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| [[File:Hummingbird hawkmoth a.jpg|right|thumb|[[Hummingbird Hawkmoth]] drinking from ''[[Dianthus]]''. [[Pollination]] is a classic example of mutualism.]]
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| '''Mutualism''' is the way two organisms of different species exist in a relationship in which each individual benefits. Similar interactions within a species are known as [[co-operation (evolution)|co-operation]]. Mutualism can be contrasted with [[interspecific competition]], in which each species experiences ''reduced'' fitness, and [[Cheating (biology)|exploitation]], or [[parasitism]], in which one species benefits at the ''expense'' of the other. Mutualism is a type of [[symbiosis]]. Symbiosis is a broad category, defined to include relationships that are mutualistic, [[parasitic]], or [[Commensalism|commensal]]. Mutualism is only one ''type''.
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| A well-known example of mutualism is the relationship between [[ungulates]] (such as [[Bovines]]) and [[bacteria]] within their [[intestines]]. The ungulates benefit from the [[cellulase]] produced by the bacteria, which facilitates [[digestion]]; the bacteria benefit from having a stable supply of [[nutrients]] in the [[host (biology)|host]] environment.
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| Mutualism plays a key part in [[ecology]]. For example, mutualistic interactions are vital for terrestrial [[ecosystem]] function as more than 48% of land plants rely on [[mycorrhizal]] relationships with [[fungi]] to provide them with inorganic compounds and trace elements. In addition, mutualism is thought to have driven the evolution of much of the biological diversity we see, such as [[flower]] forms (important for [[pollination]] mutualisms) and [[co-evolution]] between groups of species.<ref>Thompson, J. N. 2005 The geographic mosaic of coevolution. Chicago, IL: University of Chicago Press.</ref> However mutualism has historically received less attention than other interactions such as [[predation]] and [[parasitism]].<ref>Bronstein, JL. 1994. Our current understand of mutualism. [[Quarterly Review of Biology]] 69 (1): 31-51 March 1994</ref><ref>Begon, M., J.L. Harper, and C.R. Townsend. 1996. ''[[Ecology: individuals, populations, and communities]]'', Third Edition. Blackwell Science Ltd., Cambridge, Massachusetts, USA.</ref>
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| Measuring the exact [[Fitness (biology)|fitness]] benefit to the individuals in a mutualistic relationship is not always straightforward, particularly when the individuals can receive benefits from a variety of species, for example most plant-[[pollinator]] mutualisms. It is therefore common to categorise mutualisms according to the closeness of the association, using terms such as [[obligate]] and [[facultative]]. Defining "closeness," however, is also problematic. It can refer to mutual dependency (the species cannot live without one another) or the biological intimacy of the relationship in relation to physical closeness (''e.g.'', one species living within the tissues of the other species).<ref name="Ollerton06">Ollerton, J. 2006. "Biological Barter": Patterns of Specialization Compared across Different Mutualisms. pp. 411-435 in: Waser, N.M. & Ollerton, J. (Eds) Plant-Pollinator Interactions: From Specialization to Generalization. University of Chicago Press.</ref>
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| ==Types of relationships==
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| Mutualistic transversals can be thought of as a form of "biological barter"<ref name="Ollerton06" /> in which species trade resources (for example [[carbohydrates]] or inorganic compounds) or services such as [[gamete]], offspring [[Biological dispersal|dispersal]], or protection from [[predator]]s.
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| ===Resource-resource relationships===
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| Resource-resource interactions, in which one type of resource is traded for a different resource, are probably the most common form of mutualism; for example [[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>Denison RF, Kiers ET 2004. Why are most rhizobia beneficial to their plant hosts, rather than parasitic. [[Microbes and Infection]] 6 (13): 1235-1239</ref>
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| ===Service-resource relationships===
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| [[File:Impala mutualim with birds wide.jpg|thumb|250px|The [[Red-billed Oxpecker]] eats ticks on the [[impala]]'s coat]]
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| Service-resource relationships are also common.
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| [[Pollination]] in which [[nectar]] or [[pollen]] (food resources) are traded for pollen dispersal (a service) or [[ant]] protection of [[aphids]], 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.
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| [[Phagophile]]s feed (resource) on [[ectoparasite]]s, thereby providing anti-pest service, as in [[cleaning symbiosis]].
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| [[Elacatinus]] and [[Gobiosoma]], genus of [[Goby|gobies]], also feed on ectoparasites of their clients while cleaning them.<ref>{{Cite journal| author=M.C. Soares, I.M. Côté, 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 | id =10.1111/j.1469-7998.2008.00489.x| url = http://onlinelibrary.wiley.com/doi/10.1111/j.1469-7998.2008.00489.x/abstract }}</ref>
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| [[Zoochory]] is an example where animals disperse the seeds of plants. 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).
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| ===Service-service relationships===
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| [[Image:Common clownfish curves dnsmpl.jpg|thumb|right|An example of mutual symbiosis is the relationship between [[Ocellaris clownfish]] that dwell among the [[tentacle]]s of [[Heteractis magnifica|Ritteri sea anemone]]s.]]
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| 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 feed the [[symbiotic]] [[algae]] that are found in the anemone's tentacles.<ref>Porat, D. & Chadwick-Furman, N. E. 2004 Effects of anemonefish on giant sea anemones: expansion behavior, growth, and survival. [[Hydrobiologia]] 530, 513–520. {{doi|10.1007/s10750-004-2688-y}}</ref><ref>Porat, D. & Chadwick-Furman, N. E. 2005 Effects of anemonefish on giant sea anemones: ammonium uptake, zooxanthella content and tissue regeneration. Mar. Freshw.Behav. Phys. 38, 43–51. {{doi|10.1080/102362405000_57929}}</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, introducing a resource component to this service-service relationship) and competition from other plants by trimming back vegetation that would shade the acacia.
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| 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.
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| In the [[Neotropics]], the ant, ''[[Myrmelachista schumanni]]'' makes its nest in special cavities in ''[[Duroia hirsuta|Duroia hirsute]]''. 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>
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| In some of these relationships, the cost of the ant’s protection can be quite expensive. ''[[Cordia]]'' sp. trees in the [[Amazon Rainforest|Amazonian 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" />
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| ==Humans and mutualism==
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| [[Image:Backing sheep at sheepdog competition.jpg|right|upright|thumb|[[Dog]]s and [[sheep]] were among the first animals to be [[Domestication|domesticated]].]]
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| Humans also engage in mutualisms with other species, including their [[gut flora]] without which they would not be able to digest food efficiently.<ref name=Sears>{{cite journal |author=Sears CL |title=A dynamic partnership: celebrating our gut flora |journal=Anaerobe |volume=11 |issue=5 |pages=247–51 |date=October 2005 |pmid=16701579 |doi=10.1016/j.anaerobe.2005.05.001 |url=}}</ref> Apparently, [[head lice]] infestations might have been beneficial for humans by fostering an [[immune]] response that helps to reduce the threat of [[body louse]] borne lethal diseases.<ref name="mutualism">{{cite journal| last = Rozsa| first = L | coauthors =Apari, P. | url =http://www.zoologia.hu/list/Why_infest.pdf | title =Why infest the loved ones – inherent human behaviour indicates former mutualism with head lice| journal =Parasitology | volume = 139 | pages = 696–700| year =2012}}</ref>
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| Some relationships between humans and [[domestication|domesticated]] animals and plants are to different degrees mutualistic. Agricultural varieties of [[maize]] are unable to reproduce without human intervention because the leafy sheath does not fall open, and the seedhead (the "corn on the cob") does not shatter to scatter the seeds naturally.<ref>{{cite web | url=http://science.jrank.org/pages/6660/Symbiosis-Symbioses-between-humans-other-species.html | title=Symbiosis - Symbioses Between Humans And Other Species | publisher=Net Industries | accessdate=December 9, 2012}}</ref>
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| In [[traditional agriculture]], some plants have mutualist as [[Companion planting|companion plants]], providing each other with shelter, soil fertility and/or natural [[pest control]]. For example, [[bean]]s may grow up [[Maize|corn]]stalks as a trellis, while fixing nitrogen in the soil for the corn, a phenomenon that is used in [[Three Sisters (agriculture)|Three Sisters farming]].<ref>{{cite encyclopedia|last=Mt. Pleasant|first=Jane|editor=John E. Staller, Robert H. Tykot, and Bruce F. Benz|encyclopedia=Histories of maize: Multidisciplinary approaches to the prehistory, linguistics, biogeography, domestication, and evolution of maize|title=The science behind the Three Sisters mound system: An agronomic assessment of an indigenous agricultural system in the northeast|year=2006|location=Amsterdam |pages=529–537}}</ref>
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| [[Borana Oromo people|Boran]] people of [[Ethiopia]] and [[Kenya]] traditionally use a whistle to call the [[Honeyguide|honey guide]] bird, though the practice is declining. If the bird is hungry and within earshot, it guides them to a bees' nest. In exchange the Borans leave some food from the nest for the bird.<ref>{{cite book | title=Keeping All the Pieces: Perspectives on Natural History and the Environment | publisher=University of Georgia Press | author=Gibbon, J Whitfield; foreword by Odum, Eugene P. | year=2010 | location=Athens, Georgia | pages=41–42}}</ref>
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| A population of [[bottlenose dolphin]]s in [[Laguna, Brazil]] coordinates, via [[body language]], with local net-using fishermen in order for both to catch schools of [[mullet (fish)|mullet]].<ref>http://news.discovery.com/animals/whales-dolphins/helpful-dolphins-120502.htm</ref>
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| ==Mathematical modeling==
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| One of the simplest frameworks for modeling species interactions is the [[Lotka–Volterra equation|Lotka-Volterra equations]]. In this model, the change in population density of the two mutualists is quantified as:
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| :<math>
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| \cfrac{dN}{dt}=r_1 N\left(1-\cfrac{N}{K1}+\beta_{12}\cfrac{M}{K_1}\right)
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| </math>
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| :<math>
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| \cfrac{dM}{dt}=r_2 M\left(1-\cfrac{M}{K2}+\beta_{21}\cfrac{N}{K_2}\right)
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| </math>
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| where,
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| * N and M = the population density
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| * r = intrinsic growth rate of the population
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| * K = [[carrying capacity]] of its local environmental setting.
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| * β = coefficient converting encounters with one species to new units of the other
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| Mutualism is in essence the [[Logistic function|logistic growth equation]] + mutualistic interaction. The mutualistic interaction term represents the increase in population growth of species one as a result of the presence of greater numbers of species two, and vice versa.
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| In 1989, David Hamilton Wright modified the [[Lotka–Volterra equation|Lotka-Volterra equations]] by adding a new term, βM/K, to represent a mutualistic relationship.<ref name="Wright">Wright, David Hamilton. 1989. A Simple, Stable Model of Mutualism Incorporating Handling Time. [[The American Naturalist]], Vol. 134, No. 4, pp. 664-667.</ref> Wright also considered the concept of saturation, which means that with higher densities, there are decreasing benefits of further increases of the mutualist population. Without saturation, species' densities would increase indefinitely. Because that isn't possible due to environmental constraints and carrying capacity, a model that includes saturation would be more accurate. Wright's mathematical theory is based on the premise of a simple two-species mutualism model in which the benefits of mutualism become saturated due to limits posed by handling time. Wright defines handling time as the time needed to process a food item, from the initial interaction to the start of a search for new food items and assumes that processing of food and searching for food are mutually exclusive. Mutualists that display foraging behavior are exposed to the restrictions on handling time. Mutualism can be associated with symbiosis
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| ===Type II functional response===
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| In 1959, [[C. S. Holling]] performed his classic disc experiment that assumed the following: that (1), the number of food items captured is proportional to the allotted [[Search time|searching time]]; and (2), that there is a variable of [[handling time]] that exists separately from the notion of search time. He then developed an equation for the Type II [[functional response]], which showed that the feeding rate is equivalent to
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| :<math>
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| \cfrac{ax}{1+axT_H}
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| </math> | |
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| where,
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| * a = the instantaneous discovery rate
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| * x = food item density
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| * T<sub>H</sub> = handling time
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| The equation that incorporates Type II functional response and mutualism is:
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| :<math>
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| \cfrac{dN}{dt}=N[r(1-cN)+\beta M(X+M)]
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| </math>
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| where,
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| * N and M = density of the two mutualists
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| * r = intrinsic rate of increase of N
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| * c = coefficient measuring negative intraspecific interaction
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| * X = 1/a T<sub>H</sub>
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| * β = b/ T<sub>H</sub>
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| * a = instantaneous discovery rate
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| * b = coefficient converting encounters with M to new units of N
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| Rearranged:
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| :<math>
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| \cfrac{dN}{dt}=N\bigg[r(1-cN)+\cfrac{baM}{1+aT_H M}\bigg]
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| </math>
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| The model presented above is most effectively applied to free-living species that encounter a number of individuals of the mutualist part in the course of their existences. Of note, as Wright points out, is that models of biological mutualism tend to be similar qualitatively, in that the featured [[isocline]]s generally have a positive decreasing slope, and by and large similar isocline diagrams. Mutualistic interactions are best visualized as positively sloped isoclines, which can be explained by the fact that the saturation of benefits accorded to mutualism or restrictions posed by outside factors contribute to a decreasing slope.
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| ==See also==
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| * [[Arbuscular mycorrhiza]]
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| * [[Co-adaptation]]
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| * [[Co-evolution]]
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| * [[Ecological facilitation]]
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| * [[Frugivory]]
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| * [[Greater Honeyguide]] - has an interesting mutualism with humans
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| * [[Interspecific communication]]
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| * [[List of symbiotic relationships]]
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| * [[Müllerian mimicry]]
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| * [[Mutualisms and Conservation]]
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| * ''[[Mutual Aid: A Factor of Evolution]]''
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| * [[Symbiogenesis]]
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| ==References==
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| {{Reflist|2}}
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| ==Further references==
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| {{Refbegin|2}}
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| * Breton, Lorraine M., and John F. Addicott. 1992. Density-Dependent Mutualism in an Aphid-Ant Interaction. [[Ecology (journal)|Ecology]], Vol. 73, No. 6, pp. 2175–2180.
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| * Bronstein, JL. 1994. Our current understanding of mutualism. [[Quarterly Review of Biology]] 69 (1): 31-51 March 1994
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| * Bronstein JL. 2001. The exploitation of mutualisms. [[Ecology Letters]] 4 (3): 277-287
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| * Bronstein JL. 2001. The costs of mutualism. [[American Zoologist]] 41 (4): 825-839 S
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| * Bronstein JL, Alarcon R, Geber M. 2006. The evolution of plant-insect mutualisms. [[New Phytologist]] 172 (3): 412-428
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| * Denison RF, Kiers ET. 2004. Why are most rhizobia beneficial to their plant hosts, rather than parasitic? [[Microbes and Infection]] 6 (13): 1235-1239 {{ISSN|1286-4579}}
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| * DeVries, PJ; and Baker, I. 1989. Butterfly exploitation of an ant-plant mutualism: Adding insult of herbivory. Journal of the [[New York Entomological Society]] [J. N.Y. ENTOMOL. SOC.]. Vol. 97, no. 3, pp. 332–340. {{ISSN|0028-7199}}
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| * Hoeksema, J.D. & E.M. Bruna. 2000. Pursuing the big questions about interspecific mutualism: a review of theoretical approaches. [[Oecologia]] 125:321-330 {{ISSN|0029-8549}}
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| * Jahn, G.C. and J.W. Beardsley. 2000. Interactions of ants (Hymenoptera: Formicidae) and mealybugs (Homoptera: Pseudococcidae) on pineapple. Proceedings of the [[Hawaiian Entomological Society]] 34: 181-185. {{ISSN|0073-134X}}
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| * Jahn, Gary C., J. W. Beardsley and H. González-Hernández 2003. [https://scholarspace.manoa.hawaii.edu/bitstream/10125/95/1/36_9-28.pdf A review of the association of ants with mealybug wilt disease of pineapple.] Proceedings of the Hawaiian Entomological Society. 36:9-28.{{ISSN|0073-134X}}
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| * Noe, R. & P. Hammerstein. 1994. Biological markets: supply and demand determine the effect of partner choice in cooperation, mutualism and mating. [[Behavioral Ecology and Sociobiology]] 35:1-11 {{ISSN|0340-5443}}
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| * Ollerton, J. 2006. "Biological Barter": Patterns of Specialization Compared across Different Mutualisms. pp. 411–435 in: Waser, N.M. & Ollerton, J. (Eds) Plant-Pollinator Interactions: From Specialization to Generalization. University of Chicago Press. ISBN 978-0-226-87400-5
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| * Paszkowski, U. 2006. Mutualism and parasitism: the yin and yang of plant symbioses. [[Current Opinion in Plant Biology]] 9 (4): 364-370. {{doi|10.1016/j.pbi.2006.05.008}}. PMID 16713732
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| * Porat, D. & Chadwick-Furman, N. E. 2004. Effects of anemonefish on giant sea anemones:expansion behavior, growth, and survival. [[Hydrobiologia]] 530, 513–520. {{doi|10.1007/s10750-004-2688-y}}
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| * Porat, D. & Chadwick-Furman, N. E. 2005. Effects of anemonefish on giant sea anemones: ammonium uptake, zooxanthella content and tissue regeneration. Mar. Freshw. Behav. Phys. 38, 43–51. {{doi|10.1080/102362405000_57929}}
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| * Thompson, J. N. 2005. ''The Geographic Mosaic of Coevolution''. University of Chicago Press. ISBN 978-0-226-79762-5
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| * Wright, David Hamilton. 1989. A Simple, Stable Model of Mutualism Incorporating Handling Time. [[The American Naturalist]], Vol. 134, No. 4, pp. 664–667.
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| {{Refend}}
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| ==Further reading==
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| {{Commons category|Mutualism}}
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| {{Wiktionary}}
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| * Boucher, D. G., James, S. & Kresler, K. (1984) The ecology of mutualism. ''[[Annual Review of Ecology and Systematics]]'', '''13''': 315-347.
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| * Boucher, D. H. (editor) (1985) ''The Biology of Mutualism : Ecology and Evolution'' London : [[Croom Helm]] 388 p. ISBN 0-7099-3238-3
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| {{Biological interaction-footer}}
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| {{evo ecol}}
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| {{collective animal behaviour}}
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| {{modelling ecosystems|expanded=none}}
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| {{Use dmy dates|date=September 2010}}
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| {{DEFAULTSORT:Mutualism (Biology)}}
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| [[Category:Mutualism (biology)|*]]
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| [[Category:Biological interactions]]
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| [[Category:Symbiosis]]
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