Research Article |
Corresponding author: Volker Mauss ( volker.mauss@gmx.de ) Academic editor: Michael Ohl
© 2016 Volker Mauss, Alexander V. Fateryga, 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.
Citation:
Mauss V, Fateryga AV, Prosi R (2016) Taxonomy, distribution and bionomics of Celonites tauricus Kostylev, 1935, stat. n. (Hymenoptera, Vespidae, Masarinae). Journal of Hymenoptera Research 48: 33-66. https://doi.org/10.3897/JHR.48.6884
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Male and female of Celonites abbreviatus tauricus Kostylev, 1935 are redescribed and a neotype is designated. Based on morphological characters Celonites a. tauricus is synonymized with Celonites spinosus Gusenleitner, 1966 and Celonites abbreviatus invitus Gusenleitner, 1973. The taxon is hypothesized to be reproductively isolated from Celonites abbreviatus Villers, 1789 by differences in the male genitalia and in the colour pattern of the male antennae and is therefore regarded as a separate biospecies named Celonites tauricus. Celonites tauricus is allopatrically distributed with regard to C. abbreviatus and has been recorded from the Crimea, Kos, Asia Minor and Cyprus. Within this range six intraspecific taxa can be separated by morphological characters and colour patterns. Habitat, flower association, flower visiting behaviour, mate seeking behaviour and nesting of C. tauricus are almost similar to C. abbreviatus.
Hymenoptera , Vespidae , Masarinae , Celonites , taxonomy, bionomics, Palaearctic, Crimea, Asia Minor, Cyprus, flower association, Lamiaceae , female brood care
Nearly 100 years ago the first specimen of a species of the pollen wasp genus Celonites Latreille, 1802 was recorded from the Crimea. It was a single female collected by V.N. Wuczeticz on 13 June 1916 on the southern slope of Karagach in the Karadag (
In 1966 Gusenleitner described a new species belonging to the Celonites abbreviatus-complex from western Turkey, named Celonites spinosus. He argued that this taxon could not be identical with C. a. tauricus Kostylev, based on the incorrect translation or interpretation of Kostylev’s description, that antennal article A3 should not be longer than A4+A5 in C. a. tauricus. In fact, Kostylev attributed this character state explicitly to a specimen from Spain and stated that A3 is longer than A4+A5 in C. a. tauricus, which is similar to C. spinosus. Moreover, in both taxa the posterior margins of the metasomal terga II–V were described as “serrated” or “spine-like crenulated” respectively (
Another subspecies of Celonites abbreviatus named invitus was described by Gusenleitner in 1973 from central Turkey and Armenia. He separated this taxon from C. abbreviatus by the very shiny cuticula of vertex and mesosoma and the incomplete colour banding on the metasomal terga III–V. As a result of the investigation of further material, Gusenleitner in 1985 recognized C. a. invitus as an eastern form of C. spinosus and synonymized both taxa formally.
A specimen of Celonites abbreviatus tauricus was first collected from the Crimea in 1916, but none were recorded in the following 86 years. Then in 2002 two specimens were collected in the Karadag Nature Reserve (
The purpose of the study presented below was to revise the taxonomic status of Celonites abbreviatus tauricus, to designate a neotype, to describe both sexes and to summarize and discuss what is known of the distribution, geographical variation and bionomical characters of the taxon.
The specimens of Celonites studied belong to the public collections of the Taurida Academy of the Vernasdskiy Crimean Federal
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). The genitalia of all males were extracted after re-softening the specimens and were then studied in 80% ethanol. 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 A1 and the pedicellus as A2, the flagellum consists of the articles A3–A12.
Systematic bionomical observations were made on the Crimea population by A. Fateryga at Cape Aya in 2004, in the vicinity of Veseloye near Sudak in 2004, and in Lisya Bay from 2005 until 2014 (for details see material studied), and documented using a Canon Powershot A570 IS.
The Kos population was studied by V. Mauss from 25 May to 4 June 2015 (for details see material studied). Observations were made with a close-up binocular (Pentax Papilio 8.5×21) and documented with a Canon EOS 70D with a 180 mm macro-lens (scale up to 1:1, resolution 20 mega pixel) and macro flash-lights. Time intervals were measured using a digital stop-watch. Specimens of flowering plants were collected and preserved dried. The material was placed in the herbarium of the Stuttgart State
Celonites abbreviatus tauricus Kostylev, 1935, Arch. Mus. Zool. Univ. Moscou 2: 108 [Since the original type material is lost, we hereby designate a new specimen as a Neotype: 1♀ (dbM 4305), Crimea, Vicinity of Feodosiya: Lisya Bay, 44°54'N 35°09'E, leg. A. Fateryga 16.06.2010, coll.
Celonites spinosus Gusenleitner, 1966, Polskie Pismo Entomol. 36: 359–362, figs 2a, b, c [Holotype: 1♀ (dbM 4665), Turkey, Kusadasi, leg. J. Gusenleitner 11.06.1964, coll. JG]
Celonites abbreviatus invitus Gusenleitner, 1973, Boll. Mus. Civ. Stor. Nat. Venezia 24: 58, fig. 2b [Holotype: 1♀ (dbM 4662), Turkey, Gürün, 44°54'N, 35°09'E, leg. J. Gusenleitner 05.06.1970, coll. JG]
Celonites abbreviatus (Vill.) invitis Gusenl.;
Taxon 1 (Crimea, “tauricus”)
Russia: Crimea: Alushta, 44.66667°N, 34.41667°E, 06.07.1963, 1♀ IZAN; Simferopol, Marino, 44.91667°N, 34.11667°E, 31.07.2007 1♀ leg. A. Fateryga VTNU; Vicinity of Feodosiya: Karadag Nature Reserve, 44.91667°N, 35.21667°E, 07.07.2002 1♂ (dbM No. 4704) 1♀ (dbM No. 4702) leg. S. Ivanov VM; Vicinity of Feodosiya: Lisya Bay, 44.90000°N, 35.15000°E, 22.06.2003 1♀ (dbM No. 4306) leg. S. Ivanov VM 1♀ leg. S. Ivanov VTNU 1♀ leg. S. Ivanov IZAN, 06.07.2005 1♀ leg. A. Fateryga VTNU 1♀ leg. A. Fateryga IZAN, 13.06.2007 2♀♀ leg. A. Fateryga VTNU, 25.06.2009 1♀ leg. S. Ivanov VTNU 1♀ leg. S. Ivanov
Taxon 2 (Kos)
Greece: Dodekanes, Island Kos: SE Mastichari, 36.833333°N, 27.083333°E, 31.05.–01.06.2003 2♀♀ (dbM No. 3607, 3608) leg. J. Tiefenthaler
Taxon 3 (Western Asia Minor, “spinosus”)
Turkey: Ankara: 15 km S Ankara, Beynam, 39.685958°N, 32.895143°E, 23.07.1987 1♀ (dbM No. 4682) leg. Kl. Reinhold JG; Kizilcahamam N of Ankara, 40.470123°N, 32.648873°E, 09.07.2000 3♂♂ (dbM No. 3597, 3598, 3599) leg. M. Halada
Intermediate between Taxon 3 and 4
Turkey: Agri: 40 km N Muradye, 39.246612°N, 43.67878°E, 2200 m, 05.07.2000 1♀ (dbM No. 3602) leg. M. Halada
Taxon 4 (Eastern Asia Minor, “invitus”)
Iran: Elburs: 50 km S (90 km Straße) Chalus, 36.326699°N, 51.360601°E, 2800 m, 26.07.1977 1♂ (dbM No. 3656) 7♀♀ (dbM No. 3649, 3650, 3651, 3652, 3653, 3654, 3655) leg. J. Gusenleitner JG; 75 km S Chalus, 36.142606°N, 51.446689°E, 2400 m, 13.07.1977 2♀♀ (dbM No. 3648, 4664) leg. J. Gusenleitner JG.
Turkey: Adiyaman: Karadut, Nemrut Dag, 37.980373°N, 38.747183°E, 09.06.1998 1♂ (dbM No. 3430) leg. M. Halada
Intermediate between Taxon 4 and 5
Turkey: Bitlis: Nemrut Dag, 38.650366°N, 42.218203°E, 2850 m, 08.08.1986 1♀ (dbM No. 3627) leg. K. Warncke JG, 2300 m, 15.08.1991 1♀ (dbM No. 3628) leg. K. Warncke JG; Nemrut Dagi, 38.650366°N, 42.218203°E, 2800 m, 07.08.1986 1♀ (dbM No. 3626) leg. Blank JG; Tatvan, Nemrut Mt. 38.644449°N, 42.214363°E, 2000 m, 23.07.2003 1♀ (dbM No. 4544) leg. H. Özbek EY. – Hakkari: S Varegös / Mt. Sat, 37.3333°N, 44.25°E, 1700 m, 04.08.1982 1♀ (dbM No. 3633) leg. K. Warncke JG.
Taxon 5 (South-east of Taurus range)
Syria: North Syria: Marbij [Manbij], 36.528792°N, 37.935693°E, 09.05.1996 1♀ (dbM No. 3690) leg. M. Halada
Turkey: Icel: Mut, 36.6464°N, 33.4375°E, 13.06.1965 1♀ (dbM No. 4668) leg. M. Schwarz JG, 27–30.05.1967 1♀ (dbM No. 4672) leg. J. Schmidt JG, 28.05.1967 1♂ (dbM No. 4690) 1♀ (dbM No. 4673) leg. J. Gusenleitner JG, 06.06.1968 1♀ (dbM No. 4679) leg. J. Gusenleitner JG 1♀ (dbM No. 4680) leg. J. Schmidt JG, 19.05.1970 1♂ (dbM No. 4691) leg. J. Gusenleitner JG. – Şanliurfa: Birecik / Urfa, 37.066514°N, 38.104280°E, 22.05.1983 1♂ (dbM No. 3635) leg. K. Warncke JG; Urfa, 37.165902°N, 38.795883°E, 31.05.1968 1♀ (dbM No. 4669) leg. J. Gusenleitner JG; Urfa: Urfa Umgebung, 37.165902°N, 38.795883°E, 30.05.1978 1♂ (dbM 4692) 1♀ (dbM No. 4671) leg. M. Schwarz JG.
Taxon 6 (Cyprus)
Cyprus: Troodos: Mt. Olympos, 34.93°N, 32.86°E, 1900 m, 19.06.2013 7♀♀ (dbM No. 4875, 4876, 4877, 4878, 4879, 4880, 4881) leg. C. Saure CS, 20.06.2013 6♂♂ (dbM No. 4719, 4720, 4721, 4722, 4723, 4724) 4♀♀ (dbM No. 4715, 4716, 4717, 4718) leg. C. Schmid-Egger CSE.
Armenia: Erevan, Monti desertici, Aighpat 40 km SE, 23.07.1963 1♂ 1♀ leg. Giordani Soika coll. Giordani Soika det. J. Gusenleitner,
Russia: Crimea: Echkidag, 44.89667°N, 35.12167°E, 07.06.2014 field obs. A. Fateryga; Vicinity of Feodosiya: Lisya Bay, 44.90200°N, 35.15805°E, 02.07.2011 nest record A. Fateryga, 27.06.2013 28.06.2013 11.07.2013 10.06.2014 all field obs. A. Fateryga; Vicinity of Sevastopol: Cape Aya, 44.42367°N, 33.66133°E, 05.07.2004 field obs. A. Fateryga; Vicinity of Veseloye near Sudak, 44.86667°N, 34.88333°E, 02.07.2004 nest record D. Puzanov det. A. Fateryga [all Taxon 1].
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, about as long as diameter of median ocellus. In females most of these setae with tiny spherical enlargement at tip (“knob”), in males setae with distal enlargement only present in centre of frons. Cuticula of frons and clypeus dull and densely shagreened. Males with only two oval-shaped tyloids situated ventral on articles A9 and A10 of club-shaped antennae and small spine anterior at distal end of midcoxa. Posterior margin of metasomal tergum VII divided into four lobes.
Separated from Celonites abbreviatus by distinctly different colouration of club of antenna: Club has a 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 A11 fading gradually towards proximal end. In C. abbreviatus club 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 A12 remain lighter. Male genital broader than in C. abbreviatus, in dorsal view transverse width of each stipes larger than distance between dorso-medial margins of stipites. Medial process of volsella larger than in C. abbreviatus.
Female. Colour: Black. The following are weakly yellowish white (Fig.
Specimens of Celonites tauricus from populations from different geographical regions in lateral view (insect pins and in a few specimens also protruded proboscis were retouched with photo software to improve comparability of the pictures), a Crimea female dbM No. 4305 male dbM No. 4307, Kos female dbM No. 4776 male dbM No. 4790, Western Asia Minor female dbM No. 3585 male dbM No. 3586; b Eastern Asia Minor female dbM No. 3629 male dbM No. 3430, South-east of Taurus range female dbM No. 4671 male dbM No. 4690, Cyprus female dbM No. 4717 male dbM No. 4720.
Structure: Head in front view slightly longer than broad. Clypeus a little broader than long; dull with somewhat sparsely weak macropunctation; densely shagreened; covered with pale, stiff setae arising from weak macropunctures; setae about as long as diameter of median ocellus, with tiny spherical enlargement at tip (“knob”) (Fig.
Anterior margin of pronotum raised to carina distinctly present along anterior margin of pronotum, especially sharp medially (erroneously termed anterior pronotal carina by
Mesepisternum with pronounced 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 faintly shiny, finely but densely shagreened with irregularly moderately spaced micropunctation. Basal horizontal propodeal triangle laterally delimited by a perpendicular declivity, somewhat laterally produced at postero-lateral edge, 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, weakly coriaceous and covered with fine pale setae. Cuticula of sides of propodeum and metepisternum shiny, densely horizontally wrinkled. Lateral lamella broad, somewhat convex; lateral margin almost straight; 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 weakly wrinkled. Claws ventral with small tooth.
Metasomal terga with posterior two-fifth separated from anterior part by slight declivity especially laterally (Fig.
Metasomal sternum I shiny, finely shagreened, with tiny setae but without punctures. Sterna II–V posteriorly with broad stripe of asetose, translucent cuticula adjacent to posterior margin of more strongly sclerotized cuticula; small sparse band of setae along posterior sclerotized margin somewhat projecting over anterior part of translucent strip of cuticula; sclerotized cuticula shiny, finely shagreened, on posterior half of sternum II–V with dense to moderate punctation of shallow micropunctures from which short pale setae arise, becoming sparser anteriorly, on sternum II antero-laterally with a few shallow macropunctures, on sternum III–V anteriorly with moderate to sparse shallow macropunctation. Sternum VI tapering towards distal end; with outer margin raised to bulged rim, posteriorly protruded into little blunt spine; cuticula with rather narrow smooth mid-line slightly raised to weak keel at distal end, at sides with strong macropunctures from which pale setae arise.
Male. Colour: Resembles female, except as follows (Fig.
Structure: Resembles female, except as follows. Clypeus longer than broad, distal margin deeply emarginated; cuticula shiny, with moderately spaced, shallow macropunctation, interstices on apical third weakly shagreened becoming smooth distally; pale stiff setae arising from macropunctures without distal “knob”, distal ends of setae frequently curved in distal-medial direction. Frons with moderately spaced macropunctation; interstices densely obliquely shagreened; bearing pale stiff setae, mainly with curved distal end, in centre few with distal “knob” (Fig.
Male genitalia as in Figs
Male genitalia of C. tauricus (dbM No. 4307) in dorsal (left) and ventral view (right). (Setae only shown on one side of each drawing; Nomenclature follows that of
Measurements. Measurements of the exoskeleton are listed in Table
Measurements of the exoskeleton of imagines of Celonites tauricus from the Crimea (x = median; min = minimum, max = maximum; measurements were made with a Wild M3 stereomicroscope with maximum magnification 80×, maximum accuracy 0.011 mm, all distances in mm).
Parameter | Female | Male | ||||||
---|---|---|---|---|---|---|---|---|
x | min | max | n | x | min | max | n | |
lateral ocelli distance | 0.42 | 0.40 | 0.45 | 5 | 0.39 | 0.32 | 0.41 | 5 |
front./lat. ocellus distance | 0.17 | 0.14 | 0.17 | 5 | 0.14 | 0.11 | 0.14 | 5 |
compound eyes distance | 1.28 | 1.17 | 1.33 | 5 | 1.06 | 0.92 | 1.10 | 5 |
A1 length | 0.20 | 0.19 | 0.21 | 5 | 0.19 | 0.18 | 0.20 | 5 |
A3 length | 0.30 | 0.25 | 0.31 | 5 | 0.23 | 0.21 | 0.25 | 5 |
A3 width | 0.12 | 0.10 | 0.12 | 5 | 0.11 | 0.11 | 0.13 | 5 |
A4-5 length | 0.14 | 0.11 | 0.21 | 5 | 0.19 | 0.17 | 0.21 | 5 |
A8-12 length | 0.79 | 0.77 | 0.86 | 5 | 0.97 | 0.92 | 1.05 | 5 |
A8-12 width | 0.37 | 0.35 | 0.39 | 5 | 0.47 | 0.44 | 0.51 | 5 |
antennal sockets distance | 0.74 | 0.66 | 0.76 | 5 | 0.50 | 0.47 | 0.53 | 5 |
clypeus max. width | 1.12 | 1.03 | 1.14 | 5 | 0.84 | 0.80 | 0.88 | 5 |
clypeus apical width | 0.48 | 0.45 | 0.52 | 5 | 0.42 | 0.39 | 0.44 | 5 |
clypeus length | 0.95 | 0.85 | 0.97 | 5 | 0.73 | 0.68 | 0.76 | 5 |
mesonotum width | 2.81 | 2.55 | 2.90 | 5 | 2.32 | 2.16 | 2.46 | 5 |
mesoscutum length | 2.07 | 1.82 | 2.13 | 5 | 1.51 | 1.41 | 1.68 | 5 |
wing length | 5.61 | 5.11 | 5.68 | 5 | 4.90 | 4.83 | 5.04 | 3 |
R+Sc length | 3.13 | 2.85 | 3.22 | 5 | 2.60 | 1.71 | 2.80 | 5 |
number of hamuli | 10 | 8 | 12 | 5 | 10 | 8 | 10 | 5 |
femur I length | 1.50 | 1.41 | 1.55 | 5 | 1.19 | 1.14 | 1.26 | 5 |
tibia I length | 0.97 | 0.96 | 1.01 | 5 | 0.84 | 0.78 | 0.88 | 5 |
metatarsus I length | 0.58 | 0.54 | 0.61 | 5 | 0.45 | 0.41 | 0.46 | 5 |
tergum I width | 2.85 | 2.55 | 2.99 | 4 | 2.44 | 2.30 | 2.60 | 5 |
tergum I length | 1.26 | 1.12 | 1.31 | 5 | 0.99 | 0.90 | 1.07 | 5 |
tergum II width | 2.92 | 2.66 | 2.99 | 4 | 2.35 | 2.30 | 2.55 | 5 |
total length | 7.8 | 7.1 | 8.3 | 5 | 7.5 | 6.8 | 7.8 | 4 |
The geographic range of Celonites tauricus is shown in Fig.
Comparison of the morphological characters of six intraspecific taxa of Celonites tauricus from different geographic regions of its range.
Taxon No. | 1 Crimea | 2 Kos | 3 Western Asia Minor | 4 Eastern Asia Minor | 5 SE of Taurus range | 6 Cyprus |
---|---|---|---|---|---|---|
antenna | tip dark, at least A12 with distal end blackish, on ventral side markedly contrasting to adjacent light reddish brown proximal area of antennal club, on dorsal side fading more gradually into reddish brown colour of proximal area of club | tip dark, at least A12 with distal end blackish, on ventral side markedly contrasting to adjacent light reddish brown proximal area of antennal club, on dorsal side fading more gradually into reddish brown colour of proximal area of club | tip dark, at least A12 with distal end blackish, on ventral side markedly contrasting to adjacent light reddish brown proximal area of antennal club, on dorsal side fading more gradually into reddish brown colour of proximal area of club; four specimen with dorsal side dark reddish brown | tip dark, at least A12 with distal end blackish, on ventral side markedly contrasting to adjacent light reddish brown proximal area of antennal club, on dorsal side fading more gradually into reddish brown colour of proximal area of club; one specimen with dorsal side dark reddish brown | tip dark, at least A12 with distal end blackish, on ventral side markedly contrasting to adjacent light reddish brown proximal area of antennal club, on dorsal side fading more gradually into reddish brown colour of proximal area of club | tip dark, at least A12 with distal end blackish, on ventral side markedly contrasting to adjacent light proximal area of antennal club, on dorsal side fading gradually into dark reddish brown to dark brown colour of proximal area of club |
shade of light colour | faintly yellowish-white | faintly yellowish-white | faintly yellowish-white; in two specimens light yellowish | light yellowish to yellow | light yellowish to yellow | light yellow to yellow |
clypeus (female) | black; one specimen with two tiny spots | black | black | black | black with dorsal spot, that may reach the dorsal margin and can expand on ventral half of clypeus, at minimum little isolated spot | black |
tempora | short stripe to completely black | short stripe | short stripe to completely black | long stripe to completely black | long stripe | long to short stripe |
tegula | reddish brown | reddish brown | reddish brown | light yellowish to yellow with small reddish brown translucent centre or reddish brown | light yellowish to yellow; in two specimens with small reddish brown translucent centre | light yellowish to yellow with small reddish brown translucent centre |
pronotum | narrow stripe along dorsal margin, brownish interrupted in the middle of dorso-lateral margin | narrow stripe along dorsal margin, blackish interrupted in the middle of dorso-lateral margin; in one specimen brownish interrupted | narrow stripe along dorsal margin, brownish or blackish interrupted in the middle of dorso-lateral margin | continuous stripe along dorsal margin; in three specimens brownish interrupted in the middle of dorso-lateral margin | continuous stripe along dorsal margin; in two specimens brownish interrupted in the middle of dorso-lateral margin | broad continuous stripe along dorsal margin |
scutellum | medium-sized medial spot | medium-sized medial spot | medium-sized to small medial spot | medium-sized or large laterally expanding medial spot; in one specimen completely black | large laterally expanding or medium-sized medial spot | medium-sized medial spot |
metasomal terga | continuous posterior band on tergum I, posterior band on terga II–V laterally interrupted; in two specimens tergum II with continuous band | posterior band on terga I–V laterally interrupted; four specimens with continuous band on tergum I | continuous posterior band on tergum I, posterior band on terga II–V laterally interrupted; in two specimens tergum II with continuous band; in seven specimens band on tergum I interrupted; in one specimen band on tergum II continuous; in six specimens posterior band on tergum V reduced to medial stripe; in three specimens posterior band on terga III–V reduced to medial stripe | broad continuous posterior band on terga I–V; in three specimens on tergum V laterally interrupted; in three specimens on terga IV–V laterally interrupted | broad continuous posterior band on terga I–V; in two specimens on tergum V laterally interrupted; in two specimens on terga III–V laterally interrupted | broad continuous posterior band on tergum I–V; in one specimen on tergum V laterally interrupted |
metasomal tergum VI (female) | black | black | black | black | black, basally with narrow lateral stripe; in one specimen completely black | black, basally with narrow lateral stripe; in one specimen completely black |
legs | reddish brown | reddish brown | reddish brown | reddish brown | yellowish | yellowish |
size, physique | large, strong | large, strong | large, very strong to strong | medium-sized to small, more dainty | large, strong | small, more dainty |
macropunctation of terga II–IV | coarse | coarse | coarse (in specimens from Antalya very coarse) | moderate | very coarse to coarse | moderate |
Σ macropunctures along visible part of sagittal axis of tergum III | 10 | 8–10 | 8–10 | 10 | 7–8 | 10 |
sculpture of posterior margin of tergum II–IV | crenate, crenulation produced into distinct spines raised at an angle of approximately 30° distinctly projecting beyond the translucent lower posterior margin of terga (spines approximately as long as diameter of macropunctures) | crenate, crenulation consists of short ± pointed horizontal processes, posteriorly not projecting beyond the translucent lower posterior margin; in one specimen a few processes on tergum III projecting beyond posterior margin | crenate, crenulation produced into distinct spines raised at an angle of approximately 30° distinctly projecting beyond the translucent lower posterior margin of terga (spines approximately as long as diameter of macropunctures); in seven specimens spines projecting only slightly beyond posterior margin | crenate, crenulation consists of very short horizontal processes, posteriorly truncated not projecting over the translucent lower posterior margin so that the translucent margin remains completely exposed | crenate, crenulation produced into distinct spines raised at an angle of approximately 30° distinctly projecting beyond the translucent lower posterior margin of terga (spines approximately as long as diameter of macropunctures); in one specimens spines projecting only moderately beyond posterior margin | crenate, at least laterally crenulation consists of short horizontal ± pointed processes, posteriorly not projecting beyond the translucent lower posterior margin, medially processes posteriorly truncate to variable extend |
Individuals from western Asia Minor are large and strongly built (Fig.
Eight out of nine localities of Celonites tauricus in the Crimea are situated in the habitat zone of submediterranean vegetation of the south coast (Fig.
Bionomics of Celonites tauricus in the Crimea 15 Habitat 16 Female visiting flower of Teucrium chamaedrys 17 Female brushing pollen from the frons towards the mouthparts by alternating movements of the fore legs subsequent to regular flower visit at Teucrium chamaedrys 18 Nest under construction on the underside of a stone (brood cell on the left removed) 19 Old nest attached to the underside of a stone with emergence holes of Celonites imagines in the walls of three brood cells.
In Kos Celonites tauricus was found in areas from 20 to 110 m above sea level that were covered with phrygana vegetation characterized by larger patches of flowering plants of Satureja thymbra (Fig.
Bionomics of Celonites tauricus in Kos 20 Habitat (qs = potential quarry site of females) 21 Male perching on stone 22 Male visiting flower of Satureja thymbra 23 Female standing on the ground near a plant of Satureja thymbra brushing with the fore legs over her frons 24 Male visiting flower of Thymus capitatus (p = protruded proboscis).
Flower visiting records of Celonites tauricus are summarized in Table
Flower-visiting records of males and females of Celonites tauricus from different geographic regions.
plant taxon | Σ sightings of flower visiting individuals | |||||
---|---|---|---|---|---|---|
Crimea | Palandöken | Kos | ||||
♀ | ♂ | ♀ | ♂ | ♀ | ♂ | |
Lamiaceae | ||||||
Satureja thymbra L. | 52 | 11 | ||||
Teucrium chamaedrys L. | 35 | 10 | ||||
Teucrium polium L. | 1 | 2 | ||||
Thymus capitatus (L.) Hoffmanns. & Link | 1 | 6 | ||||
Thymus tauricus Klokov & Des.-Shost. | 8 | |||||
Ziziphora clinopodioides Lam. | 1 |
|||||
other plant taxa |
During flower visits at Satureja thymbra and Teucrium chamaedrys the females always stood on the lower lip of a flower and took up nectar and pollen simultaneously. The proboscis was protruded deeply into the corolla tube while the female performed at a high frequency slight back and forth movements of the anterior parts of her body, rubbing her head over the nototribic anthers (Figs
Flower visiting behaviour of Celonites tauricus at Satureja thymbra in Kos 25–26 Regular flower visit for the simultaneous uptake of nectar and pollen with the proboscis protruded into the corolla tube and the knobbed setae on the frons making contact with the nototribic anthers 25 Viewed from ventral-posterior 26 Viewed from dorso-lateral 27 Proboscis partly retracted shortly before leaving the flower, in lateral view 28–29 Female standing on the lower lip and an adjacent flower brushing pollen from her frons with alternating movements of her fore legs 30 The fore legs are brought between the mouthparts for pollen ingestion during pollen transfer from the frons by brushing movements towards the mouthparts.
On the Crimea, individual females of C. tauricus usually visited flowers over a period of 20–40 minutes. After about 10–15 minutes they regularly interrupted flower visiting and alighted on stones or grass. They remained there for several minutes repeatedly regurgitating and withdrawing again a mass of pollen and nectar that became visible as a drop of liquid between the mouthparts. This behaviour probably served to thicken the pollen and nectar mass.
During flower visits males always inserted their proboscis into the corolla tube (Figs
More than 99% of the pollen from the crops of females from Kos consisted of hexacolpat pollen grains of Lamiaceae. This pollen type occurs in several genera of Lamiaceae including Satureja and Thymus.
Three nests of Celonites tauricus were recorded in the Crimea. The first nest was investigated on 2 July 2004 in the vicinity of the village of Veseloye near Sudak. The nest site was a rocky slope with steppe vegetation predominated by Melica taurica K. Koch and Teucrium chamaedrys along with individual trees of Celtis glabrata Steven ex Planch. The nest was located in a small cavity underneath a stone situated in the shadow of one of the Celtis trees. The nest under construction contained two brood cells. The cells were placed on the underside of the stone and were made of fine clayey soil with a few tiny stones (Fig.
The second nest was studied on 2 July 2011 in Lisya Bay. The nest site was a slope covered with phrygana vegetation predominated by Elytrigia caespitosa subsp. nodosa (Nevski) Tzvelev, Atraphaxis replicata Lam., Thymus tauricus and Teucrium chamaedrys. The nest was also situated in a small cavity on the underside of a small stone. The nest was old and contained three cells covered with an additional layer of mud forming a nest covering (Fig.
The third nest was found in Lisya Bay on 11 July 2013. It was also situated on the underside of a small stone and it was also old. The nest consisted only of a single cell covered with an additional layer of mud. The cell was sealed at the apical end and had a frontal opening at the apical part, which had probably been made by an emerging imago of C. tauricus. Inside the cell were a Celonites cocoon and meconium.
Females of C. tauricus were never observed at water collection sites. Therefore the mud used for cell construction was probably made by mixing clay particles with regurgitated nectar.
Males performed patrol flights along the preferred forage plants of the females, i.e. Teucrium chamaedrys in the Crimea as well as Satureja thymbra and Thymus capitatus in Kos, and over nearby stones in a low constant flight. Patrolling was regularly interrupted by perching in the immediate vicinity of the forage plants, mainly on stones but also on the ground or on the plants themselves (Fig.
The species is univoltine. In the Crimea males were observed from 5 June to 8 July and females from 7 June to 31 July indicating slight proterandry.
The rediscovered Celonites taxon from the Crimea, assigned to C. abbreviatus tauricus, can be consistently identified as a member of the Celonites abbreviatus-complex (sensu
Within the Celonites abbreviatus-complex the members of the taxon Celonites a. tauricus from the Crimea are in the main similar to individuals of Celonites spinosus from western Asia Minor in both the colouration and morphology of the exoskeleton and in the structure of the male genitalia. Because of this high degree of similarity it is hypothesized that both taxa are not reproductively isolated from each other and thus belong to the same biospecies (sensu
Celonites tauricus resembles Celonites abbreviatus in many characters and is therefore in all probability closely related to this species, as already established by
The areas inhabited by Celonites tauricus in the Crimea and in Kos are comparable to habitats of Celonites abbreviatus in Central Europe (
The observed distribution pattern of the intraspecific taxa of C. tauricus in Asia Minor (Fig.
It can be hypothesized that C. tauricus colonized the Crimea from the west of Asia Minor, since the Crimean population of Celonites tauricus is morphologically much more similar to the spinosus-like populations of C. tauricus from the western parts of Asia Minor than to the invitus-like populations in the east. Probably, the colonization took place in a phase of low sea-level when today’s extensive western shelf of the Black Sea was situated above the level of the ancient Black Lake, resulting in a large area along the western coast of the Black Lake covered with Sub-Mediterranean vegetation that connected the Sub-Mediterranean areas on the Crimea with the north-west end of Asia Minor (
In contrast, Cyprus is a primarily oceanic island of volcanic origin, that probably has been connected to the mainland only for a period of approximately 600000 years during the Messinian Salinity Crisis of the upper Miocene (5.9–5.3 Ma BP), when a land bridge existed that joined the island with continental areas of Asia Minor and Syria (
We are greatly indebted to Sergey P. Ivanov and Dmitriy V. Puzanov (Taurida Academy of the Vernadskiy Crimean Federal University, Simferopol) for help with collecting material in the field; to Alexander V. Antropov (Zoological Museum of the Lomonosov Moscow State University, Moscow) for help with access to the collection with Kostylev’s type specimens in Moscow, to Sergey A. Belokobylskij (Zoological Institute of the Russian Academy of Sciences, Saint Petersburg) for help with access to the collection of the type specimens of Vespidae in Saint Petersburg, and James M. Carpenter (American Museum of Natural History, New York) for his consultation concerning the whereabouts of the type of C. a. tauricus and his careful review of the manuscript. Josef Gusenleitner (Linz), Erol Yildirim (Atatürk University, Department of Plant Protection, Erzurum), Christoph Saure (Berlin) and Christian Schmid-Egger (Berlin) generously provided material from their collections. Annette Rosenbauer (Stuttgart State Museum of Natural History, Stuttgart) identified the collected plants from Kos and Andreas Müller (Zürich) determined the pollen samples. We are especially grateful to Sarah Gess (Albany Museum, Grahamstown) for valuable comments on the manuscript and improvement of our English.