Harpagoxenus sublaevis

This species is a slave-maker. In Russia (Zryanin & Zryanina, 2007) it is associated with (Beibl et al., 2005; de la Mora et al., 2021 (and included references)) and  (Lenoir et al., 2001; Bauer et al., 2009; Brandt et al., 2007; Fischer-Blass et al., 2006; Foitzik et al., 2003; de la Mora et al., 2021) in boreal regions. It is also known to enslave (Foitzik et al., 2003; Guillem et al., 2014; de la Mora et al., 2021).

Identification
Pale yellowish brown to brown; head large, rectangular, with weakly concave occiput. Frontal carinae extend backward to enclose whole length of antennal scape. Antennae 11 segmented with intermediate segments strongly transverse and enlarged 4 segmented club. Eyes large, set midway at sides of head. Mesopropodeal furrow deep and distinct; propodeal spines broad and short. Femora and tibiae short and broadly rounded. Head and mesopropodeum longitudinally striate, petiole nodes and gaster smooth and shining. Whole body and appendages covered with long, acute, pale hairs. Length: 3.5-5.5 mm (Collingwood 1979).

Distribution
Pyrenees to Caucasus; northern Italy to northern Norway (Collingwood 1979); China (Xu, 2012).

Distribution based on Regional Taxon Lists
Palaearctic Region: Austria, Belarus, Belgium, Bulgaria, China, Czech Republic, Denmark, Estonia, Finland, France, Germany, Hungary, Iberian Peninsula, Italy, Lithuania, Netherlands, Norway, Poland, Romania, Russian Federation, Slovakia, Slovenia, Spain, Sweden, Switzerland, Turkey, Ukraine.

Biology
Workers are unable to forage outside the nest and are not capable of brood tending or feeding themselves. Host species workers (slaves) are needed for the continuation of the colony. In mixed colonies no host queen can survive, though unmated gynes of the host species frequently develop from raided pupae. They dealate and serve as slaves. Host brood is commonly not devoured. During raids, host species eggs are even discarded outside the host nest and are not carried to the slavemaker nest.

Harpagoxenus sublaevis is a species with queen polymorphism. Most functional queens are wingless ergatoid queens that usually look like large workers; alate queens are very rare and seem absent in northern parts of the range. A genetical mechanism is claimed to underlie this queen polymorphism.

Collingwood (1979) - In Denmark and Fennoscandia nests containing host species and inquiline are commonly found in twigs on the ground, tree stumps or under bark but in the mountains of Central Europe they occur rarely under stones.

Guillem et al., 2014 - H. sublaevis is a slave-maker, invading Leptothorax host colonies, killing the queen and enslaving the host workers. Since the parasite workers are not adapted to forage, killing the host queen means that the parasite workers need to raid neighbouring host colonies to replenish the slaves. Harpagoxenus sublaevis co-exists with three species of Leptothorax, Leptothorax acervorum, Leptothorax muscorum and Leptothorax gredleri, although the former is the most commonly recorded host. Colonies can also consist of mixed Leptothorax spp. slaves (Bauer et al., 2010).

Three colonies of Harpagoxenus sublaevis were found by breaking open dry pine twigs that contained Leptothorax acervorum workers and queens on the Hanko peninsular in Southern Finland during September 2012. Roughly every one in 15 colonies of L. acervorum found was parasitized.

Guillem et al. (2014) examined cuticular hydrocarbon (CHC) profiles between this parasite and its hosts. They found that the parasitic species had CHC profiles that were indistinguishable from that of their hosts. The level of chemical mimicry even extended to the more subtle between-colony differences in profiles. In all cases the profiles of un-parasitized colonies were similar to those that were parasitized indicating that it is the parasites that have adjusted their profile to match that of their host and not vice versa. This explains why these social parasites are fully integrated members of each colony and are treated as nest-mates. It also helps to explain why raiding parties of Harpagoxenus workers are frequently killed or driven off when trying to raid or invade new host L. acervorum colonies (Winter and Buschinger, 1986), since they are carrying their own host colony odour, which is likely to be different from that of the one they are raiding. This is why parasites continue to use a wide range of other chemical and morphological adaptations associated with their parasitic lifestyle. These include a thickened cuticle and production of appeasement or propaganda compounds (e.g. Allies et al., 1986; Lloyd et al., 1986; Ollett et al., 1987; D'Ettorre et al., 2000). These tactics allow the parasite time to make the necessary adjustments to its profile. Acquiring a host profile may be possible in just a few hours (R. Kather, pers. comm., cited in Guillem et al. (2014)).

Male
Diploid males are known to occur in this species (Fisher (1987), in Loiselle et al., 1990; Cournault & Aron, 2009).

Nomenclature

 * . Myrmica sublaevis Nylander, 1849: 33 (w.) FINLAND.
 * Type-material: syntype workers (number not stated).
 * Type-locality: Finland: Kuusamo (no collector’s name).
 * Type-depository: ZMHF.
 * [Misspelled as sublevis by Emery & Forel, 1879: 457, Dalla Torre, 1893: 64, and others.]
 * Adlerz, 1896: 62 (q.m.); Viehmeyer, 1906: 58 (q.m.); Wheeler, G.C. & Wheeler, J. 1955b: 27 (l.).
 * Combination in Tomognathus: Mayr, 1861: 56 (in key); Dalla Torre, 1893: 64;
 * combination in Harpagoxenus: Forel, 1893a: 167.
 * Status as species: Nylander, 1856b: 95; Smith, F. 1858b: 121; Mayr, 1861: 56 (in key); Meinert, 1861: 333; Roger, 1863b: 26; Mayr, 1863: 457; Dours, 1873: 170; André, 1874: 188 (in key); Emery & Forel, 1879: 457; André, 1882c: 279 (in key); Nasonov, 1889: 29; Dalla Torre, 1893: 64; Wasmann, 1894: 164; Ruzsky, 1896: 71; Wheeler, W.M. 1901c: 707; Ruzsky, 1905b: 563; Viehmeyer, 1906: 57; Emery, 1908f: 549; Bondroit, 1910: 499; Stitz, 1914: 64; Forel, 1915d: 19 (in key); Emery, 1916b: 192; Escherich, 1917: 324; Bondroit, 1918: 141; Soudek, 1922: 30; Müller, 1923b: 99; Emery, 1924d: 266; Menozzi, 1925d: 33; Karavaiev, 1927c: 269 (in key); Arnol'di, 1933b: 598 (in key); Karavaiev, 1934: 150 (redescription); Grandi, 1935: 101; Stitz, 1939: 151; Holgersen, 1940: 183; Novák & Sadil, 1941: 84 (in key); Holgersen, 1942: 6; Holgersen, 1944: 168; Bernard, 1950a: 17; Bernard, 1967: 223 (redescription); Arnol’di, 1968: 1812; Kutter, 1968a: 60; Kutter, 1968b: 204; Baroni Urbani, 1971c: 133; Collingwood, 1971: 160; Banert & Pisarski, 1972: 351; Pisarski, 1975: 24; Kutter, 1977c: 146; Arnol’di & Dlussky, 1978: 544 (in key); Collingwood, 1978: 76 (in key); Collingwood, 1979: 78; Buschinger, 1981: 213; Agosti & Collingwood, 1987a: 55; Agosti & Collingwood, 1987b: 265 (in key); Nilsson & Douwes, 1987: 58; Radchenko, 1994b: 112 (in key); Bolton, 1995b: 212; Douwes, 1995: 90; Poldi, et al. 1995: 5; Buschinger, 1997: 2; Espadaler, 1997b: 29; Gallé, et al. 1998: 215; Czechowski, et al. 2002: 58; Tinaut, Ruano & Martinez, 2005: 462; Markó, Sipos, et al. 2006: 69; Petrov, 2006: 85 (in key); Bračko, 2007: 17; Seifert, 2007: 228; Werner & Wiezik, 2007: 140; Zryanin & Zryanina, 2007: 232; Casevitz-Weulersse & Galkowski, 2009: 488; Lapeva-Gjonova, et al. 2010: 20; Boer, 2010: 48; Csösz, et al. 2011: 57; Czechowski, et al. 2012: 161; Guénard & Dunn, 2012: 43; Kiran & Karaman, 2012: 19; Xu, 2012a: 22 (redescription); Borowiec, L. 2014: 81 (see note in bibliography); Lebas, et al. 2016: 282; Radchenko, 2016: 202; Steiner, et al. 2017: 10; Werner, et al. 2018: 7.
 * Senior synonym of hirtula: Mayr, 1861: 56 (in key); Roger, 1863b: 26; Mayr, 1863: 457; Dours, 1873: 170; Emery & Forel, 1879: 457; Dalla Torre, 1893: 64; Ruzsky, 1905b: 563; Collingwood, 1971: 160; Radchenko, 2007: 32; Radchenko, 2016: 202; Seifert, 2018: 196.
 * Distribution: Austria, Belarus, Belgium, Bulgaria, Czech Republic, Denmark, Estonia, Finland, France, Georgia, Gemany, Hungary, Italy. Lithuania, Netherlands, Norway, Poland, Romania, Russia, Serbia, Slovakia, Spain, Sweden, Switzerland, Turkey, Ukraine.
 * [Note: distribution from Borowiec, L. 2014: 81.]
 * Current subspecies: nominal plus caucasicus.
 * hirtula. Myrmica hirtula Nylander, 1849: 45 (w.) FINLAND.
 * Type-material: holotype worker.
 * Type-locality: Finland: Helsingfors (= Helsinki) (no collector’s name).
 * Type-depository: ZMHF.
 * Combination in Harpagoxenus: Emery, 1924d: 266.
 * Status as species: Smith, F. 1858b: 121.
 * Subspecies of sublaevis: Emery, 1924d: 266; Stitz, 1939: 154; Bolton, 1995b: 211.
 * Junior synonym of sublaevis: Mayr, 1861: 56 (in key); Roger, 1863b: 26; Mayr, 1863: 457; Dours, 1873: 170; Emery & Forel, 1879: 457; Dalla Torre, 1893: 64; Ruzsky, 1905b: 563; Collingwood, 1971: 160; Radchenko, 2007: 32; Radchenko, 2016: 202.

References based on Global Ant Biodiversity Informatics

 * Alinvi, O., J. Bohlin and J. P. Ball. 2008. RETRACTED ARTICLE: Interspecific competition among ants in the boreal forest: Testing predictions from a linear hierarchical competition model. Insectes Sociaux 55(1):1-11.
 * 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)
 * ArtDatabanken Bugs (via GBIG)
 * Baroni Urbani C., and C. A. Collingwood. 1977. The zoogeography of ants (Hymenoptera, Formicidae) in Northern Europe. Acta Zoologica Fennica 152: 1-34.
 * Bauer, S. V. Witte, M. Bohm and S. Foitzik. 2009. Fight or flight? A geographic mosaic in host reaction and potency of a chemical weapon in the social parasite Harpagoxenus sublaevis. Behavioural Ecology and Sociobiology 64:45-56
 * Beibl, J., R.J. Stuart, J. Heinze and S. Foitzik. 2005. Six origins of slavery in formicoxenine ants. Insectes Sociaux 52:291-297
 * Bernard F. 1957. Note sur quelques Leptothorax d'Europe centrale avec description de L. carinthiacus n. sp. (Hym. Formicidae). Bull. Soc. Entomol. Fr. 62: 46-53.
 * 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.. "Notes sur les fourmis de France. II. Peuplement des montagnes méridionales." Annales de la Société Entomologique de France 115 (1950): 1-36.
 * Bezdecka P. 1996. The ants of Slovakia (Hymenoptera: Formicidae). Entomofauna carpathica 8: 108-114.
 * Bezdecka P., and K. Bezdeckova. 2012. Updated list of the ants of the Czech Republic. Pp 7-12. Bezd??ka P. & Bezd??ková K. (eds) 2012: Blanok?ídlí v ?eských zemích a na Slovensku 8, Chaloupky, 1.-3. ?ervna 2012, sborník abstrakt? z konference.  MVJ Jihlava, 37 pp.
 * Blinov V. V. 1984. New for the fauna of Byelorussia ant species. Vestsi Akademii Navuk BSSR. Seryia Biialahichnykh Navuk 1984(5): 113-115.
 * Borowiec L. 2014. Catalogue of ants of Europe, the Mediterranean Basin and adjacent regions (Hymenoptera: Formicidae). Genus (Wroclaw) 25(1-2): 1-340.
 * Borowiec M. L. 2009. New data on the occurrence of formicoxenine ants (Hymenoptera: Formicidae) in Poland. Wiadomo?ci Entomologiczne 28: 237-246.
 * Bracko G. 2007. Checklist of the ants of Slovenia (Hymenoptera: Formicidae). Natura Sloveniae 9: 15-24
 * Buschinger A. 1999. Bemerkenswerte ameisenfunde aus Sudtirol (Hymenoptera: Formicidae). Myrmecologische Nachrichten 3: 1-8.
 * Buschinger, A., W. Ehrhardt, and K. Fischer. "Doronomyrmex pacis, Epimyrma stumperi und E. goesswaldi (Hym., Formicidae) neu für Frankreich." Insectes Sociaux 28 (1981): 67-70.
 * Casevitz-Weulersse J., and C. Galkowski. 2009. Liste actualisee des Fourmis de France (Hymenoptera, Formicidae). Bull. Soc. Entomol. Fr. 114: 475-510.
 * Collingwood C. A. 1971. A synopsis of the Formicidae of north Europe. Entomologist 104: 150-176
 * Collingwood C.A. 1959. Scandinavian Ants. Entomol. Rec. 71: 78-83
 * Collingwood, C. A. 1974. A revised list of Norwegian ants (Hymenoptera: Formicidae). Norsk Entomologisk Tidsskrift 21: 31-35.
 * Collingwood, C. A.. "The Formicidae (Hymenoptera) of Fennoscandia and Denmark." Fauna Entomologica Scandinavica 8 (1979): 1-174.
 * Creighton W. S. 1927. The slave-raids of Harpagoxenus americanus. Psyche (Cambridge) 34: 11-29.
 * 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
 * Czechowski W., and W. Czechowska. 1999. New sites in Poland and notes on the biology of socially parasitic ants Formicoxenus nitidulus (Nyl.) and Harpagoxenus sublaevis (Nyl.) (Hymenoptera, Formicidae). Fragm. Faun. (Warsaw) 42: 1-6.
 * Czechowski, W., and A. G. Radchenko. "Formica lusatica Seifert, 1997 (Hymenoptera: Formicidae), an ant species new to Finland, with notes on its biology and the description of males." Myrmecologische Nachrichten 8 (2006): 257-262.
 * 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.
 * Entomological Society of Latvia. 2003. http://leb.daba.lv/Formicidae.htm (Accessed on December 1st 2013).
 * Foitzik, S., S. Bauer, S. Laurent and P.S. Pennings. 2009. Genetic diversity, population structure and sex-biased dispersal in three co-evolving species. Journal of Evolutionary Biology 22:2470-2480
 * Gallé L. 1991. Structure and succession of ant assemblages in a north European sand dune area. Holarctic Ecology 14: 31-37.
 * Gibb H., and T. Johansson. 2011. Field tests of interspecific competition in ant assemblages: revisiting the dominant red wood ants. Journal of Animal Ecology 80: 548-557.
 * Grandi G. 1935. Contributi alla conoscenza degli Imenotteri Aculeati. XV. Boll. R. Ist. Entomol. Univ. Studi Bologna 8: 27-121.
 * Hauschteck-Jungen E., and H. Jungen. 1983. Ant chromosomes. II. Karyotypes of western palearctic species. Insectes Soc. 30: 149-164.
 * Heatwole H. 1989. Changes in ant assemblages across an artic treeline. Revue d'Entomologie du Quebec 34(1-2): 10-22.
 * Heinze, J. 1995. The Origin of Workerless Parasites in Leptothorax (S. Str.) (Hymenoptera: Formicidae). Psyche 102 (3-4) :195-214
 * Holgersen H. 1940. Myrmekologiske notiser I. Nor. Entomol. Tidsskr. 5: 183-187.
 * Holgersen H. 1942. Ants of northern Norway (Hym., Form.). Tromso Mus. Årsh. 63(2): 1-34.
 * Hölzel E. 1952. Ameisen Kärntens. Carinthia II 142: 89-132.
 * IZIKO South Africa Museum Collection
 * Kupianskaia A.N. 1990. Murav'I (Hymenoptera, Formicidae) Dal'nego Vostoka SSSR (1989). Vladivostok. 258 pages.
 * Kvamme T. 1982. Atlas of the Formicidae of Norway (Hymenoptera: Aculeata). Insecta Norvegiae 2: 1-56.
 * Legakis Collection Database
 * Malozemova L. A. 1972. Ants of steppe forests, their distribution by habitats, and perspectives of their utilization for protection of forests (north Kazakhstan). [In Russian.]. Zoologicheskii Zhurnal 51: 57-68.
 * Markó B., B. Sipos, S. Csősz, K. Kiss, I. Boros, and L. Gallé. 2006. A comprehensive list of the ants of Romania (Hymenoptera: Formicidae). Myrmecologische Nachrichten 9: 65-76.
 * Markó B., S. Csősz. 2001: Nine new ant species in the Romanian fauna (Hymenoptera: Formicidae): morphology, biology and distribution.  Entomologica Romanica 6: 127-132.
 * Neumeyer R., and B. Seifert. 2005. Commented check list of free living ant (Hymenoptera: Formicidae) species of Switzerland. Bulletin de la Societe Entomologique Suisse 78: 1-17.
 * Nielsen M. G. 2011. A check list of Danish ants and proposed common names. Ent. Meddr. 79: 13-18.
 * Paukkunen J., and M. V. Kozlov. 2015. Stinging wasps, ants and bees (Hy menoptera: Aculeata) of the Murmansk region, Northwest Russia. — Entomol. Fennica. 26: 53–73.
 * Petal J. M. 1963. Faune des fourmis de la reserve de tourbiere en projet a Rakowskie Bagno pres de Frampol (voivodie de Lublin). Annales Universitatis Mariae Curie-Sk?odowska 58(7): 143-174.
 * Petrov I. Z., and C. A. Collingwood. 1992. Survey of the myrmecofauna (Formicidae, Hymenoptera) of Yugoslavia. Archives of Biological Sciences (Belgrade) 44: 79-91.
 * Radchenko A. G. 2007. The ants (Hymenoptera, Formicidae) in the collection of William Nylander. Fragmenta Faunistica (Warsaw) 50: 27-41.
 * Saaristo M. I. 1990. New provincial records for twelve ant species (Hymenoptera, Formicidae) from Finland. Entomologica Fennica 1: 191-192.
 * Salata S. 2014. Ants (Hymenoptera: Formicidae) of the National Park of the Stołowe Mts. Przyroda Sudetow 17: 161-172.
 * 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.
 * Shlyakhtenok A. S. 2007. Hymenoptera Aculeata of Raised Bogs in Belarus. Entomological Review 87(2): 136147.
 * Sonnenburg H. 2005. Die Ameisenfauna (Hymenoptera: Formicidae) Niedersachsens und Bremens. Braunschweiger Naturkundliche Schriften 7: 377-441.
 * 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.
 * Sturm P., and H. Distler. 2003. Rote liste gefahrdeter ameisen (Hymenoptera: Formicoidea) Bayerns. In: Bayrisches Landesamt für Umweltschutz (Ed.). Rote Liste gefährdeter Tiere Bayerns. 208-212.
 * Vepsalainen K., H. Ikonene, and M. J. Koivula. 2008. The structure of ant assembalges in an urban area of Helsinki, southern Finland. Ann. Zool. Fennici 45: 109-127.
 * Wegnez P., and F. Mourey. 2016. Formica uralensis Ruzsky, 1895 une espèce encore présente en France mais pour combien de temps ? (Hymenoptera: Formicidae). Bulletin de la Société royale belge d’Entomologie 152: 72-80.
 * Xu Z.-H. 2012. A newly recorded genus and species, Harpagoxenus sublaevis, from China with a key to the known species of Harpagoxenus of the world (Hymenoptera: Formicidae). Sociobiology 59:19-25.
 * Zhuytszyuan D. 2016. The ants (Hymenoptera, Formicidae) Nizhne-Svirsky reserve and their environmental features. Master's thesis Saint Petersburg State University.
 * Zryanin V. A., and T. A. Zryanina. 2007. New data on the ant fauna Hymenoptera, Formicidae in the middle Volga River Basin. Uspekhi Sovremennoi Biologii 127(2): 226-240.