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Cleaner fish

From Wikipedia, the free encyclopedia

Two bluestreak cleaner wrasses removing dead skin and external parasites from a 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 animals are typically fish of a different species,[3] but can also be 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 behaviour 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 display 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 outbreaks[2] which benefits the economy and environment by minimizing the 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]

YouTube Encyclopedic

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  • Skinning the Wolf
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  • Cleaner Wrasse Cleaning Barracuda Teeth at Kapalai Pier - Fish Behavior

Transcription

You going to the pad? I'm going to... the... where I'm estimating to be the third tarsal there. 28? - 28. We got the weight. And I think that's it. Um, I don't... want to weigh the testes. Sweet. So, should we push it over that way and then... Or, let's see how much he lays out if we flip him up. Also, see, these are... his leg is so broken. It's just flopping around all over the place. - Oh my god. Is this one broken too? I think it is. I think its femur split. Yeah, it's not like that one. He is just... ugh. Poor puppy. Obviously this is not something you wanna do at home on your kitchen table. For one thing, wolves can be protected species, you need a permit to be in possession of them. Wolves can carry roundworm and ringworm, which can be transmittable to humans, so as long as don't attempt to ingest any feces from this wolf, we should be fine. Okay, we're gonna make the first incision. I'm looking for the back of his mandible. You know, the back of your jaw where it comes together, so I can cut between it. So right now, I'm trying to get through all this underfur and find out where the skin actually starts. It's very thick. So the first series of cuts are kind of like you're testing to see how thick the skin is. But I don't wanna cut so much that I cut through all the way and reach muscle tissue. Once you cut through muscle tissue, you start to get bleeding. And I don't need blood this early in the morning. Whoa. Hey. Hey. Where did this guy come from? - I know, it's been around. Fleas! Are those fleas? Sick! This guy had been in a freezer for two weeks. And he still has live fleas living on him. So there's the first cut. So now I'm just gonna, like, I made this little hole. I'm just trying to separate the hole and like see how far the epidermal layer goes and at what point I'm going to hit fat and muscle. That's kinda where I wanna be. See the muscle layer right there? And this is the epidermal layer. This is kind of frozen congealed blood in here. See it movin' around in the veins. So, because I have that vein attached to the skin. I'm cutting through the layers of fat, so I don't puncture it. That right here is fat, that is wolf fat right there. So his throat is right here and this is his, you know, breast muscle starting to come forward, where his arm has been broken at the humerus. Someone's texting me and I don't think I will be able to get back to them. This is just the base of the subcutaneous tissue. This is the connective tissue that connects the skin to the muscle and it has this like glossiness to it. There's fat attached to the skin here, so I cut a little close there. And this is ideally what I would be doing all of the time. So you can see the top of scapula here, this is probably where it should normally be. This is the top of the scapula on the other side. So if we draw a straight line across, it's been... You know, that's about four inches right there, where it's been dislocated. So this is cool. I cut accidentally between two different muscles right here, so the same kind of like weird transparent connective tissue, this stuff right here, that attaches... I think it's the same tissue, it's a very similar connective tissue that attaches the skin to the muscle also attaches the muscles to one another. It's surprisingly strong and it's very flexible and this is what holds your insides together. So this is where I'm starting to see signs of trauma or potential trauma from when he got hit by the car 'cause you can see all these burst blood vessels in here, um, whereas like, you know, this part of the skin, it's pretty pink and consistent all the way through. You start looking in here and you're seeing these, like, bright red veins. Also this dark coloration kinda looks like a bruise, what a bruise looks like underneath the skin. Which would make sense, I mean, 'cause this is, you know, where he got the brunt of the impact, I'm thinking, from the car. Can you see all of that trauma in there? All that dark, dark, red, it's almost purple or black colored. So it's a huge bruise underneath the skin. That's obviously where he sustained the huge impact. Don't get hit by a car, you guys. Look at this, do you see this, all this blood? In that vein right there? I don't wanna pop that, it's gonna be gross if I do. But you can see, like, another indication of trauma. All of the blood is pooled under there. It's like a big zit you just wanna pop. This is the, where the semilunar notch is, of the ulna. So this is backwards, where it's supposed to be pointing that way. It's twisted, because the humerus is broken under here. I think. That's what I'm deducing right now. You guys, I'm really sorry, we just identified the wolf. I think it's Jacob. I think it's Bella's Jacob. From Twilight. I'm really sorry. We'll go back through later and cut along the inside of the limbs. I just kinda wanna fillet him open first. That made me think of McDonald's. I'm kinda hungry. So this right here, I mean, all this looks like healthy good tissue and everything and we're getting down here and all of the sudden it's green. I'm wondering if when he was hit by a car the bowels ruptured and, so, we're having some, like, seepage. So I'm gonna be really careful about how I work around here. The last thing that I wanna do is puncture that and release whatever kind of gases or bacteria have turned this green. It's not a good sign that it keeps smelling worse the further we go down here. I don't like the idea that the bowels are all bloated and distended. We just had that conversation about how roundworm and ringworm and all these other kinds of diseases and parasites can live in the bowels, so I don't want to puncture them and release any of that. It it not a healthy color. It's not a happy color. It's not a good smell coming off of it and it's not just gases being released out of the anus. 'Cause you can smell them from over there. So the tissue itself has this odor to it, which makes me think that there's bacteria, which is active and alive. I'm getting to the penis right now. I can feel the baculum right here. All dogs have a baculum which is literally the penis bone of an animal, humans don't have it anymore. Some primates do, but it is very small. So I'm gonna cut to one side of the baculum as I go down this way. Sometimes in the collection we'll also come across broken baculum. They are bones, they can break and fracture and splinter just like any other bone in your body. Think about that one, guys. I never know quite how to move around the baculum. I mean that in all seriousness, you guys. Cuz like, do you remove it, like how you do the, the other limbs? I mean, 'cause it's got, it's like that tube shape, so you have the baculum, you have the penis and you have the skin around it; do you flay it on the inside of the baculum and open it like that? Oh god. That reeks. Ugh. Let's try to describe that smell. What does it smell like? Smells like KFC if you leave it in the back of your car with a layer of water at the bottom of the bucket in hundred degree weather for eight years. I am continuously amazed by the things that nature produces. You know, the fact that some of these animals can sustain such significant injuries during their life and they don't go inside at night, they don't have health care, they don't have a doctor, like, I seriously, I feel like such a wimp, I'll stub my toe or something and I'm like: "I'm done!" "Today is over! I'm out!" - I'm sitting down for the rest of the day. That's a wrap for the day. - Yeah. And you'll see these animals and they'll have these huge compound fractures and giant radial like twists and turns and the bone is regrown together and they lose like half of the limb the forelimb, and especially like on an ungulate like a deer or an elk, they're having to walk on those things continuously for years, it takes them years to regrow that and use that limb again. So in that time they're still hunting, they're still foraging, they're probably not in a good place to be reproducing but they are evading predators And I just, I mean, that kind of thing is just so... impressive to me. It like motivates me to be a better human being. I'm not even kidding, like, if these animals can do it, - I better step up my game. I better just suck it up. This is unhealthy. This buoyancy. That's not what you want. You want things to like deflate once they die. You don't want them to inflate, 'cause that, yeah, that just implies all kinds of gases... Wow. Oh. Look at that. I just punctured the, the vein there. - Oh yeah. We got a bleeder! This is, um, the femur right here which has been broken, we have the patella, which is the knee-bone. This is the top of the tibia and then behind it is the joint bone, the fibula. You can see all the tendons running along the bottom of the foot and they all connect back here. You kinda see the system there, where he's relaxed and it pulls tight if you're pulling it like that. And I can feel his foot, like, here, his toes are relaxed and this pad is down against my palm and when I do this, I can feel these other fingers, like his... what essentially are his finger-bones, like flexing into my hand. So it's like taut and relaxed and taut and relaxed. Pretty cool, guys! High five. I just put my face right in his asshole. Something still smells bad to me. Yes, oh, here we go. There we go. If I was writing a Disney movie. This is when the female character would break into a song. I don't think Disney would make a movie about... The young, beautiful taxidermist? (singing) When I grow up (singing) I just wanna be a taxidermist. It's like a princess who doesn't want to be a princess, she wants to be a taxidermist. Whoa! - Bam! There we go, everybody. That's what the paw looks like. Sans leg bones. It's a little bit floppy. This is what goes inside of there and these still have like, the claws are attached to another third phalange, phalanges, and then they attach with this, this is the paw, this is where the pad is, right there, So you can kinda see it as it would move around. compared to that leg. You can see where the femur is like... sticking out here. It's broken. It's been broken right there. So it's just interesting to kinda... You know, play with it, and see, like, how the muscles flex in here and how they move and rotate and work together. If you could get macro shots of this, this is like a gorgeous biological landscape. I'm serious! I'm gonna be cutting at you. Here I come! My blade. Don't get your finger in the way. You're gonna make a joke about how I'm gonna cut you and it's not gonna be funny, 'cause I'm gonna freak out. - Ow! Hey! I hate that! Arrgh, you're such a jerk! I mean, you flick this and it's just like taut and this is like squishy. It smells every time you do that. What would you, how would you, I can't! We have the baculum in here and the baculum is the penis bone. We have the penis, you can see a little bit of, like, blood and urine which has come out of the tip of the penis right here. It's still in the fur. So now we're gonna remove the penis. First I have to find it. And I can feel the baculum in there, you can kind of see it. Do you wanna touch it? Are you sure? Okay. How should we remove this, curator? - Well that's new to me. You don't remove penis bones very often? Sick. I just cut through this right here? Ahh, there we go. So this is like, the sheath. I like how these guys aren't saying anything, or doing anything. They're just watching me do this. So if you can imagine an uncircumcised penis, here is how it covers the end, here's the head of the penis, this is the urethra, and it's flaccid and there's a baculum, which is a hard bone in it. This is where it attaches, this is where the testes would be. So this is, this is an entire, I mean, you can even, it looks- I mean it looks like a penis, you guys. Like a human penis. From what I've seen in textbooks. Whew, that anus! - You get a front-row seat. I do! Oh! Ugh! For the show. - Ughhh! Oh my god, that's bad. Whoo! Oh my god. I have to.. I can't sit there. Ready? One, two, three. We wanna invert all of this skin over his head like you're pulling a turtle neck from over your head. We're cutting really close to the skull, now. We're gonna get close to eyes up there. This is all cartilage right here. This is the part of the skull where my finger is right now, I'm putting it into the skull, it's what you call the external auditory meatus and that is where the cartilage of the ear connects with the skull and that's where you get the ear canal. And you wanna cut around it, so you can keep all of the ear intact with the hide. These are the masseter muscles, the big cheek muscles on the side of the face. And this right here is the eye. You can.. Or where the eyelid would be, on the inside. Right there. Um, so... You have the eyelid and the eyelashes and right here is the zygomatic and squamosal bones which make up like the occular orbital. So you wanna cut really close, so you can keep the eyelashes intact, and so then you can when you get a little hole started you can kinda poke it through and I can tell that I have the eyelids right there, that are intact. I'm gonna try and free part of the mouth. So you can start to feel, like, I can feel the molars back there, I can feel the bottom of the mandible. Cutting along the teeth on the outside of the lips. Right now I'm kind of cutting into the bone a little bit, 'cause I'd rather get a couple marks in the bone than I would screwing up the side of the mouth. Everything is- there is like no muscle in here. It's all very thin. And it's all very closely connected to the bone. And this is all the inside of his lip right here. So his lip is right here and this dark color is his gums. They're naturally dark and also because he's been bleeding on it. Okay, so we flipped him over, he's bleeding all over the table at this point. We're really close to the end, you guys. And you can feel the nasal cavity here and then the, you have cartilaginous snout, the nose in the front, just cut through that make sure to keep the lips intact, and then this is a tricky one, you wanna get the bottom lip, too, and then we're free! We did it! We did it! This is the first time I ever skinned a wolf in my life and it might be the last time. And the pelt isn't gorgeous on the inside, but the outside of it doesn't look half-bad. And then you gotta revert the head and we got, - We got the ears we got the ears Did we get the eyes? Eyes look good. - Eyes look good. Got the eyelids. You get the eyelashes. We got the nose. It still has the, a little bit of cartilage in there, too. and it retains the good shape of the snout. There you have it. Thanks for watching, guys.

Diversity and examples

Marine fish

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

Commonly studied cleaner fish are the cleaner wrasse of the genus Labroides found on coral reefs in the Indian Ocean and Pacific Ocean.[8]

Neon gobies of the genera Gobiosoma and Elacatinus provide a cleaning service similar to the cleaner wrasse, though this time on reefs in the Western Atlantic, providing a good example of convergent evolution[12] of the cleaning behaviour.

Lumpfish are utilized as salmonid cleaner fish in aquaculture, but it is unknown if they display cleaning behaviour on salmon in the wild.[13]

Brackish water 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.[15]

Freshwater fish

Cleaning has been observed infrequently in fresh waters compared to marine waters. This is possibly related to fewer observers (such as divers) in freshwater compared to saltwater.[16] One of the few known examples of freshwater cleaning is juvenile striped Raphael catfish cleaning the piscivorous Hoplias cf. malabaricus. In public aquariums, Synaptolaemus headstanders have been seen cleaning larger fish.[17][18]

Mechanisms

Facultative cleaner fish

A facultative cleaner fish does not rely solely on specialized cleaning behaviour for nutritious food.[2] Facultative cleaners can be further divided by stationary vs. wandering facultative cleaners.[1] Facultative cleaners may display cleaning behaviour through their whole life history or solely as juveniles for additional nutrients during rapid growth.[1][2] 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,[2] 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.[1] 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 a diet supplement. It has also been quantified that the blue wrasse foraging behaviour does not change in proportion to cleaning opportunities, again suggesting that the cleaning behaviour in this facultative fish is for diet supplementation and not out of necessity.[1]

Obligate cleaner fish

An obligate cleaner fish relies solely on specialized cleaning behaviour for its food.[2] 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.[1] Obligate cleaner fish may also be divided by stationary and wandering. These life history choice are made based on the amount of interspecific competition from other obligate cleaners in the area.[19] 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.[1]

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 coral reefs[1] and allow a space where cleaners have no risk of predation from larger predatory fishes, due to the mutual benefit from the cleaners' service.[8]

Interactions are begun by the client and ended by the cleaner, implying that the client is seeking out the service where the cleaner has control.[2]

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 to prevent UV damage, and open wounds can increase the risk of infection.[2] Cleaner fish maintain a balance between eating ectoparasites and mucus or tissue because of the respective nutritional benefits, sometimes despite the risk to the client.[4] 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 by the smaller cleaner.[20]

Memory

Cleaner fish (especially facultative cleaners) assess the value of possible clients when deciding whether to invest in a client or cheat and eat mucus or tissue.[1][3] Observations of cleaner and client interactions have found that cleaners may provide the client with tactile stimulation as a way to establish a relationship and gain the client's 'trust'. This interaction costs the cleaner as it is time not spent feeding.[3] This physical interaction demonstrates a cleaner fish's tradeoff. The cleaner minimizes feeding time to establish a memorable relationship with the client that also contributes to conflict management with a possibly predatory client.[3]

Neurobiology

Protein structure of non-mammalian specific hormone, vasotocin, from the posterior pituitary.

The cleaner fish neuroendocrine system has been studied specifically in reference to arginine vasotocin (AVT) and Isotocin. These are fish-specific hormones that are analogous to human hormones involved in sociality.[3] In laboratory experiments, during conditions of low AVT, cleaners are more engaged in interspecific interactions. High AVT conditions tend to show high client interactions but more instances of cheating. This implies that AVT expression acts as a switch for cleaner fish feeding behaviour, showing less client interactions (but more honest cleaning) or increased client interactions (with less honest cleaning).[3] It has also been observed that obligate cleaners have higher overall brain activity, and specifically in the cerebellum, likely related to the movements involved in cleaning.[3]

Serotonin has also been noted to influence cleaning behaviour. High serotonin increases motivation to interact with clients, and a lack of serotonin decreases client interaction and slows learning.[3]

Mimicry

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.[21] 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 then feed on the tissue and scales of client fish while posing as a cleaner.[21][22]

The presence of the cleaner mimic, P. rhinorhynchos, reduces the foraging success of the cleaner model L. dimidiatus.[22] 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.[22] 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.[23][24]

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

Implications

Salmonid aquaculture

An example of an outdoor aquaculture facility in Chile.

Aquaculture is the farming of aquatic organisms, where salmon farming is growing in the North Atlantic.[10] Cleaner fish are used to eat parasitic sea lice from salmon to reduce outbreaks which cause disease in populations. The two most commonly used cleaner fish are the lumpfish, Cyclopterus lumpus, and the ballan wrasse Labrus bergeylta.[11] Lumpfish are distributed across the Atlantic ocean, ranging from Greenland to France, Hudson's Bay to New Jersey, and in high concentrations in the Bay of Fundy and St. Pierre Coast, near Newfoundland.[25] Ballan wrasse are distributed widely across the Northeast Atlantic Ocean.[26] The switch towards lumpfish has been preferred as wrasse are less active feeders during winter months.[13]

Methods

Cleaner fish are commercially cultured and introduced into salmonid sea cages. Salmon and lumpfish are able to coexist, where the lumpfish spend a certain amount of time foraging for supplemented food and only a portion of their time delousing salmon. With significant ratios of cleaner to client, the efforts are sufficient to minimize louse outbreaks.[13][11] Sea cages are designed with additional substrate for lumpfish to attach to during periods of inactivity to minimize stress levels in the cleaner fish and maximize delousing abilities.[13]

Challenges of using cleaner fish

Sea lice causing morphological damage on cultured salmon in New Brunswick, Canada.

North Atlantic Aquaculture facilities use facultative cleaner fish (Cyclopterus lumpus, and Labrus bergeylta) in order to control the nutrients they receive during culturing, before their use in aquaculture. One of the challenges that comes along with using facultative cleaners is that parasite removal from salmon must be maximized while also balancing additional nutrients from supplemented feed to ensure the health of the cleaner fish and the safety of the salmonid clients.[1] Another challenge that arises in management of cleaner fish behaviour is balancing the number of cleaners to the number of clients. With a low cleaner-to-client ratio, the risk of lice infestation increases. With a high cleaner-to-client ratio, competition among cleaners increases and there is a higher risk of cheating and consumption of salmonid mucus and flesh thereby increasing their risk of infection.[1][11]

Minimizing disease in commercial lumpfish stocks is critical for the continuation of their usage in aquaculture. Vaccine development for the lumpfish is a current area of research as lumpfish demand is increasing in the aquaculture industry.[13] In an effort to minimize disease in the cleaner fish, commercial lumpfish stocks are supplemented with wild individuals during the breeding season to minimize inbreeding depression. The lumpfish genome has not yet been fully sequenced so subtle details between populations are not yet appreciated.[13]

Another consideration in using cleaner fish in aquaculture is minimizing escapees from sea cages. If escaped cleaner fish spawn with natural populations in the environment it may decrease the wild fishes' natural survival abilities.[13]

Environment

Cleaner fish have taken over lice-reduction strategies, which were based upon chemical delousers in the past. This decreases the amount of effluent waste affecting the surrounding wild habitats in outdoor aquaculture.[11] Introducing cleaner fish into salmonid aquaculture cages has also been found to be less stressful on salmonids than medical intervention for sea lice outbreaks.[13]

Cleaner fish in the wild contribute to the overall health of aquatic communities by reducing morphological and physiological injuries by parasites to other species of fish. Maintenance of these populations of fish help the complex web of interactions remain stable.[2]

Economic

Sea lice outbreaks are detrimental to the survival of cultured salmonids and cause the majority of revenue loss in the aquaculture business. By employing the cleaner fish instead of medical intervention for sea louse management, aquaculture farmers save money.[13]

See also

References

  1. ^ a b c d e f g h i j k l m n 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. PMID 29247694.
  2. ^ a b c d e f g h i j k 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.
  3. ^ a b c d e f g h i j Soares, Marta C. (2017). "The Neurobiology of Mutualistic Behavior: The Cleanerfish Swims into the Spotlight". Frontiers in Behavioral Neuroscience. 11: 191. doi:10.3389/fnbeh.2017.00191. PMC 5651018. PMID 29089876.
  4. ^ a b 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.
  5. ^ Grutter, A. S. (2002). "Cleaning symbioses from the parasites' perspective". Parasitology. 124 (7): 65–81. doi:10.1017/S0031182002001488. ISSN 0031-1820. PMID 12396217. S2CID 26816332.
  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". Daily Telegraph. 2010-02-26. ISSN 0307-1235. Retrieved 2019-10-28.
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External links

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