Fragmented seagrass habitats on the Mediterranean coast, and distribution and abundance of mysid assemblages

Four different habitats were sampled in order to determine the spatial and temporal distribution and abundance of mysids in a seagrass meadow ecosystem. These habitats included meadows of Posidonia oceanica and Cymodocea nodosa, the edge of P. oceanica meadow and sandy substrates. Three random sites for each habitat were sampled along the coast of Alicante at different times, using a hand-net technique (6 m²). Seven species of mysids were identified: Leptomysis posidoniae, Leptomysis buergii, Paramysis helleri, Siriella clausii, Mesopodopsis slabberi, Siriella armata and Mysidopsis gibbosa. Total abundance of mysids was highest at the edge of the P. oceanica meadow, followed by the C. nodosa meadow, sandy substrate and P. oceanica meadow, respectively (P<0.05); this pattern was found during all sampling periods except one. Three species showed persistent spatial segregation by habitat (S. clausii in Posidonia; L. posidoniae at the edge; S. armata in Cymodocea), whereas the other four did not show any clear segregation by habitat (ANOVA, P<0.01). On a site scale (km), spatial variability was very high. Multivariate analyses confirm the existence of differences in the assemblage structure among habitats, with a gradual change from the Posidonia meadow to sand. Posidonia and Cymodocea showed different mysid assemblages. Fragmented, shallow, vegetated habitat on the Mediterranean coast seems to support a higher abundance and species richness of mysids, through habitat diversification, in comparison with homogeneous habitats.     

Algal products as naturally occurring substrates for p-glycoprotein in Mytilus californianus

Mytilus californianus is a filter feeder that removes seaweed particulates, phytoplankton, and their byproducts from the water. The gills of this animal express high multixenobiotic resistance (MXR) and multixenobiotic transport activity that is related to the mammalian p-glycoprotein (p-gp). The high p-gp observed in mussel gills may provide the mussel protection from natural toxins in the diet. To test this hypothesis, extracts of various seaweeds and phytoplankton found in and around the mussel habitat were examined for presence of p-gp substrates by competition assays with a fluorescent dye substrate, rhodamine B. Accumulation of rhodamine was increased whereas dye efflux was slowed in the presence of algal extracts, indicating that p-gp substrates are present in the seaweeds Macrocystis pyrifera, Egregia menziesii, and Phyllospadix scouleri and the phytoplankton Alexandrium catanella and Pseudonitzchia australis. Initial fractionation of E. menziesii extracts by HPLC showed several distinct peaks of moderate hydrophobicity with p-gp-modulating ability. Additionally, Egregia extract showed potential as a chemosensitizer in tests with mussel (Mytilus edulis) and sea urchin (Lytechinus pictus) embryos. These data indicate that marine algae contain compounds that are substrates and/or chemosensitizers for the p-gp transporter in marine bivalves, thus providing evidence that MXR may have evolved in response to dietary pressures.     

Genetic structure of deep-water redfish, Sebastes mentella, populations across the North Atlantic

Patterns of genetic diversity of the marine deep-water redfish, Sebastes mentella Travin, from 21 sampling locations (n=46 per sample, from 1995 to 1998) were examined throughout its range in the North Atlantic. Eight microsatellite loci were used in order to: (1) define the genetic structure and consider the possible influence of dispersal, geographic distance, oceanic currents and historical factors; and (2) relate the structure to present management units and practices. Three divergent population units (mean Š=0.012) associated with distinct geographic areas were defined: Eastern (Norway and Barents Sea), Panoceanic and Western (Gulf of St. Lawrence and offshore Newfoundland). The most important observation was the lack of genetic differences (mean Š=-0.0004) and lack of genetic isolation by geographic distance (r=0.318, P=0.112) among samples within the Panoceanic zone, from the Faroe Islands to the Grand Banks (6,000 km). A combination of vicariance, historical introgression with a closely related species, S. fasciatus, and contemporary patterns of oceanic circulation are likely to have shaped the observed population structure. These results only partially support current management units, and call for more integrated practices for regulating the exploitation of S. mentella throughout its extensive range.     

Patterns of larval and early juvenile growth in a semiterrestrial crab, Armases angustipes (Decapoda: Sesarmidae): comparison with a congener with abbreviated development

In a semiterrestrial and estuarine tropical crab, Armases angustipes Dana (Grapsoidea: Sesarmidae), changes in biomass (measured as dry mass, W; carbon, C; nitrogen, N; and hydrogen, H; per individual) and relative elemental composition (C, N, H, in percent of W; C:N mass ratio) were studied during development from an early egg stage through hatching, the complete larval phase, metamorphosis and the first juvenile crab stage (CI). In the megalopa and CI, growth was measured also within the moulting cycle, and biomass and elemental composition were determined in cast exuviae. From an early egg stage to the freshly hatched larva, A. angustipes lost about 20% of W, 29% of C, 5% of N and 32% of H. Proportionally higher losses in C than in N were reflected also in a significantly decreasing C:N mass ratio (from 5.02 to 3.74). These results indicate that lipids mobilised from yolk reserves represented the principal metabolic substrate for embryonic energy production, while proteins were catabolised at a much lower rate. The present data of growth and exuviation are compared with previously published data from a congener, A. miersii Rathbun, which has an abbreviated and facultatively lecithotrophic mode of larval development (with three instead of four zoeal stages; stages I and II in principle independent of food). When growth is measured as an increase in the final (premoult) biomass of successive developmental stages, both species show an exponential pattern. Within the moulting cycles of the megalopa and the first juvenile, both species show parabola-shaped growth curves, with a rapid biomass increase in postmoult and intermoult stages, and losses in the premoult phase. Thus, the two Armases species show, in general, similar patterns of larval and early juvenile growth. However, the initial size of eggs and larvae is about four times larger in A. miersii, and its biomass remains higher throughout the period of larval and early juvenile development. A. angustipes is able to partially make up for this difference, as it has an additional zoeal stage, and its megalopa and CI stages show higher relative biomass increments (in percent of initial values). Due to this compensatory growth pattern, A. angustipes reaches in its CI stage about half the biomass of a juvenile A. miersii. When exuvial losses of megalopae and juveniles are compared between these two species, A. miersii shows higher biomass losses per individual (corresponding with its larger size), but lower relative losses (C, N, H, in percent of late premoult body mass or in percent of previously achieved growth increments). Differences in larval and early juvenile growth and in the exuvial losses of megalopae and juveniles of these two congeners are discussed in relation to their differential ecology, life history and reproductive strategy.     

Some quantitative aspects of seagrass ecology in a coastal lagoon of Mauritius

Seagrass distribution was recorded by snorkel dives on a grid of stations in the waterfront of Club Méditerranée at Mon Choisy-Trou Aux Biches lagoon (NW Mauritius) and subsequently mapped using SURFER 6 computer software. Above-ground (AG) and below-ground (BG) standing biomass in terms of dry weight (DW) and ash-free dry weight (AFDW) as well as shoot density and shoot length were monitored monthly from June1997 to May1998 in a mixed stand of Halodule uninervis and Syringodium isoetifolium (dominant) at a shallow, nearshore station in the lagoon. Measurements of physical and chemical parameters [water temperature, current speed, salinity, pH, dissolved oxygen (DO), nitrate and phosphate concentrations] were made simultaneously, as well as at a reference station (ORE) outside the coral reef. The bottom sediment was analysed for grain size and type composition. Variation patterns were examined and statistical correlations drawn to relate plant performance to the environmental variables measured. The SURFER 6 programme generated a satisfactory contour map of seagrass distribution in the lagoon with a cover range of 0-60%. The densest patches occurred adjacent to the shoreline experiencing weaker water currents (3-13 cm s^-1) rather than near the reef (5-35 cm s^-1), where seagrasses were absent. Sand (0.063-2 mm grain size) constituted 97.2% and 77.6% of the nearshore and near-reef sediment, respectively. The dominant grain types were derived from corals (about 80%) and mollusc shells (about 14%). The recorded range of total standing biomass for H. uninervis was 243.1-468.2 g DW m^-2 (326.9ᇛ.7 g) or 71.7-141.2 g AFDW m^-2 (96.8ᆨ.1 g) and for S. isoetifolium it was 271.7-758 g DW m^-2 (460.4끯.1 g) or 119-220.5 g AFDW m^-2 (155.1ᆮ.5 g), with a maximum biomass increase during September-December in both species. AG:BG biomass ratios were generally <1 and approximated 1 during the warmest months of December-February only. Mean shoot density (1,077-4,364 shoots m^-2 in the overall range of 998-4,428 shoots m^-2) and mean shoot length (10.9-20.8 cm in the overall range of 7-31 cm) in S. isoetifolium were higher than in H. uninervis (1,732-4,137 shoots m^-2 in the overall range of 1,522-4,327 shoots m^-2 and 7.9-13.7 cm in the overall range of 6-20 cm, respectively). Temperature showed strong positive correlations with total AFDW biomass of both species (r=0.755, P<0.01 for H. uninervis; r=0.679, P<0.02 for S. isoetifolium) and with DO (r=0.925, P<0.01). High DO levels (10.7-11.2 mg l^-1) coincided with optimum standing biomass at 27.2°C. Correlations were also strong with shoot density (r=0.881, P<0.01 for H. uninervis; r=0.952, P<0.01 for S. isoetifolium) and shoot length (r=0.752, P<0.01 for H. uninervis; r=0.797, P<0.01 for S. isoetifolium). Under optimal environmental conditions, nutrient inputs from surface run-off or underground freshwater seepage in the lagoon due to heavy rainfall may boost up seagrass biomass, as suggested by positive significant correlations between phosphate levels and AG AFDW biomass (r=0.63, P<0.05 for H. uninervis; r=0.65, P<0.05 for S. isoetifolium) and shoot density (r=0.6, P<0.05 for H. uninervis; r=0.687, P<0.02 for S. isoetifolium). The results generated in this study suggest local seagrass standing biomass is comparable to that reported in monospecific stands from elsewhere. Anthropogenic activities increasingly draw down the resilience of the seagrass beds around Mauritius, and preventative measures are indispensable to achieve coastal ecological stability.     

Effects of chemical cues on orientation of blue crab, <Emphasis Type="Italic">Callinectes</Emphasis><Emphasis Type="Italic">sapidus</Emphasis>, megalopae in flow: implications for location of nursery areas

Megalopae (postlarvae) of the blue crab Callinectes sapidus Rathbun use flood-tide transport (FTT) for movement into and up estuaries. Since they settle around the time of slack water at the end of flood tide during FTT, it was predicted that orientation toward primary nursery areas of aquatic vegetation occurs at this time. This study tested the hypotheses that megalopae locate nursery areas by swimming upstream in the presence of chemical odors from potential nursery areas and avoid adverse microhabitats by swimming downstream when predator or adverse environmental odors are present. Megalopae were tested in a flume where they were exposed to the sequence of cues mediating FTT (i.e. 2 psu increase in salinity followed by an increase and a decrease in current speed and turbulence). The flume contained odor water either from the developmental area (offshore water), nursery area vegetation (seagrass, Zostera marina; salt marsh cord grass, Spartina alterniflora), predators (fiddler crab, Uca pugilator; mud crab, Panopeus herbstii; grass shrimp, Palaemonetes pugio), or chemicals associated with adverse environments (ammonium). Vertical positions of premolt and intermolt megalopae were similar in water devoid of estuarine chemical cues (offshore water) and water containing seagrass odor. Upstream swimming behavior (orientation) of intermolt megalopae was also similar in these waters. However, there was an ontogenetic behavioral change, as the proportion of premolt megalopae oriented upstream generally increased as the concentration of seagrass and salt marsh cord grass odor increased and as current speed decreased. Upstream orientation of premolt megalopae in response to seagrass odor decreased significantly (i.e. downstream swimming increased) in the presence of odor from U. pugilator, P. pugio, and ammonium, but not from P. herbstii. Thus, the hypothesis was supported. These results suggest premolt megalopae orient toward nursery areas by swimming upstream in response to odors from aquatic vegetation as current speeds decrease at the end of nocturnal flood tides. Moreover, these results also indicate that megalopae may discriminate among microhabitats and avoid adverse settlement habitat, as orientation toward nursery areas is reversed by predator odors and ammonium.     

Role of the nutritive value of sediment in regulating population dynamics of the deposit-feeding polychaete Streblospio shrubsolii

The importance of changes in nutritive value of the sediment in regulating the dynamics of the small deposit-feeding polychaete Streblospio shrubsolii (Buchanam, 1890) was examined. First, fluctuations in the population dynamics of S. shrubsolii and in the nutritive content of the sediment were studied on a seasonal scale. Subsequently, a manipulative field experiment was used to test the hypothesis that, in summer, availability of food regulates aspects of the population dynamics of S. shrubsolii. In summer 1997, sharp increases in protein concentrations and in the number of individuals of S. shrubsolii were found compared to all the other seasons. Densities of S. shrubsolii also showed an increase in the subsequent season. Newly recruited juveniles (19-31 setigers) were abundant in summer and autumn 1997, whereas the number of juveniles (32-42 setigers) increased in autumn 1997. Large numbers of adults were found in summer 1997. Significant negative correlations occurred in summer 1997 between the concentrations of proteins and the total number of individuals, the number of newly recruited juveniles and the density of adults. No differences in densities of juveniles, adults or total number of individuals were found among plots before starting the experiment. After 20 days of experimental enrichment, proteins showed a tendency to increase. After 40 days of enrichment, the total number of individuals was significantly greater in the enriched plots. Large numbers of newly recruited larvae and postlarval adults were found in the enriched plots. The experiment corroborated the model that the growth of population was limited by the availability of food in summer. Thus, in this season, patches of food could have a fundamental role in determining the population dynamics of small deposit-feeders under natural conditions and over short time scales.     

<Emphasis Type="Italic">Semibalanus</Emphasis><Emphasis Type="Italic">balanoides</Emphasis> in winter and spring: larval concentration,settlement, and substrate occupancy

This study measured the progression from pelagic larvae to juvenile barnacles, and examined whether recruitment of barnacles, Semibalanus balanoides Linnaeus, at two intertidal sites in contrasting hydrodynamic regimes was determined by pre-settlement or post-settlement processes. The two sites were 1.5 km apart in the vicinity of Woods Hole, Mass., USA. Quantitative plankton samples were taken twice weekly from December 1997 to May 1998 at a nearby site as an estimate of nearshore larval abundance. The presence of S. balanoides nauplii was noted, and cyprids were enumerated and measured. Larval settlement at the two sites [Gansett Point, Buzzards Bay (GP) and Little Harbor, Vineyard Sound (LH)] was estimated from examination of replicate settlement plates exposed for 2 or 3 days throughout the settlement season, and from replicate plots on marked rock quadrats at each site. On both plates and rocks settled cyprids and metamorphs were enumerated. Space occupancy on unmanipulated rock quadrats by all stages from cyprids to adult barnacles was also examined. Settlement occurred from 2 January to 20 May, and major settlement peaks coincided with peaks in pelagic cyprid concentration at LH, but not at GP. Space occupied by juvenile barnacles was close to zero up until late February despite substantial settlement prior to that. At LH, juvenile barnacle cover was zero at the end of the observations; all settlement failed. Almost 100% of settled cyprids failed to metamorphose within 2 days from late January to late March. Then the proportion metamorphosing increased sharply coinciding with a sudden increase of 3°C in water temperature. Observed site differences in space occupancy by juvenile barnacles suggest that while cyprid supply is a necessary condition for barnacle settlement, other factors affecting metamorphosis of settled cyprids and early juvenile mortality determine recruitment.     

Spawning patterns in the leopard grouper, <Emphasis Type="Italic">Mycteroperca rosacea</Emphasis>, in comparison with other aggregating groupers

We documented the spawning patterns of the leopard grouper, Mycteroperca rosacea, from April to June 2005 in the central Gulf of California, Mexico to draw comparisons with other aggregate-spawning groupers and provide information useful for management of their fishery. Adults formed spawning aggregations of 150 to >700 individuals at specific sites, and spawning occurred daily at these sites from late April through early June. Courtship occurred throughout the day, but spawning was restricted to the evening hours. Adults spawned in groups of 6–40 fish, and pair-spawning was not observed. The group-spawning behavior of adults and the gonosomatic indices of mature males (maximum = 7.2%) suggest that sperm competition was present. The site-specificity of leopard grouper spawning aggregations and diel spawning period were typical of most aggregating groupers, and the size and structure of these aggregations was similar to other species in the genus Mycteroperca. Leopard grouper behavior patterns were unusual in that spawning aggregations persisted for extended periods, spawning was not synchronized with the lunar cycle, and adults aggregated during non-spawning periods. The extensive duration and site-specificity of spawning aggregations and the propensity of M. rosacea to form aggregations year-round increases the vulnerability of the species to overfishing. Policies that limit harvest from these aggregations are needed to improve the management of leopard grouper fisheries in the Gulf of California.     

Foraging behavior of juvenile Carcinus maenas (L.) and Cancer pagurus L.

Information concerning the way juvenile crabs choose their diet from a variety of prey types can be useful for a better understanding of community dynamics, as well as for the adequate management of natural resources. Prey size and species selection by juvenile Carcinus maenas (15-35 mm carapace width, CW) and Cancer pagurus (20-40 mm CW) feeding on four bivalves of contrasting shell morphology were investigated. When offered a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas, and Cerastoderma edule presented individually, crabs generally showed evidence of size-selective predation. Cancer pagurus selected larger mussels relative to the size of their chelae (relative prey size, RPS) than did Carcinus maenas of similar and even larger carapace width. However, the RPS of selected O. edulis and Cerastoderma edule were similar for all crabs, suggesting that certain prey features constitute effective barriers even to the powerful chelae of Cancer pagurus. When offered a wide size range of mussels and oysters simultaneously, all crabs consistently selected mussels. When offered O. edulis and Crassostrea gigas, crabs consumed both these oyster species in similar numbers. Carcinus maenas consumed similar numbers of mussels and cockles; Cancer pagurus, however, showed no preference for either prey in the smaller size classes but selected more mussels than cockles as prey increased in size. Although previous studies report that adult Carcinus maenas select prey species according to their profitability (amount of food ingested per unit of handling time, milligrams per second), consumption rates of the size classes of prey selected by juvenile shore crabs did not always parallel prey value. Although variations in crab strength can account for many of the differences between the foraging strategy of juvenile and adult C. maenas, our results suggest that juvenile crabs are less species selective than adults as a result of the restrictions imposed on small individuals that have limited access to larger prey.     

Differential effects of Aureococcus anophagefferens isolates ("brown tide") in unialgal and mixed suspensions on bivalve feeding

Picoplanktonic brown tides of Aureococcus anophagefferens have had devastating effects on production of commercially exploited bivalve populations in shallow, mid-Atlantic estuaries in the United States. The toxin produced by this alga has not been chemically characterized. This study develops a bioassay using juvenile mussels, Mytilus edulis, based on the inhibitory effect of brown tide on bivalve suspension-feeding, to compare the cellular toxicity of three Long Island, New York, clonal isolates of A. anophagefferens. Two recent (1995) isolates (CCMP 1707 and 1708) from Peconic Bay proved highly toxic and caused greater than 100-fold reduction in clearance rates (CR) of juvenile mussels in unialgal and mixed suspensions with a nutritious alga, Isochrysis galbana (clone T-iso), relative to controls. A third 1986 isolate from Great South Bay (CCMP 1784) showed no detectable toxicity in 24-h trials, and may have lost its initial potency over more than a decade of laboratory culture. Identification of a non-toxic strain provides a useful tool for future research. Cultures of the toxic isolate CCMP 1708 in late-stationary growth phase were significantly more toxic than those in early-exponential phase. The threshold concentration of toxic A. anophagefferens cells that inhibits clearance on co-occurring phytoplankton species was determined for juvenile (10-mm) hard clams, Mercenaria mercenaria. Relatively low concentrations (⣃᎒³ to 50᎒³ cells ml^-1) of isolate CCMP 1708 were sufficient to sharply reduce clam CR of I. galbana. Calculations based on these results suggest that, at peak historical densities of M. mercenaria in Great South Bay, removal of A. anophagefferens at low cell densities by suspension-feeding benthos could provide an effective top-down grazing control mechanism to prevent the initiation of brown tide in shallow, inner bays.     

Induction of larval settlement and metamorphosis of the sea urchin Strongylocentrotus intermedius by glycoglycerolipids from the green alga Ulvella lens

In aquaculture centers of the northern region of Japan, "Nami-ita" (waved polycarbonate plates), on which the green alga Ulvella lens Crouan frat. (Chaetophoraceae: Chaetophorales) was cultured, are used to promote larval settlement and metamorphosis of the sea urchin species Strongylocentrotus intermedius (A. Agassiz) and S. nudus (A. Agassiz). We investigated chemical inducer(s) for larval settlement and metamorphosis of these sea urchins with extracts of U. lens. Bioassay-guided separation of the methanol extract using a combination of column and thin-layer chromatography led to the isolation of several active compounds, the chemical structures of which were determined by spectral and chemical methods. These active compounds were identified as glycoglycerolipids, all comprising several molecular species: sulfoquinovosyl monoacylglycerols (SQMGs), sulfoquinovosyl diacylglycerols (SQDGs), monogalactosyl monoacylglycerols (MGMGs), monogalactosyl diacylglycerols (MGDGs), digalactosyl monoacylglycerols (DGMGs) and digalactosyl diacylglycerols (DGDGs). Among these glycolipids, SQMGs, MGMGs, MGDGs and DGMGs induced larval metamorphosis of the sea urchin S. intermedius. SQMGs and MGDGs induced larval metamorphosis at a concentration of 5 µg ml^-1, whereas SQDGs and DGDGs only induced larval settlement. These glycoglycerolipids are new congeners of chemical inducers to settlement and metamorphosis of planktonic larvae of sea urchins. The findings would provide a better understanding of larval settlement and metamorphosis in sea urchins.     

Evidence of different reproductive traits in the transisthmian sister species, <Emphasis Type="Italic">Alpheus saxidomus</Emphasis> and <Emphasis Type="Italic">A. simus</Emphasis> (Decapoda,Caridea, Alpheidae): description of the first postembryonic stage

The emergence of the Isthmus of Panama, approximately 3.0-3.5 million years ago, established two very different marine systems (the Caribbean and the eastern Pacific) and separated previously continuous marine populations. The geographic isolation of transisthmian sister species provides an excellent basis for the study of divergence and speciation processes. Here we describe the morphology of the first postembryonic stage of Alpheus saxidomus and A. simus, two rock-boring alpheids; the estimated time since divergence for this transisthmian pair ranges from 4.4-6.1 to 3-9 million years. The presence of a first zoeal stage in A. saxidomus, e.g., without pleopods and a telson with 7+7 setae, indicates a prolonged developmental sequence. In contrast, hatchlings of A. simus are substantially more developed and resemble juveniles. Thus, the developmental modes of A. saxidomus and A. simus are strikingly different with abbreviated, most probably direct, development in the latter species. Reduced food availability in the Caribbean compared to the Pacific coast is likely to be a possible reason for the evolution of such important differences in life history traits of the two transisthmian sister species. It is suggested that a transition from prolonged to abbreviated development evolved gradually during the estimated time since divergence; however, such a shift may have taken place within a substantially smaller time span.     

Relationships between inner ear and sagitta growth during ontogenesis of three Carapini species,and consequences of life-history events on the otolith microstructure

Three species of Carapidae have in common a tenuis larval stage, during which they settle in the lagoon and take refuge in the same species of holothuroid. From the juvenile stage, Carapus homei and C. boraborensis are commensal, whereas Encheliophis gracilis is parasitic. The aims of this study were to analyse to what extent the ontogenetic changes of the otic capsule affected the shape of the inner ear and how environmental cues, due to the above-mentioned life history and the style, could influence the structure of the sagitta. Sagittal sections revealed a three-dimensional asymmetry with a nucleus close to the proximal surface. Observations of the growth axis of the sagitta suggest it has a morphogenetic impact on the otic cavity. Each sagitta contains three main zones related to the life stages of the fish. Bands and checks were observed in the third zone in C. homei and C. boraborensis, but this pattern was less discernible in E. gracilis. These structural differences in zone 3 could be related to the commensal and parasitic life styles of these fishes. Further studies dealing with otosac features and otolith functions are suggested.     

Sibling species or poecilogony in the polychaete<Emphasis Type="Italic"> Scoloplos armiger</Emphasis>?

In marine invertebrates multiple modes of development, or poecilogony, may occur in a single species. However, after close examination, many of such putative cases turned out to be sibling species. A case in point may be the cosmopolitan orbiniid polychaete Scoloplos armiger, which inhabits marine shallow sediments. In addition to the well-known direct, holobenthic development from egg cocoons, pelagic larvae have also been described. Our culture experiments revealed a spatially segregated source of the two developmental modes. All females of an intertidal population produced egg cocoons and no pelagic larvae. All but 2 out of 15 females of an adjacent subtidal population produced pelagic larvae and no egg cocoons. Based on these results we performed a molecular genetic analysis (RAPD-PCR) on three intertidal and four subtidal populations in the North Sea. Selected samples from all sites were analysed also by the AFLP method. We found significantly higher genetic diversity within subtidal than within intertidal populations. This is consistent with a wider dispersal by pelagic larvae and a smaller effective population size when development is holobenthic. Total genetic divergence is not related to distance but to the intertidal/subtidal division. We suggest that S. armiger actually represents two sibling species.     

Cellulose digestion and phenol oxidation in coastal isopods (Crustacea: Isopoda)

In order to test three hypotheses on digestive constraints that may have affected the colonization of land by isopods, two marine isopods and one semi-terrestrial species were screened for their ability to oxidize phenolic compounds and digest cellulose in natural and artificial diets. Ligia pallasii (Isopoda: Oniscidea) and Gnorimosphaeroma oregonense (Isopoda: Sphaeromatidea) oxidized dietary phenolics, but Idotea wosnesenskii (Isopoda: Valvifera) did not, even though it feeds on seaweeds that are rich in phenolics. All three species were able to digest some cellulose, but this ability was least developed in the marine phytophagous species, Idotea wosnesenskii, and best developed in the semi-terrestrial L. pallasii. After reducing the number of endosymbiotic bacteria in the hepatopancreas (midgut digestive gland) by feeding antibiotics, cellulose digestion in L. pallasii was significantly reduced. Our results are consistent with the hypotheses that (1) the ability to oxidize phenolics is absent in phytophagous marine isopods, but present in saprophagous marine and semi-terrestrial species, (2) the ability to digest cellulose was an important pre-adaptation facilitating a fully terrestrial life-style in isopods, and (3) endosymbiotic bacteria in the hepatopancreas aid digestion in terrestrial isopods, and to a lesser degree in semi-terrestrial species, but not in marine isopods.     

Interspecific infection of aposymbiotic juveniles of <Emphasis Type="Italic">Codakia</Emphasis><Emphasis Type="Italic">orbicularis</Emphasis> by various tropical lucinid gill-endosymbionts

Previous molecular phylogenetic analyses have shown that five tropical lucinid species living in or near Thalassia testudinum seagrass beds are colonized by the same bacterial symbiont species. In addition, a new lucinid species belonging to the genus Anodontia, which inhabits reducing sediment found near seagrass beds and in mangrove swamps, has been included in the present study. Endosymbiosis in Anodontia alba was examined according to symbiont phylogenetic and gill ultrastructural analysis. Phylogenetic analysis showed that partial 16S rDNA sequences of A. alba- and Codakia orbicularis-symbionts were 100% identical at all nucleotide positions determined, suggesting that A. alba also harbors the same symbiont species as C. orbicularis (and, consequently, as C. orbiculata, C. pectinella, Linga pensylvanica and Divaricella quadrisulcata). Based on light and electron microscopy, the cellular organization of the gill filament appeared similar to those already described in other lucinids. The most distinctive feature is the lack of "granule cells" in the lateral zone of A. alba gill filaments. In order to confirm the single-species hypothesis, purified fractions of gill bacterial symbionts obtained from the gills of each of the six tropical lucinids cited above were used to infect aposymbiotic juveniles of C. orbicularis. In each case, aposymbiotic juvenile batches were successfully infected by the gill-endosymbiont fractions, whereas, during the experiments, juveniles from the negative control were still uninfected. These experimental data confirm the phylogenetic data and also demonstrate that chemoautotrophic bacterial endosymbionts from their host cells can colonize aposymbiotic juveniles. The conclusion also follows that intracellular gill-endosymbionts still have the capacity to recognize and colonize new host generations. Lucinids provide a unique model for the study of sulfide-oxidizing symbiosis, even if symbionts remain unculturable.     

Settlement patterns of newly settled plaice (Pleuronectes platessa) in a non-tidal Swedish fjord in relation to larval supply and benthic predators

In demersal fish species with a pelagic larval stage, settlement patterns may be a consequence of variations in larval supply, habitat selection at settlement, and processes acting between the time of settlement and the time of benthic sampling. This study describes temporal (1994-1998) and spatial variation in plaice (Pleuronectes platessa L.) settlement densities in four semi-isolated nursery areas with similar habitat characteristics, in the non-tidal Gullmarsfjord on the west coast of Sweden. Juvenile abundance varied by a factor of ten, both among years and among nursery grounds. For the 3 years when larval sampling was undertaken (1994-1996) and all nursery areas, there was a significant positive relationship between larval supply and juvenile abundance (linear regression: r²=0.45, n=24, P<0.001). On the southern side of the fjord, a significant positive relationship between larval and juvenile abundance was found in one area (r²=0.62, n=6, P<0.05). The absolute mortality rate of plaice after settlement was related to the initial settlement density (r²=0.95, n=20, P<0.001), and to the abundance of predatory shrimps Crangon crangon (r²=0.44, n=20, P<0.01). Plaice otoliths were found in 6% of the shrimp stomachs analysed from an area with high density (13.3 m^-2) of newly settled plaice. The present study suggests that the density of juvenile plaice was limited by larval supply to the nursery grounds. Consistency in the relative abundance of juveniles among nursery grounds between years also suggested that some nursery areas may be in the settlement shadow of others. The irregular nature of the coastline in combination with larval depletion could thereby cause small-scale (10³-10⁴ m) variation in settlement densities of the same order of magnitude as the inter-annual variability in recruitment to individual nursery grounds.     

Seasonal variation in ecophysiological patterns in macroalgae from an Arctic fjord. I. Sensitivity of photosynthesis to ultraviolet radiation

Marine macroalgae inhabiting Arctic coastal ecosystems are exposed to pronounced seasonal variations in the radiation regime, including harmful UVB radiation. This study presents the first data on the seasonal changes in the sensitivity of macroalgal photosynthesis towards UV exposure by comparing under-ice, clear-water and turbid-water conditions characteristic for late winter, spring and summer. Various brown (Laminaria saccharina, L. digitata, L. solidungula, Saccorhiza dermatodea, Desmarestia aculeata), red (Palmaria palmata, Devaleraea ramentacea) and one green macroalgal species (Monostroma aff. arcticum) were collected at the same water depth throughout the seasons in the Kongsfjord (Spitsbergen, Svalbard, Norway). Maximum quantum yield (F_v/F_m) and maximum photosynthetic electron transport rates (ETR_max) were determined immediately after collection, after 2 h exposure to artificial UV radiation and after 18 h recovery in dim white light. Photosynthesis of the studied species showed different responses depending on their morpho-functional and physiological characteristics, their life strategies, phenology and depth distribution. Within the genus Laminaria, maximum quantum yield of adult specimens of the deep-water species L. solidungula was most strongly UV sensitive. Adult L. saccharina exhibited a lower UV sensitivity than a 6-month-old specimen. Inhibition of photosynthesis after UV exposure remained at the same level throughout the study period, both in adult L. saccharina and S. dermatodea. However, adult specimens of L. saccharina collected in May showed partial recovery only, whereas photosynthesis of specimens from both species collected later recovered fully. D. aculeata exhibited a remarkable decrease of UV sensitivity during the study period. Photosynthesis of specimens collected under the ice was strongly inhibited by UV, but the degree of inhibition decreased during spring and summer. Concomitantly ETR_max values were low after UV exposure in specimens collected in June, but increased later in the season. P. palmata exhibited a relatively flexible response. Photosynthesis in specimens collected under the ice in June or in turbid water in July/August was relatively strongly inhibited; specimens collected during sunny periods and in clear water in spring showed a much lower degree of photoinhibition after UV exposure. The seasonal pattern of low/high ETR_max values in spring/summer is probably a characteristic of the life strategy of this species. The UV sensitivity of D. ramentacea exhibited a similar seasonal pattern. In M. aff. arcticum, UV sensitivity increased and ETR_max values decreased during the study period, reflecting the life strategy of this annual late winter/spring species. The physiological basis for the seasonal changes in UV sensitivity of photosynthesis is presented in a companion paper (this issue).     

Under-ice amphipods in the Greenland Sea and Fram Strait (Arctic): environmental controls and seasonal patterns below the pack ice

During three "Polarstern" cruises to the ice-covered Greenland Sea (spring 1997, summer 1994, autumn 1995) studies on the under-ice habitat (morphology, hydrography, ice-algal biomass) and on the macrofaunal, autochthonous under-ice amphipods (species diversity, abundance) were carried out in order to describe environmental controls and seasonal patterns in this community. In spring, the ice underside was rather smooth and whitish, while in summer melting structures and sloughed-off ice-algal threads were observed, in autumn detritus clumps accumulated in depressions at the ice underside. Only in summer, a thin layer of warm (up to -0.6°C) and less saline (as low as S=6.3) water was found at the ice-water interface above Polar Water. Integrated ice-algal biomass was highest during autumn (2.6 mg chl a m^-2) and lowest during summer (1.2 mg chl a m^-2). Four species of under-ice amphipods occurred in spring and summer (Apherusa glacialis, Onisimus glacialis, O. nanseni, Gammarus wilkitzkii), but only the last species was observed at the ice underside in autumn. A. glacialis and G. wilkitzkii were equally abundant in spring; A. glacialis dominated in summer. The highest total abundance of amphipods occurred during summer (31.9 ind. m^-2), compared to lower abundances in spring and autumn (5.3 and 1.1 ind. m^-2, respectively). A factor analysis revealed seasonal patterns in the data set, which mainly influenced A. glacialis, and species-specific relations between several environmental factors and the distribution of under-ice amphipods. Abundance of A. glacialis was closely related to the under-ice hydrography and ice-algal biomass, whereas the other amphipod species were more influenced by the under-ice morphology. It is therefore stated that the observed thinning of the Arctic sea ice and the resulting increased meltwater input and change in morphology of floes will have a profoundly adverse effect on the under-ice amphipods.