Habitat specific growth rates and condition indices for the sympatric soles Solea solea (Linnaeus, 1758) and Solea senegalensis Kaup 1858, in the Tagus estuary, Portugal, based on otolith daily increments and RNA-DNA ratio

Habitat specific growth rates and condition indices were estimated for Solea solea and Solea senegalensis, in two nursery areas within the Tagus estuary, at the end of the estuarine colonization process, in 2005. While in the uppermost nursery area the two species of sole live in sympatry, in the lower nursery only S. senegalensis is present. Daily increments of left lapillar otoliths were used to estimate age (in days) and determine growth rates (mm per day). Condition indices were assessed through RNA‐DNA ratio in muscle samples. Growth rates were higher for S. senegalensis (0.970and 1.180 mm per day in nursery A and B, respectively) than for S. solea (0.767 mm per day in nursery A). Growth rates of S. senegalensis from the uppermost nursery area were lower when compared to those obtained for the other nursery. The RNA/DNA condition index followed the general trend given by the growth rate estimates, i.e. values were higher for S. senegalensis than for S. solea. However, no significant differences were detected in S. senegalensis from the two nurseries. Larger variations in salinity (10‰ amplitude in the uppermost nursery vs 0.2‰ in the lower nursery) and highest pollution loads may be important factors lowering the habitat quality of the uppermost nursery in comparison to the lower nursery. The use of growth rate estimates based on otolith readings and the RNA/DNA index as tools for habitat quality assessment was discussed.     

Development and characterization of microsatellites from Senegal sole (Solea senegalensis)

To assist in the domestication of the sole fish Solea senegalensis, 15 microsatellites were developed from its genome (13 di‐ and two trinucleotide). These loci, along with three from S. solea that cross‐amplified, were characterized on a sample of 20 individuals of S. senegalensis from a hatchery stock. All loci were polymorphic displaying from two to 15 alleles/locus (mean 6.5). The results obtained from the hatchery population screening indicate that they might be useful for studies of genetic structuring and/or kinship analysis in S. senegalensis.     

Estuarine colonization, population structure and nursery functioning for 0-group sea bass (Dicentrarchus labrax), flounder (Platichthys flesus) and sole (Solea solea) in a mesotidal temperate estuary

The function of the Mondego estuary as a fish nursery habitat was investigated from June 2003 to June 2004 by comparing the timing of estuarine colonization with juveniles of sea bass Dicentrarchus labrax, flounder Platichthys flesus, and sole Solea solea, while also analysing their population structure, growth and diet composition. Differences in the onset of estuarine colonization were observed, since sole juveniles were the first to enter the estuary (in January), followed by flounder in April and sea bass in June. The estuarine population of these species consisted of several age‐groups, although the majority of individuals belonged to age‐groups 0 and 1. The growth rates determined for 0‐group fish were within the range of those reported for other European estuarine systems. Some differences were also recognized regarding the timing of estuarine colonization and the length of the growing season. Diet of 0‐group sea bass consisted mainly of Crustacea, Polychaeta and Mollusca. Flounder juveniles fed chiefly on Amphipoda (especially Corophium spp.), with Polychaeta, Isopoda and Decapoda also being common prey. The diet of 0‐group sole was dominated by Polychaeta, with Amphipoda, Mollusca and Decapoda ranking highest, with other important benthic organisms also being present. Dietary overlap among these species was relatively low.     

A review of the culture potential of <Emphasis Type="Italic">Solea solea</Emphasis> and <Emphasis Type="Italic">S. senegalensis</Emphasis>

A number of scientific studies have investigated aspects of soles(Solea soleaandS. senegalensis) ecology, population genetics and biology in their natural environment, and the species have been extensively studied in captivity during the last decade. Studies on the genetic population structure of sole indicate that several distinct breeding populations exist within its distributional range in European waters. Recent studies suggest a phylogenetic relatedness ofS. soleaand S. senegalensis, being found as closest sister lineages in most reconstructions. However, studies on molecular genetics and morphological traits give diagnostic differences that consistently lead to their taxonomic separation at the specific rank. Studies show that sole spawn readily in captivity, and the buoyant, fertilized eggs are easily collected. Stocking density during maturation should be 1–1.5kg/m², and temperature should be kept above 16°C (S. senegalensis) or between 8 and 12°C (S. solea). In nature, the onset of spawning is related to a rise in temperature occurring during spring (March–June). Salinity should be kept constant around 33–35 and the fish reared under simulated natural photoperiod (LDN). In other cultured flatfish species, a change in the photoperiod is the key environmental signal used to manipulate and control maturation, but at present time there are no published work that verifies or contradicts this for either S. senegalensisor S. solea. Studies indicate that a mixture of inert and live food may increase the weaning success of sole fry, and this can be further enhanced by using attractants in the dry feed. Future experiments are needed to determine the ideal time to commence weaning and determine the minimum duration of this period. Studies on alternative feeding strategies are also required. The effect of temperature and photoperiod on juvenile growth has not been studied systematically in neither of the two species and the relative importance of a direct photoperiod effect on growth in sole therefore remains to be defined.     

Niche overlap between juvenile flatfishes, Platichthys flesus and Solea solea, in a southern European estuary and adjacent coastal waters

The distribution and abundance patterns as well as the feeding ecology of two flatfishes, Platichthys flesus (Linnaeus, 1758) and Solea solea (Linnaeus, 1758), were studied in the Douro estuary and adjacent coastal waters (Portugal) in order to evaluate the niche overlap between these species. Fish were collected during sampling surveys conducted every 2 months between November 2000 and March 2002 using a 12‐m otter‐trawl with 18 mm stretched mesh size, at 12 sites within the estuary and six sites in the adjacent coastal area. The highest densities of P. flesus were registered in the middle estuary; a peak density of 9.12 ind 1000 m^?2 was obtained in October 2001. Solea solea densities were low compared with P. flesus; the highest values were also recorded in the middle estuary in October 2001. For both species, the proportion of juveniles was above 90% within the estuary. Highest densities of P. flesus and S. solea were recorded at sites with low water depth and high densities of crustaceans and polychaetes, the main food items in the diet of these species. High values of spatial and diet niche overlap were found between these two flatfishes (0.97 and 0.92, respectively).     

Does the flatfish community of the Mondego estuary (Portugal) reflect environmental changes?

Temporal and spatial patterns of abundance of the flatfish community in the Mondego estuary were investigated from 2003 to 2007, based on monthly beam trawl samples. During the study period there was a severe drought, with consequential reductions in river runoff. A total of eight flatfish species occurred within the estuary, with seasonal specific richness and abundance varying considerably. It was possible to identify three main groups: one composed of species present throughout the study period and in high densities: Platichthys flesus and Solea solea (maximum densities 3.7 and 4.1 Ind 1000 m^?2, respectively); a second group of less abundant but regularly present species: Scophthalmus rhombus and Solea senegalensis (maximum densities 0.2 and 0.3 Ind 1000 m^?2, respectively); and a third group of occasional species that occurred only during the drought period: Arnoglossus laterna, Buglossidium luteum, Dicologlossa hexophthalma and Pegusa lascaris. Flatfish distribution patterns varied according to the estuarine use guild: marine‐estuarine dependent fish occurred mainly in the upper reaches, while marine stragglers and marine‐estuarine opportunists occurred mostly in the downstream areas. Species with more northern latitudinal affinities were the most affected by the drought, related with a lesser extent of river plumes to the coastal area, resulting in a reduction in half of the abundance levels. However, flatfish species with more southern affinities increased in abundance during the drought, benefiting also from an increase in estuarine water temperature. Early summer salinity and precipitation values were good proxies for estimating abundance levels of P. flesus and S. solea, respectively, emphasizing the importance of hydrodynamics for the recruitment and abundance of these commercially important species.     

Comparative xenobiotic metabolism capacities and pesticide sensitivity in adults of Solea solea and Solea senegalensis

The measurement of enzymatic activities involved in xenobiotic biotransformation was carried out in adults of Solea solea and Solea senegalensis. The hepatic enzymes analysed were cytochrome P450 (CYP) related activities using eight fluorometric substrates and carboxylesterases (CbE). The conjugating activities of glutathione S-transferase (GST) and UPD-glucuronosyltransferase (UDPGT) were also assessed. Specific mammalian inhibitors were used as diagnostic tools for related activities of CYP1A (α-naphthoflavone; αNF), CYP2B6 and CYP2C19 (ticlopidine) and CYP3A4 (ketoconazole). The in vitro sensitivity to organophosphorous pesticides (OP) was tested in the S10 homogenate of brain (acetylcholinesterase-AChE) and liver (CbE). Furthermore, the pesticide chlorpyrifos oxon (CLPO) was used to explore the OP sensitivity of CbE of both species in two subcellular fractions (microsomes and cytosol), using two substrates. Overall, only two parameters confirmed species differences: EROD and cytosolic CbE being significantly elevated (p < 0.05) in the common sole, S. solea. A high inhibition of CYP1A related activities using several fluorometric substrates (ER, MR and CEC) after in vitro incubation with αNF confirmed all measure CYP1A1-related activities whereas ketoconazole was more specific for BFCOD (CYP3A4). Pesticide sensitivity was similar for brain AChE but hepatic CbE had a protective role that was species and pesticide dependent.     

Comparison between the life cycles of two Soleidae, the common sole, Solea solea, and the thickback sole, Microchirus variegatus, in the Bay of Biscay (France)

The life history of two Soleidae, the common sole, Solea solea, and the thickback sole, Microchirus variegatus, were compared in the Bay of Biscay in an attempt to set out factors which could explain settlement styles known to be different between juveniles of each species. Common sole juveniles had been shown to depend on coastal and estuarine nurseries, and the thickback sole to develop in open-sea nurseries, although the spawning grounds and spawning season of both species overlapped (offshore and at springtime, respectively). For this study, data on adult, juvenile and larva distributions were obtained from cruises carried out in the Bay of Biscay during the last decade. In addition, growth rate of larvae and planktonic interval duration, estimated by means of otolith increment analysis, were compared, as well as literature-derived information on behaviour of larvae. By comparing larval features, it appeared that thickback sole did not obtain the advantage of a slightly longer pelagic life span for an increased dispersal, due to an early shift to benthic behaviour. Other evidence was given by more specifically stated distributions that the further and deeper offshore spawning of M. variegatus, compared to S. solea, was a likely key-factor of the juvenile settlement process. This could explain why the thickback sole, contrary to the common sole, never reach coastal areas and thus settle offshore, in waters deeper than 30 m.     

Biological availability and humic properties of dissolved organic carbon in Lake Balaton (Hungary)

The nursery role of the Mondego estuary for marine fish species was studied between June 2003 and May 2004. The spatial and temporal distribution and abundance patterns of 0-group Dicentrarchus labrax (Linnaeus, 1758), Platichthys flesus (Linnaeus, 1758) and Solea solea (Linnaeus, 1758) were analyzed based on monthly sampling surveys in five stations along the estuarine gradient. Fishing took place during the night at low water of spring tides, using a 2 m beam trawl. The spatial patterns of estuarine colonization were different according to species. D. labrax showed a wider distribution, but the main nursery ground was the same as for S. solea. Highest densities of S. solea juveniles were found in oligohaline areas, with muddy bottoms and high benthic invertebrates availability, while P. flesus occurred mainly in the sandy uppermost areas. D. labrax was found in both these areas. Fish abundance in the estuary mainly reflected seasonal changes.     

Influence of sex, age and season on the feeding habits of the flatfish Solea senegalensis

Synopsis The Senegal sole Solea senegalensis is a commercially valuable fish and it is beginning to be cultured, in an extensive way, in fish farms in southern European countries. Little is known about its feeding habits, unlike other Solea species such as the common sole Solea solea. This paper studies the feeding habits of a wild population of Senegal sole. Differences according to sex, age and season were statistically analyzed. A total of 376 fish, aged between two and six years old, were studied over a single year. Twenty eight different families of prey were identified, belonging, in order of abundance, to crustaceans, polychaetes and molluscs. The Senegal sole trophic profile (preys items with a Simpson's dominance index 2) was: Glyceridae, Nephtydidae, Onuphidae, Apseudidae, Ampeliscidae and Upogebiidae. Feeding habits of Senegal sole were significantly influenced by sex, age and season, with crustaceans being the prey most affected.This paper is dedicated to the memory of Daniel Ray Ogilvie. who died 7 January 1995.     

Isolation of polymorphic DNA microsatellites in the common sole Solea vulgaris

A microsatellite‐enriched genomic library was obtained from the common sole, Solea vulgaris, and seven polymorphic dinucleotide markers were successfully optimized. These markers showed levels of expected heterozygosity ranging from 0.55 to 0.83 and allele number ranging from eight to 12. Three of these markers were also found to successfully amplify in the closely related Solea senegalensis and can be employed to define population genetic structure of the two Solea species and for inferring stock origins.     

Anomalies in the cephalic area of laboratory-reared larvae and juveniles of the common sole,Solea solea: oral jaw apparatus,dermal papillae and pigmentation

Synopsis The post-metamorphic common sole, Solea solea, as well as the other Soleidae and Cynoglossidae, are characterized by oral jaw elements without teeth on the ocular side. A relatively large proportion of late sole larvae and early juveniles, routinely reared in the laboratory, showed dentigerous and shape-modified premaxillae and mandibles on the ocular side. Abnormal jaws were associated with dermal papillae and a lack of pigmentation on parts of the ocular side of the head. These features are probably the result of suppressed epigenetic mechanisms normally active during ontogeny. Insights into the morphological evolution of the soleid jaw apparatus are discussed.     

Micro‐anatomical characterization of vertebral curvatures in Senegalese sole Solea senegalensis

The micro‐anatomical changes associated with lordotic and kyphotic vertebral curvatures (VC) in juvenile and adult Senegalese sole Solea senegalensis are described. In addition, it is demonstrated that the tissue and cellular structures of individual vertebrae can be severely affected. Two main conformations were found in deformed juvenile specimens: flattened vertebrae with dorso‐ventral compression and trapezoidal vertebrae forming concave and convex sides under compressive and tensile stresses. Histological analyses revealed the occurrence of an ectopic cartilaginous tissue within the acellular bone, both in juveniles and adults, possibly to cope with altered mechanical stress in deformed vertebrae. The results suggest that the alteration in loading to which curved vertebral columns are subjected might trigger vertebral reshaping and differentiation of cells towards this ectopic tissue. In addition, mesenchymal cells appear to play an important role in its formation. It is here proposed that the acellular bone of S. senegalensis is capable of adaptively responding to altered loading regimes at the structural level by reshaping vertebrae and at the micro‐anatomical level by recruiting chondrocyte‐like cells to areas of altered mechanical stress.     

Habitat quality of estuarine nursery grounds: Integrating non-biological indicators and multilevel biological responses in Solea senegalensis

Estuaries are highly valuable ecosystems that provide various goods and services to society, such as food provision and supporting nursery habitats for various aquatic species. Estuarine habitat quality assessment is thus critical in managing both ecological and economic value. In this work, various biological and non-biological indicators of habitat quality in estuarine nursery areas were determined, encompassing local environmental conditions, chemical contamination, anthropogenic pressures, juvenile Solea senegalensis condition, biomarkers response to contamination and juvenile density. The various indicators provided an integrated view on habitat quality and their responses were broadly concordant. Nursery quality assessment based on anthropogenic pressure indicators and fish biomarker responses were very similar, signaling nursery areas with higher anthropogenic pressure in Tejo and Ria de Aveiro estuaries. Yet, favorable environmental conditions across all sites could have contributed to lessen the potential hazardous biological effects of exposure to anthropogenic stressors, resulting in soles’ fairly good condition and generally high juvenile density. Nevertheless, a mismatch between high juvenile density and high estuarine contribution to adult coastal populations was observed in areas with higher anthropogenic pressures. Although a causal relationship cannot be established, the results emphasize the need to fully understand how the estuarine period spent in estuaries and local processes determine the quantity and quality of juveniles exported to marine adult populations, which is critical to achieve the full potential of the fish production service of estuaries and coastal stock replenishment.     

Connectivity between estuaries and marine environment: Integrating metrics to assess estuarine nursery function

Natural or anthropogenic induced variations in estuaries and the dynamics of marine fish populations potentially promote differences in connectivity between estuaries and marine areas, i.e. in their importance as nursery grounds. Within this context, an integrated assessment of the differential nursery function of the main estuaries along the Portuguese coast for commercial fish species common sole Solea solea, Senegalese sole Solea senegalensis, flounder Platichthys flesus and sea bass Dicentrarchus labrax was performed through several indicators based on available data. Contribution of individual estuaries to marine subpopulations was measured with potential metrics (juvenile density, habitat quantity, juvenile number and habitat quality within estuaries) and effective metrics (estuarine source of young adults in marine environment measured via otolith elemental fingerprints). The relationship between the two types of metrics was also assessed. Estuaries identified as important nursery and/or effective juvenile habitat (EJH) differed with species and no single estuary was best for all, highlighting species-specific regulation of nursery function. Multiple species assessment of nursery and EJH function differed among estuaries. Management and conservation of estuaries should focus on sites with higher contributions to adult subpopulations of multiple species. The importance of defining precise scientific and management objectives was emphasized by the different rankings of estuaries obtained with nursery or EJH criteria. Potential and effective contribution of estuaries were not significantly correlated, but in a quantitative analysis juvenile densities and number of juveniles seem related with effective contribution in some species. An agreement between potential and effective contributions of estuaries is concurrent with the acknowledged minor role of juvenile stage processes in regulation of recruitment to adult subpopulations.     

Identification of Microsatellite Repeats in Turbot (Scophthalmus maximus) and Dover Sole (Solea solea) using a RAPD-Based Technique: Characterization of Microsatellite Markers in Dover Sole

We have used a RAPD-based technique to identify several microsatellite repeats in turbot (Scophthalmus maximus) and Dover sole (Solea solea) and report the characterization of six novel polymorphic microsatellite markers for Dover sole. These are the first such markers to be developed for this flatfish species. They exhibit much higher levels of heterozygosity than those previously observed with allozyme loci and should prove useful in addressing population genetic questions as well as more fundamental aquaculture-related questions. Received ; accepted July 13, 1999.     

Parasitic copepods of the common sole, Solea solea (L.), from the Eastern Mediterranean coast of Turkey

There is increasing interest in the common sole, Solea solea (Linnaeus), as an alternative fish species in aquaculture in the Mediterranean region, and parasitic copepods are a potential hazard for farmed finfish. This paper provides taxonomic information on two species of sea lice (family Caligidae) collected from S. solea in eastern Mediterranean waters off the Turkish coast. Caligus brevicaudatus A. Scott, 1901 and Caligus apodus (Brian, 1924) were both found and this is the first report of C. brevicaudatus in Turkish waters. The discovery of C. apodus on S. solea is a new host record. Key diagnostic characters of both species are reported, supported by light and scanning electron microscopy observations. During a 12-month survey a prevalence of 28% was recorded for C. brevicaudatus, whereas for C. apodus peak prevalence was much lower (3%).     

Cloning,tissue and ontogenetic expression of the taurine transporter in the flatfish Senegalese sole (Solea senegalensis)

Flatfish species seem to require dietary taurine for normal growth and development. Although dietary taurine supplementation has been recommended for flatfish, little is known about the mechanisms of taurine absorption in the digestive tract of flatfish throughout ontogeny. This study described the cloning and ontogenetic expression of the taurine transporter (TauT) in the flatfish Senegalese sole (Solea senegalensis). Results showed a high similarity between TauT in Senegalese sole and other vertebrates, but a change in TauT amino acid sequences indicates that taurine transport may differ between mammals and fish, reptiles or birds. Moreover, results showed that Senegalese sole metamorphosis is an important developmental trigger to promote taurine transport in larvae, especially in muscle tissues, which may be important for larval growth. Results also indicated that the capacity to uptake dietary taurine in the digestive tract is already established in larvae at the onset of metamorphosis. In Senegalese sole juveniles, TauT expression was highest in brain, heart and eye. These are organs where taurine is usually found in high concentrations and is believed to play important biological roles. In the digestive tract of juveniles, TauT was more expressed in stomach and hindgut, indicating that dietary taurine is quickly absorbed when digestion begins and taurine endogenously used for bile salt conjugation may be recycled at the posterior end of the digestive tract. Therefore, these results suggest an enterohepatic recycling pathway for taurine in Senegalese sole, a process that may be important for maintenance of the taurine body levels in flatfish species.     

Growth variability of juvenile soles Solea solea and Solea senegalensis, and comparison with RNA : DNA ratios in the Tagus estuary, Portugal

Growth variability and condition of juvenile soles Solea solea and Solea senegalensis , were assessed through RNA : DNA estimates and compared to absolute growth rates. Higher mean cohort RNA : DNA ratios were observed for cohort I at the beginning of estuarine occurrence for both species (4·42 and 4·87, for S. solea and S. senegalensis respectively). Despite different estuarine colonization habits, no significant differences were observed between RNA : DNA monthly variation for both sole species within the same year ( P  > 0·05 for 2003 and 2004). Juvenile S. senegalensis showed significant differences between RNA : DNA ratios obtained for the two nursery areas ( P  < 0·001). The decrease of seasonal growth rates with fish age was similar to seasonal variation of mean RNA : DNA values. Thus the RNA : DNA pattern of juvenile S. solea and S. senegalensis reflected growth and estuarine colonization patterns.