Isolation and characterization of polymorphic microsatellite markers for peacock wrasse (Symphodus tinca)

Eight polymorphic microsatellite loci were isolated and characterized for the peacock wrasse (Symphodus tinca), a labrid fish inhabiting the Mediterranean and Black seas. Characterization of 35 individuals from the western Mediterranean indicated a relatively high allelic diversity (mean = 12.4, range 9–17), and observed heterozygosity ranging from 0.65 to 0.91. We found no evidence of linkage disequilibrium between pairs of loci. Two loci showed significant departure from Hardy–Weinberg equilibrium. These polymorphic markers can be useful in most basic population genetic applications.     

Acquisition of a defended resource: a benefit of group foraging for the neotropical wrasse,Thalassoma lucasanum

Synopsis The neotropical Pacific wrasse, Thalassoma lucasanum, forms groups of 30 to 300 individuals that overwhelm the defense of embryos by the Pacific sergeant major, Abudefduf troschelii, and thereby gain access to a food resource from which solitary individuals are effectively excluded by paternal defense. The duration of feeding by the wrasse groups and the fraction of embryos eaten in a nest are positively correlated with group size. The benefit of group foraging is probably derived from a reduction in the frequency with which group members experience attack by defending sergeant majors, as reflected in the positive correlation of group size with feeding duration. Large foraging groups only form when sergeant major embryos are present in nests, indicating that this is a primary benefit of participation in these groups for this wrasse. In contrast, in the Caribbean large foraging groups of the congeneric bluehead wrasse, T. bifasciatum, fail to form despite the presence of defended embryos of the Caribbean sergeant major, A. saxatilis. This might be due to the relatively low population densities of the Caribbean wrasse in comparison to those of the Pacific wrasse or to the relatively low densities of Caribbean sergeant major nests containing embryos at any time.     

Whale sharks Rhincodon typus get cleaned by the blue-streak cleaner wrasse Labroides dimidiatus and the moon wrasse Thalassoma lunare in the Philippines

Cleaning interactions are essential for healthy marine ecosystem communities. This study reports the first documentation of the whale shark Rhincodon typus cleaning behaviour in the Indo-West Pacific by two wrasse species, the blue-streak cleaner wrasse Labroides dimidiatus and the moon wrasse Thalassoma lunare in Cebu, Philippines. This study documented 36 cleaning interactions with 14 individual whale sharks. The cleaning interactions appear opportunistic rather than targeted by the sharks, unlike that observed in other species of elasmobranchs. Further work should focus on understanding the drivers of these unique cleaning interactions.     

The occurrence and distribution of peptide- or 5-HT-containing nerves in the swimbladder of four different species of teleosts (Gadus morhua,Ctenolabrus rupestris,Anguilla anguilla,Salmo gairdneri)

Summary The innervation of the swimbladder in four different teleost species has been studied by the use of immunohistochemical methods. The teleosts examined belong to two different groups regarding their swimbladder morphology: physoclists (the cod, Gadus morhua and the goldsinny wrasse, Ctenolabrus rupestris) and physostomes (the eel, Anguilla anguilla and the rainbow trout, Salmo gairdneri). Vasoactive intestinal polypeptide-like immunoreactivity was demonstrated in nerves of the swimbladder walls of all four species, and in the gas glands of the cod and the goldsinny wrasse. Substance P-like immunoreactivity was shown in swimbladders of the cod, eel and rainbow trout but not the goldsinny wrasse. Immunoreactivity to met-enkephalin antiserum was revealed in the swimbladder walls of the eel and the goldsinny wrasse, while neurotensin-like immunoreactivity was present in the goldsinny wrasse and rainbow trout swimbladders. Neurotensin-like immunoreactivity was also seen in the gas gland of the goldsinny wrasse. 5-Hydroxytryptamine immunoreactivity was found in endocrine cells in the pneumatic duct of the eel and in the swimbladder walls of the goldsinny wrasse and the rainbow trout. In conclusion, all teleosts examined showed a very close resemblance in the peptidergic/tryptaminergic innervation of the swimbladder to that of the gut, inasmuch as the immunoreactivity present in the swimbladders always occurred in the gut of the same species.     

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.     

Post-settlement emigration affects mortality estimates for two Bahamian wrasses

Understanding the dynamics of open marine populations is inherently complex, and this complexity has led to decades of debate regarding the relative importance of pre- versus post-settlement processes in structuring these populations. Movement between patches may be an important modifier of patterns established at settlement, yet local immigration and emigration have received less attention than other demographic rates. I examined loss rates from tagged populations of juvenile wrasses (yellowhead wrasse Halichoeres garnoti and bluehead wrasse Thalassoma bifasciatum) at two sites in the Bahamas. Assuming that all losses were due solely to mortality would have significantly underestimated survivorship of yellowhead wrasse by 29% and bluehead wrasse by 14%. On average, per capita mortality and emigration rates were higher for yellowhead than bluehead wrasse, but neither demographic rate differed between sites for either species. With respect to within-species density, bluehead wrasse mortality was density-dependent at the patch reef site, but mortality rates of yellowhead wrasse were consistently density-independent. Evaluating the effects of between-species density, yellowhead wrasse mortality increased with a decrease in bluehead wrasse density, but this effect was limited to the patch reef site. Emigration rates were not a function of either within-species or between-species density, but instead varied inversely with isolation distance. Numerous previous studies of coral-reef fish, conducted on patch reefs separated by only a few meters of sand and often using untagged fish, may have confounded losses due to emigration with those due to mortality. A better understanding of the factors affecting emigration in marine fishes is important to their effective management using spatial tools such as marine protected areas.     

Cleaner wrasse influence habitat selection of young damselfish

The presence of bluestreak cleaner wrasse, Labroides dimidiatus, on coral reefs increases total abundance and biodiversity of reef fishes. The mechanism(s) that cause such shifts in population structure are unclear, but it is possible that young fish preferentially settle into microhabitats where cleaner wrasse are present. As a first step to investigate this possibility, we conducted aquarium experiments to examine whether settlement-stage and young juveniles of ambon damselfish, Pomacentrus amboinensis, selected a microhabitat near a cleaner wrasse (adult or juvenile). Both settlement-stage (0 d post-settlement) and juvenile (~5 weeks post-settlement) fish spent a greater proportion of time in a microhabitat adjacent to L. dimidiatus than in one next to a control fish (a non-cleaner wrasse, Halichoeres melanurus) or one where no fish was present. This suggests that cleaner wrasse may serve as a positive cue during microhabitat selection. We also conducted focal observations of cleaner wrasse and counts of nearby damselfishes (1 m radius) to examine whether newly settled fish obtained direct benefits, in the form of cleaning services, from being near a cleaner wrasse. Although abundant, newly settled recruits (<20 mm total length) were rarely (2 %) observed being cleaned in 20 min observations compared with larger damselfishes (58 %). Individual damselfish that were cleaned were significantly larger than the median size of the surrounding nearby non-cleaned conspecifics; this was consistent across four species. The selection by settlement-stage fish of a microhabitat adjacent to cleaner wrasse in the laboratory, despite only being rarely cleaned in the natural environment, suggests that even rare cleaning events and/or indirect benefits may drive their settlement choices. This behaviour may also explain the decreased abundance of young fishes on reefs from which cleaner wrasse had been experimentally removed. This study reinforces the potentially important role of mutualism during the processes of settlement and recruitment of young reef fishes.     

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.     

Laser chasing behaviour of wild fishes exploited as a tool to compare space use between size,sex and species

The spatial extent of animal movement is a key consideration when designing conservation measures, such as marine protected areas. Methods to assess territory size in the marine environment, however, are labour intensive and/or expensive. Here, we explore a novel method to investigate the spatial ecology of territorial fishes by examining their reactions to an artificial light stimulus. During benthic towed video surveys conducted in Lyme Bay, southwest England, several species of wrasse (Labridae) have frequently been observed pursuing a laser projected onto the seabed. While the motivation behind ‘laser‐chasing’ is unclear, we quantified the spatial aspects of this behaviour by comparing chase distance and chase likelihood between and within species, to determine the potential utility of this method for investigating space use and aggression in wild fishes. Cuckoo wrasse (Labrus mixtus) were significantly more likely to display agonistic behaviour towards the laser than Goldsinny wrasse (Ctenolabrus rupestris). Goldsinny wrasse displayed a positive relationship between total length and chase‐distance, but not Cuckoo wrasse. The observed species differences may relate to behavioural factors affecting the motivation behind ‘laser‐chasing’, which is discussed. Chases by the cuckoo wrasse were significantly longer than those by Goldsinny wrasse, and these chase distances were used to estimate theoretical territory sizes for each species. To our knowledge, this is the first study to explore the spatial aspects of the reactions to an artificial stimulus by wild fishes. The potential to develop the method to directly investigate aspects of territoriality and aggression in wild fishes is discussed, including necessary further refinements and testing. Wild wrasses are increasingly exploited in Europe to provide cleaner fish for salmonid aquaculture, and we encourage the development of methods to inform spatial conservation measures for these ubiquitous and iconic species.     

Demographic history has shaped the strongly differentiated corkwing wrasse populations in Northern Europe

Understanding the biological processes involved in genetic differentiation and divergence between populations within species is a pivotal aim in evolutionary biology. One particular phenomenon that requires clarification is the maintenance of genetic barriers despite the high potential for gene flow in the marine environment. Such patterns have been attributed to limited dispersal or local adaptation, and to a lesser extent to the demographic history of the species. The corkwing wrasse (Symphodus melops) is an example of a marine fish species where regions of particular strong divergence are observed. One such genetic break occurred at a surprisingly small spatial scale (F_ST ~0.1), over a short coastline (<60 km) in the North Sea‐Skagerrak transition area in southwestern Norway. Here, we investigate the observed divergence and purported reproductive isolation using genome resequencing. Our results suggest that historical events during the post‐glacial recolonization route can explain the present population structure of the corkwing wrasse in the northeast Atlantic. While the divergence across the break is strong, we detected ongoing gene flow between populations over the break suggesting recent contact or negative selection against hybrids. Moreover, we found few outlier loci and no clear genomic regions potentially being under selection. We concluded that neutral processes and random genetic drift e.g., due to founder events during colonization have shaped the population structure in this species in Northern Europe. Our findings underline the need to take into account the demographic process in studies of divergence processes.     

Importance of sandy bottoms in coral reefs to the oscillation of daily rhythms in the tropical wrasse Halichoeres trimaculatus

Most wrasse species swim during the day and bury themselves in the sandy bottoms of shallow reefs at night. This study aimed to evaluate the importance of sandy bottoms to the day-active/night-inactive rhythmicity of the tropical wrasse Halichoeres trimaculatus. Actogram analysis revealed that fish were active during the photophase and inactive during the scotophase in aquariums with both sandy and bare bottoms. When fish were kept in aquariums with bare bottoms, rhythmicity was maintained under constant dark conditions (DD) but became obscured under constant light conditions (LL), suggesting that a day-active/night-inactive rhythmicity is regulated by the circadian system. Robust fluctuations in Period1 (wPer1) and Period2 (wPer2) expression were observed in the pectoral fin tissue under light–dark conditions (LD). Similar fluctuations in wPer1 expression persisted under DD. When fish were kept under LD conditions for 7 days and then DD for 20 days, the emergence of fish from the sandy bottom was delayed gradually. At the same time, the peak time of wPer1 expression under DD was retarded from 06:00 to 10:00. Although wPer2 expression was dampened under DD, it increased after exposing fish to light. These results suggest that wPer1 and wPer2 are differentially involved in the day-active/night-inactive rhythmicity, and that blocking light with a sandy bed at night and exposing fish to light during emergence in the morning play important roles in maintaining consistent activities in wrasse species.     

Courtship and spawning sounds in bird wrasse Gomphosus varius and saddle wrasse Thalassoma duperrey

Acoustic signals from the bird wrasse Gomphosus varius and saddle wrasse Thalassoma duperrey were recorded on coral reefs in Hawaii. Terminal phase males in both species emit two types of pulse trains (type I and type II). Type I pulses were produced during spawning and courtship, while type II pulses were associated only with courtship behaviours. Gomphosus varius type I pulses were of lower frequency than T. duperrey type I pulses (271 v. 840 Hz) and were of narrower band. Discriminant function analyses revealed interspecific differences between type I pulse trains and individual pulses of both types. This study is the first documentation of courtship and spawning sounds in sympatric labrids and shows divergence in acoustic signals.     

Spatially correlated recruitment of a marine predator and its prey shapes the large-scale pattern of density-dependent prey mortality

Patterns of predator dispersal can be critical to the dynamics of prey metapopulations. In marine systems, oceanic currents may shape the dispersal of planktonic larvae of both predators and prey, producing spatial correlations in the recruitment of both species and distinctive geographic patterns of prey mortality. I examined the potential for this phenomenon in two fishes, a wrasse and its grouper predator, at a Caribbean island where the near-shore oceanographic regime produces a temporally consistent spatial pattern of fish recruitment. I found that recruitment and adult abundance of groupers were spatially correlated with recruitment of wrasse prey. Furthermore, the local abundance of predators strongly affected the nature of density-dependent prey mortality. At sites with few predators, wrasse mortality was inversely density-dependent, while mortality was positively density-dependent at sites with higher predator densities. This phenomenon could be important to the dynamics of any metacommunity in which physical forces produce correlated dispersal.     

Unloading of homologous recombination factors is required for restoring double‐stranded DNA at damage repair loci

Cells use homology‐dependent DNA repair to mend chromosome breaks and restore broken replication forks, thereby ensuring genome stability and cell survival. DNA break repair via homology‐based mechanisms involves nuclease‐dependent DNA end resection, which generates long tracts of single‐stranded DNA required for checkpoint activation and loading of homologous recombination proteins Rad52/51/55/57. While recruitment of the homologous recombination machinery is well characterized, it is not known how its presence at repair loci is coordinated with downstream re‐synthesis of resected DNA. We show that Rad51 inhibits recruitment of proliferating cell nuclear antigen (PCNA), the platform for assembly of the DNA replication machinery, and that unloading of Rad51 by Srs2 helicase is required for efficient PCNA loading and restoration of resected DNA. As a result, srs2Δ mutants are deficient in DNA repair correlating with extensive DNA processing, but this defect in srs2Δ mutants can be suppressed by inactivation of the resection nuclease Exo1. We propose a model in which during re‐synthesis of resected DNA, the replication machinery must catch up with the preceding processing nucleases, in order to close the single‐stranded gap and terminate further resection.     

Ocean Productivity May Predict Recruitment of the Rainbow Wrasse (Coris julis)

Predicting recruitment fluctuations of fish populations remains the Holy Grail of fisheries science. While previous work has linked recruitment of reef fish to environmental variables including temperature, the demonstration of a robust relationship with productivity remains elusive. Despite decades of research, empirical evidence to support this critical link remains limited. Here we identify a consistent and strong relationship between recruitment of a temperate wrasse Coris julis, from temperate reefs in the mid-Atlantic region, with Chlorophyll, over contrasting scales, across multiple years. Additionally, we find that the correlation between Chlorophyll and recruitment is not simply masking a temperature-recruitment relationship. Understanding the potential mechanisms underlying recruitment variability, particularly as it relates to changing climate and ocean regimes, is a critical first step towards characterizing species’ vulnerability to mismatches between pulsed planktonic production and early pelagic life stages.     

A comparative study of cleaning activity of two reef fishes at Fernando de Noronha Archipelago,tropical West Atlantic

Cleaner fishes are usually classified as obligate or facultative cleaners according to their diet and the extent to which their nutritional requirements in the different ontogenetic stages are gained from cleaning. While obligate cleaners clean throughout their lives and ingest mainly food taken from the clients’ body surface, facultative cleaners clean only as juveniles and have a broader diet. In addition, some facultative cleaners may experience a relatively higher predation risk, and thus rarely interact with piscivorous fishes. Despite these acknowledged differences, there are very few studies that compare cleaning activity of obligate and facultative cleaners within the same area. Cleaning activity of the obligate cleaner goby Elacatinus cf. randalli and the facultative cleaner wrasse Thalassoma noronhanum were comparatively examined at Fernando de Noronha Archipelago, tropical West Atlantic. The client assemblage attended by the two cleaners differed, as the goby attended a slightly greater diversity of species (22), mostly piscivores and zoobenthivores, and the wrasse attended fewer species (19), mostly planktivores. Chromis multilineata was the most common client species of both cleaners, although body size (which is expected to be positively correlated to clients’ ectoparasite load) of C. multilineata individuals attended by the goby was larger than that of the individuals attended by the wrasse. Despite such differences, T. noronhanum showed a surprisingly species-rich client assemblage when compared with other cleaners of the genus Thalassoma. In addition, the frequency and time spent on cleaning interactions, as well as the number of client species attended per 10-min period, was similar for both cleaner species, which indicate that they have important yet complimentary ecological roles in the reef community at Fernando de Noronha Archipelago.     

Attack intensity by two species of territorial damselfish (Pomacentridae) as estimates of competitive overlap with two species of wrasse (Labridae)

The intensity of interspecific territorial defense should be based upon the degree of competitive overlap. We tested this relationship in two territorial Caribbean damselfish (dusky, Stegastes adustus, and longfin, S. diencaeous) with intruders being the bluehead wrasse (Thalassoma bifascatus) and the slippery dick wrasse (Halichoeres bivittatus). Based on food habits, the slippery dick and the bluehead wrasse should have the same degree of competitive overlap to the two damselfish species. We also predicted that the larger slippery dick wrasse intruder would receive more aggression than the smaller bluehead wrasse intruder. Neither damselfish species distinguished between the two wrasses suggesting that they were ecologically equivalent. We also tested size differences within both species of wrasse and found that size had no influence on the aggression in the dusky damselfish. The longfin also did not show a size preference for the bluehead but did prefer to attack the larger slippery dick wrasse. In spite of the similarities between the dusky and the longfin damselfish and the similar food habits of the bluehead and the slippery dick wrasse, our results suggest that using intensity of interspecific territorial defense alone may not be an adequate measure of competitive overlap.     

Epistatic interactions of three loci regulate flowering time under short and long daylengths in a backcross population of rice

The short-day plant rice varies greatly in photoperiod sensitivity (PS) for flowering. The hybrid F₁ from a cross between the day-neutral pure line EM93-1 and the weedy rice accession SS18-2 had stronger PS than SS18-2. Some BC₁ (EM93-1/F₁) segregates were even more sensitive to photoperiod than the F₁, as indicated by later flowering or no flowering after 250 days under a 14-h long daylength. A genome-wide scan identified the quantitative trait loci Se _7.1, Se _7.2 and Se ₈ for PS from the BC₁ population, with all alleles that inhibit flowering derived from SS18-2. These three loci regulate the time of flowering under long daylength through their main effects, and di- and trigenic epistases. Under a 10-h short daylength, the regulation is through Se _7.1 and Se ₈ main effects and digenic epistases involving all three loci. The short daylength not only nullified the main effect of Se _7.2, but also changed its epistatic effects from inhibiting flowering under long daylength to promoting flowering. The epistases indicate that genes underlying the three PS loci work in the same pathway for the control of flowering. Many non-flowered BC₁s were the trigenic heterozygote; this suggests that the three PS loci are also involved in genetic control of critical daylength.