Ontogenetic and gender-modulated behavioural rhythms in the deep-water decapods Liocarcinus depurator (Brachyura: Portunidae), Munida tenuimana and Munida intermedia (Anomura: Galatheidae)

The regulation and expression of biological rhythms with respect to sex and ontogeny in deep-water benthic decapod crustaceans constitutes an exciting field in marine biology that is far from understood. Liocarcinus depurator , Munida intermedia and Munida tenuimana are ecologically key crustacean decapod species of the Atlantic and Mediterranean shelves and slopes, and their activity rhythms in the field are poorly known. Our aim was to measure the behavioural rhythms of these species, while at the same time defining their type of displacement ( i.e. endobenthic, nektobenthic or benthopelagic). Whether gender and ontogeny modulate the rhythmic behaviour of these decapods is unknown, and we sought to clarify this issue. A temporally scheduled series of trawl hauls and light intensity measures was performed on the western Mediterranean shelf (100–110 m depth) and slope (400–430 m), close to the autumn equinox and the summer solstice. The sex and the size of animals in the catches were analysed. Catch patterns were evaluated through waveform and periodogram analyses. Liocarcinus depurator was captured at night on the shelf, whereas on the slope, animals displayed peaks both in the middle of the day and night. Size-related differences (but no gender differences) were found in its rhythmic behaviour, possibly due to intra-specific competition ( e.g. fighting) between juveniles and adults. Munida intermedia were weakly diurnal in October and both diurnal and nocturnal in June. Munida tenuimana presented no discernible rhythmicity in October, but was nocturnal in June. Both species showed no evident sex or size modulation of their behaviour. Data were interpreted assuming that all tested species present an endobenthic behaviour ( i.e. animals emerge from the substrate during the active phase of their behavioural cycle).     

Genetic diversity and population structure of three dominant myctophid fishes (<Emphasis Type="Italic">Diaphus theta,Stenobrachius leucopsarus</Emphasis>, and <Emphasis Type="Italic">S. nannochir</Emphasis>) in the North Pacific Ocean

We compared the genetic diversity of three dominant myctophid fishes in the North Pacific Ocean that have different diel vertical migration patterns on the basis of the nucleotide sequences of the mitochondrial gene for cytochrome b. No genetic structure was detected for each of these three species. The genetic diversity progressively increased for Diaphus theta, a diel migrant species showing clear diel vertical migration; Stenobrachius leucopsarus, a semi-diel migrant, in which only part of the population migrates vertically; S. nannochir, a non-diel migrant. All three species were suggested to have experienced a recent, sudden population expansion. Interspecific differences in genetic diversity might be attributable to differences in the degree of population size reduction during the glacial periods; this degree in turn corresponds to the energy demand of the fishes.     

Genetic and morphological divergence in the purple sea urchin Paracentrotus lividus (Echinodermata, Echinoidea) across the African Mediterranean coast

The present investigation focuses on population structure analysis of the purple sea urchin Paracentrotus lividus across the African Mediterranean coast, with the main aim of assessing the influence of the Siculo-Tunisian Strait on gene flow disruption in this highly dispersive echinoid species. For this purpose, patterns of morphological and genetic variation were assessed among its populations from the western and eastern Mediterranean coasts. A total of 302 specimens from seven Tunisian sites were collected and examined for morphometric variability at twelve morphometric traits. Concordant results, inferred from CDA (canonical discriminant analyses), pairwise NPMANOVA (non parametric multivariate analysis of variance) comparisons and MDS (multidimensional scaling) plot, unveiled significant inter-population differences in the measured traits among the studied populations. Furthermore, the combined use of the one way ANOSIM (analysis of similarities) and the Discriminant/Hotelling analysis allowed unravelling two morphologically differentiated groups assigned to both western and eastern Mediterranean basins. The SIMPER (similarity percentages) routine analysis showed that total dry weight, test diameter and spine length were major contributors to the morphometric separation between locations and between groups. Pattern of phenotypic divergence discerned in P. lividus across the Siculo-Tunisian Strait is interestingly in congruence with that inferred from the genetic investigation of the purple sea urchin populations from the same region based on the analysis of the mtDNA COI (cytochrome oxidase I) gene in 314 specimens from nineteen locations covering a wider geographic transect, streching westward to the Algerian coast and eastward to the Libyan littoral. The specific haplotypic composition characterizing each Mediterranean basin, as inferred from the minimum spanning network, confirmed the geographic partioning of genetic variation, as revealed by F-statistics and AMOVA (analysis of molecular variance) analyses, yielding significant genetic differentiation between eastern and western Mediterranean populations. The newly detected phylogeographic patterns, observed for the first time in P. lividus throughout the explored distribution range, suggest the involvement of different biotic and abiotic processes in shaping such variation, and provide evidence that a large and geographically exhaustive dataset is necessary to unveil phylogeographic structure within widespread marine species, previously cathegorized as panmictic in part of their distribution range.     

A cryptic species of Mytilus (Mollusca: Bivalvia) on the west coast of South Africa

A biochemical genetic study of the mussels Perna perna and Choromytilus meridionalis on the west coast of South Africa revealed the presence of an unreported mytilid mussel that had previously been mistaken, because of its shell colour polymorphism, for either P. perna or a hybrid between P. perna and C. meridionalis. The gene products of 19 protein-coding loci in P. perna. C. meridionalis and in the newly recognized mussel were examined by means of horizontal starch-gel electrophoresis. The results showed that there was no allele-frequency overlap between any of these mussel taxa, thus implying little or no genetic relatedness between them. A morphological examination showed that the previously unreported mytilid has a pitted resilial ridge (similar to that of Perna), an anterior adductor muscle (unlike either Perna or Choromytilus), and an undivided posterior foot retractor-muscle scar (unlike Perna). Such features are diagnostic of the genus Mytilus. The South African Mytilus has morphological traits that are more characteristic of M. galloprovincialis of the Mediterranean Sea than of the more cosmopolitan M. edulis. which occurs in the northern and southern Pacific and Atlantic oceans. A large heterozygosity in Mytilus sp. argues against a recent dispersal from the Mediterranean Sea via a small founder population. Rather, the presence of Mytilus sp. in South Africa may represent a relict population of a wider geographic distribution of M. galloprovincialis resulting from Pleistocene cooling. However, the warmer water at lower latitudes could have still prevented dispersal of M. edulis, a species adapted to colder waters.     

High occurrence of the elasipodid holothurian Penilpidia ludwigi (von Marenzeller, 1893) in bathyal sediment traps moored in a western Mediterranean submarine canyon

One hundred and fifty specimens of the elasipod holothurian Penilpidia ludwigi (von Marenzeller, 1893) were collected in sediment traps moored near the seabed in the La Fonera Canyon (Catalan Sea, north-western Mediterranean) and on the adjacent continental slope. These provide the third record of this apparently endemic Mediterranean species and the first record from the western Mediterranean. This was the only holothurian species trapped and the most abundant macroscopic organism found in the traps between 1200 and 1700 m depth over the whole sampling period (March–November 2001). It was particularly abundant in spring during the main flux of organic particles in the canyon. This coupling suggests that Penilpidia may aggregate at the seafloor during these events, making food availability a plausible explanation of the seasonal occurrence. Lateral transport of material re-suspended locally or up-canyon by near bottom currents appear to be the mechanism behind this uncommon occurrence, although in situ observations have recently been made on the swimming capability of this holothurian. The occurrence of benthic organisms in sediment traps set close to the seabed can provide information on bathyal benthic and benthopelagic populations.     

Reproductive biology of Parazoanthus axinellae (Schmidt, 1862) and Savalia savaglia (Bertoloni, 1819) (Cnidaria,Zoantharia) from the NW Mediterranean coast

Despite the wide distribution of zoanthids, little is known about their pattern of reproduction. Here we investigate the reproductive biology of two Mediterranean species, the common Parazoanthus axinellae (Schmidt) and the rare Savalia savaglia (Bertoloni). For both species, samples were collected during an annual cycle, from January to December 2005, in the Western Mediterranean (Ligurian Sea, Italy). Both species are gonochoric. In P. axinellae the sex‐ratio (n colonies = 30) showed a slight predominance of male colonies (M/F = 1.35), whereas in the population of S. savaglia (n colonies = 15) a predominance of females was found (M/F = 0.3). In P. axinellae the first gametocytes were visible in March, whereas in S. savaglia they became visible in May. Both species reproduce at the end of autumn when seawater temperature begins to decrease. Parazoanthus axinellae (10 m depth) spawns eggs and sperms in November, whereas S. savaglia (67 m depth) spawns in December. In P. axinellae sexes were segregated on a rocky wall, with males occurring deeper, whereas male and female colonies of S. savaglia were irregularly dispersed in the population. The maximum number of oocytes differed between the species, being higher in P. axinellae than in S. savaglia.     

Comparison of the biology, dynamics, and secondary production of Talorchestia brito (Amphipoda, Talitridae) in Atlantic (Portugal) and Mediterranean (Tunisia) populations

The biology, population dynamics, and production of Talorchestia brito were studied at two sandy beaches located on the Atlantic (Portugal) and on the Mediterranean (Tunisia) coasts, respectively. The seasonal variation in abundance and the overall densities were similar in both populations. Reproduction occurred from February to September in the Atlantic, and from March to early November in the Mediterranean. The sex ratio was male biased in the Atlantic, and female biased in the Mediterranean. Based on data from the Atlantic population, both abundance and the proportion of reproductive females were positively correlated with temperature, while the proportion of juveniles in the population was positively correlated with temperature and sediment moisture. On average, individuals from the Atlantic were larger than the ones from the Mediterranean. Life span was estimated at six to nine months in the Atlantic, and five to eight months in the Mediterranean. Talorchestia brito was shown to be a semiannual species, with iteroparous females producing two broods per year, and exhibited a bivoltine life cycle. The minimum age required for males' and females' sexual differentiation and for female sexual maturation was shorter in the Mediterranean. Growth production (P) was estimated at 0.19 g m⁻² y⁻¹ ash free dry weight (AFDW; 4.3 kJ m⁻² y⁻¹) in the Atlantic population, and 0.217 g m⁻² y⁻¹ AFDW (4.9 kJ m⁻² y⁻¹) in the Mediterranean one. Elimination production (E) was estimated at 0.35 g m⁻² y⁻¹ AFDW (7.9 kJ m⁻² y⁻¹) in the Atlantic, and 0.28 g m⁻² y⁻¹ AFDW (6.3 kJ m⁻² y⁻¹) in the Mediterranean. The average annual biomass ( ) (standing stock) was estimated at 0.032 g m⁻² in the Atlantic beach, and 0.029 g m⁻² in the Mediterranean one, resulting, respectively, in ratios of 5.9 and 7.5 and ratios of 10.8 and 9.6. Like other talitrids, T. brito exhibited geographic variation in morphometrical characteristics, sex ratio, growth rates, life span, and reproduction period, with the Atlantic population presenting a slower life history.     

Resource Partitioning Among Planktivorous Fish Larvae and Fry in a Mediterranean Coastal Lagoon

Resource partioning among the planktivorous stages of eight fish species occurring in a Mediterranean coastal lagoon was studied. Five species were migratory mullets that spawned in the sea (Liza ramada, L. aurata, L. saliens, Chelon labrosusandMugil cephalus), while the other three species were resident and spawned in the lagoon (Atherina boyeri, Cyprinus carpioandGambusia holbrooki).Mullet fry exhibited similar diets, based on the consumption of zooplanktonic Crustacea such as copepods and cladocerans, although adult chironomids were also important.Gambusia holbrookiandA. boyerifed on small prey such as copepod nauplii, copepodites and rotifers, whileC. carpioconsumed larger prey preferentially (cladocerans and copepods).According to the trophic overlap and niche width results, seven species pairs could be competing in the estuary, although further experimental evidence is still required.     

Determining the correct identity of South African Marthasterias (Echinodermata: Asteroidea)

Marthasterias glacialis are found in the cool-temperate waters of the north-eastern Atlantic Ocean, in the subtropical waters of the Mediterranean Sea and along the south-western tip of Africa. The South African Marthasterias population includes two morphotypes, a smooth, spineless rarispina form and a spiny africana form, that have been described as separate species, subspecies, or forma by various authors over the past century. To test whether these two morphotypes represent separate species, and whether either, or both, are conspecific with the North-East Atlantic species, 78 Marthasterias were collected from the Cape Peninsula of South Africa. Morphological comparisons between individuals of the two forms showed no significant clustering of samples, indicating that there is no morphological separation of the two South African forms into distinct groupings. The africana and rarispina forms were also shown to be genetically indistinguishable, using a fragment of the mitochondrial cytochrome c oxidase subunit I (COI) gene and the nuclear internal transcribed spacer 1 (ITS1). In addition, the COI sequences were also compared to those from European specimens, and phylogenetic reconstruction and intra- and interspecific levels of divergence suggested that the South African specimens form a single group that is genetically distinct from the European M. glacialis. Although the allopatric distribution, high genetic divergence (more than 3% for the COI fragment) and morphological differences suggest that the South African form should be raised to species status under the name Marthasterias africana, further work must assess an independent genetic marker (nuclear) to support raising the COI clade to species level. True M. glacialis have a spine-armament pattern of a series of three or more regular spine rows down the length of each arm, whereas M. africana are either covered in many irregularly spaced spines, or have an extraordinarily bare surface with only two spine rows per arm. Marthasterias africana may also have an actinal spine simulating the presence of a third inferomarginal spine. This work tentatively resolves the taxonomic dispute, elucidates the separation and amalgamation of the two African forms and suggests a single, uniquely South African Marthasterias species that might be distinct from the north Atlantic M. glacialis, although further analyses to test reproductive isolation between the North-East Atlantic and South African forms are required.     

Ecological niches and areas of overlap of the squat lobster ‘munida’ (Pleuroncodes monodon) and anchoveta (Engraulis ringens) off Peru

The world’s largest mono-specific fishery, the Peruvian anchovy or anchoveta (Engraulis ringens) fishery, has been the subject of many studies since the 1960s. Details of its relationship with other species have mainly focused on alternations with sardine, Sardinops sagax, and little effort has so far been paid to interactions with other species sharing the same ecosystem. This is the case for Pleuroncodes monodon, the crustacean squat lobster or ’munida’, which has become highly abundant along the Peruvian coast since the mid-1990s. Munida is now an important prey for seabirds, mammals and coastal predatory fish. Knowledge of patterns of distribution and ecological niche of munida is scarce however off Peru. Here we describe and compare spatial patterns of distribution of anchoveta and munida and their ecological niches based on data from 26 acoustic surveys performed along the Peruvian coast between 1998 and 2006. The results indicate that munida and anchoveta share ecological niches but that munida is restricted to the coldest part of the productive cold coastal waters whereas anchoveta do not present any temperature preference over a large range (14–23 °C). The recent increase in munida abundance off Peru is concomitant with colder conditions; with their onset munida extended its range from central Chile northwards. Off Peru the very shallow oxycline keeps munida from its usual bottom habitat and has forced it to adopt pelagic behaviour.     

Development of habitat models for Nephtys species (Polychaeta: Nephtyidae) in the German Bight (North Sea)

The aim of the study was the development of habitat models for Nephtys species (Polychaeta: Nephtyidae). The investigation area was the German Bight, the southeastern part of the North Sea. Models were developed based on field data collected between 2000 and 2006. In addition, data on environmental variables were retrieved from long-term monitoring data sets and from the sediment map by Figge [Figge, K., 1981. Nordsee. Sedimentverteilung in der Deutschen Bucht. Map No. 2900. Publisher: Deutsches Hydrographisches Institut, Hamburg]. The statistical modelling technique used was multivariate adaptive regression splines (MARS). Models were fitted individually for each species. Evaluation of predictive discrimination and predictive accuracy of the developed models was by calculation of the area under the receiver operating curve (AUC) or sensitivity and specificity, respectively. Habitat models with best predictive fit were selected for the presentation of habitat suitability maps.Six Nepthys species were found: Nephtys assimilis, N. caeca, N. cirrosa, N. hombergii, N. incisa and N. longosetosa. N. hombergii was most common whereas N. incisa and N. longosetosa were rare. Habitat preferences varied considerably among the species. For all investigated Nephtys species except N. longosetosa a habitat model could be developed based on four predictor variables. The habitat models with best predictive fit were those for N. cirrosa and N. hombergii. The N. caeca habitat model was of limited predictive accuracy and only accept predictive discrimination. The number of predictors as well as the relative importance of the respective predictors in the model varied among the different species. Direct comparison of most suitable habitats for the different species based on modelling revealed that in the mostly sandy regions parallel to the German coast in water depths up to 20 m an overlap between N. caeca, N. hombergii and N. cirrosa exists. In the deeper central German Bight with mostly fine sands with increased mud contents N. hombergii, N. assimilis and, at least partially and rare in numbers, N. incisa co-occur. It can be concluded that important sediment characteristics like grain size median and mud content as well as water depth and mean salinity are useful parameters to describe the habitat requirements of most Nepthys species in the German Bight. However, additional variables need to be incorporated into such analyses.     

Dioxin compounds in the deep-sea rose shrimp Aristeus antennatus (Risso, 1816) throughout the Mediterranean Sea

Polychlorodibenzo-p-dioxins (PCDDs) and polychlorodibenzofurans (PCDFs) are among the more toxic anthropogenic contaminants. They are fat-soluble and accumulate in animal tissues. Exposure to PCDD/Fs can cause several endocrine, reproductive and developmental problems in animals, including human beings. Several studies have demonstrated that fish and invertebrates living in association with sediments are exposed to and accumulate contaminants, but to date there have been no studies of PCDD/Fs contamination in deep-sea regions. Specimens of Aristeus antennatus (Risso, 1816) were collected from depths of 600–2500 m at different points in the Mediterranean Sea, from the western basin off the coast of Barcelona to the central basin off the Peloponnesian Peninsula, with otter trawl gear. Amounts of PCDD/Fs were measured in different animal tissues by high resolution gas chromatography coupled to high resolution mass spectrometry (HRGC–HRMS). This is the first study to report the presence of PCDD/Fs in deep-sea organisms dwelling at depths below 600 m. A. antennatus presented levels of PCDD/Fs of the same order of magnitude, or slightly higher, as those found in shallow-water species (Melicertus kerathurus) with respect to land-generated contamination. This highlights the widespread distribution of these pollutants and the potential threat posed to the biodiversity of fragile and vulnerable ecosystems such as the deep-sea. PCDD/F levels detected in the edible parts (muscle) of the commercial shrimp A. antennatus were clearly below the toxic limit value established by European legislation. Levels followed the trend muscle<ovary<hepatopancreas, according to the lipid content of the organs in question. The specific profile of toxic PCDD/F congeners in the studied Penaeid shrimp specimens was characterized mainly by octachlorodibenzo-p-dioxin (OCDD), 2,3,7,8-tetrachlorodibenzofuran (TCDF) and 2,3,4,7,8-pentachlorodibenzofuran (PeCDF) congeners. Moreover, non 2,3,7,8-substituted congeners bioaccumulated in this species, as observed in shallow-water crustaceans. PCDD/Fs were found in the shrimp A. antennatus throughout the Mediterranean Sea. Total PCDD/Fs burdens were higher in shrimps caught in the western Mediterranean than in those caught at eastern Mediterranean sites. There was a tendency for higher levels of PCDD/F contamination in samples obtained from deeper (2500 m) than from shallower sites (600 m).     

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.     

The paranthurid isopod crustacean Paranthura nigropunctata (Lucas, 1846): first record from the Atlantic coast of Morocco

The littoral paranthurid isopod crustacean Paranthura nigropunctata (Lucas, 1846) is recorded for the first time from the littoral of El Jadida located on the Atlantic northwest coast of Morocco. Specimens were obtained from the invasive brown seaweed Sargassum muticum and the natives Bifurcaria bifurcata and Cystoseira tamariscifolia in January 2015. This new record further confirms a significant southward distribution of P. nigropunctata and contributes to the knowledge of the biogeography of this isopod. Heretofore, the species was only known from the western and eastern Mediterranean and some Atlantic coasts. The present finding is the first of the species from Moroccan Atlantic shores, and suggests that the species may also be present in other coastal localities from Morocco and Africa. Some data on morphology, ecology and spatial distribution of the species are provided.     

Eastern and Western Poor Cod (Trisopterus minutus capelanus) Populations in the Mediterranean Sea: Evidence from Allozyme and Minisatellite Loci

Abstract. Nine allozyme and two minisatellite loci were used to investigate potential genetic differentiation among three samples of Mediterranean poor cod, Trisopterus minutus capelanus, from the Gulf of Lion, the Tuscan Archipelago and the Aegean Sea. Both types of markers showed consistent results, with F_ST values of 0.0262 and 0.0296 (P < 0.0015, after Bonferroni correction for multiple tests) for allozymes and minisatellites, respectively. Allele frequency heterogeneity tests between pairs of samples showed a clear separation between the two western Mediterranean samples (Gulf of Lion, Tuscan Archipelago) and the eastern one (Aegean Sea). The results indicate that at least two reproductively isolated populations of poor cod occur in the Mediterranean.     

Amphipoda assemblages in a disturbed area (Alicante,Spain, Western Mediterranean)

The distribution and composition of Amphipoda assemblages were analysed off the coasts of Alicante (Spain, Western Mediterranean), a disturbed area affected by several co‐occurring anthropogenic impacts. Although differences among sampled stations were mainly related to natural parameters, anthropogenic activities were linked with changes in amphipod assemblages. Expansion of the Port of Alicante, a sewage outfall and a high salinity brine discharge could be causing the disappearance of amphipods at stations closer to these disturbances. However, the completion of port enlargement works and mitigatory dilution of the brine discharge has led to the recovery of the amphipod assemblage. Among the natural parameters, depth determines the distribution of some of the species. While Siphonoecetes sabatieri was abundant at shallow stations, Ampelisca spp., Photis longipes, Pseudolirius kroyeri, Apherusa chiereghinii and Phtisica marina were more abundant at deeper stations. Grain size and percentage of organic matter also influenced amphipod distribution, resulting in changes in species composition and in the relative percentages of different trophic groups. Species such as Ampelisca brevicornis, Perioculodes longimanus, Urothoe hesperiae and Urothoe elegans were more abundant at stations with a high content of fine sand. Carnivorous species, mainly of the Oedicerotidae family, were more abundant at those stations with a low organic matter content, while detritivorous species were more abundant at stations with a higher mud content. Among 62 identified species, three were reported for the first time from the Spanish Mediterranean coast, two species were recorded for the second time and a new species of Siphonoecetes was found, Siphonoecetes (Centraloecetes) bulborostrum. These results confirm the need for further data on amphipods from the Mediterranean Spanish coast.     

Arm regeneration frequency in eight species of ophiuroidea (Echinodermata) from European sea areas

Eight ophiuroid species, six from the northern Kattegat-eastern Skagerrak and three from the northern Adriatic Sea, were examined for regeneration of arms. The species were separated into groups based upon mode of feeding and habitat. Comparison between groups collected in the northern Kattegat-eastern Skagerrak showed that infaunal suspension- and deposit-feeding species (Amphiura filiformis and A. chiajei) had significantly more scars per arm (mean number 0.78) than epibenthic suspension feeders (Ophiothrix fragilis and Ophiocomina nigra, 0.29) or epibenthic carnivores and deposit feeders (Ophiura ophiura and O. albida, 0.13). Spatial variation in arm regeneration incidence was found between sampling sites in the northern Kattegat-eastern Skagerrak for Amphiura filiformis and in the northern Adriatic Sea for Ophiothrix quinquemaculata. The ash-free dry weight (AFDW) and nitrogen (N) contents were measured in arms of six species of brittle-stars from the northern Kattegat-eastern Skagerrak. Differences between species were found, with highest concentrations of AFDW and N in Amphiura filiformis, intermediate in A. chiajei, Ophiocomina nigra and Ophiothrix fragilis, and lowest in Ophiura ophiura and O. albida. As the infaunal suspension- and deposit-feeding brittle-stars (Amphiura spp.) had the highest proportions of damaged arms and highest AFDW and N contents in their arms in this comparison, it is suggested that selective cropping of arms by demersal fish is the main cause of arm damage on Amphiura spp. in this area.     

Characterization of bacterial symbioses in Myrtea sp. (Bivalvia: Lucinidae) and Thyasira sp. (Bivalvia: Thyasiridae) from a cold seep in the Eastern Mediterranean

Cold seeps have recently been discovered in the Nile deep‐sea fan (Eastern Mediterranean), and data regarding associated fauna are still scarce. In this study, two bivalve species associated with carbonate crusts and reduced sediment are identified based on sequence analysis of their 18S and 28S rRNA‐encoding genes, and associated bacterial symbioses are investigated using 16S rRNA gene sequencing and microscopy‐based approaches. The specimens are closely related to Myrtea spinifera and Thyasira flexuosa, two species previously documented at various depths from other regions but not yet reported from the Eastern Mediterranean. Both species harbour abundant gammaproteobacterial endosymbionts in specialized gill epithelial cells. The Myrtea‐associated bacterium is closely related to lucinid symbionts from both deep‐sea and coastal species, whereas the Thyasira‐associated bacterium is closely related to the symbiont of a T. flexuosa from coastal waters off the U.K. An epsilonproteobacterial sequence has also been identified in Thyasira which could correspond to a helicoid‐shaped morphotype observed by electron microscopy, but this was not confirmed using fluorescent in situ hybridization. Virus‐like particles were observed within some symbionts in Thyasira, mostly in bacteriocytes localized close to the ciliated zone of the gill filament. Overall, results indicate that very close relatives of shallow species M. spinifera and T. flexuosa occur at cold seeps in the Eastern Mediterranean and harbour chemoautotrophic symbioses similar to those found in their coastal relatives.     

Molecular evidence for three separate cryptic introductions of the red seaweed Asparagopsis (Bonnemaisoniales,Rhodophyta) in South Africa

The red seaweed genus Asparagopsis Montagne (Bonnemaisoniales) contains two widely introduced species that are considered notorious seaweed invaders worldwide, Asparagopsis armata and A. taxiformis, both characterised by heteromorphic, diplo-haplontic life histories. To uncover cryptic introductions of Asparagopsis along the South African coastline and identify ‘Falkenbergia’ isolates (i.e. tetrasporophytic life-history phase morphologically identical between species), the mitochon-drial cox2–3 spacer was sequenced from gametophytes of Asparagopsis taxiformis from Scottburgh, KwaZulu-Natal, on the East Coast, Knysna Lagoon on the South Coast and from tetrasporophytes, otherwise unidentifiable to species level, collected from False Bay near Cape Town on the South-West Coast and Tsitsikamma on the South Coast. Only tetrasporophytes of the temperate Asparagopsis armata were encountered from the Cape Peninsula (Cape Town) probably as far east as to Port St Johns, Eastern Cape province. This is considered an introduced species, and was first collected at Kommetjie (Cape Peninsula) in 1935. Gametophytes of the warm-temperate to tropical A. taxiformis were first collected at Reunion Rocks near Durban in 1984; the KwaZulu-Natal material studied here belongs to an Atlantic Mediterranean cryptic lineage. This taxon is an ecological dominant in some intertidal and shallow subtidal areas in northern KwaZulu-Natal, and is thus considered ‘introduced’ and ‘invasive’. In contrast, A. taxiformis gametophytes, collected in Knysna Lagoon in 2008, clustered with individuals of Indo-Pacific lineage 2. The latter is considered a major invasive lineage in the western Mediterranean, but at present is categorised as introduced in South Africa. This study provides molecular evidence of three independent, cryptic introductions in South Africa, one of them probably very recent, and this is discussed with respect to potential vectors responsible for transport.     

Spatio-temporal variations in reproductive patterns and population structure of Pasiphaea multidentata (Decapoda: Caridea) in the Blanes canyon and adjacent margin, North-western Mediterranean Sea

Pasiphaea multidentata is a deep‐water caridean shrimp fished in the Mediterranean in association with the commercially exploited red‐shrimp Aristeus antennatus. A previous study describes seasonality in the reproductive pattern of P. multidentata using external morphological parametres. This study assesses the spatio‐temporal variations in the population structure, sex ratio, ovary cycle and gametogenesis of P. multidentata from three different fishing grounds in the Blanes canyon and adjacent margin (North‐western Mediterranean) over an annual cycle. The oogenetic pattern of this species is typical of a caridean shrimp. There is a pool of previtellogenic oocytes at all times that develop from the periphery of the gonad towards the centre during maturation. Previtellogenic oocytes grow to approximately 200 μm before undergoing vitellogenesis. The vitellogenic oocytes are surrounded by a monolayer of accessory cells. The maximum size observed for a mature oocyte was 1420 μm. The oocyte‐size distribution confirmed the seasonal reproductive pattern of this species; in winter, the ovaries contained mainly previtellogenic oocytes, some of which start maturing in spring, resulting in a slightly bimodal distribution. In summer, the vitellogenic oocytes reach approximately 1000 μm and in late autumn the ovaries are fully mature and ready to spawn. There were no significant differences in the reproductive and population structure patterns of P. multidentata among the three sites, suggesting that the population’s distribution is not affected by the geomorphology of the area, in particular the presence of the canyon. The populations are dominated by females at all sites and all seasons, with the arrival of juveniles in spring. The seasonal variations in the reproduction and recruitment of P. multidentata and the lack of spatial segregation within the population are discussed in terms of the species’ known biology, the effects of canyons in energy supply to the deep‐sea floor and the relationships of this species with the red‐shrimp A. antennatus.