Editing Biological interaction

{{Short description|Effect that organisms have on other organisms}}
{{Redirect|Biological relationship|family relatives|Consanguinity}}
[[File:Black Walnut middle.JPG|upright=1.35|thumb|The [[black walnut]] secretes a chemical from its roots that harms neighboring plants, an example of [[competition (biology)|competitive]] [[antagonism (phytopathology)|antagonism]].]]
In [[ecology]], a '''biological interaction''' is the effect that a pair of [[organism]]s living together in a [[Community (ecology)|community]] have on each other. They can be either of the same [[species]] (intraspecific interactions), or of different species (interspecific interactions). These effects may be short-term, or long-term, both often strongly influence the [[adaptation]] and [[evolution]] of the species involved. Biological interactions range from [[Mutualism (biology)|mutualism]], beneficial to both partners, to [[competition (biology)|competition]], harmful to both partners. Interactions can be direct when physical contact is established or indirect, through intermediaries such as shared resources, territories, ecological services, metabolic waste, toxins or growth inhibitors. This type of relationship can be shown by net effect based on individual effects on both organisms arising out of relationship.

Several recent studies have suggested non-trophic species interactions such as habitat modification and mutualisms can be important determinants of food web structures. However, it remains unclear whether these findings generalize across ecosystems, and whether non-trophic interactions affect food webs randomly, or affect specific trophic levels or functional groups.

==History==
Although biological interactions, more or less individually, were studied earlier, [[Edward Haskell]] (1949) gave an integrative approach to the thematic, proposing a classification of "co-actions",<ref>Haskell, E. F. (1949). A clarification of social science. ''Main Currents in Modern Thought'' 7: 45–51.</ref> later adopted by biologists as "interactions". Close and long-term interactions are described as [[symbiosis]];{{efn|Symbiosis was formerly used to mean a mutualism.}} symbioses that are mutually beneficial are called [[mutualism (biology)|mutualistic]].<ref>{{Cite journal |last=Burkholder |first=Paul R. |date=1952 |title=Cooperation and Conflict Among Primitive Organisms |url=https://www.jstor.org/stable/27826458 |journal=American Scientist |volume=40 |issue=4 |pages=600–631 |jstor=27826458 |issn=0003-0996}}</ref><ref>{{Cite book |last=Bronstein |first=Judith L. |url=https://books.google.com/books?id=hbgVDAAAQBAJ |title=Mutualism |date=2015 |publisher=Oxford University Press |isbn=978-0-19-967565-4 |language=en}}</ref><ref>{{Cite journal |last=Pringle |first=Elizabeth G. |date=2016-10-12 |title=Orienting the Interaction Compass: Resource Availability as a Major Driver of Context Dependence |journal=PLOS Biology |language=en |volume=14 |issue=10 |pages=e2000891 |doi=10.1371/journal.pbio.2000891 |doi-access=free |issn=1545-7885 |pmc=5061325 |pmid=27732591}}</ref>

The term symbiosis was subject to a century-long debate about whether it should specifically denote mutualism, as in [[lichen]]s or in parasites that benefit themselves.<ref>{{Cite book |last=Douglas |first=A. E. |url=https://www.worldcat.org/oclc/437054000 |title=The symbiotic habit |date=2010 |publisher=Princeton University Press |isbn=978-0-691-11341-8 |location=Princeton, N.J. |oclc=437054000}}</ref> This debate created two different classifications for biotic interactions, one based on the time (long-term and short-term interactions), and other based on the magnitude of interaction force (competition/mutualism) or effect of individual fitness, according the '''[[stress gradient hypothesis]]''' and [[Mutualism Parasitism Continuum]]. [[Evolutionary game theory]] such as [[Red Queen hypothesis|Red Queen Hypothesis]], [[Red King hypothesis|Red King Hypothesis]] or [[Black Queen hypothesis|Black Queen Hypothesis]], have demonstrated a classification based on the force of interaction is important.{{cn|date=May 2023}}

== Classification based on time of interaction ==

=== Short-term interactions ===
[[File:Osprey eating a fish.jpg|thumb|[[Predation]] is a short-term interaction, in which the predator, here an [[osprey]], kills and eats its prey.]]
Short-term interactions, including [[predation]] and [[pollination]], are extremely important in [[ecology]] and [[evolution]]. These are short-lived in terms of the duration of a single interaction: a predator kills and eats a prey; a pollinator transfers pollen from one flower to another; but they are extremely durable in terms of their influence on the evolution of both partners. As a result, the partners [[coevolution|coevolve]].<ref name="Bengtson2002"/><ref name=Lunau/>

==== Predation ====

{{main|Predation}}

In predation, one organism, the predator, kills and eats another organism, its prey. Predators are adapted and often highly specialized for hunting, with acute senses such as [[eye|vision]], [[hearing]], or [[olfaction|smell]]. Many predatory animals, both [[vertebrate]] and [[invertebrate]], have sharp [[claw]]s or [[jaw]]s to grip, kill, and cut up their prey. Other adaptations include stealth and [[aggressive mimicry]] that improve hunting efficiency. Predation has a powerful [[selection pressure|selective effect]] on prey, causing them to develop [[antipredator adaptation]]s such as [[aposematism|warning coloration]], [[alarm call]]s and other [[signalling theory|signals]], [[camouflage]] and defensive spines and chemicals.<ref>{{cite web |last1=Bar-Yam |title=Predator-Prey Relationships |url=http://necsi.edu/projects/evolution/co-evolution/pred-prey/co-evolution_predator.html |publisher=New England Complex Systems Institute |access-date=7 September 2018}}</ref><ref name="RSM2012">{{cite web |title=Predator & Prey: Adaptations |url=https://royalsaskmuseum.ca/pub/Lesson%20Plans/Resources/Predator%20and%20Prey%20Adaptations.pdf |publisher=Royal Saskatchewan Museum |access-date=19 April 2018 |date=2012 |archive-date=3 April 2018 |archive-url=https://web.archive.org/web/20180403131659/http://royalsaskmuseum.ca/pub/Lesson%20Plans/Resources/Predator%20and%20Prey%20Adaptations.pdf |url-status=dead }}</ref><ref>{{cite book |last=Vermeij |first=Geerat J. |title=Evolution and Escalation: An Ecological History of Life |url=https://books.google.com/books?id=M3pCQ6ks5PEC&pg=PR11 |year=1993 |publisher=Princeton University Press |isbn=978-0-691-00080-0 |pages=11 and passim}}</ref> Predation has been a major driver of evolution since at least the [[Cambrian]] period.<ref name="Bengtson2002">{{cite book |author=Bengtson, S. |year=2002 |contribution=Origins and early evolution of predation |title=The fossil record of predation. The Paleontological Society Papers 8 |editor-last1=Kowalewski |editor-first1=M. |editor-last2=Kelley |editor-first2=P. H. |pages=289–317 |publisher=The Paleontological Society |url=http://www.nrm.se/download/18.4e32c81078a8d9249800021552/Bengtson2002predation.pdf}}</ref>

==== Pollination ====
[[File:Hummingbird hawkmoth a.jpg|thumb|upright|[[Pollination]] has driven the [[coevolution]] of [[flowering plant]]s and their animal [[pollinator]]s for over 100 million years.]]
{{see also|Pollination|Plant-pollinator interactions}}
In pollination, pollinators including [[insect]]s ([[entomophily]]), some [[bird]]s ([[ornithophily]]), and some [[bat]]s, transfer [[pollen]] from a male flower part to a female flower part, enabling [[fertilisation]], in return for a reward of pollen or nectar.<ref name="crop_Type">{{Cite web | title=Types of Pollination, Pollinators and Terminology | work=CropsReview.Com | access-date=2015-10-20 | url=http://www.cropsreview.com/types-of-pollination.html}}</ref> The partners have coevolved through geological time; in the case of insects and [[flowering plant]]s, the coevolution has continued for over 100 million years. Insect-pollinated flowers are [[adaptation|adapted]] with shaped structures, bright colours, patterns, scent, nectar, and sticky pollen to attract insects, guide them to pick up and deposit pollen, and reward them for the service. Pollinator insects like [[bee]]s are adapted to detect flowers by colour, pattern, and scent, to collect and transport pollen (such as with bristles shaped to form pollen baskets on their hind legs), and to collect and process nectar (in the case of [[honey bee]]s, making and storing [[honey]]). The adaptations on each side of the interaction match the adaptations on the other side, and have been shaped by [[natural selection]] on their effectiveness of pollination.<ref name=Lunau>{{cite journal |last1=Lunau |first1=Klaus |title=Adaptive radiation and coevolution — pollination biology case studies |journal=Organisms Diversity & Evolution |date=2004 |volume=4 |issue=3 |pages=207–224 |doi=10.1016/j.ode.2004.02.002|doi-access= |bibcode=2004ODivE...4..207L }}</ref><ref name=Pollan2001>{{cite book |author=Pollan, Michael |title=The Botany of Desire: A Plant's-eye View of the World |publisher=Bloomsbury |isbn=978-0-7475-6300-6 |date=2001|title-link=The Botany of Desire }}</ref><ref>{{cite journal | last1=Ehrlich | first1=Paul R. |author1-link=Paul R. Ehrlich | last2=Raven | first2=Peter H. | author2-link=Peter H. Raven | year=1964 | title=Butterflies and Plants: A Study in Coevolution | journal=Evolution | volume=18 | issue=4 | pages=586–608 | doi=10.2307/2406212| jstor=2406212 }}</ref>

==== Seed dispersal ====
{{Main|Seed dispersal}}
'''Seed dispersal''' is the movement, spread or transport of [[seed]]s away from the parent plant. Plants have limited mobility and rely upon a variety of [[dispersal vector]]s to transport their propagules, including both [[abiotic]] vectors such as the wind and living ([[Biotic component|biotic]]) vectors like birds.<ref>{{Cite journal|last1=Lim|first1=Ganges|last2=Burns|first2=Kevin C.|date=2021-11-24|title=Do fruit reflectance properties affect avian frugivory in New Zealand?|url=https://doi.org/10.1080/0028825X.2021.2001664|journal=New Zealand Journal of Botany|volume=60 |issue=3 |pages=319–329|doi=10.1080/0028825X.2021.2001664|s2cid=244683146|issn=0028-825X}}</ref> Seeds can be dispersed away from the parent plant individually or collectively, as well as dispersed in both space and time. The patterns of seed dispersal are determined in large part by the dispersal mechanism and this has important implications for the demographic and genetic structure of plant populations, as well as migration patterns and species interactions. There are five main modes of seed dispersal: [[Gravitation|gravity]], wind, ballistic, water, and by animals. Some plants are [[serotinous]] and only disperse their seeds in response to an environmental stimulus. Dispersal involves the letting go or detachment of a diaspore from the main parent plant.<ref>{{Cite journal|last=Academic Search Premier|date=1970|title=Annual review of ecology and systematics|url=https://www.worldcat.org/oclc/1091085133|journal=Annual Review of Ecology and Systematics|language=English|oclc=1091085133}}</ref>

=== Long-term interactions (symbioses) ===

{{main|Symbiosis}}

[[File:Symbiotic relationships diagram.svg|thumb|upright=2|The six possible types of [[symbiosis|symbiotic relationship]], from mutual benefit to mutual harm]]

The six possible types of [[symbiosis]] are mutualism, commensalism, parasitism, neutralism, amensalism, and competition.<ref>*{{citation |last=Douglas |first=Angela |title=The Symbiotic Habit |publisher=Princeton University Press |location=New Jersey |year=2010 |isbn=978-0-691-11341-8 |pages=5–12}}</ref> These are distinguished by the degree of benefit or harm they cause to each partner.<ref>{{cite journal |last1=Wootton |first1=J.T. |last2=Emmerson |first2=M. |title=Measurement of Interaction Strength in Nature |journal=[[Annual Review of Ecology, Evolution, and Systematics]] |volume=36 |pages=419–444 |year=2005 |jstor=30033811 |doi=10.1146/annurev.ecolsys.36.091704.175535}}</ref>

==== Mutualism ====

{{main|Mutualism (biology)}}

Mutualism is an interaction between two or more species, where species derive a mutual benefit, for example an increased [[carrying capacity]]. Similar interactions within a species are known as [[Co-operation (evolution)|co-operation]]. Mutualism may be classified in terms of the closeness of association, the closest being symbiosis, which is often confused with mutualism. One or both species involved in the interaction may be [[wikt:obligate|obligate]], meaning they cannot survive in the short or long term without the other species. Though mutualism has historically received less attention than other interactions such as predation,<ref name="Begon96">Begon, M., J.L. Harper and C.R. Townsend. 1996. ''Ecology: individuals, populations, and communities'', Third Edition. Blackwell Science, Cambridge, Massachusetts.</ref> it is an important subject in ecology. Examples include [[cleaning symbiosis]], [[gut flora]], [[Müllerian mimicry]], and [[nitrogen fixation]] by bacteria in the root nodules of [[legumes]].{{cn|date=May 2023}}

==== Commensalism ====

{{main|Commensalism}}

Commensalism benefits one organism and the other organism is neither benefited nor harmed. It occurs when one organism takes benefits by interacting with another organism by which the host organism is not affected. A good example is a [[remora]] living with a [[manatee]]. Remoras feed on the manatee's faeces. The manatee is not affected by this interaction, as the remora does not deplete the manatee's resources.<ref name="Echeneid-sirenian associations, with information on sharksucker diet">{{cite journal |last1=Williams E, Mignucci, Williams L & Bonde |title=Echeneid-sirenian associations, with information on sharksucker diet |journal=Journal of Fish Biology |date=November 2003 |volume=5 |issue=63 |pages=1176–1183 |doi=10.1046/j.1095-8649.2003.00236.x |bibcode=2003JFBio..63.1176W |url=https://www.researchgate.net/publication/253117307 |access-date=17 June 2020}}</ref>

==== Parasitism ====

{{main|Parasitism}}

Parasitism is a relationship between species, where one organism, the [[parasite]], lives on or in another organism, the [[Host (biology)|host]], causing it some harm, and is [[adaptation (biology)|adapted]] structurally to this way of life.<ref>{{cite book | last=Poulin | first=Robert | author-link=Robert Poulin (zoologist) | title=Evolutionary Ecology of Parasites | publisher=Princeton University Press | year=2007 | isbn=978-0-691-12085-0 | pages=[https://archive.org/details/evolutionaryecol0000poul/page/4 4–5] | url=https://archive.org/details/evolutionaryecol0000poul/page/4 }}</ref> The parasite either feeds on the host, or, in the case of intestinal parasites, consumes some of its food.<ref>{{cite journal |title=Current usage of symbiosis and associated terminology |last1=Martin |first1= Bradford D. |last2= Schwab |first2=Ernest |year=2013 |journal=International Journal of Biology |volume=5 |issue=1 |pages=32–45 |doi=10.5539/ijb.v5n1p32|doi-access=free }}</ref>

==== Neutralism ====

Neutralism (a term introduced by [[Eugene Odum]])<ref>Toepfer, G. "Neutralism". In: ''BioConcepts''. [http://www.biological-concepts.com/views/search.php?term=1441 link].</ref> describes the relationship between two species that interact but do not affect each other. Examples of true neutralism are virtually impossible to prove; the term is in practice used to describe situations where interactions are negligible or insignificant.<ref>(Morris et al., 2013)</ref><ref>{{Cite journal |last=Lidicker |first=William Z. |date=1979 |title=A Clarification of Interactions in Ecological Systems |url=https://www.jstor.org/stable/1307540 |journal=BioScience |volume=29 |issue=8 |pages=475–477 |doi=10.2307/1307540 |jstor=1307540 |issn=0006-3568}}</ref>

==== Amensalism ====

{{further|Amensalism}}
[[File:Eucalypts inhibiting grass growth example of amensalism and allelopathy.jpg|thumb|Leaf litter from these eucalypts contains chemicals which inhibit grass growth near the trees]]
[[Amensalism]] (a term introduced by [[Edward Haskell]])<ref>Toepfer, G. "Amensalism". In: ''BioConcepts''. [http://www.biological-concepts.com/views/search.php?term=1440 link].</ref> is an interaction where an organism inflicts harm to another organism without any costs or benefits received by itself.<ref name="Willey, Joanne M. 2011">{{cite book |last1=Willey |first1=Joanne M. |last2=Sherwood |first2=Linda M. |last3=Woolverton |first3=Cristopher J. |year=2013 |title=Prescott's Microbiology |edition=9th |pages=713–38 |isbn=978-0-07-751066-4}}</ref> This unidirectional process can be based on the release of one or more chemical compounds by one organism that negatively affect another, called [[allelopathy]].<ref>{{Citation |last1=Scavo |first1=Aurelio |title=Allelopathy: Principles and Basic Aspects for Agroecosystem Control |date=2018 |work=Sustainable Agriculture Reviews 28: Ecology for Agriculture |pages=47–101 |editor-last=Gaba |editor-first=Sabrina |url=https://link.springer.com/chapter/10.1007/978-3-319-90309-5_2 |access-date=2025-04-18 |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-319-90309-5_2 |isbn=978-3-319-90309-5 |last2=Restuccia |first2=Alessia |last3=Mauromicale |first3=Giovanni |editor2-last=Smith |editor2-first=Barbara |editor3-last=Lichtfouse |editor3-first=Eric}}</ref> One example of this is the microbial production of antibiotics that can inhibit or kill other, susceptible microorganisms. Another example is leaf litter from trees such as ''[[Pinus ponderosa]]''<ref>{{Cite journal |last1=Singh |first1=Surendra P. |last2=Inderjit |last3=Singh |first3=Jamuna S. |last4=Majumdar |first4=Sudipto |last5=Moyano |first5=Jaime |last6=Nuñez |first6=Martin A. |last7=Richardson |first7=David M. |date=2018 |title=Insights on the persistence of pines (Pinus species) in the Late Cretaceous and their increasing dominance in the Anthropocene |journal=Ecology and Evolution |language=en |volume=8 |issue=20 |pages=10345–10359 |doi=10.1002/ece3.4499 |issn=2045-7758 |pmc=6206191 |pmid=30398478|bibcode=2018EcoEv...810345S }}</ref> or ''[[Eucalyptus|Eucalyptus spp.]]''<ref>{{Cite journal |last1=Chu |first1=Chaojun |last2=Mortimer |first2=P. E. |last3=Wang |first3=Hecong |last4=Wang |first4=Yongfan |last5=Liu |first5=Xubing |last6=Yu |first6=Shixiao |date=2014-07-01 |title=Allelopathic effects of Eucalyptus on native and introduced tree species |url=https://www.sciencedirect.com/science/article/abs/pii/S0378112714001492 |journal=Forest Ecology and Management |volume=323 |pages=79–84 |doi=10.1016/j.foreco.2014.03.004 |bibcode=2014ForEM.323...79C |issn=0378-1127}}</ref> preventing the establishment and growth of other plant species.

A clear case of amensalism is where [[Ungulate|hoofed mammals]] trample grass. Whilst the presence of the grass causes negligible detrimental effects to the animal's hoof, the grass suffers from being crushed. Amensalism also includes strongly asymmetrical competitive interactions, such as has been observed between the Spanish [[ibex]] and [[Weevil|weevils]] of the genus ''[[Timarcha]],'' which both feed upon the same type of shrub. Whilst the presence of the weevil has almost no influence on food availability, the presence of ibex has an enormous detrimental effect on weevil numbers, as they eat the shrub and incidentally ingest the weevils.<ref name="Gómez J., González-Megías A. 2002">{{cite journal |last1=Gómez |first1=José M. |last2=González-Megías |first2=Adela |year=2002 |title=Asymmetrical interactions between ungulates and phytophagous insects: Being different matters |journal=Ecology |volume=83 |issue=1 |pages=203–11 |doi=10.1890/0012-9658(2002)083[0203:AIBUAP]2.0.CO;2}}</ref>

==== Competition ====

{{main|Competition (biology)}}

[[File:Hirschkampf.jpg|thumb|Male-male interference competition in [[red deer]]]]

Competition can be defined as an interaction between [[organism]]s or species, in which the [[fitness (biology)|fitness]] of one is lowered by the presence of another. Competition is often for a resource such as [[food]], [[water]], or [[territory (animal)|territory]] in [[Limiting factor|limited]] supply, or for access to females for reproduction.<ref name="Begon96"/> Competition among members of the same species is known as [[intraspecific competition]], while competition between individuals of different species is known as [[interspecific competition]]. According to the [[competitive exclusion principle]], species less suited to compete for resources should either [[adaptation|adapt]] or [[extinction|die out]].<ref name=hardin60>{{cite journal |author=Hardin, Garrett |title=The competitive exclusion principle |journal=Science |volume=131 |pages=1292–1297 |issue=3409 |year=1960 |url=http://www.esf.edu/efb/schulz/seminars/hardin.pdf |doi=10.1126/science.131.3409.1292 |pmid=14399717 |bibcode=1960Sci...131.1292H |access-date=2018-10-04 |archive-date=2017-11-17 |archive-url=https://web.archive.org/web/20171117235048/http://www.esf.edu/efb/schulz/seminars/hardin.pdf |url-status=dead }}</ref><ref name=Pocheville2015>{{cite book | last=Pocheville | first=Arnaud | year=2015 | chapter=The Ecological Niche: History and Recent Controversies | chapter-url=https://www.academia.edu/6188833 | editor1-last=Heams | editor1-first=Thomas | editor2-last=Huneman | editor2-first=Philippe | editor3-last=Lecointre | editor3-first=Guillaume |display-editors=3 | editor4-last=Silberstein | editor4-first=Marc | title=Handbook of Evolutionary Thinking in the Sciences | location=Dordrecht | publisher=Springer | publication-date=2015 | pages=547–586 | isbn=978-94-017-9014-7}}</ref> This competition within and between species for resources plays a critical role in [[natural selection]].<ref name="SahneyBentonFerry2010LinksDiversityVertebrates">{{cite journal |last1=Sahney |first1=Sarda |last2=Benton |first2=Michael J. |author-link2=Michael Benton |last3=Ferry |first3=Paul A. |date=23 August 2010 |title=Links between global taxonomic diversity, ecological diversity and the expansion of vertebrates on land |journal=[[Biology Letters]] |volume=6 |issue=4 |pages=544–547 |doi=10.1098/rsbl.2009.1024 |pmc=2936204 |pmid=20106856 }}</ref>

== Classification based on effect on fitness ==

Biotic interactions can vary in intensity (strength of interaction), and frequency (number of interactions in a given time).<ref>{{Cite journal |last1=Caruso |first1=Tancredi |last2=Trokhymets |first2=Vladlen |last3=Bargagli |first3=Roberto |last4=Convey |first4=Peter |date=2012-10-20 |title=Biotic interactions as a structuring force in soil communities: evidence from the micro-arthropods of an Antarctic moss model system |url=http://dx.doi.org/10.1007/s00442-012-2503-9 |journal=Oecologia |volume=172 |issue=2 |pages=495–503 |doi=10.1007/s00442-012-2503-9 |pmid=23086506 |s2cid=253978982 |issn=0029-8549}}</ref><ref>{{Cite journal |last1=Morales-Castilla |first1=Ignacio |last2=Matias |first2=Miguel G. |last3=Gravel |first3=Dominique |last4=Araújo |first4=Miguel B. |date=June 2015 |title=Inferring biotic interactions from proxies |url=http://dx.doi.org/10.1016/j.tree.2015.03.014 |journal=Trends in Ecology & Evolution |volume=30 |issue=6 |pages=347–356 |doi=10.1016/j.tree.2015.03.014 |pmid=25922148 |bibcode=2015TEcoE..30..347M |hdl=10261/344523 |hdl-access=free }}</ref> There are direct interactions when there is a physical contact between individuals or indirect interactions when there is no physical contact, that is, the interaction occurs with a resource, ecological service, toxine or growth inhibitor.<ref>{{Cite journal |last1=Wurst |first1=Susanne |last2=Ohgushi |first2=Takayuki |date=2015-05-18 |title=Do plant- and soil-mediated legacy effects impact future biotic interactions? |url=http://dx.doi.org/10.1111/1365-2435.12456 |journal=Functional Ecology |volume=29 |issue=11 |pages=1373–1382 |doi=10.1111/1365-2435.12456 |bibcode=2015FuEco..29.1373W |issn=0269-8463}}</ref> The interactions can be directly determined by individuals (incidentally) or by stochastic processes (accidentally), for instance side effects that one individual have on other.<ref>{{Cite journal |last1=Bowman |first1=William D. |last2=Swatling-Holcomb |first2=Samantha |date=2017-10-25 |title=The roles of stochasticity and biotic interactions in the spatial patterning of plant species in alpine communities |journal=Journal of Vegetation Science |volume=29 |issue=1 |pages=25–33 |doi=10.1111/jvs.12583 |s2cid=91054849 |doi-access=free }}</ref> They are divided into six major types: Competition, Antagonism, Amensalism, Neutralism, Commensalism and Mutualism.<ref>{{Cite journal |last1=Paquette |first1=Alexandra |last2=Hargreaves |first2=Anna L. |date=2021-08-27 |title=Biotic interactions are more often important at species' warm versus cool range edges |url=http://dx.doi.org/10.1111/ele.13864 |journal=Ecology Letters |volume=24 |issue=11 |pages=2427–2438 |doi=10.1111/ele.13864 |pmid=34453406 |bibcode=2021EcolL..24.2427P |s2cid=237340810}}</ref>

==Non-trophic interactions==

{{see also|Non-trophic networks|Foundation species}}

Some examples of non-trophic interactions are habitat modification, mutualism and competition for space. It has been suggested recently that non-trophic interactions can indirectly affect [[food web]] topology and [[trophic dynamics]] by affecting the species in the network and the strength of trophic links.<ref name="Kefi2015">{{Cite journal |last1=Kéfi |first1=Sonia |author-link=Sonia Kéfi |last2=Berlow |first2=Eric L. |last3=Wieters |first3=Evie A. |last4=Joppa |first4=Lucas N. |last5=Wood |first5=Spencer A. |last6=Brose |first6=Ulrich |last7=Navarrete |first7=Sergio A. |date=January 2015 |title=Network structure beyond food webs: mapping non-trophic and trophic interactions on Chilean rocky shores |journal=Ecology |volume=96 |issue=1 |pages=291–303 |doi=10.1890/13-1424.1 |pmid=26236914|doi-access=free |bibcode=2015Ecol...96..291K }}</ref><ref name="vanderZee2016">{{Cite journal |last1=van der Zee |first1=Els M. |last2=Angelini |first2=Christine |last3=Govers |first3=Laura L. |last4=Christianen |first4=Marjolijn J. A. |last5=Altieri |first5=Andrew H. |last6=van der Reijden |first6=Karin J. |last7=Silliman |first7=Brian R. |last8=van de Koppel |first8=Johan |last9=van der Geest |first9=Matthijs |last10=van Gils |first10=Jan A. |last11=van der Veer |first11=Henk W. |date=2016-03-16 |title=How habitat-modifying organisms structure the food web of two coastal ecosystems |journal=Proceedings. Biological Sciences |volume=283 |issue=1826 |pages=20152326 |doi=10.1098/rspb.2015.2326 |pmc=4810843 |pmid=26962135}}</ref><ref name="Sanders2014">{{Cite journal |last1=Sanders |first1=Dirk |last2=Jones |first2=Clive G. |last3=Thébault |first3=Elisa |last4=Bouma |first4=Tjeerd J. |last5=van der Heide |first5=Tjisse |last6=van Belzen |first6=Jim |last7=Barot |first7=Sébastien |date=May 2014 |title=Integrating ecosystem engineering and food webs |journal=Oikos |language=en |volume=123 |issue=5 |pages=513–524 |doi=10.1111/j.1600-0706.2013.01011.x|bibcode=2014Oikos.123..513S }}</ref> It is necessary to integrate trophic and non-trophic interactions in ecological network analyses.<ref name=Sanders2014 /><ref>{{Cite journal |last1=Kéfi |first1=Sonia |author-link=Sonia Kéfi |last2=Berlow |first2=Eric L. |last3=Wieters |first3=Evie A. |last4=Navarrete |first4=Sergio A. |last5=Petchey |first5=Owen L. |last6=Wood |first6=Spencer A. |last7=Boit |first7=Alice |last8=Joppa |first8=Lucas N. |last9=Lafferty |first9=Kevin D. |last10=Williams |first10=Richard J. |last11=Martinez |first11=Neo D. |date=April 2012 |title=More than a meal... integrating non-feeding interactions into food webs |journal=Ecology Letters |volume=15 |issue=4 |pages=291–300 |doi=10.1111/j.1461-0248.2011.01732.x |pmid=22313549|doi-access=free |bibcode=2012EcolL..15..291K }}</ref><ref>{{Cite journal |last1=Pilosof |first1=Shai |last2=Porter |first2=Mason A. |last3=Pascual |first3=Mercedes |last4=Kéfi |first4=Sonia |author-link4=Sonia Kéfi |date=2017-03-23 |title=The multilayer nature of ecological networks |journal=Nature Ecology & Evolution |volume=1 |issue=4 |pages=101 |doi=10.1038/s41559-017-0101 |pmid=28812678|arxiv=1511.04453 |bibcode=2017NatEE...1..101P |s2cid=11387365 }}</ref> The few empirical studies that address this suggest food web structures (network topologies) can be strongly influenced by species interactions outside the trophic network.<ref name=Kefi2015 /><ref name=vanderZee2016 /><ref name="Christianen2017">{{Cite journal |last1=Christianen |first1=Mja |last2=van der Heide |first2=T |last3=Holthuijsen |first3=Sj |last4=van der Reijden |first4=Kj |last5=Borst |first5=Acw |last6=Olff |first6=H |date=September 2017 |title=Biodiversity and food web indicators of community recovery in intertidal shellfish reefs |journal=Biological Conservation |language=en |volume=213 |pages=317–324 |doi=10.1016/j.biocon.2016.09.028|bibcode=2017BCons.213..317C |url=https://research.rug.nl/en/publications/biodiversity-and-food-web-indicators-of-community-recovery-in-intertidal-shellfish-reefs(05a2a9ea-5041-4c44-b44b-a21ba29c4799).html }}</ref> However these studies include only a limited number of coastal systems, and it remains unclear to what extent these findings can be generalized. Whether non-trophic interactions typically affect specific species, trophic levels, or functional groups within the food web, or, alternatively, indiscriminately mediate species and their trophic interactions throughout the network has yet to be resolved. [[Sessility (motility)|sessile]] species with generally low trophic levels seem to benefit more than others from non-trophic facilitation,<ref>{{Cite journal |last1=Miller |first1=Robert J. |last2=Page |first2=Henry M. |last3=Reed |first3=Daniel C. |date=December 2015 |title=Trophic versus structural effects of a marine foundation species, giant kelp (Macrocystis pyrifera) |journal=Oecologia |volume=179 |issue=4 |pages=1199–1209 |doi=10.1007/s00442-015-3441-0 |pmid=26358195|bibcode=2015Oecol.179.1199M |s2cid=18578916 }}</ref> though facilitation benefits higher trophic and more mobile species as well.<ref name=Christianen2017 /><ref name="vanderZee2015">{{Cite journal |last1=van der Zee |first1=Els M. |last2=Tielens |first2=Elske |last3=Holthuijsen |first3=Sander |last4=Donadi |first4=Serena |last5=Eriksson |first5=Britas Klemens |last6=van der Veer |first6=Henk W. |last7=Piersma |first7=Theunis |last8=Olff |first8=Han |last9=van der Heide |first9=Tjisse |date=April 2015 |title=Habitat modification drives benthic trophic diversity in an intertidal soft-bottom ecosystem |journal=Journal of Experimental Marine Biology and Ecology |volume=465 |pages=41–48 |doi=10.1016/j.jembe.2015.01.001 |bibcode=2015JEMBE.465...41V |url=http://www.vliz.be/imisdocs/publications/83/279583.pdf }}</ref><ref name="Angelini2014">{{Cite journal |last1=Angelini |first1=Christine |last2=Silliman |first2=Brian R. |date=January 2014 |title=Secondary foundation species as drivers of trophic and functional diversity: evidence from a tree-epiphyte system |journal=Ecology |volume=95 |issue=1 |pages=185–196 |doi=10.1890/13-0496.1 |pmid=24649658|bibcode=2014Ecol...95..185A }}</ref><ref name="Borst2018">{{Cite journal |last1=Borst |first1=Annieke C. W. |last2=Verberk |first2=Wilco C. E. P. |last3=Angelini |first3=Christine |last4=Schotanus |first4=Jildou |last5=Wolters |first5=Jan-Willem |last6=Christianen |first6=Marjolijn J. A. |last7=Zee |first7=Els M. van der |last8=Derksen-Hooijberg |first8=Marlous |last9=Heide |first9=Tjisse van der |date=2018-08-31 |title=Foundation species enhance food web complexity through non-trophic facilitation |journal=PLOS ONE |volume=13 |issue=8 |pages=e0199152 |doi=10.1371/journal.pone.0199152 |doi-access=free |pmc=6118353 |pmid=30169517|bibcode=2018PLoSO..1399152B }}</ref>

==See also==
{{div col|colwidth=24em}}
* [[Altruism (biology)]]
* [[Animal sexual behaviour]]
* [[Biological pump]] – interaction between marine animals and carbon forms
* [[Cheating (biology)]]
* [[Collective animal behavior]]
* [[Detritivory]]
* [[Epibiont]]
* [[Evolving digital ecological network]]
* [[Food chain]]
* [[Kin selection]]
* [[Microbial cooperation]]
* [[Microbial loop]]
* [[Quorum sensing]]
* [[Spite (game theory)]]
* [[Swarm behaviour]]
{{div col end}}

==Notes==
{{notelist}}

==References==
{{reflist}}

==Further reading==
* Snow, B. K. & Snow, D. W. (1988). ''Birds and berries: a study of an ecological interaction''. Poyser, London {{ISBN|0-85661-049-6}}

==External links==
{{commons category|Ecological interactions}}
*[https://www.globalbioticinteractions.org/ Global Biotic Interactions (GloBI)] - Open access to finding species interaction data

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