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 global sediment mass-age distribution indicates large variations in the rates of carbonate sedimentation through time. The largest mass of carbonate deposited during the entire history of the earth was produced during the Cambrian, possibly following on an episode of phosphogenesis in the Late Precambrian. A second major episode occurred during the Late Devonian, probably reflecting the invasion of land by plants that altered the rock-weathering and soil-forming regimes. Other lesser pulses of carbonate deposition occurred in the Late Permian, Triassic, and Cretaceous. A shift in the locus of carbonate deposition from shallow waters to the deep sea occurred during the Cretaceous.

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: At Aqaba in the Northern Gulf of Aqaba, Red Sea, the stable oxygen isotope record of a Por#es lutea coral colony was studied. From a vertical and a horizontal core two parts from the base of the colony (10 years of coral growth), and two parts from the living surface (12 years of coral growth) were analysed with a monthly sampling resolution. The results show that seasonality of sea surface temperatures (SST) in the northem GulfofAqaba has increased since the early 19th century, mainly due to higher summer temper- atures. 8~80-ratios indicate that SST increased by at least 1.3~ Horizontal and vertical cores display dif- ferent variations in the modern parts, probably caused by the very shallow water depth of the youngest part of the vertical core.

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: Repeatable species dominance patterns across nutrient ratio gradients are found both in culture competition experiments and in natural waters. However, the mechanisms behind these similar patterns need not to be identical. In chemostat experiments, such patterns are caused by the two-way competitive interactions between contemporaneously occurring organisms. Nutrient competition may be important in situ too, but there is an additional effect of early successional species influencing the nutrient environment for later successional species, without being influenced themselves by their successors.

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: 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: Cyanobacteria are important constituents of most aquatic ecosystems on earth, and planktonic forms frequently show mass development in lakes and estuaries. Despite extensive scientific efforts directed towards research on cyanobacteria, a comprehensive theory explaining their success is lacking. Because cyanobacteria comprise a diverse group of organisms, it is concluded that any analysis of the conditions leading to cyanobacterial dominance in pelagic ecosystems should consider at least three different subgroups, classified based on presence or absence of (i) buoyancy regulation and (ii) the ability to fix molecular nitrogen. In this review, nine single-factor hypotheses regarding regulation of cyanobacterial development are examined against the background of physiological and ecological characteristics of these organisms. Special emphasis has been put on understanding differences in nitrogen metabolism between cyanobacteria and algae which directly relate to a hypothesis conceming inorganic nitrogen forms and cyanobacterial success previously presented by our research group. The review lends support to the theory that cyanobacteria have a low competitive ability for nitrate compared with algae and a high ability to compete for ammonium, particularly under nitrogen-limiting conditions. Finally it is concluded that to understand cyanobacterial devlopment and how high Standing Stocks in pelagic ecosystems are maintained requires parts o f all the single-factor hypotheses initially presented.

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: The density of seawater is a complex function of temperature, salinity, and pressure. Because of the non-linearity of the equation of state of seawater, the densities of sea waters having the same temperature and the same salinity differences (with respect to the mean salinity of the ocean) will vary with the mean salinity of the ocean. Although this strange property of seawater is evident in a plot of the equation of state, it has never been considered in trying to reconstruct ancient ocean circulation. These differences in the density field may have caused the ocean to respond differently to atmospheric forcing in the past. The different response may hold the key to understanding "ocean anoxic events" and episodes of large-scale burial of organic carbon and production of petroleum source rocks.

Description: The Late Cretaceous was much warmer than today. There was no significant ice at high latitudes, meridional thermal gradients were low, and continental interiors remained warm during winter. Late Cretaceous atmospheric C02 concentrations were about four times greater than today and an enhanced "greenhouse" effect contributed to the overall warmth of the Late Cretaceous. However , increases in atmospheric C02 tend to increase temperatures at all latitudes and do not explain the very low thermal gradients recognized in the geologic record. Increased poleward ocean heat transport has been cited as a mechanism for maintaining low meridional thermal gradients during the Cretaceous. However , ocean heat transport values larger than the present day are difficult to reconcile. In addition, low meridional thermal gradients suggest sluggish atmospheric circulation, implying that the advection of heat from the warm oceans into the continental interiors was limited. In general, paleoclimate simulations using Atmospheric General Circulations Models (AGCMs) have not been successful in simulating the low meridional thermal gradients and warm winter continental interiors of the Cretaceous, forcing the concept of "equability" to be questioned. Until recently, the physical effects of vegetation on pre-Quaternary climates have largely been ignored. Terrestrial ecosystems influence global climate by affecting the exchange of energy, water, and momentum between the land surface and the atmosphere. In a new approach to pre-Quaternary paleoclimate modeling, Campanian (80 Ma) climate and vegetation have been simulated using a global climate model (GENESIS Version 2.0), coupled to a predictive vegetation model (EVE), resulting in a realistic simulation of Late Cretaceous climate. The predicted distribution of Late Cretaceous vegetation played an important role in the maintenance of low meridional thermal gradients, polar warmth, and equable continental interiors. High latitude forests reduced albedo, especially during snowcovered months, and increased net surface radiation and latent heat flux.

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: 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: In this study, the impact of oceanic data assimilation on ENSO simulations and predictions is investigated. The authors’ main objective is to compare the impact of the assimilation of sea level observations and three-dimensional temperature measurements relative to each other. Three experiments were performed. In a control run the ocean model was forced with observed winds only, and in two assimilation runs three-dimensional temperatures and sea levels were assimilated one by one. The root-mean-square differences between the model solution and observations were computed and heat content anomalies of the upper 275 m compared to each other. Three ensembles of ENSO forecasts were performed additionally to investigate the impact of data assimilation on ENSO predictions. In a control ensemble a hybrid coupled ocean–atmosphere model was initialized with observed winds only, while either three-dimensional temperatures or sea level data were assimilated during the initialization phase in two additional forecast ensembles. The predicted sea surface temperature anomalies were averaged over the eastern equatorial Pacific and compared to observations. Two different objective skill measures were computed to evaluate the impact of data assimilation on ENSO forecasts. The authors’ experiments indicate that sea level observations contain useful information and that this information can be inserted successfully into an oceanic general circulation model. It is inferred from the forecast ensembles that the benefit of sea level and temperature assimilation is comparable. However, the positive impact of sea level assimilation could be shown more clearly when the forecasted temperature differences rather than the temperature anomalies themselves were compared with observations.

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: This paper presents trace element and Sm-Nd isotope data on the Mönchalpgneiss in order to compare the geochemistry of the two polymetamorphic igneous suites that comprise over 30% of the Austroalpine Silvretta nappe. The first are the so-called "Younger Orthogneisses" of the "Fliielagranitic Association" and the second are the "Older Orthogneisses" including the Mönchalpgneisswhich are associated with metagabbros, metadiorites, metatonalites and metagranitoids. U-Pb zircon results from the Mönchalpgneiss are indicative of anatectic processes in late Cambrian to Ordovician times. A volcanic-arc (VA) tectonic environment during intrusion explains the direct association of gabbroic and metagranitoid rocks in the Engadine area. This model is in line with the distribution of major, trace and rare earth elements in these anatectic rocks. However, the significance of the geochemical results remains ambiguous, since the average continental crust and paragneisses of the Silvretta also show VA-type signatures in the respective diagrams. Nd model ages on four Mönchalpgneiss whole-rock samples from the type locality are closely grouped around 1.70 Ga, which is a commonly obtained value for European continental crust. This age is interpreted to be the result of a homogeneous mixture of different crustal components

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: 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: In this paper the performance of the global coupled general circulation model (CGCM) ECHO-2, which was integrated for 10 years without the application of flux correction, is described. Although the integration is rather short, strong and weak points of this CGCM can be clearly identified, especially in view of the model's performance of the annual cycle in the tropical Pacific. The latter is simulated with more success relative to the earlier version, ECHO-I. A better representation of the low-level stratus clouds in the atmosphere model associated with a reduction in the shortwave radiative flux at the air-sea interface improved the coupled model's performance in the southeastern tropical oceans, with a strongly reduced warm bias in these regions. Modifications in the atmospheric convection scheme also eliminated the AGCM's tendency to simulate a double ITCZ, and this behavior is maintained in the CGCM simulation. Finally, a new numerical scheme for active tracer advection in the ocean model strongly reduced the numerical mixing, which seems to enhance considerably the level of interannual variability in the equatorial Pacific. One weak point is an overall cold bias in the Tropics and midlatitudes, which typically amounts to 1°C in open ocean regions. Another weak point is the still too strong equatorial cold tongue, which penetrates too far into the western equatorial Pacific. Although this model deficiency is not as pronounced as in ECHO-1, the too strong cold tongue reduces the level of interannual rainfall variability in the western and central equatorial Pacific. Finally, the interannual fluctuations in equatorial Pacific sea surface temperatures (SSTs) are too equatorially trapped, a problem that is also found in ocean-only simulations. Overall, however, the authors believe that the ECHO-2 CGCM has been considerably improved relative to ECHO-1.

Description: Abstract The famous association of metagabbros, eclogites, glaucophanites, jadeite rocks and ultramafics from the island of Syros forms a distinct lithostratigraphic or tectonic unit. It is interpreted as a high-pressure metamorphic ophiolite suite. This paper provides geochemical and Sr-isotope constraints on the geotectonic setting in which the magmatic protoliths of the Syros metabasites were formed. A compositional gap exists between the metagabbros with Mg-numbers [Mg# = Mg/(0.85Fetot + Mg) atomic ratio] of 0.75-0.88 on the one hand and eclogites and garnet-glaucophanites on the other hand with Mg# of 0.35-0.62, and maximum total iron contents of up to 18 wt.% FeOtot. Metagabbros from various localities and glaucophanites collected around Manna form geochemically coherent groups with smooth correlations between compatible as well as immobile incompatible elements and Mg#. By contrast, the behaviour of immobile incompatible elements, and to some extent also of compatible elements, is highly unsystematic in the eclogites and garnet-glaucophanites. Also, the more mobile elements display a wide scatter in all rocks. This, in conjunction with the unsystematic variation of Sr-isotopes, is thought to be due to secondary alteration. From the strong correlation of Ni and Cr with Mg# and the flat REE patterns lacking Eu-anomalies, a cumulus nature is inferred for olivine, clinopyroxene, and spinel, associated with intercumulus formation of plagioclase in the magmatic protoliths of the metagabbros. There is no direct genetic link between these rocks and the precursors of the Manna-type glaucophanites with REE characteristics typical of N- to T-type MORB. The extremely high geochemical diversity of the eclogites and garnet-glaucophanites from Syros favours individual evolution of their protoliths in small magma bodies as suggested for superferric eclogites from the Western and Ligurian Alps, as well as the ferrogabbros from the ophiolites of the Northern Apennines. From the geochemistry of the Syros metabasites along with the initial 87Sr/86Sr ratios of the metagabbros between 0.7031 and 0.7033, as well as an eNd value of 7.7 from a garnet glaucophanite, the magmatic protoliths are inferred to have formed in a back-arc setting. By analogy to the association of gabbros and ferrogabbros adjacent to the Atlantis II fracture zone of the SW Indian Ridge, we further suggest an origin at a spreading ridge in proximity to a transform fault.

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: Evolution of the Caribbean Plate can be modeled by motions about six successive rotation poles. Opening of Cayman Trough has occurred since 49.5 Ma through westward motion of the Caribbean Plate, eastern Greater Antilles and Chortis Block. Before 49.5 Ma, the eastern Greater-Antilles were west of Cuba, and the southeastern margins of Yucatan and the Nicaragua Rise (Chortis) were aligned. From 67.5 to 49.5 Ma the Caribbean Plate rotated clockwise, opening the Yucatan Basin. From 100 Ma to 67.5 Ma, the Caribbean Plate, with Cuba attached, moved along the southeastern margin of Yucatan-Chortis. At 130 Ma it was attached to northwestern South America.

Description: Microspectralphotometric examinations of various recent pollen point to a different sporopollenin chemistry of the different taxa which is distinguishable by the UV-VIS absorptionspectra. The measurement areas have a diameter of 6 micron, by which several measurements for the spectra control in one pollen grain are possible. Labaratory experiments with chemical treatments as used by palynological preparation, show a clear change in the sporopollenin chemistry whichcould be identified by the absorption spectra. Therefore a new preparation technique for fossil and recent palynomorphs has to be developed to avoid chemical changes. First investigations of recent and fossil Pinus pollen have been clone on material from various facies types and a sediment core. The variations in the absorption spectra of the pollen indicate the different chemical influence of different facies types which could be used for facies analyses. Furthermore, significant changes in the absorption spectra are useful to identify reworked pollen or pollen with a different burial history. Zusammenfassung: Vergleichende mikrospektralphotometrische Untersuchungen der UV-VIS Absorptionsspektren verschiedener rezenter Pollen-Taxa zeigen, daß die Zusammensetzung bzw. Struktur des Sporopollenins bei verschiedenen Taxa variiert und daß diese anhand der Absorptionsspektren klar unterschieden werden kann. Die gemessene Fläche auf den jeweiligen Pollen hat einen Durchmesser von 6 j.lm, so daß mehrere Kontrollmessungen an einem Pollen möglich sind. Laborversuche an rezenten Pollen zeigen, daß chemische Einflüsse (z. B. HN03, Acetolyse) deutlich meßbare Veränderungen in der Struktur des Sporopollenins verursachen. Um Veränderungen des Sporepollenins durch die chemische Probenaufbereitung auszuschließen, wurde eine neue Präparationsmethode für fossile und rezente Palynomorphen entwickelt.

Description: ana marit., 27 (3/6): 145-150, 2 figs.; Frankfurt a. M.] According to the (neo-)Darwinian theory of evolution the appearance of "design" and optimization of biological properties are the results of natural selection and not of teleology. An opti· mization by selection is only possible for entities which are able to reproduce ("units of selection"). The "unit-of-selection"-problem will be discussed in relation to the atomism-holism-continuum of ecological and evolutionary concepts which range from Dawkins' "selfish gene" to LoveJock's "Gaia". Special emphasis will be given to the discussion of the group-selection problem which seems to be the critical breakpoint along this continuum. It will be shown that optimization through natural selection is only possible for units which can multiply and preserve their identity to a sufficient degree through multiplications. Such units are called replicators. Associations, communities, ecosystems and the biosphere do not qualify as replicators.

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: 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: In Sedimenten aus flachen Bohrungen in der Nordsee wurden gasförmige Kohlenwasserstoffe gefunden. Kohlenstoff- (Methan bis Propan) und Wasserstoff-Isotopanwerte lassen auf eine Mischung von Gasen bakterieller und thermischer Entstehung schließen. Die thermischen Gase sind in größerer Tiefe aus überwiegend mariner organischer Substanz entstanden und in die oberflächennahen Sedimente migriert. [Methane through pentane have been found in sediments from shallow North Sea boreholes. Carbon (methane through propane) and hydrogen (methane) isotope ratlos indicate both bacterial and thermal origins. The gases of thermal origin are interpreted as having been generated at considerable depth from predominantly marine source rocks and then to have migrated into the overlying sediments.]

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: The chemical composition of sediments from the Reykjanes Ridge is controlled by allochthonous and autochthonous processes. The surface sediments are characterized by relatively high contents of calciumoxide and strontium. Calcium and strontium dominate in the carbonate phase due to the high content of biogenic carbonate. A high percentage of iron and manganese are bound to oxides and hydroxides. An indication of hydrothermal activity was not observed. A considerable portion of adsorbed barium is transported in clay minerals. The higher amounts of aluminium in glacial sequences indicate an enhanced input of terrigenous material; the increase of stable bonding elemen~s points toward the large influence of detrital minerals. The distinct differences in the bonding characteristics of elements in these marine sediments in comparison to fluvial and coastal - deposits could be due to the different environmental conditions.

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: The seasonal cycle over the tropical Pacific simulated by 11 coupled ocean–atmosphere general circulation models (GCMs) is examined. Each model consists of a high-resolution ocean GCM of either the tropical Pacific or near-global means coupled to a moderate- or high-resolution atmospheric GCM, without the use of flux correction. The seasonal behavior of sea surface temperature (SST) and eastern Pacific rainfall is presented for each model. The results show that current state-of-the-art coupled GCMs share important successes and troublesome systematic errors. All 11 models are able to simulate the mean zonal gradient in SST at the equator over the central Pacific. The simulated equatorial cold tongue generally tends to be too strong, too narrow, and extend too far west. SSTs are generally too warm in a broad region west of Peru and in a band near 10°S. This is accompanied in some models by a double intertropical convergence zone (ITCZ) straddling the equator over the eastern Pacific, and in others by an ITCZ that migrates across the equator with the seasons; neither behavior is realistic. There is considerable spread in the simulated seasonal cycles of equatorial SST in the eastern Pacific. Some simulations do capture the annual harmonic quite realistically, although the seasonal cold tongue tends to appear prematurely. Others overestimate the amplitude of the semiannual harmonic. Nonetheless, the results constitute a marked improvement over the simulations of only a few years ago when serious climate drift was still widespread and simulated zonal gradients of SST along the equator were often very weak.

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).