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.     

Protracted oogenesis and annual reproductive periodicity in the deep‐sea pennatulacean Halipteris finmarchica (Anthozoa,Octocorallia)

Halipteris finmarchica is one of the most common species of deep‐sea pennatulacean corals in the Northwest Atlantic; it was recently determined to act as a biogenic substrate for other species and as a nursery for fish larvae. Its reproductive cycle was investigated in colonies sampled in 2006 and 2007 along the continental slope of Newfoundland and Labrador (Canada). Halipteris finmarchica exhibits large oocytes (maximum diameter of 1000 μm), which are consistent with lecithotrophic larval development. Female potential fecundity based on mature oocytes just before spawning was ~6 oocytes · polyp^?1 (500–6300 oocytes · colony^?1); male potential fecundity was 16 spermatocysts · polyp^?1 (5500–57,400 spermatocysts · colony^?1). Based on statistical analysis of size‐probability frequency distributions, males harboured one cohort of spermatocysts that matured inside 8–11 months, whereas females harboured two distinct cohorts of oocytes; a persistent pool of small ones (≤400 μm) and a small number (~20%) of larger ones that grew from ~400 to >800 μm over a year. Despite this difference in the tempo of oogenesis and spermatogenesis, a synchronic annual spawning was detected. A latitudinal shift in the spawning period occurred from south (April in the Laurentian Channel) to north (May in Grand Banks and July–August in Labrador/Lower Arctic), following the development of the phytoplankton bloom (i.e. sinking of phytodetritus). Prolonged oogenesis with the simultaneous presence of different oocyte classes in a given polyp is likely not uncommon in deep‐sea octocorals and could hamper the detection of annual/seasonal reproduction when sample sizes are low and/or time series discontinued or brief.     

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).     

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.     

Temporal and bathymetric variation of epiphyte cover and leaf biomass in a southern Posidonia oceanica (L.) meadow: the case of Mahdia coast,Tunisia

The aim of this research was to study spatial and temporal variation in epiphyte cover and leaf biomass of Posidonia oceanica in Eastern Tunisia. Sampling was conducted at four stations on the Mahdia coast during October and December 2010, and April and August 2011, which correspond respectively to autumn, winter, spring and summer in this area. Posidonia oceanica shoots were collected at two depths (5 and 10 m). Cover of macroinvertebrates and macroalgae was estimated on adult leaves. The results showed that leaf and epiphyte biomasses vary significantly according to sampling date, with the highest values recorded in August. We found a high diversity of epiphytic assemblages on the leaves of P. oceanica with clear qualitative and quantitative dominance of Rhodophyceae compared to other groups, followed by Phaeophyceae. Most epiphyte species on the leaves of P. oceanica in Eastern Tunisia are the same as those in other parts of the Mediterranean Sea. No bathymetrical variation in the epiphytic community was found in our study area, which can be explained by the high levels of water clarity in Mahdia.     

Predation has no competition: factors influencing space and resource use by echinoids in deep‐sea coral habitats,as evidenced by continuous video transects

Predation and competition are highly influential factors determining space use in foraging animals, and ultimately contribute to the spatial heterogeneity observed within habitats. Here we investigated the influence of competition and predation on space and resource use via continuous video transect observations – a tool that has not previously been employed for this purpose. This study therefore also evaluates video data as a pragmatic tool to study community interactions in the deep sea. Observations were compiled from 15 video transects spanning five submarine canyons in the Bay of Biscay, France. Substrate choice, positioning on the coral, echinoid aggregate size, and the presence/absence of predators (e.g. fish and decapods) as well as competitors (both inter‐ and intra‐specific) were recorded. Two dominant co‐existing echinoid taxa, echinothurids and Cidaris cidaris (3188 total observations), were observed in the study. For the echinothurids, no significant trends were detected in the inter‐ and intra‐specific competition data. For Cidaris cidaris, significant shifts in substrate use were correlated to the presence of inter‐specific competitors (echinothurids), whereby an increase in dead coral substrate usage was observed. Highly significant patterns were detected amongst echinoids near fish and decapod predators. A shift to the base of the coral infra‐structure was correlated to the presence of fish, and fewer individuals were observed in the open areas of the reef and a greater number were found in the mid and top sections of the coral when in the presence of decapods. Aggregates formed irrespective of the presence of predators. Aggregations are likely to form for feeding and reproduction rather than for defensive purposes; and migration along the coral infra‐structure may be a predator‐driven behaviour as echinoids seek refuge from predators. Predation risk might play a stronger – or more detectable – role in structuring echinoid space and resource use in deep‐sea coral habitats. In addition, the study successfully detected patterns in the video data thereby demonstrating its potential usefulness for similar ecological studies on other deep‐dwelling megabenthos.     

Long‐term temporal and spatial patterns in bioeroding sponge distribution at the Abrolhos Bank,Brazil, Southwestern Atlantic

Bioeroding sponges belong to the most dominant bioeroders, significantly contributing to the erosion of coral reefs. Some species are tolerant or even benefit from environmental conditions such as ocean warming, acidification, and eutrophication. In consequence, increases in sponge bioerosion have been observed on some coral reefs over the last decades. The Abrolhos Bank is the largest coral reef system in the South Atlantic. It has been affected by sedimentation, eutrophication, overfishing, and climate change, mainly affecting coastal reefs, and at lesser intensity outer ones as well. This study aimed to describe spatial and temporal patterns in bioeroding sponge distribution in carbonate substrates in the Abrolhos Bank. Photo‐quadrats were used to compare bioeroding sponge abundance between two shallow reefs: a coastal, Pedra de Leste (PL), and an outer reef, Parcel dos Abrolhos (PAB). Each individual was delimitated over the substrate by determining the sponge surface through a line connecting the outermost papillae. The study was conducted over 6 years in 2008–2009 and 2013–2016. Four species of bioeroding sponges were identified: Cliona carteri Ridley, 1881, C. delitrix Pang, 1973, C. cf. schmidtii Ridley, 1881, and Siphonodictyon coralliphagum Rützler, 1971. The distribution and abundance of species varied between the inner and outer reefs and across the years, and displayed certain selectivity for the calcareous substrates recorded. Crustose coralline algae (CCA) were the main substrate excavated by the most abundant bioeroding species, C. carteri, and represented 70% of the substrate types occupied by this sponge (CCA, coral overgrown by CCA and plain coral). The highest abundance of bioeroding sponges observed in photo‐quadrats was 21.3 individuals/m² at the outer reefs (PAB) in 2014. The abundances or areal extents of bioeroding sponges were up to 10 times greater on the outer reefs than on the coastal ones, where sedimentation is higher and more strongly influenced by siliciclastic material. Moreover, a higher herbivorous fish biomass has been reported on outer reefs which could also influence the higher abundance of bioeroding sponges in outer reefs. During the study period of 6 years, an increase in bioeroding sponge abundance was observed at the outer reefs (PAB), with the sea surface temperature increase. As CCA have an important role in reefal cementation and carbonate production in the Abrolhos reefs, a bioerosion impact might be expected, in particular, on the outer reefs.     

Fitness‐related consequences shed light on the mechanisms of covering and sheltering behaviors in the sea urchin Glyptocidaris crenularis

We know of no comparative assessment on the benefits and costs of long‐term covering and sheltering behaviors in sea urchins. The present study investigated the long‐term effects of conditions suitable for sheltering and covering behaviors on fitness‐related traits of sea urchins Glyptocidaris crenularis. In general, conditions suitable for covering and sheltering behaviors significantly affected the fitness‐related traits of G. crenularis in a long‐term laboratory study of 31 months. Glyptocidaris crenularis kept in conditions suitable for sheltering behavior (bricks with openings) showed significantly lower test size, body weight, organ (test, lantern, gonad and gut) weights, gonad index and slower gonad development than those kept in conditions suitable for covering behavior (presence of shells) and the control conditions (without conditions for covering and sheltering). However, the index of maximum pressure resistance of the test was significantly higher in G. crenularis kept in the sheltering conditions than those in the covering and control conditions. The present study provides new insight into the mechanisms of covering and sheltering behaviors and has implications for the conservation and aquaculture of sea urchins.     

Size‐frequency distributions of Joania cordata and Argyrotheca cuneata (Brachiopoda: Megathyrididae) from the Central Tyrrhenian Sea

Size‐frequency distributions can support reliable inferences concerning population dynamics of brachiopods, but only a few data are available so far. In this study, length and width frequency distributions of dead specimens of the Recent brachiopods Joania cordata and Argyrotheca cuneata from the Marine Protected Area ‘Secche di Tor Paterno’, Central Tyrrhenian Sea, Italy (41°35′ N, 12°20′ E), are reported in order to add new data about size‐frequency distributions of brachiopods. The studied specimens came from death assemblages in the coralligenous substrate, in the Posidonia oceanica meadows, and in the sand channels. The observed patterns vary from left‐skewed (J. cordata) to right‐skewed (A. cuneata), indicating respectively a low and high mortality of smaller individuals. Significant differences between the coralligenous substrate and the P. oceanica meadow were observed for both species, revealing a variation among different habitats. All length and width distributions are clearly polymodal, but the biological meaning of the peaks is difficult to interpret, as the two species seem to have a 2‐year life span. A biometric analysis of shell sizes revealed that length and width are the most variable parameters during the growth of the animal.     

Mitochondrial DNA variation reveals distinct lineages in Penaeus semisulcatus (Decapoda,Penaeidae) from the Indo‐West Pacific Ocean

Penaeus semisulcatus, the green tiger prawn, is an ecologically and economically important penaeid shrimp in the Indo‐West Pacific region, especially in rice‐shrimp farming and capture fisheries in Bangladesh and Sri Lanka, respectively. Genetic variation and phylogeography of samples of this species from Bangladesh and Sri Lanka were studied utilizing different mitochondrial DNA markers, i.e. cytochrome oxidase subunit 1 (CO1), control region (CR) and 16S rRNA genes. No evidence of population structure was observed in Bangladesh, but distinct variations were found among the Sri Lankan samples (Φ_ST = 0.04, p = .002; F_ST = 0.07, p = .001), with the western sample differing from the northwestern and southern samples. The Bangladesh population had lower genetic diversity than two of the three Sri Lanka populations. The phylogeography of P. semisulcatus revealed two distinct mitochondrial DNA (mtDNA) lineages, one in the Western Pacific Ocean and second in the Indian Ocean. The Bangladesh samples showed highest levels of similarity with samples from Sri Lanka, India and Malaysia, with the Bangladesh and Sri Lanka populations sharing the most common recent ancestry. Among the Indian Ocean samples, high levels of variation were observed in the samples from Iran, indicating admixture of two distinct mtDNA lineages, one shared by the populations from the Bay of Bengal and the other possibly originating from Eastern Africa. The genetic and phylogeographic information obtained in this study will be useful in appropriate planning for management and conservation of shrimp fisheries in Bangladesh and Sri Lanka, and in the Indo‐West Pacific region.     

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.     

Growth and population dynamics of the non‐indigenous species Branchiomma luctuosum Grube (Annelida,Sabellidae) in the Ionian Sea (Mediterranean Sea)

The population dynamics and gametogenesis of the non‐indigenous polychaete species Branchiomma luctuosum Grube, 1869 (Annelida, Sabellidae) has been investigated at three sites in the Taranto Seas (Ionian Sea, Mediterranean Sea). The species, probably introduced from the Red Sea, has been reported in the Mediterranean Sea since 1983. The species is hermaphrodite, and the reproductive season is between June and October when the largest mean size of oocytes was recorded together with the presence of mature spermatozoa. Small oocytes are present in specimens reaching about 20–25 mm in length. Therefore oogenesis seems to begin early during the first year of life, but the first reproduction can occur when the worms attain a larger size and are at least 6 months of age. Although most of the individuals reproduce seasonally within a discrete period, some individuals can reproduce in different periods during the year, the oogenesis of individuals not being synchronous. A life span of at least 2 years is highlighted, with a faster growth rate during the first months (about 20 mm per month) decreasing to about 10 mm from the 3rd to the 8th months and slowing down again after the worm reaches 100 mm in size. Some differences in growth performance are discussed, enhanced by comparing the sites located at greater depth (5–7 m) and those located at 0.5 m depth, together with the possible interactions of this alien species with the autochthonous sabellid Sabella spallanzanii.     

Multiple paternity in the common long‐armed octopus Octopus minor (Sasaki, 1920) (Cephalopoda: Octopoda) as revealed by microsatellite DNA analysis

Multiple paternity was assessed in Octopus minor using microsatellite DNA markers. Octopus minor adults were captured in traps and kept in indoor cement ponds. The females spawned after several days. Ten broods (B1–B10), each containing 15 embryos and the maternal octopus, were maintained until the embryos reached the stage at which DNA could be extracted and used for genotyping. Multiple paternity was tested using five microsatellite DNA markers and the results proved the hypothesis that multiple paternity occurs in O. minor (observed in six out of the 10 broods). Analysis with GERUD software gave the minimum number of fathers and showed that four broods were sired by a single male, three were sired by two different males and three were sired by three different males. In brood B10, the number of fathers identified by the single‐locus method was one fewer than that by the GERUD method. Of the six multiple paternity broods, four (66.7%) showed significant skew from equal paternal contributions, which indicated that sperm competition and/or cryptic female choice may be important for post‐copulatory paternity bias in this species.