Editing Mutualism (biology) (section)

==Types==

=== Resource-resource relationships ===
Mutualistic relationships can be thought of as a form of "[[Biology|biological]] barter"<ref name="Ollerton06" /> in [[mycorrhizal]] associations between plant [[root]]s and [[fungi]], with the plant providing [[carbohydrates]] to the [[fungus]] in return for primarily [[phosphate]] but also [[nitrogenous]] compounds. Other examples include [[rhizobia]] bacteria that fix nitrogen for [[leguminous]] plants (family Fabaceae) in return for energy-containing [[carbohydrates]].<ref>{{cite journal |last1=Denison |first1=RF |last2=Kiers |first2=ET |year=2004 |title= Why are most rhizobia beneficial to their plant hosts, rather than parasitic |journal= [[Microbes and Infection]] |volume=6 |issue=13 |pages= 1235–9 |doi= 10.1016/j.micinf.2004.08.005|pmid=15488744 |doi-access=free }}</ref> Metabolite exchange between multiple mutualistic species of [[bacteria]] has also been observed in a process known as [[Syntrophy|cross-feeding]].<ref>{{Cite journal |last=Schink |first=Bernhard |date=2002-12-01 |title=Synergistic interactions in the microbial world |url=https://doi.org/10.1023/A:1020579004534 |journal=Antonie van Leeuwenhoek|volume=81 |issue=1 |pages=257–261 |doi=10.1023/A:1020579004534 |pmid=12448724 |s2cid=9310406 |issn=1572-9699}}</ref><ref>{{Cite journal |last1=Chacón |first1=Jeremy M. |last2=Hammarlund |first2=Sarah P. |last3=Martinson |first3=Jonathan N.V. |last4=Smith |first4=Leno B. |last5=Harcombe |first5=William R. |date=2021-11-02 |title=The Ecology and Evolution of Model Microbial Mutualisms |journal=Annual Review of Ecology, Evolution, and Systematics|volume=52 |issue=1 |pages=363–384 |doi=10.1146/annurev-ecolsys-012121-091753 |s2cid=239694099 |issn=1543-592X|doi-access=free }}</ref>

===Service-resource relationships===
[[File:Impala mutualism with birds wide.jpg|thumb|The [[red-billed oxpecker]] eats ticks on the [[impala]]'s coat, in a [[cleaning symbiosis]].]]

Service-resource relationships are common. Three important types are [[pollination]], [[cleaning]] symbiosis, and [[zoochory]].

In pollination, a plant trades food resources in the form of [[nectar]] or [[pollen]] for the service of pollen dispersal. However, daciniphilous ''[[Bulbophyllum]]'' orchid species trade [[sex pheromone]] precursor or booster components via floral [[synomone]]s/attractants in a true mutualistic interactions with males of [[Dacini]] fruit flies (Diptera: Tephritidae: Dacinae).<ref>See also [[Attractant]] related to synomone; and references therein</ref><ref>{{cite journal |last1=Tan |first1=K.H. |last2=Nishida |first2=R. |title=Mutual reproductive benefits between a wild orchid, ''Bulbophyllum patens'', and Bactrocera fruit flies via a floral synomone |journal=Journal of Chemical Ecology |volume=26 |issue= 2|pages=533–546 |date=2000 |doi=10.1023/A:1005477926244 |bibcode=2000JCEco..26..533T }}</ref>

[[Phagophile]]s feed (resource) on [[ectoparasite]]s, thereby providing anti-pest service, as in [[cleaning symbiosis]].
''[[Elacatinus]]'' and ''[[Gobiosoma]]'', genera of [[Goby|gobies]], feed on ectoparasites of their clients while cleaning them.<ref>{{Cite journal|author1=M.C. Soares |author2=I.M. Côté |author3=S.C. Cardoso & R.Bshary |date=August 2008| title=The cleaning goby mutualism: a system without punishment, partner switching or tactile stimulation | journal =Journal of Zoology| volume =276 |issue =3 | pages =306–312 | doi=10.1111/j.1469-7998.2008.00489.x|url=http://doc.rero.ch/record/28974/files/Soares_Marta_C._-_The_cleaning_goby_mutualism_a_system_without_20120417.pdf }}</ref>

Zoochory is the dispersal of the seeds of plants by animals. This is similar to pollination in that the plant produces food resources (for example, fleshy fruit, overabundance of seeds) for animals that disperse the seeds (service). Plants may advertise these resources using colour<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> and a variety of other fruit characteristics, e.g., scent. Fruit of the [[aardvark cucumber]] ''(Cucumis humifructus)'' is buried so deeply that the plant is solely reliant upon the [[Aardvark#Head|aardvark's keen sense of smell]] to detect its ripened fruit, extract, consume and then scatter its seeds;<ref>{{cite book|last1=van Rheede van Oudtshoorn|first1=Karen|last2=van Rooyen |first2=Margaretha W. |title=Dispersal Biology of Desert Plants|year=1998|publisher=Springer |isbn=978-3-540-64886-4 |chapter=Restriction of Dispersal Due to Reduction of Dispersal Structures |chapter-url=https://link.springer.com/chapter/10.1007/978-3-662-03561-0_6 |page=118|doi=10.1007/978-3-662-03561-0_6 |url=https://books.google.com/books?id=bdTfBfsl-DcC&q=aardvark|accessdate=2023-01-28}}</ref> ''C.&nbsp;humifructus''{{'}}s geographical range is thus restricted to that of the aardvark.

Another type is [[ant]] protection of [[aphid]]s, where the aphids trade [[sugar]]-rich [[Honeydew (secretion)|honeydew]] (a by-product of their mode of feeding on plant [[sap]]) in return for defense against [[predator]]s such as [[ladybug]]s.{{citation needed|date=June 2024}}

===Service-service relationships===
[[Image:Common clownfish curves dnsmpl.jpg|thumb|[[Ocellaris clownfish]] and [[Heteractis magnifica|Ritter's sea anemone]]s live in a mutual service-service symbiosis, the fish driving off [[butterflyfish]] and the anemone's tentacles protecting the fish from predators.]]

Strict service-service interactions are very rare, for reasons that are far from clear.<ref name="Ollerton06" /> One example is the relationship between [[sea anemone]]s and [[anemone fish]] in the family [[Pomacentridae]]: the anemones provide the fish with protection from [[predator]]s (which cannot tolerate the stings of the anemone's tentacles) and the fish defend the anemones against [[butterflyfish]] (family [[Chaetodontidae]]), which eat anemones. However, in common with many mutualisms, there is more than one aspect to it: in the anemonefish-anemone mutualism, waste [[ammonia]] from the fish feeds the [[symbiotic]] [[algae]] that are found in the anemone's tentacles.<ref>{{cite journal | last1=Porat | first1=D. | last2=Chadwick-Furman | first2=N. E. | year=2004 | title=Effects of anemonefish on giant sea anemones: expansion behavior, growth, and survival | journal=[[Hydrobiologia]] | volume=530 | issue=1–3| pages=513–520 | doi=10.1007/s10750-004-2688-y | bibcode=2004HyBio.530..513P | s2cid=2251533 }}</ref><ref>{{cite journal | last1=Porat | first1=D. | last2=Chadwick-Furman | first2=N. E. | year=2005 | title=Effects of anemonefish on giant sea anemones: ammonium uptake, zooxanthella content and tissue regeneration | journal=Mar. Freshw. Behav. Phys. | volume=38 | issue=1 | pages=43–51 | doi= 10.1080/10236240500057929| bibcode=2005MFBP...38...43P | s2cid=53051081 }}</ref> Therefore, what appears to be a service-service mutualism in fact has a service-resource component. A second example is that of the relationship between some [[ants]] in the genus ''[[Pseudomyrmex]]'' and trees in the [[genus]] ''[[Acacia]]'', such as the [[whistling thorn]] and [[bullhorn acacia]]. The [[ants]] nest inside the plant's thorns. In exchange for shelter, the ants protect acacias from attack by [[herbivores]] (which they frequently eat when those are small enough, introducing a resource component to this service-service relationship) and competition from other plants by trimming back vegetation that would shade the acacia. In addition, another service-resource component is present, as the ants regularly feed on [[lipid]]-rich food-bodies called [[Beltian bodies]] that are on the ''Acacia'' plant.<ref>{{Cite web|url=https://www2.palomar.edu/users/warmstrong/acacia.htm|title=Swollen Thorn Acacias|website=www2.palomar.edu|access-date=2019-02-22|archive-date=27 June 2018|archive-url=https://web.archive.org/web/20180627005136/http://www2.palomar.edu/users/warmstrong/acacia.htm|url-status=dead}}</ref>

In the [[neotropics]], the ant ''[[Myrmelachista schumanni]]'' makes its nest in special cavities in ''[[Duroia hirsuta]]''. Plants in the vicinity that belong to other species are killed with [[formic acid]]. This selective gardening can be so aggressive that small areas of the rainforest are dominated by ''Duroia hirsute''. These peculiar patches are known by local people as "[[devil's garden]]s".<ref name="Piper07">[[Ross Piper|Piper, Ross]] (2007), ''Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals'', [[Greenwood Press (publisher)|Greenwood Press]].</ref>

In some of these relationships, the cost of the ant's protection can be quite expensive. ''[[Cordia]]'' sp. trees in the [[Amazon rainforest]] have a kind of partnership with ''[[Allomerus]]'' sp. ants, which make their nests in modified leaves. To increase the amount of living space available, the ants will destroy the tree's flower buds. The flowers die and leaves develop instead, providing the ants with more dwellings. Another type of ''Allomerus'' sp. ant lives with the ''[[Hirtella]]'' sp. tree in the same forests, but in this relationship, the tree has turned the tables on the ants. When the tree is ready to produce flowers, the ant abodes on certain branches begin to wither and shrink, forcing the occupants to flee, leaving the tree's flowers to develop free from ant attack.<ref name="Piper07" />

The term "species group" can be used to describe the manner in which individual organisms group together. In this non-taxonomic context one can refer to "same-species groups" and "mixed-species groups." While same-species groups are the norm, examples of mixed-species groups abound. For example, zebra (''[[Equus burchelli]]'') and wildebeest (''[[Connochaetes taurinus]]'') can remain in association during periods of long distance [[wikt:migration|migration]] across the [[Serengeti]] as a strategy for thwarting predators. ''[[Cercopithecus mitis]]'' and ''[[Cercopithecus ascanius]]'', species of monkey in the [[Kakamega Forest]] of [[Kenya]], can stay in close proximity and travel along exactly the same routes through the forest for periods of up to 12 hours. These mixed-species groups cannot be explained by the coincidence of sharing the same habitat. Rather, they are created by the active behavioural choice of at least one of the species in question.<ref>{{cite journal |vauthors=Tosh CR, Jackson AL, Ruxton GD |title=Individuals from different-looking animal species may group together to confuse shared predators: simulations with artificial neural networks |journal=Proc. Biol. Sci. |volume=274 |issue=1611 |pages=827–32 |date=March 2007 |pmid=17251090 |pmc=2093981 |doi=10.1098/rspb.2006.3760 }}</ref>

===Protocooperation===
[[File:2013-07-01 19-17-28-Aphidoidea.jpg|thumb|Ants and aphids]]
Protocooperation is a form of mutualism, but the cooperating species do not depend on each other for survival. The term, initially used for intraspecific interactions, was popularized by [[Eugene Odum]] (1953), although it is now rarely used.<ref>Bronstein, J. L. (2015). The study of mutualism. In: Bronstein, J. L. (ed.). ''Mutualism''. Oxford University Press, Oxford. [https://books.google.com/books?id=hbgVDAAAQBAJ link].</ref>