Xenobiosis

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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 aquilonia Formica fusca Finland Czechowski & Vepsalainen, 1999; Kanizsai et al., 2013 Forest on rocks. Mound of ''F. lugubris''
Formica aquilonia Formicoxenus nitidulus Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007
Formica aquilonia Lasius flavus Finland Czechowski, 2004; Kanizsai et al., 2013 Different successional series of rocky habitats. Mound of ''Formica aquilonia''.
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 japonica Formica lemani Yamaoka, 1977; Kvifte and Soule, 2017 plesiobiotic
Formica japonica Tetramorium tsushimae Japan Czechowski & Yamauchi, 1998; Kanizsai et al., 2013 Urban area. In the soil.
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 sanguinea Lasius umbratus Poland Czechowski & Rotkiewicz, 1997; Kanizsai et al., 2013 Clearings in a pine forest. In the sandy soil.
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