Xenobiosis

Xenobiosis, also referred to as compound nests or guest ants, occurs when two species of ants live together in the same nest but keep their broods separate and they show minimal actual interaction with each other. Details of the relationship among the partners in a xenobiotic nest vary considerably. In some cases, the species involved are essentially equal partners with little indication of either species being a "guest" or a "host" (parabiosis and plesiobiosis). In other cases, the guest species form small colonies (up to a hundred individuals) in the nesting material or nest walls of the host and are normally much smaller than their host (cleptobiosis and lestobiosis). The guest maintains its nest well separated from the brood chambers of their host and they tend their own brood. They are dependent upon the hosts for nutrition and often shelter and some xenobiotic species may solicit regurgitated food directly from a host worker or participate in food exchange (trophallaxis) between two host workers.

The biology of Formicoxenus nitidulus provides natural history information about one representative guest ant.

Xenobiotic species generally fall into one of 4 subtypes:

• Cleptobiosis: stealing food from other species; a mutual relation in which members of one species habitually steal food from another.
• Lestobiosis: stealing brood; cleptobiosis in which covert thievery replaces aggressive plundering.
• Parabiosis: sharing nests and trails.
• Plesiobiosis: casual or regular nesting in close vicinity; casual association of two or more colonies.

Plesiobiosis

Kanizsai et al. (2013) - Plesiobiosis, the most basic form of interspecific associations in ants, denotes occasional or regular nesting of heterospecific colonies of certain species pairs in close proximity to each other without biological interdependence. Plesiobionts differ from each other both in morphology and in behaviour (e.g., in their foraging strategies), and at least one of the plesiobiotic pair is a submissive species. Recent studies on plesiobiosis have revealed that Formica fusca and Lasius flavus are two of the most frequent plesiobionts. To date, at least 48 different plesiobiotic species pairs have been recorded from various habitat types of the Holarctic region. Two main habitat properties may play a role in the forming of plesiobiosis: the scarcity of suitable nesting sites as a forcing factor and the suffcient amount of food sources available, influencing the abundance of colonies. Thus, high colony density may contribute to the formation of such associations, resulting in (1) frequent nesting in each other’s neighbourhood and (2) stronger intraspecific competition, which forces colonies into the vicinity of heterospecific nests. Plesiobiotic associations formed this way may promote persistent coexistence, leading to the formation of other types of interspecific associations (e.g., clepto- or lestobiosis).

Known Xenobiotic Ants

xenobiont	Host	Locality	Source	Notes
Aphaenogaster subterranea	Camponotus ligniperda	Hungary	Kanizsai et al., 2013	Pine forest. Under stone.
Camponotus fallax	Lasius brunneus	Poland	Czechowski, 2004; Kanizsai et al., 2013	Urban park. In decaying wood.
Camponotus femoratus	Solenopsis picea
Camponotus festinatus	Monomorium minimum	United States	Kanizsai et al., 2013; Wheeler, 1901
Camponotus festinatus	Pachycondyla harpax	United States	Kanizsai et al., 2013; Wheeler, 1901	At road. Under stone.
Camponotus herculeanus	Formica fusca	Poland	Czechowski, 2005; Kanizsai et al., 2013	Forest edge. Under wood
Camponotus herculeanus	Lasius platythorax	Finland	Czechowski, 2004; Kanizsai et al., 2013	In decaying wood.
Camponotus ligniperda	Aphaenogaster subterranea	Hungary	Kanizsai et al., 2013	Pine forest. Under stone.
Camponotus modoc	Leptothorax calderoni	United States	Kanizsai et al., 2013; Wheeler, 1917	Pine forest. In log/in stump.
Camponotus pennsylvanicus	Formica subaenescens	United States	Gaige ,1914; Kanizsai et al., 2013	Hardwood forest. Under log.
Camponotus planatus	Tapinoma litorale
Camponotus sansabeanus	Monomorium minimum	United States	Kanizsai et al., 2013; Wheeler, 1901
Camponotus sansabeanus	Pachycondyla harpax	United States	Kanizsai et al., 2013; Wheeler, 1901
Camponotus vagus	Formica fusca	Hungary	Kanizsai et al., 2013	Pine and poplar forest patches. In/under wood
Camponotus yogi	Temnothorax andrei	United States	Creighton & Snelling, 1966; Kanizsai et al., 2013	Chaparral. In living stems of ''Haplopappus pinifolius''.
Cephalotes specularis	Crematogaster ampla	Brazil	Brandao et al., 2014; Powell et al., 2014
Cephalotes varians	Tapinoma litorale
Crematogaster ampla	Cephalotes specularis	Brazil	Brandao et al., 2014; Powell et al., 2014
Crematogaster carinata	Dolichoderus bispinosus
Crematogaster carinata	Dolichoderus debilis
Crematogaster carinata	Dolichoderus inermis
Crematogaster carinata	Odontomachus panamensis
Crematogaster carinata	Solenopsis picea
Crematogaster carinata	Temnothorax subditivus
Dolichoderus bispinosus	Crematogaster carinata
Dolichoderus debilis	Crematogaster carinata
Dolichoderus inermis	Crematogaster carinata
Formica	Strumigenys pergandei	United States	Kanizsai et al., 2013; Wheeler, 1905	In soil
Formica ciliata	Formicoxenus hirticornis
Formica clara	Formicoxenus sibiricus		Seifert, 2023
Formica cunicularia	Lasius flavus	United Kingdom	Kanizsai et al., 2013; Morley, 1945	Foreshore. Under stone
Formica exsecta	Formicoxenus nitidulus		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formica fusca	Camponotus herculeanus	Poland	Czechowski, 2005; Kanizsai et al., 2013	Forest edge. Under wood
Formica fusca	Camponotus vagus	Hungary	Kanizsai et al., 2013	Pine and poplar forest patches. In/under wood
Formica fusca	Formica aquilonia	Finland	Czechowski & Vepsalainen, 1999; Kanizsai et al., 2013	Forest on rocks. Mound of ''F. lugubris''
Formica fusca	Formica lugubris	Finland	Czechowski, 2004; Kanizsai et al., 2013	Different successional series of rocky habitats. Mound of ''F. lugubris''
Formica fusca	Formica truncorum	Finland	Czechowski, 2004; Kanizsai et al., 2013	Different successional series of rocky habitats. In rock crevice
Formica fusca	Lasius flavus	Finland	Czechowski, 2004; Kanizsai et al., 2013	In different successional series of rocky habitats. Under wood; in rock crevice; under stone
Formica fusca	Lasius flavus	Italy	Balzani et al., 2022
Formica fusca	Lasius niger	United Kingdom	Kanizsai et al., 2013; Morley, 1945	under stone
Formica fusca	Lasius platythorax	Finland, Poland	Kanizsai et al., 2013; Włodarczyk et al., 2009	Forest on rocks; clearcut of managed forest. In decaying wood; in tree stump
Formica fusca	Leptothorax acervorum	Finland	Czechowski, 2004; Kanizsai et al., 2013	In different successional series of rocky habitats, in mound of ''Formica lugubris''
Formica fusca	Myrmica rubra	Finland, United Kingdom	Czechowski, W. 2003; Czechowski, 2004; Kanizsai et al., 2013; Morley, 1945	habitat foreshore, nests in decaying wood, under stone
Formica fusca	Myrmica ruginodis	Poland	Kanizsai et al., 2013; Włodarczyk et al., 2009	Clearcut of managed forest, in tree stumps
Formica fusca	Tetramorium caespitum	Poland	Kanizsai et al., 2013; Włodarczyk et al., 2009	Clearcut of managed forest, in tree stumps
Formica fuscocinerea	Lasius flavus	Poland	Czechowski & Czechowska, 2000; Kanizsai et al., 2013	Grassy mountain slope. Under stone
Formica gnava	Monomorium minimum	United States	Kanizsai et al., 2013
Formica gnava	Myrmecina americana	United States	Kanizsai et al., 2013; Wheeler, 1901
Formica integroides	Formicoxenus hirticornis
Formica lemani	Formica japonica		Yamaoka, 1977; Kvifte and Soule, 2017	plesiobiotic
Formica lugubris	Formica fusca	Finland	Czechowski, 2004; Kanizsai et al., 2013	Different successional series of rocky habitats. Mound of ''F. lugubris''
Formica lugubris	Formicoxenus nitidulus		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formica moki	Temnothorax andrei		Mann, 1911	nature of relationship unknown
Formica obscuripes	Formicoxenus diversipilosus
Formica obscuripes	Formicoxenus hirticornis
Formica occulta	Lasius latipes
Formica occulta	Lasius sitiens
Formica occulta	Myrmica hamulata
Formica pisarskii	Formicoxenus nitidulus		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formica pisarskii	Formicoxenus sibiricus		Seifert, 2023	misidentified as ''Formica pressilabris''
Formica polyctena	Formicoxenus nitidulus		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formica pratensis	Formicoxenus nitidulus		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formica ravida	Formicoxenus hirticornis
Formica ravida	Temnothorax andrei		Mackay & Mackay, 1984	nature of relationship unknown
Formica rufa	Formicoxenus nitidulus		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formica rufa	Leptothorax muscorum	Sweden	Kanizsai et al., 2013; Wheeler, 1901
Formica subaenescens	Camponotus pennsylvanicus	United States	Gaige ,1914; Kanizsai et al., 2013	Hardwood forest. Under log.
Formica truncorum	Formica fusca	Finland	Czechowski, 2004; Kanizsai et al., 2013	Different successional series of rocky habitats. In rock crevice
Formica truncorum	Formicoxenus nitidulus		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formica uralensis	Formicoxenus nitidulus		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formicoxenus chamberlini	Manica invidia		Wheeler (1904)
Formicoxenus diversipilosus	Formica obscuripes
Formicoxenus hirticornis	Formica ciliata
Formicoxenus hirticornis	Formica integroides			needs confirmation
Formicoxenus hirticornis	Formica obscuripes
Formicoxenus hirticornis	Formica ravida
Formicoxenus nitidulus	Formica aquilonia		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formicoxenus nitidulus	Formica exsecta		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formicoxenus nitidulus	Formica lugubris		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formicoxenus nitidulus	Formica pisarskii		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formicoxenus nitidulus	Formica polyctena		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formicoxenus nitidulus	Formica pratensis		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formicoxenus nitidulus	Formica rufa		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formicoxenus nitidulus	Formica truncorum		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formicoxenus nitidulus	Formica uralensis		Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formicoxenus nitidulus	Polyergus rufescens		Wilson 1971; Martin et al. 2007
Formicoxenus nitidulus	Tetramorium caespitum		Wilson 1971; Martin et al. 2007
Formicoxenus provancheri	Myrmica fracticornis
Formicoxenus provancheri	Myrmica incompleta
Formicoxenus sibiricus	Formica clara		Seifert, 2023
Formicoxenus sibiricus	Formica pisarskii		Seifert, 2023	misidentified as ''Formica pressilabris''
Lasius brunneus	Camponotus fallax	Poland	Czechowski, 2004; Kanizsai et al., 2013	Urban park. In decaying wood.
Lasius crypticus	Monomorium minimum		Mackay & Mackay, 2002
Lasius flavus	Formica aquilonia	Finland	Czechowski, 2004; Kanizsai et al., 2013	Different successional series of rocky habitats. Mound of ''Formica aquilonia''.
Lasius flavus	Formica cunicularia	United Kingdom	Kanizsai et al., 2013; Morley, 1945	Foreshore. Under stone
Lasius flavus	Formica fusca	Finland	Czechowski, 2004; Kanizsai et al., 2013	Different successional series of rocky habitats. In rock crevice
Lasius flavus	Formica fusca	Italy	Balzani et al., 2022
Lasius flavus	Formica fuscocinerea	Poland	Czechowski & Czechowska, 2000; Kanizsai et al., 2013	Grassy mountain slope. Under stone.
Lasius flavus	Lasius niger	Finland, United Kingdom	Czechowski, 2004; Kanizsai et al., 2013; Morley, 1945	Rocky outcrop; shore meadow, foreshore. In rock crevice/under stone.
Lasius flavus	Lasius platythorax	Finland	Czechowski, 2004; Kanizsai et al., 2013	Different successional series of rocky habitats. In rock crevice/under stone/overgrown soil.
Lasius flavus	Myrmica ruginodis	Italy	Balzani et al., 2022
Lasius flavus	Myrmica scabrinodis	United Kingdom	Kanizsai et al., 2013; Morley, 1945	Foreshore. Under stone.
Lasius flavus	Tetramorium caespitum	Finland	Czechowski, 2004; Kanizsai et al., 2013	Different successional series of rocky habitats. Under stone.
Lasius latipes	Formica occulta		Mackay & Mackay, 2002
Lasius latipes	Lasius sitiens		Mackay & Mackay, 2002
Lasius latipes	Myrmica hamulata		Mackay & Mackay, 2002
Lasius minutus	Pheidole picea	United States	Gaige ,1914; Kanizsai et al., 2013	Hardwood forest. In a stump.
Lasius nearcticus	Pheidole picea	United States	Gaige ,1914; Kanizsai et al., 2013	Hardwood forest. Under rock.
Lasius niger	Formica fusca	United Kingdom	Kanizsai et al., 2013; Morley, 1945	under stone
Lasius niger	Lasius flavus	Finland, United Kingdom	Czechowski, 2004; Kanizsai et al., 2013; Morley, 1945	Rocky outcrop; shore meadow, foreshore. In rock crevice/under stone.
Lasius niger	Myrmica rubra	Finland	Czechowski, 2004; Kanizsai et al., 2013	Shore meadow; at road. Under stone; between asphalt edge and grass.
Lasius platythorax	Camponotus herculeanus	Finland	Czechowski, 2004; Kanizsai et al., 2013	In decaying wood.
Lasius platythorax	Formica fusca	Finland, Poland	Kanizsai et al., 2013; Włodarczyk et al., 2009	Forest on rocks; clearcut of managed forest. In decaying wood; in tree stump
Lasius platythorax	Lasius flavus	Finland	Czechowski, 2004; Kanizsai et al., 2013	Different successional series of rocky habitats. In rock crevice/under stone/overgrown soil.
Lasius platythorax	Myrmica rubra	Finland	Czechowski, 2004; Kanizsai et al., 2013	Forest. In decaying wood
Lasius sabularum	Lasius umbratus	Poland	Borowiec, 2011; Kanizsai et al., 2013	Stand of oak trees. Under stone.
Lasius sitiens	Crematogaster punctulata
Lasius sitiens	Formica argentea
Lasius sitiens	Formica occulta
Lasius sitiens	Lasius latipes
Lasius sitiens	Monomorium minimum
Lasius sitiens	Myrmica hamulata
Lasius sitiens	Solenopsis molesta
Lasius umbratus	Formica sanguinea	Poland	Czechowski & Rotkiewicz, 1997; Kanizsai et al., 2013	Clearings in a pine forest. In the sandy soil.
Lasius umbratus	Lasius sabularum	Poland	Borowiec, 2011; Kanizsai et al., 2013	Stand of oak trees. Under stone.
Lasius umbratus	Polyergus rufescens	Poland	Czechowski & Rotkiewicz, 1997; Kanizsai et al., 2013	Clearings in a pine forest. In the sandy soil.
Leptothorax acervorum	Formica fusca	Finland	Czechowski, 2004; Kanizsai et al., 2013	In different successional series of rocky habitats, in mound of ''Formica lugubris''
Leptothorax calderoni	Camponotus modoc	United States	Kanizsai et al., 2013; Wheeler, 1917	Pine forest. In log/in stump.
Leptothorax muscorum	Formica rufa	Sweden	Kanizsai et al., 2013; Wheeler, 1901
Leptothorax muscorum	Myrmica rubra	Finland	Czechowski, 2004; Kanizsai et al., 2013	Shore meadow. Under stone
Manica invidia	Formicoxenus chamberlini		Wheeler (1904)
Megalomyrmex adamsae	Mycetomoellerius zeteki
Megalomyrmex mondabora	Cyphomyrmex cornutus	Costa Rica	Adams & Longino, 2007; Adams et al., 2015
Megalomyrmex mondaboroides	Apterostigma goniodes	Costa Rica	Longino, 2010; Adams et al., 2015
Megalomyrmex mondaboroides	Cyphomyrmex costatus	Panama	Longino, 2010; Adams et al., 2015
Megalomyrmex silvestrii	Cyphomyrmex costatus	Panama	Longino, 2010; Adams et al., 2015
Megalomyrmex symmetochus	Sericomyrmex amabilis		Liberti et al., 2015
Megalomyrmex wettereri	Paratrachymyrmex bugnioni		Brandão (2003)
Monomorium minimum	Camponotus festinatus	United States	Kanizsai et al., 2013; Wheeler, 1901
Monomorium minimum	Camponotus sansabeanus	United States	Kanizsai et al., 2013; Wheeler, 1901
Monomorium minimum	Formica gnava	United States	Kanizsai et al., 2013; Wheeler, 1901
Monomorium minimum	Lasius crypticus		Mackay & Mackay, 2002
Monomorium minimum	Pachycondyla harpax	United States	Kanizsai et al., 2013; Wheeler, 1901
Monomorium minimum	Pogonomyrmex barbatus	United States	Kanizsai et al., 2013; Wheeler, 1901
Myrmecina americana	Formica gnava	United States	Kanizsai et al., 2013; Wheeler, 1901
Myrmecina americana	Myrmica pinetorum	United States	Kanizsai et al., 2013; Wheeler, 1921
Myrmecina americana	Pheidole picea	United States	Kanizsai et al., 2013; Wheeler, 1901
Myrmecina americana	Ponera pennsylvanica	United States	Kanizsai et al., 2013; Wheeler, 1901
Myrmecina graminicola	Ponera coarctata	Hungary	Kanizsai et al., 2013	Pine forest. Under stone.
Myrmica alaskensis	Formicoxenus quebecensis
Myrmica fracticornis	Formicoxenus provancheri
Myrmica hamulata	Lasius latipes
Myrmica incompleta	Formicoxenus provancheri
Myrmica pinetorum	Myrmecina americana	United States	Kanizsai et al., 2013; Wheeler, 1921
Myrmica rubra	Formica fusca	Finland, United Kingdom	Czechowski, W. 2003; Czechowski, 2004; Kanizsai et al., 2013; Morley, 1945	habitat foreshore, nests in decaying wood, under stone
Myrmica rubra	Lasius niger	Finland	Czechowski, 2004; Kanizsai et al., 2013	Shore meadow; at road. Under stone; between asphalt edge and grass.
Myrmica rubra	Lasius platythorax	Finland	Czechowski, 2004; Kanizsai et al., 2013	Forest. In decaying wood
Myrmica rubra	Leptothorax muscorum	Finland	Czechowski, 2004; Kanizsai et al., 2013	Shore meadow. Under stone
Myrmica ruginodis	Formica fusca	Poland	Kanizsai et al., 2013; Włodarczyk et al., 2009	Clearcut of managed forest, in tree stumps
Myrmica ruginodis	Lasius flavus	Italy	Balzani et al., 2022
Myrmica scabrinodis	Lasius flavus	United Kingdom	Kanizsai et al., 2013; Morley, 1945	Foreshore. Under stone.
Odontomachus panamensis	Crematogaster carinata
Pachycondyla harpax	Camponotus festinatus	United States	Kanizsai et al., 2013; Wheeler, 1901	At road. Under stone.
Pachycondyla harpax	Camponotus sansabeanus	United States	Kanizsai et al., 2013; Wheeler, 1901
Pachycondyla harpax	Monomorium minimum	United States	Kanizsai et al., 2013; Wheeler, 1901
Paratrachymyrmex bugnioni	Megalomyrmex wettereri		Brandão (2003)
Pheidole picea	Lasius minutus	United States	Gaige ,1914; Kanizsai et al., 2013	Hardwood forest. In a stump.
Pheidole picea	Lasius nearcticus	United States	Gaige ,1914; Kanizsai et al., 2013	Hardwood forest. Under rock.
Pheidole picea	Myrmecina americana	United States	Kanizsai et al., 2013; Wheeler, 1901
Pogonomyrmex barbatus	Monomorium minimum	United States	Kanizsai et al., 2013; Wheeler, 1901
Polyergus rufescens	Formicoxenus nitidulus		Wilson 1971; Martin et al. 2007
Polyergus rufescens	Lasius umbratus	Poland	Czechowski & Rotkiewicz, 1997; Kanizsai et al., 2013	Clearings in a pine forest. In the sandy soil.
Ponera coarctata	Myrmecina graminicola	Hungary	Kanizsai et al., 2013	Pine forest. Under stone.
Ponera pennsylvanica	Myrmecina americana	United States	Kanizsai et al., 2013; Wheeler, 1901
Pseudomyrmex elongatulus	Tapinoma litorale
Sericomyrmex amabilis	Megalomyrmex symmetochus		Wheeler 1925b, Adams et al. 2013
Solenopsis picea	Camponotus femoratus	British Guiana	Weber, 1943
Solenopsis picea	Crematogaster carinata	British Guiana		Weber, 1943
Strumigenys arizonica	Trachymyrmex arizonensis
Strumigenys pergandei	Formica spp.	United States	Kanizsai et al., 2013; Wheeler, 1905	In soil.
Tapinoma litorale	Camponotus planatus
Tapinoma litorale	Cephalotes varians
Tapinoma litorale	Pseudomyrmex elongatulus
Temnothorax andrei	Camponotus yogi	United States	Creighton & Snelling, 1966; Kanizsai et al., 2013	Chaparral. In living stems of ''Haplopappus pinifolius''.
Temnothorax andrei	Formica moki		Mann, 1911	nature of relationship unknown
Temnothorax andrei	Formica ravida		Mackay & Mackay, 1984	nature of relationship unknown
Temnothorax subditivus	Crematogaster carinata
Tetramorium caespitum	Formica fusca	Poland	Kanizsai et al., 2013; Włodarczyk et al., 2009	Clearcut of managed forest, in tree stumps
Tetramorium caespitum	Formicoxenus nitidulus		Martin et al. 2007; Wilson 1971
Tetramorium caespitum	Lasius flavus	Finland	Czechowski, 2004; Kanizsai et al., 2013	Different successional series of rocky habitats. Under stone
Tetramorium tsushimae	Formica japonica	Japan	Czechowski & Yamauchi, 1998; Kanizsai et al., 2013	Urban area. In the soil.
Trachymyrmex arizonensis	Strumigenys arizonica		Ward, 1988

References

• Kanizsai, O., Lőrinczi, G., Gallé, L. 2013. Nesting associations without interdependence: A preliminary review on plesiobiosis in ants. Psyche 2013, 238602 (doi:10.1155/2013/238602).