Presence of cleaner wrasse increases the recruitment of damselfishes to coral reefs

Mutualisms affect the biodiversity, distribution and abundance of biological communities. However, ecological processes that drive mutualism-related shifts in population structure are often unclear and must be examined to elucidate how complex, multi-species mutualistic networks are formed and structured. In this study, we investigated how the presence of key marine mutualistic partners can drive the organisation of local communities on coral reefs. The cleaner wrasse, Labroides dimidiatus, removes ectoparasites and reduces stress hormones for multiple reef fish species, and their presence on coral reefs increases fish abundance and diversity. Such changes in population structure could be driven by increased recruitment of larval fish at settlement, or by post-settlement processes such as modified levels of migration or predation. We conducted a controlled field experiment to examine the effect of cleaners on recruitment processes of a common group of reef fishes, and showed that small patch reefs (61–285 m²) with cleaner wrasse had higher abundances of damselfish recruits than reefs from which cleaner wrasse had been removed over a 12-year period. However, the presence of cleaner wrasse did not affect species diversity of damselfish recruits. Our study provides evidence of the ecological processes that underpin changes in local population structure in the presence of a key mutualistic partner.     

Effects of group behavior in the predatory raid on damselfish nests by the false cleanerfish Aspidontus taeniatus

The false cleanerfish Aspidontus taeniatus, which resembles the bluestreak cleaner wrasse Labroides dimidiatus, is one of the best-known examples of mimicry in vertebrates. This mimicry system has been viewed as an aggressive mimicry to bite fish fins. However, recent field studies have reported that large individuals of the false cleanerfish often form groups and jointly raid damselfish nests to eat eggs that are guarded by their parents. The benefits of group behavior have been reported in a variety of animals. In the case of false cleanerfish, parental defense of territorial damselfishes is the main factor that constrains the availability of nutritionally valuable food resources. Here, we conducted field observations on the reefs of Okinawa, and found that the false cleanerfish formed groups of 2–12 individuals when they raided breeding nests of 13 species of damselfishes (Pomacentridae) and one species of triggerfish (Balistidae). Since the cleaner wrasse does not form such groups, the feeding groups of the false cleanerfish are assumed to reduce the effectiveness of mimicry. However, our results showed that the group behavior has two effects: a dilution effect, which reduces the risk of being attacked by egg-guarding fish, and an increase in foraging efficiency. We conclude that the false cleanerfish need to form foraging groups during egg-eating because the mimicry has no effect on parental damselfishes.     

Enhanced colonization success and competition associated with conspecifics in cleaner fish Labroides dimidiatus juveniles

The cleaner fish Labroides dimidiatus affects the abundance of many fishes, including their juveniles, yet how they affect the post-settlement processes of conspecifics remains poorly known. Using a long-term experiment, where L. dimidiatus were regularly removed from seven patch reefs (removals) for 10 years or left undisturbed (controls) on nine, the effect of conspecifics’ presence on recently settled L. dimidiatus juveniles and the relationship between juvenile and adult abundance on control reefs were examined. Repeated sampling 4–40 months after manipulating L. dimidiatus revealed that the abundance of juveniles on removals, compared with controls reefs, was 72 % (95 % CI = 65–79 %) lower; this effect did not vary over time. Thus, decreased colonization was associated with resident conspecific absence and this effect was present after 4 months. Since other studies found 4 months of cleaner absence was too short to affect client abundance—a factor also known to enhance L. dimidiatus colonization—it was likely not involved here. Over 10 years, individual control reefs were almost always occupied by at least one adult (89–100 % of times sampled); but in summer, when L. dimidiatus colonization was highest, juvenile abundance was negatively related with adult abundance at one of two sites, with the expected number of juveniles decreasing by 67 % (95 % CI = 51–83 %) with each additional adult. This suggests a deleterious effect of adults on juveniles at one site, such as competition. Enhanced colonization of juveniles associated with adult presence may partly explain the relative permanence of fish cleaning stations.     

Differential effects of native vs. invasive predators on a common Caribbean reef fish

Predators may have consumptive (lethal) and non-consumptive (sub-lethal) effects on prey. Non-consumptive effects include altered behavior and reduced growth and fecundity. Native prey may not recognize non-native predators as a threat, and therefore may suffer pronounced effects. Additionally, non-native predators may elicit different behavioral responses from prey compared to native predators. Theory predicts that consumptive effects should be greater for non-native predators (due to prey naiveté), and non-consumptive effects should be greater for native predators (due to predator recognition). To test these hypotheses, I monitored bicolor damselfish (Stegastes partitus) in the presence of invasive predatory Pacific lionfish (Pterois spp.), a native predator (graysby, Cephalopholis cruentata), and an egg predator (bluehead wrasse, Thalassoma bifasciatum). Body size and location of lionfish and graysby were monitored on reefs in the Bahamas. Bicolor fecundity was measured as the number and size of egg-masses that individual fish laid. Bicolor fecundity was negatively correlated with lionfish density but not graysby or bluehead density. Neither predator had a detectable effect on bicolor body size, but lionfish density was negatively correlated with the size of mature adult damselfish. I observed behavioral responses of bicolors to the two piscivores, to bluehead wrasse, and to two herbivorous fishes (Acanthurus coeruleus, Scarus spp.) as non-aggressive controls. Bicolors changed behavior (feeding and aggression) in the presence of all native fishes, but not in the presence of lionfish. Thus, differential effects exist between native and non-native predators, and invasive lionfish pose a non-consumptive threat to bicolor damselfish via reduced growth and fecundity.     

Cleaning interactions at the only atoll in the South Atlantic

In marine ecosystems, cleaning is a mutualistic relationship in which so-called cleaners remove ectoparasites, diseased tissue, or mucus from the body of their clients, and thus help to maintain a healthy reef community. In spite of its importance in many marine habitats, this interaction remains poorly understood, particularly at oceanic islands. Here, we present the first comprehensive study of cleaning interactions in a reef fish assemblage at Rocas, the only atoll in the South Atlantic. We recorded 318 cleaning events, in which six fish species, including two endemic ones, and two shrimp species acted as cleaners. The clients serviced by these cleaners were 21 bony fish species, one shark and one sea turtle. The cleaner wrasse Thalassoma noronhanum and the cleaner goby Elacatinus phthirophagus were the cleaners with the greatest number of events and species richness of clients. Additionally, 82% of clients in the cleaning events were non-piscivores, and the abundance of both cleaners and clients positively influenced the number of cleaning events (R² = 0.4; p < 0.001). Our results indicate that Rocas atoll has a high species richness of cleaner species despite its small size and highlight the importance of studies of cleaning symbiosis, even in isolated places with low species richness, for a better comprehension of this association in reefs.     

Long-term effects of the cleaner fish Labroides dimidiatus on coral reef fish communities

Cleaning behaviour is deemed a mutualism, however the benefit of cleaning interactions to client individuals is unknown. Furthermore, mechanisms that may shift fish community structure in the presence of cleaning organisms are unclear. Here we show that on patch reefs (61–285 m²) which had all cleaner wrasse Labroides dimidiatus (Labridae) experimentally removed (1–5 adults reef⁻¹) and which were then maintained cleaner-fish free over 8.5 years, individuals of two site-attached (resident) client damselfishes (Pomacentridae) were smaller compared to those on control reefs. Furthermore, resident fishes were 37% less abundant and 23% less species rich per reef, compared to control reefs. Such changes in site-attached fish may reflect lower fish growth rates and/or survivorship. Additionally, juveniles of visitors (fish likely to move between reefs) were 65% less abundant on removal reefs suggesting cleaners may also affect recruitment. This may, in part, explain the 23% lower abundance and 33% lower species richness of visitor fishes, and 66% lower abundance of visitor herbivores (Acanthuridae) on removal reefs that we also observed. This is the first study to demonstrate a benefit of cleaning behaviour to client individuals, in the form of increased size, and to elucidate potential mechanisms leading to community-wide effects on the fish population. Many of the fish groups affected may also indirectly affect other reef organisms, thus further impacting the reef community. The large-scale effect of the presence of the relatively small and uncommon fish, Labroides dimidiadus, on other fishes is unparalleled on coral reefs.     

Predation on crown-of-thorns starfish larvae by damselfishes

Examining the functional response of predators can provide insight into the role of predation in structuring prey populations and ecological communities. This study explored feeding behaviour and functional responses of planktivorous damselfishes when offered captive reared larvae of crown-of-thorns starfish, Acanthaster sp., with the aim of determining whether these predators could ever play a role in moderating outbreaks of Acanthaster sp. We examined predatory behaviour of 11 species of planktivorous damselfish, testing: (1) the relationship between predator size and predation rate, both within and among fish species; (2) consumption rates on larvae of Acanthaster sp. versus larvae of a common, co-occurring coral reef asteroid Linckia laevigata; (3) maximal feeding rates upon both Acanthaster sp. and L. laevigata; and (4) functional responses of planktivorous fishes to increasing densities of Acanthaster sp. Consumption rates of crown-of-thorns larvae by damselfishes were independent of predator size; however, there was a significant negative relationship between predator size and consumption rate of L. laevigata, when pooling across all predatory species. Some damselfishes, including Acanthochromis polyacanthus and Amblyglyphidodon curacao, consumed larval Acanthaster sp. at a greater rate than for L. laevigata. Most predatory species (all except A. curacao and Pomacentrus amboinensis) exhibited a Type II functional response whereby the increasing feeding rate decelerated with increasing prey density. In addition to revealing that a wide range of planktivorous fishes can prey upon larvae of Acanthaster sp., these data suggest that planktivorous damselfishes may have the capacity to buffer against population fluctuations of Acanthaster sp. Importantly, predators with Type II functional responses often contribute to stability of prey populations, though planktivorous fishes may be swamped by an abnormally high influx of larvae, potentially contributing to the characteristic population fluctuations of Acanthaster sp.     

Assemblage-level effects of the introduced peacock hind (<Emphasis Type="Italic">Cephalopholis argus</Emphasis>) on Hawaiian reef fishes

The peacock hind Cephalopholis argus (family Serranidae), locally known as ‘roi’, was introduced from French Polynesia to Hawaii in the mid-twentieth century as a food fish. However, because of its association with ciguatera fish poisoning, it is rarely fished for food. Previous research indicates that roi could have a negative impact on native reef fish assemblages because of their high densities and prey consumption rates. However, it is unclear whether roi add to the cumulative mortality of prey (predation hypothesis), or whether predation is instead compensatory (doomed surplus hypothesis). This study experimentally assessed the effects of roi on reef fish populations through a long-term (5.5 year) predator removal experiment. A Before-After-Control-Impact study design was used to assess changes in fish assemblages following the removal of roi on 1.3 ha of patch reef. Increases in the density of prey-sized fish (<15 cm TL) were observed 18 months after roi removal. However, those effects did not translate into sustained increases in prey. While increases in potential competitors, wrasses (family Labridae), particularly the piscivorous ringtail wrasse Oxycheilinus unifasciatus, were observed on roi-free reefs, the fish assemblage did not diverge substantially in composition. Native reef fish appeared to resist the potential negative impacts of predation by roi, possibly through a refuge in size for some fish families. Management to protect intact fish assemblage size-structure could serve to bolster native resistance to invading species. In considering the threats facing coral reefs, and the possible solutions, roi removal alone will not likely replenish native fishery resources.     

Aggressive mimicry of the cleaner wrasse by Aspidontus taeniatus functions mainly for small blennies

The mimic blenny Aspidontus taeniatus Quoy & Gaimard is well known for its resemblance to the juvenile and adult cleaner wrasse Labroides dimidiatus (Valenciennes) in colour and shape. As various reef fishes including piscivores actively approach the cleaner wrasse to solicit cleaning by posing, two types of benefits have been suggested for this resemblance, that is, protective mimicry and aggressive mimicry. In aggressive mimicry, the mimic blenny is supposed to have considerable opportunities to bite the fin of deceived fishes when they pose, but some studies have confirmed that fin biting does not seem to be the main feeding tactic in the blenny in nature. Here, we examined the feeding tactics including fin biting by the mimic blenny in relation to its body size in a field observational survey in the coral reefs of Sesoko Island, Okinawa, Japan. The blenny was observed feeding mainly on four food items: the tentacles of Christmas tree worms, the mantle edges of boring clams, the demersal eggs in damselfishes’ nests and the fins of fishes. The feeding frequency by fin biting significantly decreased with body size, while that by egg predation significantly increased with body size of the blenny. When predating on eggs, the blenny was vigorously attacked by egg‐guarding fish, but often succeeded in raiding their nests by forming a feeding group. When feeding by fin biting, the blenny attacked prey fish without performing any cleaning. The ratio of fin biting was considerably higher in small‐sized blennies, suggesting reliance on this feeding tactic because of a difficulty in conducting a risky egg predation. Thus, our results suggest that the mimic blenny utilizes aggressive mimicry only when it is small as an alternative feeding tactic.     

Climate-driven coral reorganisation influences aggressive behaviour in juvenile coral-reef fishes

Globally, habitat degradation is altering the abundance and diversity of species in a variety of ecosystems. This study aimed to determine how habitat degradation, in terms of changing coral composition under climate change, affected abundance, species richness and aggressive behaviour of juveniles of three damselfishes (Pomacentrus moluccensis, P. amboinensis and Dischistodus perspicillatus, in order of decreasing reliance on coral). Patch reefs were constructed to simulate two types of reefs: present-day reefs that are vulnerable to climate-induced coral bleaching, and reefs with more bleaching-robust coral taxa, thereby simulating the likely future of coral reefs under a warming climate. Fish communities were allowed to establish naturally on the reefs during the summer recruitment period. Climate-robust reefs had lower total species richness of coral-reef fishes than climate-vulnerable reefs, but total fish abundance was not significantly different between reef types (pooled across all species and life-history stages). The nature of aggressive interactions, measured as the number of aggressive chases, varied according to coral composition; on climate-robust reefs, juveniles used the substratum less often to avoid aggression from competitors, and interspecific aggression became relatively more frequent than intraspecific aggression for juveniles of the coral-obligate P. moluccensis. This study highlights the importance of coral composition as a determinant of behaviour and diversity of coral-reef fishes.     

Patterns of Local Distribution of Labroides Dimidiatus in French Polynesian Atolls

Cleaning behaviour by fishes has been described to be either facultative or specialised. If being specialised in cleaning is more advantageous than to be a facultative cleaner, cleaner fishes would prefer to settle on reefs were ecological conditions promote specialisation. To test this hypothesis, we looked at the influence of physical and ecological factors on cleaner wrasse abundance, studying variations in L. dimidiatus density between 10 atolls of French Polynesia. We used a multiple regression method based on permutations of distance matrices. Our study reveals that L. dimidiatus are significantly more numerous at sites where ecological factors, such as the species richness of the fish community, may promote their cleaning activity. Moreover, they were significantly more abundant in atolls presenting a large number of sedentary fish species, few predators and few fishes living in large groups. Finally, physical factors, such as the distance between atolls, the lagoon surface, the mean percentage of hard substrate, the mean current index, and the mean depth, play a minor role in the occurrence of L. dimidiatus.     

Cleaning interactions at the southern limit of tropical reef fishes in the Western Atlantic

Cleaning associations are one of the most dynamic and complex mutualistic interactions of reef environments and are often influenced by local conditions. In the Western Atlantic (WE) most studies concentrate in tropical areas, with little attention to subtropical areas. We examined an assemblage of cleaner fish and their clients on the rocky reefs of the coast of Santa Catarina state, South Brazil, the southern limit of tropical reef fishes in the WE. We recorded 150 cleaning interactions, in which four fish species and one shrimp species acted as facultative cleaners. The grunt Anisotremus virginicus and the angelfish Pomacanthus paru serviced most clients. Fifteen fish species acted as clients, among which the most frequent was the planktivorous grunt Haemulon aurolineatum (31%). Cleaning interactions occurred mostly (87%) with non-carnivorous clients and the number of interactions was not related to the abundance of the species involved. The absence of dedicated cleaner fishes at the study sites and the replacement of their roles by facultative cleaners may be related to local conditions, including cold currents and reduction of rock cover. Under these circumstances, clients take advantage of the services offered by facultative cleaners, a characteristic of temperate areas.     

Microhabitat characteristics of <Emphasis Type="Italic">Stegastes planifrons</Emphasis> and <Emphasis Type="Italic">S. adustus</Emphasis> territories

Stegastes adustus and Stegastes planifrons are two species of damselfishes commonly found in the Caribbean. These territorial fishes have been widely studied due to their major ecological role on coral reef in controlling the growth of macroalgae that compete with corals for space and, inversely, on their deleterious role in destroying coral tissues to impulse the development of algae. However, few studies were conducted on the biotic and abiotic components of their territories. In the present study, territory size and surfaces of benthic components (macroalgae, algal turf, massive corals, branching corals, Milleporidae, sponges, sand and rubbles) were estimated for the two species at two contrasted sites. At Ilet Pigeon site (IP), the two damselfishes were found at different depth and exhibited different territory sizes. S. adustus defended a larger territory characterized by massive corals, sand and Milleporidae, while S. planifrons territories were smaller, deeper and characterized by branching corals, sponges and rubble. At Passe-à-Colas site (PC), the two fish species coexisted in the same depth range and defended territories of similar size. Their territories presented higher proportions of macroalgae, but smaller surfaces of Milleporidae than at IP. At PC, the main difference between the two species was a higher surface of massive corals inside S. planifrons territories than S. adustus territories. Differences in microhabitat characteristics between the two Stegastes seemed mostly site related. This resulted from the high plasticity of two species, allowing them to persist on Caribbean coral reefs after the decline of most branching acroporids, their former favorite habitats.     

Experimental confirmation of aggressive mimicry by a coral reef fish

A number of potential mimetic relationships between coral reef fishes have been described, but the underlying mechanisms are poorly understood. Similarities in colour between species have often been attributed to aggressive mimicry (where predators resemble models in order to deceive prey), however this has not been tested. The fang blenny, Plagiotremus rhinorhynchos is a specialized predator that feeds on tissues of other fishes. Some individuals appear to mimic the harmless cleaner wrasse Labroides dimidiatus in order to deceive fish visiting cleaning stations, thereby increasing access to food. In this study, the ecological relationship between the mimic and model was examined at Kimbe Bay (Papua New Guinea) and the hypothesis that colour similarities represent facultative aggressive mimicry was experimentally evaluated. Some juveniles exhibited a striking resemblance to the juvenile colouration of the cleaner wrasse, but only when in close proximity to the wrasse and only when similar in size. As predicted for mimics, P. rhinorhynchos co-occurred with L. dimidiatus, but was rare relative to the model. Among site comparisons showed that the abundance of mimetic type blennies was positively correlated with the abundance of juvenile cleaner wrasses. Approximately 50% of all P. rhinorhynchos were found 1 m from the nearest L. dimidiatus, a distance significantly shorter than expected if they were not associated. A cleaner wrasse removal experiment was carried out to test whether the colour displayed by the blenny and its foraging success were contingent upon the presence of a model. In all cases, removal of the model prompted a rapid colour change to a general non-mimetic colouration in P. rhinorhynchos. Removal of L. dimidiatus also resulted in a ~20% reduction in the average foraging success of the blenny compared to controls, supporting the hypothesis that the blenny is a facultative aggressive mimic of the cleaner wrasse.     

Potential effects of fishing on cleaning interactions in a tropical reef

We investigated the effect of fishing on the behavior and abundance of the cleaner fish Labroides dimidiatus on the Great Barrier Reef, Australia, by comparing cleaner fish density (underwater visual census) and behavior (focal animal observations) at a fished and a no-take site. Cleaner fish density, as well as the number of individuals and species cleaned, did not differ significantly between sites; however, L. dimidiatus spent more time interacting (physical contact involving feeding bites) with individual clients at the no-take site, indicating that fishing can indirectly affect mutualistic interactions on tropical reefs.     

Patterns of distribution and movement of fishes, <Emphasis Type="Italic">Ophthalmolepis lineolatus</Emphasis> and <Emphasis Type="Italic">Hypoplectrodes maccullochi</Emphasis>, on temperate rocky reefs of south eastern Australia

Current ecological models predict that reef fish assemblages will be strongly influenced by habitat type. Here we test hypotheses about habitat types and abundance patterns of temperate reef fishes from broad spatial scales (100 s of km) to small spatial scales of metres to tens of metres. Habitat preferences are also described over long periods of time (22 years) for two abundant taxa. Patterns of distribution and abundance varied over ～ eight degrees of latitude (29.9–37.5°S) along the coast of New South Wales, Australia. Ophthalmolepis lineolatus (Labridae) preferred kelp and Barrens habitats and juveniles were most abundant in habitats rich in algae. This species also increased in abundance from North to South. In contrast, Hypoplectrodes maccullochi (Serranidae) were usually only found in the Barrens habitat and great variation was found among locations. Both taxa were most abundant on urchin grazed deep reefs (over 10 m deep). Habitat preferences of O. lineolatus and H. maccullochi appeared resistant to major environmental perturbations that included large El Niño events in 1991, 1998 and 2002. Home ranges of O. lineolatus varied from 52 m² to 1,660 m² and often overlapped; fish of all sizes were most abundant in algal dominated habitat. Limited movements and small home ranges (2.1–11.6 m²) combined with a strong affiliation for shelter indicated that most H. maccullochi are strongly site-attached. Habitat type is important to these taxonomically different fishes, but to varying degrees where H. maccullochi was more of a habitat specialist than O. lineolatus and would be more vulnerable to perturbations that alter Barrens. Changes in reef habitats will have a great influence on fish assemblages and this should also be considered in coastal planning (e.g. for Marine Protected Areas, MPAs) and the assessments of resistance and resilience of fishes to climate change.     

Estuarine fish behavior around slotted water control structures in a managed salt marsh

Estuaries are composed of multiple interconnected habitat types used by transient fish species during their period of estuarine residency. Structural marsh management restricts habitat connectivity and impedes the movement of fishes among these habitat types by limiting access via water control structures (WCSs) between the managed area and the rest of the estuary. While some general information on fish passage rates is available, species-specific information on passage through WCSs is lacking for salt marsh fishes. We monitored tagged fishes from March 2012 through November 2013 using passive integrated transponder antenna arrays at two identical WCSs in the Calcasieu Lake estuary, Louisiana, USA, to assess the effect of slotted WCSs on fish behavior. A total of 420 individuals of 15 species was tagged and released at the WCSs; of these, 145 individuals representing 11 species were later detected at the WCSs. Five species comprised most (93%) of the detected individuals: Elops saurus (n = 60), Mugil cephalus (n = 43), Sciaenops ocellatus (n = 20), Pogonias cromis (n = 7), and Ariopsis felis (n = 5). Passage rates were low, with most of the observed fishes (n = 80) passing only once through the structures. Other than E. saurus, which was only observed migrating out of the managed marsh, no clear pattern in swimming direction was observed for the other species. Detected species were all present primarily during the summer and fall, however, diel activity at the structures varied by species. The WCSs in our study area appeared to attract and congregate fishes, functioning more like ecological hotspots, rather than simply facilitating fish passage.     

Mesophotic reef fish assemblages of the remote St. Peter and St. Paul’s Archipelago,Mid-Atlantic Ridge,Brazil

Mesophotic reef fish assemblages (30–90 m depth) of the small and remote St. Peter and St. Paul’s Archipelago (SPSPA), Mid-Atlantic Ridge, Brazil, were characterized using remotely operated vehicles. Ordination analyses identified distinct fish assemblages in the upper (30–50 m) and lower (50–90 m) mesophotic zones, the former characterized by high abundances of species that are also abundant at euphotic reefs (Caranx lugubris, Melichthys niger, Stegastes sanctipauli and Chromis multilineata) and the latter dominated by two mesophotic specialists (Prognathodes obliquus and Chromis enchrysura). Planktivores dominated fish assemblages, particularly in the upper mesophotic zone, possibly due to a greater availability of zooplankton coming from the colder Equatorial Undercurrent in mesophotic depths of the SPSPA. Turf algae, fleshy macroalgae and scleractinian corals dominated benthic assemblages between 30 and 40 m depth, while bryozoans, black corals and sponges dominated between 40 and 90 m depth. Canonical correspondence analysis explained 74 % of the relationship between environmental characteristics (depth, benthic cover and complexity) and structure of fish assemblages, with depth as the most important independent variable. Juveniles of Bodianus insularis and adults of P. obliquus and C. enchrysura were clearly associated with branching black corals (Tanacetipathes spp.), suggesting that black corals play key ecological roles in lower mesophotic reefs of the SPSPA. Results from this study add to the global database about mesophotic reef ecosystems (MREs) and provide a baseline for future evaluations of possible anthropogenic and natural disturbances on MREs of the SPSPA.     

Cleaning up the biogeography of Labroides dimidiatus using phylogenetics and morphometrics

Cleaner fishes are some of the most conspicuous organisms on coral reefs due to their behaviour and prominent body pattern, consisting of a lateral stripe and blue/yellow colouration. All obligate cleaner fishes share this body stripe pattern, which is an important signal for attracting client fishes. However, variability in the cleaning signal of the cleaner fish Labroides dimidiatus has been documented across its range. Here, we investigate the geographic distribution of cleaner signal polymorphisms in L. dimidiatus and contrast this to phylogeographic variation in mitochondrial (mt) DNA. We used samples from 12 sites for genetic analyses, encompassing much of L. dimidiatus’ range from the Red Sea to Fiji. We obtained morphometric measures of the cleaner signal body stripe width from individuals among six of the sites and qualitatively grouped tail stripe shape. mtDNA control region sequences were used for phylogenetic and population genetic analyses. We found that body stripe width was significantly correlated with tail stripe shape and geographical location, with Indian Ocean populations differing in morphology from western Pacific populations. L. dimidiatus haplotypes formed two reciprocally monophyletic clades, although in contrast to morphology, Japanese cleaner fish fell within the same clade as Indian Ocean cleaner fish and both clade types were sympatric in Papua New Guinea. An additional novel finding of our research was that the inclusion of two closely related cleaner fish species, Labroides pectoralis and Labroides bicolor, in the phylogenetic analysis rendered L. dimidiatus polyphyletic. Overall, the findings suggest the diversity within L. dimidiatus is underestimated.     

Mesophotic depths as refuge areas for fishery-targeted species on coral reefs

Coral reefs are subjected to unprecedented levels of disturbance with population growth and climate change combining to reduce standing coral cover and stocks of reef fishes. Most of the damage is concentrated in shallow waters (<30 m deep) where humans can comfortably operate and where physical disturbances are most disruptive to marine organisms. Yet coral reefs can extend to depths exceeding 100 m, potentially offering refuge from the threats facing shallower reefs. We deployed baited remote underwater stereo-video systems (stereo-BRUVs) at depths of 10–90 m around the southern Mariana Islands to investigate whether fish species targeted by fishing in the shallows may be accruing benefits from being at depth. We show that biomass, abundance and species richness of fishery-targeted species increased from shallow reef areas to a depth of 60 m, whereas at greater depths, a lack of live coral habitat corresponded to lower numbers of fish. The majority of targeted species were found to have distributions that ranged from shallow depths (10 m) to depths of at least 70 m, emphasising that habitat, not depth, is the limiting factor in their vertical distribution. While the gradient of abundance and biomass versus depth was steepest for predatory species, the first species usually targeted by fishing, we also found that fishery-targeted herbivores prevailed in similar biomass and species richness to 60 m. Compared to shallow marine protected areas, there was clearly greater biomass of fishery-targeted species accrued in mesophotic depths. Particularly some species typically harvested by depth-limited fishing methods (e.g., spearfishing), such as the endangered humphead wrasse Cheilinus undulatus, were found in greater abundance on deeper reefs. We conclude that mesophotic depths provide essential fish habitat and refuge for fishery-targeted species, representing crucial zones for fishery management and research into the resilience of disturbed coral reef ecosystems.