ALBERT

Description: Five hormone-treated female Japanese silver eels Anguilla japonica were tagged with ultrasonic transmitters and released by submersible in the West Pacific at seamounts of the West Mariana Ridge, their supposed spawning grounds. Four eels were tracked for 60 to 423 min in the vicinity of the seamounts. They did not settle at the seamounts but swam at a mean speed of 0.37 m s-1 into open water above deep ground. Their mean swimming depth ranged from 81 to 172 m. Experiments suggest that pre-matured A. japonica migrate to their spawning grounds in temperate warm water and at shallow depths.

Description: Phosphatase (P-ase) activity was determined together with other extracellular enzyme activities, bacterial abundance and production rates during the 2 SW Monsoon process studies of the German JGOFS Arabian Sea Program. Water samples were collected along the cruise tracks from the equator to the upwelling region at the shelf edge off Oman. Depth profiles of P-ase activity were strikingly different from those of the other enzymes. While values of aminopeptidase and β-glucosidase generally decreased below the euphotic zone, P-ase increased by factors of 1 to 7. The relation between peptidase- and P-ase activity was from 4 to 21 at the surface and from 3 to 5 at 800 m depth. Because P-ase production (dissolved and cell-bound) in deep waters is mainly dependent on bacteria, P-ase activities per bacterial cell were calculated: these were, on average, 37 times higher at 800 m than at the surface. We also observed a positive correlation of P-ase activity with phosphate concentrations in the depth profiles below the euphotic zone, while this relationship was much more variable in the mixed surface layer. These observations suggest that C-limited bacteria in the deep strata did not primarily focus on the phosphate generated by their P-ase activity but on the organic C compounds which were simultaneously produced and which could probably not be taken up prior to the hydrolytic detachment of phosphate. It is hypothesised that a considerable part of the measured P-ase activity was dissolved (though it might have originated from bacteria). These enzymes may be important for the slow, but steady regeneration of phosphate and organic C in mesopelagic waters.

Description: Stomach contents of 17 sperm whales Physeter macrocephalus stranded in Scotland and Denmark during 1990-96 were analysed. All were sub-adult or adult males and stranded between November and March. They had presumably entered the North Sea during their southward migration from feeding grounds in Arctic waters. Other studies indicate that the majority of the whales were apparently healthy. The diet of these whales was found to consist almost entirely of cephalopods, principally squid of the genus Gonatus (hereafter 'Gonatus', but probably G. fabricii, an oceanic species characteristic of Arctic waters). The other prey species identified were also mostly oceanic cephalopods: the squids Histioteuthis bonnellii, Teuthowenia megalops and Todarodes sagittatus and the octopus Haliphron atlanticus. Although these results are consistent with other recent studies in the area based on single stranded whales, they differ from results of work on whales caught during commercial whaling operations in Icelandic waters (1960s to 1980s) in that little evidence of predation on fish was found in the present study. Remains of single individuals of the veined squid Loligo forbesi, the northern octopus Eledone cirrhosa and the saithe Pollachius virens provided the only possible evidence of feeding in the North Sea. We infer that sperm whales do not enter the North Sea to feed. The timing, and large and uniform sizes of the Gonatus species eaten (most had mantle lengths in the range 195 to 245 mm), as estimated from measurements of the lower beaks, and the seasonality of the strandings is consistent with the whales having fed on mature squid, possibly spawning concentrations--as has recently been reported for bottlenose whales. Assuming that the diet recorded in this study was representative of sperm whales during the feeding season, as much as 500000 t of Gonatus could be removed by sperm whales in Norwegian waters each year and up to 3 times that figure from the eastern North Atlantic as a whole. Evidence from other studies indicates that Gonatus is an important food resource for a wide range of marine predators in Arctic waters.

Description: Interpretation of diving profiles of aquatic animals would be considerably enhanced by additional behavioural information. A new sensor is presented here which records animal movements. This sensor was tested on a captive loggerhead turtle Caretta caretta which showed similar activity patterns to free-living green turtles Chelonia mydas. A computer program with user-selectable options was developed to analyse the data consistently and rapidly. Using our sensor we calculated the total resting time, which differed by less than 5% from the real resting time when the sampling interval was 2 s. The method was additionally tested for different sampling intervals to find out its applicability for field studies.

Description: Nematodes of the family Stilbonematinae are known for their highly specific association with ectosymbiotic bacteria. These worms are members of the meiofauna in marine, sulfide-rich sediments, where they migrate around the redox boundary layer. In this study, bacterial ectosymbionts of 2 species of marine nematodes, Stilbonema sp. and Laxus oneistus, were shown to be capable of the respiratory reduction of nitrate and nitrite (denitrification). The use of these alternative electron acceptors to oxygen by the bacteria allows the animals to migrate into the deeper, anoxic sediments, where they can exploit the sulfide-rich patches of the deeper sediment layers. The accumulation of thiols (sulfide, thiosulfate, sulfate and glutathione) in body tissues of the worms was determined following incubation in the presence of various electron donors (sulfide, thiosulfate) and acceptors (nitrate). In their chemoautotrophic metabolic potential, the ectosymbionts of the 2 nematode species were found to resemble the phylogenetically related, intracellular symbionts of macrofaunal hosts of deep-sea hydrothermal vents and other sulfide-rich habitats.

Description: The role of tetrathionate in the sulfur cycle of Baltic Sea sediments was investigated in different habitats and under a variety of environmental conditions. Sediment profiles were recorded with regard to numbers of thiosulfate oxidizing bacteria, concentrations of sulfur compounds, and potential rates of thiosulfate oxidation. Products of thiosulfate oxidation were quantified in incubated sediment samples and in pure cultures. Evidence was found that tetrathionate is formed within these sediments, that sulfur oxidizing bacteria are present in considerable numbers, that these bacteria are of major importance in the oxidation of reduced sulfur compounds in their habitat, and that tetrathionate is an important oxidation product of these bacteria. Thiosulfate is oxidized by bacteria isolated from these sediments to varying proportions of tetrathionate, sulfate, and also elemental sulfur. In highly sulfidic sediments and in the presence of large amounts of organic matter, tetrathionate was present in sediment horizons in which thiosulfate and elemental sulfur also accumulated. A tetrathionate cycle is proposed to be active in natural marine and brackish water sediments in which, due to combined bacterial action and chemical reactions, a net oxidation of sulfide to elemental sulfur occurs in the presence of catalytic amounts of thiosulfate and tetrathionate.

Description: Seabirds, like all marine endotherms, have to compensate for the extensive cooling effect of water when diving. Alone among them, cormorants (Phalacrocoracidae) have a wettable plumage and are predicted to require disproportionately large amounts of food to balance heat losses. These piscivorous birds are thus thought to have a detrimental impact on fish stocks. However, we show here that even in great cormorants from Greenland, which dive in water at 3 to 7°C, daily food intake is lower than for well-insulated European seabirds. Despite their wettable plumage, cormorants thus appear to manage their energy budgets in a remarkably efficient way. Nevertheless, the specific foraging strategies which enable this performance make cormorants dependent on high prey density areas, a feature that should be taken into account by future management plans.

Description: In Eckernförde Bay (western Baltic Sea) pockmark structures are induced by groundwater seeping out of the sediment. On 3 occasions in winter and spring 1993-94 we investigated the influence of groundwater on the reduction of salinity, on porewater chemistry, and on bacterial activities (methane oxidation and sulphate reduction). In 2 out of 3 sampling campaigns groundwater discharge could be detected. The concentration gradients of Cl- and SO4= are moved towards the sediment surface by the vertical advection of groundwater during seep times. Without groundwater discharge the porewater chemistry resembled the control site. Compared to the control site, the methane oxidation and sulphate reduction rates were elevated at the pockmark site, reaching maximum values of 49 and 269 µmol l-1 d-1 respectively. The groundwater venting from the pockmark had an end member composition of 80 mM Na+, 1.0 mM Ca++ and was depleted in Mg++. Due to mixing of these major cations along the groundwater/seawater interface, no CaCO3 precipitation was found around the pockmark site.

Description: Sinking velocities of more than 300 Nitzschia closterium aggregates were determined during roller table incubation using digital image analysis. To examine the influence of transparent exopolymer particles (TEP) on aggregate settling speed, 3 experiments with different ratios of TEP to cell volume concentration were conducted. The results showed that, for N. closterium aggregates without TEP, sinking velocity (U) was significantly related to the equivalent spherical diameter (ESD) of the aggregates, yielding U (cm s-1) = 1.89 (ESD, cm)0.55. The higher was the specific TEP content of an aggregate, the lower was the sinking velocity and the less pronounced was the size versus velocity relationship. Excess densities (Δρ) of aggregates were derived from velocity measurements and 3-dimensional fractal dimensions (D3) of aggregates were calculated from scaling properties of Δρ. Values for D3 never exceeded 2 and fit well to values of the 2-dimensional fractal dimension (D2) attained from image analysis.

Description: Stable carbon isotope fractionation (ε p) of 7 marine phytoplankton species grown in different irradiance cycles was measured under nutrient-replete conditions at a high light intensity in batch cultures. Compared to experiments under continuous light, all species exhibited a significantly higher instantaneous growth rate (μi), defined as the rate of carbon fixation during the photoperiod, when cultivated at 12:12 h, 16:8 h, or 18:6 h light:dark (L/D) cycles. Isotopic fractionation by the diatoms Skeletonema costatum, Asterionella glacialis, Thalassiosira punctigera, and Coscinodiscus wailesii (Group I) was 4 to 6o/oo lower in a 16:8 h L/D cycle than under continuous light, which we attribute to differences in μi. In contrast, ε p in Phaeodactylum tricornutum, Thalassiosira weissflogii, and in the dinoflagellate Scrippsiella trochoidea (Group II) was largely insensitive to daylength-related differences in instantaneous growth rate. Since other studies have reported growth-rate dependent fractionation under N-limited conditions in P. tricornutum, μi-related effects on fractionation apparently depend on the factor controlling growth rate. We suggest that a general relationship between εi and μi/[CO2,aq] may not exist. For 1 species of each group we tested the effect of variable CO2 concentration, [CO2,aq], on isotopic fractionation. A decrease in [CO2,aq] from ca 26 to 3 µmol kg-1 caused a decrease in ε p by less than 3o/oo. This indicates that variation in μi in response to changes in daylength has a similar or even greater effect on isotopic fractionation than [CO2,aq] in some of the species tested. In both groups ε p tended to be higher in smaller species at comparable growth rates. In 24 and 48 h time series the algal cells became progressively enriched in 13C during the day and the first hours of the dark period, followed by 13C depletion in the 2 h before beginning of the following light period. The daily amplitude of the algal isotopic composition (δ13C), however, was 〈=1.5o/oo, which demonstrates that diurnal variation in δ13C is relatively small.

Description: The El Niño/Southern Oscillation (ENSO) phenomenon is the strongest natural interannual climate fluctuation1. ENSO originates in the tropical Pacific Ocean and has large effects on the ecology of the region, but it also influences the entire global climate system and affects the societies and economies of manycountries2. ENSO can be understood as an irregular low-frequency oscillation between a warm (El Niño) and a cold (La Niña) state. The strong El Niños of 1982/1983 and 1997/1998, along with the more frequent occurrences of El Niños during the past few decades, raise the question of whether human-induced 'greenhouse' warming affects, or will affect, ENSO3. Several global climate models have been applied to transient greenhouse-gas-induced warming simulations to address this question4, 6, but the results have been debated owing to the inability of the models to fully simulate ENSO (because of their coarse equatorial resolution)7. Here we present results from a global climate model with sufficient resolution in the tropics to adequately represent the narrow equatorial upwelling and low-frequency waves. When the model is forced by a realistic future scenario of increasing greenhouse-gas concentrations, more frequent El-Niño-like conditions and stronger cold events in the tropical Pacific Ocean result

Description: Diverse coastal seaweed communities dominated by perennial fucoids become replaced by species-poor turfs of annual algae throughout the Baltic Sea. A large-scale field survey and factorial field experiments indicated that grazers maintain the fucoid community through selective consumption of annual algae. Interactive effects between grazers and dormant propagules of annual algae, stored in a 'marine seed bank', determine the response of this system to anthropogenic nutrient loading. Nutrients override grazer control and accelerate the loss of algal diversity in the presence but not in the absence of a propagule bank. This implies a novel role of propagule banks for community regulation and ecosystem response to marine eutrophication.

Description: An experimental laboratory set-up was used to study the influence of different grain size compositions and temperatures on the growth of benthic cyanobacteria and diatoms, and on the competition between these 2 groups. Monospecific cultures of 3 species of cyanobacteria (Merismopedia punctata, Microcoleus chthonoplastes, Oscillatoria limosa), and of 2 species of benthic diatoms (Phaeodactylum tricornutum and Nitzschia sp.) were used. The organisms were cultured in 100 ml flasks filled with medium and 3 different kinds of sediment: (1) Sand (fine sand, 63 to 200 µm), (2) Mud-I (mixed fine sand and mud 〈63 µm in the ratio 80:20 wt %), (3) Mud-II (mixed fine sand and mud in the ratio 50:50 wt %). Experimental temperatures were 10, 15 and 25°C. At 10°C and 15°C, both diatom species achieved the highest biomass on the sediments of the finest grain size (50 wt % 〈 63 µm) while cyanobacteria achieved low biomass levels. Coarsening of sediments at the same temperature levels revealed a gradually lower biomass of the diatoms. Particularly on sand, the diatoms never reached the same concentrations of chlorophyll a as on mud. The cyanobacteria, on the other hand, had the highest biomass on sand at 15°C. In the competition experiments the benthic diatom species Nitzschia sp. dominated all types of sediments at 10°C and 15°C. The experiments at 25°C were dominated by the filamentous cyanobacterium M. chthonoplastes. This indicates the importance of abiotic conditions for the distribution and abundance of benthic phototrophic micro-organisms.

Description: The multivariate patterns resulting from analyses of macrobenthic abundance data at different taxonomic levels are compared to the pattern derived from various measurements obtained through sediment profiling imagery (SPI). A time-series data set from 1 station in Kiel Bay (Western Baltic) at 22 m depth including macrobenthic and SPI replicates covering 8 yr (1989 to 1996) was analyzed by means of multidimensional scaling (MDS) ordination. The macrobenthos data showed similar patterns, and there was little information loss, with decreasing taxonomic resolution from species to phylum level. The multivariate pattern in the SPI data was not significantly correlated to any of the macrofaunal patterns. However, macrofaunal and SPI patterns seemed to be complementary since they emphasized different aspects of the long-term succession in the Southern Baltic Sea. While macrofaunal patterns were sensitive to anoxia events, changes of SPI-recorded seabed characteristics were primarily related to physical disturbances possibly due to variations in fishing intensity.

Description: Diving seabirds should evolve a variety of foraging characteristics which enable them to minimize energy expenditure and to maximize net energy gain while searching for prey underwater. In order to assess the related ecological adaptations in a marine predator, we studied the at-sea distribution and the diving behaviour of 23 cormorants Phalacrocorax carbo (Linnaeus) breeding at the Chausey Islands (France) using VHF-telemetry and data loggers recording hydrostatic pressure. Birds foraged within an area of approximately 1131 km2 situated north-east of the breeding colony. This zone represents only 25% of the maximal potentially available area that the birds may utilize considering their maximum foraging range of 35 km. Individual birds remained within restricted individual foraging areas (on average 18 and 10% of the total utilized area in 1994 and 1995, respectively) throughout the study period. Moreover, the cormorants studied conducted an average of 42 dives per foraging trip, lasting for an average of 40 s (maximum 152 s), and reached an average maximum dive depth of 6.1 m (maximum 32 m) with median descent and ascent angles calculated to be 18.7° and 20.3°, respectively. Overall, 64% of all dives were U-shaped dives and 36% V-shaped dives. We use these results to demonstrate how both specialization and opportunism may support the remarkably high foraging efficiency of this marine predator.

Description: Cephalopods play an important role in the trophic web of the Southern Ocean, but little information is available on their biology. The 2 largest sub-Antarctic seabirds, the king penguin Aptenodytes patagonicus and the wandering albatross Diomedea exulans, feed primarily on squids during the austral winter at the Crozet Islands. We examined a large number of accumulated cephalopod beaks in the stomach of these birds together with some undigested items; first, to understand how these 2 predators share the squid resource during winter, a period of supposed low food availability, and, second, to use a diving and a flying seabird as biological samplers of Southern Ocean cephalopods. Individuals of the family Onychoteuthidae formed the bulk of the squid diet, accounting for 72.6 and 57.0% of the number of lower beaks in samples from king penguins and wandering albatrosses, respectively. Seven different species were identified, the 3 main squids being Kondakovia longimana (38.8 and 28.0% by number for penguins and albatrosses, respectively), Moroteuthis ingens (13.5 and 26.2%) and M. knipovitchi (20.1 and 2.3%). Both seabirds preyed upon the same cephalopod species, but penguins primarily took small- to medium-sized juveniles (99.0% of the onychoteuthids) and albatrosses preyed on larger adult specimens (96.0%). Fresh remains indicated that adult K. longimana and M. ingens were mature individuals which, as shown by satellite tracking of albatrosses, were taken over the slope and nearby oceanic waters surrounding the archipelago. The present study indicates that mating/spawning of K. longimana and M. ingens occurs in Crozet waters during the winter months. It also extends the biogeography of K. longimana to north of the Antarctic Polar Front, in the Polar Frontal Zone, where it has not previously been recorded.

Description: The marine habitat exploited by black-browed Diomedea melanophrys and grey-headed albatrosses D. chrysostoma breeding at Campbell Island, New Zealand, was studied using satellite telemetry. Data were analysed in relation to the bathymetry and sea-surface temperature of the foraging zones. Black-browed albatrosses spent 55% of their time on the Campbell Plateau but also carried out long foraging trips to the Polar Front and Antarctic Zone at a distance of over 2000 km. They relied heavily on juvenile Micromesistius australis, a schooling fish, during foraging trips to the shelf but over oceanic waters the squid Martialia hyadesi was the main prey taken. Grey-headed albatrosses spent 71% of their time foraging over the deep waters of the Polar Frontal Zone where M. hyadesi comprised over 90% of the mass of prey taken. No satellite-tracked birds fed over the shelf, but data from the duration of foraging trips and dietary analysis suggests that shelf-feeding is important for this species. Significant inter-species differences in the time spent in neritic and oceanic zones show that black-browed albatrosses are reliant primarily on shelf resources while grey-headed albatrosses are primarily oceanic feeders. In addition, the 2 species overlapped little in the zones used over oceanic waters, with black-browed albatrosses feeding in more southerly waters than grey-headed albatrosses. However, both species feed on M. hyadesi when foraging in association with the Polar Front.

Description: The fishery for Illex argentinus in the Southwest Atlantic is subject to large inter-annual variability in recruitment strength. In this paper we attempt to build a predictive model using sea surface temperature (SST) to examine links between recruitment to the Falkland Islands fishery and environmental variability during the juvenile and adult life history stages. SST data from the National Center for Atmospheric Research (NCAR) were found to be comparable with near-surface data derived from in situ expendable bathy-thermograph (XBT) profiles in the southern Patagonian shelf. Variation in SST during the early life stages appears to be important in determining recruitment of I. argentinus. SST in the hatching grounds of the northern Patagonian shelf during the period of hatching (particularly June and July) was negatively correlated with catches in the fishery in the following season. SST anomaly data from positions in the Pacific and Southwest Atlantic were used to examine teleconnections between these areas. Links were seen at a lag of 2 yr between the Pacific and southern Patagonian shelf, and at about 5 yr between the Pacific and northern Patagonian shelf. This is consistent with SST anomalies associated with El Niño in the Pacific propagating around the globe via the Antarctic Circumpolar Wave (ACW). Predicting cold events via teleconnections between SST anomalies in the Pacific and Atlantic would appear to have the potential to predict the recruitment strength of I. argentinus in the Southwest Atlantic.

Description: Levels of genetic diversity and population differentiation were examined in temporally (1990 to 1997) and geographically separated samples of the argentine short-finned squid Illex argentinus using 7 microsatellite loci. Number of alleles (mean number of alleles per locus over all samples = 24.1) and heterozygosity (mean observed heterozygosity per sample = 0.84) were high for all samples, indicating that these loci have a greater potential utility for investigating population genetic structure than allozyme markers used in previous studies. Genetic diversity did not differ significantly between samples taken 5 yr after commencement of the fishery (1990) and those collected during a period of progressively intense fishing pressure (1994 and 1997). Several small but significant differences in between-sample genetic variation (FST) were observed, but these could not confirm the previous suggestion of cryptic species or several well-defined stocks within the fished population.

Description: The overflow and descent of cold dense water from the Denmark Strait sill-a submarine passage between Greenland and Iceland-is a principal means by which the deep ocean is ventilated, and is an important element in the global thermohaline circulation. Previous investigations of its variability-in particular, direct current measurements(1,2) in the overflow core since 1986-have shown surprisingly little evidence of long-term changes in now speed. Here we report significant changes in the overflow characteristics during the winter of 1996-97, measured using two current-meter moorings and an inverted echo sounder located at different depths in the fastest part of the now. The overflow warmed to the highest monthly value yet recorded (2.4 degrees C), and showed a pronounced slowing and thinning at its lower margin. We believe that the extreme warmth of the overflow caused it to run higher on the continental slope off east Greenland, so that the lower current meters and the echo sounder were temporarily outside and deeper than the fast-flowing core; model simulations appear to confirm this interpretation, We suggest that the extreme warmth of the overflow is a lagged response to a warming upstream in the Fram Strait three years earlier (caused by an exceptional amplification of the winter North Atlantic Oscillation). If this is so, over-now characteristics may be predictable.

Description: Epibiosis is a spatially close association between 2 or more organisms belonging to the same or different species. Through direct and indirect interactions, this association has major effects on the species involved and on community dynamics. When the effects are predominantly beneficial for epibiont and basibiont, coevolution can be expected to lead to associational specificity. Circumstantial evidence, however, suggests that many epibionts are non-specific substratum-generalists. In this arti-cle, we investigate the commonness of specificity in epibiotic associations. In a first approach, we inves-tigated the in situ recruitment preferences of potential epibionts when choosing between artificial and living substrata. After exposure for 3 wk in early summer, an early successional community had estab-lished, comprising cyanobacteria, diatoms, sesslle colonial ciliates and red algae. All species recruited on almost all substrata available. However, artificial substrata were usually preferred over living sur-faces. Consequently, the species studied are class~fied as facultative epibionts. An analysis of a list of over 2000 epibiotic associations corroborated these results, the majority of described 'epibionts' are not basibiont-specific and generally occur on non-living substrata as well. Also, basibiont species usually bear more than 1 epibiont species. Relative to each other, epibionts and basibionts are characterised by a typical set of life history traits. We conclude that specific and obligate epibionts are rare. Their scarcity is discussed in view of multilevel antifouling defences and presumptive evolutionary transi-tions from epibiosis towards endoparasitism or endosymbiosis.

Description: Diving reptiles, unlike most diving birds and mammals, return infrequently to the surface to breathe. Spending the bulk of their lives underwater, they are likely to have developed a large variety of specific behavioural patterns different from those of their warm-blooded counterparts. However, for technical reasons, underwater behaviour of these aquatic reptiles remains poorly understood. In this study green turtles Chelonia mydas nesting on Cyprus (Eastern Mediterranean) were equipped with multi-channel data loggers monitoring diving behaviour and activity (via a logger-integrated 3-D compass which served as an activity sensor) during the inter-nesting interval. Data from 2 turtles for 2 consecutive inter-nesting intervals were available for detailed dive analysis. Both turtles showed highly variable dive patterns ranging from travelling subsurface dives to specific dive types such as U- (mainly resting and foraging dives), S- (a form of energy saving swimming) and V-dives. The green turtles stayed near the coast throughout the study, dived no deeper than ca 25 m, but remained underwater for up to ca 40 min. The recordings of the activity sensor revealed high activity levels (less than 20% resting d-1) during the whole inter-nesting period which was attributed to extensive foraging. The combination of both the activity data and the dive data showed that the turtles were engaged in travelling movements for 46% of the inter-nesting time spent underwater, foraged for 34% and rested for 12% of the time. We discuss the physiological, ecological and conservation implications of these results.

Description: Serial dilution experiments were conducted on JGOFS-North Atlantic cruise of RV 'Meteor' M36/2 at a 20° W transect in June and July 1996 to assess the role of microzooplankton grazing and nitrogen supply in controlling phytoplankton stocks in the subtropical and temperate northeast Atlantic. Rates of microzooplankton grazing ranged from 0.08 d-1 at 54° N to 0.53 d-1 at 40° N and mean growth rates of phytoplankton ranged from 0.19 d-1 at 54° N to 0.75 d-1 at 40° N. Both rates were positively related to seawater temperature, whereas the apparent growth yield of phytoplankton declined with increasing temperature from 0.19 µg chl a dm-3 d-1 at 54° N to 0.01 µg chl a dm-3 d-1 at 33° N. Complete nitrogen saturation of phytoplankton growth indicated light or non-nitrogenous limitation at the nitracline at 47° N and in the deep chlorophyll maximum at 33° N, whereas in the mixed layer at 47° N and 54° N the ambient nitrogen supply was sub-saturated and yielded 63 and 39% of nitrogen- saturated growth. Nitrogen supply of phytoplankton growth was dominated by external and cellular sources in nitrate-rich waters of the mixed layer at 54° N and at the nitracline at 47° N, whereas nitrogen regeneration dominated at the nitrate-depleted surface waters at 47° N. However, in the deep chlorophyll maxima at 33° N and 40° N phytoplankton growth was primarily maintained by nitrogen regeneration, although external nitrogen was sufficiently available. The recycling efficiency of the microbial community was defined as the ratio of regenerated growth yield to herbivorous grazing loss. Efficiencies of ~100% under post-bloom situations indicated tight coupling of predation, nitrogen supply and phytoplankton growth. We suggest that microzooplankton grazing has a high potential for nitrogen supply and biomass control of phytoplankton communities during summer in the temperate and subtropical northeast Atlantic.

Description: The only known population of coelacanths, in the Comores, western Indian Ocean, is endangered by human predation. Historical catch data from Grande Comore reveal that annual catch rates increased steadily from 1954 until the 1970s. This trend was temporarily interrupted due to an international policy introducing motorized boats and promoting offshore fishing techniques. Coelacanths are only caught from traditional unmotorized outrigger canoes as an incidental by-catch of deep water line fishing. A complete survey of all motorized and unmotorized vessels in 1995 at Grande Comore in comparison to earlier years indicated that a recent decreased use of motors and increase of unmotorized canoe fishing has led to an increase in coelacanth catches. Conservation measures and strategies for reducing the fishing pressure exerted on coelacanths are discussed. The southwest coast of Grande Comore should be designated as a nature reserve and protected area where immediate protection measures should be taken, an opinion which is supported by Comorian authorities.

Description: VHF telemetry was used in November and December 1995 on 8 Humboldt penguins Spheniscus humboldti breeding at Pan de Azúcar Island (26°S, 70°W), Northern Chile, to determine at-sea behaviour of the birds. We obtained 2710 locations, 90% of which were within a radius of 20 km around the island. Mean travelling speed of the birds was 0.92 m s-1 and speed distribution showed peaks at 1.6 and 3 m s-1. Penguins travelling between foraging areas remained submerged for an average of 8.4 s between surfacings, whereas foraging dives lasted on average 61 s. The analysis of 79 complete foraging trips showed that tracks deviated from a straight course, and range (maximum distance from island) was only 0.37 times total horizontal distance swum. Birds did not forage synchronously or in the same foraging areas. However, foraging ranges were correlated between birds, indicating similar search strategies during periods of low food availability. The results obtained here via VHF telemetry agreed well with those of previous studies employing satellite transmitters and data loggers.

Description: The stable carbon isotope composition (δ13C) of particulate organic carbon (POC) was measured in 3 size fractions (POCtotal, POC〉20 µm, POC〈20 µm) during a phytoplankton spring bloom dominated by the diatom Skeletonema costatum in Lindåspollene, a land-locked fjord in southern Norway. In addition to standard parameters for characterizing the phytoplankton bloom (chlorophyll, nutrient, and POC concentrations, and species composition), simultaneous measurements of δ13C of dissolved inorganic carbon (DIC), total alkalinity and DIC concentration were obtained to determine temporal trends in dissolved carbon dioxide concentration and in carbon isotope fractionation (ε p) of the POC size fractions. The carbon isotope composition of the 〉20 µm size fraction, which was dominated by diatoms, was ca 2o/oo heavier than that of the 〈20 µm fraction, which was mainly composed of flagellates. δ13C of both size fractions increased by about 3o/oo over the course of the bloom. A 5o/oo increase in δ13C-PO Ctotal during the bloom resulted partly from a shift in the phytoplankton community from a flagellate- to a diatom-dominated one. Carbon isotope fractionation of all fractions decreased with declining CO2(aq) concentration (14 to 〉6 µmol l-1). A positive correlation between ε p and [CO2(aq)] in the diatom size fraction was obtained for the period of exponential growth. Deviation from this correlation occurred after the peak in cell density and chlorphyll a (chl a) concentration, when POC still continued to increase, and may be related to changing phytoplankton growth rates or to possible effects of nutrient (nitrate) limitation on ε p. Comparison of these results with those of previous field studies shows that, while an inverse relationship is consistently observed between ε p and the ratio of instantaneous growth rate and CO2 concentration {µi/[CO2(aq)]}, considerable scatter exists in this relationship. While this scatter may have partly resulted from inconsistencies between the different studies in estimating phytoplankton growth rate, it could also reflect that factors other than growth rate and CO2 concentration significantly contribute to determining isotope fractionation by marine phytoplankton in the natural environment

Description: A total of 354 adult European smelts Osmerus eperlanus (L.) were tested for their ability to survive the screen system of the cooling water inflow of a power plant. With increasing number of musculature parasitic third-stage larvae of Pseudoterranova decipiens, the survival rate of O. eperlanus decreased while the total number of externally visible injuries as well as the number of seriously injured specimens increased. The results indicate that even a single specimen of P. decipiens influences resistance and stamina and affects overall mortality of 7 to 20 cm long smelts. The initial effect of the parasites is to reduce swimming speed of infested fish, which leads to more frequent contact of these fish with the fine meshed screen of the cooling water inlet before they are removed by the automatic cleaning system. If the separated fishes are returned to the main stream, it becomes apparent that the cooling water inflow selectively reduces the number of living parasitised smelt in the area. Thus, the number of parasitic third-stage P. decipiens larvae in the local smelt population which are able to complete their life-cycle is also reduced. P. decipiens makes infested smelt more susceptible to negative anthropogenic influences such as cooling water intake or trawl fisheries.

Description: During a transition period from oxic to anoxic conditions in the bottom water, rates of sulfate reduction and methane production, methane fluxes, as well as concentration profiles of sulfate, sulfide and methane were measured in sediments at a central site of the Gotland Deep (Stn AL 93, 241 m depth), which is regarded as representative for the deepest part of this basin. During this period from 1993 to 1996 oxic conditions in the bottom water prevailed from spring 1994 until summer 1995 with oxygen concentrations decreasing progressively with time. In the sediments methane production occurred primarily in layers below 1 m depth and flux rates of methane to the sediment surface were characterized by a steep concentration gradient from approx. 5 mM at 4 m depth to values close to 30 μM at the surface, determined by diffusion processes and anaerobic oxidation of methane. Both processes were independent of changes at the sediment surface. Differences in the flux rates of methane between the deeper part with a mean value of 259 μmol m-2 d-1 and the upper layers with a mean of 47.7 μmol m-2 d-1 indicate that a considerable proportion of the methane is oxidized within the anoxic horizon of the sediment (71 to 86% in the layer from 40 to 70 cm). Low rates of methane production found within the top 20 cm of the sediment during periods of oxic bottom water increased after depletion of oxygen and resulted in a clear maximum of the methane concentration in the top 2 cm. Sulfate concentrations declined exponentially from values of 11.5 mM in June 1994 and 8.5 mM in October 1995 at the sediment surface to values of 2.5 mM at 20 cm depth and of less than 0.5 mM at 50 to 60 cm depth. High sulfate reduction rates (150 to 250 nmol cm-3 d-1) in the upper part of the sediment (8 to 13 cm) coincided with maxima of sulfide concentrations. During the time period of this investigation an increase of maximum sulfide concentrations in the sediment from 1 to 10 mM was measured together with decreasing oxygen concentrations in the deep water. At the same time sulfate reduction established a small but distinct maximum at the top layer of the sediment (0 to 2 cm). The relative importance of sulfate reduction and methanogenesis in the carbon budget of the Gotland Deep sediments is calculated on the basis of the actual measurements.

Description: In steady state, the export of photosynthetically fixed organic matter to the deep ocean has to be balanced by an upward flux of nutrients into the euphotic zone1. Indirect geochemical estimates2 of the nutrient supply to surface waters have been substantially higher than direct biological and physical measurements3, particularly in subtropical regions. A possible explanation for the apparent discrepancy is that the sampling strategy of the direct measurements has under-represented episodic nutrient injections forced by mesoscale eddy dynamics, whereas geochemical tracer budgets integrate fluxes over longer time and space scales. Here we investigate the eddy-induced nutrient supply by combining two methods potentially capable of delivering synoptic descriptions of the ocean's state on a basin scale. Remotely sensed sea-surface height data from the simultaneous TOPEX/Poseidon and ERS-1 satellite missions are assimilated into a numerical eddy-resolving coupled ecosystem–circulation model of the North Atlantic Ocean. Our results indicate that mesoscale eddy activity accounts for about one-third of the total flux of nitrate into the euphotic zone (taken to represent new production) in the subtropics and at mid-latitudes. This contribution is not sufficient to maintain the observed primary production in parts of the subtropical gyre, where alternative routes of nitrogen supply will have to be considered.

Description: The ability to monitor the heat content of oceans over long distances is becoming increasingly important for understanding the role of oceans in climate change, for determining the variability of the state of the oceans, for operational ocean observing systems, and for studying large-scale ocean processes such as water-mass formation. Although the properties of the upper layers of the ocean can be routinely measured on large scales by satellite remote sensing (providing altimetric and infrared data) and with expendable probes dropped from commercial vessels, the deep interior of the ocean is more difficult to monitor. Ocean acoustic tomography1 is a promising technique for such applications, as it has the potential to provide systematic, instantaneous and repeated measurements of the ocean interior over large parts of an ocean basin. Here we demonstrate the capability of this technique for measuring the heat content across an entire (albeit small) ocean basin—the western Mediterranean Sea.

Description: The effect of variable CO2 concentrations on the elemental composition (C:N:P) of marine diatoms was investigated in 2 strains of Skeletonema costatum (Grev.) Cleve. Five or 6 concentrations of dissolved molecular carbon dioxide [CO2 (aq)], ranging from 0.5 to 39 µmol l-1, were applied in dilute batch cultures. In both strains, elemental ratios were clearly dependent on [CO2 (aq)]. With decreasing CO2 concentrations, a decline in C:P and N:P and an increase in C:N was observed. The close correlation between C:P or N:P and [CO2 (aq)] corresponded to a ca 45 to 65% decrease in elemental ratios from highest (〉=30 µmol l-1) to lowest (ca 1 µmol l-1) CO2 concentrations. C:N at low [CO2 (aq)] was up to 24% higher than at high [CO2 (aq)]. To date, the elemental composition of marine phytoplankton has been considered to be independent of CO2 availability. If dependency of the C:N:P ratio on [CO2 (aq)] proves to be a general phenomenon in marine phytoplankton, changes in the elemental composition may be expected in response to the currently observed increase in partial pressure of atmospheric CO2.

Description: The ocean stores and transports vast quantities of heat, fresh water, carbon and other materials, and its circulation plays an important role in determining both the Earth's climate and fundamental processes in the biosphere. Understanding the development of climate and important biological cycles therefore requires detailed knowledge of ocean circulation and its transport properties. This cannot be achieved solely through modelling, but must involve accurate observations of the spatio-temporal evolution of the global oceanic flow field. Estimates of oceanic flow are currently made on the basis of space-borne measurements of the sea surface, and monitoring of the ocean interior. Satellite altimetry and acoustic tomography are complementary for this purpose1, as the former provides detailed horizontal coverage of the surface, and the latter the requisite vertical sampling of the interior. High-quality acoustic-tomographic2 and altimetric3 data are now available to test the combined power of these technologies for estimating oceanic flows. Here we demonstrate that, with the aid of state-of-the-art numerical models, it is possible to recover from these data a detailed spatio-temporal record of flow over basin-scale volumes of fluid. Our present results are restricted to the Mediterranean Sea, but the method described here provides a powerful tool for studying oceanic circulation worldwide.

Description: The response of epilithic microphytobenthos to in situ nutrient enrichments was studied in the Kiel Fjord, Western Baltic Sea. For this purpose an experimental setup with continuous nutrient supply was designed and installed. Experiments followed the colonization of artificial substrates and the responses of benthic algae to different concentrations and combinations of nitrogen and phosphate. They revealed nitrogen limitation of epilithic microflora from late spring to autumn, such that there was higher biovolume with increasing nitrogen concentrations. Diatoms became dominant in all experiments except one in which the rhodophyte Ceramium strictum prevailed. Species composition was altered by nutrient treatments; one to several species were highly favoured by nutrient enrichment. Consequently, diversity was negatively correlated with final yield. These findings support the hypothesis that competition is an important factor structuring microphytobenthic communities.

Description: Snails Littorina littorea with and without artificial epibionts were exposed in situ for nearly 1 mo. Measurement of individual shell length increase over this period revealed that clean snails grew 3 times faster than fouled snails. These results compare well with previously conducted flume experiments (Wahl 1996). This epibiosis effect is thought to be due to increased drag-the only feature distinguishing the 2 treatment groups-caused by the presence of epibionts on the shell. Increased drag probably entails higher energy expenditure for pedal activities (attachment and locomotion) and a reduced allocation of resources towards growth and, possibly, reproduction

Description: Stomach contents from 30 long-finned pilot whales Globicephala melas captured incidentally in the Distant Water Fleet (DWF) mackerel fishery off the northeastern United States were examined. Several methods of assessing prey importance were used in order to construct a true representation of the pilot whale diet. Separate analyses of trace (free, durable body parts from well-digested prey) and non-trace (relatively intact prey) food materials were conducted to address biases caused by differential rates of digestion and passage. Squids dominated the diet and long-finned squid Loligo pealei was the most important prey, but we noted large yearly fluctuations in prey importance. Metric multidimensional scaling analyses of trace and non-trace stomach contents of individual whales suggest that many animals were cuaght while feeding opportunistically near fishing operations, resulting in a bias of non-trace (intact) stomach contents. The diversity of prey in this study was greater than previous reports of the food habits of western North Atlantic non-finned pilot whales.

Description: Recent captures of two female giant squid ( Architeuthis ) off southern Australia have provided the first record of a mated female specimen of these almost mythical deepsea creatures. We found sperm packages (spermatophores) embedded within the skin of both ventral arms of the larger of the two specimens. It seems that male giant squids may use their muscular elongate penis to ‘inject’ sperm packages under pressure directly into the arms of females.

Description: We investigated the foraging location, diving behaviour, dietary composition, feeding rates and foraging trip durations of emperor penguins Aptenodytes forsteri raising chicks at the Auster and Taylor Glacier colonies on the Mawson Coast of Antarctica in the winter, spring and early summer of 1993, to examine seasonal changes in the penguins' foraging ecology. As day-length increased after winter, the penguins' daily swimming time increased from 7.83 ± 1.50 h in August to 12.23 ± 1.25 h in September and 12.95 ± 1.24 h in October. Accordingly, the penguins' dive rate increased from 92.7 ± 28.5 to 149.4 ± 23.4 and 161.6 ± 19.3 dives d-1 in the respective months. The birds targeted prey in the vicinity of the continental slope mainly at depths 〈100 m, although some individuals frequently hunted at depths 〉200 m, and the maximum depth achieved was 438 m. Antarctic krill Euphausia superba were the most common prey taken overall, 41% of the diet by mass, and dominated the diets between August and October. The contribution of Antarctic krill to the diet reduced over time from 68% in August to 1% in early December. In November, the glacier squid Psychroteuthis glacialis dominated the diet (47 to 63%), and in early December the diet comprised various species of fish, Trematomus species (27%), Pagothenia borchgrevinki (24%), and Pleuragramma antarcticum (8%), and squid, P. glacialis (13%) and Alluroteuthis antarcticus (9%). The birds' prey consumption rates more than doubled between late winter and early summer, from 4.0 ± 1.0 to 8.7 ± 1.7 kg d-1 spent foraging; these values are equivalent to metabolisable energy intakes of 628 ± 134 and 1422 ± 308 kJ kg-1 d-1, respectively. During brooding (late winter to early spring), females spent less time at sea than males (8.7 ± 2.7 vs 17.7 ± 3.8 d); thereafter trip durations of both sexes were similar and declined from 15-19 d in spring to 〈10 d in early summer. Between hatching and about 1 wk prior to fledging each parent fed its chick 7 or 8 times. To raise a chick, females and males consumed approximately 410 and 470 kg of prey respectively, or 880 kg for each breeding pair. Seasonal variations in the penguins' foraging were probably influenced by fluctuating sea-ice conditions, differences in the prey types available, changes in day-length toward summer, and increasing demands of the growing chicks.

Description: The nitrogen-isotope record preserved in Southern Ocean sediments, along with several geochemical tracers for the settling fluxes of biogenic matter, reveals patterns of past nutrient supply to phytoplankton and surface-water stratification in this oceanic region. Areal averaging of these spatial patterns indicates that reduction of the CO2 'leak' from ocean to atmosphere by increased surface-water stratification south of the Polar Front made a greater contribution to the lowering of atmospheric CO2 concentration during the Last Glacial Maximum than did the increased export of organic carbon from surface to deep waters occurring further north.

Description: A predictive model of carbon isotope fractionation (sigma p) and abundance (delta13C phyto) is presented under circumstances where photosynthesis is strictly based on CO2(aq) that passively diffuses into marine phytoplankton cells. Similar to other recent models, the one presented here is based on a formulation where the expression of intracellular enzymatic isotope fractionation relative to that imposed by CO2(aq) transport is scaled by the ratio of intracellular to external [CO2(aq)], ci/ce. Unlike previous models, an explicit calculation of ci is made that is dependent on ce as well as cell radius, cell growth rate, cell membrane permeability to CO2(aq), temperature, and, to a limited extent, pH and salinity. This allows direct scaling of ci/ce to each of these factors, and thus a direct prediction of sigma p and delta13C phyto responses to changes in each of these variables. These responses are described, and, where possible, compared to recent experimental and previous modeling results.

Description: The conventional model whereby plume volcanism forms linear age-progressive volcanic chains, with the youngest activity occurring nearest a spreading axis (at a 'hotspot'), has been challenged for the Easter seamount chain1–4. Whereas early work suggested the existence of a linear melting anomaly (a 'hotline')1,2, more recent studies3,4 have proposed a hotspot near Salas y Gomez island, connected with the Easter microplate spreading system by an ~800-km-long, volcanically active plume channel. Here we use geochemical, geological and geochronological data to argue that the hotspot lies close to Easter Island. Moreover, new isotopic data for lavas from the seamount chain provide evidence for bidirectional flow between the spreading axis and the plume, thus supporting geophysical and fluid-dynamical models of mantle flow in a plume/spreading axis system5–7. Material balance and flux considerations show the Easter plume to be weak and cool compared with those beneath larger features such as Iceland, Hawaii and the Galápagos islands.

Description: A knowledge of past changes in the biological productivity of the oceans is important for understanding the interactions between carbon cycling and climate. Phytoplankton productivity in today's oceans can be estimated from the concentrations of chlorophyll in sea water1, but chlorophyll is not preserved in the sediments. Existing proxies for past algal productivity do not represent total productivity; for example, biogenic opal2 reflects the contribution of only part of the phytoplankton community, and the organic carbon record can be subject to contamination from terrestrial inputs2,3. Although chlorins, the pigment-transformation products of chlorophyll, are widespread in Quaternary marine sediments, their potential as proxy measures of past variations in primary productivity has not been convincingly demonstrated. Here we report a high-resolution molecular stratigraphic record of chlorin concentrations over the past 350,000 years in a sediment core from the subtropical Atlantic continental margin. Maxima in the chlorin accumulation rate coincide with significant peaks in the accumulation rates of biogenic opal (at the end of glacial terminations) and organic carbon (between terminations). These results suggest that chlorins, unlike other proxies, can serve as a measure of total primary productivity variations.

Description: Nutrient competition experiments were performed with periphytic microalgae from the Kiel Fjord, western Baltic Sea. Diatoms were dominant competitors at high Si:N ratios (under N-limited conditions) and high Si:P ratios (under P-limited conditions). Under low silicate supply, cyanobacteria became dominant at low N:P ratios and chlorophytes at high ones. Changes in light intensity influenced the outcome of competition at the species level but not at the level of higher taxa. The nitrogen source (ammonium or nitrate) had no impact on the outcome of competition.

Description: Cod larvae from laboratory rearing experiments aged from 1 to 12 d after hatching, both fed and deprived of food, were analysed. The number of increments on the otolith and the width of these increments were determined together with the RNA/DNA ratios on the same individual larva. Alizarin marking of the otoliths was performed to confirm the formation of daily increments. Cod larvae reared at 6*C formed the first ring right after hatching and deposited increments on a daily basis. A comparison of the measurements between the right and the left lapillus showed that these can differ, if the radius is taken. The lapillae core showed especially high individual variability, whereas the sum of the increments did not differ between both lapilli. Until Day 10 after hatching, while the larvae were still feeding on their yolk, the external food situation did not affect the increment width of the lapilli or the RNA/DNA ratios. In larvae older than 10 d the width of the daily increments was dependent on the nutritional situation and RNA/DNA ratios decreased in starving larvae in comparison to feeding larvae. RNA/DNA ratios and increment widths were correlated.

Description: Mixed cultures of 4 polar diatoms regularly found in Antarctic pack ice were grown over 20 d in closed bottles at high light (200 to 250 umol photons m-2 s-1) and at 0*C in order to investigate growth physiology and biomass production under conditions simulating the sea ice habitat during summer. Species tested were: Chaetoceros cf. neogracile, Fragilariopsiscylindrus, Thalassiosiraantarctica and Porosira pseudodenticulata. Initially, all species grew exponentially, but exponential growth ceased for P. pseudodenticulata and T.antarctica after 6 d, for F.cylindrus after 8 d, and for C. cf. neogracile after 10 d. Slight increases in cell number were observed for all species 2 d later. Peak biomass amounted to 140 ug chl a (850 umol particulate organic carbon, POC) l-1. At the same time, concentrations of dissolved inorganic carbon (DIC) were reduced by 1000 uM, oxygen concentrations increased to 1400 uM, and pH increased to 10.5. At this stage, a substantial decline in plasma-containing cells was recorded for F.cylindrus. C. cf. neogracile accounted for 80%, and C. cf. neogracile and F.cylindrus accounted for 〉95% of total carbon biomass. The carbon isotope composition of POC (expressed as delta13C) increased from -24 to -9 during the experiment. Model calculations showed that diffusive uptake of dissolved CO2 satisfied cellular carbon demand for all species except P. pseudodenticulata at CO2(aq) concentrations 〉0.5 uM, whereas direct HCO3- utilization was observed for C. cf. neogracile below this concentration. Our data confirm that intense photosynthetic carbon assimilation may lead to profound chemical changes in isolated interstitial brine solutions, with significant consequences for sea ice biota. We propose that the capacity to efficiently utilize ambient DIC, possibly mediated by virtue of favorable surface to volume ratios as well as active pathways of inorganic carbon acquisition, favors growth of small diatoms, and may be an important factor driving ice algal species succession during summer blooms. Since only 2 species continued to grow in fresh medium following experimental incubation (C. cf. neogracile and P. pseudodenticulata), differential tolerance to chemical variations may influence the seeding potential of ice algae following release into the open water.

Description: A fundamental issue in marine science is the identification of the factors controlling biological uptake of CO2, in high-nitrate, low-chlorophyll regions. A recent in situ iron fertilization experiment demonstrated that iron limitation is responsible for low phytoplankton stocks in the equatorial Pacific4. Here we show that flavodoxin, a biochemical marker of iron limitation, can be used to map the degree of iron stress in natural populations. Flavodoxin assays along a 900-km east-west transect in the northeastern subarctic Pacific revealed a pronounced increase in iron stress in the region west of the 135° W meridian. Addition of dissolved iron alleviated this stress. Immunostaining of single cells from the most western station showed that flavodoxin is present specifically within the chloroplasts of diatoms. Our approach provides a rapid means of defining the extent of iron stress in the ocean5 and supports the hypothesis that diatoms are iron stressed in the northeast Pacific.

Description: Epibiosis is one of the closest interspecies associations. The presence of epibionts potentially causes a multitude of beneficial or detrimental effects for the basibiont. It has been shown previously that large epibionts may increase the risk of dislodgement of bivalves. In this study, sublethal effects of epibiont-induced drag increase are investigated. I assessed (1) the effects of common epibiont species (Balanus improvisus, Enteromorpha intestinalis, Ectocarpus sp.) on drag properties of the host (the periwinkle Littorina littorea), and (2) the long-term consequences of drag increase on growth rates of snails living in steady flow. All epibiont species increase drag on the host snail. They do so to unequal extents. This may be due to morphological and hydrodynamic differences among the epibionts. Thus, per unit volume of epibiont, the filamentous alga Ectocarpus sp, has a substantially stronger effect than the barnacles. Synergistic effects on drag increase can be observed in a mixed aufwuchs community. As compared to clean conspecifics, snails bearing artificial epibionts grow 35% more slowly when exposed to moderate, steady flow (8 cm s(-1)) for 5 mo. This difference in growth rates is enhanced when food is limited. I hypothesize that fouled snails coping with higher drag invest more energy into foot activities (muscles and mucus). As a consequence, when food is limited, growth rates decrease in fouled snails.

Description: The dynamics of methane production and oxidation in the sediments of Kiel Harbour (Baltic Sea) were studied over a period of 2 yr. Experimentally determined rates of methanogenesis varied between 13 and 82 mu mol CH4 m(-2) h(-1) integrated over the top 30 cm of sediment. The zone of maximum production was between 20 and 30 cm depth. High potential rates could be induced in sulfate-containing surface layers by the addition of molybdate, acetate or methanol, indicating competition with sulfate-reducing bacteria. The methane content in the sediment pore water increased from the surface to a seasonally varying depth of between 5 and 20 cm, where sulfate concentrations approached zero and methane concentrations of 3 to 4 mM were recorded. Aerobic methane oxidation rates in the surface sediment layer were in most cases higher than the methane flux expected to reach the sediment surface. Oxygen-depleted or anoxic conditions in the deep water, which frequently occurred during stratification periods in summer, resulted in methane gradients with increasing concentrations from the chemocline to the sediment. After recirculation of the water column in autumn in both years, unexpectedly high methane concentrations were also measured in the oxic water column of Kiel Harbour. This coincided with very reduced conditions in the surface sediment that allowed measurable methane production in the 0 to 10 cm layer.

Description: It has often been suggested that, given their large food requirements, sperm whale Physeter macrocephalus distributions should reflect the distribution of productive ocean environments, and it seems therefore that chlorophyll concentration might be a good indicator of sperm whale distribution. To examine the existence of such a relationship, and to determine over which sclaes it occurs, sperm whale density was correlated with phytoplankton pigment concentration over a range of spatial and temporal scales. Sperm whale distribution was detrmined using records of kills from 19th century Yankee whaling, and the distribution of pigment concentration from satellite colour observations averaged over 8yr interval. These measures were compared over scales of 220km square to 1780km square. The distribution of sperm whales in the temperature and tropical Pacific Ocean was associated with distributions of phytoplankton pigment over every spatial scale considered, and the coefficient of correlation increased with increasing spatial scale. However, a few exceptions to this scheme were found, implying that other factors would be of importance in some regions. This study confirmed the existence of space lag and a time lag between a peak in chlorophyll concentration and a peak in sperm whale density. It also demonstrated that over large spatial scales, and when the data are averaged over large temporal scales, chlorophyll concentration is a good indicator of sperm whale distribution and that over these scales ocean colour recorded from space could help predict areas of high or low sperm whale density.

Description: Predator data and exploratory fishing in the Scotia Sea have revealed the presence of cephalopod stocks in the Antarctic Polar Frontal Zone (PFZ). This is a vast, remote region where large epipelagic cephalopods aggregate into highly mobile schools making them difficult to locate and sample. We used satellite tagged predators and shipboard acoustics for coarse and fine scale location of cephalopods concentrations, and sampled them with commercial and scientific nets to determine the relationship between cephalopod distribution and mesoscale oceanographic features at the PFZ. Saltellite tags were attached to 9 grey-headed albatrosses Diomedea chrysostoma, breeding at Bird Island, South Georgia, to monitor foraging at sea in January-March 1994. A foraging area at the PFZ, north of South Georgia, was located, an acoustic survey undertaken and a fixed situation established where gular midwater trawl 25m² (RTM25), a horizontal multiple plankton sampler and a neuston net. Acoustic layers were targeted and the RMT25 sampled 200m layers to 1000m in daylight and darkness. Cephalopods were simultaneously recovered from food samples fed to D. chrysostoma chicks at Bird Island. Two CTD transects, approximately normal to the major current flow, were undertaken across the PFZ and remote-sensed-sea-surface temperature images from NOAA polar orbiting satellites that exploited by D. chrysostoma. The largest and most conspicious species was the ommastrphid squid Martialia hyadesi which is the most important cephalopod prey species. Net-sampled M. hyadesi had been feeding on crustaceans and mesopelagic fish. The cephalopod community was sampled in a feature, interpreted as a warm core ring, in an area characterised by mesoscale features associated with the bathymetry of the northern end of the Northeast Georgia Rise and near a gap in the Falkland Ridge. The association of these mesoscale features with the bathymetry suggests that they may be predictable foraging locations for the cephalopods and their predators.

Description: The tissue of 31 demosponge and 7 hexachnelhd species was analyzed for its composition of organic and lnorganic matter With one excephon (Hahclona cf gausaana) inorganic matter i e n~ostly the siliceous skeleton, accounted for most of the dry weight, varying between about 60 and 95%dry wt There were no general trends in the ratio of organic to inorganic matter within sponge orders or genera, and within one species, the ratio could vary between stations For one of the hexactinellids Bathydorus spmosus, several size classes were analyzed and there was no systematic change in the organic inorganic matter ratio w t h specimen size For some species the results from the Weddell Sea sponges are in good agreement with earlier data from McMurdo Sound The low organic matter content in Weddell Sea sponges mphes that sponge biomasses are much lower than hitherto assumed on the basis of then high abundances and large sizes In consequence Antarctic sponges despite their ubiquitousness, may only channel a m n o r fractlon of the general bentho-pelagic flow of matter and energy and their maln role in the ecosystem is likely structural rather than dynamic

Description: During 5 consecutive summer seasons (1990 to 1994) both Aureliaaurita medusa and mesozooplankton abundances were investigated at 2 stations in the Kiel Bight (western Baltic Sea). Stocks of medusae varied considerably between the years, with median abundance of 1 and 0.3 ind. 100 m-3 in 1990 and 1991, 3 and 4 ind. 100 m-3 in 1992 and 1994, but 9 ind. 100 m-3 in 1993. Significant differences in the mesozooplankton stock and community composition were observed in 1993 when compared with the year of rather low Aurelia abundance (1991). Total zooplankton and copepod numbers both exhibited an inverse relationship with the abundance of medusae and were thus considerably lower in 1993 than in 1991. However, not all copepod species were affected by A. aurita. Pseudo- and Paracalanus spp. and Oithonasimilis showed dramatically reduced stocks in the bloom year when compared with the medusa-poor situation, but no significant changes were found for Centropageshamatus and Acartia spp. Also other zooplankton groups with the exception of bivalve larvae were reduced by the medusae. The differential response of zooplankton to varying abundance of medusae led to a shift in the trophic structure of the zooplankton community. Fine-filter feeders and raptorial feeders were much more important in years when medusae occurred in low densities, whereas coarse-filter feeders dominated in the opposite situation.

Description: From the Gulf of Aden along a transect to the central-northern Red Sea the abundance and taxonomic composition of metazoan plankton was studied during the southwest monsoon period (summer 1987). Samples were taken with 0.055 mm mesh nets down to a maximum depth of 1050 m. In the epipelagic zone, a distinct decrease in total plankton abundance was observed from south to north, which was much more pronounced in biomass (by a factor of up to 10) as compared to numbers (by a factor of 2). This could partly be explained by differences in the taxonomic and/or size composition of the planktonic fauna. Among non-calanoid copepods, 40 out of 75 species or taxa investigated decreased in abundance from south to north. Sixteen of these species were completely absent in the central-northern area. Nineteen species or taxa, however, showed the opposite feature of a higher abundance in the central-northern Red Sea. The stations were grouped according to similarities in the taxonomic composition of non-calanoid copepods in the epipelagic zone. The following 3 geographical regions could be separated: (1) Gulf of Aden and Strait of Bab al Mandab; (2) southern Red Sea; and (3) central-northern Red Sea. In the meso- and bathypelagic zones, regional differences were not evident. The results are discussed in relation to hydrographic conditions during summer 1987.

Description: A spring investigation of the phytoplankton in the western Alboran Sea (Mediterranean) was undertaken using chlorophyll and carotenoid biomarkers to characterize the community in the water column and in drifting sediment traps set at 100 and 200 m. During 2 drifter experiments, calm and sunny conditions induced a progressive thermal stratification that reduced pigment sedimentation into deeper water and confined the phytoplankton to the surface layer, resulting in an increase in chlorophyll biomass. 19'-Hexanoyloxyfucoxanthin (prymnesiophytes) and chlorophyll b (chlorophytes, prasinophytes, prochlorophytes) were the major accessory pigments, while fucoxanthin, alloxanthin and peridinin indicated the presence of diatoms, cryptophytes and dinoflagellates, respectively. The proportional contribution of each algal group to the chlorophyll a (chl a) biomass, as derived from multiple regression analysis, revealed that prymnesiophytes, cryptophytes and the green algal group collectively accounted for at least 75% in the upper 100 m, emphasizing the importance of the nanophytoplankton. Phaeopigments, dominated by phaeophorbide a2, were the main pigments observed in sediment traps, although chl a, fucoxanthin and 19'-hexanoyloxyfucoxanthin were detected in smaller concentrations as well as traces of chlorophyll b (chl b). In deep water, fucoxanthin and 19'-hexanoyloxyfucoxanthin were the only accessory pigments present while total phaeopigment/chl a molar ratios 〉1 reflected the active transformation of fine phytogenic material at depth. High particulate organic carbon (POC)/chl a ratios (〉100 in surface water; 〉1000 in deep water) suggested that phytoplankton was a relatively small component of the total carbon biomass down the water column. Using simple budget calculations, we determined that 58 to 65% of the chl a produced in the upper 100 m accumulated in the water column over both experiments. During Expt 1, 29% of the chl a sedimented out, mostly as phaeopigment, at 100 m (24%), and 6% was degraded to colourless residues in the water column. In contrast, only 12% of the chl a sedimented in Expt 2, while 20% was degraded to colourless residues.

Description: Sediment cores of 20 cm diameter contaning the natural benthic fauna were subjected to low oxygen conditions in a laboratory microcosm system. After several days of oxic conditions ('oxic stage') the oxygen content of the water was reduced to 25% saturation for 15 d ('hypoxic stage'), followed by a 'reoxygenation stage'. Effective solute transport rates were calculated using measurements with the conservative tracer ion bromide. Profiles of oxygen and ΣCO2 were measured and molecular diffusive as well as effective fluxes, account mg for effective solute exchange, were calculated. The overall response of the benthic community was to compensate for low oxygen content of the overlying water by increased pumping activity. On average, effective diffusion coefficients (Den} were 3 times higher in hypoxia than under oxic conditions. D eff reached 1.5 x 10^-4 cm2 s^-1, a value 30 times that of molecular diffusion. During hypoxia we observed low molecular diffusive O2 flux, higher effective O2 flux, as well as an increase in ΣCO2 within the sediment. We interpret this as a shift of transport away from diffusion within the bulk sediment interstices (oxic conditions) to the advective transport pathways along burrows during hypoxia. This facilitates fast transport of oxygen and bromide along burrows and contrasts with the slower transport of CO2 from the interstices governed by molecular diffusion. In this transient situation calulations based on gradients result in an unrealistic molar ratio of fluxes(CO2/O2)as high as 11.

Description: This experimental study investigated the vertical migratory response of benthic forammifera in sediments that were initially oxygenated, to variations in dissolved-oxygen concentrations ranging from well-oxygenated to dysaerobic conditions. Two box cores, with a carpet of polychaete tubes extending above the sediment-seawater interface, were recovered from 71 m water depth in Oslofjord, southern Norway. The seawater oxygen concentration of one box core was decreased every 4 wk, to a minimum value of 〈0.2 ml 0; 1'. Aerated seawater circulation was maintained in the other (control) box core. In a time course, 2 subcores were taken from each mesocosm every 4 wk and sectioned in 0 5 cm mtervals to depths of 2.0 cm, and 1.0 cm intervals from 2.0 to 4.0 cm. The portion of the polychaete tubes extending into the overlymg water was collected and treated as an additional sample. Live foramimferal distributions were examined in the 〉 63 pm fraction of each interval shallower than 2 cm using both an ATP assay and a rose Bengal staining method, while only rose Bengal was used to assess distributions from 2 to 4 cm. Results indicate that larger numbers of live and stained forammifera are found on the polychaete tubes and in the superficial 0.5 cm of sediment after exposure to dysaerobic conditions compared to original and control abundances. After re-oxygenation for 4 mo, the foraminifera re-migrated into the sediments, exhibiting distributions similar to those of the control mesocosm. These observations suggest that some species actively migrate to a microenvironment with a particular oxygen concentration, rather than maintaining a stable position with respect to the sediment-water interface.

Description: Seismic tomography and the isotope geochemistry of Cenozoic volcanic rocks suggest the existence of a large, sheet-like region of upwelling in the upper mantle which extends from the eastern Atlantic Ocean to central Europe and the western Mediterranean. A belt of extension and rifting in the latter two areas appears to lie above the intersection of the centre of the upwelling region with the base of the lithosphere. Lead, strontium and neodymium isotope data for all three regions converge on a restricted composition, inferred to be that of the upwelling mantle.

Description: A survey of parasites in 600 short-finned squid Illex coindetii (Vérany, 1839) taken from 2 locations (north and south Galicia) off the northwesterb Iberian Peninsula revealed the presence of numerous somatoxenous helminths. Three genera of Tetraphyllidean plerocercoids were represented (prevalences: Phyllobothrium sp., 45,7%; Dinobothrium sp., 0.8%; and Pelichnibothrium speciosum, 0.001%); 1 Trypanorhynchidean metacestode was also present (Nybelinia yamagutii, 0.4%). In addition, larval nematodes of Anisakis simplex (L3) were recorded (10.6%). Abundance of infection was examined in relation to squid sex, standard length, maturity and locality. This analysis indicated that parasite infection was lower in the southern squids than in the northern squid group. Over the entire survey area, parasite infection showed a positive correlation with host life.cycle, often with the greatest number of parasites among the largest and highest maturity individuals (〉18 to 20cm; maturity stage V).

Description: THE hydrothermal circulation of sea water through permeable ocean crust results in rock–water interactions that lead to the formation of massive sulphide deposits. These are the modern analogues of many ancient ophiolite-hosted deposits1–4, such as those exposed in Cyprus. Here we report results obtained from drilling a series of holes into an actively forming sulphide deposit on the Mid-Atlantic Ridge. A complex assemblage of sulphide–anhydrite–silica breccias provides striking evidence that such hydrothermal mounds do not grow simply by the accumulation of sulphides on the sea floor. Indeed, the deposit grows largely as an in situ breccia pile, as successive episodes of hydrothermal activity each form new hydrothermal precipitates and cement earlier deposits. During inactive periods, the collapse of sulphide chimneys, dissolution of anhydrite, and disruption by faulting cause brecciation of the deposit. The abundance of anhydrite beneath the present region of focused hydrothermal venting reflects the high temperatures ( 〉 150 °C) currently maintained within the mound, and implies substantial entrainment of cold sea water into the interior of the deposit. These observations demonstrate the important role of anhydrite in the growth of massive sulphide deposits, despite its absence in those preserved on land.