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