Temporal and spatial variability in settlement of the sea urchin<Emphasis Type="Italic"> Paracentrotus lividus</Emphasis> in the NW Mediterranean

Settlement into the benthic habitat may be an important process in regulating sea urchin abundance, which potentially modifies the structure of benthic communities. Strong settlement events may increase sea urchin abundance beyond a certain threshold, leading to the formation of coralline barrens (overgrazed communities with a dominance of encrusting coralline algae). To understand the role of settlement in regulating sea urchin populations we first need to determine settlement variability. Temporal variation in settlement of the sea urchin Paracentrotus lividus was monitored at three sites in the Medes Islands, NW Mediterranean, during three settlement seasons (March 1998 through October 2000). Spatial variation in settlement was studied in 1999 at 50 sites along a gradient of exposures to waves and currents, inside and outside the archipelago, and separated by distances from tens to thousands of meters. Bathymetric distribution of settlement was also studied in 2000 at six sites at 5, 10, 15 and 20 m depths. Settlement of P. lividus occurred in a single annual peak within 3 weeks in May–June. Differences in settlement between years were more than two orders of magnitude. Spatial variability was found at all scales investigated, showing strong patchiness at the smallest spatial scales (tens of meters). Sea urchins settled preferentially at depths between 5 and 10 m. Substratum type, level of protection, and adult population densities were not significant in determining settlement. However, settlement was found to be related to the degree of exposure to waves and currents, indicating that physical processes are very important at the spatial scales investigated. This greatly variable settlement is a necessary, although not sufficient, condition to create gradients of adult P. lividus abundance. Further studies should be designed to investigate the interaction between settlement strength and post-settlement mortality.Communicated by O. Kinne, Oldendorf/Luhe     

Pelagic-benthic coupling in a subtidal system of the North-Western Mediterranean

Suspended particulate matter, zooplankton, and macrobenthos dynamics were investigated in a shallow area of the Ligurian Sea (north-west Mediterranean) characterized by wide temporal variability over an annual cycle. As indicated by multivariate analyses, the seasonal dynamics can be summarized as follows: (1) a late winter-early spring phytoplankton bloom followed by high zooplankton and macrobenthos densities during the spring months; (2) low-quality particulate suspended matter in summer, and an increase in the importance of zooplankton taxa with a wide range of feeding strategies, a decrease in macrofaunal abundance, and an increase in deposit-feeders and predators; and (3) a second phytoplankton bloom in autumn, followed by an increase in copepod density and a low macrofaunal abundance. In conclusion, pelagic and benthic communities in the coastal area of the Ligurian Sea mainly seem to be controlled bottom-up. Our results suggest that the quality of the particulate organic matter may play an important role in determining the temporal changes of both plankton and benthic assemblages, while the direct influence of other environmental features (such as sediment grain size) is relevant only for some macrobenthic taxa (e.g. crustaceans).     

Size,spatial, and bathymetrical distribution of the ascidian Halocynthia papillosa in Mediterranean coastal bottoms: benthic–pelagic coupling implications

Benthic suspension feeders are abundant in Mediterranean coastal environments, though most of them are threatened due to overexploitation, climate change impacts, and unregulated diving or fishing practices. Little is known about most of the coastal communities in terms of large-scale distribution and realistic benthic–pelagic coupling implications, which are keys to understand and manage those threatened ecosystems. The active suspension feeder Halocynthia papillosa (one of the most common ascidian species of the Mediterranean Sea) was selected as a model organism to help to understand the ecological role in benthic–pelagic coupling processes and its importance as a carbon sink (an essential ecosystem service). The spatial and bathymetrical distribution of this organism has been studied using remotely operated vehicle video transects. The species was distributed throughout the study area, with a maximum density of 4 specimens m^?2. The highest abundances and the biggest sizes were observed on the range of 20–50 m depth. The role as carbon and nitrogen sink of this suspension feeder has been quantified coupling distribution data with existing in situ studies of feeding and respiration. Along the 1.24 ha of the study area, H. papillosa yearly ingested 519.4 g C and 31.4 g N and retained 20.2 g C. As long as the physiological data are known, this new methodology could be very useful in assessing bentho–pelagic links and the capacity of being C and N sinks of a wide range of species. This new approach may be essential for the future management of benthic communities.     

Temporal patterns of settlement, recruitment and post-settlement losses in a rocky reef fish assemblage in the South-Western Mediterranean Sea

Post-settlement events can significantly alter the density distribution of settlers and subsequently the adult population structure. The temporal and inter-annual variability of settlement and the effects of mortality on recruitment were investigated across 2 years in the vicinities of Cabo de Palos–Islas Hormigas Marine Reserve by visual census and light trap sampling. Settlement was seasonal with greater species richness and abundance in summer. Although temporal synchronization was observed between larval supply and settlement, densities of settlers could not be predicted from post-larval abundances. Timing of settlement was consistent between years but with high inter-annual variation in abundance. High mortality (~80 %) and general decoupling between post-larval and settlement stages suggest that early mortality is driving such patterns. Nevertheless, indications of habitat-mediated mortality were found for benthic species such as Symphodus and Diplodus, highlighting the importance of habitat in shaping population demography.     

Population structure and connectivity of the European conger eel (Conger conger) across the north-eastern Atlantic and western Mediterranean: integrating molecular and otolith elemental approaches

Genetic variation (mtDNA) of the European conger eel, Conger conger, was compared across five locations in the north-eastern Atlantic (Madeira, Azores, South Portugal, North Portugal and Ireland) and one location in the western Mediterranean (Mallorca). Genetic diversity of conger eel was high, and differentiation among regions was not significant. Additionally, comparisons of element:Ca ratios (Sr:Ca, Ba:Ca, Mn:Ca and Mg:Ca) in otolith cores (larval phase) and edges (3?months prior to capture) among the Azores, North Portugal, Madeira and Mallorca regions for 2?years indicated that variation among regions were greater for edges than cores. Therefore, while benthic conger may display residency at regional scales, recruitment may not necessarily be derived from local spawning and larval retention. Furthermore, data from otoliths suggest a separated replenishment source for western Mediterranean and NE Atlantic stocks. The combination of genetics and otolith chemistry suggests?a population model for conger eel involving a broad-scale dispersal of larvae, with limited connectivity for benthic juvenile life stages at large spatial scales, although the existence of one or multiple spawning grounds for the species remains uncertain.     

Seasonal and annual variability in an epifaunal community in the German Bight

While the benthic infauna of the North Sea has been studied intensively over the past decades, few studies have focused on the larger mobile epifauna. Studies carried out to date have described the distribution of epifaunal communities over the whole of the North Sea, but variability within the identified communities, which occurs on a much smaller scale, has so far remained unstudied. This is the first study to describe seasonal and annual variability of an epifaunal assemblage in the German Bight area, where environmental conditions are highly variable. The benthic community was sampled with a 2 m beam trawl from 1998 to 2001. The echinoderms Ophiura albida and Asterias rubens and the crustacean Pagurus bernhardus were the dominant species caught throughout the study period. Overall the species composition of the catches was relatively consistent, while abundances of dominant species fluctuated considerably between sampling periods. Differences between sampling periods were not only influenced by the abundances of dominant species, but also by less dominant species such as Ophiura ophiura, Astropecten irregularis, Corystes cassivelaunus, Crangon crangon and Aphorrahis pespelicani. The abundances of these species varied annually and seasonally in the assemblage. Clear differences between summer and winter in the species composition, abundance and biomass were identified. Annual and seasonal changes were most likely linked to migratory movements of epifauna into and out of the area under investigation. Temporal changes in species composition and abundance correlated best with water temperatures, while the spatial distribution of the total biomass over the whole sampling period was correlated with sediment characteristics. Anthropogenic influences such as fishing activity and chronic large-scale eutrophication are thought to have influenced the community on a long-term basis, but have been considered unlikely causes for the short-term variability described by this study.Communicated by J.P. Thorpe, Port Erin     

Growth parameters of deep-water decapod crustaceans in the Northwestern Mediterranean Sea: a comparative approach

 Relative and absolute growth were studied in 17 species of deep-water decapod crustaceans, spanning nine families of six different infra-orders, in the Northwestern Mediterranean Sea. The overall maximum abundance of these species lay between 200 m and 750 m (i.e. upper- and mid-slope species). Relative and absolute growth rates were compared by contrasting the slopes of the size–weight relationships for the different species and calculating the von Bertalanffy growth-equation parameters asymptotic length (L _∞ ) and growth rate (k). The size–weight relationships differed significantly as function of the species' life habits. The results revealed a significant decrease in weight relative to size in mesopelagic species (which carry out diel vertical migrations), an almost isometric relationship between size and weight in the less mobile nektobenthic species, and a significant increase in weight relative to size in strictly benthic species. The mean allometric coefficient for each group increased significantly from mesopelagic to benthic species. However, no general trend was observed in the growth-performance index, Φ (an index used to compare absolute growth rates between species, as a function of habit and depth of maximum abundance for all species combined), suggesting that the deep-water decapod crustaceans studied have similar absolute growth rates. Nevertheless, comparison of growth-parameter and growth-performance index values within families did reveal differences. Mesopelagic species of the families Sergestoidae and Pasiphaeidae showed slightly increased growth rates with increasing depth of distribution. Nektobenthic species of the genus Plesionika followed a trend opposite to that shown by mesopelagic species, with a higher growth rate for the shallowest-dwelling species (P. heterocarpus) than the deepest-dwelling species (P. acanthonotus). Taking growth as one of the major components of an organism's energy budget, the growth rates for the decapod crustacean species in this study were significantly lower than those reported in the literature for shallow-water penaeid crustacean species (which are distributed in higher-temperature habitats than deep-water Mediterranean crustaceans) and higher than those reported for mesopelagic myctophid fish species. Hence, the well-defined growth trends shown by deep-water decapod crustacean species in the Northwestern Mediterranean Sea, compared to the less well-defined trends in the other taxa, is discussed in the framework of the overall dynamics of their ecosystem. Received: 25 May 1998 / Accepted: 27 September 1999     

Life style and genetic variation in teleosts: the case of pelagic (<Emphasis Type="Italic">Aphia minuta</Emphasis>) and benthic (<Emphasis Type="Italic">Gobius niger</Emphasis>) gobies (Perciformes: Gobiidae)

The pattern of genetic variability of two species of Mediterranean gobiids was compared, with reference to their different life history traits (Aphia minuta paedomorphic and pelagic; Gobius niger metamorphosed and benthic). The aim was to evaluate how different life histories can affect the genetic structure in these marine teleosts. The study was carried out on populations of both species sampled in the western Mediterranean and in the Adriatic Sea. Seven restriction endonucleases were used for the RFLP analysis of a mitochondrial DNA segment comprising the NADH dehydrogenase subunits 3, 4L and 4. The results highlighted two different patterns of genetic variation, a weak genetic structure in A. minuta and population subdivision in G. niger. These observations may be explained not only in terms of the different dispersal capabilities of these species, but also considering that A. minuta is an abbreviate iteroparous spawner while G. niger is a protracted iteroparous spawner. Because abbreviate iteroparity is a reproductive strategy selected in stable environments with high resource availability, Pliocene and Pleistocene climate oscillations may have represented factors that negatively influenced the reproductive success of A. minuta, producing demographic fluctuations and bottlenecks, as suggested by the mismatch distribution analysis. The weak genetic structure of A. minuta populations seems to be therefore due to a more recent re-colonization of the Mediterranean basin after a severe population decline, rather than to the high vagility of this pelagic goby.     

Heterogeneous energetic pathways and carbon sources on deep eastern Mediterranean cold seep communities

Cold seep communities in the Mediterranean Sea have only been discovered two decades ago, and their trophic ecology has been the subject of very few studies. We investigated the benthic food web of two deep chemosynthesis-based ecosystems on the Napoli and Amsterdam mud volcanoes (MVs) in the eastern Mediterranean Sea (~2,000 m depth). Seeping methane has been detected at the surface of both MVs during pioneering cruises and has been hypothesised to be assimilated by benthic fauna as observed in other oceans’ margins. Given the extreme oligotrophic character of the eastern Mediterranean Sea, we a priori expected that chemosynthetic food sources, especially methane-derived carbon (MDC), played a major trophic role in these deep seep communities relative to what has been observed in other seep systems worldwide. We aimed at unravelling the trophic relationships on Napoli and Amsterdam MVs through the analysis of carbon, nitrogen and sulphur isotopes both in the dominant benthic invertebrates including the small endofauna (300 μm < size < 1 cm) and in the sedimented organic matter. In particular, we assessed the fraction of MDC in the tissue of several heterotrophic and symbiotic species. Low mean δ³⁴S and δ¹³C values (0.4 ± 4.8‰ and −31.6 ± 5.7‰, respectively) obtained for mega- and macrofauna suggested that the investigated benthic food webs are virtually exclusively fuelled by carbon of chemosynthetic origin. A few grazer invertebrates (δ³⁴S up to 11‰) depart from this trend and could complement their diet with sedimented and decayed phytoplanktonic organic matter. Faunal δ¹³C values indicated that the oxidation of sulphur is likely the predominant energetic pathway for biosynthesis on both MVs. Nevertheless, mytilid bivalves and small capitellid, ampharetid and spionid polychaetes were ¹³C-depleted (δ¹³C < −37‰) in a way indicating they assimilated a significant portion of MDC. For these later heterotrophic species, MDC ranged between 21 and 31% (lower estimates) and 97 and 100% (upper estimates). However, our results highlighted that the origin of assimilated carbon may be complex for some symbiotic species. The vestimentiferan tubeworm Lamellibrachia sp., which exclusively depends on its sulphur-oxidising endosymbionts, showed a ~20‰ inter-individual δ¹³C variability on a very small spatial scale (<1 m) at the summit of Napoli MV. This mostly reflects the variable isotopic composition of pore-water-dissolved inorganic carbon (DIC) and evidenced that tubeworms (and subsequently their endosymbionts) uptake DIC derived from multiple methane oxidation processes in varying proportions. The lower and upper MDC estimates for the vestimentum of Napoli’s individuals were 11–38 and 21–73%, respectively. Finally, data on trophic ecology of Napoli and Amsterdam MVs clearly corroborate previous geophysical results evidencing the spatial heterogeneity of Mediterranean MV environmental conditions.     

Genetic structure of the flounders Platichthys flesus and P. stellatus at different geographic scales

The genetic structure of the flounders Platichthys flesus L. and P. stellatus Pallas was investigated on different spatial scales through analysis of allozyme variation at 7 to 24 polymorphic loci in samples collected from different regions (Baltic Sea, North Sea, Brittany, Portugal, western Mediterranean, Adriatic Sea, Aegean Sea and Japan) in 1984 to 1987. No geographic variation was evident within a region. Some pattern of differentiation by distance was inferred within the Atlantic, while the Mediterranean comprised three geographically isolated populations and was itself geographically isolated from the Atlantic (fixed allele differences at up to three loci were found among P. flesus populations from the Atlantic, the western Mediterranean, the Adriatic Sea, the Aegean Sea and also P. stellatus from the coast of Japan). Sea temperature during the reproductive period probably acts as a barrier to gene flow between populations. Genetic distances among European flounder populations (P. flesus) were higher than, or of the same magnitude as, the genetic distance between Pacific (P. stellatus) and European flounder populations, suggesting that P. flesus is paraphyletic and/or there is no phylogenetic basis to recognising P. stellatus as a different species. The divergence between P. flesus and P. stellatus was thus inferred to be more recent than the divergence between the present P. flesus populations from the NE Atlantic and eastern Mediterranean. The eastern Mediterranean populations are thought to originate from the colonisation of the Mediterranean by a proto-P. flesus/P. stellatus ancestor, whereas the present western Mediterranean population has undergone a more recent colonisation event by P. flesus. Patterns of mitochondrial DNA variation, established on a smaller array of P. flesus samples, were in accordance with the geographic patterns inferred from the allozyme survey. In addition, they supported the hypothesis of a two-step colonisation of the western Mediterranean. These results contribute to our understanding of the biogeography of the Mediterranean marine fauna, especially the group of boreal remnants to which P. flesus belongs. Received: 7 February 1997 / Accepted: 26 March 1997     

Supralittoral amphipod abundances across habitats on Mediterranean temperate beaches

Community complexity and abiotic conditions are key components of environmental heterogeneity that affect the abundance and distribution of species. In this study we evaluated how environmental conditions affect abundances of supralittoral amphipods (Talitridae) in four habitats (sandy beach, rivermouth, wrack and lakeshore), along the Italian peninsula in the Mediterranean Sea. All samplings covered a 12 month period, and used the same sampling methodology thereby enabling comparison of abundances and species composition and richness. Four species (Talitrus saltator (Montagu, 1808), Orchestia gammarellus (Pallas, 1766), O. montagui Audouin 1826, O. cf. cavimana Heller 1865) were collected in the different habitats, but most species were found or were abundant in only one of the four habitats. Abundances of talitrids (numbers per sampling hour) differed significantly among the habitats with highest abundances found in the wrack and on the riverbank in proximity to an estuary, and lowest abundances observed on four sandy beach sites. Environmental conditions (temperature, moisture, substrate penetrability) differed among the habitats and were associated with some of the among-site variability in abundances. Our findings demonstrate that talitrids thrive better in some supralittoral habitats than others, and that some habitats could be considered to be “hotspots” of talitrid ecology and biodiversity.     

Patterns of distribution and composition of sea urchin assemblages on Brazilian subtropical rocky reefs

Sea urchins are a key group of herbivores in both temperate and tropical food webs because they control macroalgal cover, and consequently influence primary productivity and phase shifts on reefs. Despite being abundant on southwestern Atlantic reefs, sea urchin distributions, and their association with abiotic and biotic variables, are poorly known. In this study, sea urchin assemblages were surveyed in 2011 at multiple depths at eight sites in Arraial do Cabo (Brazil, 22°57′S/41°01′W), with sites split between a colder, more wave-exposed location, and a warmer, more sheltered location. The influence of this large-scale physical gradient, along with changes in depth and substrate complexity, on sea urchin densities was then investigated. Predator biomass was low and did not vary significantly among sites. Among the seven species recorded, Paracentrotus gaimardi, Echinometra lucunter and Arbacia lixula were dominant. Linear mixed-effects models indicated that location was important, with mid-sized P. gaimardi individuals and A. lixula more common at cooler, exposed sites and E. lucunter more abundant at warmer, sheltered sites. Sea urchin densities typically decreased with increasing depth, probably caused by changes in factors such as light, wave exposure, and sedimentation. Substrate complexity had a positive effect on the abundance of all species, presumably because of the increased availability of refuges. Physical gradients have important consequences for urchin distributions and their ecological functions at relatively small spatial scales on these reefs, and should be incorporated into herbivore monitoring programmes. Research is also required to examine how differential sea urchin distributions affect benthic dynamics.     

Life-history and demographic spatial variation in Mediterranean populations of the opportunistic polychaete <Emphasis Type="Italic">Ophryotrocha labronica</Emphasis> (Polychaeta,Dorvilleidae)

The spatial scale of life-history and demographic variation was investigated in the opportunistic polychaete Ophryotrocha labronica La Greca and Bacci. Individuals were collected along the Italian coasts from three thermally different biogeographical regions of the Mediterranean Sea. For each region, populations from four harbours were considered, and for each harbour, two sites were examined. Life-history and demographic traits were investigated after one generation under a common garden experiment, and their variation at the three spatial scales was assessed. All the traits showed high variability with regard to site. A number of life-history and all demographic traits also varied according to the biogeographical region. Conversely, no differences were found between harbours, suggesting that geographical isolation did not contribute to phenotypic variation. Results confirmed the central role of local conditions for the evolution of life history in species colonizing heterogeneous environments, but they also pointed to the importance of large-scale factors in shaping the phenotypic responses of O. labronica, demonstrating the need for a multi-scale approach for obtaining a good measure of natural variation in widespread opportunistic species.     

Atlantic-Mediterranean and within-Mediterranean molecular variation in <Emphasis Type="Italic">Coris julis</Emphasis> (L. 1758) (Teleostei,Labridae)

Sequence variation in the mitochondrial control region was studied in the Mediterranean rainbow wrasse (Coris julis), a species with pronounced pelagic larval phase inhabiting the Mediterranean Sea and the adjacent coastal eastern Atlantic Ocean. A total of 309 specimens from 19 sampling sites were analysed with the aim of elucidating patterns of molecular variation between the Atlantic and the Mediterranean as well as within the Mediterranean Sea. Phylogeographic analyses revealed a pronounced structuring into a Mediterranean and an Atlantic group. Samples from a site at the Moroccan Mediterranean coast in the Alboran Sea showed intermediate frequencies of “Mediterranean” and “Atlantic” haplotypes. We recognised a departure from molecular neutrality and a star-like genealogy for samples from the Mediterranean Sea, which we propose to have happened due to a recent demographic expansion. The results are discussed in the light of previous studies on molecular variation in fish species between the Atlantic and the Mediterranean and within the Mediterranean.     

Vertical distribution of Calanus finmarchicus and C. helgolandicus in relation to the development of the seasonal thermocline in the Celtic Sea

The geographical distribution and annual mean abundance of Calanus finmarchicus (Gunnerus) and C. helgolandicus (Claus) in the northern North Atlantic Ocean were shown in relation to the seasonal and annual fluctuations of abundance of the species in the Celtic Sea from 1960 to 1981. These congeneric copepods, although showing allopatric distributions over most of their geographical range, have sympatric distributions in the Celtic Sea where they dominate the dry weight biomass of the plankton throughout the year. The two species respond differently to the development of the seasonal thermocline and halocline by taking up different vertical distributions in the water column. C. finmarchicus occurred in the colder, more saline water below the thermocline, while C. helgolandicus occurred in the warmer, less saline water above the thermocline. This behaviour is postulated as a mechanism by which these morphologically similar copepods more fully exploit the resources of their temporally and spatially heterogeneous environment and also minimise interspecific competition. The species have the same foraging techniques and are able to exploit the same size spectrum of particulates. The vertical depth strata in which the populations are found for most of the year in the Celtic Sea means that both species exploit the diatom bloom in early spring but, thereafter, C. helgolandicus grazes on the daily production of the autotrophs in the euphotic zone while C. finmarchicus, below the thermocline, has to rely for its food on sedimenting particulates (whole cells, detritus and faecal material). The isolating mechanisms whereby these two populations partition the habitat in the Celtic Sea are discussed.     

Population dynamics of Eudendrium racemosum (Cnidaria, Hydrozoa) from the North Adriatic Sea

Benthic suspension feeders in shallow waters develop in relation to the food availability and the variation of physical parameters giving rise to complex communities that act as a control factor on the plankton biomass. The aim of the work is to establish the role of the hydrozoan Eudendrium racemosum in the energy transfer from the plankton to the benthos in marine food chains of the North Adriatic Sea. This study highlighted that the hydroid biomass changed over time in relation to temperature and irradiance, and the highest abundance was observed during summer with about 400,000 polyps m⁻² (about 19 g C m⁻²). The population suffered an evident summer decrease in relation to a peak of abundance of its predator, the nudibranch Cratena peregrina, whose adult specimens were able to eat up to 500 polyps day⁻¹ and reached an abundance of 10 individuals m⁻². The gut content analysis revealed that the hydroid diet was based on larvae of other benthic animals, especially bivalves and that the amount of ingested preys changed during the year with a peak in summer when it was estimated an average predation rate of 13.7 mg C m⁻² day⁻¹. In July, bivalves represented over 60 % of the captured items and about 18 mg C m⁻² day⁻¹. Values of biomass of E. racemosum are the highest ever recorded in the Mediterranean Sea, probably supported by the eutrophic conditions of the North Adriatic Sea. Moreover, our data suggest that settling bivalves provide the greater part of the energetic demand of E. racemosum.     

Short- and long-term importance of small sharks in the diet of the rare deep-sea shark Dalatias licha

Knowing the trophic ecology of marine predators is essential to develop an understanding of their ecological role in ecosystems. Research conducted on deep-sea and threatened shark species is limited. Here, by combining analyses of individual stomach contents and stable isotope values, we examined the trophic ecology (dietary composition and trophic position) of the kitefin shark Dalatias licha, a deep-sea shark considered as near threatened globally and as data deficient in the Mediterranean Sea. Results revealed the importance of small sharks in the diet of the kitefin shark at short- and long-term scales, although fin-fish, crustaceans and cephalopods were also found. Predation on sharks reveals the high trophic position of the kitefin shark within the food web of the western Mediterranean Sea. Stable isotope values from liver and muscle tissues confirmed our results from stomach content analysis and the high trophic position.     

Phylogeography of Cerastoderma glaucum (Bivalvia: Cardiidae) across Europe: a major break in the Eastern Mediterranean

Molecular variation of the lagoon cockle Cerastoderma glaucum (Poiret, 1789) was examined across the species range along European coasts, from the northern Baltic to the Black Sea and the Caspian Sea. A major phylogeographic break in mitochondrial COI gene sequences (divergence 6.2%) separated a group of Ponto-Caspian and Aegean Sea haplotypes from those to the west of the Peloponnese peninsula in the Mediterranean and in the Atlantic-Baltic sector. A similar subdivision, yet not entirely geographically coincident, was found at 16 allozyme loci (Nei's distance D=0.15). The results imply a long-term isolation of populations in parts of the Eastern Mediterranean or Black Sea basins through the Pleistocene. The subdivision does not concur with previous views on the systematics of C. glaucum complex, but the pattern is notably similar to that described earlier from some fish species. Marked phylogeographic structuring was also found at lower level within the major Mediterranean-Atlantic phylogroup of C. glaucum, which was divided into six regional or local haplotype subgroups. Divergence of these groups may date back one or several major Pleistocene climatic cycles. Local mtDNA diversity was particularly high in a sample from the Ionian Sea, whose mitochondrial identity was of the Mediterranean-Atlantic type, while its nuclear characters were more strongly associated with the Ponto-Caspian type. Patterns of shallower, star-phylogeny type diversity within the Ponto-Caspian phylogroup and in the Baltic Sea area may represent more recent, post-glacial generation of variation.     

Benthic macrofaunal compositional variations in the northern Bering Sea

Point sampling of soft-sediment macrofauna provided a regional-scale analysis of species composition, diversity, and demographic distributions in the northern Bering Sea (61°0??C65°30??N Lat) in spring 2006. Taxonomic differences distinguished subregions north and south of St. Lawrence Island (P?<?0.1%). High compositional variability occurred within and between sites, with higher variability among northern sites. One to two species dominated most assemblages, with moderately common or rare species. Ampeliscid amphipods characterized the north, and bivalves the south. Bivalves (Macoma calcarea Gmelin, Nuculana radiata Krause, Ennucula tenuis Montagu) were commonly widespread, in varied abundances and sizes. Overall, the extensive soft-sediment habitat supports a varied fauna composition with location, in species dominance, abundance, and size-class. This spatially heterogeneous regional structure is important in assessing ecosystem performance related to secondary production, biogeochemistry, bioturbation, top predator feeding strategy, recruitment, and in measuring change.