Camponotus ligniperda

This is a common species present throughout Europe as well as the Caucasus and Asia Minor. Its distribution is generally more southern than that of Camponotus herculeanus (Czechowski et al. 2002). It inhabits mostly mixed and deciduous forests, but can also be found in open habitats. Nests are built mostly in dead trees or wood stumps (Marko et al., 2009). In Greece, this species is known only from mountains of mainland provinces except Thrace, and was also noted from Ionian Islands. It prefers coniferous forests and rocky alpine pastures. Nests were observed in a decaying fir trunk (Borowiec & Salata, 2021).

Identification
Collingwood (1979) - Alitrunk bright yellowish red to red; pubescence is short and sparse, usually absent on medial dorsal surfaces of the first and second gaster tergites. Length: 6-14 mm.

This species is similar in all castes to Camponotus herculeanus but distinguished by the brighter colour and more shining gaster.

Distribution
Central Spain to West Russia, Sicily to Central Sweden (Collingwood 1979).

Distribution based on Regional Taxon Lists
Palaearctic Region: Albania, Andorra, Austria, Belarus, Belgium, Bulgaria, China, Croatia, Denmark, Estonia, Finland, France, Germany, Greece, Hungary, Iberian Peninsula, Latvia, Liechtenstein, Lithuania, Luxembourg, Montenegro, Netherlands, Norway, Poland, Republic of Macedonia, Republic of Moldova, Romania, Russian Federation, Slovakia, Slovenia, Spain, Sweden, Switzerland, Turkey, Ukraine.

Biology
Collingwood (1979) - This species is characteristically found in stony banks and along the sun exposed borders of woodland, either nesting under stones or in dry stumps. It is an aggressive ant biting freely and will sometimes attack other Camponotus or Formica colonies. The larger workers bite their opponents clean through the alitrunk or crush their heads with their strong mandibles. A more xerothermic species than Camponotus herculeanus its habits are otherwise similar.

Nomenclature

 * . Formica ligniperda Latreille, 1802c: 88, pl. 1, fig. 1 (s.w.q.m.) FRANCE.
 * [Misspelled as higniperda by Donisthorpe, 1950e: 1066; misspelled as ligniperedus by Radchenko, 1997a: 557.]
 * Wheeler, G.C. & Wheeler, J. 1953e: 187 (l.); Hauschteck, 1961: 221 (k.).
 * Combination in Camponotus: Mayr, 1861: 36 (in key);
 * combination in C. (Camponotus): Forel, 1914a: 266.
 * Junior synonym of herculeanus: Fabricius, 1804: 396; Brullé, 1833: 326; Lepeletier de Saint-Fargeau, 1835: 209.
 * Subspecies of herculeanus: Forel, 1874: 39 (in key); Emery, 1878a: viii (in list); Emery & Forel, 1879: 447; Forel, 1879a: 58; Mayr, 1886d: 420; Cresson, 1887: 256; Lameere, 1892: 62; Forel, 1892i: 306; Emery, in Dalla Torre, 1893: 233 (footnote); Emery, 1893i: 674; Emery, 1896d: 372 (in list); Ruzsky, 1903b: 301; Ruzsky, 1905b: 223; Wheeler, W.M. 1905f: 402; Wheeler, W.M. 1906b: 22; Wheeler, W.M. 1906c: 325; Wasmann, 1906: 111; Emery, 1908a: 185; Wheeler, W.M. 1908f: 625; Bondroit, 1910: 487; Yano, 1910: 422; Wheeler, W.M. 1910d: 340; Wheeler, W.M. 1910g: 571; Forel, 1914a: 266; Forel, 1915d: 68 (in key); Emery, 1916b: 225; Wheeler, W.M. 1916m: 601; Wheeler, W.M. 1917a: 557; Wheeler, W.M. 1917i: 465; Escherich, 1917: 330 (in key); Menozzi, 1918: 87; Nadig, 1918: 339; Soudek, 1922: 95; Finzi, 1923: 4; Schkaff, 1925: 275; Stärcke, 1926: 119 (in key); Karavaiev, 1927c: 275 (in key); Karavaiev, 1929b: 211; Karavaiev, 1930b: 147; Soudek, 1931: 19; Gösswald, 1932: 19; Arnol'di, 1933b: 602 (in key); Zimmermann, 1935: 58; Cole, 1936a: 39; Teranishi, 1940: 25; Wesson, L.G. & Wesson, R.G. 1940: 103; Ruzsky, 1946: 69; Azuma, 1951: 88; Pisarski, 1961a: 156; Bernard, 1967: 340 (in key).
 * Status as species: Walckenaer, 1802: 159; Gravenhorst, 1807: 286; Lamarck, 1817: 95; Stephens, 1829: 356; Losana, 1834: 311; Schilling, 1839: 53; Nylander, 1846a: 898; Nylander, 1846b: 1045; Lucas, H. 1849: 302; Foerster, 1850a: 11; Schenck, 1852: 20; Mayr, 1855: 304 (redescription); Nylander, 1856b: 55; Gredler, 1858: 3; Smith, F. 1858b: 10; Mayr, 1861: 36 (in key); Meinert, 1861: 310; Mayr, 1863: 399; Roger, 1863b: 1; Walker, 1871: 9; Dours, 1873: 164; Smith, F. 1874: 402; Forel, 1874: 39; André, 1874: 176 (in key); Emery, 1878b: 44; Mayr, 1879: 645; André, 1882a: 142 (in key); Forel, 1886e: clxvii; Provancher, 1887: 228 (in key); Forel, 1889: 255; Nasonov, 1889: 10; Emery, 1892b: 161; Dalla Torre, 1893: 239; Ruzsky, 1896: 68; Forel, 1899c: 130; Forel, 1902b: 180; Ruzsky, 1902d: 5; Forel, 1909c: 105; Bondroit, 1912: 352; Stitz, 1914: 95; Bondroit, 1918: 69; Emery, 1920b: 255; Menozzi, 1922c: 143; Müller, 1923a: 73; Müller, 1923b: 159; Emery, 1925b: 73; Betrem, 1926: 216; Donisthorpe, 1927a: 8; Menozzi, 1927b: 92; Lomnicki, 1928: 10; Grandi, 1935: 102; Karavaiev, 1936: 181 (redescription); Stitz, 1939: 237; Novák & Sadil, 1941: 111 (in key); Röszler, 1942a: 54; Holgersen, 1942: 11; Holgersen, 1943b: 174 (in key); Holgersen, 1944: 179; van Boven, 1947: 181 (in key); Donisthorpe, 1950e: 1066; Röszler, 1950: 210; Röszler, 1951: 92; Chapman & Capco, 1951: 222; Consani & Zangheri, 1952: 43; Ceballos, 1956: 313; Baroni Urbani, 1964c: 163; Arnol’di, 1967: 1822; Kutter, 1968a: 60; Collingwood & Yarrow, 1969: 81; Baroni Urbani, 1971c: 176; Collingwood, 1971: 163; Banert & Pisarski, 1972: 352; Pisarski, 1975: 29; van Boven, 1977: 133; Kutter, 1977c: 205; Collingwood, 1978: 91 (in key); Collingwood, 1979: 91; Agosti & Collingwood, 1987a: 58; Agosti & Collingwood, 1987b: 283 (in key); Nilsson & Douwes, 1987: 68; Atanassov & Dlussky, 1992: 212; Bolton, 1995b: 108; Douwes, 1995: 92; Poldi, et al. 1995: 7; Radchenko, 1996b: 1202 (in key); Espadaler, 1997b: 27; Radchenko, 1997a: 557; Gallé, et al. 1998: 216; Czechowski, et al. 2002: 96; Karaman, M.G. & Karaman, 2003: 47; Csösz, & Markó, 2005: 228; Karaman, G.S. & Karaman, 2005: 58; Bračko, 2006: 146; Markó, Sipos, et al. 2006: 66; Petrov, 2006: 109 (in key); Bračko, 2007: 19; Seifert, 2007: 263; Werner & Wiezik, 2007: 143; Zryanin & Zryanina, 2007: 232; Casevitz-Weulersse & Galkowsky, 2009: 479; Lapeva-Gjonova, et al. 2010: 44; Boer, 2010: 18; Csösz, et al. 2011: 58; Karaman, M.G. 2011b: 71; Legakis, 2011: 31; Ran & Zhou, 2011: 68; Borowiec, L. & Salata, 2012: 479; Czechowski, et al. 2012: 242; Guénard & Dunn, 2012: 29; Kiran & Karaman, 2012: 7; Karaman, C. & Aktaç, 2013: 52 (in key); Borowiec, L. & Salata, 2013: 352; Borowiec, L. 2014: 36 (see note in bibliography); Bračko, et al. 2014: 18; Lebas, et al. 2016: 128; Radchenko, 2016: 330; Salata & Borowiec, 2018c: 43; Seifert, 2018: 257.
 * [Note: Seifert, 2019a: 1, reports herculeanus × ligniperda worker hybrids.]
 * Senior synonym of herculeanoligniperdus: Yasumatsu & Brown, 1951: 31; Pisarski, 1961a: 156; Bolton, 1995b: 108; Czechowski, et al. 2002: 96; Czechowski, et al. 2012: 242; Radchenko, 2016: 330.
 * Senior synonym of obsoleta: Emery, 1892b: 161; Emery, 1925b: 74; Karavaiev, 1936: 181; Bolton, 1995b: 108; Casevitz-Weulersse & Galkowsky, 2009: 479; Radchenko, 2016: 330.
 * Current subspecies: nominal plus afer, nigrescens.
 * herculeanoligniperdus. Camponotus ligniperdus var. herculeanoligniperdus Dalla Torre, 1893: 235.
 * [First available use of Camponotus herculeanus r. ligniperdus var. herculeanoligniperdus Forel, 1874: 39 (w.q.m.) SWITZERLAND; unavailable (infrasubspecific) name.]
 * As unavailable (infrasubspecific) name: Emery & Forel, 1879: 447; Forel, 1879a: 59; André, 1882a: 143; Emery, 1896d: 372 (in list); Ruzsky, 1905b: 225 (in text); Emery, 1908a: 185; Forel, 1915d: 69 (in key); Emery, 1916b: 225; Nadig, 1918: 340; Soudek, 1922: 96.
 * Subspecies of herculeanus: Ruzsky, 1896: 68; Nadig, 1918: 340; Menozzi, 1922c: 144; Karavaiev, 1926e: 192; Stärcke, 1926: 119 (in key); Novák & Sadil, 1941: 110 (in key); Holgersen, 1943b: 172 (in key).
 * Subspecies of ligniperda: Menozzi, 1922c: 144; Emery, 1925b: 74.
 * Status as species: Gösswald, 1932: 25; Stitz, 1939: 242.
 * Junior synonym of ligniperdus: Yasumatsu & Brown, 1951: 31; Pisarski, 1961a: 156; Bolton, 1995b: 108; Czechowski, et al. 2012: 233; Radchenko, 2016: 330.
 * obsoleta. Formica obsoleta Christ, 1791: 509, pl. 60, fig. 5 (q.).
 * [Unresolved junior primary homonym of obsoleta Linnaeus, 1758: 580 (Bolton, 1995b: 114).]
 * Junior synonym of ligniperda: Emery, 1892b: 161; Emery, 1925b: 74; Karavaiev, 1936: 181; Bolton, 1995b: 114; Casevitz-Weulersse & Galkowsky, 2009: 479; Radchenko, 2016: 330.

Type Material


Seifert, 2019: Latreille spent unusually much space arguing about “la torture pour les reconnoitre” of the two huge red-breasted ant species named, at that time, Formica ligniperda and Formica herculeana. There is one statement in the description that suggests non-synonymy of his ant with Camponotus herculeanus: “L’abdomen est...noir, luisant, avec le devant du premier anneau d’un rouge sanguin,...” (The abdomen is black, shining, with the anterior part of the first ring bloodred). This wording does not clearly quantify how large the red surface in front of the first gaster segment really was, but Latreille most probably meant a larger patch of a lighter red that is typical for Camponotus ligniperda (Seifert 1989, 2018). This vague indication gets some support from the climatic and geographic conditions at the type locality Brivela-Gaillarde (45.17°N, 1.53°E, 115 m), the surroundings of which hardly exceed an elevation of 500 m. If we subtract 1.0 °C of global warming from the current climate data of Brive, the mean air temperature from 1 May to 31 August should have been 17.6 °C around the year 1800; this value decreases to 15.0 °C at elevations of 500 m, and the annual precipitation was about 700 mm. These data are within the optimum of the climate niche of C. ligniperda, but represent more marginal conditions within the niche of C. herculeanus (Seifert 1989, 2017, 2018). Accordingly, we may expect C. herculeanus to have been much rarer or absent in this region during Latreille’s time.

Whatever interpretation is given, there is no definite proof which ant Latreille really meant and investigation of type specimens is required. However, according to a message from J. Casevitz-Weulersse of 13 June 2008, there are no specimens in the collection of Muséum National d’Histoire Naturelle Paris that could be considered as types. To finally settle the identity of Camponotus ligniperda, a neotype is fixed herewith at a medium-sized worker labelled “FRA: 45.3600°N, 1.9444°E/Vitrac-sur-Montane 1.8 km S/550 m, leg. Galkowski 2008.08.28” and “Neotype Camponotus ligniperda (Latr. 1802) des B. Seifert 2018”. This specimen was collected with two other workers from a nest found 38 km NE of Brive-la-Gaillarde and is stored in the.

Taxonomic Notes
Seifert (2019) found a low-level of hybridisation between C. ligniperda and Camponotus herculeanus. The frequency of hybridization between the two species is estimated for Central Europe as 0.2–1.0%. This low ratio indicates strong reproductive barriers considering syntopic occurrence at about 10% of the observation sites, a nearly complete overlap of swarming times and basically equal meteorological conditions to release swarming.

Etymology
Seifert, 2019: Two Latin spellings of the ant which Latreille had also named in French “fourmi rongé-bois” have been in use by various authors over the last 60 years: Camponotus ligniperda and C. ligniperdus. This disparity causes confusion (but is in reality not a very important issue because it does not lead to confusion with other Camponotus species). The latter spelling assumes that “ligniperda” is a female adjective attached to the female noun Formica and has to change its ending to “-us” when the species is transferred to the masculine genus Camponotus. The other spelling assumes that Latreille used “ligniperda” as a noun in apposition, which remains unchanged in combination with a genus name of any gender (§ 31.2.1. and § 31.2.2. of ICZN). This interpretation as a noun (meaning “wood destroyer”) was clearly expressed by Kutter (1977) and I consistently followed this usage throughout the last 40 years. Indirect conclusions on Latreille’s naming intention, considering the vernacular compound word “rongé-bois”, appear problematic as I received different proposals by native French speakers. The deciding point in this debate is that “ligniperda” is no accidental linguistic fault—this word really exists as a Latin noun and Pierre André Latreille, as a Latin-educated catholic priest, and Heinrich Kutter, as an old-school pharmacist educated in the 1920s, should have known this. To have this view confirmed, I asked the distinguished Latin expert Prof. Thomas Baier from the Institute of Classical Philology of the University of Würzburg. He fully concurred. This is what he wrote in a letter of 15 July 2013: “.... assessing your problem according to the rules of classical Latin, ligniperda would be a noun, which always is written ligniperda in connection with masculine and feminine, thus not being adapted in its suffix—just as you have assumed. A parallel form is parricida (murderer of relatives). Johann von Schwaben, who killed his uncle Albrecht von Schwaben around 1300, is known since then in the history books (and in Schiller’s Wilhelm Tell) as “Johannes parricida”. What applies to Swabian dukes also applies to ants, si parva licet componere magnis...”.

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