Celonites tauricus Kostylev, 1935

Mauss, Volker, Fateryga, Alexander V. & Prosi, Rainer, 2016, Taxonomy, distribution and bionomics of Celonites tauricus Kostylev, 1935, stat. n. (Hymenoptera, Vespidae, Masarinae), Journal of Hymenoptera Research 48, pp. 33-66 : 35-47

Celonites tauricus Kostylev, 1935 stat. n.

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. OLML, figs 1a, 2, 5]

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.; Schmidt 1984, Steir. Entomologenrunde 18: 89, misspelling

Material studied.

Taxon 1 (Crimea, " Celonites 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 ZIN, 27.06.2009 1♀ (dbM No. 4701) leg. A. Fateryga VM 1♀ leg. A. Fateryga VTNU, 12.06.2010 1♂ (dbM No. 4308) leg. A. Fateryga VM, 16.06.2010 1♀ (dbM No. 4305) leg. A. Fateryga VM, 09.07.2012 1♀ (dbM No. 4304) leg. A. Fateryga VM; Vicinity of Opolznevoye in the Yalta Mountain Forest Reserve , 44.40000°N, 33.93333°E, 04.07.2006 1♀ leg. S. Ivanov VTNU; Vicinity of Sevastopol: Cape Aya, 44.41667°N, 33.65000°E, 05.07.2004 1♂ (dbM No. 4705) leg. A. Fateryga VM 1♂ leg. A. Fateryga VTNU 1♂ leg. A. Fateryga IZAN 1♂ A. Fateryga KHEO 1♂ leg. A. Fateryga AA 1♀ S. Ivanov VTNU, 07.07.2004 1♀ leg. S. Naumenko DP, 08.07.2004 1♂ (dbM No. 4703) leg. A. Fateryga VM 1♂ leg. A. Fateryga ZIN 1♂ leg. A. Fateryga DP 1♀ leg. A. Fateryga VTNU, 07.2004 1♂ leg. L. Svolynskaya SM 1♀ V. Gromenko AA 1♀ V. Gromenko KHEO; Vicinity of Veseloye near Sudak, 44.86667°N, 34.88333°E, 02.07.2004 1♀ leg. D. Puzanov VTNU; Yalta, Sovetskoye, 44.51667°N, 34.18333°E, 05.06.2010 1♂ (dbM No. 4307) leg. S. Ivanov VM .

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 OLML ; 1 km SE Mastichari, Hang mit Phrygana Fragment unterhalb Kapelle, 36.841317°N, 27.085367°E, 50 m, 26.05.2015 2♀♀ (dbM No. 4776, 4777) leg. V. Mauss VM ; 2 km N Kefalos, Phrygana E des Berges Korakies, 36.765583°N, 26.960067°E, 110 m, 27.05.2015 2♀♀ (dbM No. 4778, 4797) leg. V. Mauss VM ; 4 km SW Kefalos, Phrygana W Agia Anastasia, 36.719300°N, 26.924533°E, 40 m, 31.05.2015 2♀♀ (dbM No. 4783, 4795) leg. V. Mauss VM ; 4 km SW Kefalos , Phrygana W Agia Anastasia, 36.719983°N, 26.930767°E, 90 m, 31.05.2015 1♀ (dbM No. 4782) V. Mauss VM, 02.06.2015 2♀♀ (dbM No. 4792, 4793) leg. V. Mauss VM, 04.06.2015 2♂♂ (dbM No. 4789, 4790) 3♀♀ (dbM No. 4788, 4794, 4798) leg. V. Mauss VM ; 4 km SW Kefalos, Phrygana in sandigem Trockenflusstal N Cape Kata, 36.727733°N, 26.922833°E, 20 m, 31.05.2015 1♂ (dbM No. 4785) 1♀ (dbM No. 4784) leg. V. Mauss VM, 03.06.2015 1♀ (dbM No. 4796) leg. V. Mauss VM ; 4 km W Andimachia, Phrygana Fragment nahe " Hippokrates Garden ", 36.813783°N, 27.051233°E, 60 m, 28.05.2015 1♂ (dbM No. 4780) 1♀ (dbM No. 4779) leg. V. Mauss VM, 31.05.2015 1♂ (dbM No. 4781) 1♀ (dbM No. 4791) leg. V. Mauss VM, 04.06.2015 1♂ (dbM No. 4787) 1♀ (dbM No. 4786) leg. V. Mauss VM .

Taxon 3 (Western Asia Minor, " Celonites 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 OLML. - Antalya: 100 km E Antalya: S von Taskesigi , 36.753821°N, 31.633813°E, 1998 3♂♂ (dbM No. 3586, 4511, 4512) leg. M. Halada OLML; 50 km NE Antalya , 37.183333°N, 31.183333°E, 550 m, 17.06.2001 1♀ (dbM No. 3585) leg. J. Straka OLML; Alanya , 36.558341°N, 31.997983°E, 20- 24.05.1993 1♂ (dbM No. 3431) leg. A. Kudrna OLML; Arapsuyu , 36.874761°N, 30.653459°E, 5 m, 30.06.2002 1♀ (dbM No. 4526) leg. H. Özbek EY. - Aydin: Kusadasi , 37.857802°N, 27.257981°E, 11.06.1964 3♂♂ (dbM No. 4693, 4694, 4695) 1♀ (dbM No. 4665) leg. J. Gusenleitner JG. - Bilecik: wn. Osmaneli , 40.356692°N, 30.001768°E, 14.06.1997 1♀ (dbM No. 3429) leg. M. Halada OLML. - Burdur: 28 km SEE Burdur , 37.666667°N, 30.466667°E, 1350 m, 08.07.2006 1♀ (dbM No. 3591) leg. M. Halada OLML. - Denizli: Çardak, 37.826119°N, 29.672490°E, 07.06.1964 1♂ (dbM No. 4697) leg. J. Gusenleitner JG; Pamukkale , 37.921213°N, 29.114931°E, 31.05.1966 1♀ (dbM No. 4674) leg. K. Kusdas JG. - Izmir: 10 km NE Oedemis , 38.333333°N, 28.066667°E, 1200 m, 03.07.2006 1♂ (dbM 3596) 1♀ (dbM No. 3592) leg. M. Halada OLML; Izmir, 38.417154°N, 27.129925°E, 12.06.1964 1♀ (dbM No. 4666) J. Gusenleitner JG 1♂ (dbM No. 4696) 2♀♀ (dbM No. 4670, 4681) leg. H.H.F. Hamann JG; Tar.Ars.Enst. Menemen , 38.613484°N, 27.077789°E, 02.07.1992 12♀♀ (dbM No. 4530, 4531, 4532, 4533, 4534, 4535, 4536, 4537, 4538, 4539, 4540, 4541) leg. H. Özbek EY. - Kayseri: 25 km S Kayseri , 38.540470°N, 35.490265°E, 23.07.2003 1♀ (dbM No. 3593) leg. J. Straka OLML. - Konya: 30 km S of Aksehir , 38.188770°N, 31.495138°E, 24.06.1998 2♂♂ (dbM No. 3587, 4510) 1♀ (dbM No. 3428) leg. J. Halada OLML; Alti-Napa-Baraji, 37.880339°N, 32.303139°E, 12.06.1978 1♀ (dbM No. 4675) leg. J. Schmidt JG; Sille bei Konya, 37.929344°N, 32.420692°E, 09- 17.06.1975 1♀ (dbM No. 4667) leg. J. Schmidt JG, 08.06.1978 1♀ (dbM No. 4676) leg. J. Schmidt JG, 12.06.1978 2♀♀ (dbM No. 4677, 4678) leg. J. Schmidt JG; W Seydisehir, 37.423314°N, 31.824113°E, 1800 m, 04.08.1991 1♀ (dbM No. 3632) leg. K. Warncke JG. - Nevsehir: Göreme, 38.642663°N, 34.830003°E, 1000 m, 20.07.1986 1♀ (dbM No. 4684) leg. Blank JG . - Uşak: 7 km NW Sivasli , 38.58333°N, 29.58333°E, 21.05.1983 3♀♀ (dbM No. 4687, 4688, 4689) leg. Aspöck, Rausch, Resel JG ; Between Uşak and Sivasli (= 27 km SE Usak), 38.61667°N, 29.58333°E, 830 m, 21.05.1983, 1♀ (dbM No. 4686), leg. E. Hüttinger JG. - Region unkown: Versilkogy oberhalb, 31.05.2007 1♀ (dbM No. 4725) CSE .

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 OLML . - Artvin: Yusufeli Taskiran, 40.803573°N, 41.551595°E, 1350 m, 06.07.2003, 1♂ (dbM No. 4545) leg. I. Aslan EY; Yusufeli, Morkaya , 40.729267°N, 41.602720°E, 700 m, 15.06.2010 1♀ (dbM No. 4528) leg. E. Yildirim EY . - Erzincan: W Refahiye , 39.900486°N, 38.759372°E, 2200 m, 23.08.1991 2♀♀ (dbM No. 3682, 3683) leg. Halada OLML . - Erzurum: Aribahçe, 40.073708°N, 41.167181°E, 2400 m, 20.07.2001 1♂ (dbM No. 4529) leg. E. Yildirim EY; Asagikatikli, Tortum, 40.392175°N, 41.474995°E, 26.07.1980 1♀ (dbM No. 4523) leg. E. Yildirim EY; Caliyazi-Pasinlei, 40.124501°N, 41.669960°E, 2400 m, 10.07.1997 1♀ (dbM No. 4543) leg. E. Yildirim EY; Çamlibel, Oltu, 40.488294 °N, 41.762214°E, 1700 m, 14.07.1996 1♀ (dbM No. 4521) leg. E. Yildirim EY; Dikyar / Uzundere, 40.532510°N, 41.547963°E, 08.09.1996 1♀ (dbM No. 4522) leg. I. Aslan EY; Pazaryolu, Kumaskaya, 40.531839°N, 40.751973°E, 1800 m, 20.07.2011 1♀ (dbM No. 4527) leg. S. Çoruh EY; Umudum Yayl., 40.027957°N, 41.246457°E, 2100 m, 08.08.1991 1♀ (dbM No. 4525) leg. H. Özbek EY. - Icel: Sertavul-Ort, 36.885202°N, 33.277297°E, 1300 m, 11.06.1977 1♀ (dbM No. 4685) leg. J. Schmidt JG . - Kayseri: 10 km S Kayseri , 38.616667°N, 35.516667°E, 1660 m, 20.07.2003 1♀ (dbM No. 3594) leg. J. Straka OLML . - Nigde: NO Çamardi, 37.832838°N, 34.986299°E, 2100 m, 10.08.1991 1♀ (dbM No. 4683) leg. Warncke JG . - Sivas: Gürün, 38.720560°N, 37.278278°E, 05.05.1970 1♀ (dbM No. 4662) leg. J. Gusenleitner JG, 07.06.1970 1♂ (dbM No. 4663) leg. K. Kusdas JG; 20 km E Gürün: Mezikiran Gecidi, 38.747021°N, 37.151695°E, 10.07.1997 1♀ (dbM No. 3601) leg. Ma. Halada OLML . - Tokat: Tokat, 40.324460°N, 36.553842°E, 900 m, 19.08.1992 1♀ (dbM No. 4542) leg. E. Yildirim EY .

Taxon 4 (Eastern Asia Minor, " Celonites 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 OLML . - Bayburt: Bayrampasa, Demirözü, 40.1729804°N, 39.8201781°E, 31.07.1992 1♀ (dbM No. 4524) leg. H. Bastan EY . - Bitlis: Nemrut Dag , 38.650366°N, 42.218203°E, 2300 m, 15.08.1991 1♂ (dbM No. 3630) 1♀ (dbM No. 3629) leg. K. Warncke JG 1♀ (dbM No. 3681) leg. M. Halada OLML . - Erzurum: Palandöken Dagi südwestlich Erzurum: Dutçu, Komyolu , 39.84492°N, 41.25075°E, 2190 m, 28.07.2003 1♀ (dbM No. 2795) leg. V. Mauss & E. Yildirim VM; Palandöken, 39.84492°N, 41.25075°E, 16.08.1987 1♀ (dbM No. 4519) leg. H. Özbek EY, 17.07.1990 1♀ (dbM No. 4520) leg. E. Yildirim EY; Pehlivanli, Tortum, 40.513101°N, 41.484767°E, 29.06.1990 1♀ (dbM No. 4518) leg. E. Yildirim EY . - Hakkari: Tal S Gevria-Pass, 37.499257°N, 43.958866°E, 3000 m, 04.08.1986 1♀ (dbM No. 3634) leg. K. Warncke JG. - Icel: Mut, Passhöhe, 36.630138°N, 33.466345°E, 1520 m, 09.06.1966 1♀ (dbM No. 3578) leg. K. Kusdas JG . - Kars: 50 km S Kars: Pasli , 40.287056°N, 42.964613°E, 01.07.1997 1♂ (dbM No. 3595) leg. Ma. Halada OLML . - Tunceli: Esil , 39.5667°N, 40.1883°E, 12.06.1983 1♀ (dbM 3631) leg. J. Schmidt JG .

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 OLML .

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 .

Field observations, records from literature.

Armenia: Erevan, Monti desertici, Aighpat 40 km SE, 23.07.1963 1♂ 1♀ leg. Giordani Soika coll. Giordani Soika det. J. Gusenleitner, Gusenleitner 1973 [intermediate between Taxon 4 and 5].

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].

Diagnosis.

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 .

Description.

Female. Colour: Black. The following are weakly yellowish white (Fig. 1a ): two spots on frons; small narrow streak on occiput along occipital carina at dorso-lateral corner of head (absent in two specimens); large spot on antero-dorsal angle of pronotum (humeral spot); stripe along dorso-medial (inner) margin of pronotum, slightly enlarged anteriorly, interrupted in the middle by zone of reddish-brown colour; large spot on dorsal mesopleura; median spot on scutellum of moderate size; dorsal and ventral side of propodeal lamella; continuous posterior band on tergum I extending over lateral margin on ventral part of tergum; posterior bands interrupted on each side of middle into median and two lateral markings on terga II-V (in two specimens not completely interrupted on tergum II). Reddish-brown are: distal two third of mandible; labrum; zone in the middle of whitish stripe along dorso-medial margin of pronotum; tegula except small blackish marking at antero-medial margin; distal part of femora, tibiae and tarsi; sternum I; sternum II, becoming slightly darker posteriorly. Dark to blackish brown are: strongly sclerotized parts of labio-maxillary complex; propleura; coxae, trochanters and proximal part of femora; sterna III-V. Antenna with A1-2 black, distal margin of A2 dark brown, A3 proximally dark brown distally reddish brown, A4-11 reddish brown. A12 at least with distal tip black, on ventral side markedly contrasting to adjacent light reddish brown proximal area of antennal club, dorsally extending over distal part of A11 (except in one specimen) fading gradually towards proximal end. Wings translucent blackish-brown.

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. 2 ), few somewhat shorter; some setae on ventro-lateral corners with distal ends curved towards median axis of clypeus; dorso-lateral vertical parts of clypeus more shiny not shagreened with moderately spaced micropunctures. Frons with very weak moderately spaced macropunctation; cuticula dull, densely and strongly shagreened; covered with pale outstanding knobbed setae arising from macropunctures; frontal line weak or absent, not raised to form medial carina. Vertex with close macropunctation becoming more distinct and closely reticulate behind ocelli (Fig. 3 ), where longitudinal interstices are more strongly raised forming lines; cuticula of interstices shiny, weakly longitudinally striated anterior to ocelli, completely smooth behind them; covered with short setae arising from macropunctures. Compound eye sparsely covered with small setae (Fig. 2 ). Gena narrow, preoccipital carina sharp. Antennal articles A8-12 forming ventrally flattened club (Fig. 2 ) slightly more than 2 times as long as broad (viewed dorsally).

Anterior margin of pronotum raised to carina distinctly present along anterior margin of pronotum, 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 low narrow translucent sinuate crest on humeral angle of pronotum. Pronotum with anterior side nearly vertical; dorso-medially slopes down towards mesoscutum, resulting in slight depression along the dorso-medial margin; posterior margin raised to short carina dorsally in front of upper half of tegula; cuticula shiny, with close, reticulate macropunctation, interstices smooth, rounded, without micropunctures; horizontally striated due to more strongly raised longitudinal interstices, becoming more coarsely punctured postero-ventrally with interstices raised to knife-like edges. Cutic ula of mesoscutum shiny, reticulate with close deep macropunctation and narrow, distinctly raised interstices. Postero-medial cuticula of mesoscutum and mesoscutellum with longitudinal interstices more strongly raised leading to striated appearance; finely longitudinally wrinkled, postero-medially with, in addition, moderately spaced micropunctures especially along posterior margin. Mesoscutellum laterally with distinct carina along posterior margin, carina medially increasingly reduced so that cuticula of medial lobe continues evenly into the margin. Carina along posterior margin of metanotum medially with small tooth-like projections. Axilla with short blunt lateral projection only slightly projecting over adjacent posterior end of tegula. Tegula shiny, closely covered by macropunctures except completely smooth central convex area.

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. 5 ); 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 strongly crenulated (Fig. 5 ), crenulation produced into distinct spines raised at an angle of approximately 30° projecting distinctly over the translucent lower posterior margin of terga (Fig. 5 ); cuticula with silken sheen, densely covered with coarse macropunctation, with about nine micropunctures along median axis of tergum III; 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. Tergum VI with sides converging almost as straight lines, posterior margin with curved protrusion over central two-thirds (Fig. 5 ), laterally transverse, forming distinct angle to side.

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. 1a ). White (less yellowish than female): basal spot on mandible (absent in one specimen); labrum; clypeus except small reddish brown ventral margin; sinuate band at frons laterally extending towards upper inner margin of eye where it bends ventrally into ventral half of ocular sinus (shortly to broadly interrupted medially in two specimens); humeral spot postero-laterally extended to posterior margin of pronotum (except in one specimen), dorso-medially lengthened towards enlarged anterior part of stripe along dorso-medial margin of pronotum (fused with it into complete dorso-anterior pronotum band in two specimens); stripe along lateral part of dorso-medial margin of pronotum interrupted in the middle by zone of black colour; posterior bands on terga II-VI interrupted on each side of middle into median and two lateral markings; broad band on posterior four-fifth of sternum II; (in one specimen postero-lateral spot on each side of sternum III-IV). Antenna with A1-2 black; A3-6 anteriorly with whitish stripe otherwise dark brown at basal end changing continuously into reddish-brown to orange towards A6; A7 reddish-brown to orange; antennal club (A8-12) orange, ventrally with distinct blackish marking on A12 extending on distal parts of A11 well set off from orange of adjacent ventral area, orange on dorso-posterior parts of A8-A12 can be darkened to variable extend.

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. 4 ); frontal line can be raised to form small carina or protuberance in centre of frons. Antenna with two oval shaped, perhaps sensory, depressions (tyloids) on concave, ventral side of club, situated within antennal articles A9 and A10. Midcoxa with small but distinct spine at distal end on anterior side close to anterior-medial angle. Posterior margin of tergum VII with three deep emarginations, resulting in two lateral lobes that continue laterally into lateral margins of tergum VII and two lanceolate medial projections (Fig. 6 ). Sternum VIII with distinct little spine on each side of posterior margin; even concavely emarginated, surface not much sunk in towards emargination (Fig. 7 ).

Male genitalia as in Figs 7 , 8 . Genital broad, in dorsal view distance between dorso-medial margins of stipites smaller than transverse width of adjacent part of stipes. Dorso-medial shovel-like lobe of harpide large, densely covered ventrally with long setae. Dorso-posterior margin of stipes only slightly curved, without distinct dorso-medial concavity. Stipes with dorso-medial margin running nearly parallel to sagittal plane. Sides of stipites converging continuously anteriorly towards cupula. Volsella large and broad reaching antero-medial margin of dorso-medial lobe of harpide; dorsal area with strongly sclerotized large, dark tubercles; distances between tubercles moderate; medial process large, trapeziform due to more or less truncate apex, continuing posteriorly into posterior process at a blunt angle. Sides of thyrsoi only slightly converging towards posterior (apical) end of aedoeagus. Each thyrsos continues anteriorly into apodema thyrsos with a distinct outwardly directed curve. On ventral side cupula medially clearly protruded anteriorly, completely projecting over dorsal margin of cupula in ventral view of genital capsule.

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

Geographic distribution and variation.

The geographic range of Celonites tauricus is shown in Fig. 9 . Based on morphological differences six intraspecific taxa can be separated (Table 2 , Figs 10-13 ) belonging to different geographic regions (Fig. 9 ). Distinct population groups exist in the Crimea, Kos, western Asia Minor, eastern Asia Minor up to the Elburs mountains, south-east of the Taurus range and in Cyprus.

Individuals from western Asia Minor are large and strongly built (Fig. 1a ), their metasomal terga are coarsely punctured and bear distinct pointed spines at the posterior end that project well beyond the lower posterior margin (Fig. 10 ). The specimens are dark with reduced whitish markings and reddish brown legs (Fig. 1a ). The posterior bands on the metasomal terga are laterally interrupted, usually from T2 to T5 but at least from T3 to T5. The population from the Crimea differs only slightly in that the individuals are a little less robust and the terga are somewhat less coarsely punctured. The population from Kos varies from the mainland population in that the crenulation along the posterior margin of the metasomal terga is reduced to short, weakly pointed, horizontal processes that do not project beyond the posterior margin (Fig. 12 ). The members of the populations to the south-east of the Taurus range are distinguished from individuals from western Asia Minor by their yellowish colour and considerably more extensive markings (Fig. 1b ), including a spot on the clypeus of the females. The individuals from eastern Asia Minor are smaller and daintier than individuals from the western populations. The macropunctures on their metasomal terga are much smaller and the crenulation along the posterior margins of the terga consists of short, horizontal, truncated processes (Fig. 13 ). The light colour is yellowish and the markings are more extensive than in specimens from the western populations with, at least, complete posterior bands on the terga T1 to T3 (Fig. 1b ). Populations with at least some specimens intergrading between the taxa from western and eastern Asia Minor exist in a zone extending from the Taurus range in the south-west to north-east Turkey and Armenia (Fig. 9 ). Such individuals reveal an intergrading pattern of various morphological and colour traits, but also transient characters like, for example, little spines along the posterior margins of the metasomal terga or macropunctures of intermediate size on the metasomal terga. Likewise there are populations in south-east Turkey with specimens, intergrading between the taxon from eastern Asia Minor and the taxon to the south-east of the Taurus range (Fig. 9 ), that are characterized by little spines along the posterior margins of the metasomal terga and a yellowish spot on the clypeus of the females. The specimens from Cyprus are distinguished by their small size, comparatively dark coloured dorsal side of the antennae (Fig. 1b ), deeper yellow colour, and the crenulation of the metasomal terga that, at least laterally, consists of short but more or less pointed processes.

Figures

Figure 1. 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.

Figure 2. Celonites tauricus female (dbM No. 4305) head in lateral view, clypeus and frons covered with " knobbed setae ".

Figure 7. Male genitalia in dorsal (left) and ventral view (middle) and metasomal sternum VIII of male in ventral view (right) of Celonites abbreviatus (dbM No. 3451), C. tauricus from the Crimea (dbM No. 4307) and C. tauricus from Western Asia Minor (dbM No. 3431).

Figure 8. 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 Birket-Smith (1981): ad, aedoeagus; cu, cupula; 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).

Figure 9. Geographical records of Celonites tauricus. The intraspecific taxon affiliation of the local populations is marked by different colours; a population is regarded as " intermediate " when at least one individual shows intermediate character states. [OpenStreetMap contributors]

Figures 3 - 6. Celonites tauricus 3 - 4 head in dorsal view 3 female (dbM No. 4304) 4 male (dbM No. 4307) 5 - 6 metasoma in dorsal view 5 female (dbM No. 4305) 6 male (dbM No. 4307).

Figures 10 - 13. Cuticula structure of metasomal terga II and III of females belonging to different intraspecific taxa of Celonites tauricus in dorsal view. 10 Western Asia Minor (dbM No. 4681) 11 South-east of Taurus range (dbM No. 4673) 12 Kos (dbM No. 3607) 13 Eastern Asia Minor (dbM No. 3653).