The distribution of megabenthic, invertebrate epifauna in the Balearic Basin (western Mediterranean) between 400 and 2300 m: Environmental gradients influencing assemblages composition and biomass trends

The distribution of megabenthic epifauna (invertebrates) in the Balearic Basin (western Mediterranean) has been analyzed at depths between 427 and 2265 m after compiling samplings performed in 1985–1992 and 2007–2008 with an OTSB-14 bottom trawl. 84 epibenthic taxa of invertebrates (excluded decapod crustaceans) were collected. Epibenthic assemblages were organized in five groups (n-MDS analyses) as a function of increasing depth: upper slope assemblage, U, hauls between 427 and 660 m; middle slope assemblages M1 and M2, hauls between 663–876 m and 864–1412 m, respectively; lower slope assemblages L1 and L2, hauls between 1488–1789 m and 1798–2265 m, respectively). We found significant differences in assemblage composition between all depth-adjacent pairs of groups. Trends in the distribution of biomass vs. depth and within assemblages varied when hauls taken over insular were compared to those over mainland slopes. Over insular slopes we found (n-MDS) only four distinct depth assemblages, with significant differences between all depth-adjacent group pairs, except between L1 and L2. Over the mainland slope, two peaks of biomass situated at U (427–660 m) and at L1 (1488–1789 m) were clearly identified, attributable to the echinoid Brissopsis lyrifera and holothurian Molpadia musculus at U and to the synallactid holothurian Mesothuria intestinalis at L1. The distribution of biomass vs. depth on insular slopes did not follow this pattern, showing no significant biomass peak below 1000 m and a total biomass an order of magnitude lower than adjacent to the mainland. After compiling available environmental data over the mainland slope off Barcelona, we found coincidence between the peak biomass of Mesothuria intestinalis and: i) a significant increase of labile OM (%OrgC, C/N, hydrolizable aminoacids–EHAA, and the EHAA/THAA-total hydrolizable aminoacids-ratio) over 1600 m; and ii) an increase of turbidity and T at 1500–1600 m in February 2008. We suggest that such OM inputs must likely be associated to the formation of nepheloid layers close to submarine canyons, probably associated with oceanographic processes in deep water masses in the area. This would explain why aggregations of M. intestinalis were linked to the mainland part of the Balearic basin, with highest densities located south of canyons. If hotspots of biomass as cited here for M. intestinalis are regulated by factors such as river inputs, both natural climatic changes (e.g. changes in rainfall regimes) and human impact (e.g. river damming) may affect deep-Mediterranean communities below 1000 m.     

Sponge Communities in Three Submarine Caves of the Balearic Islands (Western Mediterranean): Adaptations and Faunistic Composition

Abstract. The sponge fauna of three neighbouring submarine caves of the Island of Majorca (Balcaric Islands) is studied along a light gradient; 45 species have been recorded and systematic data are given for those showing morphological, pigmentary, or spicule differences. Adaptation to darkness and precarious trophic conditions is also discussed.     

Diet of Phycis blennoides (Gadidae) in Relation to Fish Size and Season in the Western Mediterranean (Spain)

Abstract. Despite its commercial value, little is known about the feeding ecology of the greater forkbeard, Phycis blennoides (Brünnich 1768). The stomach contents of 2631 greater forkbeard were taken at monthly intervals off the coast of the Gulf of Valencia (western Mediterranean). They were examined in order to assess the diet and to determine the fish size- and season-related dietary variation of the species. Prey items were identified to the lowest taxonomic level possible. The basic food consisted of epibenthic species, mostly Decapoda, Mysidacea and Teleostei. A change in food composition based on fish size was observed and seasonal variation in diet was also recorded. The food overlap analysis indicated a high convergence in diet among similar size groups.     

Macrofaunal density, biomass and composition of estuarine sediments and their relationship to the river plume of the Rhone River (NW Mediterranean)

This study investigates species community patterns and sediment relationships of benthic macrofauna off the Rhone river delta. Along a WE transect, changes in diversity, density, biomass and trophic structure were coupled with the mean extension of the river dilution plume. Species number and diversity were at a minimum off the river mouth, below the area of the minimum surface salinity. Density decreased by a factor 2 and biomass by a factor 5 from the fluvial to the marine system. These features are due to high overload of terrestrial organic matter in the river prodelta as evidenced by the carbon isotopic signature of surface sediment and by pigment content. On the basis of a non-metric MDS analysis and of Dufrêne and Legendre method (1997), groups of stations and characteristic species associated were identified. These species, mainly small capitellids, spionids, lumbrinerids and sternaspids, correspond to a successional dynamic in response to changes in sedimentation conditions, mainly in organic matter quality of the surface sediment. The succession observed in space was similar to described in macrotidal estuaries and off other deltaic systems all over the world and to that observed in time following the Rhone river severe flood events. Results suggest that organic matter quality is an important factor with regard to benthic macrofauna successions and recovery mechanisms following disturbances. The differences observed between the Rhone deltaic system and the general model of relations between shelf processes and discharge of large rivers are attributed to a more regular supply of organic material from terrestrial origin on the Rhone continental shelf.     

The feeding and diet of the deep-sea shrimp Aristeus antennatus off the Balearic Islands (Western Mediterranean): Influence of environmental factors and relationship with the biological cycle

Spatio-temporal variation of feeding intensity and diet in the red shrimp Aristeus antennatus was studied at two locations around the island of Mallorca (Balearic Islands, Western Mediterraean) in August, September, and November 2003, and in February, April and June 2004 at depths between 550 and 750 m. The two areas, with different oceanographic conditions, were respectively located in the northwest (Sóller) and the south (Cabrera) of Mallorca. Off Sóller, feeding intensity of A. antennatus showed a significant increase from February to April and June 2004 in all the three size-classes studied (small shrimps: CL < 30 mm; medium: CL between 30 and 40 mm; large: CL ? 40 mm). Off Cabrera, the highest fullness was recorded in November 2003 among small and medium shrimp, while only large specimens showed patterns similar to that found off Sóller. Off Sóller, the diet of both small (CL < 34 mm) and large (CL ? 34 mm) A. antennatus was mainly influenced by season, with three dietary groups corresponding to August-September 2003, to November 2003/February 2004, and to hauls from April to June 2004. Off Cabrera, hauls (representing diets) were grouped by depth, never by season. The most remarkable seasonal shift in the diet of A. antennatus off Sóller was the increase of mesopelagic prey in April-June relative to other months. In all size categories there was an increase off Sóller in the energy intake of prey ingested from February to June 2004, an increase not found off Cabrera. Degree of digestion of mesopelagic prey indicated nocturnal feeding on mesopelagic fauna. These prey probably have a shallower depth distribution at night than found in our daylight sampling, and possible migratory movements among prey and A. antennatus at night would explain the lack of correlation between prey abundance in guts and in the environment found during daylight periods for most micronekton mesopelagic prey (euphausiids, myctophids and sergestids). Off Sóller, fullness and diet were significantly linked to temporal changes in water column productivity (e.g., Chl a readings, fluorescence) and to changes in the shrimp biology (lipid content of hepatopancreas, Gonado-somatic Index, GSI). Off Cabrera, we found a higher dependence of fullness and diet with T and S, both variables in turn related to depth. The increase of stomach fullness and dietary energy intake in pre-reproductive females from February to April-June 2004 found off Sóller, coupled with the consumption of mesopelagic prey, was parallel to a significant increase of the gonad weight (GSI, fecundity) in June. Most individuals attain gonad development in the period May-June, after two months of the peak of primary production at the surface. The strong link found between pelagic resources and reproductive processes in a deep-sea species such as the shrimp Aristeus antennatus, situated near the top of the trophic web, suggests a rapid energy flow via mesopelagic fauna between surface primary production and bathyal megabenthic communities at oligotrophic insular areas. In contrast to mainland areas off the Catalan coasts submitted to the influence of submarine canyons, around the island of Mallorca the empoverishment of benthos biomass may enhance consumption of micronektonic prey and a possible accumulation of pre-reproductive females of A. antennatus in areas (e.g., steep slopes and persistent frontal systems found off Sóller) with high zooplankton aggregations.     

Deep-water stands of Cystoseira zosteroides C. Agardh (Fucales,Ochrophyta) in the Northwestern Mediterranean: Insights into assemblage structure and population dynamics

Populations dominated by Cystoseira zosteroides, an endemic and threatened Mediterranean seaweed, colonize deep-water rocky habitats down to more than 50 m depth. Assemblages dominated by this species display high algal and invertebrate species richness. Algal biomass averages 1134 g dw m⁻². Erect and turf algae account for only 25% of total algal dry weight, while encrusting corallines are responsible for the remaining 75%. Sponges, bryozoans and ascidians constitute the dominant sessile macrofauna. Cystoseira zosteroides is the dominant erect algae, with a mean biomass of 60.6 g dw m⁻², and densities ranging from 4 to 7 plants m⁻². The alien turf alga Womersleyella setacea has a biomass of 104.2 g dw m⁻² and covers most of the understory substrate. The size-frequency distribution of C. zosteroides populations shows differences over time. Mean annual growth of the main axis is around 0.5 cm and mean annual mortality rate is lower than 2%. Recruitment was almost nil during the studied period of time (10 years). Processes structuring these deep-water Cystoseira stands must be driven by episodic disturbances, after-disturbance recruitment pulses, and long periods of steady growth that last at least 10 years. However, it is also possible that recruitment is irreversibly inhibited by the alien alga W. setacea in which case these old-growth stands are faced with extinction. The highly diversified assemblages and the low growth and low mortality rates of C. zosteroides indicate high vulnerability to natural and anthropogenic disturbances, and call for effective measures to ensure their conservation.     

Crustacean decapod assemblages associated with fragmented Posidonia oceanica meadows in the Alboran Sea (Western Mediterranean Sea): composition,temporal dynamics and influence of meadow structure

The decapod assemblage associated with a Posidonia oceanica meadow located near its western limit of biogeographic distribution was studied over an annual cycle. Fauna samples were taken seasonally over a year (five replicates per season) in two sites located 7 km apart, using a non‐destructive sampling method (airlift sampler) for the seagrass. The dominant species of the assemblage, Pisidia longimana, Pilumnus hirtellus and Athanas nitescens, were associated with the protective rhizome stratum, which is mainly used as a nursery. The correlations between decapod assemblage structure and some phenological parameters of the seagrass shoots and wave height were negative or null, which reflects that species associated with the rhizome had a higher importance than those associated with the leaf stratum. The abundance and composition of the decapod assemblage as well as the ecological indexes displayed a seasonality trend with maximum values in summer‐autumn and minimum in winter‐spring, which were related to the seawater temperature and the recruitment periods of the dominant species. The spatial differences found in the structure and dynamics of the assemblages may be due to variations in the recruitment of the dominant species, probably as a result of the influence of local factors (e.g. temperature, currents) and the high dispersal ability of decapods, together with the patchy configuration and the surrounding habitats. The studied meadows are fragmented and are integrated within a mosaic of habitats (Cymodocea nodosa patches, algal meadows, rocky and sandy bottoms), which promotes the movement of individuals and species among them, maintaining a high species richness and evenness.