Editing Commensalism

{{Short description|Beneficial symbiosis between species}}
{{use dmy dates |date=May 2023}}
[[File:Echeneis naucrates 241647485.jpg|thumb|upright=1.35|[[Remora]] are specially adapted to attach themselves to larger fish (or other animals, in this case a sea turtle) that provide locomotion and food.]]

'''Commensalism''' is a long-term [[biological interaction]] ([[symbiosis]]) in which members of one [[species]] gain benefits while those of the other species neither benefit nor are harmed.<ref name=wilson1975/> This is in contrast with [[mutualism (biology)|mutualism]], in which both organisms benefit from each other; [[Symbiosis#Amensalism|amensalism]], where one is harmed while the other is unaffected; and [[parasitism]], where one is harmed and the other benefits.

The commensal (the species that benefits from the association) may obtain nutrients, shelter, support, or locomotion from the host species, which is substantially unaffected. The commensal relation is often between a larger host and a smaller commensal; the [[Host (biology)|host]] organism is unmodified, whereas the commensal species may show great structural adaptation consistent with its habits, as in the [[remora]]s that ride attached to [[sharks]] and other fishes. Remoras feed on their hosts' fecal matter,<ref>{{Cite journal|last1=Williams|first1=E. H.|last2=Mignucci-Giannoni|first2=A. A.|last3=Bunkley-Williams|first3=L.|last4=Bonde|first4=R. K.|last5=Self-Sullivan|first5=C.|last6=Preen|first6=A.|last7=Cockcroft|first7=V. G.|date=2003|title=Echeneid-sirenian associations, with information on sharksucker diet|journal=Journal of Fish Biology|language=en|volume=63|issue=5|pages=1176–1183|doi=10.1046/j.1095-8649.2003.00236.x|bibcode=2003JFBio..63.1176W |issn=0022-1112}}</ref> while [[pilot fish]] feed on the leftovers of their hosts' meals. Numerous birds perch on bodies of large mammal [[Herbivore|herbivores]] or feed on the insects turned up by grazing mammals.<ref name=pmid29576981/>

==Etymology==
The word "commensalism" is derived from the word "commensal", meaning "eating at the same table" in human social interaction, which in turn comes through [[French language|French]] from the [[Medieval Latin]] ''commensalis'', meaning "sharing a table", from the prefix ''[[wikt:com-#Latin|com-]]'', meaning "together", and ''[[wikt:mensa#Latin|mensa]]'', meaning "table" or "meal".<ref>{{OEtymD|commensalism}}</ref> Commensality, at the Universities of [[University of Oxford|Oxford]] and [[University of Cambridge|Cambridge]], refers to professors eating at the same table as students (as they live in the same "college").{{cn|date=June 2024}}

The Belgian [[Zoology|zoologist]] and [[Paleontology|paleontologist]] [[Pierre-Joseph van Beneden]] introduced the term "commensalism" in 1876.<ref>{{cite book |series=International scientific series |volume=19 |doi=10.5962/bhl.title.132633 |last=van Beneden |first=Pierre-Joseph |year=1876 |title=Animal parasites and messmates |location=London |publisher=Henry S. King |url=https://www.biodiversitylibrary.org/bibliography/132633#/summary }}</ref>

== Examples of commensal relationships ==
[[File:Rock dove - natures pics.jpg|thumb|[[Domestic pigeon|Domestic]] and [[feral pigeons]] (''Columba livia domestica'') are commensals, having lived alongside humans for thousands of years after being domesticated from the [[rock dove]] (''Columba livia''). Due to its range being expanded with human assistance, the pigeon has a [[cosmopolitan distribution]].<ref>{{Cite journal |last1=Tang |first1=Qian |last2=Low |first2=Gabriel Weijie |last3=Lim |first3=Jia Ying |last4=Gwee |first4=Chyi Yin |last5=Rheindt |first5=Frank E. |date=2018-07-21 |title=Human activities and landscape features interact to closely define the distribution and dispersal of an urban commensal |journal=Evolutionary Applications |volume=11 |issue=9 |pages=1598–1608 |doi=10.1111/eva.12650 |issn=1752-4571 |pmc=6183452 |pmid=30344630|bibcode=2018EvApp..11.1598T }}</ref> ]]
The commensal pathway was traversed by animals that fed on refuse around human habitats or by animals that preyed on other animals drawn to human camps. Those animals established a commensal relationship with humans in which the animals benefited but the humans received little benefit or harm. Those animals that were most capable of taking advantage of the resources associated with human camps would have been the 'tamer' individuals: less aggressive, with shorter [[Fight-or-flight response|fight-or-flight]] distances. Later, these animals developed closer social or economic bonds with humans and led to a domestic relationship.<ref name=zeder2012/><ref name=larson2014/>

The leap from a [[synanthrope|synanthropic]] population to a domestic one could only have taken place after the animals had progressed from anthropophily to habituation to commensalism and partnership, at which point the establishment of a reciprocal relationship between animal and human would have laid the foundation for domestication, including captivity and then human-controlled breeding. From this perspective, animal domestication is a [[coevolution]]ary process in which a population responds to selective pressure while adapting to a novel [[Ecological niche|niche]] that includes another species with evolving behaviors.<ref name=larson2014/>

===Dogs and humans===
The [[dog]] was the first domesticated animal and was domesticated and widely established across [[Eurasia]] before the end of the [[Pleistocene]], well before the cultivation of crops or the [[domestication]] of other animals.<ref name=larson2012/> The dog is often hypothesised to be a classic example of a domestic animal that likely traveled a commensal pathway into domestication. Archaeological evidence, such as the [[Bonn–Oberkassel dog]] dating to ~14,000 BP,<ref name=janssens2018/> supports the hypothesis that dog domestication preceded the emergence of agriculture <ref name=vila1997/><ref name=thalmann2013/> and began close to the [[Last Glacial Maximum]] when hunter-gatherers preyed on [[megafauna]].

The wolves more likely drawn to human camps were the less aggressive, subdominant pack members with lowered flight response, higher stress thresholds, and less wary around humans, and therefore better candidates for domestication.<ref name=zeder2012/>
Proto-dogs might have taken advantage of carcasses left on site by early hunters, assisted in capturing prey, or provided defense from large competing predators at kills.<ref name=thalmann2013/> However, the extent to which proto-domestic wolves could have become dependent on this way of life prior to domestication and without human provisioning is unclear and highly debated. In contrast, [[cats]] may have become fully dependent on a commensal lifestyle before being domesticated by preying on other commensal animals, such as rats and mice, without any human provisioning. The debate over the extent to which some wolves were commensal with humans before domestication stems from the debate over the level of human intentionality in domestication, which remains untested.<ref name=larson2014/><ref name=hulmebeaman2016/>

The earliest sign of domestication in dogs was the [[Neoteny|neotenization]] (retaining juvenile features into adulthood) of skull morphology<ref name=morey1992/><ref name=trut1999/><ref name=zeder2012/> and the shortening of snout length that results in tooth crowding, reduction in tooth size, and a reduction in the number of teeth,<ref name=turnbull1974/><ref name=zeder2012/> which has been attributed to the strong selection for reduced aggression.<ref name=trut1999/><ref name=zeder2012/> This process may have begun during the initial commensal stage of dog domestication, even before humans began to be active partners in the process.<ref name=zeder2012/><ref name=larson2014/>

A [[Mitochondrion|mitochondrial]], microsatellite, and [[Y chromosome|Y-chromosome]] assessment of two wolf populations in North America combined with satellite [[telemetry]] data revealed significant genetic and morphological differences between one population that migrated with and preyed upon caribou and another territorial ecotype population that remained in a [[Taiga|boreal coniferous forest]]. Although these two populations spend a period of the year in the same place, and though there was evidence of gene flow between them, the difference in prey-habitat specialization has been sufficient to maintain genetic and even coloration divergence.<ref name=musiani2007/><ref name=larson2014/>

A different study has identified the remains of a population of extinct [[Pleistocene]] [[Beringian wolf|Beringian wolves]] with unique mitochondrial signatures. The skull shape, tooth wear, and [[isotopic signature]]s suggested these remains were derived from a population of specialist [[megafauna]] hunters and scavengers that became extinct while less specialized wolf [[ecotype]]s survived.<ref name=leonard2007/><ref name=larson2014/> Analogous to the modern wolf ecotype that has evolved to track and prey upon caribou, a Pleistocene wolf population could have begun following mobile [[hunter-gatherer]]s, thus slowly acquiring genetic and [[Phenotype|phenotypic]] differences that would have allowed them to adapt to the [[human habitat]] more successfully.<ref name=wolpert2013/><ref name=larson2014/>

===''Aspergillus'' and ''Staphylococcus''===
{{See also|Aspergillus|Staphylococcus}}
Numerous genera of [[bacteria]] and [[fungi]] live on and in the human body as part of its natural flora. The fungal genus ''[[Aspergillus]]'' is capable of living under considerable environmental stress, and thus is capable of colonising the upper gastrointestinal tract where relatively few examples of the body's gut flora can survive due to highly acidic or alkaline conditions produced by gastric acid and digestive juices. While ''Aspergillus'' normally produces no symptoms, in individuals who are immunocompromised or suffering from existing conditions such as [[tuberculosis]], a condition called [[aspergillosis]] can occur, in which populations of ''Aspergillus'' grow out of control.{{cn|date=June 2024}}

''[[Staphylococcus aureus]]'', a common bacterial species, is known best for its numerous pathogenic strains that can cause numerous illnesses and conditions. However, many strains of ''S. aureus'' are metabiotic commensals, and are present on roughly 20 to 30% of the human population as part of the [[skin flora]].<ref name=pmid9227864/> ''S. aureus'' also benefits from the variable ambient conditions created by the body's mucous membranes, and as such can be found in the [[oral cavity|oral]] and [[nasal cavity|nasal cavities]], as well as inside the [[ear canal]]. Other ''Staphylococcus'' species including ''[[Staphylococcus warneri|S. warneri]]'', ''[[Staphylococcus lugdunensis|S. lugdunensis]]'' and ''[[Staphylococcus epidermidis|S. epidermidis]]'', will also engage in commensalism for similar purposes.{{cn|date=June 2024}}

=== ''Nitrosomonas'' spp and ''Nitrobacter'' spp ===
Commensalistic relationships between microorganisms include situations in which the waste product of one microorganism is a substrate for another species. One good example is nitrification — the oxidation of [[ammonium|ammonium ion]] to [[nitrate]]. Nitrification occurs in two steps: first, bacteria such as ''[[Nitrosomonas]]'' [[Species#Abbreviations|spp.]] and certain [[crenarchaeota|crenarchaeote]]s oxidize ammonium to [[nitrite]]; and second, nitrite is oxidized to nitrate by ''[[Nitrobacter]]'' spp. and similar bacteria. ''Nitrobacter'' spp. benefit from their association with ''Nitrosomonas'' spp. because they use nitrite to obtain energy for growth.{{cn|date=June 2024}}

Commensalistic associations also occur when one microbial group modifies the environment to make it better suited for another organism. The synthesis of acidic waste products during [[fermentation]] stimulates the proliferation of more acid-tolerant microorganisms, which may be only a minor part of the microbial community at neutral pH. A good example is the succession of microorganisms during milk spoilage.{{cn|date=June 2024}}

[[Biofilm]] formation provides another example. The colonization of a newly exposed surface by one type of microorganism (an initial colonizer) makes it possible for other microorganisms to attach to the microbially modified surface.{{cn|date=June 2024}}

=== Octocorals and brittle stars ===

In deep-sea, benthic environments there is an associative relationship between [[Octocorallia|octocorals]] and [[brittle star]]s. Due to the currents flowing upward along seamount ridges, atop these ridges there are colonies of suspension feeding corals and sponges, and brittle stars that grip tight to them and get up off the sea floor. A specific documented commensal relationship is between the ophiuran ''Ophiocreas oedipus'' Lyman and the octocoral primnoid ''Metallogorgia melanotrichos''.{{cn|date=June 2024}}

Historically, commensalism has been recognized as the usual type of association between brittle stars and octocorals.<ref>{{cite journal | url=https://pubmed.ncbi.nlm.nih.gov/21962750/ | pmid=21962750 | doi=10.1016/B978-0-12-385529-9.00002-0 | title=Biology of deep-water octocorals | journal=Advances in Marine Biology | year=2011 | volume=60 | pages=41–122 | last1=Watling | first1=Les | last2=France | first2=Scott C. | last3=Pante | first3=Eric | last4=Simpson | first4=Anne | isbn=9780123855299 }}</ref> In this association, the ophiurans benefit directly by being elevated through facilitating their feeding by suspension, while the octocorals do not seem to benefit or be harmed by this relationship.<ref>{{cite journal | jstor=24844596 | title=Size structure of dense populations of the brittle star Ophiura sarsii (Ophiuroidea: Echinodermata) in the bathyal zone around Japan | last1=Fujita | first1=Toshihiko | last2=Ohta | first2=Suguru | journal=Marine Ecology Progress Series | year=1990 | volume=64 | issue=1/2 | pages=113–122 | doi=10.3354/meps064113 | bibcode=1990MEPS...64..113F | doi-access=free }}</ref>

Recent studies in the Gulf of Mexico have suggested that there are actually some benefits to the octocorals, such as receiving a cleaning action by the brittle star as it slowly moves around the coral.<ref>{{cite journal |doi=10.3354/meps11697 |url=https://www.int-res.com/articles/meps_oa/m549p089.pdf |access-date=24 May 2023 |title=Mutualistic symbiosis with ophiuroids limited the impact of the Deepwater Horizon oil spill on deep-sea octocorals |year=2016 |last1=Girard |first1=F. |last2=Fu |first2=B. |last3=Fisher |first3=CR |journal=Marine Ecology Progress Series |volume=549 |pages=89–98 |bibcode=2016MEPS..549...89G }}</ref> In some cases, a close relationship occurs between cohabiting species, with the interaction beginning from their juvenile stages.<ref>{{cite journal |last1=Mejía-Quintero |first1=Katherine |last2=Borrero-Pérez |first2=Giomar H. |last3=Montoya-Cadavid |first3=Erika |title=Callogorgia spp. and Their Brittle Stars: Recording Unknown Relationships in the Pacific Ocean and the Caribbean Sea |journal=Frontiers in Marine Science |date=2021 |volume=8 |doi=10.3389/fmars.2021.735039 |issn=2296-7745|doi-access=free |bibcode=2021FrMaS...835039M }} [[File:CC-BY icon.svg|50px]]  Text was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/  Creative Commons Attribution 4.0 International License].</ref>

== Arguments ==

Whether the relationship between humans and some types of [[gut flora]] is commensal or [[Mutualism (biology)|mutualistic]] is still unanswered.

Some biologists argue that any close interaction between two organisms is unlikely to be completely neutral for either party, and that relationships identified as commensal are likely [[Mutualism (biology)|mutualistic]] or [[parasitism|parasitic]] in a subtle way that has not been detected. For example, [[epiphytes]] are "nutritional pirates" that may intercept substantial amounts of nutrients that would otherwise go to the host plant.<ref>{{cite book | vauthors = Benzing DH | date = 1980 | title = Biology of the Bromeliads | location = [[Eureka, California]] | publisher = Mad River Press}}{{page needed|date=September 2014}}</ref> Large numbers of epiphytes can also cause tree limbs to break or shade the host plant and reduce its rate of photosynthesis. Similarly, phoretic mites may hinder their host by making flight more difficult, which may affect its aerial hunting ability or cause it to expend extra energy while carrying these passengers.{{cn|date=June 2024}}

== Types ==

[[Image:Fly June 2008-2.jpg|right|thumb|Phoretic [[mite]]s on a fly (''[[Pseudolynchia canariensis]]'')]]

Like all ecological interactions, commensalisms vary in strength and duration from intimate, long-lived [[symbiosis|symbioses]] to brief, weak interactions through intermediaries.{{cn|date=June 2024}}

=== Phoresy ===

[[Phoresis (biology)|Phoresy]] is one animal attached to another exclusively for transport, mainly [[arthropod]]s, examples of which are [[mite]]s on [[insect]]s (such as [[beetle]]s, [[fly|flies]] or [[bee]]s), [[pseudoscorpion]]s on [[mammal]]s<ref name=jstor2388309/> or beetles, and [[millipede]]s on [[bird]]s.<ref name="Karel et al 2001"/> Phoresy can be either obligate or facultative (induced by environmental conditions).

=== Inquilinism ===
[[Image:Tillandsia bourgaei.jpg|thumb|right|200px|Inquilinism: ''[[Tillandsia bourgaei]]'' growing on an oak tree in Mexico]]

[[Inquilinism]] is the use of a second organism for permanent housing. Examples are [[epiphyte|epiphytic]] [[plant]]s (such as many [[orchid]]s) that grow on trees,<ref>{{cite encyclopedia |first=C. Michael |last=Hogan |year=2011|url=https://editors.eol.org/eoearth/wiki/Commensalism |title=Commensalism |editor-first=M. |editor-last=Mcginley |editor-first2=C. J. |editor-last2=Cleveland |encyclopedia=Encyclopedia of Earth |publisher=[[National Council for Science and the Environment]] |location=Washington DC }}</ref> or birds that live in holes in trees.

=== Metabiosis ===

Metabiosis is a more indirect dependency, in which one organism creates or prepares a suitable environment for a second. Examples include [[maggots]], which develop on and infest corpses, and [[hermit crab]]s, which use [[gastropod]] shells to protect their bodies.{{cn|date=June 2024}}

=== Facilitation ===
'''Facilitation''' or '''probiosis''' describes [[Biological interaction|species interactions]] that benefit at least one of the participants and cause harm to neither.{{cn|date=June 2024}}

=== Necromeny ===
[[Necromeny]] is one animal associating with another until the latter dies, then the former feeds on the corpse of the latter. Examples include some [[nematode]]s<ref>{{Cite journal |last=Sudhaus |first=W. |date=30 December 2010 |title=Preadaptive plateau in Rhabditida (Nematoda) allowed the repeated evolution of zooparasites, with an outlook on evolution of life cycles within Spiroascarida |url=http://www.palaeodiversity.org/pdf/03Suppl/Supplement_Sudhaus.pdf |journal=Palaeodiversity |volume=3 |pages=117–130}}</ref> and some mites.<ref>{{Cite journal |last1=Bhadran |first1=Anjitha K. |last2=Ramani |first2=N. |date=2019-10-03 |title=Relationships between phoretic mites and their carrier, the banana pseudostem weevil Odoiporus longicollis Oliver (Coleoptera: Curculionidae) |url=https://www.tandfonline.com/doi/full/10.1080/01647954.2019.1656286 |journal=International Journal of Acarology |language=en |volume=45 |issue=6–7 |pages=361–365 |doi=10.1080/01647954.2019.1656286 |bibcode=2019IJAca..45..361B |s2cid=202867426 |issn=0164-7954|url-access=subscription }}</ref><ref>{{Cite journal |last1=Al-Deeb |first1=Mohammad Ali |last2=Muzaffar |first2=Sabir Bin |last3=Sharif |first3=Eyas Mohammad |date=2012 |title=Interactions between Phoretic Mites and the Arabian Rhinoceros Beetle, Oryctes agamemnon arabicus |journal=Journal of Insect Science |language=en |volume=12 |issue=128 |page=128 |doi=10.1673/031.012.12801 |issn=1536-2442 |pmc=3637038 |pmid=23448160}}</ref>

== See also ==
* [[Mutualism (biology)|Mutualism]] – where both organisms experience mutual benefit in the relationship
* [[Parasitism]] – where one organism benefits at the expense of another organism.
* [[Parabiosis]] – where both organisms occupy the same dwelling, but do not interfere with each other
* [[Symbiosis]] – long-term interactions between different biological species, which can be mutualistic, commensal or parasitic
* [[Synanthrope]] – species commensal with humans

== References ==
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<ref name=larson2014>{{cite journal |doi=10.1146/annurev-ecolsys-110512-135813 |title=The Evolution of Animal Domestication |journal=Annual Review of Ecology, Evolution, and Systematics |volume=45 |pages=115–136 |year=2014 |last1=Larson |first1=Greger |last2=Fuller |first2=Dorian Q. |s2cid=56381833 | name-list-style = vanc }}</ref>

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<ref name=thalmann2013>{{cite journal | vauthors = Thalmann O, Shapiro B, Cui P, Schuenemann VJ, Sawyer SK, Greenfield DL, Germonpré MB, Sablin MV, López-Giráldez F, Domingo-Roura X, Napierala H, Uerpmann HP, Loponte DM, Acosta AA, Giemsch L, Schmitz RW, Worthington B, Buikstra JE, Druzhkova A, Graphodatsky AS, Ovodov ND, Wahlberg N, Freedman AH, Schweizer RM, Koepfli KP, Leonard JA, Meyer M, Krause J, Pääbo S, Green RE, Wayne RK | display-authors = 6 | title = Complete mitochondrial genomes of ancient canids suggest a European origin of domestic dogs | journal = Science | volume = 342 | issue = 6160 | pages = 871–4 | date = November 2013 | pmid = 24233726 | doi = 10.1126/science.1243650 | bibcode = 2013Sci...342..871T | hdl = 10261/88173 | s2cid = 1526260 }}</ref>

<ref name=morey1992>{{cite journal |doi=10.1016/0305-4403(92)90049-9 |title=Size, shape and development in the evolution of the domestic dog |journal=Journal of Archaeological Science |volume=19 |issue=2 |pages=181–204 |year=1992 |last1=Morey |first1=Darcy F. |bibcode=1992JArSc..19..181M | name-list-style = vanc }}</ref>

<ref name=trut1999>{{cite journal |doi=10.1511/1999.2.160 |title=Early Canid Domestication: The Farm-Fox Experiment |journal=American Scientist |volume=87 |issue=2 |pages=160 |year=1999 |last1=Trut |first1=Lyudmila | name-list-style = vanc |bibcode=1999AmSci..87.....T }}</ref>

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<ref name=wilson1975>{{cite book|last1=Wilson|first1=Edward O. | name-list-style = vanc |title=Sociobiology: The New Synthesis|publisher=Harvard University Press|year=1975|chapter=Ch.17-Social Symbiosis|pages=354|isbn=978-0-674-00089-6|chapter-url={{Google books|plainurl=yes|id=v7lV9tz8fXAC|page=354}}}}</ref>

<ref name=wolpert2013>{{cite news | last = Wolpert  | first = Stuart | name-list-style = vanc | title = Dogs likely originated in Europe more than 18,000 years ago, UCLA biologists report | newspaper = UCLA News Room | url = http://newsroom.ucla.edu/releases/dogs-likely-originated-in-europe-249325 | date = November 14, 2013 | access-date = December 10, 2014}} Statement by Wayne, R.K.</ref>

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<ref name=janssens2018>{{cite journal |doi=10.1016/j.jas.2018.01.004 |title=A new look at an old dog: Bonn-Oberkassel reconsidered |journal=Journal of Archaeological Science |volume=92 |pages=126–138 |year=2018 |last1=Janssens |first1=Luc |last2=Giemsch |first2=Liane |last3=Schmitz |first3=Ralf |last4=Street |first4=Martin |last5=Van Dongen |first5=Stefan |last6=Crombé |first6=Philippe |bibcode=2018JArSc..92..126J | name-list-style = vanc |url=https://biblio.ugent.be/publication/8550758/file/8550759.pdf |hdl=1854/LU-8550758 |hdl-access=free }}</ref>

<ref name=hulmebeaman2016>{{cite journal | vauthors = Hulme-Beaman A, Dobney K, Cucchi T, Searle JB | title = An Ecological and Evolutionary Framework for Commensalism in Anthropogenic Environments | journal = Trends in Ecology & Evolution | volume = 31 | issue = 8 | pages = 633–645 | date = August 2016 | pmid = 27297117 | doi = 10.1016/j.tree.2016.05.001 | doi-access = free | bibcode = 2016TEcoE..31..633H | hdl = 2164/6176 | hdl-access = free }}</ref>

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<ref name=jstor2388309>{{cite journal |last1=Durden |first1=Lance A. | name-list-style = vanc |date=June 1991 |title=Pseudoscorpions Associated With Mammals in Papua New Guinea |journal=Biotropica |volume=23 |issue=2 |pages=204–6 |jstor=2388309 |doi=10.2307/2388309|bibcode=1991Biotr..23..204D }}</ref>

<ref name="Karel et al 2001">{{cite journal |first1=Karel |last1=Tajovský |first2=Andrej |last2=Mock |first3=Miroslav |last3=Krumpál | name-list-style = vanc |year=2001 |title=Millipedes (''Diplopoda'') in birdsˈ nests |journal=European Journal of Soil Biology |volume=37 |issue=4 |pages=321–3 |doi=10.1016/S1164-5563(01)01108-6 |bibcode=2001EJSB...37..321T }}</ref>

}}

== External links ==
* {{Commons category-inline}}
* {{cite encyclopedia|url=https://www.britannica.com/EBchecked/topic/127789/commensalism|encyclopedia=Encyclopædia Britannica|title=Commensalism |date=29 April 2023 }}

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