Camponotus piceus

This species is mainly distributed in southern and central Europe and in the southern part of eastern Europe, but it is also present in north-west Africa, Asia Minor, Lebanon, Iran, the Caucasus, and northern Kazakshtan (Czechowski et al. 2002). It is a xerothermophilous species living mostly in open, dry grasslands, and is ground-nesting. Workers are highly polymorphic (Marko et al., 2009; Rigato & Toni, 2011). In Greece it has been collected mostly from shrubs on mountain pastures and along roadsides, also on shrubs along borders of deciduous, mixed and fir forests (Borowiec & Salata, 2021).

Identification
A member of the Camponotus lateralis species group. Seifert, 2019: The character combinations to identify this species can be derived from the key (also see Table 3 and Figures 14 and 15 from this study) and the images on AntWeb.org: CASENT0173136 (minor worker), CASENT0249995 (minor), CASENT09115597 (major, lectotype C. foveolata).

For species delimitation by exploratory and hypothesis-driven data analyses see also Camponotus candiotes and Camponotus atricolor.

Distribution
Marko et al. (2009) - It is present throughout Romania. The occurrence of C. piceus sp. 2 (Seifert 2007) can also be expected, based on preliminary distribution data.

Seifert, 2019: From Iberia to Balkans, south Central Europe north to 51.3°N. It is not known so far from Asia Minor and is apparently a western parapatric sibling species of Camponotus candiotes. There is broad range overlap with Camponotus atricolor in E Central Europe and the Balkans.

This is a common species, known from all Greek provinces (Borowiec et al., 2022).

Distribution based on Regional Taxon Lists
Palaearctic Region: Albania, Algeria, Andorra, Armenia, Austria, Balearic Islands, Belgium, Bulgaria, China, Croatia, Czech Republic, France, Georgia, Germany, Gibraltar, Greece, Hungary, Iberian Peninsula, Iran, Italy, Lebanon, Malta, Montenegro, Poland, Portugal, Republic of Macedonia, Republic of Moldova, Romania, Russian Federation , Slovakia, Slovenia, Spain, Switzerland, Turkey, Ukraine.

Other Insects

 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).

Nomenclature

 * . Formica picea Leach, 1825: 292 (w.q.m.) FRANCE.
 * Combination in Camponotus: Roger, 1863b: 1;
 * combination in C. (Myrmentoma): Menozzi, 1921: 32.
 * [Misspelled as picaea by Ruzsky, 1905b: 255.]
 * Junior synonym of lateralis: Roger, 1859: 231; Mayr, 1863: 399; Roger, 1863b: 1; Forel, 1874: 96 (in list); Emery & Forel, 1879: 448.
 * Subspecies of lateralis: Emery, 1891b: 21; Dalla Torre, 1893: 238; Ruzsky, 1905b: 255; Menozzi, 1921: 32; Menozzi, 1922b: 331; Soudek, 1922: 91; Finzi, 1923: 4; Finzi, 1924a: 14; Emery, 1924c: 170; Emery, 1925b: 120; Kuznetsov-Ugamsky, 1929b: 36; Ceballos, 1956: 313; Pisarski, 1961a: 174.
 * Status as species: Emery, 1925a: 67; Santschi, 1926f: 288; Finzi, 1927b: 52 (in key); Karavaiev, 1927a: 295; Karavaiev, 1927c: 277 (in key); Finzi, 1930d: 318; Santschi, 1931a: 12; Finzi, 1933: 165; Grandi, 1935: 102; Zimmermann, 1935: 58; Karavaiev, 1936: 190; Röszler, 1942a: 52; Röszler, 1951: 90; Consani & Zangheri, 1952: 43; Baroni Urbani, 1964b: 58; Baroni Urbani, 1964c: 162; Kutter, 1968a: 60; Cagniant, 1970c: 37; Baroni Urbani, 1971c: 193; Collingwood, 1971: 164; Baroni Urbani, 1974: 237; Pisarski, 1975: 32; Hamann & Klemm, 1976: 677; Kutter, 1977c: 207; Collingwood, 1978: 91 (in key); Agosti & Collingwood, 1987a: 59; Agosti & Collingwood, 1987b: 283 (in key); Le Moli & Rosi, 1991: 35; Atanassov & Dlussky, 1992: 222; Arakelian, 1994: 87; Bolton, 1995b: 117; Poldi, et al. 1995: 7; Radchenko, 1996b: 1198 (in key); Espadaler, 1997b: 27; Radchenko, 1997b: 706; Collingwood & Prince, 1998: 24 (in key); Gallé, et al. 1998: 216; Czechowski, et al. 2002: 99; Karaman, M.G. & Karaman, 2003: 47; Csösz, & Markó, 2005: 229; Karaman, G.S. & Karaman, 2005: 58; Bračko, 2006: 146; Markó, Sipos, et al. 2006: 66; Petrov, 2006: 71, 108 (in key); Bračko, 2007: 19; Radchenko, 2007: 37; Seifert, 2007: 270; Werner & Wiezik, 2007: 143; Zryanin & Zryanina, 2007: 233; Gratiashvili & Barjadze, 2008: 131; Casevitz-Weulersse & Galkowsky, 2009: 480; Lapeva-Gjonova, et al. 2010: 45; Boer, 2010: 20; Csösz, et al. 2011: 58; Karaman, C. et al. 2011: 189; Karaman, M.G. 2011b: 72; Legakis, 2011: 32; Ran & Zhou, 2011: 69; Borowiec, L. & Salata, 2012: 480; Czechowski, et al. 2012: 249; Guénard & Dunn, 2012: 29; Kiran & Karaman, 2012: 7; Nezhad, et al. 2012: 66; Karaman, C. & Aktaç, 2013: 50 (in key); Borowiec, L. 2014: 39; Bračko, et al. 2014: 18; Lebas, et al. 2016: 140; Radchenko, 2016: 334; Seifert, 2018: 264; Seifert, 2019b: 25.
 * Senior synonym of ebeninus: Dalla Torre, 1893: 238; Ruzsky, 1905b: 255; Soudek, 1922: 91; Emery, 1925a: 70; Emery, 1925b: 120; Baroni Urbani, 1971c: 194; Kutter, 1977c: 207; Atanassov & Dlussky, 1992: 222; Bolton, 1995b: 117; Radchenko, 1997b: 706; Czechowski, et al. 2002: 99; Radchenko, 2016: 334; Salata & Borowiec, 2018c: 43; Seifert, 2019b: 28.
 * Senior synonym of figaro: Seifert, 2019b: 28.
 * Senior synonym of foveolata: Dalla Torre, 1893: 238; Ruzsky, 1905b: 255; Soudek, 1922: 91; Emery, 1925a: 70; Emery, 1925b: 120; Baroni Urbani, 1971c: 194; Kutter, 1977c: 207; Agosti & Collingwood, 1987a: 58; Atanassov & Dlussky, 1992: 222; Bolton, 1995b: 117; Radchenko, 1997b: 706; Czechowski, et al. 2002: 99; Radchenko, 2007: 37; Legakis, 2011: 32; Kiran & Karaman, 2012: 7; Radchenko, 2016: 334; Seifert, 2019b: 28.
 * Material of the unavailable name dusmeti referred here by Seifert, 2019b: 28.
 * ebeninus. Camponotus ebeninus Emery, 1869b: 2, pl. 1, fig. 2 (w.m.) ITALY.
 * [Misspelled as eboninus by Soudek, 1922: 91.]
 * Junior synonym of merula: Emery, 1896d: 373; Ruzsky, 1905b: 255; Emery, 1916b: 226.
 * Junior synonym of foveolata: Forel, 1874: 40; Forel, 1879a: 94 (in text); Emery & Forel, 1879: 449; André, 1882a: 151; Emery, 1925a: 70.
 * Junior synonym of piceus: Dalla Torre, 1893: 238; Ruzsky, 1905b: 255; Soudek, 1922: 91; Emery, 1925a: 70; Emery, 1925b: 120; Baroni Urbani, 1971c: 194; Kutter, 1977c: 207; Atanassov & Dlussky, 1992: 222; Bolton, 1995b: 117; Radchenko, 1997b: 706; Czechowski, et al. 2002: 99; Radchenko, 2016: 334; Seifert, 2019b: 28.
 * figaro. Camponotus (Myrmentoma) figaro Collingwood & Yarrow, 1969: 84.
 * [First available use of Camponotus (Myrmoturba) lateralis subsp. piceus var. figaro Emery, 1924c: 170 (w.) SPAIN; unavailable (infrasubspecific) name.]
 * As unavailable (infrasubspecific) name: Emery, 1925a: 68; Menozzi, 1927b: 92; Santschi, 1932g: 5.
 * Status as species: Collingwood, 1978: 91 (in key); Bolton, 1995b: 99; Collingwood & Prince, 1998: 24 (in key); Borowiec, L. 2014: 31; Lebas, et al. 2016: 138.
 * Junior synonym of piceus: Seifert, 2019b: 28.
 * foveolata. Formica foveolata Mayr, 1853c: 277 (w.q.m.) HUNGARY.
 * Combination in Camponotus: Roger, 1863b: 1;
 * combination in C. (Orthonotomyrmex): Forel, 1915d: 72.
 * Junior synonym of lateralis: Mayr, 1855: 322; Nylander, 1856b: 58; Smith, F. 1858b: 12; Mayr, 1863: 399; Roger, 1863b: 1; Dours, 1873: 164; André, 1874: 201 (in list); Forel, 1874: 96 (in list).
 * Subspecies of lateralis: Forel, 1874: 40; Emery, 1878b: 46; Emery & Forel, 1879: 449; André, 1882a: 151 (in key); Forel, 1886e: clxvii; Forel, 1892i: 306; Forel, 1904f: 425; Forel, 1905b: 183; Forel, 1909c: 106; Forel, 1915d: 72 (in key); Stitz, 1917: 351.
 * Junior synonym of merula: Emery, 1896d: 373; Ruzsky, 1905b: 255 Emery, 1916b: 226.
 * Junior synonym of piceus: Dalla Torre, 1893: 238; Ruzsky, 1905b: 255; Soudek, 1922: 91; Emery, 1925a: 70; Emery, 1925b: 120; Baroni Urbani, 1971c: 194; Kutter, 1977c: 207; Agosti & Collingwood, 1987a: 58; Atanassov & Dlussky, 1992: 222; Bolton, 1995b: 100; Radchenko, 1997b: 706; Czechowski, et al. 2002: 99; Radchenko, 2007: 37; Legakis, 2011: 32; Kiran & Karaman, 2012: 7; Radchenko, 2016: 334; Seifert, 2019b: 28.

Type Material
Camponotus ebeninus. According to a message of Maria Tavano of 16 October 2013 there are no type specimens in the Emery collection of (Seifert, 2019).

Camponotus figaro: Investigated were two syntype workers labeled ‘Cordoba 29-XII-922’, ‘SYNTYPUS Camponotus lateralis piceus var. figaro Emery, 1924’, ‘ANTWEB CASENT 0905390’ from (Seifert, 2019).

Camponotus foveolata. This taxon has been described by Mayr as Formica foveolata from Blocksberg near Ofen in Hungary, from Imola in Italy and from Rauhkogel near Mödling in Austria. Herewith, a lectotype is designated in a worker labeled ‘Imola G.Mayr’, ‘z. G.Mayr.Bd.III. p.101-277’, ‘Form. foveolata det. G.Mayr’,‘Type’ and ‘Lectotype Formica foveolata Mayr, 1853 des. B.Seifert 2012’. One paralectotype male is labeled ‘Imola G. Mayr’, ‘z. G. Mayr. Bd.III. p.101-277’, ‘Form. foveolata det. G. Mayr’ and ‘Paralectotype Formica foveolata Mayr, 1853 des. B.Seifert 2012’. Both specimens are stored in (Seifert, 2019:).

Camponotus picea: This taxon has been described from Nice in southern France. The full text of the original description is ‘Capite, antennis, thorace, abdomine pedibusque piceis, glaberimis, nitentibus; geniculis tarsisque ferrugineis. Corporis longitudo. M 5 mm, g 10 mm, w 5 mm.’ Figures were not given and it seems that no later revising taxonomist has seen original material of Leach and that types do not exist. To secure the identity of this taxon, I designate herewith a neotype of C. piceus in a sample of two workers from near Nice, stored in and labeled ‘FRA: 43.799°N, 7.488°N, 90 m / Menton – 2.8 km N / leg. C. Galkowski 2011.08.10’ and ‘Neotype (top) / Camponotus piceus (Leach 1825) / des. B. Seifert 2018’. (Seifert, 2019).

Taxonomic Notes
Seifert, 2019: Emery’s original description of Camponotus ebeninus, that was collected in the vicinity of Naples, indicates a black ant with a mesosomal shape and sculpture similar to Camponotus piceus. As other blackish species are not known so far from Italy and because nobody is currently able to present counter-evidence, I synonymize it with C. piceus.

Seifert, 2019: Workers of Camponotus figaro with completely reddish pronotum (see data in section 4.3 on the low taxonomic value of color characters) are allocated to the C. piceus cluster with both p = 1.0000 if run as wild-card in a 5-class LDA considering the five black species of the group and they are in any structural character consistent with this species.

Seifert, 2019: The lectotype worker of C. foveolata is allocated to the C. piceus cluster with p = 0.9992 if run as wild-card in a 5-class LDA considering the five black species of the group (Seifert, 2019:).

Seifert, 2019: As type material for C. piceus is unavailable, it appears difficult to understand how revisers could reasonably allocate such a crude description to a certain species. There are several possible candidates for entirely blackish ants of this size from the environs of Nice with a glabrous shining surface, and apparently having no spines or dents on mesosoma (if so, Leach should have mentioned it as he did in other species descriptions). A Formica species, namely Formica gagates or Formica fusca, may be excluded because males of this subgenus do not have a clearly smaller body length than gynes. The jet black Lasius fuliginosus can be excluded too because virgin gynes do not reach 10 mm. Blackish species related to Lasius niger might roughly match the size distribution over the three castes, but ferruginous ‘knees’ (i.e., the femoratibial joint) contrasting the blackish color of femora and tibiae do not occur here as it is with glabrous, shining and jet black surfaces. It is also not very likely that Formica picea could refer to one of the two species of Proformica occurring in the vicinity of Nice (Galkowski et al. 2017) as these do not seem to have big differences in total body length between males and gynes and are more medium to dark brown in overall coloration and not glabrous. Hence, this argumentation finally points to a black species of the Camponotus lateralis group – at least there is no character in Leach’s description that is contradictory to this view. As the geographic distribution of the other blackish species of the group, namely Camponotus atricolor, Camponotus candiotes and Camponotus heidrunvogtae, is much more eastern, there is sufficient reason to maintain the name allocation as it was done by other myrmecologists in the past. To settle this point, I designate herewith a neotype of C. piceus. The neotype and another sample from the same collection are allocated to C. piceus with a mean posterior probability of p = 0.9998 if run as wild-card in a 5-class LDA considering the five black species of the group given in Tab. 3.

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