Use Of Cleaner Fish Research Articles

Cleaner Fish Do Not Impact the Pigmentation of Salmon Lice (Lepeophtheirus salmonis) in Commercial Aquaculture Cages

Salmon lice are one of the biggest challenges to sustainable salmonid aquaculture. The species display high evolutionary potential, which is evident by its development of resistance to numerous chemical compounds used for delousing. In response to this, salmon farms now use non-chemical delousing methods to minimize the damage done by salmon lice, including heavy reliance on cleaner fish. Anecdotal reports from farmers and fish health personnel in areas where cleaner fish are used have suggested that salmon lice are becoming less pigmented, potentially making them harder for cleaner fish to visually detect. This experiment investigated changes in the pigmentation of salmon lice in relation to the use of cleaner fish, louse stage and sex, temperature, preferred salmon swimming depth, daylength, and salinity. Salmon lice were sampled from snorkel cages on a commercial salmon farm where three cages were stocked with farmed lumpfish and ballan wrasse, and three cages were without cleaner fish. Water temperature, salinity, and depth were recorded using a conductivity, temperature, and depth recorder. Pigmentation was measured via photographic analysis of individual lice. Although louse pigmentation varied considerably throughout the experiment, using cleaner fish throughout a single production cycle did not reduce average louse pigmentation compared to control cages. On average, male lice were significantly darker pigmented than females, but otherwise there were no patterns in louse pigmentation in relation to life stage, salinity, temperature, or daylength. Salmon lice exhibit a high degree of evolvability and have become resistant to every chemical removal treatment developed thus far. The present data suggest that, with the densities and species of cleaner fish commonly used in commercial salmon production, there is not strong directional selection on louse pigmentation. Lice, at least with regard to visual appearance, are not likely to adapt in a way which reduces cleaner fish efficacy anytime soon.

Global, regional, and cryptic population structure in a high gene-flow transatlantic fish.

Lumpfish (Cyclopterus lumpus) is a transatlantic marine fish displaying large population sizes and a high potential for dispersal and gene-flow. These features are expected to result in weak population structure. Here, we investigated population genetic structure of lumpfish throughout its natural distribution in the North Atlantic using two approaches: I) 4,393 genome wide SNPs and 95 individuals from 10 locations, and II) 139 discriminatory SNPs and 1,669 individuals from 40 locations. Both approaches identified extensive population genetic structuring with a major split between the East and West Atlantic and a distinct Baltic Sea population, as well as further differentiation of lumpfish from the English Channel, Iceland, and Greenland. The discriminatory loci displayed ~2-5 times higher divergence than the genome wide approach, revealing further evidence of local population substructures. Lumpfish from Isfjorden in Svalbard were highly distinct but resembled most fish from Greenland. The Kattegat area in the Baltic transition zone, formed a previously undescribed distinct genetic group. Also, further subdivision was detected within North America, Iceland, West Greenland, Barents Sea, and Norway. Although lumpfish have considerable potential for dispersal and gene-flow, the observed high levels of population structuring throughout the Atlantic suggests that this species may have a natal homing behavior and local populations with adaptive differences. This fine-scale population structure calls for consideration when defining management units for exploitation of lumpfish stocks and in decisions related to sourcing and moving lumpfish for cleaner fish use in salmonid aquaculture.

First feed matters: The first diet of larval fish programmes growth, survival, and metabolism of larval ballan wrasse (Labrus bergylta)

The use of cleaner fish, such as the ballan wrasse (Labrus bergylta), is important for combatting the sea lice problem in salmonid cage farming. Ballan wrasse is the only wrasse species that is cultivated, though only about 50% of the approx. 3 million ballan wrasse used in 2020 was produced by aquaculture. The major obstacle for large scale cultivation of ballan wrasse is the difficult first feeding phase and a lack of functional feeding protocols.Like most pelagic marine fish larvae, ballan wrasse do not accept inert diets as first feed, and feeding regimes based on rotifers and Artemia (brine shrimps) usually lead to mixed results and are far from being optimal nutrition for the larvae.In a 48-day start feeding experiment, we studied the feasibility of replacing rotifers by an Experimental cirriped diet or copepod nauplii (Acartia tonsa) and replacing Artemia by nauplii of the cirriped Semibalanus balanoides. Later, all treatments received the same formulated diets. We sampled larvae at each feed transition to analyze different response variables such as growth, morphometry, gene expression, lipidomics, histology, and microbiology.We found significant differences in survival rates and growth. Larvae fed copepods as the first diet had significantly higher survival rates than larvae start-fed on either rotifers or small experimental cirripeds, and this pattern was also reflected in early growth and bone development. Gut histology at the end of the experiment (48 days after hatching) showed a more developed intestinal tissue in the larval group fed copepods first and cirripeds as the second diet compared to the other larval groups.Gene expression at day 48 post hatch still revealed pronounced differences between the larval group first fed on rotifers and larvae from the other three feeding regimes which received natural, unenriched diets. Even weeks after receiving the same formulated diets, lipidomics analyses revealed that several lipid species correlated either negatively or positively with larval growth rates or mortality.Our results are a clear indication for nutritional programming, pointing towards the importance the first diet has for the further life of a fish.

Testing a simple energy-budget model for yolk-feeding stages of cleaner fish

The use of cleaner fish is an environmentally-friendly approach to combat the salmon louse, threatening commercial salmon farming. Dynamic Energy Budget (DEB) modelling helps understand the bioenergetics of early life stages of the cleaner fish, and can thereby aid optimisation of their culturing. Here, we report on our attempts to parametrise DEBkiss models for the yolk-feeding stages of two cleaner fish species, ballan wrasse (Labrus bergylta) and lumpfish (Cyclopterus lumpus). A range of measurements was taken over early development, including biometry (using imaging), weight and composition, as well as measurements of respiration rate. Despite the previous success of applying the DEBkiss model to early life stages of Atlantic cod, the model failed to capture the patterns of yolk depletion for ballan wrasse. The main issues were related to substantial changes in the water content of both yolk and structure over development, and a stop of growth before disappearance of the yolk sac. These issues require further experimental work to address, especially more efficient proxies for the dry mass of yolk and structure, such that these compartments can be efficiently separated. Nevertheless, apart from the pattern of yolk depletion, the model provides a reasonable explanation of all traits simultaneously. This indicates that model modifications may only need to be minor. For lumpfish, the data set was quite limited for testing of the DEBkiss model, due to the opacity of the egg and the fact that there was only one time point with measurements post hatch. Nevertheless, the data are consistent with the model. The modelling results indicate that both cleaner-fish species may have very similar bioenergetic parameters (and quite similar to Atlantic cod as well); the conspicuous difference in early life history may be mainly caused by the larger yolk provisioning in the egg, and late hatching, in lumpfish. The DEBkiss model is a simple and promising tool for bioenergetics of fish early-life stages. However, its application and in-depth testing is currently limited by the difficulties of obtaining detailed measurements on these life stages.

Personality profiling may help select better cleaner fish for sea-lice control in salmon farming

Lumpfish (Cyclopterus lumpus) are increasingly being used as cleaner fish to control parasitic sea-lice in salmon farming, but cleaning rates are very variable and not all individuals eat sea-lice, which increases the risk of emaciation and has ethical and practical implications. Selecting good cleaners is a priority to make the industry more sustainable, but there is little information on what behaviours make cleaner fish effective under a commercial setting. We examined variation in lumpfish personalities according to the five-factor personality model that takes into account differences in activity, anxiety (shelter use, thigmotaxis), aggression, sociality, and boldness (neophobia). We then quantified how variation in lumpfish personalities influenced interactions with naïve Atlantic salmon (Salmo salar), without the confounding effects of variation in sea-lice loads. Variation in activity, sociality, aggression and neophobia, but not in anxiety, was repeatable. Neophilic, non-aggressive lumpfish spent more time inspecting salmon than neophobic and aggressive individuals, but salmon fled in the presence of the most active and social individuals, suggesting there may be an optimal cleaner fish personality amenable to artificial selection. The personality screening protocols developed in this study could inform a more efficient use of cleaner fish in salmon farming and reduce the number of individuals required to control sea-lice.

Prediction of goldsinny wrasse (Ctenolabrus rupestris) minimum size required to avoid escape through salmon (Salmo salar) farm nets

The salmon (Salmo salar) aquaculture industry has used various practices to treat salmon louse (Lepeophtheirus salmonis) infections, including mechanical, thermal, and chemical treatments. However, due to the environmental and fish welfare issues, and limitations associated with these practices, the use of cleaner fish has become a popular complementary delousing method. The goldsinny wrasse (Ctenolabrus rupestris) is the smallest among the delousing species used, which makes it most susceptible to escape through the salmon cage netting. Escape of goldsinny wrasse into the wild would lead to additional costs for farmers and could spread diseases and genetically contaminate wild stocks, as these cleaner fish are often transported over large distances from the capture site to the aquaculture site. The Norwegian aquaculture industry sets the goldsinny wrasse minimum size for use with different salmon cage mesh sizes using guidelines based on simple penetration tests that do not consider the potential compressibility of fish or changes in mesh state due to factors such as sea conditions and maintenance operations. Using morphological measurements of 100 fish, results of 47,800 penetration tests, and computer simulation, we predicted the minimum sizes required to avoid escape of goldsinny wrasse through and array of salmon farm nets. Our results show that the minimum stocking sizes for goldsinny wrasse in the current guidelines used by the industry may pose escape risk. Further, our findings illustrate the importance of considering fish compressibility and mesh state when estimating the fish minimum sizes required to avoid escape via cage netting mesh penetration. Finally, the results presented here provide the salmon farming industry with new guidelines that will contribute to reducing goldsinny wrasse escape from the cages and ultimately lessen the potential environmental consequences of such escape. For the two most typical mesh-size used by the aquaculture industry in Norway i.e. 30 and 50 mm and under the worst-case scenario i.e. slack state, the minimum recommended lengths for goldsinny wrasse would be ~128 (~28) and ~ 206 mm (~123 g). These sizes represent an underestimation of 16 and 24% in the sizes of goldsinny wrasse necessary to avoid escape compared to the current guidelines based on stiff meshes.

Pasteurella spp. Infections in Atlantic salmon and lumpsucker.

The use of cleaner fish as a delousing method in Norwegian salmonid aquaculture has increased tremendously over the last few years. This has led to the emergence of a new large industry of farming lumpsuckers (Cyclopterus lumpus L.). The use of lumpsuckers as cleaner fish has, however, not been problem-free. Bacterial diseases cause high mortalities with pasteurellosis as one of the major emerging diseases. During the past few years, outbreaks of pasteurellosis in farmed Atlantic salmon (Salmo salar L.) have become more frequent. This has led to an increasing concern that this disease will become common in salmon farming as well. The purpose of this study was to investigate the susceptibility of Atlantic salmon to Pasteurella spp. infection and the possibility of lumpsuckers transmitting pasteurellosis to Atlantic salmon. Atlantic salmon were experimentally challenged, either by bath or by cohabitation with challenged lumpsuckers, using two different strains of Pasteurella spp. (originating from lumpsucker and Atlantic salmon, respectively). No clinical signs of pasteurellosis were observed on any of the Atlantic salmon. The lumpsuckers were, however, equally susceptible to both isolates. In addition, clear differences in histopathological changes were observed between individuals challenged with the two isolates.

Salmon Louse (Lepeophtheirus salmonis (Krøyer)) Control Methods and Efficacy in Atlantic Salmon (Salmo salar (Linnaeus)) Aquaculture: A Literature Review

The salmon louse (Lepeophtheirus salmonis) causes problems in Atlantic salmon (Salmo salar) aquaculture in the Northern Hemisphere, because infestations can result in both a loss of production and in fish mortality. Several types of treatment have been used to control louse infestations, but these have seen varying success. The aim of this review is to examine the efficacy and safety of commonly used treatments (chemical, biological, mechanical, and preventive measures) as documented in peer-reviewed publications. Efficacy is assessed in relation to a reduction in numbers of lice, and safety is assessed as a lack of negative treatment-associated effects on fish health and welfare (Atlantic salmon and/or cleaner fish). Most chemical treatments showed decreasing efficacy over time, together with the use of increasing concentrations as a result of the development of resistance to the treatments by lice. The need for a restrictive use of pesticides to preserve treatment efficacy has been emphasized. The use of cleaner fish was suggested to be effective, with few or no negative effects towards Atlantic salmon. The use of cleaner fish would be preferable to chemical treatment if the farmed fish health and welfare criteria are met. At present, the number of peer-reviewed publications relating to other forms of treatment and prevention are sparse.

Lumpfish (Cyclopterus lumpus, Linnaeus) is susceptible to viral nervous necrosis: Result of an experimental infection with different genotypes of Betanodavirus.

In recent years, the use of cleaner fish for biological control of sea lice has increased considerably. Along with this, a number of infectious diseases have emerged. The aim of this study was to investigate the susceptibility of lumpfish (Cyclopterus lumpus) to Betanodavirus since it was detected in asymptomatic wild wrasses in Norway and Sweden. Three betanodaviruses were used to challenge lumpfish: one RGNNV genotype and two BFNNV genotypes. Fish were injected and monitored for 4weeks. Brain samples from clinically affected specimens, from weekly randomly selected fish and survivors were subjected to molecular testing, viral isolation, histopathology and immunohistochemistry. Reduced survival was observed but was attributed to tail-biting behaviour, since no nervous signs were observed throughout the study. Betanodavirus RNA was detected in all samples, additionally suggesting an active replication of the virus in the brain. Viral isolation confirmed molecular biology results and revealed a high viral titre in BFNNV-infected groups associated with typical lesions in brains and eyes of survivor fish. We concluded that lumpfish are susceptible to Betanodavirus, as proven by the high viral titre and brain lesions detected, but further studies are necessary to understand if Betanodavirus can cause clinical disease in this species.

How are the salmon lice (Lepeophtheirus salmonis Krøyer, 1837) in Atlantic salmon farming affected by different control efforts: A case study of an intensive production area with coordinated production cycles and changing delousing practices in 2013-2018.

The aims of the present study were to describe the salmon lice (Lepeophtheirus salmonis Krøyer, 1837) situation in an intensive salmon production area in mid-Norway and to consider implications of changing practices of how salmon lice infestation can be controlled. The results in this study suggest that there are steps that can be carried out to keep salmon lice under control even during years when the temperature facilitates a quick salmon lice development. The present work indicates that the use of cleaner fish can delay the time it takes adult female lice to reach 0.1 per salmon in the beginning of a production cycle. It suggests that the timing of cleaner fish deployment into salmon cages can influence its effectiveness in controlling salmon lice. It also gives caution to letting salmon lice develop unchecked, even at levels far below the current lice limit, because of the difficulties to control salmon lice when the external infection pressure is too high. This study took place during a rapid change in delousing methods, in an area with coordinated salmon production. Despite its exploratory nature, this study offers insights into the salmon lice fluctuations in relation to efforts aimed at controlling it.

Isolation of salmonid alphavirus subtype 6 from wild-caught ballan wrasse, Labrus bergylta (Ascanius).

The use of cleaner fish as a biological control for sea lice in Atlantic salmon aquaculture has increased in recent years. Wild-caught wrasse are commonly used as cleaner fish in Europe. In Ireland, samples of wrasse from each fishing area are screened for potential pathogens prior to their deployment into sea cages. Salmonid alphavirus was isolated from a pooled sample of ballan wrasse, showing no signs of disease, caught from the NW of Ireland. Partial sequencing of the E2 and nsP3 genes showed that it was closely related to the previously reported SAV subtype 6. This represents only the second isolation of this subtype and the first from a wild fish species, namely ballan wrasse.

Shelters can negatively affect growth and welfare in lumpfish if feed is delivered continuously.

Due to the socioeconomic importance of salmon farming in the North Atlantic and the economic impact of sea lice in this industry, there is high demand for novel pest control methods. One such method is the use of cleaner fish to remove the lice from the salmon. A cleaner fish that has recently gained in popularity due to its ability to work in cold water, is the lumpfish (Cyclopterus lumpus). This fish varies in efficiency, but when mortality is low and cleaning optimal, the fish are successful in keeping parasite burdens low. However, there is some concern for the welfare of lumpfish in the industry, because mortality is often high. This is sometimes attributed to inadequate feeding and shelter. Here we compare growth, body condition, and fin health of fish reared for four weeks in a crossed treatment design crossing shelter availability (shelter vs none) and feed delivery method (manual meal time feeds and continuous automated feeding). In terms of weight gain, shelter availability interacted with feeding method, with fish that had access to shelters and were fed using automated feeders gaining less weight than other fish. Fin health was not affected, but body condition was lowered both by access to shelter and being fed continuously. The results indicate a need to carefully consider how feeding method and shelter use is combined, both in cages and during rearing on land.

An overview of cleaner fish use in Ireland.

Sea lice infestations represent one of the most significant challenges facing the salmon farming industry, giving rise to lost production, additional costs of treatment and potential negative interactions with wild stocks. At present, cleaner fish, which actively remove lice from salmon, are an effective, biological, long-term option which has been adopted by many countries. In Ireland, several key studies were conducted in the 1990s on the use of wild-caught wrasse (corkwing, goldsinny and rock cook) as cleaner fish in experimental and commercial scale trials. More recently, the National University of Ireland Galway (NUIG), at their marine research facility in Carna (CRS), has undertaken applied research on ballan wrasse and lumpsucker. Currently, CRS is providing lumpsucker juveniles and research and development for the Irish salmon industry with support from BIM (Ireland's Seafood Development Agency) and Marine Harvest Ireland. There is a large amount of research currently being carried out in this area in all countries that are utilizing cleaner fish technology. The current focus in Ireland is the development of a native lumpsucker broodstock to facilitate its sustainable production. The aim of this article was to provide an overview of the research, challenges and use of cleaner fish in Ireland.

A model of the process of spillover and adaption leading to potential emergence of disease in salmon held with cleaner fish used to control lice

Sea lice control is a very important issue for sustainable salmon aquaculture, and cleaner fish (largely wrasse and lumpfish) are increasingly used as a key tool for this control. Such fish can carry pathogens, which they can potentially spread to the salmon. A modelling approach is used to help guide practice on cleaner fish use. The model simulates transmission of pathogens within and between cleaner fish and salmon populations. A function modifying the coefficient of infection allows simulation of pathogens input with cleaner fish to adaptation to salmon with passage. Adaption may require different numbers of passage events and may face obstacles that make adaption easier or more difficult, allowing the exploration of a range of potential emergent pathogens with different properties. The model includes the production cycle of salmon, whereby all salmon are periodically removed and replaced with naïve fish, but a proportion of the cleaner fish may be reused in the next production cycle. An assessment of the likely consequence of emergent diseases relative to consequence of sea lice infection is made using an analysis of literature. The modelling concludes that any emergent disease is unlikely to be of as serious consequence as sea lice already are, and therefore the risk from emergent disease is low relative to benefit of controlling lice. This risk can be further reduced by practices that limit input of infection and by restricting reused of cleaner fish in subsequent production cycle. The risk from reusing small numbers is generally substantially lower than reusing larger numbers, but avoiding reuse is more effective, especially if there is limited input of pathogens with new cleaner fish. More data as cleaner fish are increasingly used will allow more accurate assessment of risk, but the model provides a flexible framework for assessing risk of emergence of potential new pathogens and advising on good practice while information is limited and a precautionary approach is inappropriate owing to the clear benefits of effective lice control.

Characterization of IgM in Norwegian cleaner fish (lumpfish and wrasses)

The use of cleaner fish in Norwegian aquaculture has to a large extent been based on wild catches, but breeding of lumpfish and ballan wrasse is currently increasing. Due to disease problems and required vaccine development, tools to study immune responses and a better understanding of the immune system in these species is demanded. The present study comprises lumpfish (Cyclopterus lumpus) and five species of wrasses: Ballan wrasse (Labrus bergylta), rock cook (Centrolabrus exoletus), cuckoo wrasse (Labrus mixtus), corkwing wrasse (Symphodus melops), and goldsinny wrasse (Ctenolabrus rupestris). We present a comparison of the IgM sequences, phylogenetic relationship to other teleosts and characteristic features of IgM in the species studied. The lumpfish IgM heavy chain sequence was assembled from high throughput cDNA sequencing whereas the wrasse sequences were determined by molecular cloning. The secreted form of the IgM heavy chain from all species consisted of four constant Ig domains. IgM was purified from lumpfish and ballan wrasse sera by gel filtration followed by anion exchange chromatography, and polyclonal sera were produced against these proteins. Antisera against ballan wrasse IgM showed cross-reactivity to all analyzed species of wrasses, some cross-reactivity to lumpfish, very low reaction to salmon, and no reaction to cod. Anti- IgM sera against lumpfish cross-reacted to the light chain of all species studied. Wrasses and lumpfish IgM showed high binding affinities for protein A. IgM concentration in adult ballan wrasse (700–800 g) was measured by single radial immunodiffusion assay and found to be 13.4 mg/ml which is about 36% of the total protein concentration. The IgM concentration in lumpfish (600–3600 g) was estimated to 1–2.6 mg/ml, which corresponds to approximately 3% of the total protein concentration.

Aeromonas salmonicida infection levels in pre- and post-stocked cleaner fish assessed by culture and an amended qPCR assay.

Due to increasing resistance to chemical therapeutants, the use of 'cleaner fish' (primarily wrasse, Labridae, species) has become popular in European salmon farming for biocontrol of the salmon louse, Lepeophtheirus salmonis (Krøyer). While being efficient de-licers, cleaner fish mortality levels in salmon cages are commonly high, and systemic bacterial infections constitute a major problem. Atypical furunculosis, caused by Aeromonas salmonicida A-layer types V and VI, is among the most common diagnoses reached in clinical investigations. A previously described real-time PCR (qPCR), targeting the A.salmonicida A-layer gene (vapA), was modified and validated for specific and sensitive detection of all presently recognized A-layer types of this bacterium. Before stocking and during episodes of increased mortality in salmon cages, cleaner fish (primarily wild-caught wrasse) were sampled and screened for A.salmonicida by qPCR and culture. Culture indicated that systemic bacterial infections are mainly contracted after salmon farm stocking, and qPCR revealed A.salmonicida prevalences of approximately 4% and 68% in pre- and post-stocked cleaner fish, respectively. This underpins A.salmonicida's relevance as a contributing factor to cleaner fish mortality and emphasizes the need for implementation of preventive measures (e.g. vaccination) if current levels of cleaner fish use are to be continued or expanded.

A Modelling Framework for Assessing the Risk of Emerging Diseases Associated with the Use of Cleaner Fish to Control Parasitic Sea Lice on Salmon Farms.

Sea lice are the most damaging parasite of marine salmonids, both economically and in terms of potential impacts on wild fish. An increasingly widely applied control is the use of cleaner fish (CF) such as wrasse that eat lice. However, such CF can carry pathogens that may cause disease in salmon, including the potential emergence of new diseases. This is not just a theoretical risk, as demonstrated by a recent outbreak of viral haemorrhagic septicaemia in wrasse held on salmon farms in Shetland. A modelling framework is developed to identify conditions in which emergence might occur, and, from this, means of reducing risk. Diseases that might emerge easily in farmed salmon would be likely to have already done so by other routes of exposure, and if risks are very low, they would need to be greatly enhanced to become significant relative to costs of lice control. CF may most enhance risks from disease with moderate probability of emerging. Risks of emergence can be reduced by replacing wild-caught with hatchery-reared CF, minimizing mixing of CF from different sources, surveillance for clinical disease in the CF and ensuring strategic biosecurity (area management with synchronized fallowing). Reuse of CF for a second salmon production cycle may reduce costs and even probability of infection (especially from wild-caught CF), but should only be considered as part of a rigorous area management programme because the practice presents opportunities for pathogens to adapt to salmon by weakening fallowing.

Salmon lice--impact on wild salmonids and salmon aquaculture.

Salmon lice, Lepeophtheirus salmonis, are naturally occurring parasites of salmon in sea water. Intensive salmon farming provides better conditions for parasite growth and transmission compared with natural conditions, creating problems for both the salmon farming industry and, under certain conditions, wild salmonids. Salmon lice originating from farms negatively impact wild stocks of salmonids, although the extent of the impact is a matter of debate. Estimates from Ireland and Norway indicate an odds ratio of 1.1:1-1.2:1 for sea lice treated Atlantic salmon smolt to survive sea migration compared to untreated smolts. This is considered to have a moderate population regulatory effect. The development of resistance against drugs most commonly used to treat salmon lice is a serious concern for both wild and farmed fish. Several large initiatives have been taken to encourage the development of new strategies, such as vaccines and novel drugs, for the treatment or removal of salmon lice from farmed fish. The newly sequenced salmon louse genome will be an important tool in this work. The use of cleaner fish has emerged as a robust method for controlling salmon lice, and aquaculture production of wrasse is important towards this aim. Salmon lice have large economic consequences for the salmon industry, both as direct costs for the prevention and treatment, but also indirectly through negative public opinion.

Cleaner Fish Use Tactile Dancing Behavior as a Preconflict Management Strategy

The most commonly asked question about cooperative interactions is how they are maintained when cheating is theoretically more profitable [1]. In cleaning interactions, where cleaners remove parasites from apparently cooperating clients, the classical question asked is why cleaner fish can clean piscivorous client fish without being eaten, a problem Trivers [2] used to explain reciprocal altruism. Trivers [2] suggested that predators refrain from eating cleaners only when the repeated removal of parasites by a particular cleaner results in a greater benefit than eating the cleaner. Although several theoretical models have examined cheating behavior in clients [3, 4], no empirical tests have been done (but see Darcy [5]). It has been observed that cleaners are susceptible to predation [6, 7]. Thus, cleaners should have evolved strategies to avoid conflict or being eaten. In primates, conflicts are often resolved with conflict or preconflict management behavior [8]. Here, I show that cleaner fish tactically stimulate clients while swimming in an oscillating “dancing” manner (tactile dancing) more when exposed to hungry piscivorous clients than satiated ones, regardless of the client's parasite load. Tactile dancing thus may function as a preconflict management strategy that enables cleaner fish to avoid conflict with potentially “dangerous” clients.

Biting cleaner fish use altruism to deceive image-scoring client reef fish.

Humans are more likely to help those who they have observed helping others previously. Individuals may thus benefit from being altruistic without direct reciprocity of recipients but due to gains in 'image' and associated indirect reciprocity. I suggest, however, that image-scoring individuals may be exploitable by cheaters if pay-offs vary between interactions. I illustrate this point with data on cleaner-client reef fish interactions. I show the following: (i) there is strong variation between cleaners with respect to cheating of clients (i.e. feeding on client tissue instead of parasites); (ii) clients approach cleaners, that they observe cooperating with their current client and avoid cleaners that they observe cheating; (iii) cleaners that cheat frequently are avoided more frequently than more cooperative cleaners (iv) cleaners that cheat frequently behave altruistically towards their smallest client species; (v) altruistic acts are followed by exploitative interactions. Thus, it appears that cleaners indeed have an image score, which selects for cooperative cleaners. However, cheating cleaners use altruism in potentially low-pay-off interactions to deceive and attract image-scoring clients that will be exploited.