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=== Freshwater Fish ===
=== Freshwater Fish ===
Cleaning has infrequently been observed in fresh waters compared to marine waters. This is possibly related to fewer observers (such as divers) in the former habitat compared to the latter<ref>{{Cite book|title="Natural history of Amazon fishes". In Encyclopedia of Life Support Systems (ed.). Tropical Biology and Natural Resources Theme. 1.|last=Carvalho|first=L.N|publisher=Eolss Publishers|year=2007|isbn=|location=Oxford|pages=1-24}}</ref>. One of the few known examples of cleaning is juvenile [[striped Raphael catfish]] cleaning the piscivorous [[Hoplias malabaricus|''Hoplias'' cf. ''malabaricus'']]. In [[Public aquarium|public aquariums]], ''[[Synaptolaemus]]'' headstanders have been seen cleaning larger fish<ref>{{Cite web|url=https://denblaaplanet.dk/en/species/broadband-red-headstander/|title=Broadband red headstander|last=Planet|first=Den Blå|last2=Fortlingsvej 1|first2=Address: Jacob|date=2016-10-03|website=Den Blå Planet|language=en|access-date=2019-10-27|last3=Kastrup|first3=Address: 2770|last4=info@denblaaplanet.dk|first4=Send email:|last5=Phone: +45 44 22 22 44}}</ref>.
Cleaning has infrequently been observed in fresh waters compared to marine waters. This is possibly related to fewer observers (such as divers) in the former habitat compared to the latter<ref>{{Cite book|title="Natural history of Amazon fishes". In Encyclopedia of Life Support Systems (ed.). Tropical Biology and Natural Resources Theme. 1.|last=Carvalho|first=L.N|publisher=Eolss Publishers|year=2007|isbn=|location=Oxford|pages=1-24}}</ref>. One of the few known examples of cleaning is juvenile [[striped Raphael catfish]] cleaning the piscivorous [[Hoplias malabaricus|''Hoplias'' cf. ''malabaricus'']]. In [[Public aquarium|public aquariums]], ''[[Synaptolaemus]]'' headstanders have been seen cleaning larger fish<ref>{{Cite web|url=https://denblaaplanet.dk/en/species/broadband-red-headstander/|title=Broadband red headstander|last=Planet|first=Den Blå|last2=Fortlingsvej 1|first2=Address: Jacob|date=2016-10-03|website=Den Blå Planet|language=en|access-date=2019-10-27|last3=Kastrup|first3=Address: 2770|last4=info@denblaaplanet.dk|first4=Send email:|last5=Phone: +45 44 22 22 44}}</ref><ref>{{cite web | title=Broadband red headstander | url=https://denblaaplanet.dk/en/species/broadband-red-headstander/ | publisher=[[National Aquarium Denmark]] | accessdate=18 August 2018 | date=2016-10-03 }}</ref>


== Mechanisms ==
<ref>{{cite web | title=Broadband red headstander | url=https://denblaaplanet.dk/en/species/broadband-red-headstander/ | publisher=[[National Aquarium Denmark]] | accessdate=18 August 2018 | date=2016-10-03 }}</ref>

=== Facultative Cleaner Fish ===
A facultative cleaner fish does not rely solely on specialized cleaning behaviour for nutrient consumption<ref name=":12">{{Cite journal|last=Morado|first=Nadia|last2=Mota|first2=Paulo G.|last3=Soares|first3=Marta C.|date=2019|title=The Rock Cook Wrasse Centrolabrus exoletus Aims to Clean|url=https://www.frontiersin.org/articles/10.3389/fevo.2019.00182/full|journal=Frontiers in Ecology and Evolution|language=English|volume=7|doi=10.3389/fevo.2019.00182|issn=2296-701X}}</ref>. Facultative cleaners can be further divided by stationary vs. wandering facultative cleaners<ref name=":02">{{Cite journal|last=Dunkley|first=Katie|last2=Cable|first2=Jo|last3=Perkins|first3=Sarah E.|date=2018-02-01|title=The selective cleaning behaviour of juvenile blue-headed wrasse (Thalassoma bifasciatum) in the Caribbean|url=http://www.sciencedirect.com/science/article/pii/S037663571730462X|journal=Behavioural Processes|volume=147|pages=5–12|doi=10.1016/j.beproc.2017.12.005|issn=0376-6357}}</ref> .Facultative cleaners may display cleaning behaviour through their whole life history or solely during juvenile stages for additional nutrients during rapid growth<ref name=":02" /><ref name=":12" />. Examples of facultative cleaners are commonly wrasse species such as the [[Thalassoma amblycephalum|blue headed wrasse]], noronha wrasse [[Noronha wrasse|(''Thalassoma noronhanum'')]] and goldsinny wrasse ([[Goldsinny wrasse|''Ctenolabrus rupestris'']]), sharp nose [[sea perch]] in Californian waters,<ref name=":12" /> and the lumpfish (''[[Cyclopterus lumpus]]'').

Using the example of the blue wrasse from Caribbean waters, their alternative feeding strategy is described as being a generalist [[Foragers|forager]], meaning they eat a wide variety of smaller aquatic organisms based on availability<ref name=":02" />. When displaying cleaning behaviour, it has been noted that the blue wrasse inspects potential clients and only feeds on some, implying that the wrasse is seeking out a particular type of parasite as diet supplementation. It has also been quantified that the blue wrasse foraging behaviour does not change proportionate to cleaning opportunities, again promoting the idea that the cleaning behaviour in this facultative fish is for diet supplementation and not out of necessity<ref name=":02" />.

=== Obligate Cleaner Fish ===
An obligate cleaner fish relies solely on specialized cleaning behaviour for nutrient consumption<ref name=":12" />. Therefore, obligate cleaners have a higher output of cleaning on a wider range of parasites in comparison to facultative fish. To maximize nutrient consumption, obligate cleaners utilize a higher proportion of cleaning stations<ref name=":02" />. Obligate cleaner fish may also be divided by stationary and wandering and these life history choice are made based on the amount of interspecific competition from other obligate cleaners in the area <ref>{{Cite web|url=https://login.qe2a-proxy.mun.ca/login?qurl=https://link.springer.com%2farticle%2f10.1007%2fs00338-012-0933-9|title=Memorial University Libraries - Proxy Login|website=login.qe2a-proxy.mun.ca|doi=10.1007/s00338-012-0933-9|access-date=2019-11-13}}</ref> An example of an obligate cleaner is the shark nose goby (''[[Elacatinus evelynae]]'') in the Caribbean Reef, where it has been observed to perform up to 110 cleanings per day.<ref name=":02" />
[[File:Mulloidichthys_flavolineatus_at_cleaning_station.jpg|link=https://en.wikipedia.org/wiki/File:Mulloidichthys_flavolineatus_at_cleaning_station.jpg|thumb|Client ''Mulloidichthys flavolineatus'' at a cleaning station.]]

=== Cleaner Stations ===
[[Cleaning stations]] are a strategy used by some cleaner fish where clients congregate and perform specific movements to attract the attention of the cleaner fish. Cleaning stations are usually associated with unique topological features, such as those seen in the coral reef<ref name=":02" /> and allow a space where cleaners have no risk of predation from larger predatory fishes, due to the mutual benefit from the cleaners service <ref name=":43">{{Cite book|url=https://www.worldcat.org/oclc/36051279|title=The diversity of fishes|last=Helfman, Gene S.|date=1997|publisher=Blackwell Science|others=Collette, Bruce B., Facey, Douglas E.|isbn=0865422567|location=Malden, Mass.|oclc=36051279}}</ref>.

Interactions are initiated by the client and terminated by the cleaner, implying that the client is seeking out the service where the cleaner has control<ref name=":12" />.

=== Cheating ===
Cheating parasitism occurs when the cleaner eats mucus or healthy tissue from the client. This can be harmful to the client as mucus is essential for preventing UV damage, and open wounds can increase risk of infection <ref name=":12" />. Cleaner fish maintain a balance between consuming ectoparasites and mucus or tissue because of the respective nutritional benefits, sometimes despite the risk to the client <ref>{{Cite journal|last=Gingins|first=Simon|last2=Werminghausen|first2=Johanna|last3=Johnstone|first3=Rufus A.|last4=Grutter|first4=Alexandra S.|last5=Bshary|first5=Redouan|date=2013-06-22|title=Power and temptation cause shifts between exploitation and cooperation in a cleaner wrasse mutualism|url=https://royalsocietypublishing.org/doi/10.1098/rspb.2013.0553|journal=Proceedings of the Royal Society B: Biological Sciences|language=en|volume=280|issue=1761|pages=20130553|doi=10.1098/rspb.2013.0553|issn=0962-8452|pmc=PMC3652443|pmid=23615288}}</ref>. For example, the[[Caribbean cleaning goby]] (''Elacatinus evelynae'') will eat scales and mucus from the host during times of ectoparasite scarcity to supplement its diet. The [[symbiosis]] relationship between client and host does not break down because the abundance of these parasites varies significantly seasonally and spatially, and the overall benefit to the larger fish outweighs any cheating on the part of the smaller cleaner<ref>{{Cite journal|last=Cheney|first=Karen L|last2=Côté|first2=Isabelle M|date=2005-05-16|title=Mutualism or parasitism? The variable outcome of cleaning symbioses|url=http://dx.doi.org/10.1098/rsbl.2004.0288|journal=Biology Letters|volume=1|issue=2|pages=162–165|doi=10.1098/rsbl.2004.0288|issn=1744-9561}}</ref>.

=== Memory ===
Cleaner fish (especially facultative cleaners) asses value of possible clients when deciding whether to invest in a client or cheat and consume mucus or tissue <ref name=":02" /><ref name=":22">{{Cite web|url=https://www.frontiersin.org/articles/10.3389/fnbeh.2017.00191/full|title=The Neurobiology of Mutualistic Behavior: The Cleanerfish Swims into the Spotlight|last=|first=|date=|website=www.frontiersin.org|language=en|url-status=live|archive-url=|archive-date=|access-date=2019-10-27}}</ref>. Observations of cleaner and client interactions have noted that cleaners may provide the client with tactile stimulation as a way to establish a relationship and gain the clients 'trust'. This interaction is at cost to the cleaner as it is time that it is not feeding<ref name=":22" />. This physical interaction demonstrates a cleaner fish [[Trade-off|tradeoff]]. The cleaner minimizes feeding time to establish a memorable relationship with the client that also contributes to conflict management with a possible predatory client<ref name=":22" />


==Mimicry==
==Mimicry==

Revision as of 16:51, 13 November 2019

Two bluestreak cleaner wrasses removing dead skin and external parasites from the potato grouper
Video of bluestreak cleaner wrasse cleaning the gills of an elongate surgeonfish

Cleaner fish are fish that show a specialist feeding strategy[1] by providing a service to other species, referred to as clients[2], by removing dead skin, ectoparasites and infected tissue from the surface or gill chambers[2]. This example of cleaning symbiosis represents mutualism and cooperation behaviour[3], an ecological interaction that benefits both parties involved. However, the cleaner fish may consume mucus or tissue, thus creating a form of parasitism[4] called cheating. The client animal is typically a heterospecific fish [3], But can also involve aquatic reptiles (sea turtles and marine iguana), mammals (manatees and whales) or octopuses.[5][6][7] A wide variety of fish including wrasse, cichlids, catfish, pipefish, lumpsuckers and gobies display cleaning behaviors across the globe in fresh, brackish, and marine waters but specifically concentrated in the tropics due to high parasite density[2]. Similar behavior is found in other groups of animals, such as cleaner shrimps.

There are two types of cleaner fish, obligate full time cleaners and facultative part time cleaners [1] where different strategies occur based on resources and local abundance of fish.[1] Cleaning behaviour takes place in pelagic waters as well as designated locations called cleaner stations[8]. Cleaner fish interaction durations and memories of reoccurring clients are influenced by the neuroendocrine system of the fish, involving hormones Arginine Vasotocin, Isotocin and serotonin[3].

Conspicuous coloration is a method used by some cleaner fish, where they often displaying a brilliant blue stripe that spans the length of the body.[9] Other species of fish, called mimics, imitate the behavior and phenotype of cleaner fish to gain access to client fish tissue.

The specialized feeding behaviour of cleaner fish has become a valuable resource in salmon aquaculture in Atlantic Canada, Scotland, Iceland and Norway[10] for prevention of sea lice out breaks[2] which is beneficial to the economy and environment by minimizing use of chemical delousers. Specifically cultured for this job are lumpfish (Cyclopterus lumpus) and ballan wrasse (Labrus bergeylta)[11]. The most common parasites that cleaner fish feed on are gnathiidae and copepod species[1].


Diversity and Examples

Marine Fishes

The following is a selection of few of the many marine cleaner species.

Commonly studied cleaner fish are the cleaner wrasses of the genus Labroides found on coral reefs in the Indian Ocean and Pacific Ocean [12].

Neon gobies of the genera Gobiosoma and Elacatinus provide a cleaning service similar to the cleaner wrasses, though this time on reefs in the Western Atlantic, providing a good example of convergent evolution[13] of the cleaning behaviour.

Lumpfish who are utilized as salmonid cleaner fish in aquaculture, but it is unknown if they display cleaning behaviour on salmon in the wild.[14]

Brackish Freshwater Fish

Brackish water refers to aquatic environments that have a salinity in between salt and fresh water systems. Cleaning symbiosis has also been observed in these areas between two brackish water cichlids of the genus Etroplus from South Asia. The small species Etroplus maculatus is the cleaner fish, and the much larger Etroplus suratensis is the host that receives the cleaning service[16].

Freshwater Fish

Cleaning has infrequently been observed in fresh waters compared to marine waters. This is possibly related to fewer observers (such as divers) in the former habitat compared to the latter[17]. One of the few known examples of cleaning is juvenile striped Raphael catfish cleaning the piscivorous Hoplias cf. malabaricus. In public aquariums, Synaptolaemus headstanders have been seen cleaning larger fish[18][19]

Mechanisms

Facultative Cleaner Fish

A facultative cleaner fish does not rely solely on specialized cleaning behaviour for nutrient consumption[20]. Facultative cleaners can be further divided by stationary vs. wandering facultative cleaners[21] .Facultative cleaners may display cleaning behaviour through their whole life history or solely during juvenile stages for additional nutrients during rapid growth[21][20]. Examples of facultative cleaners are commonly wrasse species such as the blue headed wrasse, noronha wrasse (Thalassoma noronhanum) and goldsinny wrasse (Ctenolabrus rupestris), sharp nose sea perch in Californian waters,[20] and the lumpfish (Cyclopterus lumpus).

Using the example of the blue wrasse from Caribbean waters, their alternative feeding strategy is described as being a generalist forager, meaning they eat a wide variety of smaller aquatic organisms based on availability[21]. When displaying cleaning behaviour, it has been noted that the blue wrasse inspects potential clients and only feeds on some, implying that the wrasse is seeking out a particular type of parasite as diet supplementation. It has also been quantified that the blue wrasse foraging behaviour does not change proportionate to cleaning opportunities, again promoting the idea that the cleaning behaviour in this facultative fish is for diet supplementation and not out of necessity[21].

Obligate Cleaner Fish

An obligate cleaner fish relies solely on specialized cleaning behaviour for nutrient consumption[20]. Therefore, obligate cleaners have a higher output of cleaning on a wider range of parasites in comparison to facultative fish. To maximize nutrient consumption, obligate cleaners utilize a higher proportion of cleaning stations[21]. Obligate cleaner fish may also be divided by stationary and wandering and these life history choice are made based on the amount of interspecific competition from other obligate cleaners in the area [22] An example of an obligate cleaner is the shark nose goby (Elacatinus evelynae) in the Caribbean Reef, where it has been observed to perform up to 110 cleanings per day.[21]

Client Mulloidichthys flavolineatus at a cleaning station.

Cleaner Stations

Cleaning stations are a strategy used by some cleaner fish where clients congregate and perform specific movements to attract the attention of the cleaner fish. Cleaning stations are usually associated with unique topological features, such as those seen in the coral reef[21] and allow a space where cleaners have no risk of predation from larger predatory fishes, due to the mutual benefit from the cleaners service [23].

Interactions are initiated by the client and terminated by the cleaner, implying that the client is seeking out the service where the cleaner has control[20].

Cheating

Cheating parasitism occurs when the cleaner eats mucus or healthy tissue from the client. This can be harmful to the client as mucus is essential for preventing UV damage, and open wounds can increase risk of infection [20]. Cleaner fish maintain a balance between consuming ectoparasites and mucus or tissue because of the respective nutritional benefits, sometimes despite the risk to the client [24]. For example, theCaribbean cleaning goby (Elacatinus evelynae) will eat scales and mucus from the host during times of ectoparasite scarcity to supplement its diet. The symbiosis relationship between client and host does not break down because the abundance of these parasites varies significantly seasonally and spatially, and the overall benefit to the larger fish outweighs any cheating on the part of the smaller cleaner[25].

Memory

Cleaner fish (especially facultative cleaners) asses value of possible clients when deciding whether to invest in a client or cheat and consume mucus or tissue [21][26]. Observations of cleaner and client interactions have noted that cleaners may provide the client with tactile stimulation as a way to establish a relationship and gain the clients 'trust'. This interaction is at cost to the cleaner as it is time that it is not feeding[26]. This physical interaction demonstrates a cleaner fish tradeoff. The cleaner minimizes feeding time to establish a memorable relationship with the client that also contributes to conflict management with a possible predatory client[26]

Mimicry

Further information: Aggressive mimicry and Cleaning symbiosis
The bluestriped fangblenny is an aggressive mimic of the cleaner wrasse.

Mimic species have evolved body forms, patterns, and colors which imitate other species to gain a competitive advantage.[27] One of the most studied examples of mimicry on coral reefs is the relationship between the aggressive mimic Plagiotremus rhinorhynchos (the bluestriped fangblenny) and the cleaner wrasse model Labroides dimidiatus. By appearing like L. dimidiatus, P. rhinorhynchos is able to approach and subsequently feed on the tissue and scales of client fish while posing as a cleaner.[27][28]

The presence of the cleaner mimic, P. rhinorhynchos, has a negative impact on the foraging success of the cleaner model L. dimidiatus.[28] P. rhinorhynchos feeds by eating the tissue and scales of client fish, making client fish much more cautious while at cleaning stations. More aggressive mimics have a greater negative impact on the foraging rate and success of the cleaner fish.[28] When mimics appear in higher densities relative to cleaners, there is a significant decline in the success rate of the cleaner fish. The effects of the mimic/model ratio are susceptible to dilution, whereby an increase in client fish allows both the mimics and the models to have more access to clients, thus limiting the negative effects that mimics have on model foraging success.[29][30]

Similar species also include Plagiotremus tapeinosoma (the Mimic blenny),Aspidontus.

See also

References

  1. ^ a b c d Dunkley, Katie; Cable, Jo; Perkins, Sarah E. (2018-02-01). "The selective cleaning behaviour of juvenile blue-headed wrasse (Thalassoma bifasciatum) in the Caribbean". Behavioural Processes. 147: 5–12. doi:10.1016/j.beproc.2017.12.005. ISSN 0376-6357.
  2. ^ a b c d Morado, Nadia; Mota, Paulo G.; Soares, Marta C. (2019). "The Rock Cook Wrasse Centrolabrus exoletus Aims to Clean". Frontiers in Ecology and Evolution. 7. doi:10.3389/fevo.2019.00182. ISSN 2296-701X.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  3. ^ a b c "The Neurobiology of Mutualistic Behavior: The Cleanerfish Swims into the Spotlight". www.frontiersin.org. Retrieved 2019-10-27.{{cite web}}: CS1 maint: url-status (link)
  4. ^ Gingins, Simon; Werminghausen, Johanna; Johnstone, Rufus A.; Grutter, Alexandra S.; Bshary, Redouan (2013-06-22). "Power and temptation cause shifts between exploitation and cooperation in a cleaner wrasse mutualism". Proceedings of the Royal Society B: Biological Sciences. 280 (1761): 20130553. doi:10.1098/rspb.2013.0553. ISSN 0962-8452. PMC 3652443. PMID 23615288.{{cite journal}}: CS1 maint: PMC format (link)
  5. ^ Grutter, A. S. (2002). "Cleaning symbioses from the parasites' perspective". Parasitology. 124 (7): 65–81. doi:10.1017/S0031182002001488. ISSN 0031-1820. {{cite journal}}: no-break space character in |first= at position 3 (help)
  6. ^ Sazima, Cristina; Grossman, Alice; Sazima, Ivan (2010-02-05). "Turtle cleaners: reef fishes foraging on epibionts of sea turtles in the tropical Southwestern Atlantic, with a summary of this association type". Neotropical Ichthyology. 8 (1): 187–192. doi:10.1590/S1679-62252010005000003. ISSN 1982-0224.
  7. ^ "Manatee gets 'haircut' from gill fish". 2010-02-26. ISSN 0307-1235. Retrieved 2019-10-28.
  8. ^ Helfman, Gene S. (1997). The diversity of fishes. Collette, Bruce B., Facey, Douglas E. Malden, Mass.: Blackwell Science. ISBN 0865422567. OCLC 36051279.
  9. ^ Cheney, Karen L.; Grutter, Alexandra S.; Blomberg, Simon P.; Marshall, N. Justin (2009). "Blue and Yellow Signal Cleaning Behavior in Coral Reef Fishes". Current Biology. 19 (15): 1283–1287. doi:10.1016/j.cub.2009.06.028.
  10. ^ "Cleaner fish – what do they do?". Lochduart. 2017-06-08. Retrieved 2019-10-27.
  11. ^ Brooker, Adam J; Papadopoulou, Athina; Gutierrez, Carolina; Rey, Sonia; Davie, Andrew; Migaud, Herve (2018-09-29). "Sustainable production and use of cleaner fish for the biological control of sea lice: recent advances and current challenges". Veterinary Record. 183 (12): 383–383. doi:10.1136/vr.104966. ISSN 0042-4900.
  12. ^ Helfman, Gene S. (1997). The diversity of fishes. Collette, Bruce B., Facey, Douglas E. Malden, Mass.: Blackwell Science. ISBN 0865422567. OCLC 36051279.
  13. ^ Fenner, Robert M. (1998). The conscientious marine aquarist : a commonsense handbook for successful saltwater hobbyists. Shelburne, Vt.: Microcosm. ISBN 1890087033. OCLC 38168280.
  14. ^ Powell, Adam; Treasurer, Jim W.; Pooley, Craig L.; Keay, Alex J.; Lloyd, Richard; Imsland, Albert K.; Leaniz, Carlos Garcia de (2018). "Use of lumpfish for sea-lice control in salmon farming: challenges and opportunities". Reviews in Aquaculture. 10 (3): 683–702. doi:10.1111/raq.12194. ISSN 1753-5131.
  15. ^ "Cleaner fish – what do they do?". Lochduart. 2017-06-08. Retrieved 2019-10-28.
  16. ^ Wyman, Richard L.; Ward, Jack A. (1972-12-29). "A Cleaning Symbiosis between the Cichlid Fishes Etroplus maculatus and Etroplus suratensis. I. Description and Possible Evolution". Copeia. 1972 (4): 834. doi:10.2307/1442742. ISSN 0045-8511.
  17. ^ Carvalho, L.N (2007). "Natural history of Amazon fishes". In Encyclopedia of Life Support Systems (ed.). Tropical Biology and Natural Resources Theme. 1. Oxford: Eolss Publishers. pp. 1–24.
  18. ^ Planet, Den Blå; Fortlingsvej 1, Address: Jacob; Kastrup, Address: 2770; info@denblaaplanet.dk, Send email:; Phone: +45 44 22 22 44 (2016-10-03). "Broadband red headstander". Den Blå Planet. Retrieved 2019-10-27. {{cite web}}: |first4= has generic name (help)CS1 maint: extra punctuation (link) CS1 maint: numeric names: authors list (link)
  19. ^ "Broadband red headstander". National Aquarium Denmark. 2016-10-03. Retrieved 18 August 2018.
  20. ^ a b c d e f Morado, Nadia; Mota, Paulo G.; Soares, Marta C. (2019). "The Rock Cook Wrasse Centrolabrus exoletus Aims to Clean". Frontiers in Ecology and Evolution. 7. doi:10.3389/fevo.2019.00182. ISSN 2296-701X.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  21. ^ a b c d e f g h Dunkley, Katie; Cable, Jo; Perkins, Sarah E. (2018-02-01). "The selective cleaning behaviour of juvenile blue-headed wrasse (Thalassoma bifasciatum) in the Caribbean". Behavioural Processes. 147: 5–12. doi:10.1016/j.beproc.2017.12.005. ISSN 0376-6357.
  22. ^ "Memorial University Libraries - Proxy Login". login.qe2a-proxy.mun.ca. doi:10.1007/s00338-012-0933-9. Retrieved 2019-11-13.
  23. ^ Helfman, Gene S. (1997). The diversity of fishes. Collette, Bruce B., Facey, Douglas E. Malden, Mass.: Blackwell Science. ISBN 0865422567. OCLC 36051279.
  24. ^ Gingins, Simon; Werminghausen, Johanna; Johnstone, Rufus A.; Grutter, Alexandra S.; Bshary, Redouan (2013-06-22). "Power and temptation cause shifts between exploitation and cooperation in a cleaner wrasse mutualism". Proceedings of the Royal Society B: Biological Sciences. 280 (1761): 20130553. doi:10.1098/rspb.2013.0553. ISSN 0962-8452. PMC 3652443. PMID 23615288.{{cite journal}}: CS1 maint: PMC format (link)
  25. ^ Cheney, Karen L; Côté, Isabelle M (2005-05-16). "Mutualism or parasitism? The variable outcome of cleaning symbioses". Biology Letters. 1 (2): 162–165. doi:10.1098/rsbl.2004.0288. ISSN 1744-9561.
  26. ^ a b c "The Neurobiology of Mutualistic Behavior: The Cleanerfish Swims into the Spotlight". www.frontiersin.org. Retrieved 2019-10-27.{{cite web}}: CS1 maint: url-status (link)
  27. ^ a b Cheney, K.L. (2008). "Facultative mimicry: cues for colour change and colour accuracy in a coral reef fish", Proceedings of the Royal Society B, 275 (1631): 117–22.
  28. ^ a b c Cheney, K.L. (2012). "Cleaner wrasse mimics inflict higher costs on their models when they are more aggressive towards signal receivers"[permanent dead link‍] Biology Letters, doi:10.1098/rsbl.2011.0687
  29. ^ Cheney, K.L. (2005). "Frequency-dependent success of aggressive mimics in a cleaning symbiosis", Proceedings of the Royal Society B, 272 (1581): 2635–39.
  30. ^ Cheney, K.L. (2007). "Aggressive mimics profit from a model-signal receiver mutualism" Proceedings of the Royal Society B, 274: 1622: 2087–91. doi:10.1098/rspb.2007.0543

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