Tetramorium caespitum

This is the most common European Tetramorium species. Long believed to be introduced to North America, Wagner et al. (2017) conclusively determined the North American species is Tetramorium immigrans.

Wagner et al. (2017) - Moderately thermophilic, TAS of 465 sites 16.1 ± 2.0 °C [7.9, 21.1], different from all species except Tetramorium indocile. Most common species in most of Europe and more euryoecious than other species of complex. Most records from non-forested habitats like meadows, pastures, heaths, arid or semi-arid grasslands, vineyards, fallow grounds, ruderal areas, road embankments, rock heaps, gravel pits, river banks, but also light pine and oak forests. Urban areas like parks, pavements, and roadsides. Nest construction more flexible than in other species: in soil, under stones, rarely in dead wood; only species building soil mounds higher than 10 cm.

Pashaei Rad et al. (2018) found this species in Iran in Caspian moist littoral and moderate rainfall montane areas.

A recent revision of the Tetramorium caespitum complex (Wagner et al. 2017) showed that the group contains several cryptic species, thus all old Greek records of Tetramorium caespitum need confirmation. Borowiec & SaLata (2019c) confirmed its occurrence in Epirus, Macedonia, Peloponnese and Thessaly, and in this paper its presence is confirmed from Sterea Ellas. In Peloponnese and Aetolia-Acarnania, its nests were observed in mountain pastures and fir forest (Borowiec & Salata, 2021).

Identification
A member of the Tetramorium caespitum complex. The species can be determined with a discriminant analysis of a set of morphological measurements. See Wagner et al. (2017) and https://webapp.uibk.ac.at/ecology/tetramorium/

Blackish brown, sometimes paler; head including clypeus and mesosoma regularly longitidinally striate. Petiole and postpetiole with shallow punctures and sculpture but smooth in centre. Propodeal spines very short, broadly denticulate, petiole and postpetiole about as broad as long. Length: 2.5-4 mm (Collingwood 1979).

Siberian Species
Tetramorium sibiricum differs from Tetramorium indocile in having a longer scape, a more developed sculpture on postocular head sides, a narrower petiole and postpetiole, longer pronotal setae and a longer mesosoma and eye. The main differences to Tetramorium caespitum are the narrower petiole and postpetiole, the more developed sculpture and microsculpture on postocular head sides and on dorsum of 1st gaster tergite, the higher postpetiole, the longer scape and eye and a longer distance from spine tip to dorsocaudal corner of metapleural lobe.

Three species of the T. caespitum complex occur in Siberia west and north of the Reinig Line. This faunal divide separates East Siberian, Inner Mongolian, Chinese and Tibetan species from those of Central Siberia, West Siberia and the Turanian region (DE LATTIN, 1967).

The methodology of species identification applied by Seifert (2021) is extremely complex and data recording is by far the most time-consuming of any ant genus investigated so far by the author – a challenge even for experienced and specialized investigators having access to adequate equipment. An attempt to simplify the separation of the three West and Central Siberian species was done in writing a key that uses absolute primary data (i. e. without RAV-correction) and a strongly reduced character set.

1. Postoculo-temporal area of head with rather many costae and costulae, POTCos 11.4 ± 2.1. Anterior dorsum of 1st gaster tergite with more complex elements of stickman-like or reticulate microsculpture, MC1TG 16.6 ± 3.2. Propodeal spines and eye slightly longer. With all linear measurements given in mm, discriminant function 0.183*POTCos + 0.12*MC1TG – 34.79*FL + 46.67*EL + 17.54*ML + 28.28*MPSP – 99.07*PeW – 3.793 > 0 [error 0 % in 21 specimens] => Tetramorium sibiricum

– Postoculo-temporal area of head with fewer costae and costulae, POTCos 8.4 ± 2.4. Anterior dorsum of 1st gaster tergite with less complex elements of stickman-like or reticulate microsculpture, MC1TG 11.2 ± 3.0. Propodeal spines and eye slightly shorter. Discriminant < 0 [error 0 % in 338 specimens] => 2

2. Longest hair near anterolateral pronotal corner shorter, PnHL 0.168 ± 0.020 mm. Anterior dorsum of 1st gaster tergite with more complex elements of stickman-like microsculpture, MC1TG 13.5 ± 2.8. With all linear measurements given in mm, discriminant 0.127*POTCos – 0.173*MC1TG + 85.54*SWd – 49.08*MPSP + 62.52*MPPL + 21.985*PnHL – 9.026 < 0 [error 9.7% in 72 specimens] => Tetramorium indocile

– Longest hair near anterolateral pronotal corner longer, PnHL 0.212 ± 0.022 mm. Anterior dorsum of 1st gaster tergite with only scattered and less complex elements of stickman-like microsculpture, MC1TG 10.6 ± 2.7. Discriminant > 0 [error 1.9 % in 266 specimens] => Tetramorium caespitum

Distribution
Europe, up to 63° N, Caucasus. This species does not occur in North America (Wagner et al. 2017).

Distribution based on Regional Taxon Lists
Palaearctic Region: Afghanistan, Albania, Armenia, Austria, Balearic Islands, Belarus, Belgium, Bulgaria, Canary Islands, Channel Islands, China, Croatia, Democratic Peoples Republic of Korea, Denmark, Estonia, Finland, France, Georgia, Germany, Gibraltar, Hungary, Iberian Peninsula, Iran, Israel, Italy, Jersey, Kazakhstan, Kyrgyzstan, Latvia, Liechtenstein, Lithuania, Luxembourg, Malta, Monaco, Montenegro, Netherlands, Norway, Poland, Republic of Macedonia, Republic of Moldova, Romania, Russian Federation, Slovakia, Spain, Sweden, Switzerland, Turkmenistan, Ukraine, United Kingdom of Great Britain and Northern Ireland.

Biology
Monogynous (Seifert 2007). Hybridizes with Tetramorium immigrans (Wagner et al. 2017; Cordonnier et al. 2019).

Adult sexuals in nests on 25 June ± 14d [28 May, 19 August] (n = 67). Direct swarming behavior observed on 14 June, 15 June at 11:05 true solar time, and 30 June at 07:15 true solar time.

Collingwood (1979) - The species tends to be coastal in North Europe but also inland on heath and on the open borders of woodland, nesting in the earth and also under stones. Colonies are normally single queened, but populous with up to 10,000 or more workers. This species is moderately aggressive, living by predation on other arthropods, scavenging and also from root aphid honeydew. Seeds of various herbs and grasses are often collected into the nest. The alatae are conspicuously large compared with the workers; they are developed in the early summer and fly in late June and July.

Hybridizes with Tetramorium immigrans (Wagner et al. 2017; Cordonnier et al. 2019; Cordonnier et al. 2020).

Other Ants

 * This ant is parasitized by the workerless inquilines (de la Mora et al., 2021; Sanetra & Buschinger, 2000),  (Seifert, 2018; Wagner et al., 2017) and, the inquiline  (de la Mora et al., 2021; Novák & Sadil, 1941; Sanetra et al., 1994; Sanetra et al., 1999; Seifert, 1996; Sanetra & Buschinger, 2000; Seifert, 2018), and the slave-makers ,  (de la Mora et al., 2021; Sanetra & Buschinger, 2000; Seifert, 2018),  (de la Mora et al., 2021; Seifert, 2018) and  (de la Mora et al., 2021; Seifert, 2018).

Hemiptera

 * This species is associated with the aphids, , , , , , and  (Saddiqui et al., 2019 and included references)

Nomenclature

 *  caespitum. Formica caespitum Linnaeus, 1758: 581 (w.) EUROPE. Latreille, 1798: 50 (q.m.); Mayr, 1861: 62 (q.m.); Wheeler, G.C. & Wheeler, J. 1954d: 445 (l.); Hauschteck, 1961: 221 (k.); Imai, 1966: 119 (k.). Combination Manica: Jurine, 1807: 279; in Tetramorium: Mayr, 1855: 426. Senior synonym of fuscula: Smith, F. 1851: 118, Radchenko, 2007: 31; of modesta Foerster: Curtis, 1854: 215; Mayr, 1855: 426; of fusca: Dalla Torre, 1893: 132; of transversinodis: Brown, 1949a: 47; of immigrans: Bolton, 1979: 171; of himalayanum, indocile, transbaicalense: Radchenko, 1992b: 50; of hammi: Bolton, 1995b: 405; of jiangxiense: Wu & Wang, 1995: 82; of fusciclavum: Sanetra, Güsten & Schulz, 1999: 320. Current subspecies: nominal plus barabense, caespitomoravicum, flavidulum, japonicum, pallidum, typicum. See also: Emery, 1909d: 697; Bondroit, 1918: 107; Emery, 1925c: 177; Baroni Urbani, 1971c: 135; Kutter, 1977c: 157; Arnol'di & Dlussky, 1978: 544; Smith, D.R. 1979: 1400; Collingwood, 1979: 84; Cammaerts, Pasteels et al. 1985: 109; Kupyanskaya, 1990: 151; López, 1991a: 31; López, 1991b: 73; López, et al. 1992: 169; Radchenko, Czechowski & Czechowska, 1998: 108.
 * fusca. Formica fusca Leach, 1825: 290 (q.m.) FRANCE. [Unresolved junior primary homonym of Formica fusca Linnaeus, 1758: 580.] Junior synonym of caespitum: Dalla Torre, 1893: 132.
 * fuscula. Myrmica fuscula Nylander, 1846a: 935, pl. 18, figs. 34, 36 (w.q.m.) FINLAND. Junior synonym of caespitum: Smith, F. 1851: 118; Radchenko, 2007: 31.
 * modesta. Myrmica modesta Foerster, 1850a: 49 (w.) GERMANY. Junior synonym of caespitum: Curtis, 1854: 215; Mayr, 1855: 426.
 * himalayanum. Tetramorium caespitum subsp. himalayanum Viehmeyer, 1914b: 38 (w.q.m.) INDIA. Junior synonym of caespitum: Radchenko, 1992b: 50.
 * hammi. Tetramorium caespitum var. hammi Donisthorpe, 1915d: 178 (w.) GREAT BRITAIN. Junior synonym of caespitum: Bolton, 1995b: 408.
 * immigrans. Tetramorium caespitum var. immigrans Santschi, 1927a: 54 (w.) CHILE. Junior synonym of caespitum: Bolton, 1979: 171.
 * transbaicalense. Tetramorium semilaeve subsp. transbaicalense Ruzsky, 1936: 93 (w.) KAZAKHSTAN. Junior synonym of caespitum: Radchenko, 1992b: 50.
 * transversinodis. Myrmica (Myrmica) brevinodis var. transversinodis Enzmann, J. 1946b: 47, figs. 1,2 (w.) U.S.A. Junior synonym of caespitum: Brown, 1949a: 47.
 * fusciclavum. Tetramorium caespitum var. fusciclavum Consani & Zangheri, 1952: 42 (w.) ITALY. [First available use of Tetramorium caespitum subsp. caespitum var. fusciclava Emery, 1925c: 187; unavailable name (Bolton, 1995b: 408).] Junior synonym of caespitum: Sanetra, Güsten & Schulz, 1999: 320.
 * jiangxiense. Tetramorium jiangxiense Wang & Xiao, in Wang, M., Xiao & Wu, 1988: 269, figs. 24, 25 (w.) CHINA. Junior synonym of caespitum: Wu & Wang, 1995: 82.

Type Material
Wagner et al. (2017) - Neotype designation: Schlick-Steiner et al. 2006. Floghult Bohuslan (Sweden), 58.97° N, 11.42° E, 100 m a.s.l., leg. C.A. Collingwood, 21.VI. 2000.

Worker
Wagner et al. (2017) - Larger than most species of complex, CS = 761 ± 50 [591, 867] μm. Dark brown to blackish.

Head moderately elongate, CL / CW = 1.012 ± 0.015 [0.969, 1.043]. Eye rather small, EYE / CS = 0.171 ± 0.005 [0.158, 0.188]. Scape length moderate, SLd / CS = 0.777 ± 0.015 [0.724, 0.812]. Mesosoma long and wide, ML / CS = 1.172 ± 0.026 [1.104, 1.233], MW / CS = 0.645 ± 0.015 [0.605, 0.687].

Promesonotal dorsum convex, metanotal groove shallow. – Head dorsum and occiput with longitudinal costae and costulae, in Iberia longitudinal costae and costulae of head dorsum sometimes interrupted by smooth and shiny areas. Postoculo-temporal area of head with moderate number of longitudinal costae and costulae, POTCos = 7.45 ± 1.92 [3.38, 12.13]. Mesosoma dorsum longitudinally rugulose, in Iberia longitudinal costae and costulae sometimes interrupted by smooth and shiny areas. Lateral side of propodeum with a moderately pronounced smooth and shiny area, Ppss = 39.9 ± 20.0 [13.3, 107.7]. Dorsum of petiolar node smooth or with slightly microreticulate sculpture. General surface appearance on average moderately smooth and shiny compared with other species. – Connected stickman- like or reticulate microsculpture: small units scattered over 1st gastral tergite, MC1TG = 12.62 ± 2.31 [7.00, 19.58]. – Some workers with long c-shaped, crinkly, or sinuous hairs on ventral head posterior to buccal cavity.

Male
Wagner et al. (2017) - Paramere structure belongs to caespitum-like form: ventral paramere lobe with one or two sharp corners; without distinct emargination between paramere lobes in posterior view, both paramere lobes reduced in size; in ventro-posterior view, second corner on ventral paramere lobe missing or < 87 μm apart from first. In posterior view, typically only one sharp corner on ventral lobe.

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