Formica cunicularia Latreille, 1798 Formica fusca var. rubescens Forel, 1904 Formica cunicularia fuscoides Dlussky, 1967 A taxonomic revision of the Formica rufibarbis Fabricius, 1793 group (Hymenoptera: Formicidae). Seifert, B. Schultz, R. Myrmecologische Nachrichten 2009 12 255 272 6JH7S 72780 Latreille, 1798 Latreille 1798 Insecta Formicidae Formica Animalia Hymenoptera 261 Arthropoda species cunicularia   Formica cunicularia Latreille, 1798; France.  Formica fusca var. rubescens Forel, 1904;  Switzerland: Vaux.  Formica cunicularia fuscoides Dlussky, 1967;  Armenia: Byurakan.  Type material examined: F. cunicularia:  Neotypeworker labelled "FRA: 44.4947° N, 0.9597° E, Fumel, 120 m, in a garden, leg. Galkowski2008.07.25" and " Neotype Formica cunicularia Latreille1798, des. Seifert& Schultz2009"; SMN  Goerlitz.In case of destruction or loss of the neotypespecimen, a replacement neotypecan be designated from a series of five mounted workers from the same nest series in SMN Goerlitzand further five workers in MNHNParis. Justification of the neotypefixation: A current search in the Latreille collection of MNHNParis failed to detect a specimen interpretable as a primary type (J. Casevitz-Weu-lersse, pers. comm. 2008) and the literature gives no indication that a revisor ever has seen one. In order to establish an unambiguous standard for differentiation from similar species, we fixed a neotypein a sample from the terra ty-pica which is in agreement with the traditional morphological conception of F. cunicularia.  F. fusca var. rubescens: 4 syntypeworkers labelled by Forel himself " Vaux ", " Cotypus " and " F. fusca r. glebaria v. rubescens For", MZ Lausanne.  F. cunicularia fuscoides: Five workers from the holo-type nest, labelled "1103", " Byurakan 1800 m Armen. Dlusskij 13.VI.960" [in cyrillic letters], " Formica cunicularia kajastanica Dlussky paratypes". Serial No. 1103 definitely designates the holotypenest ( Dlussky 1967, p. 74) but a holotypecannot be identified within these five specimens. They are paratypesat least. A " F. c. kajastanica" has never been validly published. One gyne and two workers labelled "1091" and " Alagez nad Byurakanom Armeniya G. Dlusskij 13. 6.60" were also explicitly mentioned by Dlussky and have paratypestatus. Material examined: 85 samples with 307 workers were subject to a numeric analysis of 18 characters (Figs. 20, 21): Armenia (2 samples), France (12), Georgia (5), Germany (24), Great Britain (1), Greece (3), Hungary (3), Italy (10), Kazakhstan (7), Portugal (1), Russia (1), Spain (1), Switzerland (2), Turkey (11), Ukraine (2). For details, see Appendix, as digital supplementary material to this article, at the journal's web pages.  Description of worker, continental population (Tab. 1, Figs. 1, 4): medium-sized Serviformicaspecies (CS 1.365 mm); head slightly elongated (CL / CW1.4 1.131); Scape moderately long SL / CS1.4 1.073; distance of lateral ocelli moderate (OceD / CS1.4 0.164); eyes rather large (EYE / CS1.4 0.301), petiole rather wide (PEW / CS1.4 0.468). Clypeus with sharp median keel and fine longitudinal micro-carinulae. Frontal triangle finely transversely rippled and with 30 - 60 short pubescence hairs. Eyes with microsetae of 7 - 13 μ m maximum length. Total mean of unilateral setae numbers on different body parts predicted for a specimen with CS = 1.4 mm: pronotum 1.1, mesonotum 0.8, petiole scale dorsal of spiracle 0.25, flexor profile of hind tibia 0.3. Posterior margin and underside of head and dorso-lateral metapleuron as a rule without setae. Ventral coxae with long setae, dorsum of gaster with scattered, moderately long setae. Dorsal mesonotum in lateral aspect broadly rounded. Metanotal depression in larger individuals relatively deep. Propodeal dome rounded in lateral view, basal profile sometimes concave and in smaller specimens often straight. Dorsal crest of petiole in frontal view bluntly angledin smaller specimens to broadly convex in larger specimens, in some of the large individuals with straight or weekly excavate median portion. Petiole scale in lateral aspect rather thin, with convex anterior and more straight posterior profile. Gaster with transverse microripples of small average distance (RipD 4.6 μ m) and covered by dense silvery pubescence (sqPDG 3.1). Pubescence on head, meso-soma and petiole dense. Typical colour pattern: Head with exception of round reddish-yellowish spots on anterior ge-nae, dorsal promesonotum, coxae and all appendages dark brown, gaster blackish brown. Other body parts more or less reddish-yellowish. Nests with much lighter specimens having whole mesosoma, coxae and petiole uniformly reddish and such with very dark specimens having the reddish pigmentation reduced to a very small spot on frontal margin of ventrolateral mesonotum; exceptionally completely dark specimens occur. Description of worker, West Mediterranean isolated populations: We do not at this time propose these deviating and isolated populations from Corsica, Sardinia and the Sierra Nevada as heterospecific from F. cunicularia. Differences to the continental population are a significantly narrower petiole (PEW / CS1.4 0.433) and slightly longer 1 st tergite setae (GHL / CS1.4 7.24%). It seems to be the only species of the group from Corsica where F. claraand F. rufibarbishave not been reported so far.  Comments on taxonomy: The separation of F. cunicu-laria and F. clararepresents the most difficult discrimination problem within the F. rufibarbisgroup because there is a deficiency of strongly discriminating structural characters. We ran a DA for the whole Palaearctic range of F. claraexcluding material from the West Himalayas (Pakistan and India) where no ant species similar to F. cuni-culariais present. Considering the characters CS, CL / CW1.4, SL / CS1.4, OceD / CS1.4, EYE / CS1.4, PEW / CS1.4, nPN1.4, nMN1.4, nPRME1.4, nPE1.4, nHFFL1.4, RipD1.4, sqPDG1.4, PIGM 1.4, and CONT 1.4, a two-class DA separates 97.4% of 268 nest samples with p> 0.95 and the LOOCV-DA gives an error prediction of 0.4% (Fig. 14): D (15) F. clara-2.781 ± 0.864 [-4.40, -0.17] n = 183 D (15) F. cunicularia-2.642 ± 1.245 [0.17, 5.22] n = 85 The neotypesample of F. cunicularia, the syntypesample of F. fusca var. rubescensand the holotypesample of F. cunicularia fuscoidesare safely allocated to the F. cuni-culariacluster (each with p = 1.000) while syntypesample of F. clara Forel, 1886, the holotypesample of Formica lusatica Seifert, 1997and the syntypesample of F. rufi-barbis var. sinae Emery, 1925 are safely assigned to the F. claracluster (each with p = 1.000). While these type allocations allow clear nomenclatural decisions at least within this species pair, we do not fully trust any determination within the complete data set. The weak point is that intraspecific colour polymorphism and loss of pigmentation by light or storage media could possibly affect the reliability of the pigmentation characters PIGM and CONT, but just these two characters have the largest loadings (canonical correlations) in the DA. These loadings are 0.788 in PIGM 1.4 and 0.391 in CONT 1.4 but only 0.336 in EYE1.4 and 0.170 in nPN1.4, the two best structural discriminators. Another problem are the isolated West Mediterranean populations of F. cuniculariafrom Corsica, Sardinia and the Sierra Nevada which were all allocated in the DA to the F. cuniculariacluster but possibly represent a third species. We have currently no NUMO-BAT method to show this. Integrative approaches including DNA analysis could bring more clarity into this issue.  Fig. 14: Discriminant analysis of worker nest samples considering 15 characters of the Palaearctic population of F. claraand F. cunicularia. Position of type samples shown by arrows: CL - F. clara, L - F. lusatica, CU- F. cunicu-laria, R - F. fusca var. rubescens, F - F. cunicularia fus-coides, S - F. rufibarbis var. sinae.  85 samples with 307 workers were subject to a numeric analysis of 18 characters.  Armenia: Alages( typefuscoides, paratypefuscoides),  13.VI.1960[ 40.33° N, 44.25° E, coordinates estimated].  France: Breil sur Roya(No. 001),  7.V.2002[ 43.942° N, 7.518° E];  Breil sur Roya(No. 025),  8.V.2002[ 43.951° N, 7.519° E];  Breil sur Roya(No. 038),  9.V.2002[ 43.924° N, 7.484° E];  Corse(D 81)  5.VI.1985[ 42.467° N, 8.683° E];  Fumel( neotypecunicularia),  25.VII.2008[ 44.495° N, 0.960° E];  Galeria,  13.VI.1985[ 42.410° N, 8.650° E];  Labergement-Sainte-Marie,  11.VII.1990[ 46.770° N, 6.280°];  Lac de L'Oriente,  5.VII.1974[ 42.229° N, 9.058° E];  Orne, 1917 [ 48.81° N, 0.26° E, coordinates estimated];  Suartone,  6.VI.1975[ 41.5° N, 9.25° E, coordinates estimated];  Timozzo,  5.VII.1974[ 42.244° N, 9.067° E];  MNHN Paris( EY0000001658)[without date and location].  Georgia: Kazbergi(2 samples, No. 654, 644),  27.VII.1985/  8.VIII.1985[ 42.670° N, 44.580° E];  Mzcheta,  23.VII.1985[ 41.840° N, 44.720° E];  Omalo(2 samples, 650, 653m),  31.VII.1985[ 42.380° N, 45.630° E].  Germany: Burkheim,  1.V.1993[ 48.100° N, 7.600° E];   Daenkritz,  27. V.1992[ 50.770° N, 12.430° E];  Ebersbach(3 samples, No. 026, 076, 222),  24.V.1992[ 51.190° N, 14.930° E];   Foerst-gen,  8.V.1994[ 51.300° N, 14.640° E];  Griesheimer Duene,  29.IV.1993[ 49.840° N, 8.590° E];  Heilsberg,  25.VIII.1986[ 50.776° N, 11.265° E];  Isteiner Klotz,  4.V.1993[ 47.670° N, 7.530° E];  Langenhessen(3 samples, No. 053, 054, 067),  28.V.1992[ 50.770° N, 12.370° E];  Klein Schmoelen,  4.VIII.1996[ 53.123° N, 11.289° E];  Lubminer Heide,  14.VI.1998[ 54.147° N, 13.647° E];  Neubrandenburg,  20.VI.1998[ 53.596° N, 13.409° E];  Niederhohndorf(2 samples, No. 076, 084),  26.V.1992[ 50.750° N, 12.470° E];  Rengersdorf(2 samples, No. 054, 234),  20.V.1992[ 51.208° N, 14.892° E];  Rohrdorf ,  8.V.1993[ 47.717° N, 10.083° E];  Torga,  20.V.1992[ 51.208° N, 14.910° E];  Trebbichau,  31.V.1987[ 51.810° N, 12.010° E];  Vogtsburg,  3.V.1993[ 48.100° N, 7.700° E];  Waren/Feissnecksee,  27.VII.1988[ 53.490° N, 12.710°].  Great Britain: Hartland Moor,  24.X.1991[ 50.657° N, 2.092° W].  Greece: Agia Triada,  1.VI.2001[ 40.500° N, 22.882° E];  Litochoro,  29.V.2001[ 40.102° N, 22.562° E];  Nestos Delta(No. 1), 2004 [ 40.850° N, 24.800° E].  Hungary: Aggtelek(3 samples, No. 004, 005, 006),  V.1998[ 48.467° N, 20.517° E].  Italy: Baunei,  5.X.1972[ 40.120° N, 9.510° E];  Camigli-atello Silano(No. 003),  14.V.1999[ 39.376° N, 16.441° E];  Camigliatello Silano(No. 005),  14.V.1999[ 39.363° N, 16.456° E];  Camigliatello Silano(No. 023),  16.V.1999[ 39.361° N, 16.430° E];  Firenze,  20.VII.1994[ 43.783° N, 11.600° E];  Largo Arvo,  19.V.1994[ 39.230° N, 16.500° E];  Mte. Limbara,  3.V.1994[ 40.850° N, 9.180° E];  Sila Grande(No. 029),  17.V.1999[ 39.392° N, 16.550° E];  Sila Grande(No. 049),  20.V.1999[ 39.387° N, 16.559° E];  Sila Grande(No. 061b),  21.V.1999[ 39.411° N, 16.553° E].  Kazakhstan: Manrak,  27.VII.2001[ 47.317° N, 84.617° E];  Sarymobe(2 samples, No. 197a, 206),  5.VIII.2001[ 47.130° N, 82.372° E];  Saur(No. 082),  25.VII.2001[ 47.357° N, 85.518° E];  Tarbagatay(No. 160),  1.VIII.2001[ 47.784° N, 81.754° E];  Tarbagatay(No. 306),  5.VIII.2001[ 47.133° N, 82.367° E];  Tarbagatay(No. 338),  1.VIII.2001[ 47.783° N, 81.767° E].  Portugal: Porto Covo,  15.III.2002[ 37.839° N, 8.788° W].  Russia: Kursksteppe,  17.VII.1999[ 51.750° N, 36.167° E].  Spain: VeletaS,  10.V.1991[ 37.060° N, 3.360° W].  Switzerland: Pfynwald(No. g21)  16.V.1994[ 46.300° N, 7.620° E];  Vaux(cotype rubescens), [ 46.540° N, 6.470° E, coordinates estimated].  Turkey: Catak,  18.VI.1993[ 39.500° N, 43.010° E];  Dagpinar,  23.VI.1993[ 40.470° N, 43.330° E];  Dirimli Gecidi,  22.V.1993[ 36.870° N, 29.580° E];  Dranoz,  3.VII.1993[ 41.640° N, 34.840° E];  Kabali,  3.VII.1993[ 41.800° N, 35.050° E];   Oedemis(No. 870),  19.V.1993[ 38.350° N, 27.880° E];  Sac Gecidi,  22.VI.1993[ 39.800° N, 42.510° E];   Sarigoel(No. 1177),  28.VI.1993[ 41° N, 41.5° E, coordinates estimated];  Sogukpinar(2 samples, No. 1229, 1238),  6.VII.1993[ 40.060° N, 29.120° E];  Stranga Mts.,  10.V.2003[ 41.468° N, 27.375° E].  Ukraine: Jalta,  16.VIII.1995[ 44.450° N, 34.050° E]; Manhup-Kale,  12.VIII.1995[ 44.617° N, 33.867° E].  Distribution and biology: Temperate, Ponto-south-Siberian and Submediterranean species of the West Palae-arctic, occurring from southernmost England and Iberia to West Siberia ( 85° E). In northwestern Europe, it goes north to southern Sweden ( 58° N) but has not reached southern Finland so far. Having a planar to colline distribution in the northern parts of its range, it climbs up to 1800 m in the Alps, up to 2400 m in the Caucasus and up to 2000 m in the South Siberian Tarbagatay Mountains.  Moderately thermophilic. Preferred habitats are meagre and semidry grasslands on differing geological outcrop, more rarely extremely xerothermic grasslands and also open ruderal, rural or disturbed habitats, including road or railway verges. In contrast to F. rufibarbismore frequently occurring on loamy soils with more developed herb layer and less often invading the urban zone. Presence, mean and maximum nest density on 81 potentially suitable, 150- m 2-test-plots on open land in Germany 5%, 1.6 and 23.2 nests / 100 m 2 respectively. Colony foundation usually by single gynes but also pleometrotic. Nests moderately populous, usually containing 1000 - 1800 workers, sometimes weakly polygynous, but polydomous colonies unknown (as in all members of the group). Usually inhabits simple soil nests, construction of high mounds of mineral soil in meagre grassland with higher herb layer regularly observed. Usually timid and fugitive, but populous nests with large workers may be very aggressive during nest defence. Not territorial. Foraging at surface temperatures of up to 50° C, mainly on open surfaces and in the herb layer, but not avoiding bushes and trees. Zoophagous, trophobiotic and nectarivorous. Low position in dominance hierarchies of ant communities, usually inferior to even Lasiusniger (Linnaeus, 1758), whom it carefully evades thanks to superior walking speed and well-developed visual sense, thus enabling coexistence at long-term food sources. May snatch large prey items from L. niger by swift surprise attack. Favoured host species for several socially parasitic ant species. Alates occur 7 July± 12 d [ 16 June, 1 August], n = 17 ( Seifert 2007). 923905201 Switzerland Vaux 261 923905210 Armenia Byurakan 261 923905213 France SMN Galkowski, Seifert, Schultz 44.4947 Fumel 0.9597 261 Neotype 923905220 1960-06-13 Armenia 40.33 Alages 44.25 262 type 923905223 2002-05-07 France 43.942 Breil sur Roya 7.518 262 Breil Sur Roya 923905227 2002-05-08 France 43.951 Breil sur Roya 7.519 262 Breil Sur Roya 923905225 2002-05-09 France 43.924 Breil sur Roya 7.484 262 Breil Sur Roya 923905216 1985-06-05 France 42.467 Corse 8.683 262 Breil Sur Roya 923905199 2008-07-25 France 44.495 Fumel 0.96 262 Breil Sur Roya neotype 923905230 1985-06-13 France 42.41 Galeria 8.65 262 Breil Sur Roya 923905222 1990-07-11 France 46.77 Labergement-Sainte-Marie 262 Breil Sur Roya 923905219 1974-07-05 France 42.229 Lac de L'Oriente 9.058 262 Breil Sur Roya 923905208 France 48.81 Orne 0.26 262 Breil Sur Roya 923905205 1975-06-06 France 41.5 Suartone 9.25 262 Breil Sur Roya 923905206 1974-07-05 France 42.244 Timozzo 9.067 262 Breil Sur Roya 923905229 MNHN France Paris 262 EY 0000001658) Breil Sur Roya 923905207 1985-07-27 Georgia 42.67 Kazbergi 44.58 262 Breil Sur Roya 923905236 1985-07-23 Georgia 41.84 Mzcheta 44.72 262 Breil Sur Roya 923905228 1985-07-31 Georgia 42.38 Omalo 45.63 262 Breil Sur Roya 923905243 1993-05-01 Germany 48.1 Burkheim 7.6 262 Breil Sur Roya 923905233 1992-05-27 Germany 50.77 Daenkritz 12.43 262 Breil Sur Roya 923905232 1992-05-24 Germany 51.19 Ebersbach 14.93 262 Breil Sur Roya 923905234 1994-05-08 Germany 51.3 Foerst-gen 14.64 262 Breil Sur Roya 923905248 1993-04-29 Germany 49.84 Griesheimer Duene 8.59 262 Breil Sur Roya 923905246 1986-08-25 Germany 50.776 Heilsberg 11.265 262 Breil Sur Roya 923905226 1993-05-04 Germany 47.67 Isteiner Klotz 7.53 262 Breil Sur Roya 923905238 1992-05-28 Germany 50.77 Langenhessen 12.37 262 Breil Sur Roya 923905245 1996-08-04 Germany 53.123 Klein Schmoelen 11.289 262 Breil Sur Roya 923905240 1998-06-14 Germany 54.147 Lubminer Heide 13.647 262 Breil Sur Roya 923905247 1998-06-20 Germany 53.596 Neubrandenburg 13.409 262 Breil Sur Roya 923905253 1992-05-26 Germany 50.75 Niederhohndorf 12.47 262 Breil Sur Roya 923905252 1992-05-20 Germany 51.208 Rengersdorf 14.892 262 Breil Sur Roya 923905241 1993-05-08 Germany 19 47.717 Rohrdorf 10.083 262 Breil Sur Roya 923905224 1992-05-20 Germany 51.208 Torga 14.91 19 Breil Sur Roya 923905267 1987-05-31 Germany 51.81 Trebbichau 12.01 19 Breil Sur Roya 923905244 1993-05-03 Germany 48.1 Vogtsburg 7.7 19 Breil Sur Roya 923905235 1988-07-27 Germany 53.49 Waren / Feissnecksee 12.710 19 Breil Sur Roya 923905239 1991-10-24 Great Britain 50.657 Hartland Moor -2.092 19 Breil Sur Roya 923905231 2001-06-01 Greece 40.5 Agia Triada 22.882 19 Breil Sur Roya 923905237 2001-05-29 Greece 40.102 Litochoro 22.562 19 Breil Sur Roya 923905255 Greece 40.85 Nestos Delta 24.8 19 Breil Sur Roya 923905242 1998-05 Hungary 48.467 Aggtelek 20.517 19 Breil Sur Roya 923905265 1972-10-05 Italy 40.12 Baunei 9.51 19 Breil Sur Roya 923905251 1999-05-14 Italy 39.376 Camigli-atello Silano 16.441 19 Silano 923905260 1999-05-14 Italy 39.363 Camigliatello Silano 16.456 19 Silano 923905264 1999-05-16 Italy 39.361 Camigliatello Silano 16.43 19 Silano 923905256 1994-07-20 Italy 43.783 Firenze 11.6 19 Silano 923905257 1994-05-19 Italy 39.23 Largo Arvo 16.5 19 Silano 923905273 1994-05-03 Italy 40.85 Mte. Limbara 9.18 19 Silano 923905270 1999-05-17 Italy 39.392 Sila Grande 16.55 19 Sila Grande 923905250 1999-05-20 Italy 39.387 Sila Grande 16.559 19 Sila Grande 923905266 1999-05-21 Italy 39.411 Sila Grande 16.553 19 Sila Grande 923905271 2001-07-27 Kazakhstan 47.317 Manrak 84.617 19 Sila Grande 923905262 2001-08-05 Kazakhstan 47.13 Sarymobe 82.372 19 Sila Grande 923905272 2001-07-25 Kazakhstan 47.357 Saur 85.518 19 Sila Grande 923905276 2001-08-01 Kazakhstan 47.784 Tarbagatay 81.754 19 Tarbagatay 923905275 2001-08-05 Kazakhstan 47.133 Tarbagatay 82.367 19 Tarbagatay 923905261 2001-08-01 Kazakhstan 47.783 Tarbagatay 81.767 19 Tarbagatay 923905249 2002-03-15 Portugal 37.839 Porto Covo -8.788 19 Tarbagatay 923905282 1999-07-17 Russia 51.75 Kursk 36.167 19 Tarbagatay 923905268 1991-05-10 Spain 37.06 Veleta -3.36 19 Tarbagatay 923905259 1994-05-16 Switzerland 46.3 Pfynwald 7.62 19 Tarbagatay 923905254 Switzerland 46.54 Vaux 6.47 19 Tarbagatay 923905258 1993-06-18 Turkey 39.5 Catak 43.01 19 Tarbagatay 923905269 1993-06-23 Turkey 40.47 Dagpinar 43.330 19 Tarbagatay 923905277 1993-05-22 Turkey 36.87 Dirimli Gecidi 29.58 19 Tarbagatay 923905263 1993-07-03 Turkey 41.64 Dranoz 34.84 19 Tarbagatay 923905283 1993-07-03 Turkey 41.8 Kabali 35.05 19 Tarbagatay 923905274 1993-05-19 Turkey 38.35 Oedemis 27.88 19 Tarbagatay 923905280 1993-06-22 Turkey 39.8 Sac Gecidi 42.51 19 Tarbagatay 923905281 1993-06-28 Turkey 41.0 Sarigoel 41.5 19 Tarbagatay 923905278 1993-07-06 Turkey 40.06 Sogukpinar 29.12 19 Tarbagatay 923905279 2003-05-10 Turkey 41.468 Stranga Mts. 27.375 19 Tarbagatay 923905285 1995-08-16 Ukraine 44.45 Jalta 34.05 19 Tarbagatay 923905284 1995-08-12 Ukraine 44.617 33.867 19 Tarbagatay