Distribution and population structure of deep‐dwelling red coral in the Northwest Mediterranean

Commercially harvested since ancient times, the highly valuable red coral Corallium rubrum (Linnaeus, 1758) is an octocoral endemic to the Mediterranean Sea and adjacent Eastern Atlantic Ocean, where it occurs on rocky bottoms over a wide bathymetric range. Current knowledge is restricted to its shallow populations (15–50 m depth), with comparably little attention given to the deeper populations (50–200 m) that are nowadays the main target of exploitation. In this study, red coral distribution and population structure were assessed in three historically exploited areas (Amalfi, Ischia Island and Elba Island) in the Tyrrhenian Sea (Western Mediterranean Sea) between 50 and 130 m depth by means of ROV during a cruise carried out in the summer of 2010. Red coral populations showed a maximum patch frequency of 0.20 ± 0.04 SD patches·m^?1 and a density ranging between 28 and 204 colonies·m^?2, with a fairly continuous bathymetric distribution. The highest red coral densities in the investigated areas were found on cliffs and boulders mainly exposed to the east, at the greatest depth, and characterized by medium percentage sediment cover. The study populations contained a high percentage (46% on average) of harvestable colonies (>7 mm basal diameter). Moreover, some colonies with fifth‐order branches were also observed, highlighting the probable older age of some components of these populations. The Ischia population showed the highest colony occupancy, density and size, suggesting a better conservation status than the populations at the other study locations. These results indicate that deep dwelling red coral populations in non‐stressed or less‐harvested areas may diverge from the inverse size‐density relationship previously observed in red coral populations with increasing depth.     

New records of cold‐water coral sites and fish fauna characterization of a potential network existing in the Mediterranean Sea

New cold‐water coral (CWC) sites were recorded along the Apulian margin (Central Mediterranean). The species composition and depth distribution of CWCs were updated. A distribution of the CWC sites coincident with the course of the dense‐water masses that flow between the Southern Adriatic and Northern Ionian was confirmed. The faunal assemblages of five of these CWC sites were investigated and compared using experimental longlines during the spring–summer and autumn–winter seasons, between 2010 and 2014. Differences in ecological variables amongst the sites in each season were evaluated by means of a set of univariate and multivariate methods (analysis of variance, permutational multivariate analysis of variance, non‐metric multidimensional scaling). Although some differences were detected in relation to the different depths examined during spring–summer, the CWC sites showed similar features in terms of species richness and diversity as well as in the abundance of the same fish species (Galeus melastomus, Conger conger, Helicolenus dactylopterus, Merluccius merluccius, Phycis blennoides and Pagellus bogaraveo) most probably because of the distribution of adult specimens in structurally complex and heterogeneous habitats, which act as a potential ‘refuge network’ with respect to commercial fishing. The presence of maturing and mature individuals as well as post‐reproductive females indicates that these CWC sites also act as spawning areas, representing a potential ‘renewal network’ for the fish populations. The term ‘network’ used here refers to several similar subsystems (CWC sites) that play the same ecological role in a wider system (Apulian margin). These CWC communities need coherent conservation measures and management strategies according to the Ecosystem Approach to Fisheries.     

Sexual reproduction and early life‐history traits of the Mediterranean soft coral Alcyonium acaule

Understanding processes that contribute to a better comprehension of the population dynamics of long‐lived species is critical for the maintenance and potential recovery of such species. Despite the abundance of soft corals in Mediterranean rocky reefs, little information exists on their life histories and reproductive patterns. In this study, we assessed the main reproductive characteristics and early life‐history traits of the long‐lived soft coral Alcyonium acaule. The sex ratio was 1:1; the smallest fertile colonies were one finger in size (2.1 ± 0.6 cm in height), and both colony and polyp fertility increased with colony size. Likewise, the number of eggs and spermary sacs per polyp increased significantly with colony size, whereas the diameter of the female and male sexual products did not. Over 6 years of observations (2007–2012), spawning occurred primarily in July, after the seawater reached 20 °C, in a single spawning episode per year. Approximately 80% of female colonies released eggs, which were retained on the surface of the mother colony by mucous strings for up to a few days. High fertilization rates were observed during spawning in 2008 and 2009 (94.9% and 87.0%, respectively). The timing of development was ~24 h for the blastulae, ~48–72 h for the planulae and 8–22 days for metamorphosis into primary polyps. Survivorship of planulae was relatively high (~50% at 45 days after release), but only 24% of larvae metamorphosed into primary polyps, and their survivorship was moderate after 2 months (65% in 2008 and 74% in 2009). Asexual reproduction was negligible, indicating that sexual reproduction is the main mechanism supporting the maintenance and recovery of populations.     

Deep‐water Corallium rubrum (L., 1758) from the Mediterranean Sea: preliminary genetic characterisation

The precious red coral Corallium rubrum (L., 1758) lives in the Mediterranean Sea and adjacent Eastern Atlantic Ocean on subtidal hard substrates. Corallium rubrum is a long‐lived gorgonian coral that has been commercially harvested since ancient times for its red axial calcitic skeleton and which, at present, is thought to be in decline because of overexploitation. The depth distribution of C. rubrum is known to range from c. 15 to 300 m. Recently, live red coral colonies have been observed in the Strait of Sicily at depths of c. 600–800 m. This record sheds new light on the ecology, biology, biogeography and dispersal mechanism of this species and calls for an evaluation of the genetic divergence occurring among highly fragmented populations. A genetic characterization of the deep‐sea red coral colonies has been done to investigate biological processes affecting dispersal and population resilience, as well as to define the level of isolation/differentiation between shallow‐ and deep‐water populations of the Mediterranean Sea. Deep‐water C. rubrum colonies were collected at two sites (south of Malta and off Linosa Island) during the cruise MARCOS of the R/V Urania. Collected colonies were genotyped using a set of molecular markers differing in their level of polymorphism. Microsatellites have been confirmed to be useful markers for individual genotyping of C. rubrum colonies. ITS‐1 and mtMSH sequences of deep‐water red coral colonies were found to be different from those found in shallow water colonies, suggesting the possible occurrence of genetic isolation among shallow‐ and deep‐water populations. These findings suggest that genetic diversity of red coral over its actual range of depth distribution is shaped by complex interactions among geological, historical, biological and ecological processes.     

A facies of Kophobelemnon (Cnidaria,Octocorallia) from Santa Maria di Leuca coral province (Mediterranean Sea)

During a research cruise carried out in April 2010, aimed at updating the knowledge on the biodiversity of the Santa Maria di Leuca (SML) cold‐water coral province (Mediterranean Sea), a facies of the sea pen Kophobelemnon stelliferum (Muller, 1776) was found on mud‐dominated bottoms. This finding represents a new species and a new habitat record from the SML coral province as well as a new bathyal facies in the whole Central Mediterranean Sea. The colonies were collected using an epi‐benthic sledge, at depths between 400 and 470 m. A significant positive relationship between polyp number and colony length was detected. Density of the colonies ranged from 0.003 to 0.038 N·m^?2. Differences and affinities between Mediterranean and Atlantic occurrences of K. stelliferum are discussed.     

Lipid biomarkers in surface sediments from the Gulf of Genoa,Ligurian sea (NW Mediterranean sea) and their potential for the reconstruction of palaeo-environments

A series of molecular organic markers were determined in surface sediments from the Gulf of Genoa (Ligurian Sea) in order to evaluate their potential for palaeo-environmental reconstructions. Allochthonous input can be characterized by the distributions of n-C₂₉ and n-C₃₁ alkanes, n-C₂₆ and n-C₂₈ alkanols and branched glycerol dialkyl glycerol tetraethers (GDGTs), whose concentrations are generally highest near the river mouths. In the open basin however, terrestrial n-alkanes and n-alkanols may have an additional, eolian source. Autochthonous input is represented by crenarchaeol and isoprenoid GDGTs. Their concentrations are highest in the open basin showing the preference of Thaumarchaeota for oligotrophic waters. Indications of a significant degradation of sterols and C₃₇ alkenones exclude these lipids as reliable productivity proxies. Using terrestrial and aquatic lipids as end-members allows estimating the percentage of terrestrial organic matter between 20% and 58% in the coastal area decreasing to 1–30% in the deep basin. The spatial distribution of sea surface temperature (SST) estimates using the alkenone-based U^K′₃₇ index is very similar to the autumnal (November) mean satellite-based SST distribution. Conversely, TEX^H₈₆-derived SST estimates are close to winter SSTs in the coastal area and summer SSTs in the open basin. This pattern reflects presumably a shift in the main production of Thaumarchaeota from the coastal area in winter to the open basin in summer. This study represents a major prerequisite for the future application of lipid biomarkers on sediment cores from the Gulf of Genoa.     

Long‐term comparison of structure and dynamics of the red coral metapopulation of the Portofino Promontory (Ligurian Sea): a case‐study for a Marine Protected Area in the Mediterranean Sea

A large amount of data on the precious coral Corallium rubrum has attested to a dramatic shift in the size structure of its over‐harvested Mediterranean populations in recent decades. One of the main problems for the conservation of this species is the substantial lack of data concerning the time span necessary for a population subjected to pluri‐decennial harvesting pressure to return to its pristine status. Here, we present a multi‐annual data set gathered from the Marine Protected Area (MPA) of Portofino, which hosts the most important shallow‐water coral populations in the Ligurian Sea and was subjected to strong harvesting pressure from the 1950s to the early 1970s. Quantitative comparison of the population structure data recorded in 1964, 1990 and 2012 indicated a strong size increase of the colonies (from 3 to 8 g mean weight), resulting in an increase in colony biomass from 300 to the current 1500 g·m^?2. As a consequence, over the same span of time, the density of colonies has slightly decreased. The role of mass mortality phenomena, like that occurred in this area during 1999, as possible biological features triggering the switch of red coral populations from a ‘grass plain‐like’ towards a ‘forest‐like’ structure, is discussed. All these data indicate that the instigation of MPAs is a winning strategy in the conservation of this precious species and similar management plans should also be evaluated for the protection of the deep benthic communities thriving on off‐shore banks.     

Ontogenetic shifts and temporal changes in the trophic patterns of the deep‐sea red shrimp,Aristaeomorpha foliacea (Decapods: Aristeidae), in the Eastern Ionian Sea (Eastern Mediterranean)

The purpose of this paper is to provide the first detailed data concerning the diet and feeding activity of the giant red shrimp, Aristaeomorpha foliacea, in the Eastern Ionian Sea (Eastern Mediterranean), in relation to season, size class and sex. Feeding activity in A. foliacea was intense, based on its low vacuity index and high prey diversity, with a diet dominated by mesopelagic prey and less frequent occurrence of benthic taxa. Giant red shrimp displayed a highly diversified diet that exhibited slight seasonal fluctuations. The diets of both sexes consisted of 60 different prey categories belonging chiefly to three groups: crustaceans (e.g. decapods, such as Plesionika spp. and Pasiphaeidae, amphipods), cephalopods (mainly Enoploteuthidae) and fishes (Myctophidae, Macrouridae). These three prey categories accounted for 72–82% of the relative abundance and total occurrence for males and 70–88% for females, respectively. Variation in food availability, as well as increased energy demands related to gonad development and breeding activity, appear to be critical factors driving temporal changes in feeding strategy. Feeding activity increased during spring and summer, which coincides with reproductive activities (mating, gonad maturation, egg‐laying). Females seem to be more active predators than males, consuming prey with greater swimming ability. However, ontogenetic shifts in diet were also apparent, despite high dietary overlap among small, medium and large females. Large individuals, which are more efficient predators, selected highly mobile prey (e.g. fishes), whereas small individuals consumed low‐mobility prey (e.g. copepods, ostracods, tanaids and sipunculans).     

Food preferences and rhythms of feeding activity of two co‐existing demersal fish,the longspine snipefish,Macroramphosus scolopax (Linnaeus, 1758), and the boarfish Capros aper (Linnaeus, 1758), on the Mediterranean deep shelf

The feeding behaviour of two potentially competing species, the longspine snipefish, Macroramphosus scolopax, and the boarfish Capros aper was examined. While both species are very abundant along the Mediterranean coast and are regularly caught by demersal trawlers, they are of no commercial value. The diets of boarfish and longspine snipefish were investigated from samples collected between January 2001 and May 2002. Variations in the diet with fish size and season, as well as diet overlap and diversity were explored. Mysid shrimps, amphipods and gastropods were the most important food items in the diet of longspine snipefish. During ontogenetic development, M. scolopax occupies different trophic levels: the diet shifts from being predominantly composed of mysids (Anchialina agilis, Lophogaster typicus, Erythrops sp., Leptomysis spp.) in the smaller longspine snipefish [<6.5 cm total length (TL)] towards decapods (Anapagurus laevis) and amphipods (Leucothoe incisa, Eusirus longipes, Hyperidea) in the larger individuals (>6.5 cm TL). Crustacean decapods and copepods were the most important prey in the stomachs of boarfish. Mysids (Lo. typicus), euphausiids and nematodes were present in the larger individuals (>8 cm TL). A more generalist diet, still containing copepods, crustacean decapods, gastropods (Limacina retroversa) and a large variety of amphipods (e.g. Phtysica marina, Stenotoe bosphorana) and mysids (e.g. A. agilis, Leptomysis spp., Erythrops sp.), dominated the diet of C. aper between 2 and 8 cm TL. Diet overlap between longspine snipefish and boarfish was very low and the differences in stomach species diversity were explained by season and fish size.     

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.     

Patterns of Activity of the Sea Urchin Paracentrotus lividus in the Bay of Port-Cros (Var, France, Mediterranean)

Abstract. Tagged individuals of the sea urchin Paracentrotus lividus followed in Port Cros Island showed mostly nocturnal movements, with individual peaks of activity which were not coordinated for the population during the night period. The straight line distance travelled in 24 h has a mean value of 49 cm, with a range of 0 to 240 cm. Neither sea urchin size nor depth affect activity. The urchins living on rocky substrate moved significantly more than those on seagrass beds. From one day to another, the distance travelled by the population of urchins varies significantly; this is not related to season. The activity is negatively correlated with the mean wind speed; during a period of turbulence lasting several hours, movement was significantly oriented to the deep. Periods of inactivity of 1 or 2 days were observed; they are not coordinated for the individuals followed, and the relation between feeding and movement is not clear. Urchins seem to stay in a quite small area, at least for some months.     

Spatio‐temporal dynamics and biomass of Cymodocea nodosa in Bekalta (Tunisia,Southern Mediterranean Sea)

Leaf growth, biomass and production of Cymodocea nodosa were measured from October 2006 to September 2007 in Monastir Bay (Tunisia). Shoot density showed a clear seasonal pattern, increasing during spring and summer and decreasing during fall and winter. Monthly mean shoot density ranged between 633 ± 48 and 704 ± 48 shoots?m^?2. The monthly average total biomass ranged between 560 ± 37 and 646 ± 32 g dry weight (DW)?m^?2. Total biomass varied significantly among stations and sampling times but did not show seasonal variation. Leaf plastochrone intervals varied seasonally, with an annual average of 28–30 days. Leaf productivity was highest in August (2.61 g DW?m^?2?day^?1) and lowest in February (0.35 g DW?m^?2?day^?1). Annual belowground primary production varied from 263 to 311 g DW?m^?2?year^?1. Annual leaf production was approximately equal for all the stations (from 264 to 289 g DW?m^?2?year^?1). Variability in water temperature, air temperature and salinity explained the annual variability in biological characteristics. Changes in belowground and total biomass were not correlated with seasonal variability in the environmental parameters monitored. Additionally, a literature review was conducted of C. nodosa features at other Mediterranean sites, encompassing 30 studies from 1985 to 2014.     

The impact of commercial and recreational harvesting for Paracentrotus lividus on shallow rocky reef sea urchin communities in North-western Sardinia, Italy

The fishery for the edible sea urchin Paracentrotus lividus is of great importance to many European regions, although in some of them this species has shown a wide scale decline in recent years. This paper reports on direct (on P. lividus) and indirect (on the co-occurring but not fished echinoid, Arbacia lixula) effects of sea urchin harvesting in shallow rocky reefs in North-western Sardinia (Italy, central-western Mediterranean Sea), where this activity is intensively practised. Density, size (test diameter), and biomass of both species were assessed in autumn 2004 at two locations impacted by P. lividus fishery, and at two controls within an adjacent Marine Protected Area. Density of P. lividus and A. lixula was significantly greater at the controls than at the exploited locations. The average test diameter of P. lividus was also significantly larger at the controls, where large-sized specimens (i.e. >5 cm) were notably more abundant. The average size of A. lixula did not significantly differ between the impacted locations and the controls. The average biomass of P. lividus was significantly greater at the controls than at the exploited locations, whereas no differences were detected for A. lixula. These results reveal the existence of a heavy fishing impact on P. lividus in North-western Sardinia and the need for regulation of its harvesting to prevent severe direct effects on its populations. Since no indirect effects on A. lixula have been detected, it could be hypothesized that this species did not benefit from P. lividus fishery due to an only moderate competition for habitat and resources between these two echinoids. Implications for management of edible sea urchin fishery are also discussed.     

Population genetics of cold‐water coral associated Pleustidae (Crustacea,Amphipoda) reveals cryptic diversity and recent expansion off Iceland

Amphipoda are a diverse and important faunal component of deep‐sea habitats worldwide. However, relatively little is known about species that are associated with cold‐water corals, in particular concerning their genetic diversity, population genetic processes, population differentiation, and potential host specificity. We exemplarily investigate these questions studying three pleustid species—Stenopleustes aff. malmgreni (Boeck, 1871), Stenopleustes aff. nodifera (Sars, 1883) and Neopleustes aff. boecki (Hansen, 1888)—that are associated with cold‐water corals and polychaete assemblages along the Reykjanes Ridge southwest of Iceland. We sequenced mitochondrial COI and performed ddRAD to study thousands of nuclear loci. These species and a pair of morphologically cryptic species within Stenopleustes aff. nodifera were consistently delimited by COI and ddRAD. We conclude that all of our studied species are new to science, raising the number of pleustids reported from cold‐water corals to six. Two species were collected only once, and these species exhibited high levels of inbreeding, suggesting little exchange with other populations, which might be a result of strong host specificity. The other two species were wider distributed, with very low differentiation among populations, even between populations separated by 130 km. These species showed signs of recent population expansions, possibly in relation to changes in the cold‐water coral associated ecosystem (e.g., due to glacial cycles and/or volcanic or geothermal activities). They had lower levels of inbreeding and were each associated with three different host coral species, which might facilitate dispersal and gene flow. Our results show divergent population genetic patterns for the studied pleustid species. These might be attributed to the species’ host specificity and highlight important differences among species with seemingly similar life‐styles. Species diversities appear to be greatly underestimated, limiting our abilities to truly assess the vulnerability of the fauna living in these threatened cold‐water coral habitats.     

Succession patterns of polychaetes on algal‐dominated rocky cliffs (Aegean Sea,Eastern Mediterranean)

Ecological succession has been scarcely investigated on sublittoral rocky cliffs. The few relevant studies deal with the structure of the developing community and are limited to higher taxa or sessile forms. The objective of the present study was to examine succession patterns on algal‐dominated rocky cliffs both at the structural (species composition) and functional (feeding guild composition) level, using Polychaeta, a dominant taxon in this marine habitat, as a reference group. Cement panels were seasonally installed on the rocky substratum (25–30 m depth) and sampled every 3 months over a 1‐year period. Twenty‐nine polychaete species were recorded, previously reported from the surrounding benthic community, and classified into eight feeding guilds. Most species were assigned as sessile filter‐feeders; this guild dominated in abundance and biomass. A strong effect of the length of immersion and of the seasonal onset of succession on the developed communities was assessed: species composition analyses suggested convergence into a similar organization as succession proceeds, whereas the impact of starting season on succession was stronger when analysing feeding guilds. In both cases succession was faster on panels installed in winter. The main emerging patterns were in agreement with relevant surveys of the entire benthic fauna, thus supporting the efficacy of polychaetes as a surrogate group for studying ecological succession in the benthic marine environment.