The impact of intrapopulation variability in reproductive traits on population reproductive potential of Grand Bank American plaice (Hippoglossoides platessoides) and yellowtail flounder (Limanda ferruginea)

In geographically extensive fish populations the potential exists for reproductive traits to vary over the population’s range but, the impact that such intrapopulation variability has on overall population reproductive potential has not been formally assessed. Here intrapopulation spatial variability in size at maturity and fecundity are demonstrated for Grand Bank American plaice (Hippoglossoides platessoides) and yellowtail flounder (Limanda ferruginea). Recognition of intrapopulation variability in these reproductive traits coupled with spatial variability in abundance resulted in an increase in estimated population total egg production (TEP) by as much as 10¹⁴ eggs for American plaice and 10¹⁵ eggs for yellowtail flounder as compared to assessment of TEP for the population as a whole. Results highlight the need to explore variability in life history traits not only between but, also within populations and emphasizes the need for sufficient spatial coverage during sampling in order to assess the reproductive potential of fish populations.     

A comparative analysis of recruitment variability in North Atlantic flatfishes — testing the species range hypothesis

The hypothesis that recruitment variation in flatfishes should be most variable at the northern edge of the species range, least near the centre of the range, and intermediate near the southern limit was tested using stock and recruitment data generated from sequential population analysis for several different flatfish stocks involving four species (plaice Pleuronectes platessa, sole Solea vulgaris from the eastern Atlantic, American plaice Hippoglossoides platessoides, and yellowtail flounder Limanda ferruginae from the western Atlantic). Several groundfish species have been found to conform to this so-called species range hypothesis with the suggestion that density-independent processes predominate at the edges of the distributional range and density-dependent processes dominate in the centre of the range. Our results were generally inconsistent with the hypothesis: the coefficient of variation (CV) of recruitment for plaice in the eastern Atlantic was independent of latitude, the CV of recruitment for sole exhibited a dome-shaped relationship with latitude with the highest CVs occurring at the mid-point of the range, and the CV of recruitment for the western Atlantic stocks exhibited a monotonic decrease with latitude. We extended our latitudinal analyses by assessing both the degree of dependency of recruitment on spawning stock biomass and the spatial and temporal scales of variability in recruitment and pre-recruit survival for the eastern Atlantic stocks. In general our analysis revealed no evidence of a strong stock and recruitment relationship for any of the stocks examined, and previously published analyses revealed no such patterns with latitude. Analysis of both de-trended recruitment and pre-recruit survival time series over the species ranges of sole and plaice revealed strong positive correlations among adjacent stocks and inverse correlations among stocks at the extremes of the range. Recruitment variation in the flatfish stocks examined appears to be dominated by density-independent factors, operating at a local scale, on the egg and larval stages.     

Genetic structure of juvenile plaice Pleuronectes platessa on nursery grounds within the Irish Sea

As a preliminary investigation into the genetic structure of Irish Sea plaice we genotyped samples of juvenile plaice from six inshore areas within the Irish Sea across eight microsatellite loci and compared them with fish from two sites from the Dutch Wadden Sea (North Sea stocks). Genetic variability in all samples was generally low for that typically observed at microsatellite loci. The number of alleles per locus varied between two and nine (average 5.5) and the observed heterozygosity ranged from 0.080 up to a maximum of 0.909 (average 0.382). Few significant heterozygote deficits were observed, even when the data set was pooled. The majority (98%) of genetic variation present was within, rather than between, populations. None of the pairwise comparisons of population differentiation (F_ST) were significant (P>0.05) and a Bayesian analysis of population structure provided no evidence for a partitioning of the samples. Since juveniles arriving at nursery grounds in the Irish Sea are not distinct, it is likely that adult plaice form a single stock (perhaps with some weak differentiation). However, if plaice eggs and larvae do not disperse as predicted by a particle tracking model, then it is possible that the juveniles represent a mixture of several distinct stocks. Further work is therefore required to determine whether the phenotypic variation observed between female plaice from the eastern and western Irish Sea is the product of reproductive isolation or of the environment.     

Changes in the spatial structure of Grand Bank yellowtail flounder: testing MacCall's basin hypothesis

MacCall's basin model postulates that the geographic range of marine fish will co-vary with population density as a function of habitat selection. Therefore the geographic range of a stock will increase with increasing abundance, while the opposite is true of declining stocks. In this paper we investigated range contraction, and expansion, in the distribution of yellowtail flounder on the Grand Banks in relation to sediment type, temperature and depth. Yellowtail flounder were mainly distributed on gravely sand, sand-shell hash, rock-sandy sediments an to a lesser extent on rocky bottoms. As well, yellowtail flounder are highly associated with shallow, warmer waters more frequently than expected based on its occurrence in the environment. Employing a generalised additive model (GAM), we modelled the spatial distribution of yellowtail flounder in association with the environmental variables. The GAM provided a reasonable fit to the spatial distribution of yellowtail (58% overall). During periods of lower abundance, the fit of the spatial model increased, demonstrating the importance of depth and temperature in influencing the distribution of this species. We concluded that the observed range contraction of yellowtail flounder at low population levels represents selection for preferred habitats, whereas during periods of stock increase, the range of yellowtail flounder expands into less favourable habitats in support of MacCall's basin hypothesis.     

Annual and spatial variation in the abundance length and condition of juvenile turbot (Psetta maxima L.) on nursery grounds on the west coast of Ireland: 2000–2007

Turbot (Psetta maxima Linnaeus) is a high value commercially exploited marine flatfish which occurs in European waters, from the Northeast Atlantic to the Arctic Circle, the Baltic and Mediterranean Sea. In Ireland, turbot are the most valuable commercial non-quota species. Very little is known about their population dynamics in the wild, in particular during the sandy beach nursery phase of the life history. In 2000, a survey was established to assess flatfish species on nursery grounds on the west coast of Ireland. Eleven sandy beaches were assessed for 0+ turbot by beach seining, over an eight year period (2000–2007) during the months of August and September. The objective of the study was to estimate juvenile turbot abundance and size structure to determine if any spatial and annual trends existed. Large scale variability in the recruitment of fish to nursery grounds may be indicative of fluctuations in the adult stock. Turbot were found to recruit to five beaches consistently over the eight year period. Temporal and spatial variability in the relative abundance and length of turbot was discerned, with no apparent overall trend. However, certain nursery grounds were shown in most of the years examined to support higher abundances of turbot in comparison to other areas over the eight year period. Turbot abundances on nursery grounds were significantly correlated with mean spring sea temperatures during the pelagic stage. The condition of turbot did not significantly differ on an annual or spatial scale. Mean densities of 0+ turbot along the Irish coast were found to be similar and at times higher than other areas in Europe, ranging from 0.1 (± 0.3) individuals 1000 m^− 2 to 18.5 (± 6.9) individuals 1000 m^− 2. Mean turbot total length on beaches ranged from 3.8 cm (± 0.6) to 6.6 cm (± 4.3). The observed spatial and temporal variability in abundance and length highlights the need for long-term studies when assessing juvenile flatfish populations. Results from the present study have provided much needed baseline data on wild juvenile turbot populations which is severely lacking for this species both on an Irish and on a European scale.     

Responses of deep-water shrimp populations to intermediate nepheloid layer detachments on the Northwestern Mediterranean continental margin

A clear link between the distribution of intermediate nepheloid layer detachments on the Northwestern Mediterranean continental margin and the population structure of five congeneric megafaunal species of deep-water benthic shrimps inhabiting different depth ranges between 100 and 1100 m was found. The results of the multidisciplinary approach presented in this study provide evidence for the ecological conditions that affect the spatial distribution of juveniles and females of a deep-water fishery resource. The significantly higher occurrence of juveniles and females in regions where intermediate nepheloid layers detach from the seabed allows us to define these areas as potential deep-water nursery habitats. The spatial recognition of these habitats could contribute to the management of the stocks concerned and afford a basis for protecting these vulnerable habitats from fishing pressure. Overall, our findings furnish support for the hypothesis that deep-water benthic biological processes are linked to pelagic processes, suggesting that the detachments of intermediate nepheloid layers serve as one of the physical links between these two environments.     

Evidence of estuarine nursery origin of five coastal fish species along the Portuguese coast through otolith elemental fingerprints

Connectivity is a critical property of marine populations, particularly for species with segregated juvenile and adult habitats. Knowledge of this link is fundamental in understanding population structure and dynamics. Young adults of commercially important fish species Solea solea, Solea senegalensis, Platichthys flesus, Diplodus vulgaris and Dicentrarchus labrax were sampled off the Portuguese coast in order to establish preliminary evidence of estuarine nursery origins through otolith elemental fingerprints. Concentrations of Li, Na, Mg, K, Mn, Cu, Zn, Sr, Ba and Pb in the otolith section corresponding to juvenile's nursery life period were determined through laser ablation inductively coupled plasma mass spectrometry. Element: Ca ratios in coastal fish differed significantly amongst collection areas, except for Platichthys flesus, and were compared with the elemental fingerprints previously defined for age 0 juveniles in the main estuarine nurseries of the Portuguese coast. Identification of nursery estuaries was achieved for four of the species. Assigned nursery origins varied amongst species and differences in the spatial scale of fish dispersal were also found. Diplodus vulgaris was not reliably assigned to any of the defined nurseries. Overall, results give evidence of the applicability of estuarine habitat tags in future assessments of estuarine nursery role. Research developments on the links between juvenile and adult habitats should contribute for the integrated management and conservation of nurseries and coastal stocks.     

Annual variability of planktonic ostracods (Crustacea) in the South Adriatic Sea

Temporal variability and population structure of planktonic ostracods were investigated for the first time in the South Adriatic Sea during 1996. The maximal total ostracod abundance (1167 ind·100 m⁻³; 69% juveniles, 18% females and 13% males) was recorded in February. Thirteen species of marine planktonic ostracods were identified. Porroecia spinirostris and Archiconchoecia striata dominated the ostracod assemblage, accounting respectively for 62% and 18% of the total abundance. Their annual peaks were recorded during the cold season, which was the period of their intense reproduction, with favourable temperature conditions and lack of predators. The females surpassed the males in abundance in most species. The presence of the mesopelagic species indicated a strong influence of intermediate layer water masses from the Eastern Mediterranean Sea.     

Recruitment of flatfish species to an estuarine nursery habitat (Lima estuary, NW Iberian Peninsula)

One of the present concerns of fish biologists involves defining and identifying nursery habitats in the context of conservation and resource management strategies. Fish nursery studies usually report upon nursery occupation during the latter juvenile stages, despite the fact that recruitment to nurseries can start early in life, during the larval phase. Here we investigated the use of a temperate estuarine nursery area, the Lima estuary (NW Portugal), by initial development stages of flatfish species before and after metamorphosis, integrating the larval and juvenile phases. The Lima estuarine flatfish community comprised twelve taxa, seven of which were present as pelagic larvae, six as juveniles and three as adults. There was a general trend of increasing spring–summer abundance of both larvae and juveniles, followed by a sharp winter decrease, mainly of larval flatfishes. The Lima estuary was used by Solea senegalensis, Platichthys flesus and Solea solea as a nursery area, with direct settlement for the two first species. In contrast, indirect settlement was suggested for S. solea, with metamorphosis occurring outside the estuarine area. Estuarine recruitment of S. senegalensis varied between years, with young larvae occurring in the estuary throughout a prolonged period that lasted 6–9 months, corroborating the protracted spawning season. P. flesus, the second most abundant species, exhibited a typical spring estuarine recruitment, without inter-annual variations. Developed larvae arrived in the estuary during spring, whereas the 0-group juveniles emerged in the following summer period. The present study contributes new insight to our understanding of the economically important S. senegalensis, and highlights the importance of integrating the planktonic larval phase into traditional flatfish nursery studies.     

Distribution and quality of plaice (Pleuronectes platessa) nursery grounds on the Swedish west coast

Plaice (Pleuronectes platessa) has spatially restricted nursery grounds located in shallow soft bottom areas, where the nursery areas only make up a small fraction of the species distribution range. The importance of different coastal areas for recruitment is expected to depend on the quality and size of the nursery grounds. This paper describes the geographical extension of plaice nursery grounds at a regional level along the Swedish west coast. Densities of juvenile plaice were used as the response variable of habitat quality and were compiled based on records from autumn sampling in the various regions. Different aspects of nursery ground quality were related to the density of 0-group plaice, to evaluate the contribution of different factors to the geographical pattern observed. Larval supply was found to be the most important component of nursery quality showing a close relationship with the density of juvenile plaice on a regional scale. The relative contribution of plaice recruits from Swedish nursery grounds to the Skagerrak/Kattegat stock was evaluated using historical data. Swedish nursery grounds were estimated to contribute 77% of the recruits to the adult stock in the area.     

Inter-annual variability of North Sea plaice spawning habitat

Potential spawning habitat is defined as the area where environmental conditions are suitable for spawning to occur. Spawning adult data from the first quarter (January–March) of the International Bottom Trawl Survey have been used to study the inter-annual variability of the potential spawning habitat of North Sea plaice from 1980 to 2007. Generalised additive models (GAM) were used to create a model that related five environmental variables (depth, bottom temperature and salinity, seabed stress and sediment type) to presence–absence and abundance of spawning adults. Then, the habitat model was applied each year from 1970 to 2007 to predict inter-annual variability of the potential spawning habitat. Predicted responses obtained by GAM for each year were mapped using kriging. A hierarchical classification associated with a correspondence analysis was performed to cluster spawning suitable areas and to determine how they evolved across years. The potential spawning habitat was consistent with historical spawning ground locations described in the literature from eggs surveys. It was also found that the potential spawning habitat varied across years. Suitable areas were located in the southern part of the North Sea and along the eastern coast of England and Scotland in the eighties; they expanded further north from the nineties. Annual survey distributions did not show such northward expansion and remained located in the southern North Sea. This suggests that this species' actual spatial distribution remains stable against changing environmental conditions, and that the potential spawning habitat is not fully occupied. Changes in environmental conditions appear to remain within plaice environmental ranges, meaning that other factors may control the spatial distribution of plaice spawning habitat.     

Inter-season and interannual variations in fish and macrocrustacean community structure on a eastern English Channel sandy beach: Influence of environmental factors

The intertidal zone of an exposed sandy beach on the French coast of the English Channel was sampled with a 1.5 m beam-trawl over five years (2000 and 2003–2006) at weekly intervals. The fish and macrocrustacean catches were analysed to determine the inter-season and interannual variation in community structure and relate these variations to changes in the major environmental variables. Only six species (plaice Pleuronectes platessa, common goby Pomatoschistus microps, sprat Sprattus sprattus, sand eel Ammodytes tobianus, brown shrimp Crangon crangon and shore crab Carcinus maenas) from the 27 species captured can be considered as dominant species of the intertidal zone, and they accounted for >90% of total numbers. Most individuals caught were young-of-the-year or juvenile. Analysis of similarities (ANOSIM) and similarities percentage (SIMPER) indicated that inter-season variability of community structure (mean average similarity = 47%) was more pronounced than interannual variability (mean average similarity = 65%). Canonical correspondence analysis (CCA) indicates that a substantial component (32.2%) of the measured inter-season variation in community structure can be explained by environmental factors (mainly water temperature). The main inter-season changes in the abundance and community structure were due to the variation of the six key species and reflect the different times of their recruitment. During the five years of the study, the structure of the fish and macrocrustacean spring community persisted from year to year, with the dominant species reappearing consistently even though their abundances fluctuated from year to year. This interannual variation probably reflects variable recruitment success influenced by physico-chemical conditions. In spite of the considerable interannual variation (40 times) in the spring bloom of the prymnesiophyte alga Phaeocystis globosa we found no effect of this bloom on either fish and macrocrustacean species densities or diversity index.     

Ontogenetic patterns of concentration indicate lagoon nurseries are essential to common grunts stocks in a Puerto Rican bay

Estimates of abundance and size of three commercially exploited grunt species indicate ontogenetic changes in habitat utilization concentrate their juveniles within the lagoon of the Bay of La Parguera, Puerto Rico. Eleven biotopes, defined by four benthic structures (reef, mangrove, vegetation beds and unconsolidated sediments) and three geographic zones (inner lagoon, outer lagoon and bank shelf) were sampled randomly by visual surveys. French, bluestriped and white grunt (Haemulon flavolineatum, Haemulon sciurus and Haemulon plumeri) were common in the bay and appeared to exhibit similar life history patterns of cross-shelf migration and habitat selection. Recently settled grunts were dispersed over vegetated and unconsolidated soft-bottom sediments of the bay. The juvenile stage occurred in highest densities in shallow lagoon biotopes among the submerged prop-roots of mangrove stands and on inshore reefs. Length data indicates that grunts migrate offshore to adult habitat via increasingly deep reefs. Indices of biotope nursery function based on standing stock estimates of juveniles identified three biotopes, all within the inner lagoon as essential habitat for juveniles of 5–10 cm length interval. This concentration of juveniles within biotopes of the lagoon could represent a bottleneck to recruitment for grunt stocks. Evidence that quantity and quality of lagoon nurseries may limit recruitment indicates that these areas represent a key component of a marine protected area designed to restore fisheries within the bay.     

The influence of an extreme drought event in the fish community of a southern Europe temperate estuary

Between 2003 and 2006, a severe drought occurred throughout the Mondego River catchment's area, inducing lower freshwater flows into the estuary. As a consequence, both 2004 and 2005 were considered as extreme drought events. From June 2003 to June 2006, the fish assemblage of the Mondego Estuary was sampled monthly in five stations during the night, using a 2 m beam trawl. Fish abundance was standardized as the number of individuals per 1000 m² per season and the assemblage was analyzed based on ecological guilds: estuarine residents, marine juveniles, marine adventitious, freshwater, catadromous and marine species that use the estuary as a nursery area. A total of 42 species belonging to 23 families were identified, with estuarine residents and nursery species dominating the fish community. Variations in the fish community were assessed using non-metric MDS, being defined as three distinct periods: summer and autumn 2003, 2004/2005 and winter and summer 2006. The main drought-induced effects detected were the depletion of freshwater species and an increase in marine adventitious in 2004/2005, due to an extended intrusion of seawater inside the estuary and a significant reduction in abundance during the driest period of estuarine resident species. Nevertheless, from the management point of view, it could be stated that although some variations occurred due to environmental stress, the main core of the Mondego Estuary fish community remained relatively unchanged.     

Annual and spatial variation in the abundance length and condition of juvenile turbot (Psetta maxima L.) on nursery grounds on the west coast of Ireland: 2000–2007

Turbot (Psetta maxima Linnaeus) is a high value commercially exploited marine flatfish which occurs in European waters, from the Northeast Atlantic to the Arctic Circle, the Baltic and Mediterranean Sea. In Ireland, turbot are the most valuable commercial non-quota species. Very little is known about their population dynamics in the wild, in particular during the sandy beach nursery phase of the life history. In 2000, a survey was established to assess flatfish species on nursery grounds on the west coast of Ireland. Eleven sandy beaches were assessed for 0+ turbot by beach seining, over an eight year period (2000–2007) during the months of August and September. The objective of the study was to estimate juvenile turbot abundance and size structure to determine if any spatial and annual trends existed. Large scale variability in the recruitment of fish to nursery grounds may be indicative of fluctuations in the adult stock. Turbot were found to recruit to five beaches consistently over the eight year period. Temporal and spatial variability in the relative abundance and length of turbot was discerned, with no apparent overall trend. However, certain nursery grounds were shown in most of the years examined to support higher abundances of turbot in comparison to other areas over the eight year period. Turbot abundances on nursery grounds were significantly correlated with mean spring sea temperatures during the pelagic stage. The condition of turbot did not significantly differ on an annual or spatial scale. Mean densities of 0+ turbot along the Irish coast were found to be similar and at times higher than other areas in Europe, ranging from 0.1 (± 0.3) individuals 1000 m^− 2 to 18.5 (± 6.9) individuals 1000 m^− 2. Mean turbot total length on beaches ranged from 3.8 cm (± 0.6) to 6.6 cm (± 4.3). The observed spatial and temporal variability in abundance and length highlights the need for long-term studies when assessing juvenile flatfish populations. Results from the present study have provided much needed baseline data on wild juvenile turbot populations which is severely lacking for this species both on an Irish and on a European scale.     

Bloom of the Yessotoxin producing dinoflagellate Protoceratium reticulatum (Dinophyceae) in Northern Chile

Potential spawning habitat is defined as the area where environmental conditions are suitable for spawning to occur. Spawning adult data from the first quarter (January–March) of the International Bottom Trawl Survey have been used to study the inter-annual variability of the potential spawning habitat of North Sea plaice from 1980 to 2007. Generalised additive models (GAM) were used to create a model that related five environmental variables (depth, bottom temperature and salinity, seabed stress and sediment type) to presence–absence and abundance of spawning adults. Then, the habitat model was applied each year from 1970 to 2007 to predict inter-annual variability of the potential spawning habitat. Predicted responses obtained by GAM for each year were mapped using kriging. A hierarchical classification associated with a correspondence analysis was performed to cluster spawning suitable areas and to determine how they evolved across years. The potential spawning habitat was consistent with historical spawning ground locations described in the literature from eggs surveys. It was also found that the potential spawning habitat varied across years. Suitable areas were located in the southern part of the North Sea and along the eastern coast of England and Scotland in the eighties; they expanded further north from the nineties. Annual survey distributions did not show such northward expansion and remained located in the southern North Sea. This suggests that this species' actual spatial distribution remains stable against changing environmental conditions, and that the potential spawning habitat is not fully occupied. Changes in environmental conditions appear to remain within plaice environmental ranges, meaning that other factors may control the spatial distribution of plaice spawning habitat.     

Environmental control on early life stages of flatfishes in the Lima Estuary (NW Portugal)

Several flatfishes spawn in oceanic waters and pelagic larvae are transported inshore to settle in the nursery areas, usually estuaries, where they remain during their juvenile life. Nursery areas appear as extremely important habitats, not only for juveniles but also for the earlier planktonic larval fish. Yet, the majority of nursery studies tend to focus only on one development stage, missing an integrative approach of the entire early life that fishes spent within a nursery ground. Thus, the present study assessed the influence of environmental parameters on the dynamics of the larval and juvenile flatfishes, throughout their nursery life in the Lima Estuary. Between April 2002 and April 2004, fortnightly subsurface ichthyoplankton samples were collected and juveniles were collected from October 2003 until September 2005. Larval assemblages comprised nine flatfish species, while only six were observed among the juvenile assemblages. Solea senegalensis and Platichthys flesus were the most abundant species of both fractions of the Lima Estuary flatfishes. Larval flatfish assemblages varied seasonally, without relevant differences between lower and middle estuary. Platichthys flesus dominated the spring samples and summer and autumn periods were characterized by an increase of overall abundance and diversity of larval flatfishes, mainly S. senegalensis, associated with temperature increase and reduced river flow. On the contrary, during the winter abundance sharply decreased, as a consequence of higher river run-off that might compromised the immigration of incompetent marine larvae. Juvenile flatfishes were more abundant in the middle and upper areas of the estuary, but the species richness was higher near the river mouth. Sediment type, distance from the river mouth, salinity, temperature and dissolved oxygen were identified as the main environmental factors structuring the juvenile flatfish assemblages. Juveniles were spatially discrete, with the most abundant species S. senegalensis and P. flesus associated with the middle and upper estuary, while the remaining species were associated with the lower estuarine areas. The larval fraction exhibited distinct dynamics from the juvenile estuarine flatfish community. Larval flatfishes showed a strong seasonal structure mainly regulated by biological features as the spawning season and also by seasonal variations of water characteristics. On the other hand, juvenile flatfishes were markedly controlled by site specific characteristics such as sediments structure, distance from the river mouth and salinity regime. The present study emphasized the idea that the environmental control varies throughout the ontogenetic development, stressing the importance of integrating all the early life of a species in flatfish nursery studies.     

Do the shallow‐water habitats of a hypersaline tropical estuary act as nursery grounds for fishes?

Fish assemblages were studied in a hypersaline tropical estuary in Northeastern Brazil. A total of 104 species were collected by beach seine during the dry and rainy seasons of 2012. Two families, Atherinopsidae and Gerreidae, were numerically dominant in all zones of the estuary; juveniles of Atherinella brasiliensis, Eucinostomus argenteus, Lile piquitinga, Ulaema lefroyi, Lutjanus analis and Sphoeroides greeleyi contributed 77% of the total catch. The factors that best explained the fish abundance pattern were salinity and depth, which were related to spatial segregation. Species richness was lowest in the upper zone, an area with high salinity and shallow depth. The distinct roles of structurally different habitats (submerged macrophyte beds and mangrove prop‐roots) and the influence of physicochemical conditions in the middle zone of the estuary may influence taxonomic diversity. The middle zone of the estuary is the one most attractive to juveniles of reef species (Lutjanidae and Serranidae). The fish assemblages are clearly dominated by marine estuarine dependent species, highlighting the importance of the ecosystem as a nursery ground for marine species of the Brazilian coast.