by https://de.climate-data.org). At both locali-
ties, the most abundant flowering plants were Heliotropium styligerum Trautv. (Boragi-
naceae), as well as Xanthium sp. and X. spinosum L. (Asteraceae). In 2019 and 2021
neither 1. styligerum nor the wasps were found at locality 1. The third locality was
situated at 1345 ma.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 flow-
ers 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).

Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

144

‘(WO RIEPYMLIPINIEU MMM “YET [eINIeNY YIM spews ‘popnypoxo erususy pue uesy wos sazudi > Jo spro0da1 Poysty
-qnd Ajsnoraaid ‘sornyesau] Wosy spr0der sopIID uado ‘pareSnsaaur susumtdeds sapom9 ][NF) dnoss-sagudio “Dy ayi Jo siaquisw Jo uoNmNstp s1ydesSo0a5 +7 | aanBi4

sisuauawa “4D
sdaai6ns “9
snasso ‘9
IAQUBAI "9D
snudfo ‘9
sniejo "9
miBeo "9

Taxonomy, bionomics and distribution of the Celonites cyprius-group 145

Figure 18. a-e, g, h Flower visiting behaviour of Celonites ivanovi sp. nov. females at flowers of

Heliotropium styligerum and behaviours associated with it (details see text) f habitat of C. ivanovi sp. nov.
at locality 2 in the vicinity of Maydanskoye, Dagestan i habitat of C. cagrii sp. nov. near Aksar, Turkey
(aspect in autumn).

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.

146

Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

Key to the species of the Celonites cyprius-group

Male of Celonites clarus Gusenleitner, 1973 not known.

1

Larger, body length approximately 7-8 mm, more sturdily built (Fig. 1).
Crenulation of terga H-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. ......
I eNO OS, EN RR et Us De Celonites rugiceps Bischoff, 1928
Smaller, body length approximately 5-6.5 mm (Figs 1, 2). Crenulation of
terga I-IV considerably weaker, sometimes almost obsolete (Fig. 6b). Emar-
gination 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
GUMMTPOUNG CVSS Mss eS aeh sauerks eateb scurson SMR ee teres Rtn aes heed sae kcaeons lhe cured 3
Postero-lateral process of propodeum with blunt apical end, outline of emar-
gination between lateral lamella and postero-lateral process narrowly elongat-
ed (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. Ge). Head very broad, in median
[sd Gimies-as- wide asslone- (R52 1.58 EWG, 4a)e si vsloninesracooeboseleesenntetios nhead
Loe eh BCP eg Bet err Nec hot 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. Gf). Head in some species less broad (Fig. 4c) ........... 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 se-
tae, 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

Taxonomy, bionomics and distribution of the Celonites cyprius-group 147

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?ssscmese. stssrsness ct HeNtg ss nde nat ascends eae. 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 be-
tween 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 some-
times additional yellow spot close to mandible. Head very broad, in median
1.56 times as wide as long (1.55—1.56) (Fig. 4D)... eee ce eeeseeeeeeeeeeeeeeeeees
OEE. Pee eee ee, ee, ee Ne Celonites clarus Gusenleitner, 1973
Mesoscutum reticulate, with distinctly raised narrow interstices partly form-
ing 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
Blaelwar with: velloye wae cit stsous susassNuete esau Shue rauatioet aes asudtusetoesetanee 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 pos-
terior wall separated from posterior margin of pronotum by a stripe of cu-
ticula 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) vee ee eeeeees
RS at leat ogt Sake, gid 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).

148

Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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 tcontinu-
ing into bottom ribs of crenate groove. Bulged shiny rim of sternum VI forms
regular curve postero-laterally resulting in more pointed appearance of poste-
rior 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
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-cir-
cular (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: (Hit -2,) tx. areutu. Massie ectstd neta eat Celonites osseus Morawitz, 1888
Larger, body length approximately 7-8 mm, more sturdily built (Fig. 1). Ster-
num VIII semicircularly produced, not emarginated (Fig. 14a). Male genital
as in Figs 10a, 1la. 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
extendine-selerite (Pics T2 bth DoD) nsisceshensesrnrste'slentaets tater nvracarape diced 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
lorie setaex Piss 10D) D3b)e tan ieseta nscale 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,

10

1]

12

Taxonomy, bionomics and distribution of the Celonites cyprius-group 149

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 0... eee eeeeeeeeeeeeeeeeeeeeees
Yo PUNE LT SRL We RIT Celonites yemenensis Giordani Soika, 1957
Posterior lobes of sternum VII 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... eee 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-
Peteraltaneles (Pig Ie). .ctahatisvnoracssnsbleynolene Celonites osseus Morawitz, 1888
Male genital narrower at base (Figs 10e, f, lle, 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, €)... ce ceeeeeseseesseeseeseeeees 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 poste-
rior margin forming a blunt angle (Fig. 9f); posterior margin weakly con-
vex. Median lobe of tergum VII markedly produced, its posterior margin
and adjacent posterior translucent lamella entirely convex (Fig. 9d). Sternum
VII 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
J aie Pale oR A I a IL Sia a I 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 pos-
tero-distal edge distal margin bent into posterior margin at nearly right angle
(Fig. 9g); posterior margin nearly straight. Median lobe of tergum VII mod-
erately produced, its posterior margin weakly concave in middle, with adja-
cent posterior translucent lamella fairly emarginated (Fig.10e). Sternum VII
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,

150 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

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 om proxiinal,part-of AS. .n isc: welebe ict lseetotetettetva thet
ns eae oie mea Aapet th ne tani be be 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 Ce/onites 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 propor-
tions of tergum VII and sterna VII+VII 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

Taxonomy, bionomics and distribution of the Celonites cyprius-group 11

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 Miller 2014). Thus it can be hypothesized that the dis-
similarity between these traits functions as a reproductive isolation mechanism be-
tween the taxa so that they probably constitute different biospecies sensu Mayr (1967).
This hypothesis is corroborated by the measured mean interspecific genetic distance be-
tween 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 in-
traspecific 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 propo-
deum 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 un-
known 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 Ce/onites 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 mem-
bers 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 pos-
terior margins of the metasomal terga. Since the male of C. c/arus is still unknown and
genetic data are unavailable, the position of the taxon within Eucelonites remains un-
certain 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.

152 Volker Mauss et al. / Journal of Hymenoptera Research 89: 109-155 (2022)

The geographic range of Ce/onites osseus is similar to Iranian faunal elements sensu 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. Ockermiiller
kindly provided material from their private collections or public collections under
their care. The useful and encouraging reviews by Jack Neff and Andreas Miiller are
sincerely acknowledged.

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

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Supplementary material |

Supplementary material

Authors: 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.

Copyright notice: 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.

Link: https://doi.org/10.3897/jhr.89.79832.suppl1