Harpagoxenus sublaevis

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Harpagoxenus sublaevis
Conservation status
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmicinae
Tribe: Crematogastrini
Genus: Harpagoxenus
Species: H. sublaevis
Binomial name
Harpagoxenus sublaevis
(Nylander, 1849)

Harpagoxenus sublaevis casent0178588 profile 1.jpg

Harpagoxenus sublaevis casent0178588 dorsal 1.jpg

Specimen labels


This species is a slave-maker. In Russia (Zryanin & Zryanina, 2007) it is associated with Leptothorax acervorum (Beibl et al., 2005; de la Mora et al., 2021 (and included references)) and Leptothorax muscorum (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 Leptothorax gredleri (Foitzik et al., 2003; Guillem et al., 2014; de la Mora et al., 2021).

At a Glance • Dulotic  • Ergatoid queen  • Tandem running  • Diploid male  


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).


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

Latitudinal Distribution Pattern

Latitudinal Range: 69.9° to 41.38944444°.

Tropical South

Distribution based on Regional Taxon Lists

Palaearctic Region: Austria, Belarus, Belgium, Bulgaria, China, Czech Republic, Denmark, Estonia, Finland (type locality), France, Germany, Hungary, Iberian Peninsula, Italy, Lithuania, Netherlands, Norway, Poland, Romania, Russian Federation, Slovakia, Slovenia, Spain, Sweden, Switzerland, Türkiye, Ukraine.

Distribution based on AntMaps


Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.

Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.


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)).

Association with Other Organisms

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  • This species is a host for the cestode Type 2 (Choanotaenia unicoronata?) (a parasite) in South Tyrol, Italy (Buschinger, 1973; Laciny, 2021).


  • This species is a host for the gregarine Mattesia geminata (a parasite) in Germany (Buschinger & Kleespies, 1999).

Flight Period

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Source: antkeeping.info.

Life History Traits

  • Queen number: monogynous (Rissing and Pollock, 1988; Frumhoff & Ward, 1992)
  • Queen type: winged (Rissing and Pollock, 1988; Frumhoff & Ward, 1992) (queenless and queen-right worker reproduction)
  • Queen mating frequency: single (Rissing and Pollock, 1988; Frumhoff & Ward, 1992)



Images from AntWeb

Harpagoxenus sublaevis casent0178772 head 1.jpgHarpagoxenus sublaevis casent0178772 profile 1.jpgHarpagoxenus sublaevis casent0178772 dorsal 1.jpg
Worker. Specimen code casent0178772. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.


Images from AntWeb

Harpagoxenus sublaevis casent0178589 head 1.jpgHarpagoxenus sublaevis casent0178589 profile 1.jpgHarpagoxenus sublaevis casent0178589 profile 2.jpgHarpagoxenus sublaevis casent0178589 dorsal 1.jpg
Male (alate). Specimen code casent0178589. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MCZ, Cambridge, MA, USA.

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


The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • sublaevis. 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.



  • n = 20, 2n = 40 (Switzerland) (Buschinger et al., 1980; Hauschteck-Jungen & Jungen, 1983; Fischer, 1987).


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