Messor structor

Unicoloniality and lack of swarming flights reported from Retz, Austria (Schlick-Steiner et al., 2005). In Greece, Messor structor appears to be mountain species, it was collected in mountain habitats between 535 and 1340 m a.s.l., mostly pastures, often close to coniferous forests, only one locality was in shadow mountain valley, on roadsides close to deciduous forest (Salata & Borowiec, 2019); in Achaia it was observed only on alpine pastures (Borowiec & Salata, 2021).

Identification
Steiner et al. (2018) - Worker and queen. Mostly lighter than Messor ponticus and Messor muticus. Discrimination from Messor ibericus, M. ponticus, and Messor mcarthuri by long 1st funicular segment, by base of scape with rounded lobe, and by coarser sculpture of mesopleuron, from M. ponticus by abundant standing setae on side of head. Head more irregularly costate than in M. mcarthuri. Surface of 1st gastral tergite less imbricate than in M. ibericus and M. mcarthuri, similarly to M. ponticus and M. muticus.

Worker. Similarly sized as M. ibericus and M. muticus, smaller than M. ponticus and M. mcarthuri. For individuals difficult to discriminate from M. ibericus, M. ponticus, or M. muticus using qualitative morphology, linear discriminant functions based on morphometrics is available.

Queen. Larger than M. ibericus, similarly sized as the other three species. Discrimination from M. ibericus by reduced reticulation and rugosity of anepisternum and katepisternum.

Distribution based on Regional Taxon Lists
Palaearctic Region: Austria, Bulgaria, Czech Republic, France, Hungary, Romania, Slovenia.

Habitat
A range of xeric habitats and, in some areas, open well drained anthropogenic habitats (lawns, abandoned fields, etc.).

Biology
Stoyanov et al. (2015) - Messor structor are harvester ants storing seeds in underground granaries. Being major seed consumers in xeric habitats they influence the vegetation by depleting and dispersing seeds (Azcarate, Peco 2003, MacMahon et al. 2000, Schlick-Steiner et al. 2005). According to Seifert (1996), Central European populations of the harvester ant genus Messor are currently considered to constitute a single species Messor structor. It has been reported that M. structor ants live mainly in polygynous colonies and could be members of unicolonial populations (Schlick-Steiner et al. 2005). Messor structor is one of the most widespread steppe species. It is well adapted to different conditions and occurs in all of Europe. In Bulgaria this ant is found all over the country in medium-sized populations and occur in plain desolate terrains, mountainous slopes, outskirts of mixed woodland zones and in open well-drained areas such as abandoned fields, pastures, and overgrown lawns.

This study examined genetic variability among populations of Messor structor from various localities (n=36 nests) in from Southern Bulgaria using isoenzymic analysis. Three of the four enzyme loci used were found to be polymorphic. Observed and expected heterozygosities (Ho and He) ranged from 0.0 to 0.111 and from 0.168 to 0.372, respectively. Nei’s (1972) genetic distance was found to range from 0.001 (between Boliarino and Lubenova mahala) to 0.462 (between Chirpan and Merichleri). The estimated mean F-IS and F-ST values from isozyme data were 0.8738 and 0.1432, respectively. Two of the studied populations were grouped separately in dendrograms as compared to all other populations which formed a large cluster consisting of three sub-clusters.

mtDNA. Thirty-one haplotypes of COI (1375 bp) known; (GenBank).

Nomenclature

 * . Formica structor Latreille, 1798: 46 (w.m.) FRANCE.
 * Type-material: neotype worker (by designation of Steiner, Csösz, et al. 2018: 399).
 * Type-locality: France: 300 m. E Gignac, S Brive-la-Gaillarde (45°0’N, 1°28’E), 1.x.2006, FR020 (Galkowski).
 * Type-depository: NHMW.
 * [Note: Steiner, Csösz, et al. 2018: 399, record that other specimens from the neotype nest-series are deposited in NHMW (5 workers), and in HNHM (4 workers).]
 * Latreille, 1802c: 237 (q.).
 * Combination in Manica: Jurine, 1807: 279;
 * combination in Atta: Latreille, 1809: 130; Lepeletier de Saint-Fargeau, 1835: 174; Mayr, 1855: 464; Smith, F. 1858b: 162;
 * combination in Myrmica: Nylander, 1856b: 85;
 * combination in Aphaenogaster: Roger, 1863b: 29;
 * combination in A. (Messor): Forel, 1892i: 316;
 * combination in Stenamma (Messor): Emery, 1898c: 126;
 * combination in Messor: Emery, 1897f: 238; Ruzsky, 1903b: 315; Emery, 1908e: 455.
 * Subspecies of barbarus: Emery, 1908e: 455; Forel, 1910a: 9; Karavaiev, 1910b: 67; Forel, 1911d: 346; Emery, 1912f: 96; Karavaiev, 1912a: 10; Stitz, 1914: 69; Emery, 1914d: 156; Emery, 1915h: 1; Wheeler, W.M. & Mann, 1916: 170; Emery, 1916b: 143; Santschi, 1917e: 91; Escherich, 1917: 327; Menozzi, 1918: 83; Santschi, 1921a: 110; Menozzi, 1921: 26; Emery, 1921b: 210; Emery, 1921f: 73; Soudek, 1922: 34; Finzi, 1923: 3; Emery, 1924b: 12; Ceballos, 1956: 301.
 * Status as species: Latreille, 1802c: 236; Jurine, 1807: 279; Latreille, 1809: 130; Lepeletier de Saint-Fargeau, 1835: 174; Schilling, 1839: 56; Brullé, 1840: 84; Schenck, 1852: 113; Mayr, 1855: 464 (redescription); Nylander, 1856b: 85; Gredler, 1858: 30; Smith, F. 1858b: 162; Roger, 1859: 253; Mayr, 1861: 66 (in key); Smith, F. 1861a: 35; Roger, 1863b: 29; Mayr, 1863: 397; Mayr, 1867a: 93 (redescription); Emery, 1869b: 17; Walker, 1871: 10; Dours, 1873: 167; Forel, 1874: 74 (in key); André, 1874: 195 (in key); Mayr, 1877: 14; Emery, 1878b: 58; Emery & Forel, 1879: 461; Mayr, 1880: 33; Emery, 1881b: 534; André, 1883b: 353 (in key); Forel, 1886e: clxviii; Forel, 1889: 257; Nasonov, 1889: 37; Saunders, E. 1890: 204; Forel, 1892i: 316; Dalla Torre, 1893: 105; Forel, 1895d: 227; Ruzsky, 1896: 74; Emery, 1898c: 140; Ruzsky, 1902a: 235; Ruzsky, 1902d: 27; Ruzsky, 1903b: 315; Forel, 1904b: 373; Ruzsky, 1905b: 728; Forel, 1906c: 190; Forel, 1907e: 15; Forel, 1909c: 104; Forel, 1915d: 35 (in key); Bondroit, 1918: 150; Kuznetsov-Ugamsky, 1923: 256; Müller, 1923b: 60; Finzi, 1924a: 13; Lomnicki, 1925b: 2; Wheeler, W.M. 1926: 2; Donisthorpe, 1926a: 7; Menozzi, 1926b: 181; Karavaiev, 1926b: 101; Karavaiev, 1927a: 286; Kuznetsov-Ugamsky, 1927a: 92 (in key); Karavaiev, 1927c: 261 (in key); Kutter, 1927: 99; Kuznetsov-Ugamsky, 1929d: 12; Finzi, 1929: 90; Karavaiev, 1929b: 209; Finzi, 1930d: 312; Cori & Finzi, 1931: 238; Santschi, 1931a: 4; Soudek, 1931: 8; Arnol'di, 1933b: 600 (in key); Karavaiev, 1934: 108; Grandi, 1935: 100; Zimmermann, 1935: 14; Finzi, 1936: 160; Finzi, 1939c: 155; Donisthorpe, 1950e: 1059; Consani & Zangheri, 1952: 40; Bernard, 1955a: 361 (in key); Bernard, 1956b: 258; Baroni Urbani, 1964a: 2; Baroni Urbani, 1964b: 41; Baroni Urbani, 1964c: 150; Bernard, 1967: 144 (redescription); Baroni Urbani, 1968b: 436; Collingwood & Yarrow, 1969: 64; Baroni Urbani, 1971c: 66; Baroni Urbani, 1974: 232; Collingwood, 1978: 81 (in key); Schembri & Collingwood, 1981: 427; Agosti & Collingwood, 1987a: 54; Agosti & Collingwood, 1987b: 271 (in key); Dlussky, Soyunov & Zabelin, 1990: 224; Casevitz-Weulersse, 1990b: 156 (in key); Le Moli & Rosi, 1991: 30; Arakelian, 1994: 38; Bolton, 1995b: 257; Mei, 1995: 762; Espadaler, 1997b: 30; Cagniant & Espadaler, 1998: 423; Gallé, et al. 1998: 214; Czechowski, et al. 2002: 37; Markó & Csösz, 2002: 116; Karaman, M.G. & Karaman, 2003: 43; Bračko, 2006: 137; Cagnaint, 2006a: 197; Markó, Sipos, et al. 2006: 69; Petrov, 2006: 92 (in key); Schultz, R. et al. 2006: 203; Bračko, 2007: 17; Seifert, 2007: 214; Werner & Wiezik, 2007: 140, 147; Zryanin & Zryanina, 2007: 231; Wetterer, et al. 2007: 14; Gratiashvili & Barjadze, 2008: 139; Paknia, et al. 2008: 155; Borowiec, M.L. et al. 2009: 378; Casevitz-Weulersse & Galkowski, 2009: 489; Vonshak, et al. 2009: 43; Boer, 2010: 61; Lapeva-Gjonova, et al. 2010: 15; Csösz, et al. 2011: 56; Karaman, M.G. 2011b: 38; Legakis, 2011: 11; Borowiec, L. & Salata, 2012: 515; Czechowski, et al. 2012: 119; Guénard & Dunn, 2012: 45; Kiran & Karaman, 2012: 20; Borowiec, L. 2014: 112; Lebas, et al. 2016: 298; Radchenko, 2016: 171; Salata & Borowiec, 2018c: 46; Seifert, 2018: 177; Steiner, Csösz, et al. 2018: 399 (redescription); Werner, et al. 2018: 7; Salata & Borowiec, 2019a: 64; Barech, et al. 2020: 17.
 * Senior synonym of aedificator: Mayr, 1855: 464; Smith, F. 1858b: 162; Mayr, 1863: 397; Roger, 1863b: 29; André, 1874: 203 (in list); Forel, 1874: 101 (in list); Emery & Forel, 1879: 462; Dalla Torre, 1893: 105; Emery, 1921f: 73; Bolton, 1995b: 257; Czechowski, et al. 2002: 37; Czechowski, et al. 2012: 120; Radchenko, 2016: 171; Steiner, Csösz, et al. 2018: supplementary material, 6.
 * Senior synonym of darianus: Dlussky, Soyunov & Zabelin, 1990: 224; Bolton, 1995b: 257; Radchenko, 2016: 171.
 * Senior synonym of jakowlevi: Dlussky, Soyunov & Zabelin, 1990: 224; Bolton, 1995b: 257; Radchenko, 2016: 171.
 * Senior synonym of lapidum: Nylander, 1856b: 85; Smith, F. 1858b: 162; Smith, F. 1861a: 35; Mayr, 1863: 397; Roger, 1863b: 29; Forel, 1874: 100 (in list); Emery & Forel, 1879: 462; Dalla Torre, 1893: 105; Ruzsky, 1905b: 728; Emery, 1921f: 73; Bolton, 1995b: 257; Radchenko, 2016: 171; Steiner, Csösz, et al. 2018: 399.
 * Senior synonym of romanus: Dlussky, Soyunov & Zabelin, 1990: 224; Bolton, 1995b: 257.
 * Senior synonym of rufitarsis Fabricius: Nylander, 1856b: 85; Smith, F. 1858b: 162; Mayr, 1863: 397; Roger, 1863b: 29; Forel, 1874: 101 (in list); Emery & Forel, 1879: 462; Nasonov, 1889: 37; Dalla Torre, 1893: 105; Ruzsky, 1905b: 728; Emery, 1921f: 73; Agosti & Collingwood, 1987a: 54; Casevitz-Weulersse, 1990b: 154; Dlussky, Soyunov & Zabelin, 1990: 224; Atanassov & Dlussky, 1992: 115; Arakelian, 1994: 39; Bolton, 1995b: 257; Gallé, et al. 1998: 214; Czechowski, et al. 2002: 37; Gratiashvili & Barjadze, 2008: 139; Casevitz-Weulersse & Galkowski, 2009: 489; Csösz, et al. 2011: 56; Legakis, 2011: 11; Czechowski, et al. 2012: 120; Kiran & Karaman, 2012: 20; Radchenko, 2016: 171; Steiner, Csösz, et al. 2018: 399.
 * Senior synonym of subpolitus: Dlussky, Soyunov & Zabelin, 1990: 224; Bolton, 1995b: 257; Radchenko, 2016: 171.
 * Senior synonym of turanicus: Dlussky, Soyunov & Zabelin, 1990: 224; Bolton, 1995b: 257; Radchenko, 2016: 171.
 * Distribution: Austria, Bulgaria, Czech Republic, France, Greece, Hungary, Romania, Slovenia, Ukraine.
 * [Note: central Asian records from Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, Uzbekistan are probably not conspecific.]
 * Current subspecies: nominal plus tyrrhenus.
 * aedificator. Formica aedificator Schilling, 1839: 56 (w.) POLAND.
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: Poland (no further data).
 * Type-depository: unknown.
 * [Note: according to Horn & Kahle, 1936: 243, all Schilling’s material went to an insect dealer in Breslau (= Wroclaw), in 1845; it has since disappeared.]
 * [Misspelled as aedificans by Mayr, 1855: 464, Smith, F. 1858b: 162.]
 * Combination in Aphaenogaster: Roger, 1863b: 29.
 * Junior synonym of rufitarsis Fabricius: Emery, 1921b: 211; Pisarski, 1975: 16.
 * Junior synonym of structor: Mayr, 1855: 464; Smith, F. 1858b: 162; Mayr, 1863: 397; Roger, 1863b: 29; André, 1874: 203 (in list); Forel, 1874: 101 (in list); Emery & Forel, 1879: 462; Dalla Torre, 1893: 105; Emery, 1921f: 73; Bolton, 1995b: 252; Czechowski, et al. 2002: 37; Czechowski, et al. 2012: 120; Radchenko, 2016: 171; Steiner, Csösz, et al. 2018: supplementary material, 6.
 * darianus. Messor rufitarsis subsp. darianus Pisarski, 1967: 383.
 * Type-material: 6 syntype workers.
 * Type-locality: Uzbekistan (“Turkestan russe”): Syr Daria, Bogus, 20.vi.1923 (N. Kusnezov).
 * Type-depository: NHMB.
 * [First available use of Messor structor st. rufitarsis var. darianus Santschi, 1926f: 291 (w.) UZBEKISTAN; unavailable (infrasubspecific) name.]
 * [Misspelled as dariagus by He & Song, 2009: 724.]
 * As unavailable (infrasubspecific) name: Santschi, 1927d: 247; Kuznetsov-Ugamsky, 1929d: 22.
 * Material referred to structor by Kuznetsov-Ugamsky, 1927a: 92 (in key).
 * Subspecies of rufitarsis Fabricius: Tarbinsky, 1976: 54; Dlussky, 1981a: 17; He & Song, 2009: 724.
 * Status as species: Schultz, R. et al. 2006: 203.
 * Synonym of turanicus: Santschi, 1927d: 247; Pisarski, 1967: 383; Pisarski, 1970: 306; Tarbinsky, 1976: 54; Arnol'di, 1977b: 1644 (in key).
 * [Note: Santschi, Pisarski, Tarbinsky, and Arnol’di, all give darianus as senior synonym, but turanicus has priority (Bolton, 1995b: 253).]
 * Junior synonym of structor: Dlussky, Soyunov & Zabelin, 1990: 224; Radchenko, 2016: 171.
 * jakowlevi. Messor rufitarsis subsp. jakowlevi Arnol'di, 1977b: 1643 (in key).
 * Type-material: syntype workers (number not stated).
 * Type-locality: Ukraine: Crimea, Yevpatoria (V. Yakovlev).
 * Type-depository: ZMUM.
 * [First available use of Aphaenogaster (Messor) barbarus subsp. capitatus var. jakowlevi Ruzsky, 1905b: 750 (w.) UKRAINE; unavailable (infrasubspecific) name.]
 * [Misspelled as jakovlevi by Arnol’di & Dlussky, 1978: 537 (in key).]
 * As unavailable (infrasubspecific) name: Emery, 1908e: 450; Emery, 1921f: 72; Emery, 1922c: 96; Karavaiev, 1927a: 287.
 * Subspecies of rufitarsis Fabricius: Arnol’di & Dlussky, 1978: 537 (in key); Steiner, Csösz, et al. 2018: supplementary material, 8 (nomen dubium).
 * [Note: because Steiner, Csösz, et al. 2018: 399, establish rufitarsis Fabricius as a junior synonym of structor, the synonymy of jakowlevi under structor still stands.]
 * Junior synonym of structor: Dlussky, Soyunov & Zabelin, 1990: 224; Atanassov & Dlussky, 1992: 114; Bolton, 1995b: 254.
 * Material of the unavailable name salina referred here by Arnol'di, 1977b: 1643 (in key).
 * lapidum. Formica lapidum Fabricius, 1804: 407 (w.) AUSTRIA.
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: Austria: (no further data).
 * Type-depository: ZMUK.
 * Combination in Aphaenogaster: Roger, 1863b: 29.
 * Junior synonym of rufitarsis Fabricius: Emery, 1921b: 211.
 * Junior synonym of structor: Nylander, 1856b: 85; Smith, F. 1858b: 162; Smith, F. 1861a: 35; Mayr, 1863: 397; Roger, 1863b: 29; Forel, 1874: 100 (in list); Emery & Forel, 1879: 462; Dalla Torre, 1893: 105; Ruzsky, 1905b: 728; Emery, 1921f: 73; Bolton, 1995b: 255; Radchenko, 2016: 171; Steiner, Csösz, et al. 2018: 399.
 * romanus. Messor structor var. romanus Santschi, 1926f: 286 (in list).
 * Type-material: syntype workers (number not stated), syntype queen.
 * Type-locality: Romania: Bucharest (Montandon).
 * Type-depository: NHMB.
 * [First available use of Messor barbarus st. structor var. romana Santschi, 1917e: 91 (w.q.) ROMANIA; unavailable (infrasubspecific) name (Agosti & Collingwood, 1987a: 61).]
 * As unavailable (infrasubspecific) name: Emery, 1921b: 211; Emery, 1921f: 73; Santschi, 1926f: 291 (in text); Arnol’di & Dlussky, 1978: 537 (in key); Legakis, 2011: 11 (error).
 * Junior synonym of structor: Dlussky, Soyunov & Zabelin, 1990: 224; Atanassov & Dlussky, 1992: 114; Bolton, 1995b: 256.
 * rufitarsis. Formica rufitarsis Fabricius, 1804: 406 (q.) AUSTRIA.
 * Type-material: holotype (?) queen.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: Austria: (no further data).
 * Type-depository: ZMUK (according to Zimsen, 1964: 425).
 * [Note: Steiner, Csösz, et al. 2018: 399, record 2 queens in ZMUC.]
 * Combination in Aphaenogaster: Roger, 1863b: 29;
 * combination in Messor: Emery, 1921b: 211.
 * Subspecies of barbarus: Emery, 1921b: 211.
 * Subspecies of structor: Santschi, 1926f: 290; Santschi, 1927d: 247; Menozzi, 1933b: 54; Santschi, 1934d: 275; Santschi, 1939c: 6; Pisarski, 1975: 16.
 * Status as species: Röszler, 1942a: 51; Röszler, 1942b: 32; Röszler, 1951: 88; Bernard, 1955a: 361 (in key); Bernard, 1956b: 258; Baroni Urbani, 1964b: 41; Pisarski, 1967: 383; Bernard, 1967: 144 (redescription); Kutter, 1968a: 59; Pisarski, 1970: 306; Arnol’di, 1970a: 79; Tarbinsky, 1976: 54; van Boven, 1977: 88; Baroni Urbani, 1971c: 63; Kutter, 1977c: 82; Arnol'di, 1977b: 1643 (in key); Arnol'di & Dlussky, 1978: 537 (in key); Bernard, 1981b: 143.
 * Junior synonym of structor: Nylander, 1856b: 85; Smith, F. 1858b: 162; Smith, F. 1861a: 35; Mayr, 1863: 397; Roger, 1863b: 29; Forel, 1874: 101 (in list); Emery & Forel, 1879: 462; Nasonov, 1889: 37; Dalla Torre, 1893: 105; Ruzsky, 1905b: 728; Emery, 1921f: 73; Agosti & Collingwood, 1987a: 54; Casevitz-Weulersse, 1990b: 154; Dlussky, Soyunov & Zabelin, 1990: 224; Atanassov & Dlussky, 1992: 115; Arakelian, 1994: 39; Bolton, 1995b: 256; Gallé, et al. 1998: 214; Czechowski, et al. 2002: 37; Gratiashvili & Barjadze, 2008: 139; Casevitz-Weulersse & Galkowski, 2009: 489; Csösz, et al. 2011: 56; Legakis, 2011: 11; Czechowski, et al. 2012: 120; Kiran & Karaman, 2012: 20; Radchenko, 2016: 171; Steiner, Csösz, et al. 2018: 399.
 * subpolitus. Messor structor var. subpolitus Kuznetsov-Ugamsky, 1927a: 92 (in key) (w.) UZBEKISTAN, KYRGYZSTAN.
 * Type-material: syntype workers (number not stated).
 * Type-localities: Uzbekistan: Tashkent (no collector’s name, perhaps Kuznetsov-Ugamsky), Kyrgyzstan: Karakol (no collector’s name, perhaps Kuznetsov-Ugamsky).
 * [Note: original entry states, “Rare. At present only known from Tashkent and Karakol”.]
 * Type-depository: ZMUM.
 * Subspecies of structor: Kuznetsov-Ugamsky, 1929d: 15.
 * Junior synonym of clivorum: Tarbinsky, 1976: 56; Bolton, 1995b: 257.
 * Junior synonym of structor: Dlussky, Soyunov & Zabelin, 1990: 224; Radchenko, 2016: 171.
 * turanicus. Messor structor subsp. turanicus Kuznetsov-Ugamsky, 1927a: 91 (in key) (w.) UZBEKISTAN, KAZAKHSTAN.
 * Type-material: syntype workers (number not stated).
 * Type-localities: Uzbekistan: Tashkent, Samarkand, Namangan (no collector’s name, perhaps Kuznetsov-Ugamsky), Kazakhstan: Tamerlanovka Dist., Shymkent, Tschulak-kurgan, Ibata, ‘Stadt Turkestan’ (no collector’s name, perhaps Kuznetsov-Ugamsky).
 * Type-depository: ZMUM.
 * Subspecies of structor: Kuznetsov-Ugamsky, 1929d: 14.
 * Synonym of darianus: Santschi, 1927d: 247; Pisarski, 1967: 383; Pisarski, 1970: 306; Tarbinsky, 1976: 54; Arnol'di, 1977b: 1644.
 * [Note: Santschi, Pisarski, Tarbinsky, and Arnol’di all give darianus as senior synonym, but turanicus has priority (Bolton, 1995b: 258).]
 * Junior synonym of structor: Dlussky, Soyunov & Zabelin, 1990: 224; Bolton, 1995b: 258; Radchenko, 2016: 171.

Taxonomic Notes
Salata & Borowiec (2019) (misidentification):
 * As Messor cf. clivorum: Borowiec and Salata, 2018a: 7.

Description
Steiner et al. (2018) - Worker and queen. Colour: Head and mesosoma brown to brownish red, gaster brown; major workers sometimes entirely dark brown. Size: Medium. Head: Mostly costate entirely. Parallel costae below eye level. Especially in major and medium worker, vertex or even entire surface above eye level with irregular costae and costulae with microreticulation amongst. In minor and sometimes in media worker, costae reduced in postocular region; head almost smooth and frequently shining, with only fine costulae and microreticulate. Erect setae abundant on side of head from occiput to mandibular insertion. Scape: Base with rounded, laterally-downward directed lobe, very distinct in minor worker; can be reduced in major worker. 1st funicular segment long and flattened, almost as long as 2nd and 3rd segment together. Clypeus: Median notch well defined; can be shallow in minor worker. Pronotum: Middle mostly coarsely and irregularly costate throughout. Mesonotum: Coarsely and irregularly costate throughout. Mesopleuron: Coarsely sculptured with irregular transverse costae, with shallow punctuation among costae. Minor workers less costate with almost no punctuation. Propodeum: Mostly smoothly rounded; often obliquely rounded or rarely angled in major workers. Surface of 1st gaster tergite: Basis imbricate, middle with isolated snow-flakes-like structure. Covered with yellowish, thin, sparse hairs, some of which decumbent and subdecumbent in some individuals.

Queen: Metanotum: coarsely costate. Anepisternum and katepisternum: Middle shining and smooth, side with longitudinal costulae.

Male: Anepisternum: Shining, but often microreticulate. Katepisternum: Microreticulate, side with few costulae.

Type Material


Steiner et al. (2018) - 9 additional workers from the same nest series and labelled as the neotype are deposited in NHMW (5 workers) and in (4 workers). If neotype is destroyed or lost, a replacement neotype can be designated from these 9 workers.

References based on Global Ant Biodiversity Informatics

 * Abdul-Rassoul M. S., H. A. Dawah, and N. Y. Othman. 1978. Records of insect collection (Part I) in the natural history research centre, Baghdad. Bull. Nat. Hist. Res. Centre 7(2): 1-6.
 * Abril S., C. Gómez. 2012. Lista actualizada de las especies de hormigas de Menorca (Islas Baleares, España) y primera cita de Monomorium andrei Saunders, 1890 (Hymenoptera, Formicidae) de la isla Boletín de la Sociedad Entomológica Aragonesa 50: 403-407.
 * Acosta, F. J., M. D. Martínez Ibáñez, and M. A. Morales. "Contribución al conocimiento de la mirmecofauna del encinar peninsular. I." Boletín de la Asociación Española de Entomología 6 (2) (1983): 379-391.
 * Agosti, D. and C.A. Collingwood. 1987. A provisional list of the Balkan ants (Hym. Formicidae) and a key to the worker caste. I. Synonymic list. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 60: 51-62
 * Aldawood AS, Sharaf MR (2011) Monomorium dryhimi sp. n., a new ant species (Hymenoptera, Formicidae) of the M. monomorium group from Saudi Arabia, with a key to the Arabian Monomorium monomorium-group. ZooKeys 106: 4754. doi: 10.3897/zookeys.106.139
 * Andoni V. 1977. Kontribut mbi Himenopteret e familjes Formicidae te vendit tone. Buletini I Shkencave te Natyres 31(2): 93-101.
 * AntArea. Accessed on February 5th 2014 at http://antarea.fr/fourmi/
 * Antarea (Personal Communication - Rumsais Blatrix- 27 April 2018)
 * Antarea (at www.antarea.fr on June 11th 2017)
 * Antonova V., and L. Penev. 2008. Classification of assemblages of ants in the green areas in Sofia City. Acta Zoologica Bulgarica 60(2): 103-110.
 * Arakelian G. R., S. O. Movsessian, and F. A. Chubarian. 1997. Ecological and faunistic review of ants (Hymenoptera: Formicidae) - Supplementary hosts of the trematode Dicrocoelium lanceatum in Armenia. Parasitologia 31(3): 239-244.
 * Arnan, X., A. Rodrigo and J. Retana. 2006. Post-fire recovery of Mediterranean ground ant communities follows vegetation and dryness gradients. Journal of Biogeography 33(7):1246-1258
 * Asociacion Iberica de Mirmecologia. 2011. List of species collected during the Taxomara Lisboa 2010. Iberomyrmex 3: 32-33.
 * Asociacion Iberica de Mirmecologia. 2011. List of species collected during the Taxomara Lisboa 2011. Iberomyrmex 3: 30-31.
 * Asociacion Iberica de Mirmecologia. 2015. List of species collected during the Taxomara Lisboa 2015 Iberomyrmex 7: 44-45.
 * Baroni Urbani C. 1968. Studi sulla mirmecofauna d'Italia. IV. La fauna mirmecologica delle isole Maltesi ed il suo significato ecologico e biogeografico. Ann. Mus. Civ. Stor. Nat. Giacomo Doria 77: 408-559.
 * Baroni Urbani C. 1974. Studi sulla mirmecofauna d'Italia. XII. Le Isole Pontine. Fragm. Entomol. 9: 225-252.
 * Baroni Urbani, C.. "Studi sulla mirmecofauna d'Italia. II. Formiche di Sicilia." Atti dell'Accademia Gioenia di Scienze Naturali in Catania (6) 16 (1964): 25-66.
 * Barrett K. E. J. 1970. Ants in France, 1968-69. Entomologist 103: 270-274.
 * Bernadou A., V. Fourcassié, and X. Espadaler. 2013. A preliminary checklist of the ants (Hymenoptera, Formicidae) of Andorra. Zookeys 277: 13-23.
 * Bernard F. 1967. Faune de l'Europe et du Bassin Méditerranéen. 3. Les fourmis (Hymenoptera Formicidae) d'Europe occidentale et septentrionale. Paris: Masson, 411 pp.
 * Bernard F. 1967. Recherches sur les fourmis des Monts-Dore. Annales de la Station biologique de Besse-en-Chandesse : 1-11.
 * Bernard F. 1973. Tendances calcicoles ou silicicoles chez les fourmis méditerranéennes. Pp. 16-21 in: International Union for the Study of Social Insects. Congress 1973. Proceedings IUSSI VIIth International Congress, London, 10-15 September, 1973. Southampton: University of Southampton, vi + 418 pp.
 * Bernard F. 1981. Revision of the genus Messor (harvesting ants) on a biometrical basis. Pp. 141-145 in: Howse, P. E., Clement, J.-L. (eds.) Biosystematics of social insects. Systematics Association Special Volume No. 19. London: Academic Press, 346 pp.
 * Bernard, F.. "Remarques sur le peuplement des Baléares en fourmis." Bulletin de la Société d' Histoire naturelle de l' Afrique du Nord 47 (1956): 254-266.
 * Berville L., C. Santelli, J. Reybaud, M. Renucci, P. Ponel, O. Blight, and E. Provost. 2014. Suivi d’un site atelier dans le golfe de Fos: Une diversite myrmecologique insoupconne. Etudes Vauclusiennes 82: 71-78.
 * Berville L., M. Renucci, P. Vidal, and E. Provost. 2016. Peuplement myrmecologique et evaluation de l'invasion de Linepithema humile sur les iles de Marseille (Bouches-du-Rhone, France). Revue d'Ecologie (Terre et Vie) 71(3): 278-287.
 * Bonaric J. C. 1971. Contribution a l'etude systematique et ecologique des formicides du Bas-Languedoc. PhD thesis Universite des sciences et techniques du Languedoc, 175 pages.
 * Bonaric J. C. 1971. Étude systématique et écologique des fourmis de lHérault. Ann. Soc. Hortic. Hist. Nat. Hérault 111: 81-87.
 * Borowiec L. 2014. Catalogue of ants of Europe, the Mediterranean Basin and adjacent regions (Hymenoptera: Formicidae). Genus (Wroclaw) 25(1-2): 1-340.
 * Borowiec L., and S. Salata. 2012. Ants of Greece - Checklist, comments and new faunistic data (Hymenoptera: Formicidae). Genus 23(4): 461-563.
 * Borowiec L., and S. Salata. 2015. Pheidole symbioticaWasmann, 1909, an enigmatic supposed social parasite, is a nematodeinfested form of Pheidole pallidula(Nylander, 1849) (Hymenoptera: Formicidae: Myrmicinae). Sociobiology 62(2): 181-186.
 * Bracko G. 2007. Checklist of the ants of Slovenia (Hymenoptera: Formicidae). Natura Sloveniae 9: 15-24
 * Bracko, G. 2006. Review of the ant fauna (Hymenoptera:Formicidae) of Croatia. Acta Entomologica Slovenica 14(2): 131-156.
 * Bracko, G.. "Review of the ant fauna (Hymenoptera: Formicidae) of Croatia." Acta Entomologica Slovenica Vol 14 st (2006): 131-156.
 * Cagniant, H and X. Espadaler, X. 1998 [1997]. Le genre Messor au Maroc (Hymenoptera: Formicidae). Ann. Soc. Entomol. Fr. (n.s.) 33: 419-434
 * Cagniant, H. 2006. Liste actualisée des fourmis du Maroc. Myrmecologische Nachrichten 8: 193-200.
 * Cagniant, H., and X. Espadaler. "Le genre Messor au Maroc." Annales de la Société Entomologique de France (N.S.) 33 (4) (1998): 419-434.
 * Cagniant, H.. "Liste actualisee des fourmis du Maroc (Hymenoptera: Formicidae)." Myrmecologische Nachrichten 8 (2006): 193-200.
 * Casevitz-Weulersse J. 1990. Etude Systematique de la Myrmecofaune Corse (Hymenoptera: Formicidae), deuxieme partie. Bull. Mus. Natn. Hist. Nat. Paris. 4eme serie 12, section A(2): 415-442.
 * Casevitz-Weulersse J. 1990. Étude systématique de la myrmécofaune corse (Hymenoptera, Formicidae) (Première partie). Bull. Mus. Natl. Hist. Nat. Sect. A Zool. Biol. Écol. Anim. (4) 12: 135-163.
 * Casevitz-Weulersse J., and C. Galkowski. 2009. Liste actualisee des Fourmis de France (Hymenoptera, Formicidae). Bull. Soc. Entomol. Fr. 114: 475-510.
 * Casevitz-Weulersse J., and M. Prost. 1991. Fourmis de la Côte-d'Or présentes dans les collections du Muséum d'Histoire Naturelle de Dijon. Bulletin Scientifique de Bourgogne 44: 53-72.
 * Casevitz-Weulersse, J.. "Contribution a la connaisance des fourmis de la Corse (Hymenoptera: Formicidae)." These de Doctorat Museum Nat (1989): 379pp.
 * Castracani C., D. A. Grasso, A. Fanfani, and A. Mori. 2010. The ant fauna of Castelporziano Presidential Reserve (Rome, Italy) as a model for the analysis of ant community structure in relation to environmental variation in Mediterranean ecosystems. J Insect Conserv 14: 585594.
 * Ceballos, G.. "Catálogo de los Himenópteros de España.", 295-321. Vol. Catálogo de los Himenópt. Madrid: Instituto Español de Entomologia, 1956.
 * Collingwood C. A. 1956. Ant hunting in France. Entomologist 89: 106-108.
 * Collingwood C., and A. Prince. 1998. A guide to ants of continental Portugal (Hymenoptera: Formicidae). Boletim da Sociedade Portuguesa de Entomologia. Suplemento 5: 1-49.
 * Collingwood, C. A., and I. H. H. Yarrow. "A survey of Iberian Formicidae." EOS (Revista española de entomología) 44 (1969): 53-101.
 * Comín del Río, P., and Andrés de Haro. "Datos iniciales para un estudio ecológico de las hormigas de Menorca." Bolletí de la Societat d´Historia Natural de les Balears 24 (1980): 23-48.
 * Comín del Río, P., and X. Espadaler. "Ants of the Pityusic islands." In Biogeography of the Pityusic Islands, edited by H. Kuhbler, J. A. Alcover and C. Guerau, 278-231. The Hague: Junk, 1984.
 * Comín del Río, P.. "Los Formícidos de Menorca. Contribución al estudio taxonómico, geográfico y biológico." Tesina de licenciatura Universida (1977): 135 pp.
 * Comín del Río, P.. Estudio de los formícidos de Baleares: Contribución al estudio taxonómico, geográfìco y biológico. Palma de Mallorca: Universidad de las Islas Baleares, 1988.
 * Consani M., and P. Zangheri. 1952. Fauna di Romagna. Imenotteri - Formicidi. Memorie della Societa Entomologica Italiana 31: 38-48.
 * Csosz S., B. Marko, K. Kiss, A. Tartally, and L. Galle. 2002. The ant fauna of the Ferto-Hansag National Park (Hymenoptera: Formicidae). In: Mahunka, S. (Ed.): The fauna of the Fert?-Hanság National Park. Hungarian Natural History Museum, Budapest, pp. 617-629.
 * Csősz S., B. Markó, and L. Gallé. 2001. Ants (Hymenoptera: Formicidae) of Stana Valley (Romania): Evaluation of the effectiveness of a myrmecological survey.  Entomologica Romanica 6 : 121-126.
 * Csősz S., B. Markó, and L. Gallé. 2011. The myrmecofauna (Hymenoptera: Formicidae) of Hungary: an updated checklist. North-Western Journal of Zoology 7: 55-62.
 * Czechowski W., A. Radchenko, W. Czechowska and K. Vepsäläinen. 2012. The ants of Poland with reference to the myrmecofauna of Europe. Fauna Poloniae 4. Warsaw: Natura Optima Dux Foundation, 1-496 pp
 * Della Santa E. 1994. Guide pour l'identification des principales espèces de fourmis de Suisse. Miscellanea Faunistica Helvetiae 3: 1-124.
 * Della Santa E. 1995. Fourmis de Provence. Faune Provence 16: 5-37.
 * Donisthorpe, H.. "A first instalment of the ants of Turkey." Annals and Magazine of Natural History (12)3 (1950): 1057-1067.
 * Du Merle P. 1978. Les peuplements de fourmis et les peuplements d'acridiens du Mont Ventoux II. - Les peuplements de fourmis. Terre Vie 32(1): 161-218.
 * Dubovikoff D. A., and Z. M. Yusupov. 2018. Family Formicidae - Ants. In Belokobylskij S. A. and A. S. Lelej: Annotated catalogue of the Hymenoptera of Russia. Proceedingss of the Zoological Institute of the Russian Academy of Sciences 6: 197-210.
 * Dvorak, L., P. BOGUSCH, I. MALENOVSKÝ, P. BEZDÌÈKA, K. BEZDÌÈKOVÁ, K. HOLÝ, P. LIKA, J. MACEK, L. ROLLER, M. RÍHA et al. "Hymenoptera of Hády Hill, near the city of Brno (Czech Republic), collected during the Third Czech-Slovak Hymenoptera meeting." Acta Musei Moraviae, Scientiae biologicae (Brno) 93 (2008): 53-92.
 * Eidmann, H.. "Die Ameisenfauna der Balearen." Zeitschrift für Morphologie und Ökologie der Tiere 6 (1926): 694-742.
 * Eidmann, H.. "Zur Kenntnis der Insektekfauna der Balearischen Inseln." Entomologische Mitteilungen 16 (1927): 24-37.
 * Emery C. 1914. Wissenschaftliche Ergebnisse der Bearbeitung von O. Leonhard's Sammlungen. 5. Südeuropäische Ameisen (Hym.). Entomologische Mitteilungen. Berlin-Dahlem 3: 156-159.
 * Emery, C.. "Alcune formiche dell'isola di Creta." Bull. Soc. Entomol. Ital. Resoc. Adun. 26 (1894): 7-10.
 * Emery, C.. "Alcune formiche di Malta." Bollettino della Società Entomologica Italiana 56 (1924): 11-12.
 * Emery, C.. "Beiträge zur Kenntniss der palaearktischen Ameisen." Öfversigt af Finska Vetenskaps-Societetens Förhandlingar (Helsinki) 20 (1898): 124-151.
 * Emery, C.. "Beiträge zur Monographie der Formiciden des paläarktischen Faunengebietes. (Hym.) Teil III. Messor, Goniomma, Oxyopomyrmex." Deutsche Entomologische Zeitschrift 1908 (1908): 437-465.
 * Emery, C.. "Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte seconda. Formiche dell'Europa e delle regioni limitrofe in Africa e in Asia." Annali del Museo Civico di Storia Naturale 12 (1878): 43-59.
 * Emery, C.. "Escursioni zoologiche del Dr. Enrico Festa nell'Isola di Rodi. XII. Formiche." Bollettino del Museo di Zoologia ed Anatomia Comparatadella Reale Università di Torino 30 (1915): 1-7.
 * Emery, C.. "Fauna entomologica italiana. I. Hymenoptera.-Formicidae." Bollettino della Societa Entomologica Italiana 47 (1916): 79-275.
 * Emery, C.. "Viaggio ad Assab nel Mar Rosso dei Signori G. Doria ed O. Beccari con il R. Avviso "Esploratore" dal 16 novembre 1879 al 26 febbraio 1880. I. Formiche." Annali del Museo Civico di Storia Naturale 16 (1881): 525-535.
 * Espadaler X., X. Roig, K. Gómez, and F. García. 2011. Formigues de les Planes de Son i mata de València (Hymenoptera, Formicidae) Treballs de la Institució Catalana d'Història Natural 16: 609-627.
 * Espadaler, X., and L. Lopez Soria. "Rareness of certain Mediterranean ant species: fact or artifact?" Insectes Sociaux 38 (1991): 365-377.
 * Espadaler, X.. "Catalogos entomofauna aragonesa: Familia: Formicidae." Catalogos entomofauna aragonesa 13 (1997): 13-21.
 * Finzi B. 1929. Le forme italiane del genere Messor. Bollettino della Società Entomologica Italiana 61: 75-94.
 * Finzi, B.. "Formiche dell'isola d'Elba e Monte Argentario." Bollettino della Società Entomologica Italiana 56 (1924): 12-15.
 * Finzi, B.. "Risultati scientifici della spedizione Ravasini-Lona in Albania. III. Formiche." Bollettino della Società Entomologica Italiana 55 (1923): 1-4.
 * Forel A. 1892. Die Ameisenfauna Bulgariens. (Nebst biologischen Beobachtungen.). 305-318.
 * Forel A. 1904. Note sur les fourmis du Musée Zoologique de l'Académie Impériale des Sciences à St. Pétersbourg. Ezheg. Zool. Muz. 8: 368-388.
 * Forel A. 1906. Fourmis d'Asie mineure et de la Dobrudscha récoltées par M. le Dr. Oscar Vogt et Mme Cécile Vogt, Dr. méd. Ann. Soc. Entomol. Belg. 50: 187-190.
 * Forel A. 1907. Formiciden aus dem Naturhistorischen Museum in Hamburg. II. Teil. Neueingänge seit 1900. Mitt. Naturhist. Mus. Hambg. 24: 1-20.
 * Forel A. 1909. Études myrmécologiques en 1909. Fourmis de Barbarie et de Ceylan. Nidification des Polyrhachis. Bull. Soc. Vaudoise Sci. Nat. 45: 369-407.
 * Forel A. 1910. Glanures myrmécologiques. Ann. Soc. Entomol. Belg. 54: 6-32.
 * Forel A. 1911. Fourmis nouvelles ou intéressantes. Bull. Soc. Vaudoise Sci. Nat. 47: 331-400.
 * Forel, A.. "Ameisen aus den Sporaden, den Cykladen und Griechenland, gesammelt 1887 von Herrn von Oertzen." Berliner Entomologische Zeitschrift 32 (1889): 255-265.
 * Forel, A.. "Fourmis d'Espagne récoltées par M. O. Vogt et Mme Cécile Vogt, Docteurs en médecine." Annales de la Société Entomologique de Belgique 53 (1909): 103-106.
 * Forel, A.. "Nouvelles fourmis de Grèce récoltées par M. E. von Oertzen." Annales de la Société Entomologique de Belgique 30 (1886): clix-clxviii.
 * Forel, A.. "Südpalaearctische Ameisen." Mitteilungen der Schweizerischen Entomologischen Gessellschaft 9 (1895): 227-234.
 * Francois J. 1958. Contribution a l'etude ecologique des Formicides (Insectes, Hymenopteres) de la region Dijonnaise. Travaux du laboratoire de Zoologie et de la Station Aquicole Grimaldi de la Faculte des Sciences de Dijon 25, 39 pages.
 * GRETIA. 2017. Bilan annuel de l'enquete sur la repartition des fourmis armoricaines. 23 pages.
 * Galkowski C. 2011. Une liste des fourmis (Hymenoptera, Formicidae) récoltées dans la région de Grasse, avec la mention dune nouvelle espèce de la faune de France. Bulletin de la Société linnéenne de Provence, 62 : 41-44.
 * Galle L. 1972. Study of ant-populations in various grassland ecosystems. Acta Biologica Szeged 18(1-4): 159-164.
 * Galle L. 1981. The Formicoid fauna of the Hortobagy. Pp. 307-311 in: Mahunka, S. (ed.) 1981. The fauna of the Hortobágy National Park. Budapest: Akadémiai Kiadó, 415 pp.
 * Gallé L., B. Markó, K. Kiss, E. Kovács, H. Dürgő, K. Kőváry, and S. Csősz. 2005. Ant fauna of Tisza river basin (Hymenoptera: Formicidae).  In: Gallé, L. (szerk.): Vegetation and Fauna of Tisza River Basin I. Tiscia Monograph Series 7; Szeged, pp. 149-197.
 * Garcia Garcia F., and A. D. Cuesta-Esgura. 2017. First catalogue of the ants of Burgos province, Spain (Hymenoptera: Formicidae). Boletín de la Sociedad Entomológica Aragonesa 60: 245–258.
 * Gaspar C. 1968. Les fourmis de la Drome et des Basses-Alpes, en France (Hymenoptera, Formicidae). Naturaliste can. 95: 747-766.
 * Goetsch, W.. "Beiträge zur Biologie spanischer Ameisen." EOS (Revista española de entomología) 18 (1942): 175-241.
 * Gomez C. 2013. Ant species of the Tierra de pinares (Castilla y Leon, Espana) and potential recolonization sources for logged sites. Iberomyrmex 5 : 11-14.
 * Gomez K. 2017. Two species of exotic ants (Hymenoptera: Formicidae) new to Malta. Boletin de la Sociedad Entomologica Aragonesa (S.E.A.) 61: 233-235.
 * González Martín J. D., and X. Espadaler. 2011. Formicidos del País Vasco (Hymenoptera: Formicidae): Nuevas aportaciones. Heteropterus Revista de Entomologia 11(1): 109-122.
 * Gouraud C. 2015. Bilan de l’année 2014 : Atlas des fourmis de Loire-Atlantique (Hymenoptera, Formicidae). Atlas des Formicidae de Loire-Atlantique, compte rendu de la première année d’étude (2014)
 * Grandi G. 1935. Contributi alla conoscenza degli Imenotteri Aculeati. XV. Boll. R. Ist. Entomol. Univ. Studi Bologna 8: 27-121.
 * Gratiashvili N., Barjadze S. 2008. Checklist of the ants (Formicidae Latreille, 1809) of Georgia. Proceedings of the Institute of Zoology (Tbilisi) 23: 130-146.
 * Groc S., J. H. C. Delabie, R. Cereghino, J. Orivel, F. Jaladeau, J. Grangier, C. S. F. Mariano, and A. Dejean. 2007. Ant species diversity in the Grands Causses (Aveyron, France): In search of sampling methods adapted to temperate climates. C. R. Biologies 330: 913922.
 * Guillem, R.. "A survey of the ants of Minorca (Hymenoptera: Formicidae) with two new species for the island: Hypoponera punctatissima (Roger, 1859) and Temnothorax algiricus (Forel, 1894)." Boletín de la Asociación Española de Entomología 33, no. 3-4 (2010): 447-460.
 * Guénard B., and R. R. Dunn. 2012. A checklist of the ants of China. Zootaxa 3558: 1-77.
 * Gómez C., and S. Abril. 2011. Selective logging in public pine forests of the central Iberian Peninsula: Effects of the recovery process on ant assemblages Forest Ecology and Management 262: 1061-1066.
 * Hamann H. H. F., and W. Klemm. 1976. Ergebnisse der von Dr. O. Paget und Dr. E. Kritscher auf Rhodos durchgeführten zoologischen Exkursionen. XVI. Formicidae. Annalen des Naturhistorischen Museums in Wien 80: 669-679.
 * He D.-H., and L. Song. 2009. A taxon of the ant genus Messor Forel (Hymenoptera: Formicidae) from China, with description of two new species in desert region. Highlights of Sciencepaper Online 2: 720-725.
 * Hernandez Cuba O., C. Perez Banon, and M. A. Marcos Garcia. 2008. Los formicidos (Hymenoptera, Formicidae) del Parque natural de la Font Roja. Iberis 6: 9-20.
 * Hosoishi, S., S. Yamane and K. Ogata. 2010. Subterranean species of the ant genus Crematogaster in Asia (Hymenoptera: Formicidae). Entomological Science 13:345-350.
 * Huang Jian-hua, Zhou Shan-yi. 2007. A checklist of family Formicidae of China - Myrmicinae (Part II) (Insecta: Hymenoptera). Journal of Guangxi Normal University : Natural Science Edition 25(1): 91-99.
 * Huang Ren-Xing, Ouyang Tong, Wu Wei, and Fan Zhao-tian. 2004. Forty two new record species of Family Formicidae (Hymenoptera: Formicoidea) from Xinjiang, China. Entomotaxonomia 26 (2): 156-160.
 * Karaman M. G. 2009. An introduction to the ant fauna of Macedonia (Balkan Peninsula), a check list (Hymenoptera, Formicidae). Natura Montenegrina 8(3): 151-162.
 * Karaman M. G. 2011. A catalogue of the ants (Hymenoptera, Formicidae) of Montenegro. Podgorica: Catalogues 3, Volume 2, Montenegrin Academy of Sciences and Arts, 140 pp.
 * Karavaiev V. 1912. Ameisen aus Tunesien und Algerien, nebst einigen unterwegs in Italien gesammelten Arten. Rus. Entomol. Obozr. 12: 1-22.
 * Karavaiev V. 1927. Übersicht der Ameisenfauna der Krim nebst einigen Neubeschreibungen. Konowia 5: 281-303.
 * Kiran K. C. Karaman, A. Lapeva-Gjonova, and V. Aksoy. 2017. Two new species of the "ultimate" parasitic ant genus Teleutomyrmex KUTTER, 1950 (Hymenoptera: Formicidae) from the Western Palaearctic. Myrmecological News 25: 145-155.
 * Kiran K., and C. Karaman. 2012. First annotated checklist of the ant fauna of Turkey (Hymenoptera: Formicidae). Zootaxa 3548: 1-38.
 * Kiran K., and N. Aktac. 2006. The vertical distribution of the ant fauna (Hymenoptera: Formicidae) of the Samanh Mountains, Turkey. Linzer Biol. Beitr. 38(2): 1105-1122.
 * Korlevic, A.. "Prilozi fauni hrvatskih opnokrilaca." Glasn. Hrv. Narav. Dr. 5 (1890): 189-250.
 * Krausse, A. H.. "Ueber sardische Ameisen." Archiv für Naturgeschichte, Berlin A 78(7) (1912): 162-166.
 * Kugler J. 1988. The zoogeography of Israel. 9. The zoogeography of social insects of Israel and Sinai. Monographiae biologicae 62: 251-275.
 * Kuznetsov G. T. 1990. Comparative analysis of Hymenoptera (Formicidae) population on altitudinal zones of central Kopetdag. Izvestiya Akademii Nauk Turkmenskoi SSR. Seriya Biologicheskikh Nauk 1990(3): 64-67.
 * Lapeva-Gjonova, L., V. Antonova, A. G. Radchenko, and M. Atanasova. "Catalogue of the ants (Hymenoptera, Formicidae) of Bulgaria." ZooKeys 62 (2010): 1-124.
 * Legakis A. 1983. The Zoological Museum of the University of Athens 2. The collection of ants from Greece. Biologia Gallo-Hellenica 11(1): 85-87.
 * Legakis A. 2001. Ants (Hymenoptera, Formicidae) collected in the eastern Pyrenees and surrounding area, September 1999. Proceedings of the International Colloqium of the European Invertebrate Survey: Cartography and Conservation of Biodiversity Marcevol Priory, Arboussols, france 30.8-4.9.1999. OPIE.
 * Legakis Collection Database
 * Lenoir A. 1971. Les fourmis de Touraine, leur intérêt biogéographique. Cahiers des Naturalistes 27: 21-30.
 * Lomnicki, J.. "Une contribution à la connaissance de la faune des fourmis des îles Baléares." Polskie Pismo Entomologiczne - Bulletin Entomologique de Pologne 4 (1925): 1-3.
 * Marikovsky P. I. 1979. Ants of the Semireche Desert. [In Russian.]. Alma Ata: Nauka, 263 pp.
 * Markó B. 1997. Contribution to the Knowledge of the Ant-Fauna (Hymenoptera: Fromicoidea) of the Cri?ul-Repede River-Valley. In: Sárkány-Kiss, A., Hamar, J. (szerk.): The Cri?/Körös Rivers' Valleys. A study of the geography, hydrobiology and ecology of the river system and its environment. Tiscia Monograph Series 2, Szolnok-Szeged-Tîrgu Mure?, pp. 345-352.
 * Markó B., and S. Csősz. 2002. Die europäischen Ameisenarten (Hymenoptera: Formicidae) des Hermannstädter (Sibiu, Rumänien) Naturkundemuseums I.: Unterfamilien Ponerinae, Myrmicinae und Dolichoderinae. Annales Historico-Naturales Musei Nationalis Hungarici 94: 109-121.
 * Martorell, M.. "Catálogos sinonímicos de insectos encontrados en Cataluña." Barcelona (1879).
 * Martínez Ibáñez, M. D., and X. Espadaler. "Revisión de las hormigas ibéricas de la colección M. Medina y nuevos datos de distribución." Actas de las VIII Jornadas de la Asociación Española de Entomología (1986): 1022-1034.
 * Mayr, G.. "Formicina austriaca. Beschreibung der bisher im österreichischen Kaiserstaate aufgefundenen Ameisen, nebst Hinzufügung jener in Deutschland, in der Schweiz und in Italien vorkommenden Arten." Verhandlungen der Zoologisch-Botanischen Vereins in Wien 5 (1855): 273-478.
 * Mei M. 1995. Arthropoda di Lampedusa, Linosa e Pantelleria (Canale di Sicilia, Mar Mediterraneo). Hymenoptera Formicidae (con diagnosi di due nuove specie). Il Naturalista Siciliano (4)19(suppl.): 753-772.
 * Menozzi C. 1918. Primo contributo alla conoscenza della fauna mirmecologica del Modenese. Atti della Società dei Naturalisti e Matematici di Modena. (5)4: 81-88.
 * Menozzi, C.. "Zur Kenntnis der Ameisenfauna der Balearen." Zoologischer Anzeiger 66 (1926): 180-182.
 * Mokrousov M. V., and V.A. Zryanin. 2015. Materials on the early spring wasps and ants fauna of Uzbekistan (Hymenoptera: Vespomorpha: Chrysidoidea, Scolioidea, Pompiloidea, Vespoidea, Apoidea [Spheciformes], Formicoidea). Entomological research Russia and its neighboring regions 5: 36–48.
 * Müller, G.. "Le formiche della Venezia Guilia e della Dalmazia." Bollettino della Società Adriatica di Scienze Naturali in Trieste 28 (1923): 11-180.
 * Nezhad S. H., S. P. Rad, F. Firouzi, and D. Agosti. 2012. New and additional records for the ant fauna from Iran (Hymenoptera: Formicidae). Zoology in the Middle East 55: 65-74.
 * Nylander, W.. "Synopsis des Formicides de France et d'Algérie." Annales des Sciences Naturelles, Zoologie (4)5 (1856): 51-109.
 * Obregon R., J. Lopez, J. L. Reyes-Lopez. 2014. A catalogue of the ant fauna (Hymenoptera, Formicidae) of Sierra Mágina (Jaén, southern Spain). Boletín de la Sociedad Entomológica Aragonesa (S.E.A.), 54: 370374.
 * Ordóñez-Urbano C., J. Reyes-López, and S. Carpintero-Ortega. 2007. Estudio faunísidos de los formícidos (Hymenoptera: Formicidae) asociados a los bosques de ribera en la Provincia de Córdoba (España). Primeras Aportaciones. Boletin Sociedad Entomológica Aragonesa 40: 367-375.
 * Paknia O., A. Radchenko, H. Alipanah, and M. Pfeiffer. 2008. A preliminary checklist of the ants (Hymenoptera: Formicidae) of Iran. Myrmecological News 11: 151-159.
 * Paraschivescu D. 1972. Fauna mirmecologica din zonele saline ale Romaniei. Studii si Cercetari de Biologie. Seria Zoologie 24: 489-495.
 * Paraschivescu D. 1978. Elemente balcanice in mirmecofauna R. S. Romania. Nymphaea 6: 463- 474.
 * Pashaei Rad S., B. Taylor, R. Torabi, E. Aram, G. Abolfathi, R. Afshari, F. Borjali, M. Ghatei, F. Hediary, F. Jazini, V. Heidary Kiah, Z. Mahmoudi, F. Safariyan, and M. Seiri. 2018. Further records of ants (Hymenoptera: Formicidae) from Iran. Zoology in the Middle East 64(2): 145-159.
 * Passera L. 1977. Peuplement myrmecologique du cordon littoral du Languedoc-Roussillon modifications anthropiques. Vie et Milieu 27: 249-265.
 * Peru L. 1999. Quelques ajouts et corrections a un catalogue des fourmis de la region Centre (Hymenoptera, Formicidae). Bulletin de l'Entomologie Tourangelles 20(1): 9-12.
 * Petrov I. Z. 2002. Contribution to the myrmecofauna (Formicidae, Hymenoptera) of the Banat Province (Serbia). Archives of Biological Sciences, Belgrade, 54(12): 57-64.
 * Petrov I. Z. 2010. Contribution to the myrmecofauna (Hymenoptera: Formicidae) of Krusevac and its surroundings. Acta entomologica serbica, 2010, 15(1): 145-148.
 * Petrov I. Z., B. Petrov, D. Milicic, T. Karan-Znidarsic. 2007. Contribution to the Myrmecofauna (Hymenoptera: Formicidae) of East and South Serbia. Acta Zoologica Bulgarica 59(3): 295-299.
 * Petrov I. Z., and C. A. Collingwood. 1992. Survey of the myrmecofauna (Formicidae, Hymenoptera) of Yugoslavia. Archives of Biological Sciences (Belgrade) 44: 79-91.
 * Pisarski B. 1969. Beiträge zur Kenntnis der Fauna Afghanistans. (Sammelergebnisse von O. Jakes 1963-64, D. Povolny 1965, D. Povolny & Fr. Tenora 1966, J. Simek 1965-66, D. Povolny, J. Gaisler, Z. Sebek & Fr. Tenora 1967). Formicidae, Hym. Casopis Moravského Musea v Brne 54(Suppl.): 305-326.
 * Platner C., J. Pinol, D. Sanders, and X. Espadaler. 2012. Trophic diversity in a Mediterranean food webStable isotope analysis of an ant community of an organic citrus grove. Basic and Applied Ecology 13: 587596.
 * Poldi B., M. Mei, and F. Rigato. 1995. Hymenoptera, Formicidae. Vol 102. Checklist delle specie della fauna Italiana: 1-10.
 * Reznikova Z. I. 2003. Distribution patterns of ants in different natural zones and landscapes in Kazakhstan and West Siberia along a meridian trend. Euroasian Entomological Journal 2(4): 235-342.
 * Rigato F., and J. K. Wetterer. 2018. Ants (Hymenoptera: Formicidae) of San Marino. Natural History Sciences. Atti Soc. it. Sci. nat. Museo civ. Stor. nat. Milano, 5(2): https://doi.org/10.4081/nhs.2018.367
 * Rigato F., and R. Sciaky. 1989. Contributo alla conoscenza della mirmecofauna della Val Gesso (alpi Marittime) (Hymenoptera, Formicidae). Boll. Mus. Reg. Sci. Nat. Torino 7(2): 427-442.
 * Rigato F., and R. Sciaky. 1991. The myrmecofauna of the Gesso Valley (Maritime Alps) (Hymenoptera Formicidae). Ethology Ecology and Evolution Special Issue 1: 87-89.
 * Rigato S., and I. Toni. 2011. Short notes 21. Hymenoptera, Formicidae. Pp. 873-882 in: Nardi, G.; Whitmore, D.; Bardiani, M.; Birtele, D.; Mason, F.; Spada, L.; Cerretti, P. (eds.) 2011. Biodiversity of Marganai and Montimannu (Sardinia). Research in the framework of the ICP Forests network. Conservazione Habitat Invertebrati, 5. Sommacampagna, Verona: Cierre Edizioni, 896 pp.
 * Rodríguez, A.. "Contribución al conocimiento de las hormigas (Hymenoptera, Formicidae) de Sierra Morena Central." Boletín de la Asociación Española de Entomología 5 (1982): 181-188.
 * Salata S., and L. Borowiec. 2019. Preliminary contributions toward a revision of Greek Messor Forel, 1890 (Hymenoptera: Formicidae). Turkish Journal of Zoology 43: 52-67.
 * Salata S., and L. Borowiec. 2019. Preliminary division of not socially parasitic Greek Temnothorax Mayr, 1861 (Hymenoptera, Formicidae) with a description of three new species. ZooKeys 877: 81-131.
 * Sanchez-Gil Jimeno R., and J. L. Reyes-Lopez. 2016. Study of ants species of the Sierra de San Carlos del Valle (Ciudad Real) and updating the provincial check list (Hymenoptera, Formicidae). Boln. Asoc. esp. Ent. 40 (1-2): 93-109.
 * Santschi F. 1926. Travaux scientifiques de l'Armée d'Orient (1916-1918). Fourmis. Bulletin du Muséum National d'Histoire Naturelle 32: 286-293.
 * Santschi, F.. "Inventa entomologica itineris Hispanici et Maroccani, quod a. 1926 fecerunt Harald et Håkan Lindberg. Fourmis du Bassin Méditerranéen occidental et du Maroc récoltées par MM. Lindberg." Societas Scientiarum Fennica (Helsingfors) 3 (14) (1931): 1-13.
 * Saunders, E.. "Hymenoptera aculeata from Majorca and Spain. I Majorca (1901)." Transactions of the Entomological Society of London (1904): 597-598.
 * Saunders, E.. "Hymenoptera aculeata from Majorca and Spain. II Spain (1901-1902)." Transactions of the Entomological Society of London (1904): 624-660.
 * Schembri, Stephen P. and Cedric A. Collingwood. A Revision of the Myrmecofauna of the Maltese Islands. 417-442.
 * Schlick-Steiner B. C., and F. M. Steiner. 1999. Faunistisch-ökologische Untersuchungen an den freilebenden Ameisen (Hymenoptera: Formicidae) Wiens. Myrmecologische Nachrichten 3: 9-53.
 * Schultz, R., A. G. Radchenko, and B. Seifert. "A critical checklist of the ants of Kyrgyzstan (Hymenoptera: Formicidae)." Myrmecologische Nachrichten 8 (2006): 201-207.
 * Scupola A., and R. Poggi. 2014. Ricerche zoologiche della nave oceanografica "minerva" (C.N.R.) sulle isole circumsarde. XXXIII. Formiche. Estratto dagli Annali del Museo Civico di Storia Naturale “G. Doria” 106: 93-131.
 * Scupola I. 2009. Le formiche delle Isole Egadi (Sicilia) (Hymenoptera, Formicidae). Bollettino del Museo Civico di Storia Naturale di Verona Botanica Zoologia 33: 97-103
 * Seabra, A. F.. "A entomología do trigo." Arquivos da Secçao de Biología e Parasitología Coimbra (1939): 700pp.
 * Seifert B. 1994. Die freilebenden Ameisenarten Deutschlands (Hymenoptera: Formicidae) und Angaben zu deren Taxonomie und Verbreitung. Abhandlungen und Berichte des Naturkundemuseums Görlitz 67(3): 1-44.
 * Seifert B. 1998. Rote Liste der Ameisen. - in: M. Binot, R. Bless, P. Boye, H. Gruttke und P. Pretscher: Rote Liste gefährdeter Tiere Deutschlands. Bonn-Bad Godesberg 1998: 130-133.
 * Sommer F., and H. Cagniant. 1988. Peuplements de fourmis des Albères Orientales (Pyrénées-Orientales, France) (Première partie). Vie Milieu 38: 189-200.
 * Soyer B. 1951. Les fourmis des collines des environs de Marseille. Bull Soc Linn Provence 18: 3-6.
 * Steiner F. M., S. Csosz, B. Marko, A. Gamisch, L. Rinnhofer, C. Folterbauer, S. Hammerle, C. Stauffer, W. Arthofer, B. C. Schlick‐Steiner. 2018. Turning one into five: Integrative taxonomy uncovers complex evolution of cryptic species in the harvester ant Messor “structor”. Molecular Phylogenetics and Evolution https://doi.org/10.1016/j.ympev.2018.04.005
 * Steiner F. M., S. Schödl, and B. C. Schlick-Steiner. 2002. Liste der Ameisen Österreichs (Hymenoptera: Formicidae), Stand Oktober 2002. Beiträge zur Entomofaunistik 3: 17-25.
 * Stukalyuk S. V. 2016. Crematogaster subdentata Mayr, 1877, a potentially invasive species of ant (Hymenoptera, Formicidae) new to the fauna of Crimea. Entomological Review 95(8)
 * Stukalyuk S. V., S. P. Ivanov. 2013. Rhytmics of daily activity of seven species of ants (Hymenoptera, Formicidae) in multi-species assemblages in the Crimea. Optimization and Protection of Ecosystems. Simferopol: TNU 9: 150-176.
 * Suñer i Escriche, David. "Contribució al coneixement mirmecologic de Gavarres, Montgrí, Guilleríes i la Serralada Transversal." Tesis Doctoral Universida (1991): 577 pp.
 * Tinaut A. 2016. Ants of the Tejeda, Almijara and Alhama Mountains Natural Park (Andalusia, Spain) (Hymenoptera, Formicidae). Boln. Asoc. esp. Ent., 40 (1-2): 125-159.
 * Trigos Peral G., and J. L. Reyes-Lopez. 2015. New record of Temnothorax algiricus (Hymenoptera, Formicidae) from the Iberian Peninsula and ten additional new records for the ant fauna from Málaga, with preliminary list of species from Natural Park Sierras de Tejeda, Almijara y Alhama (Andalusia, Spain). Boln. Asoc. esp. Ent. 39 (3-4): 411-416.
 * Trigos Peral G., and J. Reyes Lopez. 2016. Ants as bioindicators for monitoring of urban green in the south of the Iberian Peninsula. Proposal of functional groups. Iberomyrmex 8: 37-38.
 * Trigos-Peral G., and J. L. Reyes-Lopez. 2013. Primera relacion de los formicidos (Hymenoptera, Formicidae) de la Reserva Natural Lagunas de Campillos (Malaga). Boln. Asoc. esp. Ent. 37 (3-4): 217-224.
 * Vagalinski B., and A. Lapeva-Gjonova. 2012. The ants (Hymenoptera, Formicidae) of Plana Mountain (Bulgaria). Historia naturalis bulgarica 20: 87-101.
 * Vassilev I. 1987. The ants (Formicidae, Hymenoptera) from the valley of the river Racene. Annuaire de l'Universite de Sofia Kliment Ohridski Faceulte de Biologie 78: 74-79.
 * Vesni? A. 2011: Revidirani sistematski prijegled mrava Bosne i Hercegovine. Unutar : S. Lelo (urednik),  Fauna  Bosne  i  Hercegovine    Biosistematski  pregledi.  7.  izmijenjeno  i popravljeno interno izdanje Udruenja za inventarizaciju i zatitu  ivotinja, Ilija, Kanton Sarajevo, pp: 205-207.
 * Vesnic A., and R. Skrijelj. 2014. Biodiversity and variation in species of the genus Messor Forel, 1890 (Hymenoptera, Formicidae) in the Mediterranean part of Bosnia and Herzegovina. Works of the Faculty of Forestry University of Sarajevo 1: 37-44.
 * Vogrin, V.. "Prilog fauni Hymenoptera - Aculeata Jugoslavije." Zast. Bilja 31(suppl.) (1955): 1-74.
 * Vonshak M., and A. Ionescu-Hirsch. 2009. A checklist of the ants of Israel (Hymenoptera: Formicidae). Israel Journal of Entomology 39: 33-55.
 * Wheeler, W. M.. "Ants of the Balearic Islands." Folia Myrmecologica et Termitologica 1 (1926): 1-6.
 * Zimmermann, S.. "Beitrag zur Kenntnis der Ameisenfauna Süddalmatiens." Verhandlungen der Zoologisch-Botanisch Gesellschaft in Wien 84 (1935): 1-65.
 * Zimmermann, S.. "Beitrag zur Kenntnis der Ameisenfaunader Quarnerischen Inseln." Verhandlungen der Zoologisch-Botanisch Gesellschaft in Wien 80 (1930): 45-66.
 * Zina V., M. Branco, and J. C. Franco. 2017. Ant community associated to citrus canopy in Southern Portugal over a century after the invasion by the Argentine ant. Phytoparasitica 45:183–200.
 * de Haro, Andrés, C. A. Collingwood, and P. Comín del Río. "Prospección mirmecológica por Ibiza y Formentera (Baleares)." Orsis 2 (1986): 115-120.
 * de Haro, Andrés, and C. A. Collingwood. "Formícidos de las Sierras de Prades-Montsant, Sierras de Cavalls-Alfara-Montes Blancos (Tarragona)." Boletín de la Estación Central de Ecología 10 (1981): 55-58.
 * de Haro, Andrés, and C. A. Collingwood. "Prospección mirmecológica por las sierras de Aitana-Alfaro y los cabos de la Nao-San Antonio (Alicante) y su comparación con la fauna balear y de Córcega-Cerdeña." Orsis 3 (1988): 165-172.