Camponotus lateralis

This arboreal species is present in southern and central Europe, as well as in Crimea, the Caucasus, Asia Minor, Kopet Dag (Iran) and NW Africa (Radchenko 1997a). It inhabits warm, xerotherm areas; nests are mostly built in dead wood (Marko et al., 2009; Rigato & Toni, 2011). Pashaei Rad et al. (2018) found this species in Iran on city plant in a Caspian moist littoral area.

Identification
A member of the Camponotus lateralis species group. Ionescu-Hirsch (2009) - C. lateralis is characterized by a deep metanotal groove and a flat or concave propodeal dorsum posteriorly. The petiolar scale is strongly convex anterodorsally and flat posteriorly. The body is feebly sculptured and shiny. The pilosity on the dorsum of the propodeum consists of a transversal row of six erect setae at the junction with declivity and short apressed pubescence. C. lateralis has a weak ground sculpture, is mostly shiny, and has yellowish-brown head and mesosoma, occasionally reddish-brown to dark blackish-brown, and black gaster. C. lateralis and Camponotus staryi (its sister species, according to Radchenko, 1997c) are similar, except for the propodeal dorsum being lower than the highest point of the mesonotum in C. lateralis, as opposed to a propodeal dorsum exceeding the highest point of the mesonotum in C. staryi, and a uniformly dark gaster, as opposed to a mostly dark brown gaster with the first segment ochraceous. C. lateralis is similar to Camponotus dalmaticus except for its head being paler than gaster, as opposed to the head equally dark as the gaster. C. lateralis is similar to Camponotus rebeccae in coloration but differs from it by a deep metanotal groove, as opposed to a shallow metanotal groove and by a flat or concave propodeal dorsum, as opposed to a convex propodeal dorsum in C. rebeccae.

Distribution
Ionescu-Hirsch (2009) - Southern and central Europe, Crimea, Caucasus, northwestern Morocco, and eastern Mediterranean (Radchenko, 1997c).

Marko et al. (2009) - This species occurs in every region of Romania, although data are scattered and scarce. The occurrence of C. lateralis sp. 2 (Seifert 2007) can also be expected, based on the preliminary distribution data.

Distribution based on Regional Taxon Lists
Palaearctic Region: Albania, Algeria, Andorra, Armenia, Balearic Islands, Bulgaria, Croatia, Cyprus, France , Georgia, Germany, Gibraltar, Greece, Hungary, Iberian Peninsula, Iran, Israel, Italy, Malta, Monaco, Montenegro, Morocco, Portugal, Republic of Macedonia, Romania, Russian Federation, Slovakia, Slovenia, Spain, Switzerland, Turkey, Turkmenistan, Ukraine.

Biology
The color morph of Camponotus lateralis which occurs in the West and Central Mediterranean mimics the pigmentation pattern of the aggressive and unpalatable Crematogaster scutellaris in order to achieve protective Batesian mimicry in a parabiotic association with this dominant species (Seifert, 2019).

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.

Nomenclature

 * . Formica lateralis Olivier, 1792: 497 (q.) FRANCE.
 * Mayr, 1853b: 103 (s.w.m.); Hauschteck, 1962: 219 (k.).
 * Combination in Camponotus: Mayr, 1861: 36 (in key);
 * combination in C. (Orthonotomyrmex): Forel, 1913d: 436;
 * combination in C. (Myrmentoma): Forel, 1912i: 92; Emery, 1925b: 120;
 * combination in Orthonotomyrmex: Novák & Sadil, 1941: 109 (in key).
 * Status as species: Latreille, 1798: 39; Latreille, 1802c: 172; Lepeletier de Saint-Fargeau, 1835: 217; Mayr, 1853b: 103; Mayr, 1855: 322 (redescription); Nylander, 1856b: 58; Gredler, 1858: 4; Smith, F. 1858b: 12, 52; Roger, 1859: 231; Mayr, 1861: 36 (in key); Mayr, 1863: 399; Roger, 1863b: 1; Emery, 1869b: 2; Emery, 1870: 193; Dours, 1873: 164; Forel, 1874: 40 (in key); André, 1874: 177 (in key); Mayr, 1877: 20 (in list); Emery, 1878a: viii (in list); Emery, 1878b: 45; Emery & Forel, 1879: 448; Forel, 1879a: 94; André, 1881b: 55; André, 1882a: 150 (in key); Costa, 1883: 60; Emery, 1884a: 376; Forel, 1886e: clxvii; Cresson, 1887: 255; Nasonov, 1889: 12; Forel, 1889: 256; Forel, 1890a: lxvi; Emery, 1891b: 21; Forel, 1892i: 306; Dalla Torre, 1893: 237; Forel, 1894d: 3, 41; Emery, 1896d: 373 (in list); Ruzsky, 1902d: 7; Ruzsky, 1905b: 251; Emery, 1908d: 24; Forel, 1909c: 106; Bondroit, 1910: 487; Karavaiev, 1910b: 12; Forel, 1911d: 354; Krausse, 1911: 18; Krausse, 1912b: 166; Karavaiev, 1912a: 20; Emery, 1914d: 159; Forel, 1915d: 71 (in key); Emery, 1916b: 226; Stitz, 1917: 350; Escherich, 1917: 330 (in key); Bondroit, 1918: 78; Santschi, 1919e: 247; Menozzi, 1921: 32; Menozzi, 1922b: 331; Soudek, 1922: 90; Müller, 1923a: 73; Müller, 1923b: 163; Finzi, 1923: 4; Lomnicki, 1925b: 3; Emery, 1925a: 68; Emery, 1925b: 120; Santschi, 1925g: 359; Wheeler, W.M. 1926: 5; Donisthorpe, 1926b: 18; Karavaiev, 1926e: 193; Menozzi, 1926b: 182; Santschi, 1926f: 288; Karavaiev, 1927a: 295; Wheeler, W.M. 1927g: 117; Finzi, 1927b: 52 (in key); Santschi, 1929e: 163; Finzi, 1930c: 24; Finzi, 1930d: 318; Cori & Finzi, 1931: 240; Santschi, 1931a: 12; Gösswald, 1932: 25; Santschi, 1932g: 4; Finzi, 1933: 165; Grandi, 1935: 102; Zimmermann, 1935: 59; Stitz, 1939: 254; Novák & Sadil, 1941: 109 (in key); Bernard, 1945: 137; Donisthorpe, 1950e: 1067; Consani & Zangheri, 1952: 43; Ceballos, 1956: 312; Bernard, 1956b: 260; Bernard, 1959: 351; Pisarski, 1961a: 170; Baroni Urbani, 1964b: 57; Baroni Urbani, 1964c: 162; Cagniant, 1964: 92; Bernard, 1967: 344 (redescription); Baroni Urbani, 1968b: 486; Cagniant, 1968a: 146; Kutter, 1968a: 60; Baroni Urbani, 1969a: 334; Collingwood & Yarrow, 1969: 84; Cagniant, 1970c: 37; Baroni Urbani, 1971c: 191; Collingwood, 1971: 164; Baroni Urbani, 1974: 238; Pisarski, 1975: 32; Kutter, 1977c: 207; Collingwood, 1978: 91 (in key); Schembri & Collingwood, 1981: 438; Agosti & Collingwood, 1987a: 58; Agosti & Collingwood, 1987b: 283 (in key); Kugler, J. 1988: 259; Dlussky, Soyunov & Zabelin, 1990: 136; Le Moli & Rosi, 1991: 35; Atanassov & Dlussky, 1992: 225; Cagniant & Espadaler, 1993: 436 (in key); Arakelian, 1994: 86; Bolton, 1995b: 107; Poldi, et al. 1995: 7; Radchenko, 1996b: 1197 (in key); Cagniant, 1996b: 92; Espadaler, 1997b: 27; Radchenko, 1997b: 706; Collingwood & Prince, 1998: 24 (in key); Gallé, et al. 1998: 216; Karaman, M.G. & Karaman, 2003: 47; Bračko, 2006: 145; Cagniant, 2006a: 194; Markó, Sipos, et al. 2006: 66; Petrov, 2006: 108 (in key); Bračko, 2007: 19; Seifert, 2007: 268; Werner & Wiezik, 2007: 156; Gratiashvili & Barjadze, 2008: 131; Casevitz-Weulersse & Galkowsky, 2009: 479; Vonshak, et al. 2009: 39; Ionescu-Hirsch, 2010: 81; Lapeva-Gjonova, et al. 2010: 44; Csösz, et al. 2011: 58; Karaman, C. et al. 2011: 189; Karaman, M.G. 2011b: 70; Legakis, 2011: 31; Borowiec, L. & Salata, 2012: 477; Kiran & Karaman, 2012: 7; Karaman, C. & Aktaç, 2013: 51 (in key); Borowiec, L. 2014: 35 (see note in bibliography); Bračko, et al. 2014: 18; Lebas, et al. 2016: 138; Radchenko, 2016: 333; Borowiec, L. & Salata, 2018: 5; Salata & Borowiec, 2018c: 43; Seifert, 2018: 262; Seifert, 2019b: 17.
 * Senior synonym of armouri: Collingwood, 1978: 73; Bolton, 1995b: 107; Radchenko, 2016: 333; Seifert, 2019b: 17.
 * Senior synonym of axillaris: Mayr, 1855: 322; Nylander, 1856b: 58; Smith, F. 1858b: 12; Mayr, 1863: 399; Roger, 1863b: 1; André, 1874: 201 (in list); Forel, 1874: 97 (in list); Emery & Forel, 1879: 448; Dalla Torre, 1893: 237; Emery, 1896d: 373; Emery, 1925b: 120; Baroni Urbani, 1971c: 192; Bolton, 1995b: 107; Radchenko, 2016: 333; Seifert, 2019b: 17.
 * Senior synonym of balearis: Collingwood, 1978: 73; Bolton, 1995b: 107; Radchenko, 2016: 333; Seifert, 2019b: 17.
 * Senior synonym of bicolor Latreille: Mayr, 1853b: 102; Mayr, 1855: 322; Nylander, 1856b: 58; Smith, F. 1858b: 12; Mayr, 1863: 399; Roger, 1863b: 1; André, 1874: 201 (in list); Forel, 1874: 96 (in list); Emery & Forel, 1879: 448; Dalla Torre, 1893: 237; Emery, 1896d: 373; Forel, 1915d: 71 (in key); Soudek, 1922: 90; Emery, 1925b: 120; Baroni Urbani, 1971c: 192; Bolton, 1995b: 107; Radchenko, 2016: 333.
 * Senior synonym of hemipsila: Forel, 1894d: 5; Emery, 1896d: 373; Emery, 1925b: 120; Bolton, 1995b: 107; Radchenko, 2016: 333.
 * Senior synonym of kosswigi: Radchenko, 1997b: 706; Kiran & Karaman, 2012: 7; Radchenko, 2016: 333; Seifert, 2019b: 18.
 * Senior synonym of melanogastes: Mayr, 1853b: 102; Mayr, 1855: 322; Nylander, 1856b: 58; Smith, F. 1858b: 12; Mayr, 1863: 399; Roger, 1863b: 1; André, 1874: 201 (in list); Forel, 1874: 96 (in list); Emery & Forel, 1879: 448; Dalla Torre, 1893: 237; Emery, 1896d: 373; Forel, 1915d: 71 (in key); Soudek, 1922: 90; Emery, 1925b: 120; Pisarski, 1975: 32; Bolton, 1995b: 107; Casevitz-Weulersse & Galkowsky, 2009: 479; Radchenko, 2016: 333; Seifert, 2019b: 17.
 * Senior synonym of merula: Seifert, 2019b: 17.
 * Senior synonym of pallidinervis: Roger, 1861b: 163; Mayr, 1863: 399; Emery & Forel, 1879: 448; Dalla Torre, 1893: 203 (footnote), 237; Emery, 1925b: 120; Bolton, 1995b: 107; Radchenko, 2016: 333.
 * Senior synonym of purius: Seifert, 2019b: 17.
 * [Note: appropriate subgenus is not mentioned by the authors, but as they stress its similarity with barbatus the subgenus is assumed to be C. (Tanaemyrmex).]
 * armouri. Camponotus (Myrmentoma) lateralis subsp. armouri Wheeler, W.M. 1926: 5 (w.q.m.) SPAIN (Balearic Is).
 * Subspecies of lateralis: Ceballos, 1956: 312.
 * Synonym of balearis: Santschi, 1936c: 207.
 * [Note: Santschi gives balearis as senior synonym, but armouri has priority (Bolton, 1995b: 86).]
 * Junior synonym of lateralis: Collingwood, 1978: 73; Bolton, 1995b: 86; Radchenko, 2016: 333; Seifert, 2019b: 17.
 * axillaris. Formica axillaris Spinola, 1808: 243 (w.) ITALY.
 * Junior synonym of lateralis: Mayr, 1855: 322; Nylander, 1856b: 58; Smith, F. 1858b: 12; Mayr, 1863: 399; Roger, 1863b: 1; André, 1874: 201 (in list); Forel, 1874: 97 (in list); Emery & Forel, 1879: 448; Dalla Torre, 1893: 237; Emery, 1896d: 373; Emery, 1925b: 120; Baroni Urbani, 1971c: 192; Bolton, 1995b: 87; Radchenko, 2016: 333; Seifert, 2019b: 17.
 * balearis. Camponotus (Myrmentoma) lateralis var. balearis Santschi, 1929e: 164.
 * [First available use of Camponotus lateralis st. spissinodis var. balearis Santschi, 1925g: 360, fig. 3b (w.) SPAIN (Balearic Is, and mainland); unavailable (infrasubspecific) name.]
 * Subspecies of lateralis: Santschi, 1932c: 72; Santschi, 1936c: 207.
 * Synonym of armouri: Santschi, 1936c: 207.
 * [Note: Santschi, 1936c: 207, gives balearis as senior synonym, but armouri has priority (Bolton, 1995b: 87).]
 * Junior synonym of lateralis: Collingwood, 1978: 73; Bolton, 1995b: 107; Radchenko, 2016: 333; Seifert, 2019b: 17.
 * bicolor. Formica bicolor Latreille, 1798: 43 (w.) FRANCE.
 * [Unresolved junior primary homonym of Formica bicolor Fabricius, 1793: 351 (Emery, 1921a: 26; Bolton, 1995b: 88).]
 * Subspecies of lateralis: Forel, 1874: 40.
 * Junior synonym of lateralis: Mayr, 1853b: 102; Mayr, 1855: 322; Nylander, 1856b: 58; Smith, F. 1858b: 12; Mayr, 1863: 399; Roger, 1863b: 1; André, 1874: 201 (in list); Forel, 1874: 96 (in list); Emery & Forel, 1879: 448; Dalla Torre, 1893: 237; Emery, 1896d: 373; Forel, 1915d: 71 (in key); Soudek, 1922: 90; Emery, 1925b: 120; Baroni Urbani, 1971c: 192; Bolton, 1995b: 88; Radchenko, 2016: 333.
 * hemipsila. Formica hemipsila Foerster, 1850b: 492 (q.) ALGERIA.
 * Status as species: Mayr, 1863: 416; Dalla Torre, 1893: 233.
 * Subspecies of maculatus: Emery, 1891b: 20; Emery, in Dalla Torre, 1893: 233, 241 (footnote).
 * Junior synonym of lateralis: Forel, 1894d: 5; Emery, 1896d: 373; Emery, 1925b: 120; Bolton, 1995b: 103; Radchenko, 2016: 333.
 * kosswigi. Camponotus (Orthonotomyrmex) kosswigi Donisthorpe, 1950c: 68 (w.) TURKEY.
 * [Species described but name of species not printed: Donisthorpe, 1950b: 61.]
 * Status as species: Donisthorpe, 1950e: 1067; Bolton, 1995b: 107.
 * Junior synonym of lateralis: Radchenko, 1997b: 706; Kiran & Karaman, 2012: 7; Radchenko, 2016: 333; Seifert, 2019b: 18.
 * melanogastes. Formica melanogastes Latreille, 1802c: 171, pl. 7, fig. 39 (w.) FRANCE.
 * Status as species: Schenck, 1852: 127.
 * Junior synonym of lateralis: Mayr, 1853b: 102; Mayr, 1855: 322; Nylander, 1856b: 58; Smith, F. 1858b: 12; Mayr, 1863: 399; Roger, 1863b: 1; André, 1874: 201 (in list); Forel, 1874: 96 (in list); Emery & Forel, 1879: 448; Dalla Torre, 1893: 237; Emery, 1896d: 373; Forel, 1915d: 71 (in key); Soudek, 1922: 90; Emery, 1925b: 120; Pisarski, 1975: 32; Bolton, 1995b: 111; Casevitz-Weulersse & Galkowsky, 2009: 479; Radchenko, 2016: 333; Seifert, 2019b: 17.
 * merula. Formica merula Losana, 1834: 313, pl. 36, fig. 2 (w.) ITALY.
 * [Misspelled as morula by Soudek, 1922: 91.]
 * Bondroit, 1918: 77 (s.q.m.).
 * Combination in Camponotus: Dalla Torre, 1893: 238;
 * combination in C. (Orthonotomyrmex): Müller, 1923b: 162;
 * combination in C. (Myrmentoma): Emery, 1925b: 120;
 * combination in Orthonotomyrmex: Novák & Sadil, 1941: 109 (in key).
 * Subspecies of lateralis: Emery, 1896d: 373 (in list); Emery, 1898c: 125; Ruzsky, 1902d: 7; Ruzsky, 1905b: 255; Karavaiev, 1912b: 596; Emery, 1914d: 159; Emery, 1916b: 226; Menozzi, 1918: 87; Santschi, 1919e: 247; Ruzsky, 1925b: 42.
 * Subspecies of piceus: Karavaiev, 1927a: 295.
 * Status as species: Mayr, 1855: 371 (footnote); Bondroit, 1918: 77; Müller, 1923a: 73; Müller, 1923b: 162; Ruzsky, 1926: 108; Cori & Finzi, 1931: 240; Soudek, 1931: 18; Novák & Sadil, 1941: 109 (in key); Bernard, 1959: 351; Bernard, 1967: 344 (redescription); Collingwood & Yarrow, 1969: 84; Arnol'di & Dlussky, 1978: 552.
 * Synonym of piceus: Emery, 1896d: 373 (in list); Ruzsky, 1905b: 255; Bernard, 1967: 344; Collingwood, 1978: 73.
 * [Note: Emery, Ruzsky, Bernard, and Collingwood give merula as senior synonym, but piceus has priority.]
 * Junior synonym of piceus: Dalla Torre, 1893: 238; Soudek, 1922: 91; Emery, 1925a: 70; Finzi, 1927b: 52; Kuznetsov-Ugamsky, 1929b: 36; Baroni Urbani, 1964b: 58; Baroni Urbani, 1971c: 194; Collingwood, 1971: 164; Kutter, 1977c: 207; Atanassov & Dlussky, 1992: 222; Arakelian, 1994: 87; Bolton, 1995b: 111; Radchenko, 1997b: 707; Czechowski, et al. 2002: 99; Radchenko, 2007: 37; Werner & Wiezik, 2007: 143; Gratiashvili & Barjadze, 2008: 131; Casevitz-Weulersse & Galkowsky, 2009: 480; Legakis, 2011: 32; Radchenko, 2016: 334.
 * Junior synonym of lateralis: Seifert, 2019b: 17.
 * pallidinervis. Formica pallidinervis Brullé, 1833: 326 (m.) GREECE.
 * Combination in Camponotus: Roger, 1863b: 1.
 * Junior synonym of aethiops: Roger, 1863b: 1.
 * Junior synonym of lateralis: Roger, 1861b: 163; Mayr, 1863: 399; Emery & Forel, 1879: 448; Dalla Torre, 1893: 203 (footnote), 237; Emery, 1925b: 120; Bolton, 1995b: 116; Radchenko, 2016: 333.
 * purius. Camponotus (Myrmentoma) lateralis var. purius Santschi, 1929e: 163 (w.q.m.) ALGERIA.
 * Subspecies of lateralis: Bolton, 1995b: 119.
 * Junior synonym of lateralis: Seifert, 2019b: 17.

Taxonomic Notes
Camponotus armouri: Examined was one worker syntype from, labeled ‘Barranco de San Juan, Minorca 8-24 – 25 W.M. Wheeler’, ‘M.C.Z. CoType 1-3 21538’. It is a typical Camponotus lateralis morph 1 and is allocated to this cluster with p = 1.000 if run as wild-card in a discriminant analysis considering all investigated characters and all entities given in Tab. 2 (Seifert, 2019).

Formica axillaris: According to the shape characters reported, this name belongs to the C. lateralis group. Furthermore, Spinola’s statements on color (‘Caput rubrum...Thorax niger... Abdomen nigrum’) allow the conclusion that it may belong to the color morph of C. lateralis mimicking Crematogaster scutellaris (Seifert, 2019).

Camponotus balearis: Investigated were three syntype workers from, labeled ‘Type’, ‘Baleares Ekker’, and ‘Camponotus (Myrmentoma) lateralis Ol v. balearis Sant’. All types are typical Camponotus lateralis morph 1 and each is allocated to this cluster with p=1.000 if run as wild-card in discriminant analysis considering all investigated characters and all entities given in Tab. 2 (Seifert, 2019).

Formica bicolor: The full text of the original description states ‘Rouge, luisante, rase. Yeux et abdomen noirs. Écaille épaisse, entière.’ Considering that the terra typica is France, this allows only to conclude on an ant of the subfamily Formicinae (Seifert, 2019).

Formica hemipsila: This taxon was described on the basis of a gyne from Alger/Algeria. Types most certainly do not exist. The description ‘Kopf...Mittelleib roth...nur Mittelbrust und Schildchen ganz schwarz’ may suggest a Camponotus related to the C. lateralis group (Seifert, 2019).

Camponotus kosswigi: The lecto- and paratype, stored in, were collected in Erbeyli / W Turkey, 14.V.1947. Pictures of syntypes in antweb.org under CASENT0903594 (major worker) and CASENT0903595 (minor worker) strongly suggest a synonymy with Camponotus lateralis morph 1 based on diagnostic setae and shape characters. A synonymy with Camponotus lateralis morph 2 and Camponotus rebeccae is excluded by the large scape base index SCI which is about 1.23 in the specimen with CW = 1.46 mm and 1.37 in the specimen with CW = 0.96 mm (Seifert, 2019).

Formica melanogastes: The shape and pigmentation characters reported suggest that it may belong to the color morph of C. lateralis mimicking Crematogaster scutellaris (Seifert, 2019).

Formica merula: It appears unlikely that a type specimen of Camponotus merula could ever be identified in a collection as the original description gives no type locality and its text and figure do not allow a reasonably safe allocation to a certain species. A total body length of 6 mm with a slender overall body shape and the ‘bilobe’ mesosoma (‘... bilobo, col lobo anteriore piu grande subrotonde...’) suggests a member of the Camponotus lateralis group. The chestnut-brown color (‘colore castagno intenso’) and occurrence on poplar trees plagued by aphids (‘..d‘onde percorre specialmente i pioppi dagli afidi travagliati.’) point to C. lateralis and make a synonymization with Camponotus piceus unlikely. As nobody is currently able to present counter-evidence, I synonymize C. merula with C. lateralis (Seifert, 2019).

Formica pallidinervis: This taxon has been described based on a male from Peloponnesus in Greece. The whereabouts of the type are unknown and the original description is insufficient – the more as males are generally poorly studied. The only possible conclusions seems to be that it is an ant of the subfamily Formicinae (Seifert, 2019).

Camponotus lateralis purius: Investigated were two worker syntypes from Algeria, stored in, mounted on the same pin and labeled ‘Camponotus lateralis Ol. / v. purius Sant / SANTSCI det. 1928’, ‘Alger Bequaert’, ‘Type’, ‘CASENT 09111698’ [major worker], ‘ANTWEB CASENT 0911699’ [minor worker]. Considering all characters and running the syntypes as wild-cards in a 4-class LDA considering the four species presented in Tab. 2, they are allocated to the C. lateralis cluster with posterior probabilities of 0.9998 and 0.9974. A wild-card run of the syntypes within the C. lateralis cluster, allocated the major to morph 1 with p=0.998 but the minor to morph 2 with p = 0.986 (Seifert, 2019).

Worker
Ionescu-Hirsch (2009) - TL = 3.9–7.0, HL = 0.98–1.80, HW = 0.84–1.68, EL = 0.26–0.37, SL = 1.02–1.29, ML = 1.46–2.23, PW = 0.64–1.07, mTbL = 0.70–1.04, hTbL = 0.96–1.45 (n = 5).

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