Formica clara clara Forel

Seifert, B. & Schultz, R., 2009, A taxonomic revision of the Formica rufibarbis Fabricius, 1793 group (Hymenoptera: Formicidae)., Myrmecologische Nachrichten 12, pp. 255-272 : 263-265

Formica clara clara Forel , 1886

Formica rufibarbis var. clara Forel , 1886; Syria: Damascus .

Formica rufibarbis var. caucasica Wheeler , 1913; Caucasus . [first available use of Formica rufibarbis ssp. clara var. caucasica Ruzsky, 1905 ]

Formica lusatica Seifert, 1997 ; Germany: Förstgen .

Type material examined: F. clara : 9 syntype workers labelled " F. rufibarbis v. clara Forel, Damas (Lortet) "; MHN Genéve.

F. rufibarbis var. caucasica : One worker lectotype labelled " Form. rufib. clara, var. caucasica , Kavkaz Ruzsky " and " Lectotype des.", ZMLSU Moskva. The specimen is badly damaged: pilosity and pubescence of dorsal body surfaces is almost completely torn off and both scapes are lacking. By subjective guess, based upon the ventral pilosity situation and overall impression, the specimen belongs to Formica clara .

F. lusatica : Holotype plus 5 paratype workers labelled " Kr. Niesky Foerstgen- 1.5 W 1994.06.19-88 " and " Formica lusatica Seifert Holotyp " / " Formica lusatica Seifert Para-typ ", SMN Görlitz .

Material examined: 199 samples with 734 workers were subject to a numeric analysis of 18 characters (Figs. 22, 23): Azerbaijan (1 sample) , Bulgaria (1) , China (18) , Cyprus (1) , Czech Republic (1) , Finland (8) , France (1) , Georgia (2) , Germany (40) , Greece (3) , Hungary (1) , India (14) , Iran (26) , Kazakhstan (22) , Kyrgyzstan (27) , Pakistan (3) , Russia (2) , Slovakia (1) , Switzerland (1) , Syria (3) , The Netherlands (1) , Turkey (22) . For details, see Appendix, as digital supplementary material to this article, at the journal's web pages.

Description of worker (Tab. 1, Figs. 2, 5, 12, 13): larger than F. cunicularia (CS 1.441 mm), head slightly longer (CL / CW1.4 1.138), distance of lateral ocelli moderate (OceD / CS1.4 0.162); eye smaller than in F. cunicula-ria (EYE / CS1.4 0.290), petiole wide (PEW / CS1.4 0.467). Scape rather long (SL / CS1.4 1.080). Clypeus with sharp median keel and fine longitudinal microcarinulae. Frontal triangle finely transversely rippled and with 35 - 55 short pubescence hairs. Eyes with microsetae of 10 - 15 μ m maximum length. Total mean of unilateral setae numbers on different body parts predicted for a specimen with CS = 1.4 mm: pronotum 3.2 (W-population: 3.6, E-population: 2.2), mesonotum 1.4 (W: 1.7, E: 0.8), petiole dorsal of spiracle 0.8 (W: 0.9, E: 0.6), flexor profile of hind tibia 0.6. Posterior margin and underside of head and dorsolat-eral metapleuron as a rule without setae. Ventral coxae with long setae. Dorsum of gaster normally with rather long setae, rarely these are missing. Dorsal mesonotal profile broadly rounded. Metanotal depression in larger individuals relatively deep. Propodeal dome in profile rectangular to round, the basal part sometimes rather linear to concave, whole propodeum flatter in smaller specimens. Dorsal crest of petiole in frontal view normally broadly convex, in smaller specimens sometimes bluntly angled, in larger specimens occasionally with weakly excavate median portion. Petiole scale in lateral aspect rather thin, with convex anterior and more straight posterior profile. Gaster with transverse mi-croripples of small average distance (RipD 4.4 μ m) and covered by dense silvery pubescence (sqPDG 3.1). Pubescence on head, mesosoma and petiole dense. Typical size-dependent variation of colour pattern in the northern population ( F. lusatica ): smaller specimens: area between frontal carinae, posterior vertex, dorsal promesonotum, coxae, and all appendages dark brown, all other body parts light reddish; large specimens: whole dorsal head more or less reddish, whole mesosoma and petiole light reddish, but coxae remaining dark brown; gaster in all size-classes dark brown. Colouration in the southern population ( F. clara ) on average significantly lighter, with predominance of yellowish reddish pigmentation, though specimens with the whole body being more or less dark brown may occur. The type series of F. clara represents a colour extreme: all body parts pale yellowish red with exception of antennal funicu-lus and a patch on dorsocaudal head where brown colour components are added and the gaster which is light brown with a yellowish tinge or even yellowish red (reminiscent of Formica truncorum ).

Comments on taxonomy: The distinction from the similar species F. cunicularia and the unambiguous position of the type samples of F. cunicularia , F. fusca var. ru-bescens, F. cunicularia fuscoides , F. clara , F. rufibarbis var. sinae and F. lusatica within a DA has already been shown above (see also Fig. 14). We only comment here on the synonymisation of F. lusatica with F. clara .

The extreme colour differences and some morphometric deviation between the Syrian type series of F. clara and the northern F. lusatica population has suggested hetero-specificity in a time when the geographic variation of both taxa over their whole Palaearctic range has been poorly known ( Seifert 1997). This situation has changed now - 200 nest samples from Eurasia with recording of the complete character set and 62 samples with incomplete character set became available during the last years. Considering all sixteen structural characters and two pigmentation characters, a principal component analysis gave no suggestion on a possible grouping according to geography. We also could not demonstrate two entities by starting with the hypothesis that only material from Syria, Lebanon and Iran represented F. clara , then iteratively running discriminant analyses and forming new hypotheses after each run. The circuit system stabilised at insufficient parameters: only 83.1% of determinations achieved p> 0.95, the error indication by LOOCV was 2.6% and 21% of the samples from Germany and Finland were determined as F. clara which should not be true if the latter one is a separate southern species. The assumed differences between northern and southern populations most probably represent intraspecific clinal variation. Indeed there are detectable some highly significant geographic trends: CS and CL / CW1.4 increase with growing geographical latitude while GHL / CS1.4 falls (all regressions with p <0.0001).

199 samples with 734 workers were subject to a numeric analysis of 18 characters. Azerbaijan: Mingacevir , 1.VI.2006 [40.743° N, 47.114° E] . Bulgaria: Obsor 1.VIII.1979 [42.820° N, 27.880° E] . China: Boro-Horo (No. 115), 18.VII.2006 [43.838° N, 85.358° E] ; Boro-Horo (No. 186), 22.VII.2006 [43.964° N, 85.180° E] ; glacier No. 1 (No. 036), 28.VIII.2004 [43.588° N, 87.198° E] ; glacier No. 1 (No. 038), 28.VIII.2004 [43.559° N, 87.198° E] ; Halamuyan Zhan vall. (3 samples, No. 221, 222, 227), 23.IX.2004 [43.859° N, 88.175° E] ; Tjanchi (No. 255), 26. IX.2004 [43.937° N, 88.106° E] ; Tjanchi (No. 257), 26.IX.2004 [43.936° N, 88.106° E] ; Tjanchi (No. 262), 26.IX.2004 [43.936° N, 88.105° E]; Tjanchi E (2 samples, No. 246, 247), 25.IX.2004 [43.953° N, 88.201° E]; Tjanchi S (No. 170), 19.IX.2004 [43.872° N, 88.141° E]; Urumqi (No. 002a), 1.IX.2003 [43.800° N, 87.583° E]; Urumqi (No. 111), 17.VII. 2006 [43.764° N, 87.617° E]; Urumqi (No. 141a), 17.IX.2004 [43.765° N, 87.611° E]; Urumqi (No. 245), 24.VII.2006 [43.768° N, 87.621° E] ; NW China, 1991, [leg. Heatwole , exact location not available] . Cyprus: Limassol , 29.III.1999 [34.9° N, 32.9° E, coordinates estimated] . Czech Republic: Chomutov , 2007 [50.568° N, 13.565° E] . Finland: Luumäki (2 samples, No. 24, 34), 12.VII.1996 [60.913° N, 27.380° E] ; Sandvik (4 samples, No. 16, 30, 136, 160), 10.VII. 1996 [60.280° N, 22.210° E] ; Stormälo (2 samples, No. 20, 39), 10.VII.1996 [60.250° N, 22.150° E] . France: Font-Romeu , 24.VI.1999 [42.500° N, 2.080° E] . Georgia: Tbilisi (No. 660), 21.VII.1985 [41.810° N, 44.790°E] ; Tbilisi (No. 661), 20.VII.1985 [41.688° N, 44.808° E] . Germany: Förstgen (6 samples, No. 88, type lusatica, A1, 12, 59, 94, 127), 19.VI. 1994 [51.300° N, 14.640° E] ; Gartz , 5.VIII.1987 [53.210° N, 14.380° E] ; Halbendorf (3 samples, No. 041, 094, 12), 16. V.1992 / 30.VII.1992 [51.290° N, 14.530° E] ; Halbendorf (No. 095), 16.V.1992 [51.300° N, 14.530° E] ; Hiddensee , IX. 2000 [54.157° N, 13.816° E] ; Ingelheim , 2001 [49.970° N, 8.050° E] ; Jänkendorf (5 samples, No. g2, g13, g19, 76, 208), 2.VIII.1992 [51.270° N, 14.800° E] ; Jänkendorf (No. 85), 2.VIII.1992 [51.280° N, 14.790° N] ; Klotzsche , 18.VIII.1992 [51.090° N, 13.780° E] ; Leutra (3 samples, No. g3, g5, g31), 19.VII.1993 [50.870° N, 11.570° E] ; Lodenau , 18.IX.1992 [51.360° N, 14.960° E] ; Münsingen, 31.VII.2008 [48.426° N, 9.577° E] ; Münsingen, 29.IX.2008 [48.447° N, 9.532° E] ; Oberbergen (3 samples, No. 6, 11, 143), 2.V.1993 [48.110° N, 7.660° E] ; Peenemünde, 5.V.2001 [54.157° N, 13.816° E] ; Rothenburg (3 samples, No 41, 72, 103), 10.VII.1993 [51.350° N, 14.970° E] ; Rotschönberg, 16.V.1993 [51.2° N, 13.5° E, coordinates estimated] ; Sasbachwalden (4 samples, No. g27, 65, 86, g89), 30.IV.1993 [48.610° N, 8.160° E] ; Schönecken, 1.V.1990 [50.165° N, 6.475° E] ; Wüstermarke, 25.VIII.1984 [51.820° N, 13.600° E] . Greece: Elis (2 samples), 16.VII. 1996 / 6.VIII.1996 [37.900° N, 21,410° E] ; Nestos Delta (No. 2), 2004 [40.850° N, 24.800° E] . Hungary: Hortobagy , V. 1998 [47.583° N, 21.167° E] . India: Fagu (2 samples, No. 385, 389), 29.IX.1996 [31.127° N, 77.332° E] ; Jari (2 samples, No. 490, 491), 3.X.1996 [32° N, 77.3° E, coordinates estimated] ; Kalath (No. 440, 446), 2.X.1996 [32.2° N, 77.2° E, coordinates estimated] ; Kasol (3 samples, No. 472, 476, 483), 3.X.1996 [32.020° N, 77.320° E] ; Rothang La Pass (2 samples, No. 433, 458), 1./ 2.X.1996 [32.346° N, 77.217° E] ; Spiti vall. , 15.VII.1994 [32.083° N, 78.217° E] ; Theog , 29.IX.1996 [31.142° N, 77.393° E] ; Yourdu , 16.VII.1980 [33.42° N, 75.77°E, coordinates estimated] . Iran: Alam kola forest , 22.VI. 2008 [36.550° N, 51.801° E] ; Astaneh , 2005 [37.267° N, 49.667° E] ; Baba Aman , 22.X.2004 [37.467° N, 57.317° E] ; Babolsar , 2005 [36.700° N, 52.633° E] ; Bojnourd (2 samples), III.2004, 24.VII.2004 [37.500° N, 57.317° E] ; Chabokar , 3.VII.2008 [36.950° N, 50.550° E] ; Dimeh , 8.VIII.1973 [32.483° N, 50.267° E] ; Firozeeh , 2.VI.2005 [37.483° N, 57.267° E] ; Ganznameh , 4.VII.1974 [34.733° N, 48.500° E] ; Ghaemshahr , 3.VI.2008 [36.371° N, 52.850°E] ; Golestan N.P. (2 samples, forest and canyon), 14.V.2007 [37.367° N, 55.817° E] ; Gorgan , 11.V.2006 [36.767° N, 54.367° E] ; Hamseh-kola , 19.VII.1973 [36.500° N, 52.517° E] ; Khoshyelagh , 10.VI.2007 [36.783° N, 55.350° E] ; Kuhrang , 19.VI.1974 [32.457° N, 50.133° E] ; Mamraz dasht , 23.VI.2008 [36.539° N, 51.853° E] ; Noor , 3.VI.2005 [36.555° N, 51.950° E] ; Shahrood , 7.V.2008 [36.418° N, 54.961° E] ; Sisangan park , 24.VI.2008 [36.576° N, 51.813° E] ; Tehran , Avadj , 27.VII.1973 [35.633° N, 49.217° E] ; Tehran , Delichal , 24.VII.1973 [35.667° N, 52.500° E] ; Tehran , Feruzkuh , 24.VII.1973 [35.717° N, 52.667° E] ; Tehran , Shemshak , 27.VIII.1973 [36.017° N, 51.483° E] ; 2007/12 [without location]. Kazakhstan: Aksu Canyon , 31.VII.1998 [42.332° N, 70.375° E] ; Almaty/Medeo (2 samples), 30.VIII.2006, X.2006 [43.518° N, 77.059° E] ; Dshungarsky Alatao , 8.VIII.2001 [45.248° N, 80.288° E] ; Dzhabagly , 2.VIII.1998 [42.408° N, 70.583° E] ; Kalichan , 26.VII.2001 [47.964° N, 85.061°E] ; Karaganda , 4.IX.2006 [49.833° N, 73.167° E] ; Kenbirlik , 26.VII.2001 [47.682° N, 84.961° E] ; Kyrgysky Alatao , 3.VIII.1998 [42.781° N, 71.226° E] ; Manrak (No. 115), 27.VII.2001 [47.323° N, 84.622° E] ; Manrak (No. 122), 27.VII.2001 [47.321° N, 84.617° E] ; Saisan (No. 050), 23.VII.2001 [47.466° N, 84.878° E] ; Saisan (No. 113), 27.VII.2001 [47.467° N, 84.873° E] ; Sarymobe (No. 186), 4.VIII.2001 [47.119° N, 82.371° E] ; Sarymobe (No. 190a), 4.VIII.2001 [47.127° N, 82.369° E] ; Saur (No. 075b), 24.VII.2001 [47.294°N, 85.618° E] ; Saur (No. 089), 25.VII. 2001 [47.357° N, 85.518° E] ; Saur , Matagul , 28.VII.2001 [47.049° N, 84.915° E] ; Tarbagatay (No. 023), 20.VII.2001 [47.274° N, 80.808° E] ; Tarbagatay (No. 025), 20.VII.2001 [47.276° N, 80.806° E] ; Tarbagatay (No. 034), 22.VII.2001 [47.517° N, 82.726° E] ; Tarbagatay (No. 182), 4.VIII.2001 [47.070° N, 82.313° E] . Kyrgyzstan: Abshir Say (No. 049), 18.VII.2004 [40.086° N, 72.347° E] ; Abshir Say (No. 057), 18.VII.2004 [40.093° N, 72.350°] ; Abshir Say (No. 060), 18. VII.2004 [40.101° N, 72.361° E] ; Ananjevo , 29.VII.2000 [42.709° N, 77.694° E] ; Bishkek , 15.VII.2004 [42.882° N, 74.591° E] ; Chodsha-Ata , 26.VII.1998 [41.757° N, 71.973° E] ; Dshan-Bulak, 24.VII.2005 [42.393° N, 78.308° E] ; Kara-kol, 28.VII.2000 [42.424° N, 78.437° E] ; Karavshin vall., 24.VII.2004 [39.781° N, 70.412° E] ; Karavshin vall., 25.VII. 2004 [39.780° N, 70.412° E] ; Kysyl-Alai, 26.VII.1999 [39.851° N, 73.291° E] ; Moldotoo (2 samples, No. 025, 026), 16.VII.2000 [41.643° N, 75.036° E] ; Naryn vall., 21.VII.2000 [41.496° N, 76.424° E] ; Naryn vall., 21.VII.2005 [41.495° N, 76.425° E] ; Ottuc (No. 066), 22.VII.2000 [42.314° N, 76.314° E] ; Ottuc (No. 070), 22.VII.2000 [42.315° N, 76.315° E] ; Sary-Tshelek (3 samples, No. 091, 092, 093), 25.VII.1998 [41.812° N, 71.959° E] ; Sousamyr vall. (2 samples, No. 009, 021), 17/ 18.VII.1998 [42.207° N, 73.625° E] ; Toktogul, 29.VII.1999 [41.880° N, 72.947° E] ; Tos Kol (4 samples, No. 068, 071, 076, 078), 23./ 24.VII.1998 [41.874° N, 71.979° E] . Pakistan: Kawai, 15.VI.1977 [34.7° N, 73.5° E, coordinates estimated] ; Naran S, 31.V.1983 [34.9° N, 73.65° E, coordinates estimated] ; Naran, 1.VI.1983 [34.8° N, 73.5° E, coordinates estimated] . Russia: Alagir, 1907 [43° N, 44.22° E, coordinates estimated] ; Kursk steppe, 17.VII.1999 [51.750° N, 36.167° E] . Slovakia: Vidova, 15.IV.2005 [48.570° N, 20.458° E] . Switzerland: Pfynwald (No. 41), 16.V.1994 [46.300° N, 7.620° E] . Syria: Ar Rastan, 18.III.1979 [34.917° N, 36.750° E] ; Damascus, before 1886 [33.5° N, 36.3° E, type clara, leg. Lortet, coordinates estimated] ; Pac de Homs, [34.75° N, 36.75° E, leg. Kerville, coordinates estimated] . The Netherlands: Roer, 26.VI.2000 [51.160° N, 6.120° E] . Turkey: Aksaray, 2.VI.1993 [38.23° N, 34° E, coordinates estimated] ; Ankara, [39.92° N, 32.85° E, leg. Kerville, coordinates estimated] ; Arsdahan, 24.VI.1993 [41.130° N, 43.010° E] ; Arslanköy, 30.V.1993 [37.170° N, 34.330° E] ; Aydogdu (2 samples, No. 1149, 1150), 26.VI.1993 [40.680° N, 42.410° E] ; Borcka Civan, 29.VI.1993 [41.330° N, 41.690° E] ; Budakli, 17.VI.1993 [38.420° N, 42.420° E] ; Catak, 18.VI.1993 [39.500° N, 43.010° E] ; Eregli, 3.VI.1993 [37.630° N, 34.080° E] ; Kabali, 3.VII.1993 [41.800° N, 35.050° E] ; Karakurt, 17.VI.1986 [40.160° N, 42.610° E] ; Kizikadag, 21.V.1993 [36.880° N, 29.910° E] ; Marmaris, 18.VIII.1992 [36.78° N, 28° E, coordinates estimated] ; Mersin NW, 29.V.1993 [37.020° N, 34.330° E] ; Ödemis (No. 866), 19.V.1993 [38.420° N, 27.890° E] ; Ovitdagi Gecidi, 30.VI.1993 [40.770° N, 40.420° E] ; Pinahisar, 10.V.2003 [41.411° N, 27,237° E] ; Saklikent, 27.V.1993 [36.840° N, 30.250° E] ; Samailli, 20.V.1993 [38.080° N, 28.290° E] ; Sarigöl (No. 1179), 28.VI.1993 [41° N, 41.5° E, coordinates estimated] ; Ziyarettepesi Gecidi, 9.V.1997 [38.886° N, 36.824° E] .

Distribution and biology: Shows the widest geographical range among all species of the F. rufibarbis group. Inhabits the temperate, submeridional and meridional zones of the Palaearctic, occurs between 2° and 88° E (as opposed to 120° E in F. clara sinae stat. n.) and 31° and 61° N (in Finland). Having a planar to colline distribution in the northern parts of its European range, but climbing up to 3500 m at 31° N in the Himalayas. Primarily a species of the dry steppes. Most thermophilic of the European species of the F. r u f ib a r b i s group ( Seifert 1997, 2007) and in Central Europe the rarest of these species, with only regional occurrence in warm regions below 700 m, occurring here mainly on very xerothermic sandy and limestone grasslands, also on ruderal grassland, generally preferring open land with patchy herb layer. In sandy areas of the Lausitz (Germany) outcompeting F. rufibarbis . In the southern parts of the range also in more moist places, city parks and semi-shaded woodland. Presence, mean and maximum nest density on 81 potentially suitable, 150- m 2-test-plots on open land in Germany 15%, 0.21 and 3.7 nests / 100 m 2 respectively. Nests often rather populous, monogynous to weakly poly-gynous, with aggressive workers which are, at least in the northern range, notably larger than those of F. cunicularia and F. rufibarbis . Big nests defend territories. Resistance against social parasites certainly stronger than in F. cuni-cularia ( Seifert 1997, 2007; Czechowski & Radchenko 2006). However, main host of Polyergus rufescens (Latreille, 1798) in Central Asia where F. cunicularia and F. rufibarbis are rare or absent. Alates occur in Central Europe 14 July ± 16 d [29 June, 20 July], n = 14.