Habitat characteristics as determinant of the structure and spatial distribution of epibenthic and demersal communities of Le Danois Bank (Cantabrian Sea, N. Spain)

The purpose of this paper is to detect and describe general trends in the spatial distribution of epibenthic and demersal communities of Le Danois Bank (El Cachucho) in relation to the environmental variables that characterize their habitat by means of multivariate ordination. Data were derived from two multidisciplinary surveys carried out in October 2003 and April 2004 included in the ECOMARG project. The surveys were focused on the study of the physical scenario, including both geological (seabed characterization from bathymetry and backscatter data) and hydrographic features, and the different compartments of the benthic fauna (endobenthic, epibenthic, suprabenthic and demersal communities). For the present study, epibenthos and demersal species were sampled using two different gears, a 3.5 m beam trawl and a Porcupine 39/52 type baca otter trawl respectively. The total species richness combining both samplers ascended to 221, including 71 species of fishes, 65 crustaceans, 35 molluscs, 29 echinoderms, 10 cnidarians and 5 sponges.Multivariate methods were used for the study of the characteristics of communities and habitats. Hierarchical cluster analysis was applied to calculate and visualize similarities between samples in terms of species composition. To assess the amount of variation of faunal densities related to a set of eight environmental variables, a redundancy analysis (RDA) was used. The set of environmental variables used were: depth, near-bottom temperature and salinity, sedimentary typology (dry weight percentages of coarse sands, medium and fine sands, silt and organic matter) and seafloor reflectivity. Using the spatial distribution of the ranges of depth and seafloor reflectivity that characterize the habitats of the faunal assemblages we defined the spatial distribution of the different communities.The multivariate analysis of 18 beam-trawl samples and 15 otter trawl samples showed the existence of 4 main assemblages associated with the more characteristic habitats of the area. The Callogorgia–Chimaera community live mainly in the areas at the top of the Bank (425–550 m depth), where the sediment coverage is lower and there exists a high presence of rocky outcrops. The Gryphus–Galeus community were found in the areas at the top of the Bank covered by medium and fine sand sediments with low organic matter content. The Phormosoma–Trachyrincus community live on the sedimentary terraces that characterize the Bank break (550–800 m) and the Pheronema–Deania community occupy all the deeper (800–1050 m) sedimentary grounds of the inner basin.     

Dynamics of suprabenthos-zooplankton communities around the Balearic Islands (western Mediterranean): Influence of environmental variables and effects on the biological cycle of Aristeus antennatus

Dynamics of suprabenthos and zooplankton were analyzed in two areas located in the NW (off Sóller harbour) and S (off Cabrera Archipelago) of Mallorca (Balearic Islands, western Mediterranean) at depths ranging between 135–780 m. Four stations situated respectively at 150 m (shelf-slope break), and at bathyal depths of 350, 650 and 750 m were sampled at bi-monthly intervals during six cruises performed between August 2003 and June 2004. Suprabenthos showed maximum biomass in both areas from late spring to summer (April to August), while minimum biomass was found in autumn (September–November). Though variable, temporal dynamics of zooplankton showed peaks of biomass in late winter and summer (February and June), while minimals occurred in autumn (August–September) and, at bathyal depths, in April. Suprabenthos (abundance; MDS analyses) showed a sample aggregation as a function of depth (3 groups corresponding to the shelf-slope break, upper slope — over 350 m; and the middle, deeper part of the slope — over 650–750 m), without any separation of hauls by season. By contrast, zooplankton samples were separated by season and not by depth. There was evidence of three seasonal groups corresponding to summer (June 2004–August 2003), autumn–winter (September and November 2003, February 2004), and spring (April 2004), being especially well established off Sóller. In general, suprabenthos was significantly correlated with the sediment variables (e.g. total organic matter content (% OM), potential REDOX), whereas zooplankton was almost exclusively dependent on Chl a at the surface, which suggests two different food sources for suprabenthos and zooplankton. The increase of suprabenthos abundance in April–June was paralleled by a sharp increase (ca. 2.8 times) in the %OM on sediment during the same period, coupled ca. 1–2 months of delay with the peak of surface Chl a recorded in February–March (from satellite imagery data). Suprabenthos biomass was also correlated with salinity close to the bottom, suggesting a link between suprabenthos abundance and changes in the oceanographic condition of water masses close to the bottom. It is suggested that a higher suprabenthos biomass recorded off Sóller in comparison to that off Cabrera in June could, in turn, be related to a seasonal inflow of Levantine Intermediate Water (LIW) in April–June in this area at mid bathyal depths (350–650 m). This trend would be based on: 1) it was evident only at mid-slope depths between 350–750 m, coinciding with the LIW distribution, and 2) it was not recorded among zooplankton (collected throughout the water column). The possible effect of the fluctuations of suprabenthos and zooplankton on higher trophic levels has been explored studying the diet and food consumption rates of the red shrimp Aristeus antennatus, as indicator species by its dominance in bathyal communities. A. antennatus increased its food consumption from February to April–June 2004 off Sóller, which in the case of large (CL > 40 mm) specimens was found in both areas. In addition, there was a shift of diet from winter to spring–early summer. In this last period, A. antennatus preyed upon euphausiids and mesopelagic decapods and fish, while benthos (e.g. polychaetes and bivalves) decreased in the diet. This indicates an increase in the food consumption and probably in the caloric content of the diet in pre-spawning females in April–June 2004, which is synchronized with the period when gonad development begins in A. antennatus females (May–June). Anyway, macrozooplankton, and not suprabenthos, is crucial as a high energetic food source in the coupling between food intake and reproduction in the red shrimp.     

The influence of oceanographic scenarios on the population dynamics of demersal resources in the western Mediterranean: Hypothesis for hake and red shrimp off Balearic Islands

The aim of the present paper is to study the relationships between some climatic indices and parental stock, recruitment and accessibility to trawl fishery of hake (Merluccius merluccius) and red shrimp (Aristeus antennatus) off Balearic Islands (western Mediterranean). Available annual catch per unit effort, recruitment and spawning stock biomass have been used as biological data. As environmental data, the meso-scale IDEA index and the large-scale North Atlantic Oscillation (NAO) and Mediterranean Oscillation (MO) indices have been used. To analyze possible links between these indices with the population dynamics of demersal resources, two non-linear approaches have been applied: (i) stock–recruitment relationships from Ricker and Beverton–Holt models, by sequentially incorporating environment factors; (ii) generalized additive modelling, both classical general and threshold non-additive models were considered. The latter simulate an abrupt change in explicative variables across different phases (time periods or climatic index values). The results have shown that two oceanographic scenarios around the Balearic Islands, associated with macro and meso-scale climate regimes, can influence the population dynamics of hake and red shrimp. This is especially true for recruitment, which seems to be enhanced during low NAO and IDEA indices periods. During these periods, colder-than-normal winters generate high amounts of cold Western Mediterranean Intermediate Waters (WIW) in the Gulf of Lions, which flow southwards and reach the Balearic Islands channels in spring, increasing the productivity in the area. This oceanographic scenario could also be favourable to the distribution of hake on the fishing grounds where the trawl fleet targets this species, increasing its accessibility to the fishery. Both spawning stock and abundance of red shrimp seems to be also enhanced by high MO index periods, which could reflect the increased presence of the saline and warm Levantine Intermediate Waters (LIW) in the study area, extending over the fishing grounds of this species. The proposed interactions can be useful to assess and manage these important demersal resources.     

Population dynamics of the red shrimp Aristeus antennatus in the Balearic Islands (western Mediterranean): Short spatio-temporal differences and influence of environmental factors

The red shrimp Aristeus antennatus is one of the target species of the bottom trawl fishery of the Balearic Islands. The objective of the present paper is to study the short spatial and temporal differences of this important economic resource between two different locations off Mallorca (Cabrera: CA; Sóller: SO), where a fleet mobility pattern has been detected, and to study the influence of environmental conditions on this species. Six simultaneous bottom-trawl and oceanographic surveys were carried out at these two locations in order to collect data from the demersal species, hydrography (temperature and salinity), trophic resources and sediment characteristics. The commercial fleet from both locations was monitored by monthly on-board sampling, log-books and daily landings obtained from sales slips. Additional data was obtained from other fishing surveys. Short spatial and temporal differences have been detected between both locations. The population at CA was more demographically homogeneous, while that at SO showed important variations, like high abundance of juveniles recruiting to fishing grounds in autumn–winter and high abundance of large females during summer. Several differences have also been found in the biology of the species between locations, such as males were more abundant in SO than in CA. Also, the reproductive period started sooner in SO than in CA, and the condition of pre-spawning females was better in SO. The percentage of total lipids in the hepatopancreas was minimal during the spawning period, showing their importance as a reserve of energy for the ovary ripening. Water masses could play an important role in these differences, the characteristics of water masses being more stable in CA than in SO. Red shrimp adult females seemed to be more correlated with the warmer and more saline Levantine Intermediate Waters, while juveniles (males and females) and adult males were more correlated with the colder Western Mediterranean Deep Waters, detected only in SO during autumn–winter. Two different hypothesis of mobility patterns for the species are discussed in relation to these observed differences.     

Isotopic composition of carbon and nitrogen of suprabenthic fauna in the NW Balearic Islands (western Mediterranean)

Stable isotope (δ¹³C and δ¹⁵N) analyses were performed on suprabenthic fauna collected in the western Mediterranean (NW Balearic Islands), at depths ranging between 350 and 780 m. Samples were collected seasonally at bi-monthly intervals during six cruises performed between August 2003 and June 2004, using a Macer-GIROQ suprabenthic sledge (0.5 mm mesh size). Twenty-four separate species (5 mysids, 12 amphipods, 2 cumaceans, 2 isopods, 1 euphausiid, 1 decapod and 1 fish) and bulk copepods were analyzed on a seasonal basis for stable carbon and nitrogen isotopes. Stable nitrogen isotope ratios (δ¹⁵N) ranged from 2.3‰ (the amphipod Lepechinella manco in September 2003) to 13.0‰ (the amphipod Rhachotropis caeca in August 2003). δ¹³C values ranged from − 24.2 (the cumacean Campylaspis sulcata in June 2004) to − 16.1 (the amphipod Bruzelia typica in November 2006). Both δ¹³C and δ¹⁵N values suggest that there are three trophic levels within the suprabenthic community. However, considering the bathymetric range of the species, the results suggest that the deepest assemblage supported only two trophic levels. The stable isotope ratios of suprabenthic fauna displayed a continuum of values and confirmed a wide spectrum of feeding types (from filter-feeders to predators). In general, and in spite of the poor knowledge about diets available for most suprabenthic species, higher δ¹⁵N were found for carnivorous amphipods (e.g. Rhachotropis spp., Nicippe tumida) consuming copepods. Low overlap for δ¹³C and δ¹⁵N values was observed, though δ¹⁵N values where less variable than δ¹³C, which suggests high resource partitioning in this assemblage. Seasonal variations in isotopic composition for both δ¹³C and δ¹⁵N were low (less than 1‰ and 3‰, respectively) and variable depending on species. Low correlations between δ¹³C and δ¹⁵N of suprabenthic fauna were found for all periods studied, though increasing from February 2004 to June 2004 (after the main peak of primary production in surface). C:N ratio (indicator of lipid content) showed higher values in summer than in winter. This suggests that lipid content may explain the seasonal patterns of δ¹³C variability and, due to the increase of storage products in phytoplankton and zooplankton, it possibly indicates the peak of primary production at the surface.     

Seasonal and short spatial patterns in European hake (Merluccius merluccius L.) recruitment process at the Balearic Islands (western Mediterranean): The role of environment on distribution and condition

This study evaluates the link between the recruitment process of European hake (Merluccius merluccius L.) of the Balearic Islands (western Mediterranean) and the environmental and physiological conditions. Spatio-temporal variation of abundance and condition of fish were evaluated at two locations each with different oceanographic conditions, one in the north (Sóller, SO) and another in the south (Cabrera, CA) of Mallorca Island. Environmental variables explored were hydrography, sediment characteristics, phytoplankton pigment concentration (ppc) and the trophic resources of hake. Individuals were divided in three life stages: recruits, post-recruits and young adults. Hepatosomatic index (HSI), relative condition index (K_n), gonadosomatic index (GSI) and fullness index (FI) were analysed for the three life stages. Recruitment starts in February with the incorporation of smaller hakes, and it can be followed through spring and early summer with a peak in April. However, some spatial heterogeneity in the recruitment process has been found between north and south of the Island. The main pulse of recruitment occurred at a different time in the two areas. Spatial heterogeneity was also consistent with the condition of hake recruits, with higher values of K_n and HSI at SO than at CA. Maximum values of K_n were found in February at SO and in April at CA, coinciding with the start of the different recruitment pulses to the fishing grounds. Post-recruits and young adults also showed higher condition at SO than at CA. The arrival in spring of the Western Winter Intermediate Waters (WIW) drives the spatial-temporal variation in abundance and condition of hake. Ppc was highly correlated with recruit abundance with a time lag of two months, while for post-recruits the time lag was three months. The observed differences in the condition of hake between areas could be a consequence of the fact that the waters to the north of Mallorca are comparatively more under the seasonal influence of WIW which is formed in more productive areas. Thus, this study characterises the short temporal and spatial variability in the hake recruitment process off the Balearic Islands, both in terms of abundance and fish condition. This pattern is explained on the basis of the mesoscale environmental variability observed between north and south of Mallorca and the ecological adaptive strategy of recruiting in the optimal environmental season.     

Evidence of episodic changes in deep-sea mesozooplankton abundance and composition in the Levantine Sea (Eastern Mediterranean)

Mesozooplankton samples from the 4270-m deep Ierapetra Basin in the oligotrophic Levantine Sea have revealed a strong temporal variability in abundance and composition hitherto unknown for the deep sea pelagic zone. The phenomenon reflected by a survey in June 1993 as compared to January 1987 is assessed by (i) the increase of the mesozooplankton standing crop throughout the water column, (ii) the absence of a decline in mesozooplankton abundance with increasing depth below 1000 m, (iii) the outstanding dominance of two interzonal calanoid copepods at bathypelagic and abyssopelagic depths and (iv) a substantial faunal change due to the codominance of Calanus helgolandicus, which was found for the first time in the area. The underlying hydrographical factors are addressed in the light of the Eastern Mediterranean Transient (EMT). Apart from a possible long-term faunal change, the EMT may exaggerate episodic plankton blooms and surface-abyssopelagic coupling in space and time.