Dulosis

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Dulosis is the presence of permanent parasitism with slavery.

The biology of Temnothorax muellerianus and Polyergus rufescens provide representative accounts of dulosis.

Note that slave-making ants show wide variation in the degree of degeneracy of the worker caste. At one end of the spectrum (facultative dulosis), represented by Formica sanguinea, workers are self-sufficient, conduct all of the custodian tasks of the colony on their own and can easily survive without the support of slaves. In intermediate forms, termed "obligatory slavemakers", workers have a very limited behavioral repertory, but are behaviorally and often morphologically specialized for fighting and slave raiding. They are completely helpless without their slaves and are permanently dependent on regular replenishment of their slave stock through raids on host species colonies. At the other extreme, the term "degenerate slavemaker" has been coined for species that have secondarily lost their worker caste, or reduced its number, and hence can no longer conduct slave raids. Examples of degenerate slavemakers are Temnothorax kraussei (workers reduced to a few, or completely absent), Temnothorax corsicus and Temnothorax adlerzi (both workerless; all three belong to former genus Myrmoxenus), and Temnothorax brunneus (workerless, former genus Chalepoxenus).

Degenerate slavemakers are distinct from workerless inquilines in that the queens of the degenerate slavemakers kill the host colony queen by throttling (former Myrmoxenus species) or stinging (former Chalepoxenus species), while in workerless inquilines the queens live in queenright host colonies. Evolutionarily, degenerate slavemaking seems to have arisen fewer times than workerless inquilinism, with degenerate slavemakers most closely related to other dulotic species (thus previously being placed together in genera such as Chalepoxenus, Myrmoxenus or currently in Strongylognathus), while workerless inquilines have tended to arise independently of each other and are (generally) more closely related to their hosts than to other inquilines (for example Leptothorax kutteri and Leptothorax pacis with Leptothorax acervorum as the host species).

Known Slave-Makers

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Slave Maker Host Locality Source Notes
Formica adamsi alpina Formica neorufibarbis Mackay and Mackay 2002
Formica aserva Formica neorufibarbis Wheeler and Wheeler, 1986
Formica aserva Formica podzolica Ruano et al., 2019; Savolainen & Deslippe, 2001; de la Mora et al., 2021
Formica creightoni Formica neogagates J. Trager (pers. comm.)
Formica creightoni Formica pallidefulva J. Trager (pers. comm.)
Formica densiventris Formica argentea
Formica densiventris Formica biophilica doubtful, possible misidentification of host species
Formica densiventris Formica neogagates
Formica densiventris Formica neorufibarbis
Formica densiventris Formica subaenescens host given as ''Formica fusca'', but likely ''Formica subaenescens'', needs confirmation
Formica densiventris Formica subpolita
Formica densiventris Formica subsericea host possibly ''Formica podzolica'', needs confirmation
Formica fossaceps Formica montana
Formica gynocrates Formica argentea Bono et al., 2006; de la Mora et al., 2021 host as ''Formica'' sp. cf. ''argentea''
Formica gynocrates Formica lasioides Snelling & Buren, 1985 uncommon
Formica gynocrates Formica neogagates Snelling & Buren, 1985 uncommon
Formica gynocrates Formica perpilosa S. Cover, MCZbase
Formica gynocrates Formica vinculans Snelling & Buren, 1985
Formica impexa Formica subaenescens
Formica pergandei Formica biophilica
Formica pergandei Formica difficilis
Formica pergandei Formica dolosa
Formica pergandei Formica fusca
Formica pergandei Formica glacialis Apple et al., 2014; de la Mora et al., 2021
Formica pergandei Formica incerta
Formica pergandei Formica neogagates Mackay and Mackay, 2002
Formica pergandei Formica obscuriventris
Formica pergandei Formica pallidefulva
Formica pergandei Formica podzolica
Formica pergandei Formica querquetulana J. Trager, pers. comm.
Formica pergandei Formica subsericea
Formica puberula Formica subpolita Mackay & Mackay, 2002
Formica puberula Formica altipetens Mackay & Mackay, 2002
Formica puberula Formica bradleyi Mackay & Mackay, 2002
Formica puberula Formica densiventris Mackay & Mackay, 2002
Formica puberula Formica hewitti Mackay & Mackay, 2002
Formica puberula Formica lasioides Mackay & Mackay, 2002
Formica puberula Formica lepida Mackay & Mackay, 2002
Formica puberula Formica montana Mackay & Mackay, 2002
Formica puberula Formica neoclara Mackay & Mackay, 2002
Formica puberula Formica neorufibarbis Mackay & Mackay, 2002
Formica puberula Formica occulta Bono et al., 2006; de la Mora et al., 2021
Formica puberula Formica pallidefulva Mackay & Mackay, 2002
Formica puberula Formica subaenescens Mackay & Mackay, 2002 reported as ''Formica fusca''
Formica rubicunda Formica altipetens
Formica rubicunda Formica argentea
Formica rubicunda Formica aserva
Formica rubicunda Formica bradleyi doubtful, slave maker possibly ''Formica pergandei''
Formica rubicunda Formica fossaceps
Formica rubicunda Formica fusca
Formica rubicunda Formica lasioides
Formica rubicunda Formica lepida
Formica rubicunda Formica montana
Formica rubicunda Formica neoclara
Formica rubicunda Formica neogagates
Formica rubicunda Formica neorufibarbis
Formica rubicunda Formica obscuriventris
Formica rubicunda Formica obtusopilosa
Formica rubicunda Formica pallidefulva
Formica rubicunda Formica subsericea
Formica rubicunda Myrmica possibly a prey item rather than a slave
Formica sanguinea Formica candida
Formica sanguinea Formica cinerea
Formica sanguinea Formica cunicularia Ruano et al., 2019; Seifert, 2018; de la Mora et al., 2021
Formica sanguinea Formica fusca Ruano et al., 2019; Seifert, 2018; de la Mora et al., 2021
Formica sanguinea Formica gagatoides
Formica sanguinea Formica hayashi Japanese Ant Image Database
Formica sanguinea Formica japonica Ruano et al., 2019; de la Mora et al., 2021; Japanese Ant Image Database
Formica sanguinea Formica lemani Ruano et al., 2019; de la Mora et al., 2021; Japanese Ant Image Database
Formica sanguinea Formica picea
Formica sanguinea Formica polyctena Seifert, 2018; de la Mora et al., 2021
Formica sanguinea Formica pratensis Ruano et al., 2019; Seifert, 2018; de la Mora et al., 2021 probable ephemeral host
Formica sanguinea Formica rufa Ruano et al., 2019; Seifert, 2018; de la Mora et al., 2021 probable ephemeral host
Formica sanguinea Formica rufibarbis
Formica sanguinea Formica transkaucasica
Formica sanguinea Myrmica jessensis Hayashida, 1966
Formica subintegra Formica glacialis Apple et al., 2014; de la Mora et al., 2021
Formica subintegra Formica pallidefulva Mackay & Mackay, 2002
Formica subintegra Formica podzolica Savolainen & Deslippe, 2001; de la Mora et al., 2021
Formica wheeleri Formica altipetens
Formica wheeleri Formica bradleyi
Formica wheeleri Formica fusca probably ''Formica subaenescens'' rather than ''F. fusca''
Formica wheeleri Formica lasioides
Formica wheeleri Formica lepida
Formica wheeleri Formica neogagates Mackay and Mackay, 2002
Formica wheeleri Formica neorufibarbis
Harpagoxenus canadensis Leptothorax canadensis Beibl et al., 2005; de la Mora et al., 2021
Harpagoxenus canadensis Leptothorax muscorum Québec, Canada Smith, 1939
Harpagoxenus sublaevis Leptothorax acervorum Beibl et al., 2005; de la Mora et al., 2021 (and included references)
Harpagoxenus sublaevis Leptothorax gredleri Foitzik et al., 2003; Guillem et al., 2014; de la Mora et al., 2021
Harpagoxenus sublaevis 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
Myrmecocystus depilis Myrmecocystus mimicus Holldobler et al., 2011; Kronauer et al., 2003; de la Mora et al., 2021
Polyergus bicolor Formica neorufibarbis
Polyergus bicolor Formica subaenescens Trager, 2013; de la Mora et al., 2021
Polyergus breviceps Formica altipetens Trager, 2013; Torres & Tsutsui, 2016; de la Mora et al., 2021
Polyergus breviceps Formica canadensis Trager, 2013; de la Mora et al., 2021
Polyergus breviceps Formica montana Trager, 2013; de la Mora et al., 2021
Polyergus breviceps Formica neoclara Trager, 2013; de la Mora et al., 2021
Polyergus breviceps Formica occulta Mackay and Mackay, 2002; Trager, 2013; de la Mora et al., 2021; Bono et al., 2006; Bono et al., 2007; Johnson, 2000; Johnson et al., 2005 may be ''Polyergus mexicanus'' or ''Polyergus topoffi''
Polyergus longicornis Formica dolosa Goodloe & Sanwald, 1985; MacGown & Brown, 2006; King & Trager, 2007; Trager, 2013; de la Mora et al., 2021 slave-maker misidentified as ''Polyergus lucidus'' in some publications
Polyergus lucidus Formica incerta Goodloe & Sanwald, 1985; Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica accreta Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica aerata
Polyergus mexicanus Formica argentea Moffett, 2010; Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica cf. occulta Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica fusca Trager, 2013; Sulger et al., 2014; de la Mora et al., 2021 probably ''Formica subaenescens'', not ''Formica fusca''
Polyergus mexicanus Formica hewitti Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica manni Wheeler & Wheeler, 1986; Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica microphthalma Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica neoclara Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica neogagates Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica neorufibarbis Mackay and Mackay, 2002; Trager, 2013; de la Mora et al., 2021 as ''Polyergus breviceps''
Polyergus mexicanus Formica occulta
Polyergus mexicanus Formica pacifica Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica podzolica Savolainen & Deslippe, 2001; Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica subpolita
Polyergus mexicanus Formica subsericea Trager, 2013; de la Mora et al., 2021
Polyergus mexicanus Formica vinculans Trager, 2013; de la Mora et al., 2021
Polyergus montivagus Formica pallidefulva King & Trager, 2007; Trager, 2013; de la Mora et al., 2021
Polyergus nigerrimus Formica candida Trager, 2013; de la Mora et al., 2021
Polyergus nigerrimus Formica kozlovi
Polyergus oligergus Formica archboldi King & Trager, 2007; Trager, 2013; de la Mora et al., 2021
Polyergus ruber Formica biophilica Trager, 2013; de la Mora et al., 2021
Polyergus rufescens Formica cinerea Trager, 2013; Seifert, 2018; de la Mora et al., 2021
Polyergus rufescens Formica clara Trager, 2013; Seifert, 2018; de la Mora et al., 2021
Polyergus rufescens Formica cunicularia D'Ettorre et al., 2000; Mori et al., 2000; Visiccio et al., 2000; Visiccio et al., 2001; Romani et al., 2006; Seifert, 2018; Trager, 2013; de la Mora et al., 2021
Polyergus rufescens Formica fusca Trager, 2013; Seifert, 2018; de la Mora et al., 2021
Polyergus rufescens Formica fuscocinerea Seifert, 2018; de la Mora et al., 2021
Polyergus rufescens Formica gagates Trager, 2013; Seifert, 2018; de la Mora et al., 2021
Polyergus rufescens Formica gerardi Seifert, 2018; de la Mora et al., 2021
Polyergus rufescens Formica rufibarbis Mori et al., 2000; Trager, 2013; Seifert, 2018; de la Mora et al., 2021
Polyergus rufescens Formica tianshanica Seifert & Schultz, 2009; Seifert, 2018; de la Mora et al., 2021
Polyergus samurai Formica fukaii Trager, 2013; de la Mora et al., 2021 rare host
Polyergus samurai Formica glabridorsis Trager, 2013; de la Mora et al., 2021
Polyergus samurai Formica hayashi Trager, 2013; de la Mora et al., 2021 uncommon host
Polyergus samurai Formica japonica Tsuneoka, 2008; Tsuneoka & Akino, 2009; Tsuneoka & Akino, 2012; Trager, 2013; de la Mora et al., 2021 most common host
Polyergus sanwaldi Formica dolosa Goodloe and Sanwald, 1985; Goodloe, Sanwald and Topoff, 1987; Trager, 2013; de la Mora et al., 2021
Polyergus topoffi Formica foreliana Trager, 2013
Polyergus topoffi Formica gnava Trager, 2013; de la Mora et al., 2021
Polyergus topoffi Formica subcyanea
Polyergus vinosus Formica moki Trager, 2013; de la Mora et al., 2021
Rossomyrmex anatolicus Proformica korbi Tinaut et al., 2010; Ruano et al., 2013; de la Mora et al., 2021
Rossomyrmex minuchae Proformica longiseta Ruano & Tinaut, 1999; D'Ettorre & Heinze, 2001; Errard et al., 2006; Ruano et al., 2013; de la Mora et al., 2021
Rossomyrmex proformicarum Proformica epinotalis D'Ettorre & Heinze, 2001; Ruano et al., 2013; de la Mora et al., 2021
Rossomyrmex quandratinodus Proformica sp. Tinaut et al., 2010; Ruano et al., 2013; de la Mora et al., 2021
Strongylognathus alpinus Tetramorium alpestre Seifert, 2018; de la Mora et al., 2021
Strongylognathus alpinus Tetramorium impurum Sanetra & Buschinger, 2000; Wagner et al. 2017; Seifert, 2018; de la Mora et al., 2021
Strongylognathus christophi Tetramorium cf. caespitum Sanetra & Buschinger, 2000; de la Mora et al., 2021
Strongylognathus christophi Tetramorium cf. impurum Sanetra & Buschinger, 2000; de la Mora et al., 2021
Strongylognathus destefanii Tetramorium cf. impurum Sanetra & Buschinger, 2000; de la Mora et al., 2021
Strongylognathus destefanii Tetramorium cf. semilaeve Sanetra & Buschinger, 2000; de la Mora et al., 2021
Strongylognathus huberi Tetramorium caespitum Sanetra & Buschinger, 2000; Seifert, 2018; de la Mora et al., 2021
Strongylognathus huberi Tetramorium impurum Sanetra & Buschinger, 2000; Seifert, 2018; de la Mora et al., 2021
Strongylognathus huberi Tetramorium semilaeve Seifert, 2018; de la Mora et al., 2021
Strongylognathus italicus Tetramorium caespitum Seifert, 2018; de la Mora et al., 2021
Strongylognathus italicus Tetramorium impurum Sanetra & Buschinger, 2000; de la Mora et al., 2021
Strongylognathus koreanus Tetramorium tsushimae Japan Collingwood, 1976
Strongylognathus kratochvili Tetramorium caespitum Seifert, 2018; de la Mora et al., 2021
Strongylognathus kratochvili Tetramorium moravicum Seifert, 2018; de la Mora et al., 2021
Strongylognathus silvestrii Tetramorium lucidulum Sanetra & Buschinger, 2000; de la Mora et al., 2021
Strongylognathus silvestrii Tetramorium semilaeve Sanetra & Buschinger, 2000; de la Mora et al., 2021
Strongylognathus testaceus Tetramorium alpestre Seifert, 2018; de la Mora et al., 2021
Strongylognathus testaceus Tetramorium caespitum
Strongylognathus testaceus Tetramorium impurum Wagner et al. 2017
Temnothorax algerianus Temnothorax curtulus Buschinger et al., 1990
Temnothorax algerianus Temnothorax gentilis Buschinger et al., 1990
Temnothorax algerianus Temnothorax monjauzei Buschinger et al., 1990
Temnothorax algerianus Temnothorax spinosus Algeria, Morocco Buschinger et al., 1990 the most common slave
Temnothorax americanus Temnothorax ambiguus
Temnothorax americanus Temnothorax curvispinosus
Temnothorax americanus Temnothorax longispinosus
Temnothorax bikara Temnothorax makora Japan Japanese Ant Image Database
Temnothorax bikara Temnothorax spinosior Japan Japanese Ant Image Database
Temnothorax brunneus Temnothorax marocanus Morocco Buschinger & Douwes, 1990; Beibl et al., 2005; de la Mora et al., 2021
Temnothorax brunneus Temnothorax rottenbergii Morocco Buschinger & Douwes, 1990
Temnothorax duloticus Temnothorax ambiguus
Temnothorax duloticus Temnothorax curvispinosus
Temnothorax duloticus Temnothorax longispinosus
Temnothorax gordiagini Temnothorax lichtensteini Buschinger et al., 1984
Temnothorax gordiagini Temnothorax nylanderi Buschinger et al., 1984
Temnothorax gordiagini Temnothorax serviculus Buschinger et al., 1984
Temnothorax gordiagini Temnothorax unifasciatus Buschinger et al., 1984
Temnothorax kraussei Temnothorax recedens Buschinger et al., 1984
Temnothorax kutteri Temnothorax aveli Acosta Salmerón et al., 1983 as rabaudi, rare host
Temnothorax kutteri Temnothorax aveli Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri Temnothorax aveli Cagniant, 1973 as berlandi, rare host
Temnothorax kutteri Temnothorax exilis Acosta Salmerón et al., 1983 rare host
Temnothorax kutteri Temnothorax exilis Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri Temnothorax exilis Espadaler and Restrepo, 1983 as specularis, fairly common host
Temnothorax kutteri Temnothorax niger Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri Temnothorax niger Cagniant, 1973 rare host
Temnothorax kutteri Temnothorax racovitzai Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri Temnothorax racovitzai Buschinger et al., 1988 as massiliensis, most common host
Temnothorax kutteri Temnothorax racovitzai Cagniant, 1973 rare host
Temnothorax kutteri Temnothorax recedens Espadaler & Restrepo, 1983; Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri Temnothorax unifasciatus Beibl et al., 2005; Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri Temnothorax unifasciatus Espadaler & Restrepo, 1983 rare host
Temnothorax muellerianus Temnothorax affinis Buschinger et al., 1988 rare host
Temnothorax muellerianus Temnothorax exilis Crete Salata, Borowiec & Trichas, 2018
Temnothorax muellerianus Temnothorax flavicornis Buschinger et al., 1988 rare host
Temnothorax muellerianus Temnothorax luteus Buschinger et al., 1988 as Leptothorax tristis
Temnothorax muellerianus Temnothorax nigriceps Buschinger et al., 1988 common host
Temnothorax muellerianus Temnothorax recedens Buschinger et al., 1988 common host
Temnothorax muellerianus Temnothorax semiruber Crete Salata, Borowiec & Trichas, 2018
Temnothorax muellerianus Temnothorax tuberum Buschinger et al., 1988 rare host
Temnothorax muellerianus Temnothorax unifasciatus Buschinger et al., 1988 most common host
Temnothorax pilagens Temnothorax ambiguus
Temnothorax pilagens Temnothorax longispinosus
Temnothorax ravouxi Temnothorax affinis southern Europe Czechowski & Czechowska, 2000 relatively rare
Temnothorax ravouxi Temnothorax albipennis Czechowski & Czechowska, 2000
Temnothorax ravouxi Temnothorax nadigi Poland Czechowski & Czechowska, 2000
Temnothorax ravouxi Temnothorax nigriceps Czechowski & Czechowska, 2000
Temnothorax ravouxi Temnothorax tuberum eastern Europe Czechowski & Czechowska, 2000
Temnothorax ravouxi Temnothorax unifasciatus mainly western Europe Czechowski & Czechowska, 2000
Temnothorax tamarae Temnothorax crasecundus Georgia Csösz et al., 2015
Temnothorax tamarae Temnothorax sp. NYL2 Georgia Gratiashvili et al., 2014 an undescribed species closely related to ''Temnothorax nylanderi''
Temnothorax tamarae Temnothorax unifasciatus Georgia Gratiashvili et al., 2014 needs confirmation
Temnothorax tramieri Temnothorax spinosus Algeria, Morocco Cagniant, 1983

Taxonomic Distribution

Subfamily placement (in red) of species known to be dulotic.




Leptanillinae



Martialinae








Proceratiinae




Amblyoponinae



Apomyrminae






Ponerinae




Agroecomyrmecinae



Paraponerinae








Dorylinae






Aneuretinae



Dolichoderinae





Myrmeciinae



Pseudomyrmecinae







Formicinae - 3 genera




Myrmicinae - 3 genera




Ectatomminae



Heteroponerinae










Known Hosts of Slave-Makers

Click here to show/hide known hosts of slave-makers.
Host Slave Maker Locality Source Notes
Formica accreta Polyergus mexicanus Trager, 2013; de la Mora et al., 2021
Formica aerata Polyergus mexicanus
Formica altipetens Formica puberula Mackay & Mackay, 2002
Formica altipetens Formica rubicunda
Formica altipetens Formica wheeleri
Formica altipetens Polyergus breviceps Trager, 2013; Torres & Tsutsui, 2016; de la Mora et al., 2021
Formica archboldi Polyergus oligergus King & Trager, 2007; Trager, 2013; de la Mora et al., 2021 as ''Polyergus lucidus'' in early records
Formica argentea Formica densiventris
Formica argentea Formica gynocrates Bono et al., 2006; de la Mora et al., 2021 host as ''Formica'' sp. cf. ''argentea''
Formica argentea Formica rubicunda
Formica argentea Polyergus mexicanus Moffett, 2010; Trager, 2013; de la Mora et al., 2021
Formica aserva Formica rubicunda doubtful, needs confirmation
Formica biophilica Formica densiventris doubtful, possible misidentification of host species
Formica biophilica Polyergus ruber Trager, 2013; de la Mora et al., 2021
Formica bradleyi Formica puberula Mackay & Mackay, 2002
Formica bradleyi Formica rubicunda doubtful, slave maker possibly ''Formica pergandei''
Formica bradleyi Formica wheeleri
Formica canadensis Polyergus breviceps Trager, 2013; de la Mora et al., 2021
Formica candida Formica sanguinea
Formica candida Polyergus nigerrimus Trager, 2013; de la Mora et al., 2021
Formica cinerea Formica sanguinea
Formica cinerea Polyergus rufescens Trager, 2013; Seifert, 2018; de la Mora et al., 2021
Formica clara Polyergus rufescens Trager, 2013; Seifert, 2018; de la Mora et al., 2021
Formica cunicularia Formica sanguinea Ruano et al., 2019; Seifert, 2018; de la Mora et al., 2021
Formica cunicularia Polyergus rufescens Romani et al., 2006; D'Ettorre et al., 2000; Mori et al., 2000; Visiccio et al., 2000; Visiccio et al., 2001; Romani et al., 2006; Seifert, 2018; Trager, 2013; de la Mora et al., 2021
Formica difficilis Formica pergandei
Formica dolosa Formica pergandei
Formica dolosa Polyergus longicornis Goodloe & Sanwald, 1985; MacGown & Brown, 2006; King & Trager, 2007; Trager, 2013; de la Mora et al., 2021 slave-maker misidentified as ''Polyergus lucidus''
Formica dolosa Polyergus sanwaldi Goodloe and Sanwald, 1985; Goodloe, Sanwald and Topoff, 1987; Trager, 2013; de la Mora et al., 2021
Formica foreliana Polyergus topoffi Trager, 2013
Formica fossaceps Formica rubicunda
Formica fukaii Polyergus samurai Trager, 2013; de la Mora et al., 2021 a relatively rare host
Formica fusca Formica sanguinea Ruano et al., 2019; Seifert, 2018; de la Mora et al., 2021
Formica fusca Polyergus mexicanus Trager, 2013; Sulger et al., 2014; de la Mora et al., 2021
Formica fusca Polyergus rufescens Trager, 2013; Seifert, 2018; de la Mora et al., 2021
Formica fuscocinerea Polyergus rufescens Seifert, 2018; de la Mora et al., 2021
Formica gagates Polyergus rufescens Trager, 2013; Seifert, 2018; de la Mora et al., 2021
Formica gagatoides Formica sanguinea
Formica gerardi Polyergus rufescens Seifert, 2018; de la Mora et al., 2021
Formica glabridorsis Polyergus samurai Trager, 2013; de la Mora et al., 2021
Formica glacialis Formica dakotensis J. Trager, pers. comm. smetimes as a cohabiting inquiline
Formica glacialis Formica pergandei Apple et al., 2014; de la Mora et al., 2021
Formica glacialis Formica subintegra Apple et al., 2014; de la Mora et al., 2021
Formica gnava Polyergus topoffi Trager, 2013; de la Mora et al., 2021
Formica hayashi Polyergus samurai Trager, 2013; de la Mora et al., 2021 a relatively rare host
Formica hewitti Formica puberula Mackay & Mackay, 2002
Formica hewitti Polyergus mexicanus Trager, 2013; de la Mora et al., 2021
Formica incerta Formica pergandei
Formica incerta Polyergus lucidus Goodloe & Sanwald, 1985; Trager, 2013; de la Mora et al., 2021
Formica japonica Formica sanguinea Ruano et al., 2019; de la Mora et al., 2021
Formica japonica Polyergus samurai Tsuneoka, 2008; Tsuneoka & Akino, 2009; Tsuneoka & Akino, 2012; Trager, 2013; de la Mora et al., 2021
Formica kozlovi Polyergus nigerrimus
Formica lasioides Formica gynocrates Snelling & Buren, 1985 uncommon
Formica lasioides Formica puberula Mackay & Mackay, 2002
Formica lasioides Formica rubicunda
Formica lasioides Formica wheeleri
Formica lemani Formica sanguinea Ruano et al., 2019; de la Mora et al., 2021
Formica lepida Formica puberula Mackay & Mackay, 2002
Formica lepida Formica rubicunda
Formica lepida Formica wheeleri
Formica manni Polyergus mexicanus Wheeler & Wheeler, 1986; Trager, 2013; de la Mora et al., 2021
Formica microphthalma Polyergus mexicanus Trager, 2013; de la Mora et al., 2021
Formica moki Polyergus vinosus Trager, 2013; de la Mora et al., 2021
Formica montana Formica fossaceps
Formica montana Formica puberula Mackay & Mackay, 2002
Formica montana Formica rubicunda
Formica montana Polyergus breviceps Trager, 2013; de la Mora et al., 2021
Formica neoclara Formica puberula Mackay & Mackay, 2002
Formica neoclara Formica rubicunda
Formica neoclara Polyergus breviceps Trager, 2013; de la Mora et al., 2021
Formica neoclara Polyergus mexicanus Trager, 2013; de la Mora et al., 2021
Formica neogagates Formica creightoni J. Trager (pers. comm.)
Formica neogagates Formica densiventris Mackay and Mackay 2002
Formica neogagates Formica gynocrates Snelling & Buren, 1985 uncommon
Formica neogagates Formica pergandei Mackay and Mackay 2002
Formica neogagates Formica rubicunda
Formica neogagates Formica wheeleri Mackay and Mackay 2002
Formica neogagates Polyergus mexicanus Trager, 2013; de la Mora et al., 2021
Formica neorufibarbis Formica adamsi alpina Mackay and Mackay 2002
Formica neorufibarbis Formica adamsi Mackay and Mackay 2002
Formica neorufibarbis Formica aserva Wheeler and Wheeler 1986
Formica neorufibarbis Formica densiventris
Formica neorufibarbis Formica puberula Mackay & Mackay, 2002
Formica neorufibarbis Formica rubicunda
Formica neorufibarbis Formica wheeleri
Formica neorufibarbis Polyergus bicolor
Formica neorufibarbis Polyergus mexicanus Mackay and Mackay 2002; Trager, 2013; de la Mora et al., 2021 as ''Polyergus breviceps''
Formica obscuriventris Formica pergandei
Formica obscuriventris Formica rubicunda
Formica obtusopilosa Formica rubicunda
Formica occulta Formica puberula Bono et al., 2006; de la Mora et al., 2021
Formica occulta Polyergus breviceps Mackay and Mackay, 2002; Trager, 2013; de la Mora et al., 2021; Bono et al., 2006; Bono et al., 2007; Johnson, 2000; Johnson et al., 2005
Formica occulta Polyergus mexicanus Trager, 2013; de la Mora et al., 2021 as ''Formica'' cf. ''occulta''
Formica pacifica Polyergus mexicanus Trager, 2013; de la Mora et al., 2021
Formica pallidefulva Formica creightoni J. Trager (pers. comm.)
Formica pallidefulva Formica pergandei
Formica pallidefulva Formica puberula Mackay & Mackay, 2002
Formica pallidefulva Formica rubicunda
Formica pallidefulva Formica subintegra Mackay and Mackay, 2002
Formica pallidefulva Polyergus montivagus King & Trager, 2007; de la Mora et al., 2021 slave-maker misidentified as ''Polyergus lucidus'' in some publications
Formica pergandei Formica puberula Mackay & Mackay, 2002
Formica perpilosa Formica gynocrates S. Cover, MCZbase
Formica picea Formica sanguinea
Formica podzolica Formica aserva Ruano et al., 2019; Savolainen & Deslippe, 2001; de la Mora et al., 2021
Formica podzolica Formica pergandei
Formica podzolica Polyergus mexicanus Trager, 2013; de la Mora et al., 2021
Formica polyctena Formica sanguinea Seifert, 2018; de la Mora et al., 2021
Formica pratensis Formica sanguinea Ruano et al., 2019; Seifert, 2018; de la Mora et al., 2021 probable ephemeral host
Formica querquetulana Formica pergandei J. Trager, pers. comm.
Formica rufa Formica sanguinea Ruano et al., 2019; Seifert, 2018; de la Mora et al., 2021 probable ephemeral host
Formica rufibarbis Formica sanguinea
Formica rufibarbis Polyergus rufescens Mori et al., 2000; Trager, 2013; Seifert, 2018; de la Mora et al., 2021
Formica subaenescens Formica densiventris
Formica subaenescens Formica dirksi Mackay and Mackay 2002
Formica subaenescens Formica impexa
Formica subaenescens Formica pergandei
Formica subaenescens Formica rubicunda
Formica subaenescens Formica wheeleri
Formica subaenescens Polyergus bicolor Trager, 2013; de la Mora et al., 2021
Formica subaenescens Polyergus mexicanus
Formica subcyanea Polyergus topoffi
Formica subpolita Formica densiventris
Formica subpolita Formica puberula Mackay & Mackay, 2002
Formica subpolita Polyergus mexicanus
Formica subsericea Formica densiventris
Formica subsericea Formica pergandei
Formica subsericea Polyergus mexicanus Trager, 2013; de la Mora et al., 2021
Formica tianshanica Polyergus rufescens Seifert, 2018; de la Mora et al., 2021
Formica transkaucasica Formica sanguinea
Formica vinculans Formica gynocrates Snelling & Buren, 1985
Formica vinculans Polyergus mexicanus Trager, 2013; de la Mora et al., 2021
Leptothorax acervorum Harpagoxenus sublaevis Beibl et al., 2005; de la Mora et al., 2021 (and included references)
Leptothorax canadensis Harpagoxenus canadensis Beibl et al., 2005; de la Mora et al., 2021
Leptothorax gredleri Harpagoxenus sublaevis Foitzik et al., 2003; Guillem et al., 2014; de la Mora et al., 2021
Leptothorax muscorum Harpagoxenus canadensis Québec, Canada Smith, 1939
Leptothorax muscorum Harpagoxenus sublaevis 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
Myrmecocystus mimicus Myrmecocystus depilis Holldobler et al., 2011; Kronauer et al., 2003; de la Mora et al., 2021
Myrmica Formica rubicunda
Proformica epinotalis Rossomyrmex proformicarum D'Ettorre & Heinze, 2001; Ruano et al., 2013; de la Mora et al., 2021
Proformica korbi Rossomyrmex anatolicus Tinaut et al., 2010; Ruano et al., 2013; de la Mora et al., 2021
Proformica longiseta Rossomyrmex minuchae Ruano & Tinaut, 1999; D'Ettorre & Heinze, 2001; Errard et al., 2006; Ruano et al., 2013; de la Mora et al., 2021
Temnothorax affinis Temnothorax muellerianus Buschinger et al., 1988 rare host
Temnothorax affinis Temnothorax ravouxi southern Europe Czechowski & Czechowska, 2000
Temnothorax albipennis Temnothorax ravouxi Czechowski & Czechowska, 2000
Temnothorax ambiguus Temnothorax americanus
Temnothorax ambiguus Temnothorax duloticus
Temnothorax ambiguus Temnothorax pilagens
Temnothorax aveli Temnothorax kutteri Acosta Salmerón et al., 1983; Cagniant, 1973; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021 rare host
Temnothorax bernardi Temnothorax algerianus Buschinger et al., 1990
Temnothorax crasecundus Temnothorax tamarae Georgia Csösz et al., 2015
Temnothorax curtulus Temnothorax algerianus Buschinger et al., 1990
Temnothorax curvispinosus Temnothorax americanus
Temnothorax curvispinosus Temnothorax duloticus
Temnothorax exilis Temnothorax kutteri Acosta Salmerón et al., 1983; Espadaler & Restrepo, 1983; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021 fairly common host
Temnothorax exilis Temnothorax muellerianus Crete Salata, Borowiec & Trichas, 2018
Temnothorax flavicornis Temnothorax muellerianus Buschinger et al., 1988 rare host
Temnothorax gentilis Temnothorax algerianus Buschinger et al., 1990
Temnothorax lichtensteini Temnothorax gordiagini Buschinger et al., 1984
Temnothorax longispinosus Temnothorax americanus
Temnothorax longispinosus Temnothorax duloticus
Temnothorax longispinosus Temnothorax pilagens
Temnothorax luteus Temnothorax muellerianus Buschinger et al., 1988 as Leptothorax tristis
Temnothorax marocanus Temnothorax brunneus Morocco Buschinger & Douwes, 1990; Beibl et al., 2005; de la Mora et al., 2021
Temnothorax monjauzei Temnothorax algerianus Buschinger et al., 1990
Temnothorax nadigi Temnothorax ravouxi Poland Czechowski & Czechowska, 2000
Temnothorax niger Temnothorax kutteri Cagniant, 1973; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021 rare host
Temnothorax nigriceps Temnothorax muellerianus Buschinger et al., 1988 common host
Temnothorax nigriceps Temnothorax ravouxi Czechowski & Czechowska, 2000
Temnothorax nylanderi Temnothorax gordiagini Buschinger et al., 1983 ''T. nylanderi'' as the host of ''Temnothorax gordiagini'' had been suggested by Soudek, 1925, (cited in Buschinger et al., 1983) though this has never been confirmed and was probably a misidentification
Temnothorax racovitzai Temnothorax kutteri Buschinger et al., 1988; Cagniant, 1973; Beibl et al., 2007; de la Mora et al., 2021 most common host
Temnothorax recedens Temnothorax kraussei
Temnothorax recedens Temnothorax kutteri Espadaler & Restrepo, 1983; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021
Temnothorax recedens Temnothorax muellerianus Buschinger et al., 1988 common host
Temnothorax rottenbergii Temnothorax brunneus Morocco Buschinger & Douwes, 1990
Temnothorax semiruber Temnothorax muellerianus Crete Salata, Borowiec & Trichas, 2018
Temnothorax serviculus Temnothorax gordiagini Buschinger et al., 1984
Temnothorax spinosus Temnothorax algerianus Algeria, Morocco Buschinger et al., 1990 the most common host
Temnothorax spinosus Temnothorax tramieri Algeria, Morocco Cagniant, 1983
Temnothorax tuberum Temnothorax muellerianus Buschinger et al., 1988 rare host
Temnothorax tuberum Temnothorax ravouxi eastern Europe Czechowski & Czechowska, 2000
Temnothorax unifasciatus Temnothorax gordiagini Buschinger et al., 1984
Temnothorax unifasciatus Temnothorax kutteri Espadaler & Restrepo, 1983; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021 rare host
Temnothorax unifasciatus Temnothorax muellerianus Buschinger et al., 1988 common host
Temnothorax unifasciatus Temnothorax ravouxi mainly western Europe Czechowski & Czechowska, 2000
Temnothorax unifasciatus Temnothorax tamarae Georgia Gratiashvili et al., 2014 needs confirmation
Tetramorium alpestre Strongylognathus alpinus Seifert, 2018; de la Mora et al., 2021
Tetramorium alpestre Strongylognathus testaceus Seifert, 2018; de la Mora et al., 2021
Tetramorium caespitum Strongylognathus huberi Sanetra & Buschinger, 2000; Seifert, 2018; de la Mora et al., 2021
Tetramorium caespitum Strongylognathus italicus Seifert, 2018; de la Mora et al., 2021
Tetramorium caespitum Strongylognathus kratochvili Seifert, 2018; de la Mora et al., 2021
Tetramorium caespitum Strongylognathus testaceus
Tetramorium impurum Strongylognathus alpinus Sanetra & Buschinger, 2000; Wagner et al. 2017; Seifert, 2018; de la Mora et al., 2021
Tetramorium impurum Strongylognathus huberi Sanetra & Buschinger, 2000; Seifert, 2018; de la Mora et al., 2021
Tetramorium impurum Strongylognathus italicus Sanetra & Buschinger, 2000; de la Mora et al., 2021
Tetramorium impurum Strongylognathus testaceus Wagner et al., 2017
Tetramorium lucidulum Strongylognathus silvestrii Sanetra & Buschinger, 2000; de la Mora et al., 2021
Tetramorium moravicum Strongylognathus kratochvili Seifert, 2018; de la Mora et al., 2021
Tetramorium semilaeve Strongylognathus huberi Seifert, 2018; de la Mora et al., 2021
Tetramorium semilaeve Strongylognathus silvestrii Sanetra & Buschinger, 2000; de la Mora et al., 2021

References