Long-term trend in substratum occupation by a clonal, carbonate bryozoan in a temperate rocky reef in times of thermal anomalies

Studies over time provide opportunities to detect variations in the spatial and temporal patterns of clonal organisms and measure changes on their population dynamics related to extreme events. We assessed population dynamics for a bryozoan species dominating a subtidal rocky reef at Tino Island, in the eastern Ligurian Sea (NW Mediterranean). Using 9 years of annual photosurveys (1997–2005), rapid decline in Pentapora fascialis colony cover was shown at 11 and 22 m depths following the anomalous warming events in 1999 and 2003. An 86 % reduction in live colony portion was found after the 1999 warming event (2.3 °C higher than normal), with larger colonies being most affected. Effects from the 2003 event were delayed, and gradual cover decline occurred during the following 2 years. At the “Shallow” photostations, none of the larger colonies (>1,000 cm²) survived after the first cover decline. Availability of new substrate after the 1999 disturbance resulted in enhanced recovery through new colony production. At the “Deep” photostations, the population structure did not change over the duration of the monitoring period showing the same monomodal structure and same dominant size class (50–500 cm²). In the 4 years following the first cover decline, the deeper population regained colony cover to levels similar to pre-disturbance level, showing a good resilience. This 9-year monitoring analysis provided the temporal resolution needed to detect changes occurring in the P. fascialis population and will contribute to the assessment of long-term changes on benthic populations suffering during recent decades from dramatic increases in extreme events.     

Aspects of the biology of the bryozoan Pentapora fascialis in the northwestern Mediterranean

Colonies of the bryozoan Pentapora fascialis (Pallas) were monitored photographically from January 1993 to March 1994, in two quadrats at depths of 11 and 22 m at Tino Island (Ligurian Sea, Italy). Colony size, shape, growth and mortality were analysed. Zoarial growth was measured as change in the two-dimensional area of substratum covered per unit time. Colonies ranged in size from less than 30 to 2760 cm². Colony shape was normally depressed globular, with an elliptic or subcircular perimeter, but tended to elongate in crowded situations. Although extremely variable in small colonies, growth was inversely related to colony size and was maximum in spring. Highest net growth rate was 211% in 11 months. No recruitment was observed in the quadrats during the study period. Partial mortality in colonies was low and was mainly due to overgrowth by algae and to siltation. Physical disturbance by an exceptionally severe winter storm in December 1993 caused total mortality of almost all colonies in the shallow quadrat. Received: 24 March 1997 / Accepted: 22 November 1997     

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     

Genetic relationships between marine and marginal-marine populations of Cerithium species from the Mediterranean Sea

Results of isozyme electrophoresis were used to explore the genetic relationships between several Mediterranean morphs of Cerithium (Gastropoda: Prosobranchia), for which taxonomy is currently uncertain because of high intraspecific variability and low interspecific differentiation. The large species, classically known as C. vulgatum Bruguière, 1789 was identified at four sites (two in the French Mediterranean and two in southern Spain). Two different larval types were found in the French sites, but poecilogony could not be demonstrated. Individuals collected from harbours were not genetically distinct from open-sea populations of classic C. vulgatum. However, a population in the Embiez lagoon (French Mediterranean) which morphologically resembles C. vulgatum did display distinct genetic traits, supporting its status as a separate species. Of the small Cerithium species usually known as C. rupestre, two sympatric species (C.“rupestre” Risso, 1826 and C. lividulum Risso, 1826) were distinguished. Genotype frequencies within the analysed populations revealed much heterozygote deficiency. F _ST values (fixation index measuring the effects of population subdivision) suggest a higher genetic differentiation for C. lividulum populations than for C. vulgatum populations. We assume that a high larval dispersal capability (via planktotrophy) allows a high gene flow between populations of C. vulgatum. Received: 24 November 1998 / Accepted: 24 September 1999     

Population dynamics of the invasive bryozoan Membranipora membranacea along a 450-km latitudinal range in the subarctic northwestern Atlantic

Thermal environment is often regarded as a key determinant of distribution limits in marine invertebrates and hence may represent one of the most important barriers to invasion by non-indigenous species. For the first time in the subarctic northwestern Atlantic, we investigated variation in the timing and magnitude of settlement, recruitment, and colony cover of the recently (early 2000s) introduced bryozoan Membranipora membranacea on the kelp Saccharina longicruris and how this variation relates to changes in sea temperature (thermal histories) across eight sites spanning a 450-km latitudinal range between southwestern Newfoundland and southeastern Labrador, Canada. We show that (1) up to 61 % of the variation in settlement, recruitment, and colony cover was explained by sea temperature alone, with highest and lowest abundances at warmest and coldest sites, respectively; (2) between-site differences in rates of sea cooling explained 85 % of the variation in settler abundance; (3) varying the temporal window over which data were aggregated increased the explanatory power of sea temperature to as much as 98 % for settlement and recruitment, and 86 % for colony cover; (4) exposure to waves and surface area of colonies improved relationships between sea temperature and settlement and recruitment by up to 11 %; and (5) recruit abundance was a strong predictor of colony cover, explaining as much as 89 % of the variation. Consistently low abundances of settlers and recruits at the northern tip of Newfoundland and southern tip of Labrador suggest that M. membranacea is nearing its northern distribution limit in the northwestern Atlantic. Our findings extend knowledge of population dynamics of M. membranacea in the northwestern Atlantic, while highlighting the complexity of the interactions between physical and biological factors and processes that affect population dynamics in invertebrates with planktonic larvae in predominantly cold marine habitats.     

The influence of seawater temperature on zooid size and growth rate in Pentapora fascialis (Bryozoa: Cheilostomata)

Zooid size variation and growth banding in the bryozoan Pentapora fascialis, collected from the South of Great Britain to the South of Italy, were investigated in order to test their relationship with the variation of seawater temperature. Zooid length appeared to be more sensitive than zooid area and width to temperature variation. The longest zooids were budded in localities that typically experience low seasonal variation in temperature (3.2–7.5°C) and low mean annual temperature (about 11°C). Zooid length demonstrated the strongest relationship with the range of seasonal variation in temperature rather than the mean annual temperature. Growth bands produced during colder months were less high than those produced during warmer months. Measurements of these alternating bands were used to calculate growth rate of the laminae, ranging from 20.7 at Plymouth to 29.3 mm year⁻¹ at Tino Island. Significant differences in zooid length were found between the two bands, with longer zooids within the band developed in colder months and shorter zooids in bands produced in the warmer period. We conclude that zooid size can be used as an environmental proxy of the seasonal temperature regimes experienced by the species.     

Population structure of the common sole (<Emphasis Type="Italic">Solea solea</Emphasis>) in the Northeastern Atlantic and the Mediterranean Sea: revisiting the divide with EPIC markers

Spatial and temporal population genetic structures of the common sole, Solea solea, were studied in Northeastern Atlantic and Mediterranean Sea populations, using three polymorphic exon-primed intron-crossing (EPIC) markers. Results demonstrated significant multilocus differentiation among Eastern Mediterranean and a group composed by Western Mediterranean and Atlantic populations (θ = 0.150, P < 0.001), but also suggested unrecorded genetic differentiation of the Adriatic Sea population. No pattern of isolation-by-distance was recorded across the range covered by sampling, from the Kattegat to the Aegean Sea. Conversely to genetically structured Mediterranean populations, Atlantic populations ranging from Denmark to Portugal could be considered as representative of the same panmictic unit (θ = 0.009, not significant). Results further demonstrated stability of multilocus genetic structure among temporarily replicated cohort samples [0+, 1+, subadults] from several coastal and estuarine locations from Bay of Biscay, excepted for the amylase locus Am2B3-2 at one location (Pertuis d’Antioche). Despite coherence of such observed patterns of multilocus differentiation with previous allozymic surveys in sole, and with patterns generally obtained for other marine fish species, single-locus results from EPICs indicated divergent coalescence schemes supporting a complex response to ecology and history of sole’s populations. Results stress the use of nuclear genes such as EPIC markers to investigate population structure, but also historical, demographic, and possibly selective processes in marine fishes.     

Spatial–temporal variability of epibenthic assemblages on subtidal biogenic reefs in the northern Adriatic Sea

Spatial distribution and temporal variation of epibenthic assemblages of coralligenous biogenic rocky outcrops occurring in the northern Adriatic Sea (45°04′–45°24′N; 12°23′–12°43′E) were investigated by photographic sampling from 2003 to 2006 at 12 randomly selected sites. The dominant reef-forming organisms were the encrusting calcareous algae (Lithophyllum stictaeforme, Lithothamnion minervae and Peyssonnelia polymorpha), while the main bioeroders were boring sponges (Cliona viridis, C. celata, C. thoosina, C. rhodensis, Piona vastifica) and the bivalve Gastrochaena dubia. Composition of the assemblages varied thorough years and among sites. Spatial heterogeneity, at local and regional scale, prevailed over temporal variation. This variability was related both to the geo-morphological features of the outcrops and to environmental variables. Sites clearly differed in the percent cover of reef builder and bioeroder species while only limited temporal variation within site was found. Some taxa revealed complex intra-site temporal trends. These results provide valuable information on the diversity and variability of epibenthic assemblages of the northern Adriatic coralligenous reefs, essential for the management and conservation of these unique biogenic habitats.     

Phylogeography of two species of <Emphasis Type="Italic">Lysidice</Emphasis> (Polychaeta,Eunicidae) associated to the seagrass <Emphasis Type="Italic">Posidonia oceanica</Emphasis> in the Mediterranean Sea

The aim of this work was to define the phylogeographic patterns of the two species of polychaete Eunicidae, Lysidice ninetta Audouin and Milne Edwards and Lysidice collaris Grube, both associated as sheath borers to the Mediterranean seagrass Posidonia oceanica, and with reference to their different origin and their actual geographic distribution. L. ninetta is distributed in the Atlantic and in the Mediterranean Sea while L. collaris is a tropical species, whose introduction into the Mediterranean Sea through the Suez Canal (lessepsian migrant) has been hypothesized in recent years. The two species have been often confused in the past, although they appear morphologically distinct. They share the same microhabitat (Posidonia sheaths) and they co-exist along a broad bathymetric range (1–30 m). Several populations for both taxa were sampled all along the coast of the Mediterranean basin. A variable no coding region of nuclear DNA (rDNA, ITS1) and a portion of a more conservative coding region of mitochondrial DNA (sub-unit one of citochrome oxidase, COI) were used as molecular markers. Both markers confirmed the separation between the two species. Low intraspecific polymorphism was present in L. collaris, together with absence of phylogeographic structure. In L. ninetta, instead, the presence of intraspecific cryptic lineages, sympatric in some sites, was recorded. Clustering of single populations in the two main clades was not always consistent between markers. The mitochondrial COI region showed more resolution at the given spatial scale. Our results suggest that Lysidice collaris could be recently introduced into the Mediterranean Sea from one or more separate events. On the other hand, for L. ninetta one could presume a re-colonization of the Mediterranean basin from the Atlantics, after the Messinian crisis (dry-out of the Mediterranean Sea, 5.5 my) with the subsequent separation of intraspecific lineages. The phylogeographic patterns of both Lysidice spp. are disjoined with respect to that of the host plant, P. oceanica. The obtained results suggest that environmental constraints and evolutionary history of these polychaetes and their host plant act in different ways to determine their actual genetic spatial structure.     

Survey of deep-dwelling red coral (<Emphasis Type="Italic">Corallium rubrum</Emphasis>) populations at Cap de Creus (NW Mediterranean)

The distribution and population structure of the eurybathic gorgonian Corallium rubrum were studied off Cap de Creus (Costa Brava, Northwestern Mediterranean Sea). Red coral is endemic to the Mediterranean Sea and the adjacent NE Atlantic coast, where it has been over exploited for centuries. This study presents, the first quantitative data on the spatial distribution and structure of a population extending between 50 (common SCUBA limits) and 230 m depth, and compared it with shallow populations previously studied in the same area. Different remotely operated vehicles (ROV) and two methodological approaches were employed during four cruises between 2002 and 2006: 1-Extensive surveys: sea to coast transects in which red coral density and patch frequency were recorded; 2-Intensive surveys, in which parameters describing colony morphology were recorded. Most of the hard substrate between 50 and 85 m depth was inhabited by red coral colonies, showing a patch frequency of 8.3 ± 7.9 SD patches per 100 m-transect (total transect area: 34 m²), and within-patch colony densities of 16–376 colonies m⁻² (mean of 43 ± 53 colonies m⁻²). Below 120 m depth red coral was less abundant, and rather than forming dense patches as in shallow water, isolated colonies were more common. The population structure differed between sites that are easily accessible to red coral fishermen, and remote ones (both at similar depth, 60–80 m), as colonies in easily accessible locations were smaller in height and diameter, and showed a less developed branching pattern. At shallower locations (10–50 m depth) the population structure was significantly different from those at deeper locations, due to the heavy harvesting pressure they are exposed to in the shallows. Twenty-five to forty-six percentage of the deeper colonies were taller than 6 cm, while only 7–16% of the shallow water colonies exceeded 6 cm colony height. Forty-six to seventy-nine percentage of the colonies in deeper waters were large enough to be legally harvested, while only 9–20% of the shallow water colonies met the 7 mm legal basal diameter to be collected. The branching pattern was also better developed in deeper colonies, as up to 16% of the colonies showed fourth order branches, compared to less than 1% of the shallow water colonies (of which 96% consisted of only one single branch). The results thus confirm that C. rubrum populations above 50 m depth are exposed to a higher harvesting intensity than deeper populations in the same area.     

Genetic variability and relationships among geographically widely separated populations of Petitia amphophthalma (Polychaeta: Syllidae). Results from RAPD-PCR investigations

An analysis of the population genetics of the meiofaunal polychaete Petitia amphophthalma Siewing, 1956, in which the RAPD-PCR method was applied to 103 individuals from eight populations, some of them very far apart (Atlantic: Florida, Tenerife, France; Mediterranean: two Greek islands, Tunisia; Red Sea: Egypt), gave closely reproducible results. In the band patterns produced with 13 decamer primers, a total of 195 genetic characters was detected. The data were evaluated by a number of methods, including the cluster programs UPGMA, WPGMA and neighbour-joining. The detected genetic distances between the populations vary between 58.9 and 66.6, but 97% of the genetic characters, although polymorphic, are found in at least two populations and usually in all the others as well. Phenograms of the analyses find four population clusters [Florida, France (Atlantic), the Mediterranean and Tenerife]. They are, however, not completely congruent and show low bootstrap values at the junction points of the clusters (with the exception of the Tenerife cluster). Mediterranean P. amphophthalma form a cohesive population, although within it the genetic distances are graded in parallel with the geographic distances between the sites. The colonization of Tenerife, an island of relatively recent volcanic origin, can be taken as evidence that this meiofaunal species can become dispersed not only along coastlines but also across expanses of open water. However, the severely restricted variability of these populations implies that in this case a founder effect has operated, and that transport over open water is not a routine event but extremely rare. The absence of the species on the Australian coast and, for instance, on the Galapagos Islands indicates that there has been no continuous gene flow across the oceans. The idea that all the populations investigated belong to one cosmopolitan species is discussed. Received: 17 October 1997 / Accepted: 15 April 1998     

A pelagic bryozoan from Antarctica

Spherical or sub-spherical bryozoan colonies were collected from the surface waters of a coastal polynya in the southeastern Weddell Sea near Halley Station in February 1992. These are the first truly pelagic marine bryozoan colones yet recorded. The collection site is the edge of the Brunt ice shelf, which is between 150 and 250 m thick in this area, and the depth of water to the seabed is -400 m. The colonies were hollow, composed of a single layer of autozooids, and appeared complete and undamaged. They were between 5.0 and 23.0 mm in diameter, were brownish in colour in life, and pale yellowish brown after preservation. Light and scanning electron microscope investigations of the colonies indicated that they belonged to the genus Alcyonidium, and they are here compared with A. flabelliforme Kirkpatrick, a known antarctic benthic species. Both a pelagic existence and hollow spherical colony form are new attributes for the phylum Bryozoa. However, because of the plasticity of form of species belonging to the genus Alcyonidium, these is not enough evidence for the introduction of a new species at this time. The colonies found may represent a previously undescribed juvenile stage of a known Alcyonidium species. It is postulated that these colonies may obtain nutrition from the often abundant populations of ice algae present in the lower layers of permanent sea-ice.     

Phylogeographic patterns of the mysid Mesopodopsis slabberi (Crustacea, Mysida) in Western Europe: evidence for high molecular diversity and cryptic speciation

The phylogeographic patterns among populations of Mesopodopsis slabberi (Crustacea, Mysida), an ecological important mysid species of marine and estuarine habitats, were analysed by means of DNA sequencing of a fragment of the mitochondrial cytochrome c oxidase subunit I (COI) and the 16S ribosomal RNA genes. Samples of M. slabberi collected from five Atlantic and two Western Mediterranean populations were investigated. Very high levels of within-population molecular diversity were observed in all samples (mean h=0.807 and π=0.0083), with exception of the Mediterranean Ebro population which contained only one haplotype. Differentiation among populations was high, and a clear phylogeographic break was observed between the Atlantic and Mediterranean populations. Moreover, a strong differentiation was detected between both populations in the Western Mediterranean Sea (Alicante and Ebro delta), while two divergent lineages occurred in sympatry within the Atlantic Mondego estuary. The high congruence between both the COI and 16S rRNA sequence data, the reciprocal monophyly of the different mitochondrial clades and the levels of nucleotide divergence between them suggest the presence of a complex of cryptic species within M. slabberi. Estimations of divergence time between the different mitochondrial lineages indicate that a split occurred during the late Miocene/early Pliocene. Such a divergence could be concordant with vicariant events during sea-level drops within the Mediterranean region at that time. However, within the Mediterranean Sea, the potential of divergence through ecological diversification cannot be ruled out.     

Effects of duration of larval swimming period on early colony development in Bugula stolonifera (Bryozoa: Cheilostomata)

Laboratory experiments with larvae of the cheilostome bryozoan Bugula stolonifera Ryland, 1960 assessed the time to settlement in the presence of a constantly available polystryrene substrate, the development of competence for metamorphosis, and the effects of the duration of swimming period on early colony development. Sexually mature colonies of B. stolonifera were collected on 11 and 18 September 1987; 2 and 18 August, 1988; and 6, 12, 19, and 26 September 1988, from Eel Pond (Woods Hole, Massachusetts, USA) and were maintained at 20°C. In the presence of a constantly available substrate, cumulative percent settlement curves were sigmoid, with 75% of larvae settled in 3.2±0.5 h. Typically, 50% of the larvae settled in less than 3 h and 95% settled in 6.1±1.2 h. The number of settled individuals that developed feeding ancestrulae by 3 d and the number that developed first-feeding autozooids by 6 d was assessed as a function of duration of larval swimming. Individuals which were kept swimming for 8 and 10 h after hatching developed significantly more slowly to the ancestrula and autozooid stages in 13 out of 14 experiments than did larvae that swam 2 or 6 h. This is the first report for any bryozoan that prolongation of the larval free-swimming period affects the rate of colony development.     

Genetic structure of populations of two species of Chthamalus (Crustacea: Cirripedia) in the north-east Atlantic and Mediterranean

Protein electrophoresis on starch gels was used to investigate population genetic structure of the barnacles Chthamalus montagui Southward and C. stellatus (Poli) over their north-east Atlantic and Mediterranean ranges. In each species, a single locus exhibited marked differentiation of allele frequencies between Atlantic and Mediterranean localities; in C. stellatus, genetic differentiation between the two basins had not previously been noted. In both species, mean heterozygosity per locus appeared higher in the Mediterranean samples than in the Atlantic, and Mediterranean populations had more alleles at the loci studied. Possible explanations for the differentiation between the Atlantic and the Mediterranean populations are discussed. Received: 30 May 1996 / Accepted: 17 September 1996     

Genetic variability in the marine gastropods Patella coerulea and Patella aspera: patterns and problems

Allozymic variation encoded by 15 gene loci was compared and contrasted in naturally coexisting populations of two marine gastropods (Patella coerulea and Patella aspera) collected along the rocky beaches of the Mediterranean sea of Israel in June, 1979. The genetic variability did not differ consistently or significantly between the two species, but in all the populations studied, P. aspera was more heterozygous than P. coerulea. These results are discussed in terms of the potential ecological factors affecting this genetic differentiation.     

Size at maturity,fecundity and reproductive potential of a protected population of the spiny lobster<Emphasis Type="Italic"> Palinurus elephas</Emphasis> (Fabricius, 1787) from the western Mediterranean

Protected lobster populations are expected to contribute to the replenishment of fished populations through increased egg production. We studied the reproductive biology and egg production potential of a population of the spiny lobster Palinurus elephas protected from fishing since 1990 in the Columbretes Islands Marine Reserve (western Mediterranean). An index of spawning potential was derived to compare egg production potential in the Reserve and in western Mediterranean exploited populations. Females' physiological maturity (ability to reproduce) and functional maturity (ability to mate and bear eggs) occurred at a carapace length (CL) of 76–77 mm. Males' physiological maturity occurred at a slightly larger size, 82.5 mm CL. In the Reserve, P. elephas' individual fecundity increases linearly with body size up to the females' maximum size, although maximum reproductive yield (eggs per body gram) was reached at intermediate sizes. Size-specific fecundity in the protected population was similar to that of lightly fished populations off Ireland and greater than that of western Mediterranean exploited populations. The female size class of 105–110 mm CL contributed most to egg production in the protected population and is well above the minimum landing size (MLS) for western Mediterranean fisheries. Newly mature females (below MLS) generate a very small fraction (1%) of the egg production from the Reserve. Given the pattern of exploitation in western Mediterranean fisheries, egg production potential depends more on the quantity than on the mean size and fecundity of the available females. The role of the greater availability of large males for mating in unfished populations is discussed in terms of the females' individual fecundity and mating success.     

Genetic structure and allozyme variation of sea bass (Dicentrarchus labrax and D. punctatus) in the Mediterranean Sea

This paper reports data on 28 allozyme loci in wild and artificially reared sea bass (Dicentrarchus labrax) samples, originating from either coastal lagoon or marine sites in the Mediterranean Sea. F _ST analysis (θ estimator) indicated strong genetic structuring among populations; around 34% of the overall genetic variation is due to interpopulation variation. Pairwise θ estimates showed that, on average, the degree of genetic structuring was much higher between marine populations than between samples from lagoons. Six polymorphic loci showed differences in allele frequencies between marine and lagoon samples. Multivariate analyses of individual allozymic profiles and of allele frequencies suggested that different arrays of genotypes prevail in lagoons compared to marine samples, particularly at those loci that, on the basis of previous acclimation experiments, had been implicated in adaptation to freshwater. On the other hand, variation at “neutral” allozyme loci reflects to a greater extent the geographic location of populations. Allozyme differentiation was also studied in a D. labrax population from the Portuguese coast. Average genetic distance between this population and the Mediterranean populations was quite high (Nei's D = 0.236) and calls into question the taxonomic status of the Portuguese population. Finally, genetic relationships between D. labrax and D. punctatus were evaluated. Average Nei's D was 0.648, revealing high genetic differentiation between the two species, even for two sympatric populations of these species in Egypt; thus gene flow was not indicated between species. Received: 24 October 1996 / Accepted: 27 November 1996     

Temporal and spatial variation in protein,carbohydrate, and lipid levels in <Emphasis Type="Italic">Corallium rubrum</Emphasis> (Anthozoa,Octocorallia)

Red coral (Corallium rubrum) is an important component of the Mediterranean coralligenous habitat. The present study analyzes the relationship between energy storage molecule concentration (i.e. tissue balance of protein–carbohydrate–lipid levels) and the gonadal output in two populations located at different depths (16–18 m and 40–45 m depth) through a seasonal cycle (2002–2003). In June 2003, another six populations have been sampled to compare medium-scale geographical variation. The colonies sampled were less than 6 cm high, thus representing the majority of the population in the studied area. There was no clear relationship between energy storage (i.e. carbohydrate and lipid levels) and gonadal output, nor in the seasonal cycle or the spatial variation. A clear-cut seasonal trend of energy storage was detected only in the shallower population. The water temperature was warmer at 20 m depth and the water transparency (Secchi disk) was also higher (lower seston concentration) in 2003 when the same periods in 2002 and 2003 were compared (May–August). Carbohydrates seemed to be more sensitive to food constraints than lipids. The partial mortality detected in summer 2003 in the sampling area appeared to be recorded in the carbohydrate and lipid levels of this species, especially in the shallow population. The biochemical levels prove to be a useful tool to detect unusual environmental conditions that may cause partial or total mortality in benthic suspension feeders.     

Allozyme variation in European populations of the oyster Ostrea edulis

Variations at 22 enzyme coding loci were surveyed in 11 populations of the oyster Ostrea edulis L., which were sampled between 1988 and 1990 along the Atlantic and Mediterranean coasts of Europe. Atlantic oyster beds suffered a steady decline during the last century, and restocking of beds with oysters of foreign origin has probably resulted in a high degree of interbreeding of natural oyster stocks from all Atlantic Europe. Our study confirms the low levels of genetic variability previously reported for the oyster populations from the Atlantic coasts, and extends it to the Mediterranean coasts. The locus arginine-kinase (ARK ^*) exhibited a high degree of interpopulation differentiation (F _ST=0.289), resulting from extensive variation in gene frequencies along a geographical cline. However, the overall genetic differentiation between populations was slight, and similar to that reported for other local populations of bivalves (mean genetic distance between populations is 0.010, mean F _ST=0.062). A general pattern of increasing differentiation along the coastline in an Atlantic-mediterranean direction emerged; but genetic differentiation among the Atlantic populations was not significantly lower than that observed among the Mediterranean populations. This and other results suggest that the effects of extensive transplantation of oysters among various areas in Europe are detectable only in some particular localities. The geographical distribution of low-frequency alleles suggests a restriction to gene flow outwards from the Mediterranean Sea, across the Straits of Gibraltar.