identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
F52B87F65E316149FF74D870FA0D1DD9.text	F52B87F65E316149FF74D870FA0D1DD9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica rufa LINNAEUS 1761	<div><p>Key to the workers of the Palaearctic Formica rufa group:</p> <p>This key can certainly solve a good number of determinations in a rather simple way but the best results are achieved when own discriminant functions are run using the data of SI1 as reference. In contrast to Tables 1 - 5 and for more simplicity, this key works with primary, not allometrically corrected data, and all linear measurements are given in mm. Because of the enormous intraspecific and intranidal variation, nest sample means should be considered in critical cases. The percentage of reddish pigmentation follows a positive allometric trend and is of rather little indicative value. Figures 5-20 with z-stacking photos of the species from different viewing positions are presented after the key.</p> <p>1a With view on the swivelling plane of the first seg- ment of antennal funiculus, ratio of median length of second funiculus segment against its maximum width ≥ 2.0 (movement within only a swivelling plane is defined by the hinge joint of distal scape with the first funiculus segment). Lateral clypeus deeply depressed, as result anterior portion of lat- eral clypeus forming a bead. Medium to large-sized workers usually with whole surface of head, meso- soma and petiole light reddish brown. Formica truncorum group...................................................... 2</p> <p>1b Ratio of median length of second funiculus segment against its maximum width &lt;2.0. Lateral clypeus less deeply depressed and anterior portion of lateral clypeus not forming a bead. Percentage of surface with dark or blackish brown pigmentation usually larger.......................................................................... 5</p> <p>2a Iberian Peninsula and Pyrenees............................... 3</p> <p>2b Outside Iberian Peninsula and Pyrenees................. 4</p> <p>3a Gula and dorsal mesosoma without or with very few setae; nGu + nPn + nPr + nMet &lt;30................................................................... Formica dusmeti</p> <p>3b Gula and dorsal mesosoma with more setae; nGu + nPn + nPr + nMet&gt; 30.......... Formica frontalis</p> <p>4a More hirsute on all body positions. Discriminant 76.03 * EyeHL + 0.073 * nCH-0.057 * nGu + 0.08 * nMet + 12.36 * mPnHL - 14.75 * MetHL - 1.33&gt; 0 (error 0% in 92 individuals, Tab. 5)................................................................ Formica truncorum</p> <p>4b Less hirsute on all body positions. Often a disparity between rather few setae on dorsal mesosoma and rather many setae on gula is visible. Discriminant &lt;0 (error 0% in 70 individuals, Tab.5). Only E Tibet and China................................... Formica sinensis</p> <p>5a Palaearctic with exception of Russian Far East, NE China, Korea, Japan.................................................. 6</p> <p>5b Russian Far East, NE China, Korea, Japan........... 12</p> <p>6a Hind margin of head without or only occasional small setae, nCH 0 - 1; if nCH is slightly larger, then scape slender with SL / Smax 10.08 ± 0.39. Eyes with only short microsetae, EyeHL 0.020 ± 0.004. Mesopleuron with rather few setae, nMes 10.5 ± 7.4. Discriminant 0.024 * nCH + 0.08 * nMes - 0.046 * nPr - 11.451 * SL + 72.20 * Smax + 62.96 * EyeHL + 3.879 &lt;0 (error 0% in 114 nest sample means)........................................................................ 7</p> <p>6b Hind margin of head usually with many setae, nCH&gt; 1; if nCH is near zero (occasional in F. aquilonia), then scape thickset with SL / Smax 9.27 ± 0.34. Eyes with longer microsetae, EyeHL 0.030 ± 0.008. Mesopleuron with rather many setae, nMes 21.6 ± 10.4. Discriminant&gt; 0 (error 1.7% in 180 nest sample means)........................................................... 8</p> <p>7a Weakly haired; nest means: nGu 0.1 - 3.0, GuHL 0.007 - 0.097, nPn 0.1 - 5.6, mPnHL 0.006 - 0.055, nPr 0 - 5.2, nCH 0 - 0.9 (microsetae)...................................................................... Formica polyctena</p> <p>7b Moderately hairy; nest means: nGu 1.9 - 6.6, GuHL 0.096 - 0.197, nPn 5.8 - 16.0, mPnHL 0.047 - 0.083, nPr 4.4 - 11.2, nCH 0 - 1.2 (microsetae).................................................... Formica polyctena × rufa</p> <p>7c More strongly haired; nest means: nGu 5.1 - 11.0, GuHL 0.155 - 0.224, nPn 12.5 - 45.0, mPnHL 0.061 - 0.102, nPr 8.5 - 25.1, nCH 0 - 3.6 (setae small).............................................................. Formica rufa</p> <p>8a Longest propodeo-metapleural hair below level of propodeal spiracle shorter: MetHL 0 - 0.142. With maximum CL in focal plane, contour of head from median occiput to anterior eye margin with fewer setae: nCH 1.3 - 12.3. In doubtful cases: discriminant 0.0503 * nCH + 22.213 * MetHL -3.481 &lt;0 (error 0% in 75 nest samples)............................................................................. Formica aquilonia</p> <p>8b Longest propodeo-metapleural hair below level of propodeal spiracle longer: MetHL 0.134 - 0.237. With maximum CL in focal plane, contour of head from median occiput to anterior eye margin with more setae: nCH 5.2 - 65.2. Discriminant&gt; 0 (error 0% in 305 nest samples)............................................ 9</p> <p>9a Scape more compact: SL / Smax 8.45 - 10.07. Frons appears at low magnification not perfectly matt, with a mild silky shine. This overall impression is produced by a weaker microsculpture with more longitudinal and less reticulate elements, particularly along the frontal line and anteriolaterally from mid ocellus. With all measurements in mm, discriminant 2.524 - 3.89 * CW - 12.25 * SL + 5.889 * PeW + 117.36 * Smax&gt; 0 (error 0% in 291 nest samples for whole Palaearctic range). Boreo-montane species............................................................. 10</p> <p>9b Scape slender: SL / Smax 9.94 - 11.74. Frons matt. This overall impression is produced by a stronger more reticulate microsculpture. Discriminant &lt;0 (error 0% in 94 nest samples for whole Palaearctic range). More xerothermous woodland and woodland-steppe habitats................ Formica pratensis</p> <p>10a Only W Alps, ranging east to approximately 11° E. Pronotal setae shorter: mPnHL 0.061 - 0.093. Propodeo-metapleural area below level of propodeal spiracle with fewer and shorter setae: nMet 3.3 - 9.7, MetHL 0.130 - 0.174. Discriminant 48.8 * mPnHL + 16.6 * MetHL - 0.100 *nCH + 0.087 * nMet - 0.072 * nSc - 5.535 &lt;0 (error 0% in 70 nest sample means of four individuals, function only valid for Central Europe)................................................................................................ Formica paralugubris</p> <p>10b Widely distributed. Pronotal setae longer: mPnHL 0.079 - 0.127; nMet 6.5 - 14.8, MetHL 0.153 - 0.225. Discriminant&gt; 0 (error 3.1% in 98 nest sample means of four individuals; function only valid for Central Europe........................................................ 11</p> <p>11a Whole Palaearctic. Population from the Alps poorly separable from F. helvetica sp.n. Discriminant 36.64 * Smax + 0.128 * nMet + 0.110 * nSc + 0.068 * nCH - 53.54 * mPnHL - 3.224&gt; 0 (error 20.5% in 380 individuals and 14.6% in 89 nest samples)..................................... Formica lugubris</p> <p>11b Only known from a local population in Mingèr Valley in the Eastern Swiss Alps. Separation from F. lugubris very weak. Discriminant 36.64 * Smax + 0.128 * nMet + 0.110 * nSc + 0.068 * nCH - 53.54 * mPnHL - 3.224 &lt;0 (error 13.3% in 30 individuals and 11.1% in nine nest samples)..................................................................... Formica helvetica sp.n.</p> <p>12a Gula, mesopleuron, propodeum and metapleuron with many setae; nGu + nMes + nPr + nMet = 38 - 110. Discriminant 0.092 * nGu + 0.048 * nMes + 0.235 * nMet + 0.051 * nPr - 3.8554&gt; 0 (error 4.9% in 161 individuals and 0% in 44 nest samples)......................................................... Formica lugubris</p> <p>12b Gula, mesopleuron, propodeum and metapleuron with fewer setae; nGu + nMes + nPr + nMet = 3 - 46. Discriminant &lt;0 (error 0.9% in 424 individuals and 0% in 95 nest samples)............................................ 13</p> <p>13a Gular setae much longer, GuHL 0.217 ± 0.021; setae in all other body positions longer. Mesopleuron with fewer setae, nMes 6.6 ± 2.1. Discriminant 20.041 * GuHL + 30.455 * mPnHL + 2.661 * MetHL - 0.147 * nMes - 4.466&gt; 0 (error 0% in 38 individuals). Ussuri region....... Formica kupyanskayae</p> <p>13b Gular setae much shorter, GuHL 0.098 ± 0.033; setae in all other body positions shorter. Mesopleuron with more setae, nMes 13.3 ± 4.9. Discriminant &lt;0 (error 0% in 386 individuals)........................... 14</p> <p>14a Dorsal plane of scape and hind margin of head with rather many setae, nSc 10.5 ± 4.3, nCH 9.8 ± 4.2. Metapleuron with rather long setae, MetHL 0.133 ± 0.029. Discriminant 6.56 * PeW - 11.58 * Smax - 3.155 * CW + 0.013 * nCH + 0.357 * nSc + 3.505 * MetHL - 1.224&gt; 0 (error 9.8% in 41 individuals and 0% in 10 nest sample means). Ussuri and Sichote Alin range.................. Formica ussuriensis sp.n.</p> <p>14b Dorsal plane of scape and hind margin of head with very few setae, nSc 0.3 ± 1.3, nCH 4.3 ± 3.8. Metapleuron with short setae, MetHL 0.074 ± 0.047. Discriminant &lt;0 (error 1.0% in 102 individuals and 0% in 31 nest sample means)............................................................... Formica aquilonia</p></div> 	http://treatment.plazi.org/id/F52B87F65E316149FF74D870FA0D1DD9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E256152FC9FD94EFA0C1D19.text	F52B87F65E256152FC9FD94EFA0C1D19.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica meridionalis NASSONOV 1889	<div><p>Formica meridionalis NASSONOV, 1889</p> <p>Formica rufa var. meridionalis NASSONOV, 1889 [identification by DLUSSKY (1967)]</p> <p>This taxon was described from Kharkiv / E Ukraine and was synonymized by DLUSSKY (1967), who investigated types in the collection of ZMLU Moskva and stated these to be in all characters consistent with F. rufa. This is credible considering G.M. Dlussky’s good knowledge of wood ants and the species spectrum present near Kharkiv.</p> </div>	http://treatment.plazi.org/id/F52B87F65E256152FC9FD94EFA0C1D19	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E256152FC9FDFEEFA111E18.text	F52B87F65E256152FC9FDFEEFA111E18.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica obscurata SANTSCHI 1925	<div><p>Formica obscurata SANTSCHI, 1925</p> <p>Formica rufa var. obscurata SANTSCHI, 1925 [type investigation]</p> <p>This taxon was described from Vernon / France (49.09°N, 1.49° E, 19 m). Investigated were two type workers from NHM Basel labelled “type”, “ F. rufa L. v. obscura Sant type SANTSCHI det. 1925”, “France Vernon (Eure) G. d. Kerville ”; the specimen with CW = 1768 μm is addition- ally labelled with “ANTWEB CASENT 0912252 ”. If run as wild-card in an LDA, the type sample is allocated with p = 0.8677 to Formica rufa, with p = 0.1321 to Formica polyctena × rufa, and with p = 0.0001 to F. polyctena.</p> </div>	http://treatment.plazi.org/id/F52B87F65E256152FC9FDFEEFA111E18	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E256152FC9FDB2EFA101A39.text	F52B87F65E256152FC9FDB2EFA101A39.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica piniphila Schenck 1852	<div><p>Formica piniphila SCHENCK, 1852</p> <p>Formica piniphila SCHENCK, 1852 [description and zoogeography]</p> <p>This taxon was described from Hessen-Nassau. Types were not available. SCHENCK (1852) reported for the worker “mesosoma always homogenously covered by setae, eyes bare, mesosoma with two small, pale blackish spots, the latter often missing” and for the gyne “scutellum and gaster brilliantly shiny.” As SCHENCK (1852) correctly described differential characters of Formica polyctena, Formica pratensis, and Formica truncorum and because no other Formica rufa group species are expectable for Hessen-Nassau, the synonymy with F. rufa is obvious.</p> </div>	http://treatment.plazi.org/id/F52B87F65E256152FC9FDB2EFA101A39	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E256152FF5EDB7FFC4E1859.text	F52B87F65E256152FF5EDB7FFC4E1859.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica rufa LINNAEUS 1761	<div><p>Formica rufa LINNAEUS, 1761</p> <p>Formica rufa LINNAEUS, 1761</p> <p>[original description; YARROW (1954, 1955), photo of type specimen]</p> <p>Each of the two first descriptions of this ant(LINNAEUS 1758, 1761) are contradictory within themselves. The morphological description of the worker states in 1758: “Thorace compresso toto ferrugineo, capite abdominique nigris.” In 1761, the same statements are repeated and a supplementation is added: “Corpus fuscum. Thorax ferrugineus, compressus, squama intergerina ferruginea, acuminata.” This agrees with the condition in Camponotus herculeanus or Camponotus ligniperda, which both occur in Sweden. In contradiction to the morphological description, LINNAEUS stated “habitat in Europae acervis-acerosis sylvaticis” (LINNAEUS 1758) and “Piss Myror. SueciStackMyra...Habitat ubique in sylvis, acervos e foliolis acerosis exstruens.” (LINNAEUS 1761). It is obvious from these data that Linnaeus in both publications more likely intended to give the name F.rufa to those most abundant (“Habitat ubique in sylvis”), needle-thatch building (“Stack-Myra... acervos e foliolis acerosis exstruens.”) and acid squirting (“Piss Myror”) woodland ants. The description of the gyne presented in 1761 reads as follows: “Corpus nigricans. Caput subtus ferrugineum. Thorax ferrugineus dorso fusco. Abdomen segmentis quatuor, primo antice ferrugineo.” This description does not tell much but “Caput subtus ferrugineum” speaks against the two Swedish Camponotus species and is in agreement with the situation in ants of the F. rufa group. According to YARROW (1954), F. rufa group specimens were represented in the collection of the Linnaean Society London by a single worker bearing the label “rufa ex descr.”, two unlabelled winged females, and three unlabelled males. YARROW (1954) published a lectotype fixation in the better preserved of the two winged F. rufa group gynes but he made no statements on its characters. The reasons why he did not fix a lectotype in the only specimen labelled “rufa ex descr.” were not explained. I could not investigate the lectotype.As YARROW (1955) showed a rather good knowledge on the separation of the gynes of F. rufa, Formica aquilonia, Formica lugubris, and Formica pratensis, it appears most probable that he had a specimen at hand which indeed belonged to F. rufa as it is characterized here. Yet, as YARROW (1955) did not separate F. rufa and Formica polyctena, I inspected the picture of the lectotype (specimen number 2870) presented on the homepage of the Linnaean Society (http://linnean-online.org/16186/; retrieved on 31 October 2020). It shows a brilliantly shiny first gaster tergite, a shiny scutellum, and a massive, thickset body. This overall impression corresponds to the gyne morph of monogynous F. rufa and likely precludes the specimen representing F. polyctena. In order to unambiguously dissolve the confusion with Linnaeus’ descriptions, YARROW (1954) argued that “The Commission should use their plenary power to place F. rufa L., 1758 on the list of permanently rejected names and to place instead F. rufa L., 1761 on the Official List of Specific Names in Zoology”. This proposal was accepted by the International Commission of Zoological Nomenclature in an opinion published 2 October 1956.</p> </div>	http://treatment.plazi.org/id/F52B87F65E256152FF5EDB7FFC4E1859	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E256152FC9FDE6EFAA61C98.text	F52B87F65E256152FC9FDE6EFAA61C98.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica rufotruncicola WASMANN 1891	<div><p>Formica rufotruncicola WASMANN, 1891</p> <p>Formica rufa var. rufotruncicola WASMANN, 1891 [description of types by BETREM (1960)]</p> <p>This taxon was described from Panheel near Roermond / Netherlands on the basis of workers collected from a nest in April 1889. WASMANN (1891a) gave no description of structural characters. However, as WASMANN (1891b) reported on the mixed colouration of the worker population of exactly this nest, F. rufotruncicola WASMANN, 1891 is no nomen nudum. The data reported by BETREM (1960) on seven investigated syntype workers from NHM Maastricht indicate that it is a red colour variant of F. rufa.</p> </div>	http://treatment.plazi.org/id/F52B87F65E256152FC9FDE6EFAA61C98	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E256157FC9FDD6EFF751B59.text	F52B87F65E256157FC9FDD6EFF751B59.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica rufa subsp. emeryi STITZ 1939	<div><p>Formica rufa emeryi STITZ, 1939</p> <p>Formica rufa ab. emeryi STITZ, 1939 [description and zoogeography]</p> <p>STITZ (1939) made the first available use of F. rufa subsp. rufa ab. emeryi KRAUSSE, 1926. This taxon was described from near Eberswalde / Germany as specimens found within the nests of F.rufa showing a colouration as in Formica pratensis but with missing setae on eyes and hind tibia. Considering the species spectrum occurring in the vicinity of Eberswalde, these statements make a synonymy with F. rufa most likely. Types are unknown.</p> <p>All material examined. Numeric phenotypical data were recorded in 61 nest samples with 331 workers and 29 gynes; for details, see SI1, SI2, and SI3. The total number of mounted samples stored in SMN Görlitz and investigated either subjectively or by partial or complete numeric recording of the phenotypical characters used here was 103. These included 547 workers and 98 gynes and originated from Austria (two samples), Belgium (four), Bulgaria (six), Croatia (two), Finland (five), France (two), Germany (61), Greece (three), Italy (one), Norway (one), Poland (one), Russia (one), Slovenia (one), Spain (five), Sweden (four), and Switzerland (four). Character recording in ethanol-stored material according to the former investigation protocol of SEIFERT (1991) had been done until the year 1993 in further 196 nest samples with about 1600 workers, largely from Germany and Russia.</p> <p>Geographic range. From Iberia east to Baikal region; probably a little more widely distributed than Formica polyctena due to the larger potency for long-range single-queen flight dispersal and socially parasitic colony foundation (SEIFERT 1991, 2018). In Europe between 40.5° N (Spain), 63.5° N (Sweden), and 64.8° N (Finland). A morphologically aberrant population exists in Asia Minor and Caucasus (here rare). The altitudinal distribution in European mountains is not well known. According to credible reports, it ascends in the Southern Alps (46° N) to 1500 m and in Asia Minor (40° N) to 1900 m.</p> <p>Diagnosis of worker (Tab. 1, key). Large; mean and maximum CS over all social types 1891 and 2274µm: Scape rather long and slender, SL / CS 1750 0.939, SL / Smax 10.10. Setae on eyes short, EyeHL 1750 22µm. Setae on posterior margin of head usually missing and, if present, rather short, nCH 1750 0.81, OccHL 1750 23 µm (a population in the montane region of Blanský Les, S Bohemia – labelled “ nespori ” in the collection of SMN Görlitz – has above-average values of nCH and OccHL but does not form a cluster sufficiently separate from Formica rufa when all characters are considered). Gular, pronotal, mesopleural and propodeal setae always present and rather long, nGu 1750 6.3, GuHL 1750 188 µm, nPn 1750 20.2, mPnHL 1750 81 µm, nMes 1750 15.4, nPr 1750 12.9; setae on metapleuron few or absent and of medium length, nMet 1750 1.9, MetHL 1750 144 µm. Pigmentation: Head with genae and its ventral surfaces always light reddish, dorsal head caudad from about transverse level of eye centers, the area between frontal carinae and surface along the frontal carina usually blackish brown; mesosoma light reddish, often with a medium-sized dark brown patch on dorsal pronotum. Specimens with nearly all surfaces of head, the mesosoma, and frontal face of first gaster tergite reddish, reminiscent of the condition in Formica truncorum, may occur, most frequently in large workers of the most hairy phenotypes.</p> <p>Diagnosis of gyne (Tab. 6, Fig. 1). Medium-sized, CS 2140 µm; scape rather long and slender, SL / CS 0.868, SL / Smax 9.13. Setae on eyes short, EyeHL 25µm; setae on posterior margin of head always missing; gula without or with single setae of up to 280 µm length; pronotum without or single short setae of up to 44 µm length; meso- and metapleuron and frontal face of first gaster tergite without setae, if single setae are present these may have 200 - 244µm length; ventral surface of first gaster sternite usually with numerous long setae, nSt 18.8, StHL 350µm. Pubescence distance and distance of foveolae on paramedian dorsum of first gaster tergite rather high, sqPDG 9.00, FodG 57µm. Large parts of median and paramedian scutellum perfectly smooth and brilliantly shiny. Dorsum of gaster viewed at lower magnification always shiny. Pigmentation of head similar to worker; mesonotum, scutellum, and metanotum blackish brown; gaster black.</p> <p>Taxonomic comments and clustering results. The nomenclatoric separation of Formica rufa and Formica polyctena is maintained here for pragmatic reasons but this decision is problematic according to the data presented in the next paragraph. This pragmatism follows a functional argument: In their pure expression, F. rufa and F. polyctena represent most different mor- phologies, types of ecological adaptation, dispersal, and reproduction strategies, which call for a different naming. Several hundred papers have been published so far using the name F. polyctena. Giving up the name F. polyctena would mean a loss of information and would complicate communication about biological issues.</p> <p>A broad study considering 432 nest samples with 6100 worker ants and eight NUMOBAT characters and integrating intranidal phenotype composition as well as</p> <p>topographic, ecological, and biological information (SEIFERT 1991) provided evidence for frequent hybridization of Formica rufa and Formica polyctena. The adaptive advantage of the hybrid for conditions of fragmented forest systems hypothesized by the same author was made credible through mathematic modelling by HÖFENER &amp; al. (1996). The presence of the supposed hybrid cluster in East Germany was later convincingly confirmed by a study integrating NUMOBAT characters and nuclear DNA data (SEIFERT &amp; al. 2010), with a high agreement of classifications provided by morphometrics and nuDNA (Fig. 21). Data on microsatellite DNA of GYLLENSTRAND &amp; al. (2004) and SEIFERT &amp; al. (2010) suggest that backcrossing of the hybrid and introgression occur mainly with the F. polyctena parent. This biased gene flow towards F. polyctena was also to be expected on the basis of differential queen acceptance and mating behaviour of the species (GÖSS- WALD 1942, SEIFERT 1991).</p> <p>Hybrid frequencies of up to 28% for particular regions sum up to 6 - 8% over the whole European range and cause a dilemma in taxonomic decision making. A blind NC-clustering study of morphological data, ignoring any other source of information and the possibility of hybrid occurrence, provided misleading results (Fig. 22). The figure considers 169 nest samples with 960 workers from entire Europe and the characters CS, CL / CW 1750, SL / CS 1750, nCH 1750, OccHL 1750, nGu 1750, GuHL 1750, nPn 1750, mPnHL 1750, nMes 1750, nMet 1750, MetHL 1750, and nPr 1750. Assuming K = 2 (two species and no hybrids present), the disagreement of the exploratory data analyses with the controlling LDA was 4.1% in NC-Ward, 5.9% in NCpart.hclust, and 1.2% in NC-part.kmeans. This means an average error rate of 3.7% and suggests acceptance of heterospecificity of Formica rufa and Formica polyctena following the &lt;4% threshold recommended for NC-clustering by SEIFERT (2020a).Explicitly, NC-clustering confirms the F. polyctena cluster and a collective cluster formed by F. rufa and the hybrids. This clustering is largely explained by the high similarity of F.rufa and the hybrids in nGu 1750 and GuHL 1750 and a strong dissimilarity of F. polyctena regarding these characters. Assuming K = 3 (two parental species and a hybrid cluster), the disagreement is 13.6% in NC-Ward, 15.4% in NC-part.hclust, and 10.6% in NC-part. kmeans. Out of 53 hybrid samples and as mean of the three methods, 16.4% were allocated to the F. rufa and 2.5% to the F. polyctena cluster. These data show that the different algorithms of NC clustering cannot clearly expose hybrid samples and are likely to suggest a stronger divergence of the parental clusters than really given. A clear demonstration of hybrids with gaps to the parental clusters may be achieved in a vectorial space, either by running a PCA of RAV-corrected data or checking suspicious samples as wild-cards in an LDA. Yet, this proves only true when no or very few backcrosses are in the material (SEIFERT 1984, 1999, 2006, KULMUNI &amp; al. 2010, BAGHERIAN YAZDI &amp; al. 2012, SEIFERT 2019a, b). In the present case, the LDA of the 169 samples (Fig. 23) does not show obvious gaps separating the hybrid from the parental clusters, which may be explained by numerous backcrosses. The reader should be aware that the frequency of hybrids in this data set is about fivefold larger than the average figure expected for random sampling all over the European range. For hybridization with Formica lugubris, see section “Hybrids Formica lugubris × rufa ” (p. 174).</p> <p>Biology. See the condensed information in SEIFERT (2018).</p></div> 	http://treatment.plazi.org/id/F52B87F65E256157FC9FDD6EFF751B59	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E206157FC9FD8EEFB8F1AD8.text	F52B87F65E206157FC9FD8EEFB8F1AD8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica gaullei BONDROIT 1917	<div><p>Formica gaullei BONDROIT, 1917</p> <p>Formica gaullei BONDROIT, 1917 [type investigation]</p> <p>This taxon was described using workers collected at <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=2.112&amp;materialsCitation.latitude=48.925" title="Search Plazi for locations around (long 2.112/lat 48.925)">Mesnil-le-Roi</a> near St. Germain-en-Laye (48.925° N, 2.112° E, 65 m). Investigated were three worker syntypes collected in 1889 by de Gaulle at Mesnil-le-Roi and stored in MSNB Bruxelles. This sample showed the following posterior probabilities if run as wild-cards in an LDA: Formica polyctena 0.9976, F. polyctena × rufa 0.0024, Formica rufa 0.0002.</p> </div>	http://treatment.plazi.org/id/F52B87F65E206157FC9FD8EEFB8F1AD8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E206157FCF9DBDFFA0C1B99.text	F52B87F65E206157FCF9DBDFFA0C1B99.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica polyctena FOERSTER 1850	<div><p>Formica polyctena FOERSTER, 1850</p> <p>Formica polyctena FOERSTER, 1850 [description, zoogeography]</p> <p>This ant was described from Stolberg near Aachen / Germany where it formed a polydomous colony. FOERSTER (1850)’s description of setae condition and surface structures in the worker and gyne allows a clear differentiation from any Formica rufa group species present in this region.</p> </div>	http://treatment.plazi.org/id/F52B87F65E206157FCF9DBDFFA0C1B99	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E206157FF5ED83FFB7218B9.text	F52B87F65E206157FF5ED83FFB7218B9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica polyctenaxrufa	<div><p>Formica polyctena × rufa</p> <p>– hybrids and backcrosses</p> <p>All material examined. The full set of numeric phenotypical data was recorded in 55 nest samples with 345 workers and 16 gynes; for details, see SI1, SI2, and SI3. The total number of mounted samples stored in SMN Görlitz and investigated either subjectively or by partial or complete numeric recording of the phenotypical characters used here was 68. These included 453 workers and 18 gynes and originated from Austria (one sample), Bulgaria (three), Czechia (one), Finland (two), Germany (29), Great Britain (20), Poland (two), Russia (two), Sweden (one), and Switzerland (six). Character recording in ethanol-stored material according to the former investigation protocol of SEIFERT (1991) was done until the year 1993 in further 98 nest samples with about 2700 workers largely from Germany and Russia.</p> <p>Geographic range. Hybrids are expected to occur wherever the parental species are in contact and hybrid frequency is estimated over the whole range as 6 - 8%. However, there are big regional differences in hybrid frequency. In Britain, where the typical Formica polyctena and Formica rufa are absent, 95% of all samples are phenotypically intermediate and the whole population is supposed to consist of hybrids. The British hybrids are on average smaller and have longer gular setae than the continental hybrids (Tab. 1). Yet, describing them as a separate hybridogenous species is not justified as they cannot be separated from continental hybrids by any form of exploratory or hypothesis-driven data analysis. Anyway, it would be interesting to study their nuclear genome for genetic divergence during the time after the formation of the English Channel 7500 b.p.</p> <p>Diagnosis of worker (Tab. 1, key). Both the continental and the British population are in nearly all characters intermediate between the parental species (Tab. 1). Identification is in most cases possible by dis- criminant functions if sufficiently large nest samples are considered.</p> <p>Diagnosis of gyne (Tab. 6). Number and length of setae on average lower than in Formica rufa but on individual level often inseparable from either parental species.</p> <p>Taxonomic comments and clustering results. This issue was thoroughly discussed in section “ Formica rufa LINNAEUS, 1761 ” (p. 152).</p> <p>Biology. A brief report on the relations between the hybrid and the parental genotypes as well as on the biological properties and the probable adaptive advantage of the hybrid is given in SEIFERT (2018).</p></div> 	http://treatment.plazi.org/id/F52B87F65E206157FF5ED83FFB7218B9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E206156FC9FD9AEFF751F39.text	F52B87F65E206156FC9FD9AEFF751F39.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica minor GoSSWALD 1951	<div><p>Formica minor GÖSSWALD, 1951</p> <p>Formica minor GÖSSWALD, 1951 [description]</p> <p>This taxon was described from near Würzburg / Germany. GÖSSWALD (1951) apparently did not define type specimens but everything he reported in his lengthy treatise makes clear that he meant Formica polyctena.</p> <p>All material examined. The full set of numeric phenotypical data was recorded in 58 nest samples with 314 workers and 33 gynes; for details, see SI1, SI2, and SI3. The total number of mounted samples stored in SMN Görlitz and investigated either subjectively or by partial or complete numeric recording of the phenotypical characters used here was 69. These included 329 workers and 32 gynes and originated from Belgium (one), Finland (10), France (one), Germany (45), Poland (one), Russia (four), and Switzerland (seven). Character recording in ethanol-stored material according to the former investigation protocol of SEIFERT (1991) was done until the year 1993 in further 176 nest samples with about 1800 workers largely from Germany and the Moscow region.</p> <p>Geographic range. Whole range apparently similar to that of Formica rufa: Iberia to Lake Baikal. The clearly confirmed occurrence in Europe extends between 42° N und 61° N; absent from British Isles, Asia Minor and Caucasus. The northern distributional border in Fennoscandia and Siberia and the upper altitudinal limit in Central European mountains are not exactly known because of frequent confusion with Formica aquilonia and occurrence of F. aquilonia × polyctena hybrid populations. The putative northern limit in Finland is at 63° N or along the -10 °C January isotherm. Natural distribution in the Giant Mountains (Czechia) up to 800 m (here artificially introduced at 1020 m), in the Alps ascending to 1200 m at least.</p> <p>Diagnosis of worker (Tab. 1, key). Clearly smaller than Formica rufa, mean and maximum CS over all social types 1669 and 2067µm. Scape rather long and slender, SL / CS 1750 0.932, SL / Smax 1750 9.97. Setae on eyes short, EyeHL 1750 17µm; on posterior margin of head nearly always missing and, if present, usually of minute size; gular, pronotal, mesopleural and propodeal setae sparse and rather short, nGu 1750 1.46, GuHL 1750 51 µm, nPn 1750 2.5, mPnHL 1750 30 µm, nMes 1750 5.9, nPr 1750 2.3; setae on metapleuron usually absent. Pigmentation as in F. rufa but percentage of dark pigmentation on mesosoma on average slightly higher due to smaller size and positive allometry of reddish pigmentation.</p> <p>Diagnosis of gyne (Tab. 6, Fig. 2). Rather small; mean and maximum CS 2037 and 2165 µm; scape rather long and slender, SL / CS 0.864, SL / Smax 9.17. Setae on eyes short, EyeHL 21 µm; setae on posterior margin of head and gula always missing; pronotum bare, exceptionally without single short setae of up to 42 µm length; meso- and metapleuron and frontal face of first gaster tergite without setae, if single setae are present these may have 60 - 80 µm length; ventral surface of first gaster sternite with fewer setae than in Formica rufa, nSt 6.2, StHL 122 µm. Pubescence distance and distance of foveolae on paramedian dorsum of first gaster tergite high, sqPDG 12.2, FodG 58µm. Shiny surface of scutellum usually restricted to a small median stripe. Dorsum of gaster viewed at lower magnification shiny but usually less than in F. rufa, which is caused by very faint transverse microripples. In some specimens the microripples may be stronger developed somewhat reminiscent of the situation in Formica pratensis. Colouration as in F. rufa.</p> <p>Taxonomic comments and clustering results. For separation from Formica rufa and hybrids / backcrosses Formica polyctena × rufa, see section “ Formica rufa LINNAEUS, 1761 ” (p. 152) and for separation from Formica aquilonia and hybrids / backcrosses F. aquilonia × polyctena, see section “ Formica aquilonia × polyctena – hybrids and backcrosses” (p. 156).</p> <p>Biology. See the condensed information in SEIFERT (2018).</p></div> 	http://treatment.plazi.org/id/F52B87F65E206156FC9FD9AEFF751F39	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E216156FF5EDC5FFE3B1E58.text	F52B87F65E216156FF5EDC5FFE3B1E58.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica aquiloniaxpolyctena YARROW 1955	<div><p>Formica aquilonia × polyctena</p> <p>– hybrids and backcrosses</p> <p>Article 23.8 of ICZN regulates that a species-group name established for an animal later found to be a hybrid must not be used as the valid name for either of the parental species, even if it is older than all other available names for them. This excludes Formica major NYLANDER, 1849 and Formica constricta KARAVAJEV, 1926 to be considered as synonyms of Formica polyctena FOERSTER, 1850 or F. aquilonia YARROW, 1955.</p> </div>	http://treatment.plazi.org/id/F52B87F65E216156FF5EDC5FFE3B1E58	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E216156FF74DD2EFC301E78.text	F52B87F65E216156FF74DD2EFC301E78.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica major NYLANDER 1849	<div><p>Formica major NYLANDER, 1849</p> <p>Formica major NYLANDER, 1849 [type investigation]</p> <p>This taxon was described from the environs of Helsingfors (Helsinki). Three syntype workers on one pin labelled “Zool. Mus. H:fors Spec. typ. No 5423 Formica major Nyl. “, “major Nyl piniphila Schenck rufa auctt nec Bondr.”, “Mus. Hels. N:o 2676” do not carry an information on the sampling locality but the specimens fully match the morphological description of NYLANDER (1849). Considering their numeric data and by subjective impression, the type series of F. major cannot be hybrids Formica polyctena × rufa but represents either F. polyctena, Formica aquilonia × polyctena hybrids or backcrosses. This morphology-based idea is fully in line with the existence of large hybridization and introgression zone in southern Finland shown by nuDNA data (BERESFORD &amp; al. 2017). Run as wild-cards in a two-class LDA considering the characters CS, CL / CW 1750, SL / CS 1750, nCH 1750, OccHL 1750, nGu 1750, GuHL 1750, nPn 1750, mPnHL 1750, nMes 1750, nMet 1750, MetHL 1750, nPr 1750, SL / Smax 1750, EyeHL 1750, and nSc 1750, the type sample is allocated with p = 0.4867 to F. polyctena and with p = 0.5133 to the hybrid cluster. The marginal position in the hybrid cluster suggests F. major to represent a backcross of a F. aquilonia × polyctena hybrid with F. polyctena. A PCA plot considering the characters CS, SL / CS 1750, OccHL 1750, GuHL 1750, SL / Smax 1750, EyeHL 1750, and nSc 1750 provides the same impression (Fig.24). These investigations considered 27 nest samples and 148 workers of F. aquilonia × polyctena hybrids or backcrosses and 57 nest samples with 217 workers of F. polyctena. See also section “ Formica aquilonia YARROW, 1955 ” (p. 158) and Figure 25.</p> </div>	http://treatment.plazi.org/id/F52B87F65E216156FF74DD2EFC301E78	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E216158FC9FDD0EFE5B1B59.text	F52B87F65E216158FC9FDD0EFE5B1B59.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica constricta KARAVAJEV 1926	<div><p>Formica constricta KARAVAJEV, 1926</p> <p>Formica rufa var. constricta KARAVAJEV, 1926 [type investigation]</p> <p>Investigated were two syntype workers on one pin labelled „ Akmolin. ob. Kokchetav. g. bl. Borovoye. Bej-Bijenko”, “3269. coll Karavaiev”, “Form. (Form.) rufa v. constricta Karavaiev typ” (Karavajev’s handwriting), “ Syntypus Formica rufa constricta Karaw. ” (label of Radchenko) and three syntype workers on another pin labelled “3269. coll Karavaiev”, “ Syntypus Formica rufa constricta Karaw. ” (label of Radchenko); depository SIZ Kiev. The type locality Borovoye is situated in the geographic zone (see below) where hybrids between Formica aquilonia and Formica polyctena do regularly occur. The status of the types was tested using the same 16 characters as in the analysis in the previous section. If run as wild-cards in a three-class LDA and comparing F. aquilonia (class 1), hybrids and backcrosses F. aquilonia × polyctena (class 2) and F. polyctena (class 3), the type series is allocated to these classes with p = 0.0152, p = 0.8019, and p = 0.1829, respectively (Fig.25). These investigations considered 75 nest samples with 345 workers of F. aquilonia, 27 nest samples and 148 workers of F. aquilonia × polyctena hybrids or backcrosses, and 57 nest samples with 217 workers of F. polyctena. Under this setting, the type series of Formica major was allocated in a wild-card run to the three classes with p = 0.0021, p = 0.5244, and p = 0.4735, respectively.</p> <p>All material examined. The full set of numeric phenotypical data was recorded in 27 nest samples with 148 workers and 18 gynes. These originated from Finland (13), Kazakhstan (one), Mongolia (one), and Russia (12). For details, see SI1, SI2, and SI3.</p> <p>Geographic range. The geographic range where hybrids Formica aquilonia × polyctena are known to occur corresponds to the transition zone between temperate and boreal climate and ranges from 19.8° E (Aaland Isles / Finland) to 107° E (Baikal Region and Bogdkhan NP / Mongolia). With increasing continentality, this zone moves</p> <p>south: <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.0&amp;materialsCitation.latitude=53.2" title="Search Plazi for locations around (long 104.0/lat 53.2)">It</a> extends in Finland between 59.8 and 63.3° N, in West Siberia (about 62° E) between 53.1 and 60.4° N, and in Central Siberia (about 104° E) between 47.8 and 53.2° N. Hybrid occurrence and introgression should also occur in the Alps somewhere in the submontane or montane zone.</p> <p>Diagnosis of worker (Tab. 2, key). The hybrids are intermediate in all characters in which the parental species show notable differences (Tab.2).</p> <p>Diagnosis of gyne (Tab. 6). Except for one specimen, all examined gynes are from a single supercolony near Tvärminne / Finland in which the genetic structure was thoroughly investigated by KULMUNI &amp; al. (2010) and KULMUNI &amp; PAMILO (2014). The data in Table 6 are thus biased to a local situation and probably not representative for the Palaearctic hybrid population. Yet, the intermediate position of the hybrids becomes obvious in characters showing the most obvious differences of the parental species: SL / CS, SL / Smax, FodG, and GuHL.</p> <p>Taxonomic comments and clustering results. The earliest indications of this hybridization came from a West Siberian sample (SaNo 156, Yekaterinburg-1998-U23) combining a clear Formica polyctena phenotype with a mtDNA of Formica aquilonia (GOROPASHNAYA &amp; al. 2004) and observation of viable laboratory crosses (SORVARI 2006). In the time since then, hybridization of F. aquilonia and F. polyctena has been confirmed and thoroughly investigated by morphometric analyses and investigation of nuDNA (KULMUNI &amp; al. 2010, BERESFORD &amp; al. 2017). A nuDNA study of the latter authors in 17 sites within an area of 3000 km ² in South Finland revealed extremely frequent hybridization and introgression between F. aquilonia and F. polyctena. This investigation showed that nest populations with a hybrid history but having developed their nuclear DNA close to the situation in the F. aquilonia parent preferentially had the mtDNA of F. polyctena, whereas a majority of those with a nuDNA approaching F. polyctena had the mtDNA of F. aquilonia.</p> <p>Hybrids in the Irkutsk Region (Central Siberia) seem to form an own self-sustaining population – at least they are no result of a very recent hybridization because Formica polyctena is not confirmed so far to occur there and because its presence is unlikely for climatic reasons (mean January temperature -21°C).</p> <p>Results of clustering worker samples are commented above (Figs. 24, 25). Due to introgression, it is not possible to demonstrate three separate clusters. A PCA of the gynes, considering all 24 characters presented in Table 6, provided a rather good separation of the three entities (Fig. 26). However, there are no backcrosses in the gyne hybrid cluster in which 94% of specimens came from a single, isolated supercolony.</p></div> 	http://treatment.plazi.org/id/F52B87F65E216158FC9FDD0EFE5B1B59	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2F6158FF5ED83FFC4A1C99.text	F52B87F65E2F6158FF5ED83FFC4A1C99.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica aquilonia YARROW 1955	<div><p>Formica aquilonia YARROW, 1955</p> <p>Formica aquilonia YARROW, 1955 [description, photo of holotype, zoogeography]</p> <p><a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=-4.347&amp;materialsCitation.latitude=56.667" title="Search Plazi for locations around (long -4.347/lat 56.667)">This</a> taxon was described from Black Wood of Rannoch, Pertshire, Scotland (56.667° N, 4.347° W). YAR- ROW (1955)’s description of gynes and workers, the pictures of the holotype gyne in AntWeb (ANTWEB 2021) (CASENT0903277), and the geographic position of the type locality unquestionably indicate the identity of this taxon.</p> <p>All material examined. Numeric phenotypical data were recorded in 81 nest samples with 381 workers and 30 gynes. These originated from Austria (17 samples), Czechia (three), Finland (24), Scotland (one), Mongolia (five), Norway (three), Russia (19), Sweden (one), and Switzerland (eight). For details, see SI1, SI2, and SI3. The total number of samples numerically or subjectively investigated was 130.</p> <p>Geographic range. Eurosiberian-boreomontane. Continuous range from Northern Ireland and Scotland to <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=9.0&amp;materialsCitation.latitude=63.0" title="Search Plazi for locations around (long 9.0/lat 63.0)">East Siberia</a> (131° E), in Fennoscandia between 56.3 and 71° N, and in Siberia between 47.5 and 63° N. The montane range in Europe extends from SE to NW over the <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=9.0&amp;materialsCitation.latitude=63.0" title="Search Plazi for locations around (long 9.0/lat 63.0)">Rila Mountains, NW</a> Carpathians, Bohemian Forest, and the Eastern Alps westward to 9° E. In the Alps ascending to 2400 m. Main distribution in the Alps within the autochthonous distributional area of Larix (EICHHORN 1964).</p> <p>Diagnosis of worker (Tab. 2, key). Small; mean and maximum CS over all social types 1575 and 1902 µm. Scape short and rather thickset, SL / CS 1750 0.908, SL / Smax 1750 9.25. Setae on eyes rather short, EyeHL 1750 24 µm; setae on dorsal plane of scape usually absent or few, nSc 1750 usually 0 - 2; head margin behind eyes with few short setae which usually concentrate at the occipital corners, nCH 1750 5.1, OccHL 1750 64 µm; gular, pronotal, and propodeal setae sparse and rather short, nPn 1750 7.9,mPnHL 1750 42µm, nPr 1750 5.8; seta on lateral mesopleuron more numerous but on lateral metapleuron absent or very few and of moderate length, nMes 1750 14.7, nMet 1750 1.8, MetHL 1750 86µm.</p> <p>Diagnosis of gyne (Tab. 6, Fig. 8). Small; mean and maximum CS 2015 and 2173 µm. Scape short and thickset, SL / CS 0.810, SL / Smax 8.32. Setae on eyes rather short, EyeHL 29 µm; head margin behind eyes with very few short setae which usually concentrate at the occipital corners, nCH 2.1, OccHL 30 µm; gular, pronotal, mesopleural, and metapleural setae and those on frontal face of first gaster tergite few and rather short, nGu 3.1, GuHL 51 µm, PnHL 43 µm, nMes 2.5, nMet 1.4, MetHL 30 µm, nGfr 8.1, GfrHL 64 µm. Margin of petiole scale above spiracle with few short setae. Pigmentation without peculiarities. Dorsum of gaster shiny but less than in Formica rufa; foveolae on first gaster tergite more dense, FodG 27.7 µm.</p> <p>Taxonomic comments and clustering re-sults. Results of clustering are shown and commented in section “ Formica aquilonia × polyctena – hybrids and backcrosses” (p. 156). Frequent hybridization and introgression raise the question if F. aquilonia and Formica polyctena can be considered as separate species. One option would be to reduce them to subspecies with differing climatic adaptations – boreo-montane and frost-hardy in F. aquilonia, and temperate-planar-colline and less frosthardy in F. polyctena. I advocate here, for operational and pragmatic reasons, to stay with a nomenclatorial treatment as different species. Reticulate evolution in the Formica rufa group as a whole already produces a diffi- cult taxonomic situation which would be further complicated if we abandon the parsimonious binary naming. A third, radical solution, synonymizing F. aquilonia with F. polyctena and then, as a logical consequence (see section “ Formica rufa LINNAEUS, 1761 ”, p. 152), synonymizing these two taxa also with F. rufa, causes more problems than it solves. Speaking only of F. rufa would cause a loss of information on the structure of biodiversity and on the natural history of its elements. For hybridization with Formica lugubris and Formica paralugubris, see sections “Hybrids Formica aquilonia × lugubris ” (p.152) and “Hybrids Formica aquilonia × paralugubris ” (p.156).</p> <p>Habitat and biology. See the species profile in SEIFERT (2018).</p></div> 	http://treatment.plazi.org/id/F52B87F65E2F6158FF5ED83FFC4A1C99	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2F615BFCF9DFFFFA0C1F78.text	F52B87F65E2F615BFCF9DFFFFA0C1F78.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica ussuriensis Seifert 2021	<div><p>Formica ussuriensis sp.n.</p> <p>Etymology. Referring to the region of the river Ussuri in which the species occurs.</p> <p>Type material. Holotype worker plus four paratype workers labelled “ RUS: Ussuri: 43.24N, 133.39E 770m, ober. Povarotnaja-Tal, lehmiger Hangschutt, Sandboden mit Pioniervegetation, 1999.09.14 -050 (L.Kanter)”; 5 paratype workers labelled “ RUS: <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=133.21&amp;materialsCitation.latitude=43.01" title="Search Plazi for locations around (long 133.21/lat 43.01)">Ussuri</a>: 43.01N, 133.21E, 850m; S- Mt. Lysaya Sopka, dunkle Fichten-Tannen-Taiga, 1999.09.03 -037 (L.Kanter)”; 5 paratype workers labelled “ RUS: Ussuri: 43.24N, 133.39E, 555 m; ober. Povaratnaya-Tal; sandige Flussterrasse, 1999.09.14 -051 (L.Kanter)”; depository SMN Görlitz.</p> <p>All material examined. Numeric phenotypical data were recorded in 10 nest samples with 41 workers and three gynes. All these originated from Russian Far East. For details, see SI1, SI2, and SI3.</p> <p>Geographic range. Known are eight sites in the Ussuri River / Sichote Alin region situated at altitudes between 270 and 850 m, and latitudes between 43.0 and 48.3° N.</p> <p>Diagnosis of worker (Tab. 2, Figs. 5 and 6, key). S imilar to Formica aquilonia; slightly smaller, mean and maximum CS over all social types 1537 and 1808 µm. Scape as short but less thickset than in F. aquilonia, SL / CS 1750 0.903, SL / Smax 1750 9.51. Petiole on average wider than in any other species, PeW / CS 1750 0.519. Setae on eyes rather short, EyeHL 1750 26 µm; setae on dorsal plane of scape much more numerous than in F. aquilonia, nSc 1750 10.8; setae on head margin behind eyes more numerous than in F. aquilonia and slightly longer, nCH 1750 10.9, OccHL 1750 80 µm; gular, pronotal, propodeal, and metanotal setae sparse and short as in F. aquilonia with exception of clearly longer metanotal setae, nGu 1750 5.6, GuHL 1750 117µ m, nPn 1750 7.1, mPnHL 1750 47µ m, nPr 1750 3.5; mMet 1750 3.8, MetHL 1750 141µm; setae on lateral mesopleuron as numerous as in F. aquilonia, nMes 1750 15.4. Pigmentation without specific characters.</p> <p>Diagnosis of gyne (Tab. 6, Fig. 7). Clearly larger than Formica aquilonia, mean and maximum CS 2225 and 2357 µm. Hind margin and sides of head more linear, as result head shape more trapezoid. Scape short and thickset, SL / CS 0.806, SL / Smax 8.27. Setae on eyes rather short, EyeHL 33µm; setae on head margin behind eyes much more numerous than in F. aquilonia and slightly longer, nCH 15.0, OccHL 1750 65 µm; gular setae much more numerous and longer than in F. aquilonia, nGu 10.8, GuHL 107 µm; setae on metapleuron and frontal face of first gaster tergite much more numerous and much longer than in F. aquilonia, nMet 12.2, MetHL 167µm, nGfr 26.3, GfrHL 257µm. Distance of pubescence and of foveolae on paramedian surface of first gaster tergite as in F. aquilonia, sqPDG 10.12, FodG 25.3 µm. Pigmentation without diagnostic characters.</p> <p>Taxonomic comments and clustering results. The species is similar to Formica aquilonia and probably closely related. The geographically closest finding of F. aquilonia in the south of Amur Oblast’ (49.16° N, 130.67° E) is situated about 340 km west of the next known site of Formica ussuriensis at 48.22° N, 135.06° E. Accordingly, occurrence of hybridization and introgression has to be checked in the region of the lower course of river Amur. Yet, the data currently available provide clearly separate clusters in exploratory data analyses. Considering the characters CL / CW 1750, SL / Smax 1750, nSc 1750, nCH 1750, OccHL 1750, nMet 1750, MetHL 1750, and nPr 1750, 10 worker nest samples of F. ussuriensis were separated from 75 Panpalaearctic samples of F. aquilonia by NC-Ward, NC-part.kmeans, NC-NMDS-kmeans, and a PCA with an error rate of 0%, whereas NC-part.hclust misclassified 2.4% of the samples (Fig. 27). Using the same character system, an LDA classified 98.7% of 386 worker individuals in agreement with the classification provided by four ex- ploratory data analyses. Regarding gynes, the first factor of PCA considering the characters CS, EyeHL, nCH, nMet, and MetHL scored 2.663 ± 0.221 (2.413, 2.832) in three gynes of F. ussuriensis but -0.275 ± 0.515 (-0.913, 0.784) in 29 gynes of F. aquilonia. A full separation by the PCA was also given when all 24 characters shown in Table 6 were considered unselectively.</p> <p>Habitat and biology. It formed monodomous and polydomous colonies and was found in often very dark spruce (Picea), fir (Abies), and broadleaf forests. These biomes have a boreal to subboreal character due to the influ- enceofcoldwatersofthePacificOcean.Thisleadstoashiftof biomes to more southern latitudes and lower altitudes than observed in comparable biomes in Europe. The 10 known sites are situated at altitudes of 622 ± 213 (270, 850) m.</p> </div>	http://treatment.plazi.org/id/F52B87F65E2F615BFCF9DFFFFA0C1F78	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2C615AFCF9DC1FFD5818D9.text	F52B87F65E2C615AFCF9DC1FFD5818D9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica pratensis RETZIUS 1783	<div><p>Formica pratensis RETZIUS, 1783</p> <p>Formica pratensis RETZIUS, 1783 [concept of FOREL (1874), present neotype designation]</p> <p>Type specimens are unknown and the original description of RETZIUS (1783), reporting only “rufa, capite abdomineque nigris, petiolo abdominis squamifero”, does not allow an identification even of the genus. The tradition which ant has to be named as F. pratensis was founded by FOREL (1874) who reported for the worker a large extension of the dark patch on dorsal mesosoma, a longer pilosity compared with Formica truncoru m, presence of setae on legs and eyes, and, for the gyne, a completely matt, densely pubescent gaster. This character combination excludes any of the other seven species occurring in Switzerland. FOREL (1874) also reported as main habitat grassland and wood margins, which probably prompted him to assign A.J. Retzius’ name to this ant. In order to stabilize the nomenclature, a neotype was fixed from a nest sample con- taining 18 workers, labelled “GER: 51.4048° N, 14.8746° E, Daubitz- 3.3 km ENE, 162 m, flacher HÜgel, R.Schultz 1999.06.30 -050” and “ Neotype Formica pratensis Retzius 1783 des. B. Seifert 2020 ”; depository SMN Görlitz.</p> </div>	http://treatment.plazi.org/id/F52B87F65E2C615AFCF9DC1FFD5818D9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2D615AFC9FDAAEFC261C39.text	F52B87F65E2D615AFC9FDAAEFC261C39.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica angusticeps STARCKE 1947	<div><p>Formica angusticeps STÄRCKE, 1947</p> <p>Formica rufa var. angusticeps STÄRCKE, 1947 [description, zoogeography]</p> <p>The taxon was described from the Netherlands. BOL- TON (1995) considered the name as available. The title of the paper “De boreale form van de roode boschmier (Formica rufa rufa)” suggests that STÄRCKE (1947) intended to introduce the new name at infrasubspecific rank. Yet, the main text did not make it clear what his intention was. If the name is available, we can assume a synonymy with Formica pratensis based on the following argumentation. The type locality is in Hoge Veluwe (51.08° N, 5.83° E, 38m) – a sand dune area with interspersed moister parts. We have only four species of the F. rufa group potentially occurring in that region: F. rufa, Formica polyctena, F. pratensis, and Formica truncorum. Formica truncorum is extremely rare in the Netherlands and STÄRCKE (1947) would have noted the diagnostic pigmentation. Therefore, he would have referred to one of the other three species. The reported presence on extensor profile of tibiae of 5 - 11 setae which are erected by 30 - 45° clearly speaks against F. rufa, F. polyctena, or F. polyctena × rufa. STÄRCKE (1947) gave no data on pilosity on back of head but he compared his F. angusticeps with specimens of “ F. rufa rufa ” from the Norwegian coast near the Lofoten (68° N) and of “ F. rufa alpina Santschi ” from the high Alps. For zoogeographical and morphological reasons, his specimens from the Lofoten obviously belonged to Formica lugubris and those from the high Alps to either F. lugubris or Formica paralugubris. These data implicate F. angusticeps to have a rich overall pilosity as it is typical for F. pratensis and to be hairier than in the hairiest F. rufa phenotypes.</p> </div>	http://treatment.plazi.org/id/F52B87F65E2D615AFC9FDAAEFC261C39	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2D615AFF74DE91FD301F58.text	F52B87F65E2D615AFF74DE91FD301F58.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica cordieri BONDROIT 1917	<div><p>Formica cordieri BONDROIT, 1917</p> <p>Formica cordieri BONDROIT, 1917 [type investigation]</p> <p>This taxon was described by BONDROIT (1917) as gyne in the key. No type locality, collector, nor date was given. Examined were three type specimens from MSNB Bruxelles: one gyne labelled “Orne-6-Longny Collection E.Cordier”, “ Formica cordieri Type Bondr.”; one gyne labelled “Hte Savoie de Gaulle”, “ Formica v. cordieri Type Bondr.”, and one gyne labelled “Sayat P. de D.”, “ Formica v. cordieri Type Bondr.”. The synonymy with Formica pratensis is obvious: All three gynes belong to the hairy N-morph as defined by SEIFERT (1992). For separation of N- and P-morphs, see also below and Figure 29.</p> </div>	http://treatment.plazi.org/id/F52B87F65E2D615AFF74DE91FD301F58	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2D615AFF74DC2EFEF21E19.text	F52B87F65E2D615AFF74DC2EFEF21E19.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica grouvellei BONDROIT 1918	<div><p>Formica grouvellei BONDROIT, 1918</p> <p>Formica grouvellei BONDROIT, 1918 [type investigation]</p> <p>Investigated was the type gyne from MSNB Bruxelles labelled “Digne Grouvelle”, “ F.rufa var. grouvellei Type Bondr.”. The type belongs to the hairy N-morph as defined by SEIFERT (1992).</p> </div>	http://treatment.plazi.org/id/F52B87F65E2D615AFF74DC2EFEF21E19	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2D615AFF74DBAEFEB51DF9.text	F52B87F65E2D615AFF74DBAEFEB51DF9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica nigricans BONDROIT 1912	<div><p>Formica nigricans BONDROIT, 1912</p> <p>Formica pratensis var. nigricans BONDROIT, 1912 [photos of lectotype, zoogeography]</p> <p>This is the first available use of Formica rufa pratensis var. nigricans EMERY, 1909. EMERY (1909) reported as collecting sites “eine südliche Form; aus den Seealpen und aus Spanien, auch im Apennin (Vallombrosa)”. A lectotype was fixed in worker specimen stored in MHN Genève labelled “Cotypus”, “ Formica rufa pratensis var. nigricans Em Vallombrosa ” (C. Emery’s handwriting), “ v. nigricans Em ” (A. Forel’s handwriting), and “ MHNG ENTO 00085011”. The characters revealed in the photos in combination with zoogeography clearly indicate that F. pratensis var. nigricans is a junior synonym of F. pratensis and not of Formica lugubris. The type locality Vallombrosa (43.73° N, 11.56° E, 950 m) is situated in a geographic region, as it is with the whole Apennine, where boreomontane species of the F. lugubris species complex did not occur in the times of C. Emery (BARONI URBANI 1971). All populations of boreomontane wood ants from the Apennine mountains known today go back to a massive artificial introduction performed during the years 1959 - 67 (e.g., PAVAN 1959).</p> </div>	http://treatment.plazi.org/id/F52B87F65E2D615AFF74DBAEFEB51DF9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2D615AFC9FDF4EFB4C1FF9.text	F52B87F65E2D615AFC9FDF4EFB4C1FF9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica pratensoide GOSSWALD 1951	<div><p>Formica pratensoides GÖSSWALD, 1951</p> <p>Formica minor ssp. pratensoides GÖSSWALD, 1951 [description, zoogeography]</p> <p>This taxon was described from near Würzburg (Germany): “Revierförsterei Irtenberg, Forstamt Waldbrunn”. GÖSSWALD (1951) reported a large, clearly demarcated black patch on promesonotum of the workers and gynes having 110 - 115% of the size of Formica rufa and a completely matt gaster surface. It becomes obvious from his lengthy treatise that GÖSSWALD (1951) studied a polygynous-polydomous colony of Formica pratensis. Such colony types up to true supercolonial conditions have been repeatedly observed in forests of the planar and colline zone of Germany (SEIFERT 1992, 2018).</p> </div>	http://treatment.plazi.org/id/F52B87F65E2D615AFC9FDF4EFB4C1FF9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2D615DFC9FDC8EFEBD1839.text	F52B87F65E2D615DFC9FDC8EFEBD1839.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica staerckei BETREM 1960	<div><p>Formica staerckei BETREM, 1960</p> <p>Formica nigricans var. staerckei BETREM, 1960 [description, zoogeography]</p> <p>It was described from a gyne collected by E. Wasman on 30 May 1885 near Castle Exaten at Baexem (Netherlands). BETREM (1960), using low-resolution microscopes, mentioned hairy eyes, absence of long hairs on head, a “practically hairless” mesosoma, and a matt surface of scutellum and gaster. This corresponds to the P-morph of Formica pratensis. The type specimen should exist in NHM Maastricht but was not available for investigation.</p> </div>	http://treatment.plazi.org/id/F52B87F65E2D615DFC9FDC8EFEBD1839	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2D615AFF74DD6EFC6D19D9.text	F52B87F65E2D615AFF74DD6EFC6D19D9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica thyssei STARCKE 1942	<div><p>Formica thyssei STÄRCKE, 1942</p> <p>Formica pratensis ab. thyssei STÄRCKE, 1942 [type investigation]</p> <p>This taxon was described from the Netherlands in a gyne collected at Eerbeek op de Veluwe, June 1916, leg. F. T. Valck Lucassen. This type from NBC Leiden was in- vestigated and belongs to the less hairy P-morph as defined by SEIFERT (1992).</p></div> 	http://treatment.plazi.org/id/F52B87F65E2D615AFF74DD6EFC6D19D9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E2A615FFF74DB4EFF751FB9.text	F52B87F65E2A615FFF74DB4EFF751FB9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica nigropratensis BETREM 1962	<div><p>Formica nigropratensis BETREM, 1962</p> <p>Formica nigropratensis BETREM, 1962 [unnecessary replacement name]</p> <p>This new name is an objective junior synonym as it was referred by BETREM (1962) simultaneously to the available names Formica thyssei STÄRCKE, 1947, Formica pratensoides GÖSSWALD, 1951, and Formica staerckei BETREM, 1960.</p> <p>All material examined. Numeric phenotypical data were recorded in 96 nest samples with 331 workers and 21 gynes. These originated from Bulgaria (seven samples), Czechia (one), Georgia (one), Finland (two), France (six), Germany (37), Hungary (two), Italy (one), Kazakhstan (four), Kyrgyzstan (three), Poland (one), Romania (two), Russia (21), Sweden (seven), and Switzerland (one). For details, see SI1, SI2, and SI3. Character recording in ethanol-stored material according to the former investigation protocol of SEIFERT (1992) had been done until the year 1991 in 224 nest samples with 1756 workers and 295 gynes originating from Europe.</p> <p>Geographic range. <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.0&amp;materialsCitation.latitude=42.2" title="Search Plazi for locations around (long 104.0/lat 42.2)">Continuously</a> distributed through the temperate and submeridional zones of the <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.0&amp;materialsCitation.latitude=42.2" title="Search Plazi for locations around (long 104.0/lat 42.2)">Palaearctic</a>, from Spain (9° W) to <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.0&amp;materialsCitation.latitude=42.2" title="Search Plazi for locations around (long 104.0/lat 42.2)">Irkutsk</a> (104° E) at least. <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.0&amp;materialsCitation.latitude=42.2" title="Search Plazi for locations around (long 104.0/lat 42.2)">In Europe</a> from 37° N (S Spain) to 63.9° N (<a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.0&amp;materialsCitation.latitude=42.2" title="Search Plazi for locations around (long 104.0/lat 42.2)">Fennoscandia</a>). In the Alps ascending to 1500 m (46.0° N), in the Pyrenees and Bulgaria to 1800 m (42° N), and in the Tian Shan to 2100m (42.2° N). Reports from higher elevations in Europe should be checked for confusion with Formica lugubris.</p> <p>Diagnosis of worker (Tabs. 3 and 4, Figs. 9 and 10, key). Dimorphic, with P and N morphs frequently occurring within the same nest. According to the data of SEIFERT (1992), then measured in ethanol-stored specimens, the less hairy P-morph had nCH 17.9 ± 5.7, OccHL 103 ± 25 µm, nHT 17.8 ± 5.7, and CS 1820 ± 220 (1050, 2250) µm in 962 workers, and the hairier N-morph nCH 28.2 ± 6.4, OccHL 132 ± 20 µm, nHT 25.1 ± 5.1 and CS 1770 ± 220 (1040 - 2180) µm in 794 workers. Large species; mean and maximum CS over all social types and both morphs in dry mounted specimens (with slight bias to selecting larger specimens) 1819 and 2239 µm. Head elongated, CL / CW 1750 1.111. Scape much longer and slender than in Formica lugubris, SL / CS 1750 0.927, SL / Smax 1750 10.66. Petiole scale clearly narrower than in F. lugubris, PeW / CS 1750 0.453. Setae number and length extremely variable but even in the least hairy phenotypes larger than in Formica rufa; separation from F. lugubris by seta characters impossible due to extreme setae polymorphism also in this species; for variance of setae data, see Tables 3 and 4. All body surfaces except the frontal triangle matt due to developed microsculpture. The blackish patch on promesonotum is often larger than on average seen in other species and is often sharply demarcated from the reddish surface.</p> <p>Diagnosis of gyne (Tab. 6; Figs. 4, 11, 12). Dimorphic, P- and N-morphs frequently occurring within the same nest, and more clearly separable than workers. According to the data of SEIFERT (1992), then measured in ethanol-stored specimens, the less hairy P-morph had nCH 0.2 ± 0.8, OccHL 30 ± 20µm, nHT 0.6 ± 1.0, and CS 2290 ± 80 (2090, 2511) µm in 172 gynes, and the hairier N-morph nCH 16.2 ± 14.1, OccHL 218 ± 82 µm, nHT 8.4 ± 4.3, and CS 2250 ± 70 (2040 - 2400) µm in 123 gynes. Large; mean and maximum CS over all social types and both morphs in 21 mounted specimens 2296 and 2432 µm. Head moderately elongated, CL / CW 1.024. Scape much longer and more slender than in species related to Formica rufa, Formica aquilonia, or Formica lugubris, SL / CS 0.853, SL / Smax 9.75. Petiole scale relatively narrow, PeW / CS 0.628. Strong pilosity dimorphism: setae number, distribution, and length extremely variable (SEIFERT 1992). Setae on eyes always present and rather long to very long, EyeHL 46 - 86µm. The least hairy gynes of the P-morph have no setae on posterior margin of head, scape, scutellum, propodeum, petiole scale above spiracle, frontal face of first gaster tergite, and extensor profile of hind tibiae. The hairiest gynes of the N-morph have an extremely rich pilosity on nearly all body surfaces with seta length reaching 453µm on scutellum and 432µm on frontal face of first gaster tergite.All body surfaces, with exception of the frontal triangle, matt due to developed microsculpture. At magnifications&gt; 100x, the dorsum of gaster tergites shows strong transverse microripples and a dense pubescence; sqPDG 4.0 ± 0.5.</p> <p>Taxonomic comments and clustering results. Combining big size, a long and slender scape, hairy eyes, and dense transverse microripples on gaster tergites, gynes are easily separable from any species except for the East Palaearctic sister species Formica kupyanskayae (for identification, see section “ Formica kupyanskayae BOLTON, 1995 ”, p. 163). Separation of Formica pratensis workers from those of Formica lugubris is safely possible throughout the Palaearctic range using the characters CS, CL / CW 1750, SL / CS 1750, SL / Smax 1750, PeW / CS 1750, nSc 1750, nCH 1750, OccHL 1750, mPnHL 1750, nMet 1750, and MetHL 1750. Exploratory data analyses considering these characters could clearly distinguish 225 nest samples with 1059 workers of F. lugubris from 77 nest samples with 266 workers of F. pratensis. Classification errors were 0.6% in NC-part.hclust, 0% in both NC-part.kmeans and NC-Ward (Fig. 28), and 2% in a PCA. Separation on individual level by an LDA was also very strong with only 1.6% misclassification in 1325 worker individuals.Yet, the separation of the two clusters in two-dimensional plots of LDA and PCA was not strong enough to allow conclusions on recent hybrid samples. For repeated hybridization of the two species in the past, see section “Hybrids Formica pratensis × lugubris ” (p. 174).</p> <p>The strong dimorphism in Formica pratensis is most apparent in gynes. A re-analysis of the data of SEIFERT (1992) consisting of the 11 characters head width and number and maximum length of setae on posterior</p> <p>margin of head, scutellum, extensor part of hind tibia, propodeum, and frontal face of first gaster tergite was performed here. As result, the setae-reduced P-morph and the hairy N-morph can be clearly clustered in a PCA after logarithmic transformation of the raw data (Fig. 29). The low percentage of doubtful (or intermediate) specimens is also indicated by the LDA that classified only 2.4% of 295 gynes with posterior probabilities &lt;0.95. Clear arguments against considering the P-and N-morph as separate species are provided by the presence of both morphs in 21.6% of 37 nest samples and simultaneous observation of sexuals of both morphs at the same mating places (SEIFERT 1992). The same author also showed that phenotype dimorphism is correlated with ecological adaptations. He explained the demonstrated statistical differences in geographical distribution along a thermal gradient by selection of genotypes with differing climatic adaptations. According to SEIFERT (1992), the P-morph differs from the N-morph by the following traits: It constructs clearly flatter mounds for equal insolation conditions and goes to higher altitudes and latitudes. In Germany, the P-morph is rarer than the N-morph in dry habitats with typical Mediterranean elements but is more frequent than the N-morph on loamy soils.</p> <p>Biology. See the species profile given by SEIFERT (2018).</p></div> 	http://treatment.plazi.org/id/F52B87F65E2A615FFF74DB4EFF751FB9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E28615EFF5EDCDFFEFB1D58.text	F52B87F65E28615EFF5EDCDFFEFB1D58.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica kupyanskayae Bolton 1995	<div><p>Formica kupyanskayae BOLTON, 1995</p> <p>Formica kupyanskayae BOLTON, 1995 [type investigation]</p> <p>Replacement name for Formica opacus KUPYANSKAYA, 1980 that is a junior primary homonym of F. opaca NY- LANDER, 1856 (now in Camponotus MAYR, 1861).</p> <p>Type material. Five paratype workers and two paratype gynes from the holotype sample labelled “ Primorye, Anisimovka, 12.7.1975 Kupyanskaya ”, “Paratypus Formica opaca Kupyanskaya ”; depository SMN Görlitz.</p> <p>All material examined. Numeric phenotypical data were recorded in 10 nest samples with 41 workers and two gynes plus one isolated gyne. All these originated from Russian Far East. For details, see SI1, SI2, and SI3.</p> <p>Geographic range. Known are 10 sites in the Ussuri River / Sichote Alin region and one site in Sakhalin. These eleven sites are situated at altitudes of only 180 ± 150 [40, 530] m which is clearly lower than in sympatric Formica ussuriensis sp.n. (ANOVA F 1,19 30.6, p &lt;0.001).</p> <p>Diagnosis of worker (Tab. 2, key). Large, mean and maximum CS over all social types 1745 and 2068 µm. Scape rather long but less slender than in Formica pratensis, SL / CS 1750 0.917, SL / Smax 1750 9.78. Petiole width medium, PeW / CS 1750 0.476. Setae on eyes long, EyeHL 1750 35µm; setae on dorsal plane of scape absent or very few, nSc 1750 0.68; setae on head margin behind eyes less numerous than in F. pratensis but rather long, nCH 1750 5.8, OccHL 1750 118 µm; gular setae always present and long, nGu 1750 7.0, GuHL 1750 217µm; pronotal setae moderately numerous and rather long, nPn 1750 18.9, mPnHL 1750 98µm, number of mesopleural, propodeal, and metanotal setae clearly smaller than in F. pratensis but of comparable length, nMes 1750 6.6, nPr 1750 9.0; mMet 1750 3.4, Methl 1750 198 µm. Pigmentation without specific characters.</p> <p>Diagnosis of gyne (Tab. 6, Fig. 13). In many characters similar to Formica pratensis. Very large, mean and maximum CS 2348 and 2386 µm. Scape very long and slender, SL / CS 0.903, SL / Smax 9.66, absolute scape length&gt; 2090 µm, exceeding data in other species. Setae on eyes long, EyeHL 44 µm; setae on head margin behind eyes much fewer and shorter than in F. pratensis, nCH 4.8, OccHL 1750 87µm; gular setae always present and long, nGu 9.3, GuHL 258µm; setae on pronotum very few and short, PnHL 56 µm; setae number on mesopleuron, metapleuron, petiole above spiracle, frontal face of first gaster tergite, and flexor profile of hind tibia lower than in F. pratensis, nMes 2.7, nMet 4.8, nPe 0.2, nGfr 2.3, nHT 2.5; setae length on metapleuron and frontal face of first gaster tergite shorter than in F. pratensis, MetHL 125 µm, GfrHL 44 µm. All body surfaces matt; dorsum of first gaster tergite with dense pubescence, well-developed transverse microripples, and narrowly spaced foveolae as bases of pubescence hairs, sqPDG 3.6, FodG 20.5 µm. Most surfaces light reddish brown; posterior dorsum of head and mesonotum dark reddish brown; scutellum, metanotum, and gaster blackish brown, except for light reddish brown frontal face of first gaster tergite.</p> <p>Taxonomic comments and clustering results. Formica kupyanskayae is obviously the closest known relative of Formica pratensis but well separable. The 10 nest samples of F. kupyanskayae can be clearly separated from 76 nest samples of F. pratensis by all five exploratory data analyses considering the characters CS, CL / CW 1750, SL / CS 1750, nCH 1750, OccHL 1750, mPnHL 1750, nMet 1750, MetHL 1750, PeW / CS 1750, SL / Smax 1750, and nSc 1750 (Fig.30). The classification error by an LDA using the same character set was 0.7% in 301 worker individuals. Gynes of the two species can be strongly separated by a PCA considering the characters SL / CS, GfrHL, nPe, nMet, and ML / CS.</p> <p>-</p> <p>-</p> <p>Habitat and biology. There are no data on habitat selection available except for one record reporting a probably monogynous nest found in a bushy grassland with groups of trees. This selection of an open habitat and distribution at low altitudes suggests that it is a rather thermophilous species – another character it shares with Formica pratensis.</p> </div>	http://treatment.plazi.org/id/F52B87F65E28615EFF5EDCDFFEFB1D58	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E29615EFF74DD2EFA891A38.text	F52B87F65E29615EFF74DD2EFA891A38.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica congerens NYLANDER 1846	<div><p>Formica congerens NYLANDER, 1846</p> <p>Formica congerens NYLANDER, 1846 [type investigation]</p> <p>NYLANDER (1846) described this species from the island Mjölön near Helsinki. Five syntypes from FMNH Helsinki were investigated. These were two worker syntypes on the same pin labelled “H:Fors \ W. Nyland. \ Coll. Nyland \ congerens Nyl. \ Mus. Zool. H:fors Spec. typ. No 5023 Formica congerens Nyl. ”, ZM Helsinki. In the upper specimen, petiole and gaster are missing. The lower specimen with CS 2060µm is designated herewith as lectotype and the upper one with CS 2152 µm is the paralectotype. When run as wild-cards in the two-class LDA described in section “ Formica pratensis RETZIUS, 1783 ” (p. 159) and separating Formica lugubris and F. pratensis, the lectotype and paralectotype are classified as F. lugubris with posterior probabilities of p = 0.9921 and p = 1.0000. Three designated syntypes (now paralectotypes) on a second pin, with the lower specimen broken in two parts and badly glued on a card board, are labelled “ Mus. Zool. H:fors Spec. typ. No 5022 Formica congerens Nyl. ”. There is no label indicating the sampling site. The two undamaged specimens were clearly classified as F. pratensis with posterior probabilities of 0.9998 and 0.9944.</p> </div>	http://treatment.plazi.org/id/F52B87F65E29615EFF74DD2EFA891A38	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E29615EFF5EDE3FFF7C1E59.text	F52B87F65E29615EFF5EDE3FFF7C1E59.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica lugubris ZETTERSTEDT 1838	<div><p>Formica lugubris ZETTERSTEDT, 1838</p> <p>Formica lugubris ZETTERSTEDT, 1838 [diagnosis of YAR- ROW (1955)]</p> <p>The taxon was described from a male from Ofotford (Norway, 68.43° N, 17.03° E). ZETTERSTEDT (1838) ’s description matches all three species expected to occur at the Ofotford: F. lugubris, Formica aquilonia, and Formica truncorum. YARROW (1955), who was the first after 120 years to revive F. lugubris from synonymy and to give it a status of a good species, did not explain on which basis he identified J.W. Zetterstedt’s male. All later authors (including me) followed YARROW (1955) ’s diagnosis based on workers and gynes and kept silent regarding the male problem. J.W. Zetterstedt’s type male is, according to YARROW (1955), stored in ZMLU Lund. If still present there, its identity can only be determined after a thorough, broad-based study on male characters at least in the Fennoscandian population of F. lugubris, F. aquilonia and F. truncorum – at least it remains to be investigated if the differential characters proposed in the keys of KUTTER (1977) and CZECHOWSKI &amp; al. (2012) for the Central European population do apply to the boreal populations of these species.</p> </div>	http://treatment.plazi.org/id/F52B87F65E29615EFF5EDE3FFF7C1E59	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E29615EFC9FD94EFB261CD8.text	F52B87F65E29615EFC9FD94EFB261CD8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica nylanderi BONDROIT 1920	<div><p>Formica nylanderi BONDROIT, 1920</p> <p>Formica nylanderi BONDROIT, 1920 [type investigation]</p> <p>Investigated were one syntype gyne labelled “Grindelwald Suisse Chabanaud coll. de Gaulle ”, “ Type ”, “ Formica v. nylanderi Type Bondr.” and one syntype gyne labelled “Lautaret 15 VII ”, “ Type ”, “ Formica rufa v. Nylanderi Type Bondr.”; depository MSNB Bruxelles.</p> <p>Phenotypic variation in Formica lugubris gynes can be classified into several morphs the data of which are given in Table 7 (see also SEIFERT 2018). In both type gynes of F. nylanderi, data of OccHL, GuHL, PnHL, and MetHL are clearly above the upper extremes known from Formica paralugubris and morph A3 of Alpine F.lugubris but they may belong to Formica helvetica sp.n. or morph A1 of Alpine F.lugubris. Running the types as wild-cards against a sample of 28 gynes of F. helvetica and 36 gynes of morph A1, and considering the five characters ML / CS, PeW / CS, nCH, nMes, and nPe, the gyne from Grindelwald is allocated to morph A1 with p = 0.9928 and the gyne from Col de Lautaret with p = 0.9976. As the specimen from Grindelwald achieved a higher posterior probability in a wild-card run when all 20 characters were considered, it is designated herewith as lectotype.</p> </div>	http://treatment.plazi.org/id/F52B87F65E29615EFC9FD94EFB261CD8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E29615EFC9FDFAEFCFE1ED9.text	F52B87F65E29615EFC9FDFAEFCFE1ED9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica rufopratensis FOREL 1874	<div><p>Formica rufopratensis FOREL, 1874</p> <p>Formica rufa var. rufopratensis FOREL, 1874 [type investigation and zoogeography]</p> <p>FOREL (1874) did not mention a type locality, and his description of worker and gyne may refer to Formica lugubris, Formica paralugubris, and Formica helvetica sp.n. A smaller set of morphological data available for two supposed type gynes in MNH Genève from the locality Barberine (France, 46.050° N, 6.943° E) allow to exclude F. paralugubris but cannot separate the other two species. As F. helvetica sp.n. is considered to be restricted to East Switzerland and unlikely to occur farther west, I synonymize F. rufopratensis with F. lugubris. Alternatively, if the Barberine specimens are not recognized as types, F. rufopratensis has to be listed up under incertae sedis.</p> </div>	http://treatment.plazi.org/id/F52B87F65E29615EFC9FDFAEFCFE1ED9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E166161FF74DAF1FC951DB8.text	F52B87F65E166161FF74DAF1FC951DB8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica santschii WHEELER 1913	<div><p>Formica santschii WHEELER, 1913</p> <p>Formica rufa var. santschii WHEELER, 1913 [type investigation]</p> <p>Formica santschii is WHEELER (1913) ’s replacement name for F. rufa var. alpina SANTSCHI, 1911, which is a junior primary homonym of Formica adamsi var. alpina WHEELER, 1909. SANTSCHI (1911) described his F. rufa var. alpina from the Alps north of Sondrio. Investigated were two syntype workers on separate pins stored in NHM Basel and labelled in F. Santschi´s handwriting. The larger specimen with CW 1754 µm is depicted in AntWeb (ANTWEB 2021) under CASENT0912249 and is labelled “Valteline. Galli-Valerio”, “ Formica alpina € Sant.”, “type”. The smaller, damaged specimen with CW = 1685 µm on the other pin is labelled “Valteline. Galli-Valerio”, “ Formica alpina € Sant.” By morphology and geography, the F. santschii types could either belong to Formica lugubris or Formica paralugubris. The specimens were run as wild-cards in an LDA collecting 369 workers of the F. lugubris morphs A1 and A3 from the Alps in class 1, and 298 workers of F. paralugubris in class 2. The eleven characters available in the F. santschii types were CS, CL / CW 1750, SL / CS 1750, SL / Smax 1750, PeW / CS 1750, nSc 1750, nCH 1750, OccHL 1750, mPnHL 1750, nMet 1750, and MetHL 1750. With these data, the larger syntype worker was allocated to class 1 with p = 0.9674 but the smaller one with p = 0.9551 to class 2. It appears questionable if the two workers mounted on separate pins belong to the same nest sample. In order to save a stable nomenclature, a lectotype of F. santschii is fixed by present decision in the undamaged larger specimen labelled “CASENT0912249”.</p> </div>	http://treatment.plazi.org/id/F52B87F65E166161FF74DAF1FC951DB8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E166160FF74DECEFF751D59.text	F52B87F65E166160FF74DECEFF751D59.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica unicolor RUZSKY 1926	<div><p>Formica unicolor RUZSKY, 1926</p> <p>Formica pratensis ssp. unicolor RUZSKY, 1926 [description, diagnosis of DLUSSKY (1967)]</p> <p>This taxon was described on the basis of a single worker collected at Surgut (West Siberia, 62.25° N, 73.42° E) on 21 June 1913. A synonymy with Formica lugubris appears possible from description. DLUSSKY (1967), who might have seen the type specimen, interpreted this taxon as a light coloured F. lugubris without a dark patch on dorsal mesosoma. Such specimens are more frequent in Siberia than in Europe.</p> <p>All material examined. Numeric phenotypical data were recorded in 224 nest samples with 1108 workers and 109 gynes. These originated from Austria (seven samples), Bulgaria (seven), Czechia (four), Finland (42), France (28), England (two), Germany (18), Italy (15), Mongolia (two), Norway (10), Russian West Siberia (seven) Russian East Siberia (15), Sweden (24), and Switzerland (44). For details, see SI1, SI2, and SI3.</p> <p>Geographical range. Eurosiberian, boreo-montane. The huge boreal range extends from the British Isles east to Kamchatka. In Fennoscandia from S Sweden (55.5° N) to <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=132.79&amp;materialsCitation.latitude=43.17" title="Search Plazi for locations around (long 132.79/lat 43.17)">North Cape</a> (71° N). <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=132.79&amp;materialsCitation.latitude=43.17" title="Search Plazi for locations around (long 132.79/lat 43.17)">The</a> northern distribution in <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=132.79&amp;materialsCitation.latitude=43.17" title="Search Plazi for locations around (long 132.79/lat 43.17)">Siberia</a> is limited by 65° N. <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=132.79&amp;materialsCitation.latitude=43.17" title="Search Plazi for locations around (long 132.79/lat 43.17)">Occurrence</a> at low latitudes and altitudes in the Ussuri region (43.17° N, 132.79° E, 350m) is explained by influence of cold water of the West Pacific. The montane-subalpine population in Europe extends over the Pyrenees, Massif Central, Alps, Vosges, Schwarzwald, Bayrischer Wald, the whole Carpathians, Stara Planina, Vitosha, Rila, Pirin, and Rhodopes. In the Alps occurring at altitudes between 550 and 2510 m. Under influence of Atlantic climate descending to 250m (Vosges, France) and 50 m (Ireland).</p> <p>Diagnosis of worker (Tab. 3 and 4, Figs. 14 and 15, key). Extremely polymorphic, with regional differences shown in Table 3. Rather large, mean and maximum CS over all morphological and social types 1805 and 2184 µm. Scape shorter and much less slender than in Formica pratensis, SL / CS 1750 0.894, SL / Smax 1750 9.30. Petiole wider than in F. pratensis, PeW / CS 1750 0.489. Setae on eyes long, EyeHL 1750 33 µm; setae condition on dorsal plane of scape most variable, nSc 1750 usually 0 - 5 but in Fennoscandian Hippie-morph (SEIFERT 2003) up to 40. Setae on head margin behind eyes always numerous but most variable in length, nCH 1750 25.0; average OccHL 1750 usually 115µm but up to 235 µm in Fennoscandian Hippie-morph. Gular setae always numerous and long, nGu 1750 15.1, GuHL 1750 188 µm. Pronotal setae numerous but most variable in length, average mPnHL 1750 usually 84 - 112µ m but in Fennoscandian Hippie-morph up to 176µm. Number and length of mesopleural, propodeal, and metanotal setae usually large, nMes 1750 28.1, nPr 1750 21.7; mMet 1750 10.3, MetHL 1750 182 µm. Pigmentation without specific characters. The dark patch on mesosoma is usually large with diffuse margins in European popu- lations but may also be reduced.</p> <p>Diagnosis of gyne (Tab. 7, Fig. 3, key). Extremely polymorphic. Mean number and length of setae is lowest in morph A3 from the Alps (SEIFERT 2018) and highest in the Fennoscandian Hippie-morph (SEIFERT 2003). The remaining morphs, such as the most abundant morphs A1 from the Central European mountains or morph N1 from Fennoscandia, are intermediate. The numeric data given in the following summarize data of 109 gynes from the whole geographic range and of any social and morphological type. Rather large, CS 2213 ± 80 (1996, 2382) µm. Head short, CL / CW 0.998 ± 0.020 (0.950, 1.052). Scape short and thickset, SL / CS 0.789 ± 0.026 (0.708, 0.857), SL / Smax 8.13 ± 0.37 (7.12, 9.04). Eyes always with setae, the longest occur in the Fennoscandian Hippie-morph, EyeHL 48 ± 18 (30, 151) µm. Dorsal plane of scape with most variable setae numbers, which are highest in Fennoscandian Hippie-morph, nSc 2.1 ± 3.5 (0, 21). Posterior margin of head always with setae, but these are most variable in length and number, nCH 23.2 ± 8.8 (10.5, 65.0), OccHL 156 ± 91 (40, 320) µm. Underside of head, all surfaces of mesosoma, and petiole scale always with setae of most variable length and number, nGu 18.3 ± 6.7 (8, 36), GuHL 244 ± 111 (49, 425) µm, PnHL 208 ± 93 (34 - 411), MetHL 260 ± 97µm (40, 375), nPe 15.5 ± 6.5 (2,33). Dorsalsurfaceofgasterappearsatlowermagnifica- tion more or less shiny. Transverse microripples on dorsum of first gaster tergite usually very weak or absent but occa- sionally more developed, then approaching the situation in Formica pratensis. Foveolae and pubescence on dorsum of first gaster tergite on average more densely packed than in Formica rufa or Formica polyctena, FodG 25.2 ± 5.7 (17.9, 50.8) µm, sqPDG 4.84 ± 1.14 (3.13, 8.06) µm.</p> <p>Taxonomic comments and clustering results. The European population is extremely polymorphic, being a mixture of (a) sympatrically occurring most different phenotypes, as for example observed in Fennoscandia (SEIFERT 2003) or in the Alps (SEIFERT 2018), and (b) of deviating but rather monomorphous regional populations, such as found in the Pyrenees or the Balkan mountains. Clustering of morphological data by exploratory data analyses and assessment of putative clusters by discriminant analyses did not allow to give one of these entities a taxonomic significance (for the special case of Formica helvetica sp.n., see section “ Formica helvetica sp.n. ”, p. 166). Considering this very complicated structure, it appears astonishing that the separation from Formica pratensis all over the Palaearctic range was such clear in workers and gynes (see section “ Formica pratensis RETZIUS, 1783 ”, p. 159 or SEIFERT &amp; GOROPASHNAYA 2004), and that also Formica paralugubris was sufficiently separable in both castes (see section “ Formica paralugubris SEIFERT, 1996 ”, p. 167, and SEIFERT 2016a). Sections “Hybrids Formica aquilonia × lugubris ” (p.173), “Hybrids Formica pratensis × lugubris ” (p.174) and “Hybrids Formica lugubris × rufa ” (p.174) report on hybridization of F. aquilonia × lugubris, F. pratensis × lugubris, and Formica rufa × lugubris.</p> <p>Biology. See the species profile given by SEIFERT (2018).</p></div> 	http://treatment.plazi.org/id/F52B87F65E166160FF74DECEFF751D59	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E176163FF5EDE3FFC951F38.text	F52B87F65E176163FF5EDE3FFC951F38.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica helvetica Seifert 2021	<div><p>Formica helvetica sp.n.</p> <p>Formica lugubris -A2 (BERNASCONI &amp; al. 2011)</p> <p>Formica lugubris, morph A2 (SEIFERT 2018)</p> <p>Etymology. Referring to Helvetia, the Latin name of Switzerland.</p> <p>Type material. Holotype worker plus four paratype workers labelled “SWI: 46.7218° N, 10.2988° E, Scuol, Pinus forest, 1767m, polydomous, Bernasconi 2005.05.13 - MIN7 ”; another nine nest samples from the same supercolony with 47 paratype workers and 30 paratype gynes collected by C. Bernasconi in the years 2005 - 2008, sample numbers MIN8, MIN11, MIN13, MIN15, MIN17, MIN18, MIN20, MIN35, and MIN36; depository SMN Görlitz.</p> <p>All material examined. Numeric phenotypical data were recorded in nine nest samples with 30 workers and 28 gynes. All originated from the supercolony of the type locality. For details, see SI1, SI2, and SI3.</p> <p>Geographic range. Only known from the type locality in Mingèr Valley in the Eastern Swiss Alps at altitudes between 1700 and 2000 m.</p> <p>Diagnosis of worker (Tab. 3, Figs. 16 and 17). Small, as it is expected for a supercolonial social phenot ype, mean and maximum CS 1663 and 1966µm. Scape as short as but on average less thickset than in Alpine Formica lugubris, SL / CS 1750 0.904, SL / Smax 1750 9.47. Petiole width and setae number as in Alpine F. lugu- bris but setae length on average larger, EyeHL 1750 37µm, OccHL 1750 132µm, mPnHL 1750 111µm, MetHL 1750 188 µm.</p> <p>Diagnosis of gyne (Tab. 7). Size slightly smaller than in Alpine Formica lugubris, mean and maximum CS 2126 and 2278 µm. Head short, CL / CW 0.999. Scape on average longer and less thickset than in morph A3 of Alpine F. lugubris, SL / CS 0.797, SL / Smax 8.41. Eyes always with rather long setae, EyeHL 44µm. Dorsal plane of scape usually without or very few setae, nSc 0.4. Posterior margin of head always with setae, these on average longer than in morph A3 of Alpine F. lugubris, nCH 18.5, OccHL 179µm. Setae number comparable with morph A1 and A3 of Alpine F. lugubris but seta length larger than in morph A3, nGu 17.2, GuHL 303µm, PnHL 236µm, MetHL 287µm, nPe 13.7. Dorsal surface of gaster appears at lower magnification more or less shiny. Dorsum of first gaster tergite usually with weak transverse microripples and with foveolae and pubescence being on average more densely packed than in Formica rufa or Formica polyctena, FodG 26.1 µm, sqPDG 4.84 µm.</p> <p>Taxonomic comments and clustering results. Formica helvetica sp.n. is the first ant species I describe as new without having a sufficiently clear mor- phological diagnosis. This decision is certainly a risk, considering the patchwork situation in Alpine Formica lugubris populations. However, this taxonomic act is intended as a constructive, positively provocative hypothesis for future ant students to re-investigate the case with more advanced methods. The recognition as species here is largely based on an apparently clear clustering by nuclear DNA and moderate support by gyne morphology. BERNASCONI &amp; al. (2011) investigated nine microsatellite markers in seven Formica rufa group species in and around the Swiss National Park in East Switzerland. They showed a clear difference between F. helvetica sp.n. and F. lugubris (in Mingèr Valley represented by morph A1) and five other F.rufa group species sympatrically occurring in the area. The genetic distance between F. helvetica sp.n. and F. lugubris (Fst = 0.101) was comparable with those between Formica aquilonia and Formica paralugubris (Fst = 0.117), or between F. lugubris and F. aquilonia (Fst = 0.130) but lower than between monodomous and polydomous populations of F. lugubris in another area of the Swiss National Park (BERNASCONI &amp; al. 2005). The mtDNA of F. helvetica sp.n. clustered with that of F. paralugubris and F. aquilonia, which may suggest a hybridogenous evolutionary history, but also that there was no influx of genes by immigration of F. lugubris gynes.</p> <p>The morphological clustering of workers failed when the whole Alpine population from the French Jura to Eastern Austria was considered. Running an LDA with the characters CS, CL / CW 1750, SL / CS 1750, nSc 1750, nCH 1750, OccHL 1750, mPnHL 1750, and nMet 1750, all nine worker nest samples of Formica helvetica were allocated to the same cluster, but this cluster was shared with nine Formica lugubris samples of the morph A1 from nine different lo- calities. This meant a classification error of 9.3% in a total of 97 worker samples. The misplaced F. lugubris samples came from all over the Alps and included a population from Grande Rolat in Swiss Jura which had been intensively studied biologically, genetically and morphologically over many years.</p> <p>The situation in gynes appears better, but the low sample size of only 28 specimens in Formica helvetica sp.n. required a strong character reduction, performed by a stepwise LDA. Using the eight characters SL / CS, SL / Smax, PeW / CS, ML / CS, nSc, nMes, GuHL, and nHT, the nest sample means of the LDA scores provide a rather good separation of F. helvetica sp.n. (Fig. 31), but the figure also shows that the first component of the PCA (calculated from the same eight characters as the LDA) does not expose F. helvetica but separates instead the Formica lugubris morphs A1 and A3.</p> <p>Habitat and biology. The supercolony stretched along a transect of about 1.6k m length within a Pinus forest on limestone ground. The social structure is comparable with that of Formica aquilonia, with mating intranidally or within the colony borders and high genetic viscosity.</p> </div>	http://treatment.plazi.org/id/F52B87F65E176163FF5EDE3FFC951F38	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E146162FF5EDC5FFCEC1A39.text	F52B87F65E146162FF5EDC5FFCEC1A39.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica paralugubris SEIFERT 1996	<div><p>Formica paralugubris SEIFERT, 1996</p> <p>Formica paralugubris SEIFERT, 1996 [type investigation]</p> <p>This taxon was described from the Swiss Jura <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=6.192&amp;materialsCitation.latitude=46.537" title="Search Plazi for locations around (long 6.192/lat 46.537)">Mountains</a> (46.537° N, 6.192° E, 1450 m). <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=6.192&amp;materialsCitation.latitude=46.537" title="Search Plazi for locations around (long 6.192/lat 46.537)">The</a> holotype gyne is labelled “SWI: Jura: 1994.06, Le Brassus-5SSW, Chalet a Roch Field Stat., nest G5” and depicted in <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=6.192&amp;materialsCitation.latitude=46.537" title="Search Plazi for locations around (long 6.192/lat 46.537)">AntWeb</a> (ANTWEB 2021) under the unique specimen identifier FOCOL0762. Investigated was all type material, consisting of five gynes and 34 workers from the nests G1-G5 of the holotype supercolony, collected in the years 1993 and 1994. Depository SMN Görlitz.</p> <p>All material examined. Numeric phenotypical data were recorded in 73 nest samples with 355 workers and in 53 gynes. These originated from Austria (six samples), Canada (two), France (two), Germany (four), Italy (three), and Switzerland (58).</p> <p>Geographical range. Its natural range is rather small and extends over the montane-subalpine zone of the Jura Mountains and western Alps between 6° E and 11.5° E with a small exclave in the southern Schwarzwald / Germany. In the Alps, it ascends to 2300 m. A colony artificially introduced to Quebec / Canada in 1971 showed continuous growth to supercolonial size over 34 years (SEIFERT 2016a). Artificial introductions of wood ants to at least 42 localities over entire Italy south to Sicily and west to Sardinia were performed in the years 1959 - 1967 (e.g., PAVAN 1959). In that time, the transferred ants were classified as Formica lugubris. However, it is very likely that the vast majority of these introductions really involved Formica paralugubris as it was confirmed for five sites in the North Apennine (MASONI &amp; al. 2019).</p> <p>Diagnosis of worker (Tab. 4, key). Minimum size, mean and maximum CS 1680 and 2020µ m.Head rather short, CL / CW 1750 1.091. Scape rather short and thickset, SL / CS 1750 0.902, SL / Smax 1750 9.22. Eyes always with long microsetae, EyeHL 1750 34µm. Setae number on dorsal plane of scape variable but on average higher than in Alpine Formica lugubris, nSc 1750 5.2. Posterior margin and underside of head always with conspicuous setae, nCH 1750 24.9, OccHL 1750 108 µm, nGu 1750 14.2, GuHL 1750 164µm. Mean length of pronotal setae, number and length of metapleural setae on average lower than in morph A1 of Alpine F. lugubris, mPnHL 1750 78 µm, nMet 1750 7.7, MetHL 1750 154 µm. Workers of morph A3 of Alpine F. lugubris are similar in the pilosity condition but have a much larger size, a larger head length index, and a shorter scape.</p> <p>Diagnosis of gyne (Tab. 7). On average smaller than morph A1 and A3 of Alpine Formica lugubris, mean and maximum CS 2095 and 2238 µm. Scape longer than in morph A3 of Alpine F. lugubris and very thickset, SL / CS 0.805, SL / Smax 7.97. Eyes always with conspicuous microsetae, EyeHL 41 µm. Setae number on dorsal plane of scape variable but on average higher than in morph A1 and A3 of Alpine F. lugubris, nSc 6.4. Posterior margin and underside of head always with conspicuous setae, the length of which is lower than in morph A1 but larger than in morph A3 of Alpine F. lugubris, nCH 23.8, OccHL 117 µm, nGu 16.7, GuHL 128 µm. Pronotal setae shorter than in morph A1 of Alpine F. lugubris, mPnHL 88 µm. Petiole setae fewer and metapleural setae shorter than in morph A1 of Alpine F. lugubris but more numerous and longer than in morph A3 of Alpine F. lugubris, nPe 9.2, MetHL 110 µm. Dorsal surface of gaster appears at lower magnification more or less shiny. Dorsum of first gaster tergite usually with weak transverse microripples and with foveolae and pubescence on average more densely packed than in Formica rufa or Formica polyctena, FodG 21.1 µm, sqPDG 4.62 µm.</p> <p>Taxonomic comments and clustering results. Considering the extreme polymorphism in Alpine Formica lugubris and the presence of another similar sympatric species Formica helvetica sp.n., the separation of Formica paralugubris in both workers and gynes should be problematic. I combined 98 nest samples with 409 workers of Alpine F. lugubris morphs A1 and A3 and of F. helvetica sp.n. in class 1, and 70 nest samples with 323 workers of F. paralugubris in class 2. A two-class LDA considering the characters CS, CL / CW 1750, SL / CS 1750, SL / Smax 1750, PeW / CS 1750, nSc 1750, nCH 1750, OccHL 1750, mPnHL 1750, nMet 1750, and MetHL 1750 classified all samples of F. paralugubris and 96 samples of the collective cluster correctly. This means a classification error of 1.2% within a total of 168 nest samples. A plot of the first and second factors of a PCA supported the existence of two main clusters class 1 and class 2, and disagreed in 3.0% of the samples with the LDA (Fig.32). The exploratory data analyses NC-part.kmeans, NC-Ward, and NC-NMDS.kmeans suggested two clusters and disagreed with the final species hy- pothesis by 3.6, 4.1, and 3.0%. The clustering by NC-part. hclust was ignored as it splitted into seven clusters and exposed 8.9% indeterminate samples (outliers). As result, we have a sufficiently good separation of F. paralugubris workers by morphology. The distinction of F.paralugubris gynes from those of Alpine F.lugubris morphs A1 and A3, and of F. helvetica sp.n. by a principal component analysis appears also rather clear when the eight characters CS, SL / CS, SL / Smax, PeW / CS, ML / CS, nSc, nCH, and OccHL are considered (Fig.33). Section “Hybrids Formica aquilonia × paralugubris ” (p.175) discusses the situation in hybrids F. aquilonia × paralugubris.</p> <p>Biology. See the species profile given by SEIFERT (2018).</p></div> 	http://treatment.plazi.org/id/F52B87F65E146162FF5EDC5FFCEC1A39	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E156162FC9FDFAEFAAE1E78.text	F52B87F65E156162FC9FDFAEFAAE1E78.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica truncicola NYLANDER 1846	<div><p>Formica truncicola NYLANDER, 1846</p> <p>Formica truncicola NYLANDER, 1846 [type investigation]</p> <p>Investigated were four type specimens from ZM Helsinki: one worker syntype pinned together with an alate gyne, both with detached heads glued on a separate cardboard, labelled “Kuusamo \ W. Nyland. \ Coll. Nyland \ Mus. Zool. H:fors Spec. typ. No 5026 Formica truncicola Nyl. ”; one alate gyne on another pin with same labels but “...Spec. typ. No 5024”; one male on a third pin with same labels but “...Spec. typ. No 5025”. All specimens fully match the established conception Formica truncorum.</p> </div>	http://treatment.plazi.org/id/F52B87F65E156162FC9FDFAEFAAE1E78	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E156165FC9FDD0EFC951B19.text	F52B87F65E156165FC9FDD0EFC951B19.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica truncicolopratensis FOREL 1874	<div><p>Formica truncicolopratensis FOREL, 1874</p> <p>Formica rufa var. truncicolopratensis FOREL, 1874 [photo of type]</p> <p>Forel described the worker and gyne but did not mention a type locality. The taxon was synonymized by all previous authors with Formica truncorum. If a worker specimen in MHN Genève, labelled “ Typus ”, “F. truncicolo-pratensis Mt Ce..ere” and “ANTWEB CASENT 0911086” can be recognized as true type, the synonymy with F. truncorum appears likely after inspecting the z-stack photos shown in AntWeb (ANTWEB 2021) under the specimen identifier CASENT0911086.The photos suggest shiny circular lateral clypeal depressions and the length of the second funiculus segment to be more than twice its width as it is typical for F. truncorum. The more developed dark pigmentation is not contradictory as such specimens do occasionally occur over the whole range of the species.</p> </div>	http://treatment.plazi.org/id/F52B87F65E156165FC9FDD0EFC951B19	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E156162FCF9D95FFAEC1CD9.text	F52B87F65E156162FCF9D95FFAEC1CD9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica truncorum FABRICIUS 1804	<div><p>Formica truncorum FABRICIUS, 1804</p> <p>Formica truncorum FABRICIUS, 1804 [type investigation]</p> <p>The taxon was described from Moravia / Czechia from a nestinadeadtreetrunk.Investigatedweretwotypespecimens from ZM Copenhagen: one worker labelled “ Formica truncorum 403.31 Kiel ” (a permanent loan from ZM Kiel) and with a handwritten fragmentary label “ trunca fusca ”, and a strongly damaged alate gyne labelled “ Formica truncorum 403.31 Kiel ” but without a second label. Both specimens are fully consistent with the established conception of F. truncorum. A synonymy with Formica frontalis is excluded by zoogeography and the higher setae numbers on scape and metapleuron. Data of the worker are: CS 1957 µm, CL / CW 1.046, SL / CS 0.978, PeW / CS 0.483; nGu 29, OccHL 124 µm, GuHL 220 µm, mPnHL 81.6 µm, nPr 44.5, nMet 24.5, MetHL 170 µm (setae data of damaged body parts were estimated and are not given here). The gyne fragment, consisting of the dorsal mesosoma with wings, petiole, both hindlegs, and one foreleg, is a typical reddish F. truncorum with all surfaces covered by a profuse, thin, and very long pilosity. The wings are notably infuscated, matching the original description.</p> </div>	http://treatment.plazi.org/id/F52B87F65E156162FCF9D95FFAEC1CD9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E126165FF74DE0EFE811CD8.text	F52B87F65E126165FF74DE0EFE811CD8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica approximans WHEELER 1933	<div><p>Formica approximans WHEELER, 1933</p> <p>Formica truncorum var. approximans WHEELER, 1933 [type investigation]</p> <p>The taxon was described from Eastern Tomb / China. Investigated were six worker syntypes from MCZ Harvard, labelled “Eastern Tomb July 16, 30 Chi Ho”, “Gift of W.M.Wheeler ”, “ M.C.Z. CoType 1.-6 21720”, and “ Syntypes Formica truncorum var. approximans Wheeler ” (upper side) “S P Cover IX-2006 ” (underside). The specimens fully correspond to the East Palaearctic population of F. truncorum. For a more detailed argumentation in favor of this synonymization, see below under taxonomic comments.</p> </div>	http://treatment.plazi.org/id/F52B87F65E126165FF74DE0EFE811CD8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E126165FF74DFAEFE1D1E38.text	F52B87F65E126165FF74DFAEFE1D1E38.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica finzii STITZ 1939	<div><p>Formica finzii STITZ, 1939</p> <p>Formica truncorum var. finzii STITZ, 1939 [description and zoogeography]</p> <p>This taxon was described in a gyne from near Eberswalde (Germany) under the unavailable name Formica rufa ssp. truncicola var. finzii KRAUSSE, 1926. Types are unknown. Within the four species known to occur in the vicinity of Eberswalde, the description matches the situation in F. truncorum.</p> </div>	http://treatment.plazi.org/id/F52B87F65E126165FF74DFAEFE1D1E38	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E126165FF74DD4EFC2319D8.text	F52B87F65E126165FF74DD4EFC2319D8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica menozzii STITZ 1939	<div><p>Formica menozzii STITZ, 1939</p> <p>Formica truncorum var. menozzii STITZ, 1939 [description and zoogeography]</p> <p>This taxon was described from near Eberswalde (Germany) under the unavailable name Formica rufa ssp. truncicola var. menozzii KRAUSSE, 1926. Types are unknown. Within the four species known to occur in the vicinity of Eberswalde, the description matches the situation in F. truncorum.</p> </div>	http://treatment.plazi.org/id/F52B87F65E126165FF74DD4EFC2319D8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E126165FC9FDAAEFC231B38.text	F52B87F65E126165FC9FDAAEFC231B38.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica staegeri STITZ 1939	<div><p>Formica staegeri STITZ, 1939</p> <p>Formica truncorum var. staegeri STITZ, 1939 [description and zoogeography]</p> <p>This taxon was described from near Eberswalde (Germany) under the unavailable name Formica rufa ssp. truncicola var. staegeri KRAUSSE, 1926. Types are unknown. Within the four species known to occur in the vicinity of Eberswalde, the description matches the situation in F. truncorum.</p> </div>	http://treatment.plazi.org/id/F52B87F65E126165FC9FDAAEFC231B38	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E126165FF74D86EFE811D78.text	F52B87F65E126165FF74D86EFE811D78.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica yessensis WHEELER 1913	<div><p>Formica yessensis WHEELER, 1913</p> <p>Formica truncorum var. yessensis WHEELER, 1913 [type investigation]</p> <p>The taxon was described by FOREL (1901) under the infrasubspecific name Formica rufa r. truncorum var. yessensis FOREL, 1901 from Sorachi, some 100 km ENE of Sapporo, Hokaido. Investigated was one type worker from MHN Genève labelled by A. Forel “ Formica truncicola v. yessensis For Type ” and carrying a printed label “Sorachi Prov. Ishikari Yesso V. 1899. Mus.em. 24.XI.1899 ”. The specimen shows below average setae length on eyes and no setae on dorsum of scape, and extensor profile of hind tibia but is in any character within the range of variation known from the Palaearctic population of F. truncorum. For a more detailed argumentation in favor of this synonymization, see below under taxonomic comments.</p> </div>	http://treatment.plazi.org/id/F52B87F65E126165FF74D86EFE811D78	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E126164FC9FD84EFBC41AD8.text	F52B87F65E126164FC9FD84EFBC41AD8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica stitzi STITZ 1939	<div><p>Formica stitzi STITZ, 1939</p> <p>Formica truncorum var. stitzi STITZ, 1939 [description and zoogeography]</p> <p>This taxon was described from near Eberswalde (Germany) under the unavailable name Formica rufa ssp. truncicola var. stitzi KRAUSSE, 1926. Types are unknown. Within the four species known to occur in the vicinity of Eberswalde, the description matches the situation in F. truncorum.</p> <p>All material examined. Numeric phenotypical data were recorded in 50 samples with 92 workers and in 22 gynes. These originated from Belarus (one sample), China (seven), Czechia (one), Denmark (one), Finland (two), France (three), Germany (12), Japan (two), Kazakhstan (seven), Kyrgyzstan (11), Norway (one), Russia (two), and Ukraine (one). For details, see SI1, SI2, and SI3. The total of samples investigated either numerically or subjectively was 136.</p> <p>Geographical range. <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=148.0&amp;materialsCitation.latitude=42.0" title="Search Plazi for locations around (long 148.0/lat 42.0)">Eurosiberian</a>, temperate to boreal; from Netherlands and E France (5° E) to Yakutsk at least (62° N, 130° E). In NE China, the <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=148.0&amp;materialsCitation.latitude=42.0" title="Search Plazi for locations around (long 148.0/lat 42.0)">Ussuri region</a>, Sakhalin, Hokkaido, the northern half of Honshu, and the Western Kurils (148° E), it is replaced by the weakly deviating East Asian population. The southern and northern distributional limits in Europe are 42° N (Bulgaria) and 71° N (North Cape). These are in Central Siberia, along the 100 th degree of longitude, 46° N and 67° N. In Europe occurring from the planar to montane zone, in the Alps ascending to 1800m. Rare in the Crimea and Caucasus (DLUSSKY 1967). A population isolated from the Eurosiberian one is found in the Central Asian mountains – in Dzungarian Alatau, Tian Shan (here at 43° N up to 2700 m), and Pamir south to Karakoram (35° N).</p> <p>Diagnosis of worker (Tab. 5, Fig. 18, key). Polymorphic, with regional differences shown in Table 5. Medium-sized, mean and maximum CS over all morphological and social phenotypes 1754 and 2177µm. Head moderately elongated, CL / CW 1750 1.099. Middle part of lateral clypeus more deeply depressed than in the Formica rufa species complex, anterolateral clypeus as a result forming a bead; median clypeal carina blunt or absent. Scape long and very slender, SL / CS 1750 0.985, SL / Smax 1750 10.87. Second and third segment of antennal funiculus more slender than in the species treated above, IF2 2.15 ± 0.09 (n = 70). Petiole scale narrow, PeW / CS 1750 0.442. Eyes always with microsetae, EyeHL 1750 40 µm. Dorsal plane of scape usually with more setae than in other species, nSc 1750 10.3. Except for the Hippie-morph of Formica lugubris, setae number on each place of the body larger than in any species of the F. rufa group, nCH 1750 42.5, nGu 1750 40.5, nPn 1750 81.6, nMes 1750 35.8, nPr 1750 46.5, nMet 1750 17.6. Hind margin of head usually with very long setae, but setae length on other body parts lower than in the most hairy morphs of Formica pratensis and F. lugubris, OccHL 1750 136µm, GuHL 1750 187µm, mPnHL 1750 91 µm, MetHL 1750 142 µm. Dorsum of head, in addition to the other elements of microsculpture, with deeper and broader microfoveolae which are usually the base of setae. Typical pigmentation in medium-sized to large workers: whole head, mesosoma, petiole, and frontal part of first gaster segment light orange brown; dark brown or blackish brown patches on vertex and dorsal mesosoma may occur in some samples.</p> <p>Diagnosis of gyne (Tab. 8, Fig. 19, key). On average rather small but extremely size-polymorphic; minimum, mean, and maximum CS over all social phenotypes 1737, 2002, and 2256 µm. Head capsule in dorsal view appears trapezoidal, with more or less linear, frontad converging sides. Middle part of lateral clypeus more deeply depressed than in the Formica rufa species complex, anterolateral clypeus as a result forming a bead; median clypeal carina absent. Scape long and slender, SL / CS 0.903, SL / Smax 9.63. Petiole scale relatively narrow, PeW / CS 0.597. Setae on whole body very numerous, very thin, and usually very long. EyeHL 95 µm, nSc 28.0, nCH 62.6, OccHL 215µm, nGu 65.1, GuHL 267µm, PnHL 242 µm, nMet 38.0, MetHL 249 µm, nPe 15.0. Dorsum of first gaster tergite moderately shiny, with dense transverse microripples, rather dilute pubescence (sqPDG 9.76 µm), and rather large, deep, and widely spaced microfoveolae. The latter may be occasionally absent. Light reddish pigmentation component on all body surfaces more developed than in the F. rufa species complex.</p> <p>Taxonomic comments and clustering results. Formica truncorum, as any species of the F. truncorum species complex, is rarely confused with other species of the Formica rufa group. Occasional confusion with Formica pratensis is possible due to high similarity in pilosity data and most of the body shape data and due to variation in pigmentation in both species. However, the separation by exploratory and hypothesis-driven data analyses shows an error &lt;1% on the nest sample level in material from the whole Palaearctic range. Furthermore, the shape of clypeus and funiculus segments is usually diagnostic. In gynes, the separation from any species of the F. rufa species complex is also clear in both exploratory and hypothesis-driven data analyses. Less hirsute specimens from Kyrgyzstan, formerly identified as „ Formica cf. frontalis “ (SCHULTZ &amp; al. 2006), were re-classified in this study as F. truncorum. The current data give no indication that F. frontalis does occur outside of Iberia.</p> <p>The Panpalaearctic population of Formica truncorum cannot reasonably be subdivided into clusters of separate taxonomic identity based on the morphological data available at present. On worker individual level, the West and Central Palaearctic population differs from the East Palearctic one by smaller SL / CS 1750, nPn 1750, and nMet 1750 (Tab.5; ANOVA F 1,90&gt; 15, p &lt;0.001). Yet, it was not possible by any tested exploratory data analysis to cluster the East Palaearctic population separately – neither on individual nor on nest sample level. As a consequence, Formica yessensis WHEELER, 1913 and Formica approximans WHEELER, 1933 are synonymized here with F. truncorum. A subdivision within the East Palearctic population is also not possible. Using nuDNA (microsatellites), IMAI &amp; al. (2016) investigated the Japanese-Korean population. They could not show genetic differences between populations they had pre-determined as “ F. yessensis ” and “ F. truncorum ” based on subjective assessment of setae numbers on hind tibia. IMAI &amp; al. (2016) reported a “robust” genetic indication for the presence of only a single Japanese-Korean population and a “fragile” morphological classification. For separation from Formica frontalis and Formica sinensis, see section “ Formica frontalis SANTSCHI, 1919 ” (p. 171) and “ Formica sinensis WHEELER, 1913 ” (p. 172).</p> <p>Biology. See the species profile given by SEIFERT (2018). Gyne size polymorphism with large disperser gynes having more fat and glycogen and larger flight muscles than smaller inbreeding gynes is more strongly expressed in Formica truncorum than in any other species of the Formica rufa group.</p> </div>	http://treatment.plazi.org/id/F52B87F65E126164FC9FD84EFBC41AD8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E136167FCF9D9BFFF5C1D18.text	F52B87F65E136167FCF9D9BFFF5C1D18.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica dusmeti EMERY 1909	<div><p>Formica dusmeti EMERY, 1909</p> <p>Formica rufa ssp. dusmeti EMERY, 1909 [conception of TINAUT &amp; MARTINEZ-IBANEZ (1998)]</p> <p>The original description reports as type locality and collector: “Peñalosa in Spanien; 3 Exemplare wurden mir von Herrn Dusmet mitgeteilt.” According to TINAUT &amp; MARTINEZ-IBANEZ (1998), there is obviously a misspelling of the site as the two worker specimens in the Dusmet collection in MNCN Madrid bear labels indicating “Peñalara” as the collecting site. Furthermore, TINAUT &amp; MARTIN- EZ-IBANEZ (1998) quoted F. dusmeti to be frequent at Peñalara (Sierra de Peñalara) but to be absent from Peñalosa and surrounding areas in the province of Cordoba.</p> <p>All material examined. Numeric phenotypical data were recorded in three samples with seven workers from Spain. For details, see SI1, SI2, and SI3.</p> <p>Geographical range. According to TINAUT &amp; MARTINEZ-IBANEZ (1998) and ESPADALER &amp; GOMEZ (2000) restricted to Iberia with main occurrence in the northern part of the Peninsula. The altitude of seven sites was 1573 ± 261 m, which is higher than in Formica frontalis.</p> <p>Diagnosis of worker (Tab. 5, key). Medium-sized, mean and maximum CS 1706 and 2074 µm. Head elongated, CL / CW 1750 1.113. Clypeal structure as in Formica truncorum. Scape very long and slender, SL / CS 1750 1.001, SL / Smax 1750 11.08. Petiole scale narrow, PeW / CS 1750 0.438. Eyes without or with only short microsetae, EyeHL 1750 14 µm. Dorsal plane of scape completely without setae, nSc 1750 0.0. Setae number on each place of the body strongly reduced, clearly smaller than in Formica frontalis, nCH 1750 0.0, nGu 1750 0.0, nPn 1750 0.3, nMes 1750 2.4, nPr 1750 0.8, nMet 1750 0.4; when present, setae are short or of medium length, GuHL 1750 156 µm, mPnHL 1750 18µm, MetHL 1750 28µm. Typical pigmentation in medium-sized to large workers: whole head, mesosoma, petiole, and frontal part of first gaster segment light or- ange brown.</p> <p>Diagnosis of gyne. Unknown to me.</p> <p>Taxonomic comments and clustering results. I maintain here the concept of TINAUT &amp; MAR- TINEZ-IBANEZ (1998), who separated Formica dusmeti and Formica frontalis workers based on strong differences in setae numbers on pronotum, hind margin, and underside of head. This is basically confirmed by the data given in Table 5. Using these data, the first principal component provided a clear separation of seven F. dusmeti and 25 F. frontalis workers. However, the sample size is low, and considering the extreme intraspecific variation of pilosity data known in several species of the Formica rufa group, a thorough morphometric and genetic study is desired. Xavier Espadaler (pers. comm. July 2020) commented that some Portuguese and West Spanish populations of F. dusmeti and F. frontalis appeared to be intermediate in pilosity data.</p> <p>Biology. TINAUT &amp; MARTINEZ-IBANEZ (1998) report as main habitat conifer woodland, but it has also been found in open situations under large rocks and stone plates. Alates were observed between 14 June and 4 August.</p></div> 	http://treatment.plazi.org/id/F52B87F65E136167FCF9D9BFFF5C1D18	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E106166FF5EDE7FFE531B99.text	F52B87F65E106166FF5EDE7FFE531B99.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica frontalis SANTSCHI 1919	<div><p>Formica frontalis SANTSCHI, 1919</p> <p>Formica truncorum var. frontalis SANTSCHI, 1919 [type investigation]</p> <p>Investigated was the lectotype worker (by present designation), labelled in Santschi’s handwriting “ Formica truncorum F. v. frontalis type Sants”, “ Espagne Pozuelo de Calatrava (de la Fuente)”; depository NHM Basel. The original description reports as type locality nothing but “ Pozuelo de Calatrava (De la Fuente)”. Accordingly, a worker specimen depicted in AntWeb (ANTWEB 2021) under CASENT0912253, labelled “SIERRA DE QUADAR- RAMA Dusmet” and “type” has no type status.</p> <p>All material examined. Numeric phenotypical data were recorded in 11 samples with 26 workers and six gynes from Spain. For details, see SI1, SI2, and SI3.</p> <p>Geographical range. According to TINAUT &amp; MARTINEZ-IBANEZ (1998) and ESPADALER &amp; GOMEZ (2000) rather homogenously distributed over entire Iberia with the altitude of 15 sites being 1160 ± 450 m, which is lower than in Formica dusmeti.</p> <p>Diagnosis of worker (Tab. 5, key). Medium-sized, mean and maximum CS over all morphological and social phenotypes 1792 and 2045 µm. Head moderately elongated, CL / CW 1750 1.102. Clypeal morphology as in Formica truncorum. Scape long and slender, SL / CS 1750 0.993, SL / Smax 1750 10.60. Petiole scale narrow, PeW / CS 1750 0.450. Eyes without or with short microsetae, EyeHL 1750 16 µm. Dorsal plane of scape without or with only single setae, nSc 1750 0.7. Setae number on each place of the body smaller than in F. truncorum but setae length equal or even larger, nCH 1750 14.5, nGu 1750 19.5, nPn 1750 59.5, nMes 1750 26.1, nPr 1750 29.2, nMet 1750 10.2, OccHL 1750 114 µm, GuHL 1750 204 µm, mPnHL 1750 97µm, MetHL 1750 161 µm. Dorsal surface of head usually without the deep and broad microfoveolae characteristic for F. truncorum. Pigmentation as in the latter.</p> <p>Diagnosis of gyne (Tab. 8). Only specimens from a single supercolony were available. These were much larger than Formica truncorum gynes of the polygynous to supercolonial social form, CS 2106 µm. Head capsule in dorsal view less trapezoidal than in F. truncorum, clypeal shape as in that species. Scape rather long and slender, SL / CS 0.872, SL / Smax 9.30. Petiole wider than in F. truncorum, PeW / CS 0.686. Setae on eyes short, EyeHL 31 µm. Setae on dorsum of scape usually absent, nSc 0.2. Setae on remaining parts of body present, less numerous than in F. truncorum but of similar length, nCH 23.4, OccHL 202µm, nGu 34.8, GuHL 287µm, PnHL 255 µm, nMet 26.3, MetHL 252µm, nPe 10.8. Dorsum of first gaster tergite moderately shiny, with dense transverse microripples and dilute pubescence (sqPDG 12.25µm), microfoveolae often absent. Pigmentation as in F. truncorum.</p> <p>Taxonomic comments and clustering results. The geographic ranges of Formica frontalis and Formica truncorum are disjunct, and the morphological distinctness of the two species is strong enough to consider them as heterospecific. The eleven Iberian nest samples of F. frontalis can be separated from 37 Panpalaearctic nest samples of F. truncorum using the characters SL / CS 1750, PeW / CS 1750, EyeHL 1750, nCH 1750, nMet 1750, and MetHL 1750 (Fig. 34). The classification error by an LDA was 4.2% in 118 worker specimens. It is worth mentioning that gynes from highly polygynous or supercolonial F. truncorum populations have a much smaller size than the investigated gynes from the F. frontalis supercolony at Puerto de Navacerrado (SaNo. 149 and 150), which provides additional support for heterospecificity.</p> <p>Biology. Compared with Formica dusmeti, TINAUT &amp; MARTINEZ-IBANEZ (1998) reported a more frequent occurrence in Quercus than in conifer forests, which is probably not due to a preference of broad-leafed woodland but rather a consequence of the lower altitudinal range. The species is also found in open treeless habitats with low shrubs. Nest construction is similar to Formica truncorum, not showing the large regular mounds seen in the Formica rufa species complex. The above-ground part is usually a rather flat accumulation of plant material, or the nests are completely under stones around which some plant material is deposited.</p> </div>	http://treatment.plazi.org/id/F52B87F65E106166FF5EDE7FFE531B99	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E116166FF5ED8FFFEFD1CF9.text	F52B87F65E116166FF5ED8FFFEFD1CF9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica sinensis WHEELER 1913	<div><p>Formica sinensis WHEELER, 1913</p> <p>Formica truncicola var. sinensis WHEELER, 1913 [type investigation]</p> <p>This taxon was described from the vicinity of Chongging (W China, 29.53° N, 106.52° E). Investigated were 32 syntype workers on 10 pins which were probably from the same sample as concluded from the consistent morphology. A lectotype plus two paralectotype workers on the same pin are labelled “ Lectotype (top specimen) Formica truncicola var. sinensis Wheeler 1913 desig. B. Seifert 2006 ”, “Chun King, China, 1909”, “Am. Mus. Nat. Hist. Dept. Invert. Zool. No. 22594” and “ Formica truncicola var. sinensis Whlr. ”; depository AMNH New York. 26 paralectotype workers on eight pins in AMNH New York and three paralectotype workers in SMN Görlitz are all labelled “Chun King, China, 1909” and “Am. Mus. Nat. Hist. Dept. Invert. Zool. No. 22594”. In the key of the original description (p. 391) and on the syntype labels, WHEELER (1913) used F. sinensis as trinomen but later in the heading of the main text (p. 437) as quadrinomen. Herewith, by majority indication, I suppose that WHEELER (1913) did not intend to introduce this taxon in infrasubspecific rank. Otherwise, WU (1990) would be the first who made the name available.</p> </div>	http://treatment.plazi.org/id/F52B87F65E116166FF5ED8FFFEFD1CF9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E116166FF74DF8EFF781ED9.text	F52B87F65E116166FF74DF8EFF781ED9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica wongi Wu 1990	<div><p>Formica wongi WU, 1990</p> <p>Formica wongi WU, 1990 [type investigation]</p> <p><a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=126.34&amp;materialsCitation.latitude=43.7" title="Search Plazi for locations around (long 126.34/lat 43.7)">This</a> taxon was described from near Yongji, province Jilin (NE China, 43.70° N, 126.34° E) in material collected 6 October 1983. Investigated were five paratype workers from the holotype sample from RIFCAF Beijing (now in SMN Görlitz) labelled in handwritten Chinese. The label was confirmed by different native Chinese speakers to name the type locality and it contained the sequence “ 1983.X.6 ” and “ Formica wongi Wu ”. The type series consists of unusually small workers of apparently a colony in foundation and its higher SL / CS of 1.018, and reduction of reddish pigment is explained by the normal allometric trends we observe in all Formica rufa group ants. For investigation results of the type series, see below under taxonomic comments and clustering results.</p> </div>	http://treatment.plazi.org/id/F52B87F65E116166FF74DF8EFF781ED9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
F52B87F65E116169FC9FDAF1FAA41DB8.text	F52B87F65E116169FC9FDAF1FAA41DB8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Formica delinghaensi CHANG & HE 2002	<div><p>Formica delinghaensis CHANG &amp; HE, 2002</p> <p>Formica delinghaensis CHANG &amp; HE, 2002 [zoogeography and description]</p> <p>This taxon was described from Delingha, Qinghai Province / China (37.32° N, 97.22° E, 2750 m). CHANG &amp; HE (2002) separated F. delinghaensis from Formica wongi by a brighter colour and a shorter scape (SL / CL 0.89 - 1.00). This scape length ratio is just the typical situation in Formica sinensis (SL / CL in 70 specimens 0.944 ± 0.027). Furthermore, only a single Formica rufa group species was discovered in E Tibet, Sichuan, and Gansu during seven collecting trips performed by A. Gebauer, D. Wrase, M. Schülke, I. Kabak, B. Seifert, R. Schultz, V. Assing, and A. Pütz in the years 1990 - 2012, and this species was F. sinensis.</p> <p>All material examined. Numeric phenotypical data were recorded in 29 samples with 70 workers and seven gynes from China. For details, see SI1, SI2, and SI3.</p> <p>Geographical range. Formica sinensis is found in the Chinese provinces Qinghai, Gansu, Sichuan, Chong- ging, and, as a seemingly disjunct population, in the province Jilin. Eleven samples from Quinghai and Gansu, from latitudes of 33 - 38° N, were found at elevations of 2480 ± 276 [2080, 2862] m and 11 samples from Sichuan, from latitudes between 28 - 32° N, at elevations of 3334 ± 542 [2700, 4130] m. This indication of high-altitude distribution may be partially misleading as the entomologists were not much motivated to sample in anthropogenically affected landscapes at lower elevations. A much lower alti- tudinal limit is indicated by the findings from <a href="http://tb.plazi.org/GgServer/search?materialsCitation.longitude=126.3&amp;materialsCitation.latitude=43.7" title="Search Plazi for locations around (long 126.3/lat 43.7)">Chongging</a> (29.53° N, 106.52° E, 1500m), Chincheng Shan (30.90° N, 103.55° E, 975 m), and Yongij (43.7° N, 126.3° E, 560m).</p> <p>Diagnosis of worker (Tab. 5, Fig. 20, key). Medium-sized, mean and maximum CS over all morphological and social phenotypes 1783 and 2134 µm. Head elongated CL / CW 1750 1.113. Clypeal morphology as in Formica truncorum. Scape very long and slender, SL / CS 1750 0.997, SL / Smax 1750 11.18. Petiole scale very narrow, PeW / CS 1750 0.430. Eyes with no or only short microsetae, EyeHL 1750 5µm. Dorsal plane of scape always without setae, nSc 1750 0.0. Setae number on each place of the body smaller than in F. truncorum but setae length on underside of head and metapleuron comparable: nCH 1750 8.2, nGu 1750 26.1, nPn 1750 26.6, nMes 1750 24.2, nPr 1750 20.5, nMet 1750 7.7, OccHL 1750 114 µm, GuHL 1750 184µm, mPnHL 1750 70 µm, MetHL 1750 140 µm. Dorsal surface of head without the deep and broad microfoveolae characteristic for F. truncorum. Pigmentation as in the latter.</p> <p>Diagnosis of gyne (Tab. 8). Rather large, mean CS 2192 µm. Head capsule in dorsal view less trapezoidal than in Formica truncorum, clypeal shape as in that species. Scape length and slenderness largest in all species considered here, SL / CS 0.921, SL / Smax 10.30. Petiole width as in F. truncorum, PeW / CS 0.589. Setae on eyes fully absent, EyeHL 0 µm. Setae on dorsum of scape always absent, nSc 0.0. Setae on hind margin of head much less numerous and shorter than in F. truncorum, nCH 11.4, OccHL 163µm. Gular setae less numerous than in F. truncorum but of similar length, nGu 33.0, GuHL 250µm. Setae on pronotum, metapleuron and petiole scale less numerous and shorter than in F. truncorum, PnHL 194 µm, nMet 22.3, MetHL 215 µm, nPe 11.2. Cuticular surface of head very homogenous, without pits or foveolae. Dorsum of first gaster tergite weakly shiny, with dense transverse microripples and dilute pubescence (sqPDG 10.14µm) but very homogenous microsculpture, microfoveolae often absent. Pigmentation as in F. truncorum.</p> <p>Taxonomic comments and clustering results. WU (1990) separated his new taxon Formica wongi from Formica sinensis because of absence of setae from pronotum and from the first three gaster tergites and the “rather dull body”. As there are always setae on the first three gaster tergites even in the least hairy species of the Palaearctic Formica rufa group, it appeared unclear if WU’ s material really belonged to this species group. The investigation of five syntypes had the following results:</p> <p>Very small body size for a species of the Formica truncorum species complex (CS = 1214, 1251, 1326, 1490, 1563µm) – this suggests a colony shortly after foundation. Ablation of pilosity on many areas of surface is confirmed by presence of numerous basal pits of setae. Anterior face of first tergite with numerous standing setae; most setae on exposed dorsal surfaces of first three tergites torn off, the few remaining are pasted flat to surface; in shel- tered surface dints numerous and long setae are present. Large parts of cuticular surface are polluted (“rather dull body”) and setae ablations are possibly due to attempts of mechanical cleaning prior preparation. Standard setae numbers and lengths were estimated in the three largest workers by scrutinizing the cuticular surface at magnifi- cations of 360 × for basal pits of setae and measuring the length of both standing setae and those glued flat to the surface. The sample means of the three largest workers are: CS 1460 µm, CL / CW 1750 1.119, SL / CS 1750 1.067, SL / Smax 1750 11.67, PeW / CS 1750 0.397, EyeHL 1750 7.3µm, nSc 1750 0.0, nCH 1750 5.1, OccHL 1750 105.6 µm, nGu 1750 31.1, GuHL 1750 130.0µm, nPn 1750 5.4, mPnHL 1750 57.1 µm, nMes 1750 22.5, nPr 1750 24.8, nMet 1750 11.4, and MetHL 1750 114.0 µm. All these data indicate a typical series of F. sinensis.</p> <p>The separation of Formica sinensis and Formica truncorum was very clear in any exploratory and hypothesis-driven data analysis considering the 17 characters mentioned above (Fig.35). The classification error in an LDA was 0% in 70 and 92 individual workers of F. sinensis and F. truncorum, respectively. Wild-card runs allocated all type specimens of F. truncorum, Formica truncicola, Formica approximans, and Formica yessensis to the F. truncorum cluster and all type specimens of F. sinensis and Formica wongi to the F. sinensis cluster. The classification errors in NC-Ward and NC-part.hclust were 0% and 1.6% in NC-part.kmeans.</p> <p>Biology. The data collected for this revision show the following aspects: Formica sinensis is the only Formica rufa group species present in large areas of China and thus without direct competitors in a rather broad niche space. It occurs in coniferous, mixed, and broad-leafed woodland of natural or anthropogenous origin – at higher elevations preferentially in woodland with low canopy closure or on clearings. Social types vary from monodomous colonies to true supercolonies with large mounds. Six observations of alates occurred between 26 June and 5 August.</p> </div>	http://treatment.plazi.org/id/F52B87F65E116169FC9FDAF1FAA41DB8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Seifert, Bernhard	Seifert, Bernhard (2021): A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) - the famous mound-building red wood ants. Myrmecological News 31: 133-179, DOI: 10.25849/myrmecol.news_031:133, URL: http://zoobank.org/0e55c0d7-531a-48d7-a078-148b96bd461d
