yaa

JHR 66: 55-70 (2018) JOURNAL OF ‘revere cveraceioar
doi: 10.3897/jhr.66.29795 RESEARCH ARTICLE ME Hymenoptera

http://jhr.pensoft.net The Inerational Society of ymenoptersts. RESEARCH

Identity and distribution of Celonites hermon
Gusenleitner, 2002 (Hymenoptera, Vespidae, Masarinae)
from the Middle East with a description of the hitherto

unknown male

Volker Mauss!, Rainer Prosi?

I Staatliches Museum fir Naturkunde, Abt. Entomologie, Rosenstein 1, D-70191 Stuttgart, Germany 2. Lerchen-
strafse 81, D-74564 Crailsheim, Germany

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

Academic editor: M@. OA/ | Received 16 September 2018 | Accepted 23 October 2018 | Published 31 October 2018
http://zoobank.org/ADA1A7/C2-3976-4FA 1-A63C-8D6EDB8EA71C

Citation: Mauss V, Prosi R (2018) Identity and distribution of Celonites hermon Gusenleitner, 2002 (Hymenoptera,
Vespidae, Masarinae) from the Middle East with a description of the hitherto unknown male. Journal of Hymenoptera
Research 66: 55-70. https://doi.org/10.3897/jhr.66.29795

Abstract

Based on the morphological examination of the exoskeleton and the male genitalia of newly discovered
specimens from Syria the status and taxonomic position of Celonites hermon Gusenleitner, 2002 is re-
investigated. The hitherto unknown male is formally described. Celonites hermon is a member of the
C. abbreviatus-complex. It shares several probably derived characters with C: andreasmuelleri, from which
it is considered to be reproductively isolated by differences in the male genitalia, the form of sternum VII
and tergum VII as well as the colour pattern of the male antennae. Therefore C. hermon is hypothesized to
represent a separate biospecies. It has been recorded only from a small area in the mountain ranges along

the Lebanese section of the Dead Sea Transform in Syria and Israel.

Keywords

Pollen wasps, biogeography, taxonomy, Palaearctic, Israel, Syria

Copyright Volker Mauss, Rainer Prosi. 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.

56 Volker Mauss & Rainer Prosi / Journal of Hymenoptera Research 66: 55—70 (2018)

Introduction

The genus Celonites is very distinct from other pollen wasps in the wide horizontal
lamellae on the propodeum and the acute sides of the metasoma (Richards 1962).
Both characters are related to the outstanding ability of the imagines to roll up
(Gess and Gess 2010). Approximately 46 species of the genus (including six well
separated subspecies) have been recognized in the Palaearctic 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). With-
in the Palaearctic Celonites abbreviatus-group (sensu Richards 1962) the Celonites
abbreviatus-complex constitutes a well defined monophylum characterized by the
existence of knobbed setae at the front of the head forming a pollen collecting ap-
paratus (Mauss 2013). All members of this species-complex for which flower visits
have been recorded were observed to visit nototribic flowers of Lamiaceae (Beqg-
uaert 1940, Schremmer 1959, Miller 1996, Mauss 2006, Mauss 2013, Mauss et al.
2016). During pollen uptake from these flowers a female would stand on the lower
lip of the corolla, while she performs at a high frequency slight back and forth move-
ments, rubbing her head over the nototribic anthers. In this manner pollen grains
are removed from the anthers with the stiff knobbed setae on frons and clypeus of
the female (Schremmer 1959, Miiller 1996, Mauss 2006, Mauss et al. 2016). The
species-complex includes six nominate species, namely Celonites abbreviatus (Vil-
lers), C. andreasmuelleri Mauss, C. hermon Gusenleitner, C. mayeti Richards, C.
persicus Richards and C. tauricus Kostylev (= C. spinosus Gusenleitner) (Mauss 2013,
Mauss et al. 2016).

Celonites hermon was described from a single female collected by Bytinski-Salz
at Mount Hermon in the Golan Mountains by Gusenleitner in 2002. He separated
the specimen from the “very similar” Celonites abbreviatus because of its extremely
long proboscis that reaches the distal end of the abdomen (Gusenleitner 2002).
As already pointed out by Mauss (2013) proboscis length can not be examined
in most Celonites specimens, since the mouthparts are usually retracted, while the
protruded proboscis of the type specimen is an exception. This may have misled
Gusenleitner (2002) into believing, that the length of the proboscis is outstanding
in C. hermon. In fact, the careful investigation by Schremmer (1961) had already
demonstrated that the proboscis of C. abbreviatus clearly exceeds the tip of the
metasoma. Moreover, Mauss (2013) referred to a female of C. abbreviatus in his
collection with protruded mouthparts extending beyond the end of the metasoma
as well. Therefore C. Hermon does not distinctly differ from C. abbreviatus in the
character that mainly substantiated its description and was considered to be a junior
synonym of C. abbreviatus.

However, Mauss (2013) avoided formal synonymy of both taxa as the situation
appeared to be more complicated. The club of the antenna of the type of C. hermon
is completely orange, which is also the case in specimens of C. abbreviatus from the

Identity and distribution of Celonites hermon Gusenleitner, 2002... 7

Balkan and Aegean populations whereas the antennae of Central and West European
specimens of C. abbreviatus are much darker suggesting that the eastern populations
might belong to a different subspecies or even species that could be identical with C.
hermon. Subsequently Mauss et al. (2016) demonstrated that Celonites tauricus differs
distinctly from C. abbreviatus in that the likewise orange antennal club of C. tauricus
has always a dark tip, which is possibly related to different male signals during mat-
ing behaviour. Since the tip of the antennal club of the holotype of Celonites hermon
is completely orange, it can not be identical with C. tauricus. The distribution area
of Celonites abbreviatus extends from Northwest Africa over the Iberian Peninsula,
Southern and Central Europe eastwards to the Balkan and the western Aegean. Ce-
lonites tauricus is allopatrically distributed with regard to Celonites abbreviatus and
has been recorded from the Crimea, Kos, Rhodes, Cyprus and Asia Minor (Mauss
et al. 2016, Mauss unpubl.) (Fig. 11), separating the locus typicus of C. hermon by
more than 1000 km from the eastern margin of the distribution range of C. abbre-
viatus. Therefore C. hermon could be hypothesized to represent a relict population
of C. abbreviatus in the Golan Mountains that was separated from the Balkan and
Aegean populations of C. abbreviatus as a result of range expansion of C. tauricus. An
alternative hypothesis is that Celonites hermon is an indigenous biospecies. In this case
one would expect closer relationship to other members of the C. abbreviatus-complex
from geographically adjacent areas in the Middle East, that is Celonites tauricus or
Celonites andreasmuelleri.

Based on new material of Celonites hermon, the aim of this study is to reinvestigate
the status of the taxon carefully, to characterize its taxonomic position within the genus
Celonites and to describe the newly discovered male.

Material and methods

The specimens of Celonites studied belong to the public collection of the Upper Aus-
trian State Museum (Linz, Austria) (OLML), as well as to the private collection of J.
Gusenleitner (Linz, Austria) (JG). The specimens were investigated under a WILD
M3 stereo microscope (maximum magnification 80 times). Measurements of the
exoskeleton were made using an ocular micrometer (highest resolution 0.011 mm).
Distances between the ocelli, the compound eyes and the mesonotum width were
measured according to Eck (1978). The genitalia of all males were extracted after
re-softening the specimens and were then studied in 80% ethanol or 85% glycerol.
Nomenclature of male genitalia follows that of Birket-Smith (1981). Drawings were
made with a drawing tube (WILD Type 308700). Micro-photos were taken with a
Leica IC 80 HD camera mounted on a Leica MS 5 stereomicroscope. Multifocus-
pictures were generated with Leica Application Suite (LAS) software. The scapus of
the antenna is referred to as antennal article Al and the pedicellus as A2, the flagellum
consists of the articles A3—A12.

58 Volker Mauss & Rainer Prosi / Journal of Hymenoptera Research 66: 55—70 (2018)

Systematics

Celonites hermon Gusenleitner, 2002

Figs 1-9

Material studied. Israel: Mt. Hermon, 33.416111N, 35.857500E, 1900 m a.s.L.,
17.07.1971, 12. (holotype, dbM No. 3579), leg. Bytinski-Salz, coll. JG. Syria: 40
km NE of Damascus, 33.416667N, 35.866667E, 13.05.1996, 499 (dbM No. 3684,
3685, 3686, 3687), 23. ¢ (dbM No. 3688, 3689), leg. Mi. Halada, coll. OLML; West
Syria, Tartus env., 34.900000N, 35.883333E, 25.05.1996, 222 (dbM No. 3691,
3692), leg. Mi. Halada, coll. OLML.

Diagnosis. Pale markings light yellowish to yellow without reddish tinges (Fig. 1).
Immediate tip of antennal club (A8—12) and especially ventral side of A12 orange to
orange-brown, not markedly contrasting in colour from adjacent light ventral side of
antennal club. Axilla of mesoscutellum with short blunt lateral projection that only
slightly projects over adjacent posterior part of tegula. Frons and clypeus covered with
pale, stiff pollen collecting setae (Fig. 3). In females most, in males at least some of
these setae with tiny spherical enlargement at tip (“knob”). Diameter of median ocel-
lus distinctly larger than diameter of lateral ocelli, ocelli larger than in C. abbreviatus
and C. tauricus. Median ocellus circular, lateral ocelli more or less obliquely oval
(Figs 4, 7). Vertex falling away obliquely towards the occipital carina immediately
behind ocelli (Figs 4, 7).

Females with head in front view more elongated in relation to its length. Clypeus
with lateral margins more continuously rounded leading to elongated appearance
(Fig. 2). Cuticula on frons and at least dorsal half of clypeus densely and strongly sha-
egreened leading to a conspicuous dull region covered with knobbed pollen-collecting
setae, changing gradually into the adjacent smooth, shiny cuticula on vertex and
less densely shagreened cuticula ventrally on clypeus (Figs 2, 4). Knobbed pollen
collecting setae on frons distinctly longer and relatively thinner in comparison with
Celonites abbreviatus and C. tauricus, up to 1.4 times as long as diameter of median
ocellus, with diameter almost equal along most of length, only weakly broadened
towards base (Figs 3, 4).

Males with only two oval-shaped tyloids situated ventrally on articles A9 and A10
of club-shaped antennae. Midcoxa with a small but distinct spine at distal end, on
anterior side of midcoxa close to its antero-medial angle. Tergum VII markedly lobed
posteriorly (Fig. 6). Sternum VIII only shallowly emarginate, surface not much sunk
in towards emargination (Fig. 8). Tergum VII with medial lobe distinctly emargin-
ate leading to quadrilobed appearance of posterior margin (Fig. 6). Emargination of
posterior margin of sternum VII medially angled, laterally with a transverse section
well set off by protruded edge with distinct little spine on each side of posterior mar-
gin (Fig. 8). Posterior margin of stipes of male genitalia with dorso-medial concavity
(Figs 9, 10). Stipes in dorsal view elongated, with medial and lateral margin running
more or less parallel towards posterior end (Figs 9, 10). Posterior process of volsella

Identity and distribution of Celonites hermon Gusenleitner, 2002... 59

female

Figure |. Imagines of Celonites hermon in lateral view (female dbM No. 3692, male dbM No. 3689).

conically rounded, proximally continuing into shaft of volsella. Medial process of vol-
sella set off distinctly from shaft by longitudinal fold (Figs 9, 10).

Description of the male. Colour (Fig. 1): Black, changing into blackish-brown
on strongly sclerotized parts of labio-maxillary complex, sterna, coxae, trochanters
and femora. The following are light yellowish: large basal spot on mandible; labrum;
central area of clypeus except small lateral and ventral margin; broad parallel-sided
band at front of frons, shortly interrupted medially on the supra-antennal area, later-
ally extending obliquely towards upper inner margin of eye where it terminates with
straight truncated end; small spot on ocular sinus; narrow streak on tempora along
occipital carina at dorso-lateral corner of head; large spot on antero-dorsal angle of
pronotum (humeral spot); stripe along dorso-medial (inner) margin of pronotum,
posteriorly small becoming distinctly broader antero-medially where it continues
into humeral spot in one specimen (Fig. 7); large spot on dorsal mesopleura; tegula
except small black marking along anterior part of medial margin; medium-sized me-
dian spot on scutellum; propodeal lamella; distal part of femora, tibiae and tarsi
(weakly mixed with reddish-brown in parts especially distally); continuous posterior
bands on terga I-VI, somewhat smaller on each side of middle (Fig. 6), extending
over lateral margin on ventral part of tergum; continuous broad posterior band on
sternum II; small spots postero-laterally on sternum III. Reddish-brown are: distal
two thirds of mandible; ventral margin of clypeus; claws. Antenna with Al—2 black;
A3-6 anteriorly with light yellowish stripe otherwise blackish to dark orange-brown;
A7-A12 orange-brown, posteriorly with blackish marking becoming lighter distally,
anteriorly and on ventral side markedly orange without black. Wings translucent,
weakly yellowish-brown.

Structure: Head in front view as long as broad. Clypeus a little broader than long,
distal margin deeply emarginate; cuticula shiny and smooth with close macropuncta-
tion becoming less distinct towards distal margin; pale stiff setae arising from macro-
punctures without distal “knob”; lateral setae with distal ends frequently curved in dis-
tal-medial direction. Frons diagonally striated with moderately spaced to close macro-
punctation, interstices smooth; bearing pale stiff setae, up to 0.22 mm long, 1.1 times
as long as diameter of median ocellus, laterally some with curved distal end, in centre
mainly with tiny distal “knob”; frontal line in one specimen raised to form a small pro-

Volker Mauss & Rainer Prosi / Journal of Hymenoptera Research 66: 55—70 (2018)

Figures 2—6. Head of female of Celonites hermon (dbM No. 3685) 2 in frontal view 3 in lateral view show-

ing pollen collecting apparatus with “knobbed” setae on frons and clypeus 4 in dorsal view 5—6 Metasoma

of Celonites hermon in dorsal view 5 female (dbM No. 3685) 6 male (dbM No. 3689).

Identity and distribution of Celonites hermon Gusenleitner, 2002... 61

0.5 mm

Figures 7—9. Head of male Celonites hermon in dorsal view (dbM No. 3456) 8 Metasomal sternum VIII
of male of C. fermon in ventral view (dbM No. 3689) 9 Male genital of Celonites hermon (dbM No. 3689)

a in dorsal b ventral view.

tuberance in centre of frons. Median ocellus circular, lateral ocelli more or less obliquely
oval; diameter of median ocellus distinctly larger than diameter of lateral ocelli. Vertex
falling away obliquely towards occipital carina immediately behind ocelli (Fig. 7); cu-
ticula with close macropunctation becoming more distinct and closely reticulate behind
ocelli, where longitudinal interstices are more strongly raised forming lines; cuticula of
interstices shiny and smooth; covered with short setae arising from macropunctures.
Compound eye sparsely covered with small setae. Gena very narrow; preoccipital carina
sharp. Antenna with articles A8-A12 forming ventrally flattened club about 2.0 times as
long as broad (viewed dorsally) (Fig. 1); two oval-shaped, perhaps sensory, depressions
(tyloids) on concave, ventral side of club, situated within antennal articles A9 and A10.

62 Volker Mauss & Rainer Prosi / Journal of Hymenoptera Research 66: 55—70 (2018)

Anterior margin of pronotum raised to carina especially sharp medially (erroneously
termed anterior pronotal carina by Mauss 2013). Short anterior pronotal carina (sensu
Carpenter 1988) distinctly present at antero-ventral angle of pronotum running parallel
to anterior margin, preceding crenate groove; distance between anterior pronotal carina
and anterior margin of pronotum about width of fore metatarsus. Posterior pronotal
carina forms narrow translucent sinuate crest on humeral angle of pronotum. Posterior
margin of pronotum raised to short carina dorsally in front of tegula. Cuticula of prono-
tum shiny, with close to reticulate macropunctation, interstices smooth, rounded, with
very sparse uneven micropunctation; laterally fairly horizontally striated due to slightly
more raised interstices. Cuticula of mesoscutum and mesoscutellum shiny, reticulate
with close, deep macropunctation and raised interstices. Postero-medial cuticula of mes-
oscutum and especially of mesoscutellum with longitudinal interstices becoming nar-
rower and more strongly raised to sharp edges leading to distinctly striated appearance;
interstices smooth, laterally on mesoscutellum somewhat finely longitudinally wrinkled.
Mesoscutellum laterally with distinct carina along posterior margin, carina medially re-
duced to a few small tooth-like projections along posterior margin. Carina along poste-
rior margin of metanotum medially with small tooth-like projections. Axilla with short
blunt lateral projection only slightly projecting over adjacent, somewhat emarginate,
posterior part of tegula. Tegula shiny, closely covered by macropunctures except less
densely punctured or completely smooth central convex area. Mesepisternum with pro-
nounced epicnemial carina deflexed backwards to run transversely in front of mid coxa;
cuticula shiny, with close macropunctation; horizontally striated by raised interstices;
area ventral to scrobal groove coarsely punctured with some interstices strongly raised
to knife-like edges forming coarse honeycomb-like sculpture. Process at mesepisternal
scrobal groove of moderate size; cuticula on posterior side shiny in centre with irregularly
moderately spaced distinct micropunctation, ventrally densely longitudinally wrinkled.
Horizontal propodeal triangle laterally delimited by a perpendicular declivity, somewhat
laterally produced at postero-lateral edge of propodeal triangle, posteriorly bordered by
serrated carina; cuticula shiny, coarsely punctured, interstices almost knife-like. Posterior
surface of propodeum striated by strong vertical cuticula-folds; cuticula shiny, without
punctuation, weakly coriaceous and covered with short fine pale setae. Cuticula of sides
of propodeum and metepisternum shiny, densely horizontally wrinkled. Lateral lamella
broad and somewhat convex; lateral margin slightly convex; posterior margin straight,
not crenate; medially where lamella joins central part of propodeum with a rounded
emargination, ventro-medial edge of which produced to a small blunt protrusion; dorsal
cuticula of lamella shiny, with moderately spaced macropunctation, interstices medially
smooth, dorso-laterally shagreened. Midcoxa with small but distinct spine at distal end
on anterior side close to anterio-medial angle. Claws ventrally with small tooth.

Metasomal terga with dark anterior part continuing into posterior pale part by
slight declivity; postero-lateral corners slightly produced; posterior margin of tergum
I weakly crenulated, crenulation not produced into spines and not projecting over
smooth translucent posterior margin of tergum; posterior margin of terga II-VI crenu-
lated, crenulation produced into short partially posteriorly truncated spines, not or only
slightly projecting over smooth translucent lower posterior margin of terga (Fig. 6);

Identity and distribution of Celonites hermon Gusenleitner, 2002... 63

cuticula with silken sheen, densely covered with moderately coarse macropunctation,
diameter of punctures and distance between them fairly larger on posterior pale part of
tergum, with about seven macropunctures along median axis of tergum II]; single thin
seta arises from bottom of each macropuncture, only slightly protruding over rim of
puncture; interstices finely shagreened, moderately covered with very tiny, decumbent,
pale setae, all setae orientated towards caudal end. Posterior margin of tergum VII with
two lateral deep emarginations, resulting in two lateral lobes that continue laterally
into margins of tergum and single medial lobe divided into two lanceolate more or
less pointed medial projections of variable length (Fig. 6). Metasomal sternum I shiny,
finely shagreened, with tiny setae but without punctures. Sterna II-VI posteriorly
with broad stripe of asetose, translucent cuticula adjacent to posterior margin of more
strongly sclerotized cuticula; small sparse band of setae along posterior sclerotized mar-
gin somewhat projecting over anterior part of translucent stripe of cuticula; sclerotized
cuticula shiny, close to anterior margin faintly shagreened, on posterior half of sternum
II-VI medially and laterally with sparse punctuation of shallow micropunctures from
which short pale setae arise, on sternum II antero-laterally with a few small shallow
macropunctures from which short pale setae arise, on sternum HI—VI anteriorly with
moderate to sparse shallow macropunctation. Sternum VIII with distinct little spine
on each side of posterior margin; emargination of posterior margin medially angled,
laterally with a transverse section well set of by protruded edge (Fig. 8).

NK

S& Late |
i
}

ha

tl

i
AWA
ci

ec
N

sp

“at
ie 7

Si eee

cu

\— J

cu Zz

0.5 mm

Figure 10. Male genital of Celonites hermon (dbM No. 3689) in dorsal (left) and ventral view (right).
Setae only shown on one side of each drawing. Nomenclature follows that of Birket-Smith (1981): ad, ae-
doeagus; at, apodema thyrsos; cu, cupula; dc, dorso-medial concavity of the posterior margin of the stipes;
ha, harpide; mp, medial process of volsella; pp, posterior process of volsella; sl, shovel-like dorso-medial

lobe of harpide; sp, stipes; ty, thyrsos; vo, volsella.

64 Volker Mauss & Rainer Prosi / Journal of Hymenoptera Research 66: 55—70 (2018)

Male genitalia as in Figs 9-10. Dorso-medial shovel-like lobe of harpide large,
densely covered ventrally with long setae. Dorso-posterior margin of stipes with dor-
so-medial concavity angled. Stipes in dorsal view elongated, with medial and lateral
margin running more or less parallel towards posterior end. Sides of stipites converg-
ing continuously anteriorly towards cupula. Volsella moderately large and broad,
not reaching antero-medial margin of dorso-medial lobe of harpide; dorsal area with
strongly sclerotized, large, dark tubercles; distances between tubercles moderate; plane
medial process curved more or less hook-like towards posterior end, set off distinctly
from shaft by longitudinal fold; posterior process conically rounded, proximally con-
tinuing into shaft of volsella. Sides of thyrsoi more or less parallel only slightly con-
verging towards posterior (apical) end of aedoeagus. Each thyrsos continues anteriorly
into apodema thyrsos with a distinct outwardly directed curve. On ventral side cupula
medially protruded towards anterior end projecting over dorsal margin of cupula in
ventral view of genital capsule.

Measurements. Measurements of the exoskeleton are summarized in Table 1.

Floral association. Unknown.

Distribution. Celonites hermon is known only from three localities in the ranges
of the Lebanon and Anti-Lebanon Mountains along the Lebanese section of the Dead
Sea Transform (Fig. 11).

Table |. Measurements of the exoskeleton of imagines of Celonites hermon (x = median; min = minimum,
max = maximum; measurements were made with a Wild M3 stereomicroscope with maximum magnifica-

tion 80x, maximum accuracy 0.011 mm, all distances in mm).

Female Male
Parameter F :

x min max n x min max n
lateral ocelli distance 0.46 0.42 0.47 7 0.43 0.42 0.45 2
median/lateral ocellus distance 0.17 0.15 0.18 ys 0.15 0.15 0.15 2
compound eyes distance 1.28 1.22 1.36 7 1.16 1.16 1.16 2
Al length 0.18 0.17 0.20 i 0.19 0.19 0.19 2
A3 length 0.24 0.22 0.26 7 0.24 0.23 0.24 2
A3 width 0.11 0.11 0.11 7 0.13 0.13 0.13 2
A4—5 length 0.20 0.19 0.20 yf 0.21 0.21 0.21 2
A8-12 length 0.77 0.73 0.87 i 1.08 1.08 1.08 2
A8—12 width 0.40 0.37 0.42 7 0.53 0.52 0.55 2
antennal sockets distance 0.81 0.76 0.84 Ti 0.61 0.59 0.62 2
clypeus maximum width 1.11 1.04 1.16 7 0.99 0.98 1.00 2
clypeus apical width 0.52 0.51 0.61 f 0.46 0.44 0.48 2
clypeus length 0.90 0.84 1.38 7 0.79 0.76 0.81 2
mesonotum width 2.74 2.55 2.80 6 2.60 2.49 2.72 2
mesoscutum length 1.96 1.88 2.10 6 1.96 1.88 2.04 2
wing length 5.54 525 5.68 7 5.47 5.40 5.54 2
R+Sc length 2.94 2.81 3.13 7 2.88 2.85 2.90 2
number of hamuli 12 10 14 vA 10 10 10 2
femur I length 1.39 1.34 1.48 6 1.33 1.31 1.34 2
tibia I length 0.95 0.87 0.98 7 0.89 0.88 0.89 2
metatarsus I length 0.56 0.55 0.57 7 0.50 0.46 0.53 2
tergum I width 2.74 2.58 2.81 7 2.65 2.60 2.69 2
tergum I length 1.00 0.95 1.14 7 1.04 0.94 1.14 2
tergum II width 2.77 2.63 2.90 7 2.67 2.60 2.74 2
total length ve) 6.7 ED 7 73. 7.0 7.5 2

Identity and distribution of Celonites hermon Gusenleitner, 2002... 65

© Celonites andreasmuelleri. ©
© Celonites hermon

© Celonites tauricus

a U 2 Sp EA

Figure I 1. Geographic distribution of the species of the Celonites abbreviatus-complex occurring in
the Middle East. Records of Celonites andreasmuelleri taken from Mauss 2013 and Mauss unpublished,
records of Celonites tauricus according to Mauss et al. 2016 and Mauss unpublished.

Key to separate Celonites hermon from its next relatives C. abbreviatus, C. tauricus
and C. andreasmuelleri

For illustrations of the morphological characters of C. abbreviatus, C. tauricus and C.
andreasmuelleri see Mauss (2013) and Mauss et al. (2016).

1 | ey et og F2\ Ft PROM Ges A pcm le MEA IM an RU GR et OE MA RRR oT eas EE eR At at tI 2
= Ae ee Ne eS poe Sel, DLE. Ro OL BM ar Sl 5
2 Cuticula on frons with a dull, densely shagreened, semi-circular central area

covered with knobbed pollen-collecting setae, contrasting distinctly to the
adjacent smooth, shiny cuticula. Clypeus and vertex much more shiny. Terga
of metasoma with extensive yellowish-white and reddish markings...............
PEON ok RI Ce ENS A Celonites andreasmuelleri Mauss, 2013
_ Cuticula on frons and at least dorsal half of clypeus densely and strongly sha-
greened leading to a conspicuous dull region covered with knobbed pollen-
collecting setae, changing gradually into the adjacent smooth, shiny cuticula
on vertex (Fig. 2). Terga of metasoma with yellowish-white to yellow mark-
ings without reddish tinge (exception being made for C. abbreviatus from

66

Volker Mauss & Rainer Prosi / Journal of Hymenoptera Research 66: 55—70 (2018)

the Peloponnesus with small reddish markings close to the lateral margin of
SESE a9 ala et inn esti Maat tn tash ane cantata hoe kat taas ute ane Sareea 3
Pollen collecting knobbed setae on frons longer and thinner, up to 1.4 times
as long as diameter of median ocellus (Figs 3, 4). Ocelli larger, with diameter
of median ocellus distinctly larger than diameter of lateral ocelli (Fig. 4).
Lateral ocelli more or less obliquely oval (Fig. 4). Vertex falls away obliquely
towards the occipital carina immediately behind the ocelli (Fig. 4). Mesos-
cutum shiny with dense, moderately coarse punctures separated by rounded,
not knife-like interstices... Celonites hermon Gusenleitner, 2002
Pollen collecting knobbed setae on frons shorter and more robust, not more
than 1.0 times as long as diameter of median ocellus. Ocelli smaller, with me-
dian ocellus only slightly larger than lateral ocelli. Lateral ocelli more evenly
rounded. Vertex continues more or less horizontally behind ocelli towards
occipital carina. Mesoscutum less shiny, with dense, moderately coarse punc-
tures separated by knife-like interstices, especially anteriorly resulting in a
mMOLeCOMP-HKE Ap PeatkAaNeen.W eccesen ten cusecessvasroscecsnengucverge sated carte Satie 4.
Antennal club (A8—12) with dark tip, that is, at least distal end of A12 is
blackish markedly contrasting on ventral side to adjacent light reddish brown
area of antennal club. On dorsal side blackish marking extends usually over
distal parts of All fading gradually towards proximal end (for differences
between geographic populations see Mauss et al. 2016) oc. ceeeeeeeeeeeeeeeees
ee ee ane eco ntti Ne eee ee a Celonites tauricus Kostylev, 1935
Antennal club (A8-12) of individuals from Balkan populations completely
orange, in some dark coloured individuals from western populations club
becomes darker dorsally, while immediate tip and especially ventral side of
tema Me Huer e285 Mia elce ses Celonites abbreviatus (Villers, 1789)
Tergum VII trilobed. Posterior emargination of sternum VII medially bent
at an angle of about 135°. Laterally the posterior, angled margin of sternum
VUI continues directly into the lateral surface of the sternum without a
prominent edge or a distinct spine, so that even a small postero-lateral trans-
verse section is lacking. Genitalia as in Mauss 2013: fig. 26. Metasoma black
with extensive yellowish-white and reddish markings... ees eeeeseeseeeeeeees
ba i vB is Pi BPs dh bach «nA oateucl A 2 Celonites andreasmuelleri Mauss, 2013
Tergum VII quadrilobed, since the median lobe is medially distinctly emar-
ginate (Fig. 6). Sternum VIII different, with a distinct little spine on each side
of the posterior margin (Fig. 8). Genitalia different. Metasoma black with
extensive yellowish-white or yellow but without reddish markings (excep-
tion being made for C. abbreviatus from the Peloponnesus with small reddish
tines close-to. thre’ lateral margins Of theyerga)+.c.,,..resosesssspeesusoensees spnbrseat ones 6
Posterior margin of stipes of male genitalia with dorso-medial concavity
(Figs 9, 10). Stipes in dorsal view elongated, with medial and lateral margin
running more or less parallel towards posterior end (Figs 9, 10). Posterior
process of volsella conically rounded. Medial process of volsella set off dis-

Identity and distribution of Celonites hermon Gusenleitner, 2002... 67

tinctly from shaft by longitudinal fold (Figs 9, 10). Ocelli larger. Vertex fall-
ing away obliquely towards occipital carina immediately behind ocelli..........
wy Pa yet eh ab eae deh nba Noort Ei Pate Celonites hermon Gusenleitner, 2002
= Posterior margin of stipes of male genitalia straight, without dorso-medial con-
cavity. Stipes in dorsal view shorter, with medial and lateral margin diverging
continuously towards posterior end. Posterior process of volsella flat. Medial
process of volsella continues gradually into shaft. Ocelli smaller. Vertex run-
ning more or less horizontally towards occipital carina behind ocelli ............. 7
7 Antennal club (A8—12) orange, ventrally with distinct blackish marking on
A12 extending on distal parts of All well set off from orange of adjacent
ventral area, orange on dorso-posterior parts of A8—A12 can be darkened to
variable extend. Clypeus white to whitish-yellow. Emargination of sternum
VIII more evenly concave. Male genital broader. Volsella strongly sclerotized,
with a larger medial process ...........seee Celonites tauricus Kostylev, 1935
— Antennal club (A8-12) of individuals from Balkan populations completely
orange, in some dark coloured individuals from western populations club be-
comes darker dorsally, while immediate tip and especially ventral side of A12
remain lighter. Clypeus yellow, except in one rare variety from Switzerland
in which it is nearly white. Emargination of sternum VIII medially angled.
Genital less broad. Volsella less sclerotized, with a smaller medial process......
Rac reran elie, een Rccrae aa Ne Dai Wn Rate: Celonites abbreviatus (Villers, 1789)

Discussion

Status and taxonomic position of Celonites hermon

Celonites hermon can be assigned to the subgenus Ce/lonites s.str., the Celonites ab-
breviatus-group and finally the C. abbreviatus-complex without contradiction as it
shares all characters of these taxa established by Richards (1962) and Mauss (2013).
Among the members of the C. abbreviatus-complex Celonites hermon resembles C.
andreasmuelleri most of all in the comparatively thin pollen collecting knobbed se-
tae that are of equal diameter along most of their length, the enlarged ocelli, with a
distinctly large circular median ocellus and smaller oval shaped lateral ocelli and the
characteristic form of the head, that falls away obliquely towards the occipital carina
immediately behind the ocelli. Moreover, in the male genitalia the posterior margin
of the stipes has a characteristic dorso-medial concavity and the medial and lateral
margin of the stipes are running more or less parallel towards the posterior end.
These distinctive similarities are probably synapomorphic characters of both taxa, as
they are not present in other members of the C. abbreviatus-complex and even the
whole C. abbreviatus-group.

On the other hand, Celonites hermon is lacking some of the potential autapomor-
phies of C. andreasmuelleri recognized by Mauss (2013), i.e. the characteristic semi-

68 Volker Mauss & Rainer Prosi / Journal of Hymenoptera Research 66: 55—70 (2018)

circular cuticula structure on the frons of the females, as well as the differing form of
sternum VIII and tergum VII and the considerably larger shovel-like dorso-medial
lobe of the harpide in the males of C. andreasmuelleri. Moreover, both taxa differ in
colour, the colour pattern on the ventral side of the tip of the antennal club and the
form of the volsella. The distinct differences in the male genitalia, the form of sternum
VIII and tergum VI as well as the colour pattern on the ventral side of the antennal
club that has been assumed by Mauss et al. (2016) to be associated with mate recogni-
tion, lead to the conclusion that both taxa are probably reproductively isolated and
represent two separate biospecies (sensu Mayr 1967).

The hypotheses proposed by Mauss (2013) that Celonites hermon is identical with
C. abbreviatus, is rejected. Both species differ clearly in form and length of the pollen
collecting knobbed setae, the size of the ocelli and the shape of the vertex. Moreover,
there are distinct differences in the male genitalia. The misinterpretation of the mor-
phological characters of the type by Mauss (2013) resulted to some extend from the
mediocre condition of the female specimen, of which the setae on frons and vertex are
stuck together to a high degree by remnants of a viscous liquid, giving them a shorter
and stouter appearance and partly covering the exoskeleton.

Biogeography

Celonites hermon has been recorded only from a small area in the mountain ranges
along by both sides of the Lebanese section of the Dead Sea Transform (Fig. 11). Since
C. hermon differs significantly from C. abbreviatus in various morphological characters,
it cannot represent a relict population of C. abbreviatus as implied by Mauss (2013).
Moreover, the morphological similarities between C. hermon and C. andreasmuelleri
indicate a close relationship between them and a more distant relationship to C. ab-
breviatus and C. tauricus. Therefore, it can be hypothesized that C. hermon and C.
andreasmuelleri originated from a common stem species that inhabited the area along
the Dead Sea Transform. Geographic isolation and succeeding speciation may have led
to the recent allopatric distribution with C. hermon being restricted to the northern
Lebanese section of the Dead Sea Transform and C. andreasmuelleri to the southerly
adjacent area of the Jordan Rift Valley (Fig. 11).

Acknowledgements

Fritz Gusenleitner (Upper Austrian State Museum, Linz) and Josef Gusenleitner (Linz)
generously provided the investigated specimens of Celonites hermon. We are especially
grateful to Jim Carpenter (American Museum of Natural History, New York) for valu-
able comments on the manuscript and improvement of our English and Rainer Neu-
meyer and Andreas Miiller for their careful reviews.

Identity and distribution of Celonites hermon Gusenleitner, 2002... 69

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