Crematogaster scutellaris

An extremely common, arboreal species reported from the Maghreb to western Russia (Rigato & Toni, 2011).

Identification
A member of the Crematogaster scutellaris group.

Head shining yellowish red contrasting with the brown alitrunk and dark gaster. Antennae 11 segmented. Length: 3.5-5.0 mm (Collingwood 1979).

Distribution
Karaman (2010) - Crematogaster scutellaris is distributed in Italy, Austria, France, Iberian Peninsula and south Germany (Hölzel, 1966; Collingwood & Yarrow, 1969; Collingwood, 1978; Poldi et al., 1995; Seifert, 1996; Heller, 2004). The species is also found in Croatia on the island of Korčula (Bračko, 2006), and in the southeast Bosnia and Herzegovina on Baba Mtn (Fahringer, 1922). Cobelli (1906) cites Crematogaster scutellaris for Slovenia based on his collection of workers. However, the color pattern of the workers, reddish-brown head, thorax and petiole, and black abdomen, matches the description of C. schmidti. Petrov (2008) cite C. scutellaris for Montenegro, providing no locality data.

Crematogaster schmidti and C. scutellaris are allopatric species. The transgression zone between the two species is the region from Slovenia (east slopes of Julian Alpes) to the Adriatic coast of Croatia (Dalmatia), and the southeast Bosnia and Herzegovina. The two species do not hybridize because no intermediate forms have been found in the transgression zone [the territory of former Yugoslavia] (Atanassov & Dlusskij, 1992). Based on the current distribution of C. schmidti and C. scutellaris, we are expecting C. scutellaris to be found in Slovenia, and rejecting the possibility of the same species being present in Montenegro.

Confirmed findings of this species are from the western Mediterranean to the Western Balkans (Croatia) (Borowiec & Salata, 2017) and it is apparently absent from Bulgaria (Lapeva-Gjonova & Antonova, 2022).

Distribution based on Regional Taxon Lists
Palaearctic Region: Andorra, Austria, Balearic Islands, Croatia, Denmark, France, Georgia, Germany, Gibraltar, Greece, Hungary, Iberian Peninsula, Israel, Italy, Malta, Montenegro, Portugal, Republic of Macedonia, Romania, Slovenia, Spain, Switzerland, Tunisia, Turkey, United Kingdom of Great Britain and Northern Ireland.

Biology
Collingwood (1979) - This arboricolous species is frequently introduced with cork from South Europe or North Africa and has on occasion established itself temporarily in and around warehouses and cork factories in England.

Frizzi et al. (2015) - Crematogaster scutellaris is commonly found in both natural and human-managed ecosystems across the Mediterranean Basin. Nests are excavated in tree trunks or logs and can host up to several thousand workers. This is a dominant species that influences ant community dynamics and arthropod communities. Previous studies suggested C. scutellaris is monogynous, has polydomous nests and exhibits inter-nest aggressiveness which varies with distance. This study examined nestmate recognition in a large olive orchard near Florence, Italy, from March 2009 to September 2010. Nests were located by banging on the trunk of trees. Resident colonies would respond with a defensive swarm of ants that would emerge from their nest holes. Genetic, chemical (CHCs) and spatial characteristics (from 5 to 175 m distances) were examined and behavioral experiments (5 x 5 battles) performed for a range of within and between colony analyses. Polydomy was confirmed by genetic analysis. Many nests were spread across more than a single tree. Genetic analysis supported there being a single queen per colony, with a core nest containing the queen and a network of satellite nests. Genetic differences between nests were strong but were not correlated with distance. This implies a long distance spatial dispersal strategy during mating and/or prior to colony foundation. Cuticular chemical profiles varied significantly among both nests and colonies. This appeared to be explained by a combination of genetic and environmental factors. Aggression between colonies was uneven, occurring in ~50% of the staged tests. Aggressiveness increased with both spatial proximity and similarity in cuticular hydrocarbon profiles among colonies, suggesting a “nasty neighbor” effect.

Bordoni et al. (2018) experimentally showed how Crematogaster scutellaris workers exposed to the same pathogen as their mother exhibited increased survival compared to control workers. These results suggest social immunity, passed on from one generation to the next, can be important for social organisms. "They "found, for the first time in ants, that founding queens exposed to a non-lethal dose of an entomopathogenic fungus produce offspring with an increased resistance to the same pathogen. This phenomenon did not reveal apparent costs to foundresses and workers, at least with respect to their survival and reproductive output."

Guiliani et al. (2019) observed this species foraging on extrafloral nectaries of the invasive Reynoutria x bohemica (Polygonaceae) in Tuscany. The habitats examined were river banks and disturbed habitats.

Association with Other Organisms

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

Nomenclature

 * . Formica scutellaris Olivier, 1792: 497 (w.) FRANCE.
 * Type-material: syntype workers (number not stated).
 * Type-locality: France: Provence (no collector’s name).
 * Type-depository: MNHN.
 * Mayr, 1855: 470 (q.m.).
 * Combination in Myrmica: Schenck, 1852: 132;
 * combination in Crematogaster: Mayr, 1855: 469;
 * combination in C. (Acrocoelia): Emery, 1922e: 143;
 * combination in C. (Crematogaster): Bolton, 1995b: 166.
 * Status as species: Latreille, 1798: 48; Latreille, 1802c: 261; Schenck, 1852: 132; Mayr, 1855: 469 (redescription); Nylander, 1856b: 100; Gredler, 1858: 30; Smith, F. 1858b: 135; Roger, 1859: 259; Mayr, 1861: 74 (in key); Mayr, 1863: 404; Roger, 1863b: 36; Emery, 1869b: 23; Dours, 1873: 169; Forel, 1874: 69 (in key); André, 1874: 201 (in key); Emery, 1878a: x (in list); Emery, 1878b: 58; Emery & Forel, 1879: 464; Emery, 1880: 398; André, 1883b: 392 (in key); Forel, 1886e: clxviii; Provancher, 1887: 244; Cresson, 1887: 262; Forel, 1889: 257; Saunders, E. 1890: 205; Emery, 1891b: 13; Dalla Torre, 1893: 85; Medina, 1893: 105; Forel, 1894d: 24; Forel, 1895d: 227; Saunders, E. 1896: 42; Forel, 1902a: 154; Ruzsky, 1902d: 31; Forel, 1904b: 372; Forel, 1904c: 6; Ruzsky, 1905b: 488; Forel, 1907a: 22; Emery, 1908d: 23; Forel, 1911d: 340; Emery, 1912e: 652; Krausse, 1912b: 164; Stitz, 1914: 73; Emery, 1914d: 157; Forel, 1915d: 32 (in key); Donisthorpe, 1915d: 337; Emery, 1916b: 155; Menozzi, 1918: 85; Bondroit, 1918: 113; Santschi, 1919e: 244; Menozzi, 1921: 27; Müller, 1921: 48; Emery, 1922e: 143; Finzi, 1922: 120; Menozzi, 1922b: 327; Müller, 1923b: 70; Finzi, 1924a: 13; Emery, 1924b: 12; Emery, 1924c: 165; Santschi, 1925g: 348; Lomnicki, 1925b: 1; Donisthorpe, 1926a: 7; Wheeler, W.M. 1926: 3; Karavaiev, 1926b: 109; Menozzi, 1926b: 182; Stärcke, 1926: 84 (in key); Karavaiev, 1927a: 289; Kutter, 1927: 99; Donisthorpe, 1927b: 388; Menozzi, 1927b: 90; Kutter, 1928: 66; Finzi, 1930d: 313; Cori & Finzi, 1931: 238; Santschi, 1931a: 5; Santschi, 1932c: 70; Finzi, 1933: 164; Menozzi, 1934: 159; Grandi, 1935: 100; Zimmermann, 1935: 22; Stitz, 1939: 128; Finzi, 1940: 160; Novák & Sadil, 1941: 82 (in key); Bernard, 1950a: 4; Donisthorpe, 1950e: 1061; Consani & Zangheri, 1952: 40; Ceballos, 1956: 304; Bernard, 1956b: 258; Bernard, 1959: 346; Fromantin & Soulié, 1961: 95; Baroni Urbani, 1964a: 4; Baroni Urbani, 1964b: 44; Baroni Urbani, 1964c: 154; Cagniant, 1964: 88; Cagniant, 1966b: 279; Bernard, 1967: 164 (redescription); Baroni Urbani, 1968b: 449; Kutter, 1968a: 59; Baroni Urbani, 1969a: 332; Collingwood & Yarrow, 1969: 65; Cagniant, 1970a: 418; Baroni Urbani, 1971c: 79; Baroni Urbani, 1974: 233; Bolton & Collingwood, 1975: 3 (in key); Aktaç, 1977: 121; Kutter, 1977c: 88; Collingwood, 1978: 82 (in key); Collingwood, 1979: 66; Schembri & Collingwood, 1981: 430; Agosti & Collingwood, 1987a: 54; Agosti & Collingwood, 1987b: 272 (in key); Casevitz-Weulersse, 1990b: 137; Le Moli & Rosi, 1991: 30; Bolton, 1995b: 162; Mei, 1995: 762; Poldi, et al. 1995: 4; Espadaler, 1997b: 29; Collingwood & Prince, 1998: 14 (in key); Gallé, et al. 1998: 214; Cagniant, 2005: 10; Bračko, 2006: 136; Cagniant, 2006a: 198; Petrov, 2006: 94 (in key); Bračko, 2007: 17; Seifert, 2007: 215; Radchenko, 2007: 33; Karaman, M.G. 2008: 20 (in key), Gratiashvili & Barjadze, 2008: 138; Casevitz-Weulersse & Galkowski, 2009: 487; Lapeva-Gjonova, et al. 2010: 18; Boer, 2010: 46; Csösz, et al. 2011: 57; Legakis, 2011: 14; Borowiec, L. & Salata, 2012: 490; Kiran & Karaman, 2012: 18; Borowiec, L. 2014: 68 (see note in bibliography); Lebas, et al. 2016: 272; Steiner, et al. 2017: 8; Salata & Borowiec, 2018c: 44; Seifert, 2018: 178.
 * Senior synonym of corsica: Casevitz-Weulersse, 1990b: 137; Bolton, 1995b: 162; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * Senior synonym of degener: Casevitz-Weulersse, 1990b: 137; Bolton, 1995b: 162; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * Senior synonym of grouvellei: Bernard, 1967: 164; Bolton, 1995b: 162; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * Senior synonym of haematocephala: Roger, 1859: 259; Mayr, 1863: 405; Roger, 1863b: 36; Forel, 1874: 100 (in list); Emery & Forel, 1879: 464; Emery, 1891b: 13; Dalla Torre, 1893: 85; Emery, 1922e: 143; Fromantin & Soulié, 1961: 95; Bolton, 1995b: 162.
 * Senior synonym of hybrida: Baroni Urbani, 1964a: 4; Baroni Urbani, 1971c: 80; Bolton, 1995b: 162.
 * Senior synonym of lichtensteini: Bernard, 1967: 164; Bolton, 1995b: 162; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * Senior synonym of rediana: Mayr, 1855: 469; Nylander, 1856b: 100; Smith, F. 1858b: 135; Mayr, 1861: 72 (in key); Mayr, 1863: 405; Roger, 1863b: 36; André, 1874: 204 (in list); Forel, 1874: 100 (in list); Emery & Forel, 1879: 464; Emery, 1891b: 13; Dalla Torre, 1893: 85; Ruzsky, 1905b: 488; Emery, 1922e: 143; Fromantin & Soulié, 1961: 95; Baroni Urbani, 1971c: 79; Bolton, 1995b: 162.
 * Senior synonym of rubriceps: Mayr, 1855: 469; Nylander, 1856b: 100; Smith, F. 1858b: 135; Mayr, 1861: 72 (in key); Mayr, 1863: 405; Roger, 1863b: 36; Dours, 1873: 169; André, 1874: 204 (in list); Forel, 1874: 100 (in list); Emery & Forel, 1879: 464; Emery, 1891b: 13; Dalla Torre, 1893: 85; Ruzsky, 1905b: 488; Emery, 1922e: 143; Fromantin & Soulié, 1961: 95; Baroni Urbani, 1971c: 79; Bolton, 1995b: 162; Radchenko, 2007: 33.
 * Senior synonym of ruficeps: Mayr, 1855: 469; Nylander, 1856b: 100; Smith, F. 1858b: 135; Mayr, 1861: 72 (in key); Mayr, 1863: 405; Roger, 1863b: 36; André, 1874: 204 (in list); Forel, 1874: 100 (in list); Emery & Forel, 1879: 464; Emery, 1891b: 13; Dalla Torre, 1893: 85; Ruzsky, 1905b: 488; Wheeler, W.M. 1911f: 158; Emery, 1922e: 143; Fromantin & Soulié, 1961: 95; Baroni Urbani, 1971c: 79; Bolton, 1995b: 162.
 * Material of the nomen nudum capiterubro referred here by Baroni Urbani, 1971c: 80.
 * Distribution: Algeria, Andorra, Austria, Bosnia, Bulgaria, Croatia, France (+ Corsica), Goergia, Germany, Gibraltar, Greece, Hungary, Israel, Italy (+ Sardinia, Sicily), Libya, Malta, Morocco, Netherlands, Portugal, Spain (+ Balearics), Switzerland, Tunisia, Turkey.
 * [Note: recorded distribution after Borowiec, L. 2014: 68, who claims that some localities may be based on misidentifications.]
 * Current subspecies: nominal plus alii, nigra (unresolved junior homonym), tenuispina.
 * corsica. Crematogaster scutellaris var. corsica Santschi, 1921b: 435 (w.) FRANCE (Corsica).
 * Type-material: syntype workers (number not stated).
 * Type-locality: France: Corsica (Desanti).
 * Type-depository: NHMB.
 * Subspecies of scutellaris: Santschi, 1937e: 312 (in key); Baroni Urbani, 1971c: 82.
 * Junior synonym of scutellaris: Casevitz-Weulersse, 1990b: 137; Bolton, 1995b: 151; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * degener. Crematogaster scutellaris var. degener Santschi, 1937e: 310 (w.) TUNISIA, FRANCE (Corsica).
 * Type-material: syntype workers (number not stated).
 * Type-localities: Tunisia: Sousse, 6.x.1910 (F. Santschi), Tunisia: Hammamet, viii.1918 (F. Santschi), 1 worker France: Corsica (J. Perez).
 * Type-depository: NHMB.
 * Subspecies of scutellaris: Baroni Urbani, 1971c: 82.
 * Junior synonym of scutellaris: Casevitz-Weulersse, 1990b: 137; Bolton, 1995b: 151; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * grouvellei. Crematogaster scutellaris var. grouvellei Bondroit, 1918: 115 (w.q.) FRANCE.
 * Type-material: syntype workers, syntype queens (numbers not stated).
 * Type-locality: France: Hyères (A. Grouvelle).
 * Type-depository: ISNB (perhaps also in MNHN).
 * Subspecies of scutellaris: Emery, 1922e: 143; Santschi, 1937e: 312 (in key); Fromantin & Soulié, 1961: 96.
 * Junior synonym of scutellaris: Bernard, 1967: 164; Bolton, 1995b: 154; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * haematocephala. Formica haematocephala Leach, 1825: 291 (w.q.m.) FRANCE.
 * Type-material: syntype worker(s), syntype queen(s), syntype male(s) (numbers not stated).
 * Type-locality: France: vic. Nice (W.E. Leach).
 * Type-depository: BMNH.
 * [Note: depository according to Horn & Kahle, 1935: 151.]
 * Combination in Crematogaster: Roger, 1863b: 36.
 * Junior synonym of scutellaris: Roger, 1859: 259; Mayr, 1863: 405; Roger, 1863b: 36; Forel, 1874: 100 (in list); Emery & Forel, 1879: 464; Emery, 1891b: 13; Dalla Torre, 1893: 85; Emery, 1922e: 143; Fromantin & Soulié, 1961: 95; Bolton, 1995b: 154.
 * hybrida. Crematogaster scutellaris var. hybrida Zimmermann, 1935: 22.
 * Type-material: syntype workers (number not stated).
 * Type-locality: Italy: Palmanova (Friuli) (no collector’s name).
 * Type-depository: MSNG.
 * [First available use of Crematogaster scutellaris subsp. schmidti var. hybrida Emery, 1916b: 157 (w.) ITALY; unavailable (infrasubspecific) name (Agosti & Collingwood, 1987a: 61).]
 * As unavailable (infrasubspecific) name: Emery, 1922e: 144; Santschi, 1937e: 312 (in key).
 * Junior synonym of scutellaris: Baroni Urbani, 1964a: 4; Baroni Urbani, 1971c: 80; Bolton, 1995b: 154.
 * lichtensteini. Crematogaster scutellaris var. lichtensteini Bondroit, 1918: 114 (w.) FRANCE.
 * Type-material: syntype workers (number not stated).
 * Type-locality: France: Montpellier (J.-L. Lichtenstein).
 * Type-depository: ISNB (perhaps also in MNHN).
 * Subspecies of scutellaris: Emery, 1922e: 143; Santschi, 1937e: 312 (in key); Fromantin & Soulié, 1961: 96.
 * Junior synonym of scutellaris: Bernard, 1967: 164; Bolton, 1995b: 156; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * rediana. Myrmica rediana Gené, 1841: 399 (w.) ITALY.
 * Type-material: syntype workers (number not stated).
 * Type-locality: Italy: (no further data).
 * Type-depository: unknown.
 * [Note: Horn & Kahle, 1935: 88, say that Gené non-coleopteran material is in “Sammlg. an C. Bassi (Mailand)”.]
 * Combination in Acrocoelia: Mayr, 1853b: 103;
 * combination in Crematogaster: Roger, 1863b: 36;
 * Status as species: Mayr, 1853b: 103.
 * Junior synonym of scutellaris: Mayr, 1855: 469; Nylander, 1856b: 100; Smith, F. 1858b: 135; Mayr, 1861: 72 (in key); Mayr, 1863: 405; Roger, 1863b: 36; André, 1874: 204 (in list); Forel, 1874: 100 (in list); Emery & Forel, 1879: 464; Emery, 1891b: 13; Dalla Torre, 1893: 85; Ruzsky, 1905b: 488; Emery, 1922e: 143; Fromantin & Soulié, 1961: 95; Baroni Urbani, 1971c: 79; Bolton, 1995b: 161.
 * rubriceps. Myrmica rubriceps Nylander, 1849: 44 (w.) ITALY (Sicily).
 * Type-material: 2 syntype workers (Radchenko, 2007: 33).
 * Type-locality: Italy: Sicily, Messina (C. Zeller).
 * Type-depository: ZMHF.
 * Combination in Crematogaster: Roger, 1863b: 36.
 * Junior synonym of scutellaris: Mayr, 1855: 469; Nylander, 1856b: 100; Smith, F. 1858b: 135; Mayr, 1861: 72 (in key); Mayr, 1863: 405; Roger, 1863b: 36; Dours, 1873: 169; André, 1874: 204 (in list); Forel, 1874: 100 (in list); Emery & Forel, 1879: 464; Emery, 1891b: 13; Dalla Torre, 1893: 85; Ruzsky, 1905b: 488; Emery, 1922e: 143; Fromantin & Soulié, 1961: 95; Baroni Urbani, 1971c: 79; Bolton, 1995b: 161; Radchenko, 2007: 33.
 * ruficeps. Acrocoelia ruficeps Mayr, 1853a: 147 (w.) ITALY.
 * Type-material: syntype workers (number not stated).
 * Type-localities: Italy: S Tyrol, Zenoburg, nr Meran (G. Mayr ?), Italy: S Tyrol, Botzan (= Bolzano) (Gredler), Italy: Venedig (= Venice), Lido I. (G. Mayr).
 * Type-depository: NHMW.
 * [Misspelled as rubriceps by Roger, 1863b: 36.]
 * Combination in Crematogaster: Roger, 1863b: 36.
 * Junior synonym of scutellaris: Mayr, 1855: 469; Nylander, 1856b: 100; Smith, F. 1858b: 135; Mayr, 1861: 72 (in key); Mayr, 1863: 405; Roger, 1863b: 36; André, 1874: 204 (in list); Forel, 1874: 100 (in list); Emery & Forel, 1879: 464; Emery, 1891b: 13; Dalla Torre, 1893: 85; Ruzsky, 1905b: 488; Wheeler, W.M. 1911f: 158; Emery, 1922e: 143; Fromantin & Soulié, 1961: 95; Baroni Urbani, 1971c: 79; Bolton, 1995b: 161.

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