Social Parasitism

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Social parasitism is the coexistence of two or more ant species in one nest or colony. It involves a parasitic species which is dependent on one or several host species. The relationship can be obligatory or facultative, permanent or temporary. These relationships can take many forms and have been classified in various ways.

Holldobler & Wilson (1990), following a suggestion of Wasmann (1891), distinguished between "compound nests" and "mixed colonies".

  • Compound nests (xenobiosis) involve two species of ants living together in the same nest but keeping their broods separate. One species lives in the walls or chambers of the nests of the other and moves freely among its hosts, obtaining food from them by one means or another, usually by soliciting regurgitation. The details of specific relationships vary with the species involved, and can broadly be classified into a series of essentially continuous types.
    • Plesiobiosis. In this most rudimentary association, different ant species nest very close to each other, but engage in little or no direct communication.
    • Cleptobiosis. Some species of small ants build nests near those of larger species and either feed on refuse in the host kitchen middens or rob the host workers when they return home carrying food.
    • Lestobiosis. Certain small species, most belonging to Solenopsis and related genera, stay in the walls of large nests built by other ants or termites and enter the nest chambers of their hosts to steal food and prey on the inhabitants.
    • Parabiosis. In this peculiar form of symbiosis, two or more species use the same nest and sometimes even the same odour trails, but they keep their brood separate.
  • Mixed colonies comprise temporary parasites, slave-makers (dulosis) and inquilines, where the host workers care for the parasite brood, at least temporarily.

As an alternative, Buschinger (2009), in his review of social parasitism, proposed the following classification:

  • Xenobiosis (guest ants, sometimes called cleptobiosis or kleptobiosis) - The biology of Formicoxenus nitidulus provides natural history information about one representative guest ant.
  • Temporary parasitism (occurring together only during colony foundation) - The biology of Lasius umbratus provides natural history information about one representative temporarily parasitic ant.
  • Permanent parasitism with slavery (dulosis) - The biology of Temnothorax muellerianus and Polyergus rufescens provide representative accounts of dulosis. Note that slave-making species range from being largely self-sufficient and easily able to survive without the support of slaves (facultative dulosis, as in Formica sanguinea), to having a very limited behavioral repertory and apparently completely helpless without their slaves (referred to as degenerate slavemakers) (Holldobler & Wilson, 1990).

"Obligatory slavemakers" are completely and permanently dependent on regular replenishment of their slave stock by raiding host species colonies. The term "degenerate slavemaker" has been coined for species that belong to a monophylum of obligatory slavemaking species but have secondarily abandoned the slavemaker worker caste, or reduced its number, hence can‘t conduct slave raids. Examples are Temnothorax kraussei (reduced slavemaker workers to a few, or zero), Temnothorax corsicus and Temnothorax adlerzi (both workerless; all three belong to former genus Myrmoxenus), and Temnothorax brunneus (workerless, former genus Chalepoxenus). The distinction is necessary since the queens of the degenerate slavemakers have retained the colony foundating behavior of their actively dulotic relatives, i.e. kill the host coloy queen by throttling her to death (former Myrmoxenus) or stinging (former Chalepoxenus). This is different from workerless inquilines whose queens live in queenright host species colonies, and who most probably derive directly from an independent species (example Leptothorax kutteri and Leptothorax pacis with Leptothorax acervorum as host species).

  • Permanent parasitism without slavery (inquilinism) - The biology of Tetramorium inquilinum (as Teleutomyrmex schneideri) provides natural history information about one representative inquiline ant.

Parasite Evolution

Understanding how ants evolved to be parasites of other ants is an important, long-standing question.Hölldoble rand Wilson (1990) stated the following "Social parasitism in ants is complicated . . . the source of the complexity is first the large number of ant species that have entered into some form of parasitic relationship with each other. Second, at least two and possibly three major evolutionary routes lead to the ultimate stage of permanent, workerless parasitism. Finally, no two species are exactly alike in the details of their parasitic adaptation."

This section needs to be expanded!

Chemical Deception Among Social Parasites

Guillem et al. (2014) examined cuticular hydrocarbon (CHC) profiles among the parasites Camponotus universitatis, Harpagoxenus sublaevis‎‎ and Strongylognathus testaceus and their hosts. They found that the parasitic species had CHC profiles that were indistinguishable from that of their hosts, even when the parasite is using more than one host species. 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.

They also noted that in some species, for example Harpagoxenus sublaevis (Winter and Buschinger, 1986), raiding workers are frequently killed or driven off when trying to raid or invade new host colonies, 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)).

Parasitic Ant Species

This information has been modified from Buschinger (2009). Please cite the original publication as the source for these data.

dDul = degenerate dulosis; Dul = dulosis, slave-maker; In = inquilinism; Tp = temporary parasitism; Xen = xenobiosis, guest ant.

Sortable table
Genus / Species Reference Subfamily Type Range Remarks
Acromyrmex ameliae De Souza, Soares & Della Lucia, 2007 Myrmicinae In Brazil social parasite with workers
Acromyrmex insinuator Schultz, Bekkevold & Boomsma, 1998 Myrmicinae In Panama "Incipient" social parasite
Bothriomyrmex Emery, 1869 Dolichoderinae Tp Old World, Australia Ca. 38 species + many subspecies
Bothriomyrmex decapitans Santschi, 1911 Dolichoderinae Tp N-Africa
Camponotus universitatis Forel, 1890 Formicinae In S-Europe
Cataglyphis hannae Agosti, 1994 Formicinae In N-Africa
Ectatomma parasiticum Feitosa and Fresneau, 2008 Ectatomminae In Mexico Host E. tuberculatum
Formica dirksi Wing, 1949 Formicinae In N-America
Formica rufa Linnaeus, 1761 Formicinae Tp Eurasia Facultative Tp
Formica talbotae Wilson, 1977 Formicinae In N-America
Formica (Coptoformica) spp. Müller, 1923 Formicinae Tp Eurasia Facultative Tp
Formica (Raptiformica) sanguinea Latreille, 1798 Formicinae Dul Eurasia Facultative Dul
Formicoxenus Mayr, 1855 Myrmicinae, Formicoxenini Xen holarctic 7 species
Formicoxenus diversipilosus (Smith, M.R. 1939) Myrmicinae, Formicoxenini Xen N-America worker-queen intermediates
Formicoxenus nitidulus (Nylander, 1846) Myrmicinae, Formicoxenini Xen Eurasia
Formicoxenus provancheri (Emery, 1895) Myrmicinae, Formicoxenini Xen N-America
Formicoxenus quebecensis Francoeur, 1985 Myrmicinae, Formicoxenini Xen N-America
Harpagoxenus canadensis Smith, 1939 Myrmicinae, Formicoxenini Dul N-America
Harpagoxenus sublaevis (Nylander, 1849) Myrmicinae, Formicoxenini Dul Eurasia queen polymorphism
Lasius fuliginosus (Latreille, 1798) Formicinae Tp Eurasia Hyperparasite
Lasius reginae Faber, 1967 Formicinae Tp Europe
Lasius umbratus (Nylander, 1846) Formicinae Tp Eurasia
Leptothorax faberi Buschinger, 1983 Myrmicinae, Formicoxenini In Canada possible inquiline
Leptothorax goesswaldi Kutter, 1967 Myrmicinae, Formicoxenini In Europe Host-queen-intolerant
Leptothorax kutteri Buschinger, 1966 Myrmicinae, Formicoxenini In Europe Host-queen-tolerant
Leptothorax pacis (Kutter, 1945) Myrmicinae, Formicoxenini In Europe Host-queen-tolerant
Leptothorax paraxenus Heinze and Alloway, 1992 Myrmicinae, Formicoxenini In N-America
Leptothorax wilsoni Heinze , 1989 Myrmicinae, Formicoxenini In N-America
Megalomyrmex mondabora Brandão, 1990 Myrmicinae * Central and S-America Agro-Predation
Megalomyrmex symmetochus Wheeler, 1925 Myrmicinae Xen? Panama
Monomorium inquilinum DuBois, 1981 Myrmicinae ? Mexico
Monomorium pergandei (Emery, 1893) Myrmicinae In USA
Monomorium talbotae DuBois, 1975 Myrmicinae In USA, Michigan
Myrmecocystus spp. Wesael, 1838 Formicinae Dul? N-America Intraspecific!
Myrmica hirsuta Elmes, 1978 Myrmicinae In Europe
Myrmica karavajevi (Arnol'di, 1930) Myrmicinae In Eurasia
Myrmica laurae (Emery, 1907) Myrmicinae In Europe
Myrmica lemasnei Bernard, 1967 Myrmicinae In Europe
Myrmica myrmicoxena Forel, 1895 Myrmicinae Europe
Myrmica nefaria Bharti, 2012 xxxx Myrmicinae ? Europe
Myrmica semiparasitica Francoeur, 2007 Myrmicinae Tp Canada Not confirmed
Myrmica vandeli Bondroit, 1920 Myrmicinae Tp? Europe Not confirmed
Nylanderia deceptrix Messer, Cover & LaPolla, 2016 Formicinae In North America
Plagiolepis spp. Mayr, 1861 Formicinae In Europe, N-Africa Many independent
Plagiolepis ampeloni (Faber, 1969) Formicinae In S-Europe (= Aporomyrmex FABER, 1969)
Plagiolepis grassei Le Masne, 1956 Formicinae In S-Europe Few own workers
Plagiolepis regis Karavaiev, 1931 Formicinae In Turkestan (= Paraplagiolepis FABER, 1969)
Plagiolepis xene Staercke, 1936 Formicinae In S-Europe (= Paraplagiolepis FABER, 1969)
Pogonomyrmex anergismus Cole, 1954 Mymicinae In N-America
Pogonomyrmex colei Snelling, 1982 Mymicinae In N-America
Polyergus Latreille, 1804 Formicinae Dul Holarctic 6 spp., several sspp.
Polyergus breviceps Emery, 1893 Formicinae Dul N-America
Polyergus lucidus Mayr, 1870 Formicinae Dul N-America
Polyergus nigerrimus Marikovsky, 1963 Formicinae Dul E-Asia
Polyergus rufescens (Latreille, 1798) Formicinae Dul Europe
Polyergus samurai Yano, 1911 Formicinae Dul Japan
Polyrhachis lama Kohout, 1994 Formicinae Xen SE-Asia
Polyrhachis lamellidens Smith, 1874 Formicinae Tp? SE-Asia ??
Polyrhachis loweryi Kohout, 1990 Formicinae Xen Australia
Pheidole spp. Westwood, 1839 Myrmicinae In World-wide Many independent
Pseudoatta argentina Gallardo, 1916 Myrmicinae In S-America
Pseudomyrmex inquilinus Ward, 1996 Pseudomyrmecinae In S-America
Rossomyrmex minuchae Tinaut, 1981 Formicinae Dul Spain, Turkey
Rossomyrmex proformicarum Arnol'di, 1928 Formicinae Dul Central to East Asia
Solenopsis enigmatica Deyrup and Prusak, 2008 Myrmicinae Tp? West Indies Not confirmed
Solenopsis fugax Latreille, 1798 Myrmicinae Facultative Xen Eurasia Xenobiotics colonies are common.
Strongylognathus spp. Mayr, 1853 Myrmicinae Dul, dDul Eurasia Ca. 25 species
Strongylognathus afer Emery, 1884 Myrmicinae Dul N-Africa
Strongylognathus karawajewi Pisarski, 1966 Myrmicinae dDul Europe Host-queen-tolerant
Strongylognathus minutus Radchenko, 1991 Myrmicinae Dul? Europe
Strongylognathus pisarskii Poldi, 1994 Myrmicinae Dul? Europe
Strongylognathus potanini Radchenko, 1995 Myrmicinae Dul? China
Strongylognathus testaceus (Schenck, 1852) Myrmicinae dDul Europe Host-queen-tolerant
Strongylognathus tylonus Wei, Xu and He, 2001 Myrmicinae Dul? China
Temnothorax spp. (= Epimyrma, = Myrmoxenus Ruzsky, 1902) Myrmicinae, Formicoxenini Dul, dDul Europe, N-Africa Ca. 12 species
Temnothorax adlerzi (Douwes, Jessen and Buschinger, 1988) Myrmicinae, Formicoxenini dDul Greece Workerless (= Myrmoxenus)
Temnothorax algerianus (Cagniant, 1968) Myrmicinae, Formicoxenini Dul N-Africa (= Myrmoxenus)
Temnothorax bernardi (Espadaler, 1982) Myrmicinae, Formicoxenini Dul S-Europe (= Myrmoxenus)
Temnothorax birgitae (Schulz, 1994) Myrmicinae, Formicoxenini dDul Tenerife Workerless (= Myrmoxenus)
Temnothorax corsicus (Emery, 1895) Myrmicinae, Formicoxenini dDul S-Europe Workerless (= Myrmoxenus)
Temnothorax gordiagini (Ruzsky, 1902) Myrmicinae, Formicoxenini Dul SE-Europe (= Myrmoxenus)
Temnothorax kraussei (Emery, 1915) Myrmicinae, Formicoxenini dDul S-Europe Workers few to none (= Myrmoxenus)
Temnothorax ravouxi (André, 1896) Myrmicinae, Formicoxenini Dul Europe (= Myrmoxenus)
Temnothorax spp. (= Chalepoxenus Menozzi, 1923) Myrmicinae, Formicoxenini Dul, dDul S-Europe to Turkmenistan Ca. 9 species
Temnothorax americanus (Emery, 1895) Myrmicinae, Formicoxenini Dul N-America (= Protomognathus)
Temnothorax brunneus Cagniant, 1985 Myrmicinae, Formicoxenini dDul N-Africa Workerless (= Chalepoxenus)
Temnothorax duloticus (Wesson, 1937) Myrmicinae, Formicoxenini Dul N-America
Temnothorax kutteri Cagniant, 1973 Myrmicinae, Formicoxenini Dul S-Europe (= Chalepoxenus)
Temnothorax minutissimus (Smith, 1942) Myrmicinae, Formicoxenini In N-America Host-queen-tolerant
Temnothorax muellerianus (Finzi, 1922) Myrmicinae, Formicoxenini Dul S-Europe (= Chalepoxenus)
Temnothorax pilagens Seifert et al., 2014 Myrmicinae, Formicoxenini Dul N-America
Tetramorium atratulum (Schenck, 1852) Myrmicinae In Europe Queen-intolerant (= Anergates atratulus)
Tetramorium buschingeri (Lapeva-Gjonova, 2017) Myrmicinae In Bulgaria (= Teleutomyrmex)
Tetramorium inquilinum Ward, Brady, Fisher & Schultz, 2014 Myrmicinae In Europe Host-queen-tolerant (= Teleutomyrmex schneideri Kutter, 1950)
Tetramorium kutteri (Tinaut, 1990) Myrmicinae In S-Europe Host-queen-tolerant (= Teleutomyrmex)
Tetramorium microgyna Bolton 1980 Myrmicinae In southern Africa
Tetramorium seiferti (Kiran and Karaman, 2017) Myrmicinae In Turkey (= Teleutomyrmex)
Vollenhovia nipponica Kinomura and Yamauchi, 1992 Myrmicinae In Japan