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

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
Formica densiventris Formica argentea
Formica densiventris Formica fusca
Formica densiventris Formica neogagates
Formica densiventris Formica neorufibarbis
Formica densiventris Formica subpolita
Formica densiventris Formica subsericea
Formica pergandei Formica biophilica
Formica pergandei Formica difficilis
Formica pergandei Formica dolosa
Formica pergandei Formica fusca
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 subsericea
Formica rubicunda Formica altipetens
Formica rubicunda Formica argentea
Formica rubicunda Formica aserva
Formica rubicunda Formica bradleyi
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
Formica sanguinea Formica cinerea
Formica sanguinea Formica cunicularia
Formica sanguinea Formica fusca
Formica sanguinea Formica gagatoides
Formica sanguinea Formica lemani
Formica sanguinea Formica picea
Formica sanguinea Formica rufibarbis
Formica sanguinea Formica transkaucasica
Formica subintegra Formica pallidefulva Mackay and Mackay 2002
Formica wheeleri Creighton Formica altipetens
Formica wheeleri Creighton Formica bradleyi
Formica wheeleri Creighton Formica fusca
Formica wheeleri Creighton Formica lasioides
Formica wheeleri Creighton Formica lepida
Formica wheeleri Creighton Formica neogagates Mackay and Mackay, 2002
Formica wheeleri Creighton Formica neorufibarbis
Harpagoxenus canadensis Leptothorax muscorum
Harpagoxenus sublaevis Leptothorax acervorum
Harpagoxenus sublaevis Leptothorax gredleri Guillem et al., 2014
Harpagoxenus sublaevis Leptothorax muscorum
Polyergus bicolor Formica neorufibarbis
Polyergus bicolor Formica subaenescens
Polyergus breviceps Formica altipetens
Polyergus breviceps Formica canadensis
Polyergus breviceps Formica montana
Polyergus breviceps Formica neoclara
Polyergus breviceps Formica occulta Mackay and Mackay, 2002
Polyergus longicornis Formica dolosa
Polyergus lucidus Formica incerta
Polyergus mexicanus Formica accreta
Polyergus mexicanus Formica aerata
Polyergus mexicanus Formica argentea
Polyergus mexicanus Formica cf. occulta
Polyergus mexicanus Formica fusca
Polyergus mexicanus Formica hewitti
Polyergus mexicanus Formica manni
Polyergus mexicanus Formica microphthalma
Polyergus mexicanus Formica neoclara
Polyergus mexicanus Formica neogagates
Polyergus mexicanus Formica neorufibarbis Mackay and Mackay, 2002 as ''Polyergus breviceps''
Polyergus mexicanus Formica occulta
Polyergus mexicanus Formica pacifica
Polyergus mexicanus Formica podzolica
Polyergus mexicanus Formica subpolita
Polyergus mexicanus Formica subsericea
Polyergus mexicanus Formica vinculans
Polyergus montivagus Formica pallidefulva
Polyergus nigerrimus Formica candida
Polyergus nigerrimus Formica kozlovi
Polyergus oligergus Formica archboldi
Polyergus ruber Formica biophilica
Polyergus rufescens Formica cinerea
Polyergus rufescens Formica clara
Polyergus rufescens Formica cunicularia Romani et al., 2006
Polyergus rufescens Formica fusca
Polyergus rufescens Formica gagates
Polyergus rufescens Formica gerardi
Polyergus rufescens Formica rufibarbis
Polyergus samurai Formica fukaii
Polyergus samurai Formica glabridorsis
Polyergus samurai Formica hayashi
Polyergus samurai Formica japonica
Polyergus sanwaldi Formica dolosa Goodloe and Sanwald, 1985; Goodloe, Sanwald and Topoff, 1987
Polyergus topoffi Formica foreliana
Polyergus topoffi Formica gnava
Polyergus topoffi Formica subcyanea
Polyergus vinosus Formica moki
Rossomyrmex minuchae Proformica longiseta Ruano & Tinaut, 1999
Strongylognathus alpinus Tetramorium alpestre Seifert, 2018
Strongylognathus alpinus Tetramorium impurum Wagner et al. 2017; Seifert, 2018
Strongylognathus huberi Tetramorium caespitum Seifert, 2018
Strongylognathus huberi Tetramorium impurum Seifert, 2018
Strongylognathus huberi Tetramorium semilaeve Seifert, 2018
Strongylognathus italicus Tetramorium caespitum Seifert, 2018
Strongylognathus kratochvili Tetramorium caespitum Seifert, 2018
Strongylognathus kratochvili Tetramorium moravicum Seifert, 2018
Strongylognathus testaceus Tetramorium alpestre Seifert, 2018
Strongylognathus testaceus Tetramorium caespitum
Strongylognathus testaceus Tetramorium impurum Wagner et al. 2017
Temnothorax algerianus Temnothorax curtulus gentilis Buschinger et al., 1990
Temnothorax algerianus Temnothorax curtulus 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 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 kutteri Temnothorax aveli Acosta Salmerón et al., 1983 as rabaudi, rare host
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 Espadaler and Restrepo, 1983 as specularis, fairly common host
Temnothorax kutteri Temnothorax niger Cagniant, 1973 rare host
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
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

Known Hosts of Slave-Makers

Host Slave Maker Locality Source Notes
Formica accreta Polyergus mexicanus
Formica aerata Polyergus mexicanus
Formica altipetens Formica rubicunda
Formica altipetens Formica wheeleri
Formica altipetens Polyergus breviceps
Formica archboldi Polyergus oligergus
Formica argentea Formica densiventris
Formica argentea Formica rubicunda
Formica argentea Polyergus mexicanus
Formica aserva Formica rubicunda
Formica biophilica Formica densiventris
Formica biophilica Polyergus ruber
Formica bradleyi Formica rubicunda
Formica bradleyi Formica wheeleri
Formica canadensis Polyergus breviceps
Formica candida Polyergus nigerrimus
Formica cinerea Formica sanguinea
Formica cinerea Polyergus rufescens
Formica clara Polyergus rufescens
Formica cunicularia Formica sanguinea
Formica cunicularia Polyergus rufescens Romani et al., 2006
Formica difficilis Formica pergandei
Formica dolosa Formica pergandei
Formica dolosa Polyergus longicornis
Formica dolosa Polyergus sanwaldi Goodloe and Sanwald, 1985; Goodloe, Sanwald and Topoff, 1987
Formica foreliana Polyergus topoffi
Formica fossaceps Formica rubicunda
Formica fukaii Polyergus samurai a relatively rare host
Formica fusca Formica sanguinea
Formica fusca Polyergus rufescens
Formica gagates Polyergus rufescens
Formica gagatoides Formica sanguinea
Formica gerardi Polyergus rufescens
Formica glabridorsis Polyergus samurai
Formica gnava Polyergus topoffi
Formica hayashi Polyergus samurai a relatively rare host
Formica hewitti Polyergus mexicanus
Formica incerta Formica pergandei
Formica incerta Polyergus lucidus
Formica japonica Polyergus samurai
Formica kozlovi Polyergus nigerrimus
Formica lasioides Formica rubicunda
Formica lasioides Formica wheeleri
Formica lemani Formica sanguinea
Formica lepida Formica rubicunda
Formica lepida Formica wheeleri
Formica manni Polyergus mexicanus
Formica microphthalma Polyergus mexicanus
Formica moki Polyergus vinosus
Formica montana Formica rubicunda
Formica montana Polyergus breviceps
Formica neoclara Formica rubicunda
Formica neoclara Polyergus breviceps
Formica neoclara Polyergus mexicanus
Formica neogagates Formica densiventris Mackay and Mackay 2002
Formica neogagates Formica pergandei Mackay and Mackay 2002
Formica neogagates Formica rubicunda
Formica neogagates Formica wheeleri Mackay and Mackay 2002
Formica neogagates Polyergus mexicanus
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 rubicunda
Formica neorufibarbis Formica wheeleri
Formica neorufibarbis Polyergus bicolor
Formica neorufibarbis Polyergus mexicanus Mackay and Mackay 2002 as ''Polyergus breviceps''
Formica obscuriventris Formica pergandei
Formica obscuriventris Formica rubicunda
Formica obtusopilosa Formica rubicunda
Formica occulta Polyergus breviceps Mackay and Mackay, 2002
Formica occulta Polyergus mexicanus
Formica pacifica Polyergus mexicanus
Formica pallidefulva Formica pergandei
Formica pallidefulva Formica rubicunda
Formica pallidefulva Formica subintegra Mackay and Mackay, 2002
Formica pallidefulva Polyergus montivagus
Formica picea Formica sanguinea
Formica podzolica Formica aserva
Formica podzolica Formica pergandei
Formica podzolica Polyergus mexicanus
Formica rufibarbis Formica sanguinea
Formica rufibarbis Polyergus rufescens
Formica subaenescens Formica densiventris
Formica subaenescens Formica dirksi Mackay and Mackay 2002
Formica subaenescens Formica pergandei
Formica subaenescens Formica rubicunda
Formica subaenescens Formica wheeleri
Formica subaenescens Polyergus bicolor
Formica subaenescens Polyergus mexicanus
Formica subcyanea Polyergus topoffi
Formica subpolita Formica densiventris
Formica subpolita Polyergus mexicanus
Formica subsericea Formica densiventris
Formica subsericea Formica pergandei
Formica subsericea Polyergus mexicanus
Formica transkaucasica Formica sanguinea
Formica vinculans Polyergus mexicanus
Leptothorax acervorum Harpagoxenus sublaevis
Leptothorax canadensis Harpagoxenus canadensis
Leptothorax gredleri Harpagoxenus sublaevis Guillem et al., 2014
Leptothorax muscorum Harpagoxenus sublaevis
Myrmica Formica rubicunda
Proformica longiseta Rossomyrmex minuchae
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 rare host
Temnothorax bernardi Temnothorax algerianus Buschinger et al., 1990
Temnothorax crasecundus Temnothorax tamarae Georgia Csösz et al., 2015
Temnothorax curtulus gentilis Temnothorax algerianus Buschinger et al., 1990
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 fairly common host
Temnothorax exilis Temnothorax muellerianus Crete Salata, Borowiec & Trichas, 2018
Temnothorax flavicornis Temnothorax muellerianus Buschinger et al., 1988 rare host
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 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 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., 1984
Temnothorax racovitzai Temnothorax kutteri Buschinger et al., 1988; Cagniant, 1973 most common host
Temnothorax recedens Temnothorax kutteri Espadaler & Restrepo, 1983; Buschinger et al., 1988
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 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 caespitum Strongylognathus testaceus
Tetramorium impurum Strongylognathus alpinus Wagner et al. 2017
Tetramorium impurum Strongylognathus testaceus Wagner et al., 2017

References

  • Alloway, T. 1980. The origins of slavery in leptothoracine ants (Hymenoptera: Formicidae). The American Naturalist 115(2): 247-261.
  • Buschinger, A. 1990. Sympatric speciation and radiative evolution of socially parasitic ants ‐ Heretic hypotheses and their factual background. Journal of Zoological Systematics and Evolutionary Research 28: 241-260 (DOI 10.1111/j.1439-0469.1990.tb00379.x).
  • Buschinger, A., Ehrhardt, W. & Winter, U. 1980. The organization of slave raids in dulotic ants — A comparative study (Hymenoptera; Formicidae). Zeitschrift für Tierpsychologie 53: 245-264. (DOI 10.1111/j.1439-0310.1980.tb01053.x).
  • Ruano, F., Sanllorente, O., Lenoir, A. & Tinaut, A. 2013. Rossomyrmex, the slave-maker ants from the arid steppe environments. Psyche 2013: Article ID 541804 (DOI 10.1155/2013/541804).
  • Stuart, R.J., Alloway, T.M. 1985. Behavioural evolution and domestic degeneration in obligatory slave-making ants (Hymenoptera: Formicidae: Leptothoracini).

Animal Behaviour 33: 1080-1088 (DOI 10.1016/S0003-3472(85)80166-4).