Cross‐amplification of 10 new isolated polymorphic microsatellite loci for red mullet (Mullus barbatus) in striped red mullet (Mullus surmuletus)

Ten polymorphic dinucleotide microsatellite loci were isolated and characterized for the red mullet (Mullus barbatus). Allele variability was tested on both the red mullet and its congener the striped red mullet (Mullus surmuletus). Characterization of 30 individuals of both species from the western Mediterranean showed moderate to high allelic diversity ranging from two to 26 alleles per locus (mean 10.9). Three loci showed departures from Hardy–Weinberg proportions. No evidence of significant association between genotypes at pairs of loci was observed. These polymorphic loci could be suitable for population genetic assessments of both species.     

Phylogenetic relationships among four species and a sub-species of Mullidae (Actinopterygii; Perciformes) based on mitochondrial cytochrome B, 12S rRNA and cytochrome oxidase II genes

DNA sequence comparisons of three mitochondrial DNA genes were used to reveal phylogenetic relationships among four species and a sub-species of Mullidae family. This is the first report using mitochondrial DNA sequence data to infer intraspecific relationship among different populations of Mullus barbatus and Mullus surmuletus; phylogenetic relationships between M. barbatus and its sub-species; M. barbatus ponticus. Cytochrome b, 12S ribosomal RNA, and cytochrome oxidase II regions of 242 individuals belonging to species M. barbatus, M. surmuletus, Upeneus moluccensis, Upeneus pori and sub-species M. barbatus ponticus were sequenced and phylogenetic trees were constructed using four different algorithms. The phylogenetic trees constructed support the existing taxonomical data of two mullid genera (Mullus, Upeneus). Molecular data shows no significant difference between same species of different geographical populations. The results suggest that the molecular difference is not large enough between M. barbatus and M. barbatus ponticus to consider them as sub-species.     

First Microsatellite Loci of Red Mullet (<Emphasis Type="Italic">Mullus barbatus</Emphasis>) and Their Application to Genetic Structure Analysis of Adriatic Shared Stock

In order to study the genetic structure of the Adriatic shared stock of red mullet (Mullus barbatus), we developed a set of dinucleotide microsatellite markers. A dinucleotide-enriched genomic library was obtained, and 6 polymorphic dinucleotide loci were successfully optimized. The markers showed high expected heterozygosity (from 0.68 to 0.92) and allele number (from 12 to 33); thus they appear to be suitable for detecting genetic differences in the population of red mullet. Four Adriatic samples were subsequently analyzed for microsatellite variation, and the results showed subtle but statistically significant genetic differentiation, indicating that the Adriatic red mullet may group into local, genetically isolated populations. No correlation between geographic distance and genetic differentiation was observed. In addition, the evidence of recent bottlenecks in the Adriatic samples indicates that the observed population subdivision might reflect random local allelic variations, generated by reproductive success, survival rates, or fishing pressure.     

Aggregation dynamics and foraging behaviour of striped red mullet Mullus surmuletus in the western Mediterranean Sea

Habitat‐related heterogeneity of striped red mullet Mullus surmuletus heterospecific foraging assemblages was examined off the coast of Spain. Video‐based focal‐follows conducted on 122 M. surmuletus assemblages (446 total individuals) revealed an array of attendant species (n = 7) with composition linked to benthic habitat complexity; bare sandy substrata were characterized by homospecific groups of M. surmuletus, while habitats with rock and vegetation attracted a variety of scrounging labrids and sparids. Although the nature of the relationship between M. surmuletus and attendants requires further exploration, the present study indicates that substratum composition can be a driving factor explaining the dynamics of this heterospecific assemblage.     

How an invasive alga species (<Emphasis Type="Italic">Caulerpa taxifolia</Emphasis>) induces changes in foraging strategies of the benthivorous fish <Emphasis Type="Italic">Mullus surmuletus</Emphasis> in coastal Mediterranean ecosystems

The foraging behaviour variability of three striped red mullet (Mullus surmuletus L. 1758) populations with respect to the vegetation cover was examined along the French Mediterranean coasts. We tested both the carrying capacity of different habitats and the hypothesis of a food segregation by the invasive alga Caulerpa taxifolia (Vahl) C. Agardh, on the functioning of benthic fish populations. The results indicated a significant site effect on M. surmuletus foraging behaviour and movements. The vegetation cover may play an important role in modifying the striped red mullet cost of foraging (sampling effort) and thus its strategy of prey capture. As long as the cover of marine phanerogams and/or macrophyte algae (Caulerpaceae) increases, the foraging budget and the distances covered in search of prey decrease significantly. Similarly, the striped red mullet increases the prospected sample periods to augment its foraging success, swimming above the bottom. These changes are related to the development of a dense superficial network of plagiotropic rhizomes and stolons, and to the reduction of space between fronds that limits the accessibility to resources and may increase intra-specific food competition. The role of C. taxifolia does not differ from that of other marine phanerogams but induces significant changes in the structure of Mullidae populations at the local level. The rapid expansion of Caulerpaceae in the Mediterranean Sea could constitute a real threat for the balance of the marine coastal biodiversity and the ecology of M. surmuletus, which is considered a flagship species for coastal Mediterranean demersal fisheries.     

Difference in diet and size‐related trophic level in two sympatric fish species,the red mullets Mullus barbatus and Mullus surmuletus,in the Gulf of Lions (north‐west Mediterranean Sea)

Stable‐isotope ratios (δ¹⁵N and δ¹³C) and diet of the red mullets Mullus barbatus and Mullus surmuletus were analysed in two zones differently subjected to the Rhône River runoff in the Gulf of Lions (north‐west Mediterranean sea) in May and October 2004. δ¹⁵N and δ¹³C increased significantly with fish size in M. barbatus in both zones and seasons, whereas no significant trend was evidenced in M. surmuletus. A clear size‐related shift in diet was observed in M. barbatus, with an increase in polychaete and shrimp consumption with size and a decrease in small crustacean consumption. In M. surmuletus, a diet shift was observed only between medium and large individuals. Large M. surmuletus preyed on shrimps, polychaetes, bivalves, ophiurids and amphipods, and ingested prey of lower mean mass than M. barbatus of similar size. Difference in size‐related increase in δ¹⁵N between the two mullid fish species were related to difference of trophic level of their main prey. Sub‐surface deposit‐feeding polychaetes, carnivorous polychaetes, shrimps and brachyurans presented higher δ¹⁵N values than bivalves, small crustaceans and ophiurids. The lower δ¹³C values observed in M. barbatus compared to M. surmuletus were related to a higher consumption of sub‐surface polychaetes in the former species. Significantly, lower δ¹³C were recorded in fishes collected off the Rhône River, particularly in spring, suggesting an influence of river inputs as a source of particular organic matter for mullids in this zone after the flooding season. Thus, these closely related sympatric fish species displayed diet divergences that were reflected in their stable isotopic signatures.     

Oocytes development,maturity classification,maturity size and spawning season of the red mullet (Mullus barbatus barbatus Linnaeus, 1758)

The present study was designed to identify maturity of female red mullet, Mullus barbatus barbatus, using microscopic criteria, offering an accurate determination of the ovarian cycle and estimation of maturity size. Histological analysis of 499 females from 13 monthly samplings was applied (Thermaikos Gulf, N. Aegean Sea, Greece). Oocyte development was divided into five stages, with a mean oocyte size range from 56 to 363 μm in diameter. Ovarian maturity was defined by the maturation stage of the most advanced oocytes and divided into four phases. By using the monthly changes in the percentage frequency of ovarian phases, the ovarian cycle of M. barbatus was divided into three periods: a long period of early oogenesis (November to February), a short period of vitellogenesis (February to April) and a spawning period from April through June with peak activity in May. Both gonadosomatic (up to 7.5%) and hepatosomatic indices (up to 2%) can be used together to predict the spawning period. Red mullet should be classified as a multiple spawning species. Females reached 50% maturity (Lm₅₀) at 11.4 cm and 95% maturity (Lm₉₅) at 15.5 cm TL. The Lm₅₀ values for female red mullet populations across the Mediterranean Sea tended to show differences ranging from 11 to 14.4 cm FL or 11.4–14 cm TL. The current European Union fisheries management plan stipulates a minimum landing size (MLS) of 11 cm TL for Mullidae spp. fisheries in the Mediterranean. This MLS is at the lower part of the Lm₅₀ range, thus allowing an extensive removal of immature juveniles. A revision of this MLS value is recommended to help ensure that red mullet stocks remain within safe biological limits.     

Gut of goatfishes,a heterogeneous biotope for intestinal mesoparasites: variations in pyloric caeca number and growth models of colonizable digestive surface area

Fish organs constitute biotopes for various parasites. In the case of gills or digestive tracts, hosts of different ages and sizes lead to a heterogeneity in these biotopes at the fish population level. The digestive tracts of goatfishes Mullus surmuletus and M. barbatus harbour several species of helminths that colonize the pyloric caeca and intestine. The structural heterogeneity of this biotope is analysed by study of variations in caeca number and colonizable surface area. Caeca number varies between host species and among individuals of the same population. Colonizable intestinal surface area also varies between goatfish species and increases with fish size, but differently according to the intestinal parts considered. These variations were studied, using image-processing techniques, and analysed quantitatively. Theoretical growth models are proposed and some consequences for parasite ecology discussed.     

Morphological variability of the shape of striped red mullet Mullus surmuletus in relation to stock discrimination between the Bay of Biscay and the eastern English Channel

Truss analysis and length measurements were made on 168 striped red mullet Mullus surmuletus. Multivariate statistical analyses with principal component analysis and partial redundancy analysis (pRDA) were used on these measurements to evaluate the influence of maturity, sex and geographical area distribution on body shape. Truss measurements were important to quantify and discriminate changing body shape, presumably due to changing environmental conditions. Sexual dimorphism was not observed and juveniles could be distinguished from adults based on their body shape. More importantly, M. surmuletus occurring in different geographical areas could be differentiated using this method. Based on pRDA, a significant difference of head morphological dimensions was observed between populations occurring in the eastern English Channel and those occurring in the Bay of Biscay, suggesting that fish from these areas could represent two subpopulations.     

Sperm motility of the marine teleosts Boops boops, Diplodus sargus, Mullus barbatus and Trachurus mediterraneus

The spermatozoa of Boops boops, Diplodus sargus, Mullus barbatus, and Trachurus mediterraneus were motile in sea water, and in electrolyte solutions (NaCl) and non-electrolyte solutions (glucose) with an osmolality of 600–1000 mosmol kg^?1. Their mean motility rate 10 s after initiation was about 80%, while about 10% of the motile spermatozoa moved non-linearly, 45% linearly, and 45% circularly. The average path swimming velocity was significantly higher in M. barbatus (about 90 μm s^?1) than in the other species (70 μm s^?1). The number of motile spermatozoa decreased to 0% within 50 s after initiation of motility in T. mediterraneus, within 90 s in M. barbatus . In B. boops and D. sargus about 90% of the spermatozoa stopped movement during the first 90 s of the motility period, while the rest remained motile for 2–3 h. Motility of B. boops and D. sargus spermatozoa was reversibly suppressed in the seminal plasma, and in electrolyte and non-electrolyte solutions of 100–200 mosmol kg^?1. The trigger for motility activation was hyperosmolality (700–1000 mosmol kg^?1). Motility of M. barbatus and T. mediterraneus sperm was only partly suppressed in the seminal plasma since freshly collected semen contained about 25–50% locally motile spermatozoa. When sperm was activated immediately after collection with electrolyte and non-electrolyte solutions of 700–1000 mosmol kg^?1 spermatozoa moved progressively. The motility of those spermatozoa which had not yet been motile after collection was completely and reversibly suppressed in M. barbatus at osmolalities of 1200 mosmol kg^?1, and at osmolalities of 100–200 mosmol kg^?1 in T. mediterraneus . Therefore two triggers were necessary for initiation of motility. The nature of the first trigger was uncertain, the second trigger was a switch to hypoosmolality in M. barbatus and to hyperosmolality in T. mediterraneus . The sperm organisation of B. boops, D. sargus, M. barbatus and T. mediterraneus revealed species-specific parameters which could not be related with the sperm motility behaviour.     

Spatial graphs highlight how multi-generational dispersal shapes landscape genetic patterns

Current approaches that compare spatial genetic structure of a given species and the dispersal of its mobile phase can detect a mismatch between both patterns mainly due to processes acting at different temporal scales. Genetic structure result from gene flow and other evolutionary and demographic processes over many generations, while dispersal predicted from the mobile phase often represents solely one generation on a single time-step. In this study, we present a spatial graph approach to landscape genetics that extends connectivity networks with a stepping-stone model to represent dispersal between suitable habitat patches over multiple generations. We illustrate the approach with the case of the striped red mullet Mullus surmuletus in the Mediterranean Sea. The genetic connectivity of M. surmuletus was not correlate with the estimated dispersal probability over one generation, but with the stepping-stone estimate of larval dispersal, revealing the temporal scale of connectivity across the Mediterranean Sea. Our results highlight the importance of considering multiple generations and different time scales when relating demographic and genetic connectivity. The spatial graph of genetic distances further untangles intra-population genetic structure revealing the Siculo-Tunisian Strait as an important corridor rather than a barrier for gene flow between the Western- and Eastern Mediterranean basins, and identifying Mediterranean islands as important stepping-stones for gene flow between continental populations. Our approach can be easily extended to other systems and environments.     

Factors influencing dolphin depredation in coastal fisheries of the northern Aegean Sea: Implications on defining mitigation measures

Dolphin interactions with coastal fisheries are of major concern, reportedly leading to gear damage, which increases the cost of coastal fishing globally and in the Mediterranean Sea. The aim of this study was to determine the effect that gear, target species, mesh size, depth, soaking duration, fishing area, and season have on net depredation frequency and to offer insights on possible mitigation solutions. From November 2013 to February 2016 we monitored 107 active coastal fishers in 22 ports of the northern Aegean Sea coastline, identified the main target species of the fishery and recorded the damages on gill nets and trammel nets caused by dolphins, mainly the common bottlenose dolphin (Tursiops truncatus). Quasi-binomial generalized linear models were used to determine the relationship between the examined factors and depredation frequency. The analysis revealed that the gears mostly depredated were gill nets and trammel nets with small mesh sizes, mainly targeting surmullet (Mullus surmuletus), red mullet (Mullus barbatus), common sole (Solea solea), European hake (Merluccius merluccius), and caramote prawn (Melicertus kerathurus). The probability of depredation was also significantly dependent on the fishing area.     

Morphological variation between red mullet populations in Greece

Morphometric variation in 15 characters of the red mullet Mullus barbatus samples from seven Greek localities was examined using multivariate analysis. ANOVA, stepwise discriminant analysis and cluster analysis revealed a rather high morphological variability between the populations studied. The discriminant analysis revealed that about 80% of the examined fish could be correctly classified into the seven areas. This variability could be explained in terms of genetic structuring of the populations and/or environmental conditions prevailing in each geographic area in combination with fish migration and egg/larvae transportation from one area to another. The results suggest that multivariate analysis when combined with other important biological parameters of red mullet may have important implications for the management of the Greek demersal fish resources.     

Age and Growth of the Red Mullet Mullus barbatus in the North Aegean Sea

Journal of Ichthyology - The age and growth of Mullus barbatus L. 1758, sampled around the Gökçeada Island from September 2014 to June 2015 was determined. A lifetime of 0–VII years...     

Sorting grid trials to improve size selectivity of red mullet (Mullus barbatus) and annular sea bream (Diplodus annularis) in Turkish bottom trawl fishery

Sorting grids with two different bar spacings (12 mm and 14 mm) were tested to improve size selectivity of the commercially important fish species, red mullet (Mullus barbatus) and annular sea bream (Diplodus annularis), in Turkish bottom trawl fishery. Fishing trials were carried out with R/V ‘Egesüf’ between April and May 2003 in Izmir Bay, Aegean Sea, using a traditional bottom trawl. Selectivity data were collected by the top cover method and analyzed by means of a logistic equation with the maximum likelihood method. Selectivity parameters for individual hauls were obtained with the software program cc 2000 . Mean selectivity was also estimated and compared using the EC Model software that takes between‐haul variations into account. The codend catch size as an additional explanatory variable was used in the comparison. The Kolmogorov–Smirnov (K–S) test was also applied to detect differences between length‐frequency distributions in the upper and lower bags of the 12 and 14 mm bar spacings for red mullet and annular sea bream. The mean L₅₀ values of red mullet were 8.7 and 10.0 cm with the 12 and 14 mm grids, respectively. The L₅₀ value of 14 mm was comparable with the L₅₀ value of the codend mesh size; however, the 12 mm value was rather low. The mean L₅₀ values of 12 and 14 mm bar spacings were 8.8 and 10.4 cm for annular sea bream, respectively; the value of 14 mm bar spacing was very close to 50% size at sexual maturity of the species (10.5 cm). The K–S test indicated length distributions of red mullet and annular sea bream in the 12 and 14 mm upper and lower bags as significantly different (P < 0.05). These results show that improving the size selectivity in a multi‐species fishery using a single selective device is rather difficult. However, higher size selectivity can be obtained when considering the minimum landing size or the 50% sexual maturity size for a given species.     

Environmental influences on the spatial distribution of European hake (Merluccius merluccius) and red mullet (Mullus barbatus) in the Mediterranean

Generalised additive models (GAMs) were used to test the hypotheses that red mullet (Mullus barbatus) and hake (Merluccius merluccius) abundances are related to the bathymetry, spatial location and the temperature-based variability of the NE Mediterranean. Data were collected during a 2-year period (1996–1997) of quasi-synoptic sampling using demersal trawl surveys in the northern Aegean Sea. The results of this study supported these hypotheses. It was found that geographic position and sea bottom characteristics influence the temporal distribution patterns of red mullet and hake. Data are presented that reveal species-specific aggregation patterns and an important habitat utilisation of the area. The modelled species’ abundances showed a strongly non-linear dependence on the explanatory covariates. This study, although not directly dealing with spawning, also provided evidence that red mullet and hake demonstrate environmental and bathymetric preferences in the months posterior to spawning. Mean red mullet abundance was consistently highest in areas with warmer bottom waters in the east and west than in the centre, these areas having shallower depths (35–60 m) and bottom temperatures around 19°C. The red mullet appeared to avoid the cold bottom waters (<16°C) of the deeper regions. Results indicated that areas with the highest hake abundances were located in waters of ∼160 m having bottom temperature of ∼16°C and avoided the shallower waters (<70 m) regardless of their bottom temperature. The present findings also suggested that colder bottom waters at all depths were associated with lower-than-average hake abundance. Red mullet and hake preferences for areas with specific bathymetric and ocean environmental conditions are believed to be linked to the oceanography of the NE Mediterranean ecosystem.     

Behavioural response of Mullus surmuletus to habitat modification by the invasive macroalga Caulerpa taxifolia

Densities of red mullet Mullus surmuletus have declined since 1996 at Cap Martin, Alpes‐Maritimes, France, following colonization by Caulerpa taxifolia . Foraging M. surmuletus were rarely observed over C. taxifolia or the seagrass Posidonia oceanica , but were observed over bare sand and endemic macroalgae. Within colonized sites (where the mean cover of C. taxifolia was 30–100%), fish were concentrated on areas with low cover of C. taxifolia (<21%). A difference in the frequency of observations of foraging groups (17%) was observed between sites with and without C. taxifolia . The observed patterns were probably due to a physical barrier to foraging that is presented by meadows of C. taxifolia . Changes to habitat structure that occur when the substratum is colonized by C. taxifolia influence the accessibility to benthic food resources, and consequently the foraging activities of fish that feed on benthic invertebrates. The decline in density of M. surmuletus at Cap Martin could be the result of fish emigration from colonized sites (unfavourable to the species) to uncolonized sites.     

Modern state of the blunt-snouted mullet <Emphasis Type="Italic">Mullus barbatus ponticus</Emphasis> population dwelling in the coastal zone near Sevastopol

The existence of the independent Sevastopol group of the blunt-snouted mullet Mullus barbatus ponticus described by Danilevskii (1939) that differs from the Caucasian one by a smaller length of the head, barbels, and pectoral fins has been supported. It was found that, during the maximum anthropogenic impact on the Black Sea ecosystem, the state of M. barbatus ponticus deteriorates, which was manifested in a decrease in the rate of growth, a high value of liver index, and a decrease in fatness. From 2003 to 2007, a tendency for an improvement in the state of M. barbatus ponticus of the Sevastopol school was revealed that manifested itself in an increase in the numbers of fish arriving to the coastal zone for spawning and feeding and in the restoration of the age composition typical of this species.