ALBERT

Description: Latitudinal declines of species richness from the tropics to the poles represent a general spatial pattern of diversity on land. For the marine realm, the generality of this pattern has frequently been questioned. Here, I use a database with nearly 600 published gradients (198 of which were marine) to assess whether there is a marine latitudinal diversity gradient of similar average strength and slope as that for terrestrial organisms. Using meta-analysis techniques, I also tested which characteristics of organisms or habitats affected gradient strength and slope. The overall strength and slope of the gradient for marine organisms was significantly negative and of similar magnitude compared to gradients for terrestrial organisms. Marine gradients were on average stronger as well as steeper than freshwater gradients. Latitudinal gradients were clearly a regional phenomenon, with stronger gradients and steeper slopes for diversity assessed on regional than on local scales. The gradient parameters differed also between oceans and between different habitats, with steeper gradients related to the pelagial rather than the benthos. There were on the other hand no significant differences between hemispheres and between different gradient ranges, although such differences have often been presumed. The most important organismal characteristic related to gradient structure was body mass, with significant gradients related to large organisms. A significant increase in gradient strength with increasing trophic level was observed. The meta-analysis also revealed strongest gradients for nekton and mobile epifauna, whereas the gradients were weak for sessile epifauna and for infauna. In conclusion, marine biota reveal a similar overall decline in diversity with latitude to that observed in terrestrial realms, but the strength and slope of the gradient are clearly subject to regional, habitat and organismal features.

Description: Four in situ experiments were conducted to examine the potential top-down and bottom-up control of epibenthic ciliate communities. The experiments were run in the littoral of Lake Erken and at a brackish water site on the island of Väddö on the Baltic coast of Sweden, during the spring of 2000. The experimental manipulations were the presence/absence of the natural macrozoobenthos grazer community, cross-classified with the presence/absence of additional nutrients. Epibenthic ciliates responded to both manipulation of grazers and resources, but the response was group specific. Total ciliate abundance decreased when macrozoobenthos (largely chironomids, gastropods, trichopteran larvae, isopods and amphipods) were removed, thus excluding a direct predation effect of the macrozoobenthos community on ciliates. Total ciliate biomass, but not abundance, tended to increase in the presence of additional nutrients; an effect weakly dependent on season and site. The disparity between effects of nutrients on biomass and abundance was due to effects on heterotrichs, a group of large but relatively rare algivorous ciliates. The manipulations altered the ciliate community composition, and between lakes there were differences in species richness and diversity and experiments. However, neither the removal of macrozoobenthos nor the addition of nutrients changed species richness or diversity. This runs counter to work with other taxonomic groups, which shows maximal diversity at an intermediate level of resources or predation. This can only be partially explained by the lack of direct predation effects and the open nature of the experimental system.

Description: The role of transparent exopolymer particles (TEP) and dissolved organic carbon (DOC) for organic carbon partitioning under different CO2 conditions was examined during a mesocosm experiment with the coccolithophorid Emiliania huxleyi. We designed 9 outdoor enclosures (similar to11 m(3)) to simulate CO2 concentrations of estimated 'Year 2100' (similar to710 ppm CO2), 'present' (similar to410 ppm CO2) and 'glacial' (similar to190 ppm CO2) environments, and fertilized these with nitrate and phosphate to favor bloom development. Our results showed fundamentally different TEP and DOC dynamics during the bloom. In all mesocosms, TEP concentration increased after nutrient exhaustion and accumulated steadily until the end of the study. TEP concentration was closely related to the abundance of E. huxleyi and accounted for an increase in POC concentration of 35 2 % after the onset of nutrient limitation. The production of TEP normalized to the cell Abundance of E. huxleyi was highest in the Year 2100 treatment. In contrast, DOC concentration exhibited considerable short-term fluctuations throughout the study. In all mesocosms, DOC was neither related to the abundance of E. huxleyi nor to TEP concentration. A statistically significant effect of the CO2 treatment on DOC concentration was not determined. However, during the course of the bloom, DOC concentration increased in 2 of the 3 Year 2100 mesocosms and in 1 of the present mesocosms, but in none of the glacial mesocosms. It is suggested that the observed differences between TEP and DOC were determined by their different bioavailability and that a rapid response of the microbial food web may have obscured CO2 effects on DOC production by autotrophic cells.

Description: Chromophoric dissolved organic matter (CDOM) represents the optically active fraction of the bulk dissolved organic matter (DOM) pool. Recent evidence pointed towards a microbial source of CDOM in the aquatic environment and led to the proposal that phytoplankton is not a direct source of CDOM, but that heterotrophic bacteria, through reprocessing of DOM of algal origin, are an important source of CDOM. In a recent experiment designed at looking at the effects of elevated pCO2 on blooms of the coccolithophorid alga Emiliania huxleyi, we found that despite the 3 different pCO2 levels tested (190, 414 and 714 ppm), no differences were observed in accumulation of CDOM over the 20 d of incubation. Unlike previous mesocosm experiments where relationships between CDOM accumulation and bacterial abundance have been observed, none was observed here. These results provide some new insights into the apparent lack of effect of pCO2 on CDOM accumulation in surface waters, and question the previously proposed mechanisms and rates of CDOM production in natural phytoplankton blooms.

Description: This contribution aims to report the reflections we had with the scientific community during two international workshops on reference materials for stable isotopes in Davos (2002) and Nice (2003). After evaluating the isotopic homogeneity of some existing reference materials, based on either certificates, literature data or specific inter-laboratory rounds, we confirm these as primary reference materials or propose new ones relative to which stable isotope compositions should be reported. We propose DSM-3 for Mg, NIST SRM 915a for Ca, L-SVEC for Li and NBS28 for Si. Cadmium does not yet have a well identified delta zero material, although three commercial mono-elemental Cd solutions have yielded the same isotopic composition relative to one another. In order to scale the linearity of any mass spectrometer, some secondary reference materials are also proposed: Cambridge-1 solution for Mg, the “Münster-Cd” and JEPPIM Cd solutions for Cd and the “Big Batch” silicate for Si. The team from Nancy propose to prepare a mixed spike solution for Li isotopes. Well-characterised natural samples such as ocean or continental waters, diatoms, sponges, rocks and minerals are needed to validate the entire analytical procedure, particularly to take into account the effect of sample mineralisation and of chemical manipulations for elemental separation prior to analysis.

Description: Ecological stoichiometry describes the biochemical constraints of trophic interactions emerging from the different nutrient content and nutrient demand of producers and consumers, respectively. Most research on this topic originates from well-mixed pelagic food webs, whereas the idea has received far less attention in spatially structured habitats. Here, we test how light as well as grazing and nutrient regeneration by consumers affects growth and biomass of benthic primary producers. In the first laboratory experiment, we manipulated grazer presence (two different snail species plus ungrazed control), in the second experiment we factorially combined manipulation of grazer presence and light intensity. We monitored snail and periphyton biomass as well as dissolved and particulate nutrients (nitrogen and phosphorus) over time. Grazers significantly reduced algal biomass in both experiments. Grazers affected periphyton nutrient content depending on the prevailing nutrient limitation and their own body stoichiometry. In the nitrogen (N-) limited first experiment, grazers increased N both in the periphyton and in the water column. The effect was stronger for grazers with lower N-content. In the phosphorus (P-) limited second experiment, grazers increased the P-content of the periphyton, but the grazer with lower N-content had additionally positive effects on algal N. Light reduction did not affect periphyton biomass, but increased chlorophyll-, N- and P-content of the periphyton. These experiments revealed that the indirect effects of grazers on periphyton were bound by stoichiometric constraints of nutrient incorporation and excretion.

Description: Ecosystem resistance to a single stressor relies on tolerant species that can compensate for sensitive competitors and maintain ecosystem processes, such as primary production. We hypothesize that resistance to additional stressors depends increasingly on species tolerances being positively correlated (i.e. positive species co-tolerance). Initial exposure to a stressor combined with positive species co-tolerance should reduce the impacts of other stressors, which we term stress-induced community tolerance. In contrast, negative species co-tolerance is expected to result in additional stressors having pronounced additive or synergistic impacts on biologically impoverished functional groups, which we term stress-induced community sensitivity. Therefore, the sign and strength of the correlation between species sensitivities to multiple stressors must be considered when predicting the impacts of global change on ecosystem functioning as mediated by changes in biodiversity.

Description: Recent experiments, mainly in terrestrial environments, have provided evidence of the functional importance of biodiversity to ecosystem processes and properties. Compared to terrestrial systems, aquatic ecosystems are characterised by greater propagule and material exchange, often steeper physical and chemical gradients, more rapid biological processes and, in marine systems, higher metazoan phylogenetic diversity. These characteristics limit the potential to transfer conclusions derived from terrestrial experiments to aquatic ecosystems whilst at the same time provide opportunities for testing the general validity of hypotheses about effects of biodiversity on ecosystem functioning. Here, we focus on a number of unique features of aquatic experimental systems, propose an expansion to the scope of diversity facets to be considered when assessing the functional consequences of changes in biodiversity and outline a hierarchical classification scheme of ecosystem functions and their corresponding response variables. We then briefly highlight some recent controversial and newly emerging issues relating to biodiversity-ecosystem functioning relationships. Based on lessons learnt from previous experimental and theoretical work, we finally present four novel experimental designs to address largely unresolved questions about biodiversity-ecosystem functioning relationships. These include (1) investigating the effects of non-random species loss through the manipulation of the order and magnitude of such loss using dilution experiments; (2) combining factorial manipulation of diversity in interconnected habitat patches to test the additivity of ecosystem functioning between habitats; (3) disentangling the impact of local processes from the effect of ecosystem openness via factorial manipulation of the rate of recruitment and biodiversity within patches and within an available propagule pool; and (4) addressing how non-random species extinction following sequential exposure to different stressors may affect ecosystem functioning. Implementing these kinds of experimental designs in a variety of systems will, we believe, shift the focus of investigations from a species richness-centred approach to a broader consideration of the multifarious aspects of biodiversity that may well be critical to understanding effects of biodiversity changes on overall ecosystem functioning and to identifying some of the potential underlying mechanisms involved.

Description: Marine and fisheries scientists are increasingly using metapopulation concepts to better understand and model their focal systems. Consequently, they are considering what defines a metapopulation. One perspective on this question emphasizes the importance of extinction probability in local populations. This view probably stems from the focus on extinction in Levins' original metapopulation model, but places unnecessary emphasis on extinction–recolonization dynamics. Metapopulation models with more complex structure than Levins' patch-occupancy model and its variants allow a broader range of population phenomena to be examined, such as changes in population size, age structure and genetic structure. Analyses along these lines are critical in fisheries science, where presence–absence resolution is far too coarse to understand stock dynamics in a meaningful way. These more detailed investigations can, but need not, aim to assess extinction risk or deal with extinction-prone local populations. Therefore, we emphasize the coupling of spatial scales as the defining feature of metapopulations. It is the degree of demographic connectivity that characterizes metapopulations, with the dynamics of local populations strongly dependent upon local demographic processes, but also influenced by a nontrivial element of external replenishment. Therefore, estimating rates of interpopulation exchange must be a research priority. We contrast metapopulations with other spatially structured populations that differ in the degree of local closure of their component populations. We conclude with consideration of the implications of metapopulation structure for spatially explicit management, particularly the design of marine protected area networks.

Description: We fertilised 12 mesocosms with NE Atlantic phytoplankton with different Si:N ratios (0:1 to 1:1). After 1 wk, we added mesozooplankton, mainly calanoid copepods at natural densities to 10 of the mesocosms; the remaining 2 mesocosms served as controls. A trend of increasing diatom dominance with increasing Si:N ratios and species-specific correlations of diatoms to Si:N ratios were not changed by the addition of mesozooplankton. Large unicellular and chain-forming diatoms, thin-walled dinoflagellates (Gymnodiniales) and ciliates were reduced by copepod grazing while armoured dinoflagellates remained unaffected. Nanoplanktonic flagellates and diatoms profited from the addition of copepods, probably through release from ciliate grazing.

Description: Herbivory can greatly modify benthic community structure by affecting the distribution of algal species. To deter herbivores, algae have developed several mechanisms, including the induction of chemical and morphological defenses, which may be influenced by nutrient availability. We tested 4 red (Chondrophycus flagellifera, Cryptonemia seminervis, Osmundaria obtusiloba, Pterocladiella capillacea), 4 brown (Dictyota menstrualis, Lobophora variegata, Sargassum vulgare, Stypopodium zonale), and 1 green (Codium decorticatum) algae for inducible defenses following exposure to direct consumption by an amphipod community dominated by Elasmopus brasiliensis. In addition, the effects of water-borne cues from nearby grazed conspecifics and non-grazing consumers on the induction of defenses were examined in C. decorticatum under natural and enhanced (200% natural) nutrient levels. Induction of defense was assessed in choice-feeding assays, using live algae or artificial food containing non-polar extracts of amphipod-exposed (treated) and non-exposed (control) algae. Palatability levels, estimated as the relative difference in wet mass due to consumption in feeding assays between grazer-exposed and control plants, declined significantly in 3 species after the acclimatization period. Tissue from the directly consumed red alga P. capillacea (live alga) was significantly less palatable than tissue from the control plants. Likewise, a significant effect was observed in the brown alga L. variegata. Similar, although not statistically significant, trends were observed in 6 other species. For the green alga C. decorticatum, nutrient enrichment did not affect induction of defenses by herbivores, yet unfertilized plants were more palatable than fertilized conspecifics.

Description: Connell¹s intermediate disturbance hypothesis (IDH) postulates that diversity peaks at an intermediate level of disturbance frequency or intensity. To test the validity of this concept for the species-poor marine hard-bottom community of the Western Baltic, we chose an in situ experimental approach. Undisturbed fouling communities of 2 different successional stages, 3 and 12 mo old, were submitted to various levels of emersion intensities, defined as time spans of continuous exposure to the air d-1. Disturbance levels ranged from 0.25 h up to 12 h of daily exposure. The study on 3 mo old communities was repeated in 2 subsequent years, 1999 and 2000. Species richness, evenness and diversity (Shannon index) were recorded to measure the effect of intensity treatments on community structure. The IDH was confirmed in the first year, when diversity was found to peak at intermediate disturbances. However, for communities of both successional stages, diversity-disturbance relationships were U-shaped or not significant in the second year. This ambiguous picture basically confirms the validity of the mechanisms proposed by the IDH, but shows that their forcing can be masked by fluctuations in environmental parameters, such as climatic conditions. An extension of the model is proposed, that considers diversity enhancement under extreme conditions due to a disturbance-induced change in community structure. Furthermore we discuss a conceptual linkage of the IDH to the multiple stable-state hypothesis. Finally, we found community stability not to be positively correlated with community age and complexity.

Description: Some marine mammals may increase their underwater locomotor efficiency by taking down little air for dives and descending passively, although at the point of maximum depth they presumably have to use energy to counteract the downthrust, to stop themselves sinking further. Birds, having considerable quantities of body-associated air, would appear not to have this option. However, measurements of locomotor activity and inspiratory behaviour of free-living, diving penguins has revealed that birds regulated the inspired air volume so that upthrust, primarily derived from depth-related changes in air volume, was minimal and constant at the preferred foraging depth. Although this results in minimized costs of travel, it means total body oxygen stores have to vary with depth, something that helps explain why dive duration is so closely correlated with depth in birds.

Description: We determined key chemical parameters and thiosulfate oxidation in low-temperature hydrothermal fluids from the North Fiji Basin. In addition, the bacterial diversity (with the main emphasis on sulfur-oxidizing bacteria) was investigated. The hydrothermal fluids had low concentrations of sulfide (up to 50.0 µM) and increased counts of both total bacteria and sulfur-oxidizing bacteria compared to ambient seawater. Pure cultures of bacteria were isolated from these fluids on media suited for autotrophic, and potentially heterotrophic, sulfur-oxidizing bacteria. Evidence for the abundance of α- and γ -Proteobacteria was obtained from identification of isolated pure cultures. A large number of 16S rDNA sequences of these groups were retrieved from environmental DNA. Representatives of the Cytophaga-Flavobacterium phylum were found by denaturing gradient gel electrophoresis (DGGE) and 16S rDNA sequence information of DGGE bands, although these bacteria could not be isolated with the media used in this study. Evidence for the presence of chemolithoautotrophic sulfur-oxidizing bacteria was found by analysis of environmental DNA, using 16S rDNA-specific primers of various groups of chemotrophic sulfur bacteria. They were isolated in low numbers compared to chemoheterotrophic and mixotrophic sulfur-oxidizing bacteria. In addition, the formation of tetrathionate as major oxidation product of thiosulfate added to hydrothermal fluid samples and to pure cultures of new isolates indicates the importance of chemoheterotrophic sulfur-oxidizing bacteria within the warm vent waters investigated during this study.

Description: Three simple fisheries indicators are presented: (i) percentage of mature fish in catch, with 100% as target; (ii) percent of specimens with optimum length in catch, with 100% as target; and (iii) percentage of ‘mega-spawners‘ in catch, with 0% as target, and 30–40% as representative of reasonable stock structure if no upper size limit exists. Application of these indicators to stocks of Gadus morhua, Sardinella aurita and Epinephelus aeneus demonstrate their usefulness. It is argued that such simple indicators have the potential to allow more stakeholders such as fishers, fish dealers, supermarket managers, consumers and politicians to participate in fisheries management and eventually hold and reverse the global pattern of convenience overfishing, which is defined here as deliberate overfishing sanctioned by official bodies who find it more convenient to risk eventual collapse of fish stocks than to risk social and political conflicts.

Description: The epibacterial chemical defense of the marine sponge Suberites domuncula was explored by screening sponge extract, sponge primmorph (3-D aggregates containing proliferating cells) extract and sponge-associated as well as primmorph-associated bacteria for antibacterial activity. 16S rDNA sequencing revealed that the antimicrobially active bacteria belonged to the a- and γ-subdivisions of Proteobacteria ( α-Proteobacterium MBIC 3368, Idiomarina sp. and Pseudomonas sp., respectively). Moreover, a recombinant perforin-like protein was cloned from S. domuncula that displayed strong antibacterial activity. Based on these observations, it is proposed that the sponge may be provided with a direct (by producing antibacterial metabolites) as well as an indirect (with the help of associated bacteria) epibacterial defense.

Description: There is mounting molecular evidence that bacteria belonging to the phylum Planctomycetes are abundant in marine sponges including members of the genus Aplysina. In an attempt to culture planctomycete bacteria from Aplysina sponges, 116 bacterial strains were isolated on selective oligotrophic media. Screening of the strain collection by fluorescence in situ hybridization with the planctomycete-specific probe Pla46 yielded 3 positive candidates. Nearly complete sequencing of the respective 16S rRNA genes revealed that the isolates were affiliated with 2 distinct clusters of the genus Pirellula: 1 isolate was obtained from a Mediterranean sponge, 1 from a Caribbean sponge and a third from Caribbean seawater. To our knowledge this is the first report of cultured Planctomycetes from marine sponges. The isolates grew slowly on oligotrophic media and failed to grow on nutrient-rich media. Pirellula sp. Strain 797 was pink-pigmented while the other 2 isolates, 16 and 81, were non-pigmented. Transmission electron microscopy revealed a pear- or droplet-shaped cell morphology that is characteristic of the genus Pirellula. The application of strain-specific oligonucleotide probes to sponge tissue cryosections showed that the isolates contribute only a minor fraction to the total microbial community that is associated with Aplysina spp. sponges

Description: Conceptual models predict counteractive effects of herbivores and nutrient enrichment on plant diversity and reversed effects of grazers under different nutrient regimes. I tested these hypotheses in 11 field experiments with periphyton communities in three different aquatic habitats (a highly eutrophic lake, an meso-eutrophic lake, and an meso-eutrophic part of the Baltic Sea coast) and in different seasons. Grazer access and nutrient supply were manipulated in a factorial design. Species richness and evenness were chosen as response variables. Both manipulated factors had significant and contrasting effects on diversity, with variable effect strength between sites and seasons. From the two aspects of diversity, evenness well reflected the changes in community composition. Fertilization tended to increase the dominance of few species and thus to decrease evenness, whereas grazers counteracted these effects by removing dominant life forms. The response of species richness was not as expected, since grazers decreased richness throughout, whereas nutrients had weaker effects but tended to increase richness. Species richness rather reflected changes in periphyton architecture. Grazers reduced algal richness presumably by co-consumption of rare species in the tightly connected periphyton assemblages, whereas enrichment may increase richness by providing more structure via increased dominance of filamentous species. Although grazer and nutrient effects on richness and evenness were opposing, there was no change in the effect of one factor by manipulation of the other.

Description: In January and February 1998, when an unprecedented fourth repetition of the zonal hydrographic transect at 24.5°N in the Atlantic was undertaken, carbon measurements were obtained for the second time in less than a decade. The field of total carbon along this section is compared to that provided by 1992 cruise which followed a similar path (albeit in a different season). Consistent with the increase in atmospheric carbon levels, an increase in anthropogenic carbon concentrations of Full-size image (〈1 K) was found in the surface layers. Using an inverse analysis to determine estimates of absolute velocity, the flux of inorganic carbon across 24.5° is estimated to be −0.74±0.91 and Full-size image (〈1 K) southward in 1998 and 1992, respectively. Estimates of total inorganic carbon flux depend strongly upon the estimated mass transport, particularly of the Deep Western Boundary Current. The 1998 estimate reduces the large regional divergence in the meridional carbon transport suggested by previous studies and brings into question the idea that the tropical Atlantic constantly outgasses carbon, while the subpolar Atlantic sequesters it. Uncertainty in the carbon transports themselves, dominated by the uncertainty in the total mass transport estimates, are a hindrance to determining the “true” picture. The flux of anthropogenic carbon (C★ANTH) across the two transects is estimated as northward at 0.20±0.08 and Full-size image (〈1 K) for the 1998 and 1992 sections, respectively. The net transport of C★ANTH across 24.5°N is strongly affected by the difference in concentrations between the northward flowing shallow Florida Current and the mass balancing, interior return flow. The net northward transport of C★ANTH is opposite the net flow of total carbon and suggests, as has been found by others, that the pre-industrial southward transport of carbon within the Atlantic was stronger than it is today. Combining these flux results with estimates of atmospheric and riverine inorganic carbon input, it is determined that today's oceanic carbon system differs from the pre-industrial system in that today there is an uptake of anthropogenic carbon to the south that is advected northward and stored within the North Atlantic basin.

Description: Exchanges of water south of Africa between the South Indian Ocean and the South Atlantic Ocean are an important component of the global thermohaline circulation. Evidence exists that the variability in these exchanges, on both meso- and longer time scales, may significantly influence weather and climate patterns in the southern African region and the significance of these regional ocean–atmosphere interactions is discussed. Observations of the inter-ocean exchange are limited and it is necessary to augment these with estimates derived from models. As a first step in this direction, this study uses an eddy-permitting model to investigate the heat and volume transport in the oceanic region south of Africa and its variability on meso, seasonal and inter-annual time scales. On the annual mean, about Full-size image (〈1 K) (standard deviation Full-size image (〈1 K)) of heat flows west into the South Atlantic across 20°E (longitude of Cape Agulhas, the southernmost point of Africa), with just over Full-size image (〈1 K) (standard deviation Full-size image (〈1 K)) flowing north into the South Atlantic across 35°S. The seasonal variations in this transport are about 10% at 35°S in the South Atlantic and around 20% through 20°E; the model value of Full-size image (〈1 K) for summer (standard deviation ranging from Full-size image (〈1 K) in January to Full-size image (〈1 K) in March) appears consistent with respective estimates of 0.51 and Full-size image (〈1 K) derived from two WOCE summer cruises southwest of Cape Town to 45°S in 1990 and 1993. Volume transports of the Agulhas Current section through 35°S in the SW Indian Ocean range from 58 to Full-size image (〈1 K) in summer/autumn to 64–Full-size image (〈1 K) in winter/spring. The model results suggest that the inter-ocean exchange south of Africa is highly variable on seasonal through to interannual scales. If this variability is also the case in the real ocean (and the limited observations suggest that this is so), then there are likely to be significant implications for climate.

Description: A set of histological characteristics to judge ovarian development was established and used to elaborate morphological criteria of 10 maturity stages of Baltic cod Gadus morhua sampled throughout the annual cycle to represent different macroscopic maturity stages. The applied characteristics confirmed most stages of the macroscopic scale, but the separation of late immature and resting mature females remained imprecise. Atretic vitellogenic oocytes or encapsulated residual eggs identified the resting condition morphologically, but not all ovaries with visible signs of previous spawning showed such features. One ovarian stage that was previously classified as ‘ripening’ was changed to ‘spawning’, owing to the prevalence of hydrated eggs and empty follicles. Ovaries with malfunctions were defined by a separate stage. Macroscopic criteria were revised by comparing the gross anatomy of ovaries with their histology. Female length and gonado-somatic index supported stage definitions, but substantial variation in Fulton's condition factor and the hepato-somatic index rendered these of little use for this purpose. The time of sampling influenced staging accuracy. A female spawner probability function based on the proportion of ripening and ripe specimens in early spring seems to be the most appropriate method to estimate spawner biomass and reproductive potential.

Description: Somatic growth and nucleic acid content were studied in North Sea houting Coregonus oxyrhinchus larvae fed exclusively on dry diets at 2 temperatures (8.4 and 17.5°C) during a 32 d rearing experiment. The higher temperature enhanced growth significantly. Mean dry weights at the end of the experiment were 3.6 mg (SD = 1.07, range 1.4 to 5.7 mg) and 31.5 mg (SD = 21.9, range 3.6 to 96.0 mg), and mean standard lengths were 17.7 ± mm (SD = 1.6, range 11.8 to 19.6 mm) and 25.5 mm (SD = 4.2, range 17.0 to 35 mm), respectively. Significant responses to temperature were also found in the nucleic acids. However, these differences were not as remarkable and gave indications that differences in protein growth between treatments was based on protein biosynthesis being driven by the activity of the ribosomes, rather than their number. The use of the degree-day approach to normalize the data clearly showed the temperature-dependence of somatic growth. Only small differences in growth and nucleic acid content were observed in the comparable range of the first 280 degree-days in fed fish. In non-fed fish, the starving potential was very similar (approx. 350 degree-days). Shifts in growth pattern from predominantly hyperplasia to predominantly hypertrophy were detected, with ongoing growth at both temperatures.

Description: Pelagic and demersal juvenile Baltic cod Gadus morhua L. were collected on the slope and the top of Rønne bank in the Baltic Sea during 2 cruises in November and December 1998. The objective of this study was to evaluate distinct changes in otolith increment width observed in demersal juveniles by comparison with laboratory-reared individuals, and to investigate the factors determining variation in these increments. The different increment-width patterns were identified with a method based on the widths of consecutive increments. Otolith increment widths of juvenile cod were found to be highly variable within and between individuals, in both the experimental and the field samples. The first change in increment pattern observed in the field samples was related to settling. The formation periodicity of increments within the different pattern intervals was confirmed with a growth model based on otolith growth rates of juvenile cod reared in the laboratory under different conditions. In this model, otolith growth rate was expressed as a function of rearing temperature and fish dry weight. Otolith growth of the field samples was calculated using ambient temperatures obtained from a 3D-circulation model. The best fit to observed otolith growth rates was obtained under the assumption that fish on the slope performed daily vertical migrations between the warm surface layer and the cold bottom layer. The data suggested that fish stayed in the surface layer during the first increment-pattern interval, performed vertical migrations during the second interval, and stayed in association with the seafloor in the subsequent interval, corresponding to the time after the breakdown of the thermocline.

Description: During peak spawning of sprat Sprattus sprattus in the Baltic Sea in May–June egg specific gravity averaged ±s.d. 1·00858 ± 0·00116 g cm−3 but was significantly higher in the beginning and significantly lower towards the end of the spawning season. A close relationship was found between egg diameter and egg specific gravity (r2 = 0·71). This relationship, however, changed during the spawning season indicating that some other factor was involved causing the decrease in specific gravity during the spawning period. The vertical egg distribution changed during the spawning season: eggs were distributed mainly in the deep layers early in the season, occurred in and above the permanent halocline during peak spawning, and above the halocline towards the end of the spawning season. Consequently, poor oxygen conditions in the deep layers and low temperatures in layers between the halocline and the developing thermocline may affect egg development. Thus, opportunities for egg development vary over the spawning season and among spawning areas, and depending on frequency of saline water inflows into the Baltic Sea and severity of winters, between years

Description: The active channel–levee system of the middle Bengal Fan was studied by a combined analysis of Parasound echosounder and Hydrosweep swathsounder data. The channel is characterized by highly variable sinuosities. Compared to other mud-rich submarine fans, an exceptionally low channel slope is found. The system can be subdivided into inner and outer zones of significantly different depositional architecture. The inner zone consists of the active channel and sharply separated vertical blocks, which are characterized by parallel, distinct reflectors and planforms of bends. These blocks are interpreted as abandoned channel segments (cut-off loops). The outer zones represent undisturbed levees, which are constructed of parallel and wedge-shaped sedimentary units. The wedge-shaped units, varying significantly in thickness and lateral extent, are found at the outer convex arcs of active and abandoned channel loops caused by overspilling of channelized turbidity currents at sharp bends. The parallel units are the deposits of turbidity currents, which spread their sediments over wide areas as their size significantly exceeds the cross-section of the channel. The complex vertical and horizontal distribution of partially small sedimentary units suggests a more complicated deposition in time and space as hitherto reported from other submarine fans. Within the inner zone, more than 20 cut-off loops were identified over a channel length of 90 km. In contrast to most other large mud-rich submarine fans, channel avulsion within the active channel–levee system is a frequent process. In particular, a temporal succession of at least 4 cut-off loops was reconstructed in the southern study area, indicating channel avulsion on average every 750 years. Channel avulsion seems to be a repetitious process caused by erosion through turbidite currents in a highly sinuous channel. Compared to other submarine fans, no morphological parameter shows a remarkable difference except the channel slope, which is significantly smaller than, for example, on Amazon, Congo and Mississippi fans. The interaction between this low channel slope and the flow parameter of the turbidity currents is most likely the reason for the instability of the active channel planform, leading to an exceptionally large number of meander loop breaches and cut-off loops.

Description: Combining in-situ Lagrangian intermediate depth velocity measurements from the KAPEX (Cape of Good Hope Experiments) float program with sea-surface height data, this study reviews the inter-ocean exchange mechanisms around southern Africa. In the southeastern Cape Basin, a highly energetic field of coexisting anticyclonic and cyclonic eddies is documented. Agulhas Rings of typically 200 km diameter are observed to merge, split, deform, and to reconnect to the Agulhas Retroflection. Concomitant, slightly smaller cyclones are observed to drift across the northwestward migration path of the Agulhas Rings. These cyclones, with typical diameters of 120 km, are formed within the Cape Basin along the African shelf, inshore of the Agulhas Current, and in the subantarctic region south of Africa. The data suggest the annual formation of 3–6 long-lived Agulhas Rings that eventually cross 5°E longitude, while approximately twice the number of rings occur in the southeastern Cape Basin. Within this region, cyclones outnumber anticyclones by a factor of 3:2. Both cyclones and anticyclones extend through the upper thermocline into the intermediate depth layer. Mean drifts of anticyclones are 3.8±1.2 cm s−1 to the northwest, while cyclones follow a west–southwestward route at 3.6±0.8 cm s−1. Transport estimates suggest that the intermediate depth layer in the southeastern Cape Basin is primarily supplied from the east (approximately 9 Sv), with minor direct inflow from the Atlantic to the west and south. Cyclone/anticyclone interaction is surmised to result in vigorous stirring and mixing processes in the southeastern Cape Basin, which necessitates a review of the traditional concept of Indo-Atlantic inter-ocean exchange. We propose to limit the concept of “isolated Agulhas Rings embedded in a sluggish Benguela Drift” to the northwestern Cape Basin and beyond, while linking this regime to the Agulhas Retroflection proper through a zone of turbulent stirring and mixing in the southeastern Cape Basin, named for the first time the “Cape Cauldron” hereinafter.

Description: At Hydrate Ridge (HR), Cascadia convergent margin, surface sediments contain massive gas hydrates formed from methane that ascends together with fluids along faults from deeper reservoirs. Anaerobic oxidation of methane (AOM), mediated by a microbial consortium of archaea and sulfate-reducing bacteria, generates high concentrations of hydrogen sulfide in the surface sediments. The production of sulfide supports chemosynthetic communities that gain energy from sulfide oxidation. Depending on fluid flow, the surface communities are dominated either by the filamentous sulfur bacteria Beggiatoa (high advective flow), the clam Calyptogena (low advective flow), or the bivalve Acharax (diffusive flow). We analyzed surface sediments (0 to 10 cm) populated by chemosynthetic communities for AOM, sulfate reduction (SR) and the distribution of the microbial consortium mediating AOM. Highest AOM rates were found at the Beggiatoa field with an average rate of 99 mmol m-2 d-1 integrated over 0 to 10 cm. These rates are among the highest AOM rates ever observed in methane-bearing marine sediments. At the Calyptogena field, AOM rates were lower (56 mmol m-2 d-1). At the Acharax field, methane oxidation was extremely low (2.1 mmol m-2 d-1) and was probably due to aerobic methane oxidation. SR was fueled largely by methane at flow-impacted sites, but exceeded AOM in some cases, most likely due to sediment heterogeneity. At the Acharax field, SR was decoupled from methane oxidation and showed low activity. Aggregates of the AOM consortium were abundant at the fluid-impacted sites (between 5.1 × 1012 and 7.9 × 1012 aggregates m-2) but showed low numbers at the Acharax field (0.4 × 1012 aggregates m-2). A transport-reaction model was applied to estimate AOM at Beggiatoa fields. The model agreed with the measured depth integrated AOM rates and the vertical distribution. AOM represents an important methane sink in the surface sediments of HR, consuming between 50 and 100% of the methane transported by advection.

Description: A compilation of δ44/40Ca (δ44/40Ca) data sets of different calcium reference materials is presented, based on measurements in three different laboratories (Institute of Geological Sciences, Bern; Centre de Géochimie de la Surface, Strasbourg; GEOMAR, Kiel) to support the establishment of a calcium isotope reference standard. Samples include a series of international and internal Ca reference materials, including NIST SRM 915a, seawater, two calcium carbonates and a CaF2 reference sample. The deviations in δ44/40Ca for selected pairs of reference samples have been defined and are consistent within statistical uncertainties in all three laboratories. Emphasis has been placed on characterising both NIST SRM 915a as an internationally available high purity Ca reference sample and seawater as representative of an important and widely available geological reservoir. The difference between δ44/40Ca of NIST SRM 915a and seawater is defined as -1.88 O.O4%o (δ44/42CaNISTSRM915a/Sw= -0.94 0.07%o). The conversion of values referenced to NIST SRM 915a to seawater can be described by the simplified equation δ44/40CaSa/Sw=δ44/40CaSa/NIST SRM 915a - 1.88 (δ44/42CaSa/Sw=δ44/42CaSa/NIST SRM 915a - 0.94). We propose the use of NIST SRM 915a as general Ca isotope reference standard, with seawater being defined as the major reservoir with respect to oceanographic studies.

Description: The combined analysis of hydrographic, kinematic, and dynamic data collected during the 1997–1999 KAPEX (CAPe of Good Hope EXperiments) reveals a quasi-stationary meandering pattern of the Agulhas Retroflection Current east and upstream of the Southwest-Indian Ridge. The current meanders between 38°S and 40°S in a spatially and temporally continuous fashion and has a core width of approximately Full-size image (〈1 K) with an associated transport of Full-size image (〈1 K) in the upper Full-size image (〈1 K). Peak surface velocities decrease from Full-size image (〈1 K) near the Agulhas Retroflection to Full-size image (〈1 K) around 32°E. Meander troughs (northward extremes) are found predominantly near 26.8°E, 32.6°E and 38.9°E, while crests (southward extremes) are located with high probability near 29.7°E, 35.5°E and 42.9°E, resulting in a typical wavelength of Full-size image (〈1 K). Cold eddies are shed along the northern boundary of the current from meander troughs into the recirculation regime between the Agulhas Current proper and the Agulhas Return Current. Strongest cyclonic eddies are preferably shed in austral autumn. The cyclonic eddies so formed propagate westward at an average phase-speed of Full-size image (〈1 K), with, however, a variability of at least the same magnitude. Subsequently, the cyclones are absorbed by the next meander trough located upstream and to the west of the shedding trough.

Description: Although the 'intermediate disturbance hypothesis' (IDH) was originally postulated for marine hard-bottom communities, it has rarely been tested for this community type. We experimentally examined the effects of ambient plus artificially enhanced UV-B radiation (eUV-BR) along an intensity gradient on the abundance of individual species, species composition, and diversity of a macrobenthic community in the western Baltic, Germany. Plots were either exposed each day for 0.5, 1, 2, 4, and 8 h to the maximum expected future UV-BR or were left untreated (= ambient irradiance). Species recruitment and succession on artificial substrata were followed for 5 mo. Transient unimodal patterns for species richness and diversity H' were produced along the UV-BR gradient early in succession. The density of the competitively superior mussel Mytilus edulis and the abundance of the foliose green alga Ulvopsis grevillei were positively related to UV-BR exposure; suggesting conditions different from those predicted by the IDH-generated unimodal patterns. Protective shading of substratum by U. grevillei and the resultant mitigation of the UV-BR impact for succeeding colonizers may be important in this context. Species composition differed persistently among eUV-BR treatments. In contrast to mussels and U. grevillei, the abundance of red algae was adversely affected by eUV-BR along the disturbance gradient. Our results suggest that expected future UV-BR levels will have limited influence on the diversity of shallow-water macrobenthic fouling assemblages in the western Baltic, whereas their species composition may be affected over longer periods.

Description: Dilution experiments were used to examine growth and grazing mortality rates among Prochlorococcus and Synechococcus populations in the Sargasso Sea and California Current. In these experiments, deviation from linearity in the relationship between dilution and net growth rate was significant in a large number of cases. An alternative, more conservative approach for estimating growth and grazing mortality rates (independent of the shape of this relationship) was therefore employed. Growth rates estimated by this approach ranged from 0.32 to 0.76 and 0.37 to 0.67 d-1 for Prochlorococcus and Synechococcus, respectively. Grazing mortality rates ranged from 0.25 to 0.85 and 0.13 to 0.51 d-1, respectively. Cell-cycle-based growth rate estimates were consistent with these dilution-based rates. Nutrient amendment had little affect on picocyanobacterial growth rates, but did stimulate grazing mortality (and in some cases changed the apparent functional response of the grazer community) in a number of experiments. We hypothesize that improved food quality in nutrient-replete picoplankton cells may be responsible for these changes. Diel patterns of picocyanobacterial abundance in the Sargasso Sea experiments suggest that grazing activity varied strongly over the diel cycle, with low grazing activity during the first half of the light period. Growth rate and abundance were not positively correlated among or within picocyanobacterial groups, as might be expected if physiologically mediated controls were the dominant forces regulating these populations.

Description: Age, growth and maturity parameters were examined for the southern Australian ommastrephid squid Nototodarus gouldi. Squid were obtained from the fishing ports of: Ulladulla, New South Wales; Port Lincoln, South Australia; Lakes Entrance, Victoria; and Hobart, Tasmania. Squid were collected during 2 seasonal periods: summer/autumn-caught (warm-season squid) and spring-caught (cool-season squid) over 2 consecutive years (2000, 2001). N. gouldi is a sexually dimorphic species, with females generally reaching larger sizes than males. Initial genetic analysis has found only a single species in Australian waters. Statolith ageing revealed that squid completed their life cycle in 〈1 yr, and appear to hatch throughout the year. Trends in size, growth and maturity varied considerably between sites, seasons and years. Squids hatched in summer/autumn grew consistently faster than squid that hatched in winter/spring, presumably due to the influence of temperature on growth. Squid in 1999/2000 also grew faster than squid in 2000/2001. Growth of female squid in winter correlated with sea surface colour (SSC) during peak hatch periods, but the SSC relationship did not exist for males. Ulladulla squid were generally smaller, younger, had smaller gonads than most other squid and were possibly a smaller morph of the species. Tasmania and Lakes Entrance tended to have larger older individuals with larger gonads, while Port Lincoln was variable and intermediate. However, during spring 2001 both Tasmania and Port Lincoln had individuals that were much smaller than those of the other seasons for these sites, and were more like those from Ulladulla. Trends in age of mature individuals showed considerable variability (over 100 d from youngest to oldest) and there appeared to be a cline across all sites and seasons. Arrow squid appear to reveal marked plasticity in age, growth and maturity parameters, but currently the extent to which the environment or genetics control plasticity is unclear.

Description: Rings shed at the Agulhas retroflection are an integral part of interoceanic exchange south of Africa. There is clear evidence of westward ring translation from the northern Cape Basin across the South Atlantic Ocean. Early ring development and translation from the southern to the northern Cape Basin, however, are obscured by an intensely variable kinematic field close to the spawning site. In this study unique in situ observations, obtained in March to September 1997, are analyzed to improve the understanding of the early development of a juvenile Agulhas Ring. In March the ring was surveyed near 37°S, 16°E, approximately 4 months after its generation. Its strength and size were in the upper range typical for Agulhas Rings, and its trapping depth extended down to at least 1600 dbar according to geostrophic velocities and RAFOS trajectories in the ring. Between March and September the ring propagated in a general northwestward direction; however, RAFOS trajectories and MODAS sea-surface steric height fields revealed a large variability of the translation speed (Full-size image (〈1 K) to more than Full-size image (〈1 K)) and direction. In September 1997, the mature ring was examined near 31°S, 9°E. By this time, its available heat and salt anomaly were reduced by about 30% and its available potential energy was reduced by about 70%. This indicates that a significant loss of the ring characteristics occurred on the way from the southern to the northern Cape Basin. One-third of this loss is due to changes at intermediate depth (between 800 and Full-size image (〈1 K)).

Description: The phylogenetic relationship of sulphur-oxidising endosymbiotic bacteria from bivalves of the families Vesicomyidae (Calyptogena sp. C1, Calyptogena sp. C3), Solemyidae (Acharax sp.) and Thyasiridae (Conchocele sp.) from cold-seep habitats were determined by 16S rDNA nucleotide sequence analyses. The endosymbiotic bacteria form distinct groups within the gamma-Proteobacteria and are well separated from each other and from free-living sulphur-oxidising bacteria of the genera Beggiatoa, Halothiobacillus and Thiomicrospira. The endosymbiotic bacteria of Acharax sp. from cold seeps off Oregon, Indonesia and Pakistan have sequences highly similar to each other but quite distinct from other thiotrophic endosymbionts. This includes endosymbionts from Solemya spp., to which they are distantly related. Symbiotic bacteria of Conchocele sp. from a cold seep in the Sea of Okhotsk are similar to those of Bathymodiolus thermophilus and related species, as shown by their overall sequence similarity and by signature sequences. The endosymbiotic bacteria of Calyptogena spp. from cold seeps off Oregon and Pakistan are closely related to those of other vesicomyids. Endosymbiont species found off Oregon corresponded to 2 different clusters of Calyptogena spp. symbionts in the same samples. The results corroborate the hypothesis of a monophyletic origin of the symbionts in vesicomyid clams, and support the existence of deeply branching groups in solemyid symbionts and of divergent lines and distribution for thyasirid symbionts. The results also indicate that certain symbiont species cluster according to the depth distribution of their hosts, and that in consequence host species together with their symbionts may have undergone depth-specific adaptation and evolution.

Description: New technology has allowed inter-dive oxygen-loading behaviour to be assessed in free-living Magellanic penguins Spheniscus magellanicus and shows, via pre-dive hyperventilation and oxygen loading, that birds predict both their upcoming dive performance in terms of duration and depth, and foraging success based on the number of prey they are likely to catch. Thus wild penguins make decisions based on complex environmental parameters and this leads to particularly efficient time use as expected according to optimized foraging.

Description: Observations from cruises in the Arabian Sea and data from satellites are interpreted using different realizations of a multi-level primitive equation model and an eddy-permitting reduced-gravity shallow water model of the Indian Ocean. The focus is on the interannual circulation variability of the Arabian Sea, and especially of the meridional location of the Great Whirl (GW). The results suggest that the variability in the western Arabian Sea is not only due to the interannual variability in the wind field, but that a substantial part is caused by the chaotic nature of the ocean dynamics. Decreasing the friction coefficient from 1000 to 500m2s-1 in a 19o numerical reduced-gravity model, the variance of the GW location increases dramatically, and the mean position moves southward by one degree. In the eddy-permitting experiments analyzed, both mechanisms appear to determine the GW location at the onset of the GW dynamics in late summer.

Description: Sea-surface height data acquired by the TOPEX/POSEIDON satellite over the Arabian Sea from October 1992 to October 1998 are analyzed. Strong seasonal fluctuations are found between 61 and 101N, which are mainly associated with westward propagating annual Rossby waves radiated from the western side of the Indian subcontinent and that are continuously forced by the action of the wind-stress curl over the central Arabian Sea. An analysis of hydrographic data acquired during August 1993 and during January 1998 at 81N in the Arabian Sea reveals the existence of first- and second-mode annual Rossby waves. These waves, which can be traced as perturbations in the density fields, have wavelengths of 12�103 and 4.4�103km as well as phase velocities of 0.38 and 0.14 m/s, respectively. The waves are associated with a time-dependent meridional overturning cell that sloshes water northward and southward. Between 581 and 681E in the central Arabian Sea, we found a Rossby-wave induced transport in the upper 500m of about 10 Sv southward in August 1993 and northward in January 1998. Below 2000 m, there was still a northward transport of 3.2 Sv in August 1993 and a southward transport of 4.8 Sv in January 1998. A comparison of steric height differences between August 1993 and January 1998 calculated from the observed density fields as well as calculated from the reconstructed density fields using first- and second-mode annual Rossby waves agree quite well with the corresponding sea-surface height differences. Implications resulting from the reflection of annual Rossby waves, like fluctuations of the western boundary currents, are discussed.

Description: Phytoplankton (〉100 µm) abundance was studied in the open waters of the Gulf of Aqaba during the summer stratification period of 1996. A succession took place among the major phytoplankton groups, with diatom numbers decreasing throughout the summer. The diazotrophic cyanobacteria Trichodesmium spp. became more prominent as the stratification period progressed; 5 Trichodesmium species were identified: T. thiebautii, T. erythraeum with tuft-shaped colonies and Trichodesmium sp. with puff-shaped colonies were common at ~102 colonies m-3 throughout the stratification period, whereas T. tenue and T. hildebrandtii were more rare. A bloom of T. thiebautii and T. erythraeum with 〉106 tuft colonies m-3 was observed in coastal waters of the Gulf during fall 1997. Tuft-shaped colonies were dominant near the surface, while puff-shaped colonies of Trichodesmium sp. were mainly found in the bottom half of the photic zone. These depth distributions were maintained for more than 2 mo, suggesting that the 2 colony types occupied distinct niches. Puff-shaped colonies were found to have higher chlorophyll a contents than tufts, but their photosynthetic activities were not significantly different. Fatty acid analysis of dominant plankton species yielded new trophic relationships for Trichodesmium spp. The Trichodesmium spp.-specific fatty acid C22:2 ω6 was found in Macrosetella gracilis (the sole copepod to graze on Trichodesmium spp.) and in chaetognaths, suggesting that these carnivorous zooplankton fed on M. gracilis. Furthermore, this fatty acid was observed in the filter-feeding Salpa maxima, which was abundantly present in the Gulf of Aqaba during June 1997.

Description: The combined and interactive effects of climatic and ecological factors are rarely considered in marine communities. We designed a factorial field experiment to analyze (1) the interactive effects of ambient UV radiation and consumers; and (2) the effects of photosynthetically active radiation (PAR 400 to 700 nm), UVA (320 to 400 nm) and UVB (280 to 320 nm) radiation on a marine hard-bottom community in Nova Scotia, NW Atlantic. Species recruitment and succession on ceramic tiles were followed for 5 mo. We found strong negative UV effects on biomass and cover of the early colonizing macroalga Pilayella littoralis, whereas UVB was more harmful than UVA radiation. Consumers, mainly gammarid amphipods, increased P. littoralis biomass when UV was excluded, probably through fertilization. These initially strong and interacting UV and consumer effects on total biomass and cover diminished as species succession progressed. Species diversity was not affected by experimental treatments, but significant shifts in species composition occurred, especially at the recruitment stage. Red algae were most inhibited by UV, whereas sedentary invertebrates and some brown algae tended to increase under UV exposure. Consumers suppressed green and filamentous brown algae, but favored the other groups. Again, these effects diminished during the later stages of succession. We conclude that UV radiation can be a significant structuring force in early successional benthic communities, and that consumers can mediate its effects.

Description: Between 1991 and 1999, carbon measurements were made on twenty-five WOCE/JGOFS/OACES cruises in the Pacific Ocean. Investigators from 15 different laboratories and four countries analyzed at least two of the four measurable ocean carbon parameters (DIC, TAlk, fCO2, and pH) on almost all cruises. The goal of this work is to assess the quality of the Pacific carbon survey data and to make recommendations for generating a unified data set that is consistent between cruises. Several different lines of evidence were used to examine the consistency, including comparison of calibration techniques, results from certified reference material analyses, precision of at-sea replicate analyses, agreement between shipboard analyses and replicate shore based analyses, comparison of deep water values at locations where two or more cruises overlapped or crossed, consistency with other hydrographic parameters, and internal consistency with multiple carbon parameter measurements. With the adjustments proposed here, the data can be combined to generate a Pacific Ocean data set, with over 36,000 unique sample locations analyzed for at least two carbon parameters in most cases. The best data coverage was for DIC, which has an estimated overall accuracy of ∼3 μmol kg−1. TAlk, the second most common carbon parameter analyzed, had an estimated overall accuracy of ∼5 μmol kg−1. To obtain additional details on this study, including detailed crossover plots and information on the availability of the compiled, adjusted data set, visit the Global Data Analysis Project web site at: http://cdiac.esd.ornl.gov/oceans/glodap.

Description: The effect of methane released from decomposing surficial gas hydrates (SGH) on standing stocks and activities of the small-sized benthic biota (SSBB; i.e. bacteria, fungi, protozoa, and meiobenthic organisms) was studied at about 790 m water depth, at the Hydrate Ridge, Cascadia subduction zone. Presence of SGH and elevated sulfide concentrations in the sediment were indicated by extensive bacterial mats of Beggiatoa sp. and clam fields of the bivalve mollusc Calyptogena sp. Vertical and horizontal distribution patterns of the SSBB biomass were derived from DNA and total adenylate (TA) sediment assays. Potential bacterial exoenzymatic hydrolytic activity was measured using fluorescein-di-acetate (FDA) as substrate. Estimates of chemoautotrophic production of particulate organic carbon (POC.) were determined by 14CO2 uptake incubations. Inventories of chl a and pheopigments were determined as parameters of surface water primary produced POC input. Average SSBB biomass in clam field sediments integrated over the upper 10 cm (765.2 gC m-2, SD 190.1) was 3.6 times higher than in the adjacent control sites (213 gC m-2, SD 125). Average SSBB biomass in bacterial mat sediments, which were almost devoid of eukaryotic organisms 〉 31 µm, was 209 gC m-2 (SD 65). Significant correlations between FDA, DNA and plant pigments imply that productivity of the SSBB at SGH sites is only partially uncoupled from the primary production of the surface water. Areal estimates of autotrophic Corg production at control sites, bacterial mat sites and in clam field sites were 5.7, 59.7 and 190.0 mgC m-2 d-1, respectively. Based on different models predicting vertical POC fluxes from surface water primary production and water depth, these autotrophic POC productions account for 5 to 17% (controls), 35 to 68% (bacterial mats), and 63 to 87% (clam fields) of the bulk POC (sum of allochthonous POC input through the water column and sedimentary autochthonous autotrophic POC production) provided at the various sites. At SGH sites inventories of chl a and pheopigments, integrated over the upper 10 cm of the sediment, were half of that found at the control sites. This might be due to enhanced degradation of phytodetritally associated organic matter. The resulting low molecular weight organic carbon compounds might stimulate and fuel sulfate reduction, which is conducted in a microbial consortium with anaerobic methane consuming archaea. This syntrophic consortium might represent a prominent interface between gas hydrate derived carbon and allochthonous Corg flow. We infer that degradation kinetics of SGH is affected by, e.g., seasonally varying input of allochthonous organic matter.

Description: The bottom and deep circulation in the Somali Basin are investigated on the basis of hydrographic and direct velocity profiles from three shipboard surveys carried out during the southwest monsoon in 1995 and of velocity time series from the WOCE mooring array ICM7. The inflow of bottom water into the Somali Basin through the Amirante Passage drives a thermohaline circulation, which may be modulated by the monsoon wind forcing. Details of the abyssal circulation have been discussed controversially. Deep velocity records from the mooring array in the northern Somali Basin are dominated by fluctuations with periods of 30–50 days and amplitudes above Full-size image (〈1 K). Despite this strong variability annual record averages indicate the existence of a deep western boundary current (DWBC) below Full-size image (〈1 K) at the base of the continental slope south of Socotra Island as part of a cyclonic bottom circulation. The southwestward DWBC transport off Socotra Island is estimated to Full-size image (〈1 K). The bottom and deep water exchange between the Somali and Arabian Basin north of 7°N is estimated from two cross-basin geostrophic velocity sections referenced by vertically averaged LADCP currents. For the bottom water, an eastward transport into the Arabian Basin of Full-size image (〈1 K) and Full-size image (〈1 K) was determined in June and August, respectively, while for the deep-water layer above Full-size image (〈1 K) eastward transports of Full-size image (〈1 K) in June and Full-size image (〈1 K) in August were obtained.

Description: The sediment beneath and at various distances from the fish farm cages in Cephalonia bay (Eastern Mediterranean) was investigated seasonally through sediment profiling imagery (SPI) as well as through monitoring of geochemical variables and macrofaunal assemblages. The SPI images (SPI) repeatedly showed the same qualitative pattern along the benthic enrichment gradient with readily identifiable attributes such as depth of dark sediment, signs of outgassing and bioturbation marks. Quantitative comparisons showed that a large number of SPI attributes showed significant positive or negative correlation with geochemical and biological attributes describing the effect of fish farming on the seabed. All multivariate patterns obtained through the analysis of SPI attributes were highly correlated to those obtained from standard multivariate analysis of macrofauna during the respective seasons. It is argued that SPI provides an integrated assessment of the sedimentary conditions and therefore may be used as a complement of or even a substitute for standard sampling methods when mapping the effects of aquaculture on silty substrates

Description: Sediment incubation microcosms, multitrap apparatus and water column variables have been employed to describe the dynamic changes in benthic-pelagic coupling between nutrient pools in the Pontevedra ria, NW Spain, during spring and summer 1998. A comparison of the chemical characteristics of suspended and bed sediment together with sediment nutrient effluxes revealed that particulate organic nitrogen and carbon were progressively depleted upon transit through the nutrient pools. The main fate of particulate organic nitrogen reaching the bed sediment is denitrification, although resuspension is also important. An estimate of net denitrification (Dnet) was made at 2 muddy sites in the ria with a mass balance at the benthic boundary layer. First-order approximations calculate Dnet to be 178 and 182 µmolN m-2 h-1 at the 2 stations, and agree well with previous modelling estimates. Denitrification is highest when upwelling relaxes and the flux of organic matter to the sediment increases. Regular inputs of offshore water ensure water renewal and re-oxygenation of bottom waters, thus preventing anoxia, particularly in the summer. With upwelling, large quantities of ammonium are effluxed to the water column (250 µmolNH4+ m-2 h-1), probably as a result of the bed sediment resuspension engendered by upwelling and stirring of phytodetrital fluff held in suspension as neutrally buoyant material above the sediment surface. We hypothesise that hydrodynamical processes play an important role in determining the quantity of nutrients remineralised in the Pontevedra ria and, in the case of nitrogen, the rate of denitrification at the benthic boundary layer.

Description: Gas hydrates occur at the sediment surface on the southern summit of Hydrate Ridge, Cascadia convergent margin. The hydrates are found in mounds several meters in diameter and up to 2 m high, and are covered by sediment and mats of the filamentous sulfur-oxidizing bacteria Beggiatoa. The mounds are surrounded by vesicomyid clams (Calyptogena pacifica, C. kilmeri), which in turn are encircled by solemyid bivalves (Acharax sp.). The zonation pattern of 3 species (Calyptogena spp. and Acharax sp., which harbor chemoautotrophic bacteria in their gills, and the chemoautotrophic Beggiatoa), is also reflected in a change in the entire community structure. Beggiatoa, Calyptogena spp. and Acharax sp. are shown to be characteristic species for the different communities. The Beggiatoa community directly overlaying the gas hydrates consists of seep endemic species in high densities: gastropods (Provanna laevis, P. lomana, Pyropelta corymba, Hyalogyrina sp. nov.), bivalves (Nuculana sp. nov.) and polychaetes (Ampharetidae, Polynoidae, Dorvilleidae). Based on pooled samples, the rarefaction curves show a decrease in species diversity in the Beggiatoa and Calyptogena communities. The hydrogen sulfide gradients in the porewater of sediments below the different communities dominated by either Beggiatoa, Calyptogena spp. or Acharax sp. vary by 3 orders of magnitude. The diffusive sulfide flux based on the measured sulfide concentration gradients is highest in Beggiatoa sp. communities (23 ± 13 mol m-2 yr-1), slightly less in Calyptogena communities (6.6 ± 2.4 mol m-2 yr-1), and low in Acharax communities (0.05 ± 0.05 mol m-2 yr-1). The difference in the sulfide environment is a factor influencing the distribution patterns of the chemoautotrophy-dependant and heterotrophic species at the deep-sea sediments containing gas hydrate.

Description: A colony of Humboldt penguins Spheniscus humboldti in central Chile was monitored from August 1995 to July 2000 to determine patterns of breeding and colony attendance and how these were affected by climatic (rainfall) and oceanographic (El Niño) factors. Nests were periodically checked for contents and roosting birds were counted from vantage points. Two main breeding events were observed: between August and January (spring event) and between April and June (autumn event). Whereas the spring event regularly produced offspring, the autumn event was systematically affected by rains, causing considerable nest desertion. Adults were present in the colony from August to May, abandoning the colony during winter after the nests were flooded. Juveniles occurred only between November and March. Adults moulted mainly in February, while juveniles moulted in January. During the 1997/98 El Niño episode, the number of breeding pairs was 55 to 85% lower than the mean, the onset of nesting was delayed, and abnormally heavy rainfall flooded nests. While the number of breeding pairs was significantly related to sea surface temperature anomalies (SSTA), breeding success was not. The attendance of adults and juveniles at the colony during El Niño was 25 and 73% lower, respectively, than the mean attendance. This 2-peak breeding strategy of Humboldt penguins appears to have evolved in response to the more favourable oceanographic and climatic conditions of Perú, where breeding is continuous and not interrupted by rains. Although less productive, the species probably maintains its autumnal breeding in central Chile because this provides additional offspring to supplement those regularly produced during the spring event.

Description: Zooplankton grazing on bacterio- and phytoplankton was studied in the Gulf of Aqaba and the Northern Red Sea during Meteor Cruise Me 44-2 in February-March 1999. Protozoan grazing on bacterioplankton and autotrophic ultraplankton was studied by the Landry dilution method. Microzooplankton grazing on phytoplankton 〉6 µm was studied by incubation experiments in the presence and absence of microzooplankton. Mesozooplankton grazing was studied by measuring per capita clearance rates of individual zooplankton with radioactively labelled food organisms and estimating in situ rates from abundance values. Protozoan grazing rates on heterotrophic bacteria and on algae 〈6 µm were high (bacteria: 0.7 to 1.1 d-1, ultraphytoplankton: 0.7 to 1.3 d-1), while grazing rates on Synechococcus spp. were surprisingly low and undetectable in some experiments. Mesozooplankton grazing was weak, cumulative grazing rates being ca. 2 orders of magnitude smaller than the grazing rates by protozoans. Among mesozooplankton, appendicularians specialised on smaller food items and calanoid copepods on larger ones.

Description: Factors responsible for aggregations of jumbo flying squid Dosidicus gigas, an important component of the marine food web and target of commercial fisheries off the Costa Rica Dome in the Eastern Tropical Pacific Ocean (ETP), were examined during 2 years of different extreme oceanographic conditions: fall 1997 El Niño and fall 1999 La Niña. A high abundance of squid occurred in association with the well-developed countercurrent ridge (upwelling) off the Costa Rica Dome during fall 1997, but not during fall 1999, when the countercurrent ridge was less developed. Two features of the well-developed countercurrent ridge were considered important for the occurrence of high jumbo flying squid concentrations. Firstly, subsurface chlorophyll a (chl a) maxima were formed along the countercurrent ridge, resulting in integrated chl a concentrations in the upper 100 m being relatively high considering the generally low productivity of the ETP during an El Niño event. Secondly, a strong salinity front formed along the North Equatorial Countercurrent, which is possibly responsible for retention of jumbo flying squid in the ridge. Large yellowfin tuna Thunnus albacares, which mainly feed on micronekton (small fishes, cephalopods and swimming crabs), as do jumbo flying squid, were also more highly concentrated along the countercurrent ridge during 1997 than during 1999. It was noted that skipjack tuna Katsuwonus pelamis and small yellowfin tuna, which mainly feed on zooplankton, were associated with the equatorial ridge in the ETP, indicating that prey faunal components may also play an important part in the close association of jumbo flying squid with the countercurrent ridge.

Description: The differences in the water mass distributions and transports in the Arabian Sea between the summer monsoon of August 1993 and the winter monsoon of January 1998 are investigated, based on two hydrographic sections along approximately 8°N. At the western end the sections were closed by a northward leg towards the African continent at about 55°E. In the central basin along 8°N the monsoon anomalies of the temperature and density below the surface-mixed layer were dominated by annual Rossby waves propagating westward across the Arabian Sea. In the northwestern part of the basin the annual Rossby waves have much smaller impact, and the density anomalies observed there were mostly associated with the Socotra Gyre. Salinity and oxygen differences along the section reflect local processes such as the spreading of water masses originating in the Bay of Bengal, northward transport of Indian Central Water, or slightly stronger southward spreading of Red Sea Water in August than in January. The anomalous wind conditions of 1997/98 influenced only the upper 50–100 m with warmer surface waters in January 1998, and Bay of Bengal Water covered the surface layer of the section in the eastern Arabian Sea. Estimates of the overturning circulation of the Arabian Sea were carried out despite the fact that many uncertainties are involved. For both cruises a vertical overturning cell of about 4–6 Sv was determined, with inflow below 2500 m and outflow between about 300 and 2500 m. In the upper 300–450 m a seasonally reversing shallow meridional overturning cell appears to exist in which the Ekman transport is balanced by a geostrophic transport. The heat flux across 8°N is dominated by the Ekman transport, yielding about –0.6 PW for August 1993, and 0.24 PW for January 1998. These values are comparable to climatological and model derived heat flux estimates. Freshwater fluxes across 8°N also were computed, yielding northward freshwater fluxes of 0.07 Sv in January 1998 and 0.43 Sv in August 1993. From climatological salinities the stronger freshwater flux in August was found to be caused by the seasonal change of salinity storage in the Arabian Sea north of 8°N. The near-surface circulation follows complex pathways, with generally cyclonic-circulation in January 1998 affected at the eastern side by the Laccadive High, and anticyclonic circulation in August 1993.

Description: Three View the MathML sourceacoustic transceivers were deployed on either side of the eastern entrance of the Strait of Gibraltar during April–May 1996 to determine the feasibility of using acoustic methods to make routine, rapidly repeated, horizontally integrated measurements of flow and temperature in straits. Reciprocal transmissions between the transceivers were used to test the feasibility of using traditional ray differential travel times to monitor the component of flow along the acoustic paths. Transmissions directly across the Strait were used to test the feasibility of using horizontal arrival angle fluctuations and acoustic intensity scintillations to monitor the flow perpendicular to the acoustic path. The geometry was selected to provide ray paths that only sample the lower-layer Mediterranean water, so that the feasibility of monitoring the Mediterranean outflow using the various methods could be evaluated. The acoustic scintillation method did not yield useful current estimates, but the experimental parameters were not optimized for this approach. Since the low-frequency variability in log-amplitude was found to be highly correlated at receivers View the MathML source apart, it is possible that acoustic scintillation measurements using different receiver spacings and more rapid sampling might yield better results. The horizontal deflection method gave encouraging results at the time of neap tides, but less so during spring tides. For this approach, both theoretical estimates and measured phase differences between the horizontally separated receivers suggest that internal-wave-induced horizontal arrival angle fluctuations may fundamentally limit the precision with which arrival angles can be measured. Further work is needed to determine if a smaller horizontal spacing and higher signal-to-noise ratios would yield better results. Reciprocal travel time measurements diagonally across the Strait performed the best of the three methods, giving absolute flow estimates consistent with those derived from current-meter data. The fractional uncertainty variance for the lower layer tidal transport from a single tomographic path was estimated to be 0.017 (i.e. 98% of the a priori tidal transport variance was resolved). The spatial scales of the sub-tidal flow are thought to be significantly shorter than those of the tidal flow, however, which means that a more elaborate monitoring network is required to achieve the same performance for sub-tidal variability. Finally, sum travel times from the reciprocal transmissions were found to provide good measurements of the temperature and heatcontent in the lower layer.

Description: The late Tertiary tectonic and geological evolution of the Southern Andes at 33–34°S has been strongly controlled by the Challenger–Juan Fernández–Maipo (CH–JF–M) structural system. The present configuration of a flat slab between 28–33°S may be explained by a series of favorable conditions evolving with time since the breakup of the Farallon Plate at 25 Ma. The dramatic shift of the pole of rotation and the rapid eastward propagating rift along the Challenger Fracture Zone induced an unbalanced slab pull force, south of the CH–JF–M, that may have triggered the detachment of the subducting slab. The upwelling of a warmer asthenospheric material and the partial melts of the slab are likely consequences that are consistent with the anhydrous tholeiitic late Oligocene volcanism and the anomalous adakite-type magmatism of the early Miocene, respectively. The present seismogenic zone across the CH–JF–M tectonic boundary shows a continuity for more than 600 km along the flat slab segment, in contrast with the much shorter slab southward. Such a tectonic configuration is probably a quasi-steady condition since 25 Ma. Gravity modeling along the JF chain and the broadly located focal mechanisms at the locus of the already subducted JF chain indicate a thick (〉25 km), wide (100 km), and continuous belt of lighter oceanic crust, which is the major contribution to the positive buoyancy of the slab. The decoupling of the subducted slab at intermediate depths further contributes to the flattening of the slab, focusing the buoyancy forces associated with the thickened oceanic crust along the already subducted JF chain. The absolute plate reconstruction during the Miocene shows that the JF collision against the margin migrates southward, in agreement with geological and tectonic observations that further support the causative relationship between the flattening of the slab and the subduction of the JF chain. Preliminary deformation models for the indentation of the JF ridge against the continental lithosphere is consistent with the particular east–west trend of the Maipo deformation zone, connecting in this way the CH–JF–M major tectonic boundary in the ocean-continent lithosphere system.

Description: In this paper, different types of new data sets (hydrography, Lagrangian and Eulerian current measurements) and Quasi-Geostrophic model results, obtained after altimetric data assimilation, are used to study the structure and variability of the Azores Current and Frontal system. The Azores Current was observed to transport between 11.0 and 18.0 Sv eastwards and the associated Counter Current some 2.0–10.0 Sv to the west, resulting in a net value of about 8–9 Sv. Furthermore, both data and model results revealed a meandering Azores Current, where some freely rotating eddies were also identified. These hydrographic and Lagrangian results exhibit space and time scales that agree fairly well with the dynamics shaped by baroclinic instability. Current meters moored across the Azores Current system delineated a mean Counter Current flowing westwards with a maximum subsurface core of about 2.0 cm/s at a mean depth of 800 dbar. This is in excellent agreement with previous studies, which explains this Azores Counter Current as the rectification process of the geostrophic turbulence occurring north of the main Azores Current stream. A new scheme is proposed for the formation mechanism of the freely rotating cyclonic eddies observed south of the Azores. It is shown that the north–south contrast thickness of the 17–19°C water layer across the Azores Front decreases downstream. This will create, in turn, a downstream increase of the most unstable wavelength, in a non-linear baroclinic instability context. As a consequence, both large cyclonic and anticyclonic features are able to form at the eastern side of the Azores Current (around 19°W), while at the western side (around 35°W) only large anticyclones will survive (western-generated cyclones will be small enough to be quickly dissipated). This means that the eastern cyclones of the Azores Current may live longer than the shorter western ones. However, because longer-lived cyclonic eddies propagate westwards with a mean speed of 2.5 km/day, they may be observed south of Azores and of the main stream several months later, although they were not formed there.

Description: We examined how seabirds might be used to study marine environmental variables, which necessitates knowing location and the value of the variable to be studied. Five systems can potentially be used for determination of location: VHF (Very High Frequency) telemetry, PTT (Platform Terminal Transmitters) telemetry, GLS (Global Location Service) geolocation methods, dead reckoning and GPS (Global Positioning System), each with its own advantages with respect to accuracy, potential number of fixes and size. Temperature and light were used to illustrate potential difficulties in recording environmental variables. Systems currently used on seabirds for measurement of temperature respond slowly to environmental changes; thus, they may not measure sea surface temperature adequately when contact periods with water bodies are too short. Light can be easily measured for light extinction studies, but sensor orientation plays a large role in determining recorded values. Both problems can be corrected. The foraging behaviour of seabirds was also examined in order to identify those features which would be useful for determination of marine environmental variables at a variety of spatial and temporal scales. Area coverage by birds is highly dependent on breeding phase and tends to be concentrated in areas where prey acquisition is particularly enhanced. The identification of these sites may be of particular interest to marine biologists. 'Plungers' and 'divers' are potentially most useful for assessment of variables deeper within the water column, with some divers spending up to 90% of their time sub-surface. Few seabirds exploit the water column deeper than 20 m, although some divers regularly exceed 50 m (primarily penguins and auks), while 2 species dive in excess of 300 m. The wide-ranging behaviour of seabirds coupled, in many instances, with their substantial body size makes them potentially excellent carriers of sophisticated environmental measuring technology; however, the ethical question of how much the well-being of birds can, and should, be compromised by such an approach needs to be carefully considered.

Description: To study the EasternBoundaryCurrentsystem off Northwest Africa in detail several CTD/ADCP-sections and long-term mooring work were carried out in the channel between Lanzarote and Africa. The observations are compared with a fine-resolution model, which was developed in the framework of the CANIGO project. The water masses, which are observed in this area, are characterised and classified in density ranges. The current field shows a high spatial and temporal variability with maximum velocities of about 35 cm/s. Seasonal means as well as currents averaged across the channel are only a few cm/s. In the surface water a steady southward flow in the middle of the channel indicates the CanaryCurrent in this area. During fall a strong northward current is observed close to the African shelf. Though the CanaryCurrent strengthens during summer and fall due to an increase of the trade winds, the transport in the channel decreases or turns northward during that time due to the enhanced poleward current at the eastern side. A northward undercurrent with a mean velocity of +2.3 cm/s is observed at the African slope in 950 m depth. The poleward transport of AAIW increases during fall and a strong influence of relatively fresh AAIW is observed during that time. Most of the observations fit well to the results of the CANIGO model, but the occurrence of MW at the bottom of the channel and the corresponding southward flow cannot be resolved by the model.

Description: Light abundance is a major prerequisite for primary production in pelagic ecosystems, influencing the evolution of the marine environment. Realistic simulations of planktonic ecosystems therefore require an appropriate representation of the underwater light field. Taking a look at the different biogeochemical models discussed in literature, one finds a variety of descriptions for the distribution of light, or more specific, photosynthetically available radiation (PAR) in the water column. This paper compares the effect of different parameterizations of PAR on the primary production and phytoplankton evolution at the European Station for Time-Series in the Ocean Canary Islands (ESTOC) station (29°10′N, 15°30′W) north of Gran Canaria, Canary Islands. Observations from two cruises in 1997 are used to illustrate the winter and spring situation at the time-series site. Four alternative PAR descriptions are used in a one-dimensional coupled biological–physical model of the upper ocean driven by daily forcing fields over a 5-year period. The biological model is a simple nitrate-phytoplankton–zooplankton-detritus model. Although the different descriptions are found to have only a small effect (±3%) on the annual primary production, we observe significant changes in the vertical distribution of simulated phytoplankton. The large variation (±32%) in the near-surface chlorophyll contents will be of particularly crucial importance when using satellite ocean-color sensors for model validation and parameter estimation. For future three-dimensional biogeochemical models, a computationally efficient and accurate parameterization of the light field will be particularly relevant.

Description: We report how different zooplankton groups (doliolids, cladocerans and copepods) are able to influence the coastal pelagic food web, including the microbial food web, in waters of the NW Mediterranean. We studied the effect of grazing and of grazing-induced nutrient recycling mediated by different types of zooplankton grazing on a natural phytoplankton community. Experiments were conducted in semicontinuous 2-stage chemostats. The 1st stage vessels contained seawater from Blanes Bay, Spain (NW Mediterranean) including its natural phytoplankton community; the 2nd stage vessels contained the same seawater and copepods, cladocerans or doliolids. At daily intervals we transferred part of the medium from the 2nd to the 1st stage flasks, which contained ungrazed algae and excreted nutrients. In this way, the zooplankton could influence phytoplankton dynamics both by selective grazing and by differential excretion of limiting nutrients. In the 2nd stage flasks grazing changed the algal community composition. Doliolids and cladocerans promoted the growth of large algae and copepods shifted the size spectrum towards small sizes. This effect was transferred to the 1st stage flasks. Doliolids, cladocerans and copepods also affected the microbial food web in different ways. Size-selective grazing led to differences in the nanoplankton concentrations. These in turn affected bacterial concentrations in a trophic cascade. The potential to modify a given algal population increased with increasing selectivity of the grazer.

Description: Both surface texture and littorinid grazing are known to influence the establishment of many shallow-water benthic hard-bottom communities. However, the effects of these factors and their interactions have not yet been investigated in a quantifiable manner. This investigation aims to assess the interactive effect of both factors in a strictly standardized manner. Natural recruitment by diatoms, the barnacle Balanus improvisus and the tube-building polychaete Polydora sp. was monitored under a 2-factorial treatment: grazing by the periwinkle Littorina littorea (Factor 1; 2 levels: I or no snails per plate) on artificial recruitment plates of different initial surface rugosities (Factor 2; 5 levels: smooth, 0.1, 0.5, 1 and 5 mm rugosity elements), In the absence of grazers, barnacle recruitment decreased with increasing initial rugosity, polychaete recruitment peaked at intermediate rugosities, and diatoms exhibited contrasting recruitment patterns in an in vitro and an in situ experiment. When preferred recruitment sites coincided for Polydora and B. improvisus, a competition for space could be inferred from a negative correlation between the 2 species. However, when the overlap of requirements weakened on the 5 rugosities, the relationship was positive, but was not statistically significant. Grazing efficiency by L. littorea depended on initial rugosity, generally showing minimum values on intermediate rugosities which is attributable to a mismatch between radula dimensions and surface structures in these rugosity classes. Additionally, grazing effects tended to increase with higher prey densities. As all factors-initial rugosity, grazing, colonizer species-interact with each other, the outcome of recruitment under combined factors is difficult to predict from single factor effects.

Description: The living coelacanth Latimeria chalumnae has a unique position in world biodiversity which raises important questions about conservation and ethics. Some relevant details of coelacanth biology are summarized, including those obtained by direct observation from submersibles. The importance of the coelacanth for evolutionary theory and palaeontology is shown to be paralleled in cultural, literary and artistic areas of human heritage. Threats to the Comoran coelacanths from artisanal fishing are described and conservation measures discussed in relation to local customs and economies as well as the promotion of tourism to spread a new awareness and concern for coelacanths worldwide.

Description: Filamentous sulfur-oxidizing bacteria and geochemical parameters of sediments at the Makran accretionary wedge in the northeastern Arabian Sea off Pakistan were studied. The upper continental slope between 350 and 850 m water depth, which is in the center of the oxygen-minimum zone, is characterized by numerous sites of small-scale seeps of methane- and sulfide-charged porewater. White bacterial mats with diameters 〈1 m were discovered at the surface of these sites using a photo-TV sled. Seep sediments, as well as non-seep sediments, in the vicinity were characterized by the occurrence of the bacterium Thioploca in near-surface layers between 0 and 13 cm depth. Thioploca bundles were up to 20 mm in length and contained up to 20 filaments of varying diameters, between 3 and 75 µm. Up to 169 ind. cm-2 were counted. Maximum numbers occurred in the top 9 cm of sediment, which contained very low concentrations of soluble sulfide (〈0.2 µM) and high amounts of elemental sulfur (up to 10 µmol cm-3). Moderate sulfate reduction activity (between 20 and 190 nmol cm-3 d-1) was detected in the top 10 cm of these sediments, resulting in a gradual downcore decrease of sulfate concentrations. CO2 fixation rates had distinct maxima at the sediment surface and declined to background values below 5 cm depth. The nutritional implications of the distinct morphology of Thioploca and of the geochemical setting are discussed and compared to other sites containing Thioploca communities.

Description: Natural marine bacteria populations collected from nearshore waters produce different types of siderophores depending on the degree of iron limitation. These siderophores can facilitate iron uptake in the marine diatom Phaeodactylum tricornutum. Water samples from 15 stations along the Italian coast of the northwest Adriatic Sea were collected and filter fractionated (3.0, 0.8 and 0.2 µm). Siderophore production in the fractions was determined using cross-feeding experiments with siderophore-auxotrophic bacteria. At most stations sampled, bacteria collected in the 3.0 and 0.8 µm filters produced siderophores which stimulated growth in Morganella morganii, the indicator strain for α-keto/ α-hydroxy acids. The results suggest that MGF (ŒMorganella-Growth Factor¹) production is common among filamentous and appendaged bacteria or strains associated with particles. Natural bacteria populations grown in iron-deficient media stimulated growth of all the indicator strains in the cross-feeding tests. Examples of known MGF which supply iron to M. morganii were tested for their ability to act as iron source for the marine diatom P. tricornutum. Iron uptake from 55Fe-MGFs was measured in P. tricornutum cells grown in Fe-sufficient and Fe-deficient media. Unchelated iron (55FeCl3 ) and 55FeEDTA were used as controls. The uptake of iron from the 55Fe-MGF and 55FeCl3 by Fe-deficient cells was higher (109 to 150 pgFe mg-1) than from 55FeEDTA (34 pgFe mg-1). Similarly, Fe-sufficient P. tricornutum took up iron from the 55Fe-MGF and 55FeCl3 to the same extent (~50 pgFe mg-1) while minimal uptake (8 pgFe mg-1) was measured from FeEDTA. In growth experiments where iron-deficient diatom cells were incubated in media containing different sources of iron, e.g. FeCl3, Fe-MGF and FeEDTA, a greater increase in number was observed in cells supplied with Fe-MGF. Further experiments also show that the uptake of Fe from MGF was enhanced by light and that a reduction step was involved in the uptake process. MGF also promoted the uptake of colloidal ferrihydrites. This study gives further evidence that siderophores produced by bacteria can be utilized by phytoplankton as an iron source. We therefore suggest that these substances play an important role in increasing the availability of iron to phytoplankton in coastal waters and thus are major factors defining the chemistry of iron in the marine environment.

Description: Marine angiosperms, or seagrasses, continue to be a major focus of marine biologists because of their important ecological role in many coastal ecosystems. Seagrass population biology could benefit from a population genetic perspective because genetic data enable the extraction of useful demographic information such as isolation and gene flow between demes. Moreover, population genetic processes may contribute to the growing ecological risks of local population extinction. Progress in seagrass genetics is partly driven by novel genetic markers which detect variation at the DNA level and overcome the limited polymorphism of allozymes. Key results of studies in the past decade, mostly using RAPD and microsatellites, were (1) considerable genetic and genotypic (clonal) diversity is present in several species in contrast to earlier notions of low polymorphism detected at allozyme loci, and (2) genetic differentiation among populations seems to be the rule despite earlier reports of genetic uniformity. Pronounced genetic structure was detected between populations of 4 species examined thus far (Posidonia oceanica, P. australis, Zostera marina, Thalassia testudinum). The FST estimates varied widely and ranged from 0.01 to 0.623 across studies and species. Genetic differentiation at a systematic range of scales was only studied in eelgrass Zostera marina, where it was positively correlated with geographic distance. The high polymorphism of RAPD or microsatellite markers will allow the augmention of indirect estimates of gene flow by methods detecting individual immigration events through paternity analysis or assignment tests. Important conservation related issues such as the level of inbreeding and the effective population size have also been obtained from genetic marker data, but results are too scarce at the moment to allow generalizations. In Zostera marina and Posidonia australis, several population genetic attributes such as clonal diversity, mating system and effective population size varied among populations within species, highlighting that there is no Œtypical¹ population. An important gap in our knowledge is whether the effects of natural population fragmentation and patchiness enhance the genetic isolation of populations due to anthropogenic disturbances. It is also unclear whether genetic differentiation displayed at marker loci are correlated with fitness-related plant traits, and whether genetic or genotypic diversity is important for medium- to long-term meadow persistence. An assessment of the genetic and genotypic diversity at marker loci should be combined with experiments on the ecological plasticity and reaction norms of genotypes composing the populations in question. This way, the role of genetic diversity for seagrass population maintenance and growth in the face of changing environmental conditions can be evaluated.

Description: Previous work has shown that during early summer, the partial pressure of CO2 (pCO(2)) in surface waters north of about 45 degreesN in the Atlantic exhibits widespread undersaturation. In many areas. this follows after a "spring bloom" of phytoplankton, at which time, nutrient concentrations and pCO(2) decrease sharply from their winter surface values. As part of OMEX I, the late summer distribution of surface water pCO(2) was surveyed in the northeastern Atlantic on cruises of R/V Poseidon and R/V Belgica in 1995. The pattern of the surface distribution of the sea-air pCO(2) difference (Delta pCO(2)) measured on these ship surveys was generally iri accord with that observed in this area in early to mid-summer of 1981. The greatest CO2 undersaturation (-95 mu atm) during our surveys was observed near the west coast of Iceland, with Delta pCO(2) increasing to about -60 mu atm away from the coast. In shelf waters south of Ireland, the pCO(2) was relatively higher than in surface waters of the open ocean adjacent to the Celtic Shelf margin, but the Celtic Shelf waters were still undersaturated relative to the atmospheric CO2 concentration. Because of the variation of wind speed, the synoptic distribution of air-sea CO2 flux, derived from the transfer velocity and Delta pCO(2), does not resemble the distribution of Delta pCO(2) itself. The sharp increase in wind speed at about 53 degreesN, 20 degreesW during the R/V Poseidon survey produces an order of magnitude rise in the estimated air-sea flux of CO2, to a level of about 10-14 mol m(-2) a(-1). The overall synoptic picture appears to be one of moving centers of higher air-sea fluxes that occur where storms pass over regions of surface water pCO(2) undersaturation.

Description: Assimilation experiments with data from the Bermuda Atlantic Time-series Study (BATS, 1989¯1993) were performed with a simple mixed-layer ecosystem model of dissolvedinorganic nitrogen (N), phytoplankton (P) and herbivorous zooplankton (H). Our aim is to optimize the biological model parameters, such that the misfits between model results andobservations are minimized. The utilized assimilation method is the variational adjoint technique, starting from a wide range of first-parameter guesses. A twin experiment displayedtwo kinds of solutions, when Gaussian noise was added to the model-generated data. The expected solution refers to the global minimum of the misfit model-data function, whereasthe other solution is biologically implausible and is associated with a local minimum. Experiments with real data showed either bottom-up or top-down controlled ecosystemdynamics, depending on the deep nutrient availability. To confine the solutions, an additional constraint on zooplankton biomass was added to the optimization procedure. Thisinclusion did not produce optimal model results that were consistent with observations. The modelled zooplankton biomass still exceeded the observations. From the model-datadiscrepancies systematic model errors could be determined, in particular when the chlorophyll concentration started to decline before primary production reached its maximum. Adirect comparision of measured 14C-production data with modelled phytoplankton production rates is inadequate at BATS, at least when a constant carbon to nitrogen C : N ratio isassumed for data assimilation.

Description: Phylogenetic analyses, using 482 bp of the mitochondrial 16S rDNA and 461 bp of the control region of 16 Diplodus species and Oblada melanura, Pagellus bogaraveo and Pagellus acarne, all close relatives of Diplodus, identified the two representatives of Pagellus as the sister group of Diplodus. Oblada melanura was confirmed as the sister taxon of D. puntazzo, despite its different dental morphology and ecology. Within the genus Diplodus, three clades were identified, the first containing D. annularis and D. bellottii, the second D. vulgaris and D. prayensis, and the third comprising three subclades. These were formed by O. melanura clustering with D. puntazzo, D. fasciatus with D. cervinus, and by the Diplodus sargus sub-species assemblage which also included the West Atlantic taxa D. argenteus, D. bermudensis, D. holbrooki, and the Red Sea endemic D. noct. All members of the D. sargus assemblage were genetically closely related. Among them, D. sargus lineatus from the Cape Verde islands was resolved as most ancestral branch, pointing to the possibility that the diversification and spread of the D. sargus assemblage originated in this region. The hypothesis of stepwise speciation following colonization events within the D. sargus complex is fully supported by phylogenetic reconstruction.

Description: In the framework of the Ocean Margin Exchange project, a multi-disciplinary study has been conducted at the shelf edge and slope of the Goban Spur in order to determine the spatial distribution, quantity and quality of particle flux, and delineate the transport mechanisms of the major organic and inorganic components. We present here a synthesis view of the major transport modes of both biogenic and lithogenic material being delivered to the open slope of the Goban Spur. We attempt to differentiate between the direct biogenic flux from the surface mixed layer and the advective component, both biogenic and lithogenic. Long-term moorings, instrumented with sediment traps, current meters and transmissometers have yielded samples and near-continuous recordings of hydrographic variables (current direction and speed, temperature and salinity) and light transmission for a period of 2.5 years. Numerous stations have been occupied for CTD casts with light transmission and collection of water samples. The sedimenting material has been analysed for a variety of marker compounds including phytoplankton pigments, isotopic, biomineral and trace metal composition and microscopical analyses. These samples are augmented by seasonal information on the distribution and composition of fine particles and marine snow in the water column. The slope shows well-developed bottom nepheloid layers always present and intermediate nepheloid layers intermittently present. Concentrations are mainly in the range 50–130 mg m−3 in nepheloid layers and 6–25 mg m−3 in clear water. A seasonal variability in the concentration at the clear water minimum is argued to be related to seasonal variations in vertical flux and aggregate break-up in transit during summer months. It is suggested that the winter sink for this seasonal change in particulate matter involves some re-aggregation and scavenging, and some conversion of particulate to dissolved organic matter. This may provide a slow seasonal pump of dissolved organic carbon to the deep ocean interior. Differences in trapped quantities at different water depths are interpreted as due to lateral flux from the continental margin. There is a major lateral input between 600 and 1050 m at an inner station and between 600 and 1440 m at an outer one. The transport is thought to be related to intermediate nepheloid layers, but those measured are too dilute to be able to supply the flux. Observed bottom nepheloid layers are highly concentrated very close to the bed (up to 5 g m−3), with a population of large aggregates. Some of these are capable of delivering the flux seen offshore during intermittent detachment of nepheloid layers into mid-water. Concentrated bottom nepheloid layers are also able to deliver large particles with unstable phytoplankton pigments to the deep sea floor in a few tens of days. Calculated CaCO3 fluxes are adjusted for dissolution, which is inferred from Ca/Al ratios to be occurring in the CaCO3-saturated upper water column where up to 80% of the CaCO3 resulting from primary production is dissolved.

Description: Nitrogen fixation rates and related plankton parameters were determined at 8 stations in the Baltic proper and Mecklenburg Bay in 1997 and 1998. Nitrogen fixation was measured with 15N-tracer method in unenriched samples. Measurable nitrogen fixation rates were found from July to October, with highest rates in August when heterocystous cyanobacteria formed blooms. Nitrogen fixation rates measured during a moderate bloom in 1998 were related to biomass of heterocystous and coccoid (non-heterocystous) cyanobacteria and primary production. The size fraction 〈10 µm contributed significantly to total nitrogen fixation both during day and night. It is discussed whether this may be due to small, non-heterocystous cyanobacteria, which were abundant in summer and autumn. They may separate photosynthesis and nitrogen fixation temporally (day and night) and may be especially responsible for the high nitrogen fixation rates observed in the dark. As the fraction of pico- and nanoplankton was not considered in earlier studies, a new budget of nitrogen fixation in the Baltic proper has been estimated. On average, daily nitrogen fixation rates of 2.5 mmol N m-2 d-1 (in July/August 1997/1998) and mean annual nitrogen fixation of 125 mmol N m-2 yr-1 were estimated for the Baltic proper. The high variability is discussed. For summer 1998, a budget of the nitrogen cycle at the main station in the Gotland Sea is give

Description: Small cetaceans are susceptible to incidental mortality in the various forms of gillnet fisheries throughout their range. Research conducted since 1994 has shown that acoustic alarms (pingers) emitting high-frequency pulsed sounds effectively reduce the number of harbor porpoise Phocoena phocoena casualties in sink gillnets. However, the mechanisms behind the effects of pingers were still not understood. Until now, advantages and risks associated with their widespread use could not be evaluated. Here we present the results of 2 field experiments: (1) theodolite-tracking of harbor porpoises exposed to a single PICE-pinger in Clayoquot Sound, Vancouver Island, Canada and (2) herring Clupea harengus capture rates in surface gillnets equipped with and without acoustic alarms (Dukane Netmark 1000, Lien, PICE) in the Baltic Sea herring fishery at Rügen Island, Germany. Our results show that harbor porpoises do not seem to react to an experimental net in their foraging area (n = 172 groups, median group size = 2 porpoises). Porpoise distance from the mid-point of the net was distributed around a median of only 150 m (range 4 to 987 m). A net equipped with an acoustic alarm, however, was avoided (n = 44 groups) within audible range (distance distribution median = 530 m, range 130 to 1140 m). The porpoises were thus effectively excluded from the ensonified area. Herring, one of the main prey species of harbor porpoises, were not affected by the acoustic alarms tested (n = 25407 fish captured). The advantages and risks of using acoustic alarms to mitigate by-catch are discussed.

Description: The carbon and nitrogen content of transparent exopolymer particles (TEP) was determined and related to the concentration of TEP as quantified by a colorimetrical method. TEP were produced in the laboratory from dissolved precursors by laminar or turbulent shear. Dissolved precursors were obtained by 0.2 µm filtration from diatom cultures, with or without nutrient reduction, and from natural diatom populations. The relationship between carbon and TEP was significant, linear and species-specific. Carbon concentration of TEP derived from this relationship concurred with previous findings. Shortage of silicic acid or nitrate in the culture media had no effect on the carbon content of TEP. Molar C:N ratios of TEP were above the Redfield ratio, with a mean value of 26. It is suggested that the nitrogen fraction of TEP can be explained by adsorption of dissolved organic nitrogen (DON) onto TEP. Based on the newly established relationship, concentrations of TEP-derived carbon (TEP-C) were calculated for the Baltic Sea, the coastal Pacific, the North East Atlantic and the Northern Adriatic Sea.

Description: The benthic diagenetic model OMEXDIA has been used to reproduce observed benthic pore water and solid phase profiles obtained during the OMEX study in the Goban Spur Area (N.E. Atlantic), and to dynamically model benthic profiles at site OMEX III (3660-m depth), with the sediment trap organic flux as external forcing. The results of the dynamic modelling show that the organic flux as determined from the lowermost sediment trap (400 metres above the bottom) at OMEX III is insufficient to explain the organic carbon and pore water profiles. The best fitting was obtained by maintaining the seasonal pattern as observed in the traps, while multiplying the absolute values of the flux by a factor of 1.85. The “inverse modelling” of diagenetic processes resulted in estimates of total mineralisation rate and of degradability of the organic matter at the different stations. These diagenetic model-based estimates are used to constrain the patterns of lateral and vertical transports of organic matter. Using the observed degradability as a function of depth, we show that the observed organic matter fluxes at the different depths are consistent with a model where at all stations along the gradient the same vertical export flux occurs at 200 m, and where organic matter sinks with a constant sinking rate of around 130 m d−1. If sinking rates were higher, in the order of 200 m d−1, the observations could be consistent with an off-slope gradient in export production of approximately a factor of 1.5 between the shallowest and deepest sites. The derived high degradability of the arriving organic matter and the consistency of the mass fluxes at the different stations exclude the possibility of a massive deposition, on the margin, of organic matter produced on the shelf or shelf break. However, other hypotheses to explain the patterns found in the sediment trap data of both OMEX and other continental margin study sites also suffer from different inconsistencies. Further, close examination of the flow patterns at the margin will be needed to examine the question.

Description: Starting with the North Atlantic Bloom Experiment in 1989 oceanographers from 2 variety of countries and scientific disciplines have studied biogeochemical processes in the North Atlantic ocean as a contribution to the Jomt Global Ocean Flux Study (JGOFS). The papers collected in this special issue of Deep-Sea Research Part II are a contribution to the oncjoing international sqnthesis of this decade long effect. In the introduction we give an overview on the major results presented by the individual papers.

Description: An unusually large number of replicated macrofaunal samples (70) was taken from the Western Baltic in May 1995 for a ringtest in an ICES/HELCOM intercalibration exercise. This data set was employed in this study in order to investigate the performance of numerical methods used for predicting species richness and to assess the accuracy of the estimates of abundance and diversity currently used in benthic ecology. The results of this study indicate that: (1) more than 10 replicates are required in order to include in the data set more than two-thirds of the species found in 70 replicates, and more than 53 replicates are required in order to include 95% of the species; (2) estimates of average abundance and of average Shannon-Wiener diversity index using 5 replicates could result in less than 40% error; this could be less than 30% for 10 replicates and less than 5% for 70 replicates; (3) both types of species-richness predictions (jackknife estimate and S 0) increased with increasing number of samples used in the calculations, indicating that their ability to assess overall species richness in the community is rather limited; in particular, it is shown that jackknife overestimates and S0 slightly underestimates species richness. Different configurations of the S0 method were tested in order to optimize its performance, and it was found that both truncation and increasing sampling lag result in increased and stabilized estimates of species richness.

Description: We provide an overview of the role of biological processes in the Benthic boundary layer (BBL) and in sediments on the cycling of particulate organic material in the Goban Spur area (Northeast Atlantic). The benthic fauna, sediment and BBL characteristics were studied along a transect ranging from 208 to 4460 m water depth in different seasons over 3 years. Near-bottom flow velocities are high at the upper part of the slope (1000–1500 m), and high numbers of filter-feeding taxa are found there such that organic carbon normally passing this area during high flow conditions is probably trapped, accumulated, and/or remineralised by the fauna. Overall metabolism in shelf and upper slope sediments is dominated by the macrofauna. More than half of the organic matter flux is respired by macrofauna, with a lower contribution of metazoan meiofauna (4%) and anoxic and suboxic bacterial mineralisation (21%); the remainder (23%) being channelled through nanobiota and oxic bacteria. By its feeding activity and movement, the macrofauna intensely reworks the sediments on the shelf and upper slope. Mixing intensity of bulk sediment and of organic matter are of comparable magnitude. The benthos of the lower slope and abyssal depth is dominated by the microbiota, both in terms of total biomass (〉90%) and carbon respiration (about 80%). The macrofauna (16%), meiofauna (4%) and megafauna (0.5%) only marginally contribute to total carbon respiration at depths below 1400 m. Because large animals have a lower share in total metabolism, mixing of organic matter within the sediments is reduced by a factor of 5, whereas mixing of bulk sediment is one to two orders of magnitude lower than on the shelf. The food quality of organic matter in the sediments in the shallowest part of the Goban Spur transect is significantly higher than in sediments in the deeper parts. The residence time of mineralisable carbon is about 120 d on the shelf and compares well with the residence time of the biota. In the deepest station, the mean residence time of mineralisable carbon is more than 3000 d, an order of magnitude higher than that of biotic biomass.

Description: Bottom-tethered sediment traps deployed in the deep eastern North Atlantic between 54°N 20°W and 33°N 20°W (L1, L2, L3), at the European continental margin at 49°N (OMEX) and off the Canary Islands (ESTOC) were investigated for the determination of 230Th trapping efficiencies. The ratios of 230Th flux measured in the traps (Fa) to the expected 230Th flux from the production rate of 230Th in the overlying water column (Fp) ranged between 0.09 and 1.26. For the traps with deployment periods 〉300 days the interannual variation of Fa/Fp ratios (different years but same location and water depth) were up to 10%, suggesting that the average 230Th flux to the sediment traps did not vary significantly. The influence of lateral advection on the 230Th flux was taken into account either by applying a mass balance of 230Th and 231Pa or by assuming a constant removal rate of 230Th from the water column, an assumption based on similar 230Th concentration-depth profiles observed at most locations investigated. 230Th trapping efficiencies were between 9 and 143%, showing a trend of increasing efficiencies with increasing water depth. No relation was found between current velocities and 230Th trapping efficiencies. Our investigations suggest that the observation of constant or even increasing particle flux rates with increasing water depths in several sediment trap arrays investigated may be a result of sediment trap biases. The correction for the trapping biases is important for the understanding of the regional differences in the particle flux in the eastern North Atlantic.

Description: A geochemical model of the Peru Basin deep-sea floor, based on an extensive set of field data as well as on numerical simulations, is presented. The model takes into account the vertical oscillations of the redox zonation that occur in response to both long-term (glacial/interglacial) and short-term (El Niño Southern Oscillation (ENSO) time scale) variations in the depositional flux of organic matter. Field evidence of reaction between the pore water NO3− and an oxidizable fraction of the structural Fe(II) in the clay mineral content of the deep-sea sediments is provided. The conditions of formation and destruction of reactive clay Fe(II) layers in the sea floor are defined, whereby a new paleo-redox proxy is established. Transitional NO3− profile shapes are explained by periodic contractions and expansions of the oxic zone (ocean bottom respiration) on the ENSO time scale. The near-surface oscillations of the oxic–suboxic boundary constitute a redox pump mechanism of major importance with respect to diagenetic trace metal enrichments and manganese nodule formation, which may account for the particularly high nodule growth rates in this ocean basin. These conditions are due to the similar depth ranges of both the O2 penetration in the sea floor and the bioturbated high reactivity surface layer (HRSL), all against the background of ENSO-related large variations in depositional Corg flux. Removal of the HRSL in the course of deep-sea mining would result in a massive expansion of the oxic surface layer and, thus, the shut down of the near-surface redox pump for centuries, which is demonstrated by numerical modeling.

Description: Grazing experiments were conducted with natural mesozooplankton from Kiel Bight, Germany, using radioactive labelled phytoplankton cultures and seston size fractions. The results of experiments using phytoplankton cultures indicated that bivalve veligers performed highest clearance of particles within a size range of 4.7 to 6.3 µm, whereas optimum particle size for copepods was 15 µm. The results of experiments using labelled natural seston size fractions identified bivalve veligers and appendicularians as those responsible for the removal of particles within the smallest size class (〈2 µm). Seston size fractions larger than 5 µm were mainly cleared by copepods and nauplii. As particle size increased, the contribution of copepod clearance to total zooplankton clearance within size classes increased from 57% (〈5 µm size class) to more than 81% (30 to 100 µm size class). When the nauplii clearance rates were included, the total copepod clearance accounted for 90 to 97.6% of the total volume cleared of particles bigger than 10 µm. Despite low abundances of bivalve veligers and appendicularians in Kiel Bight at the time of the experiment, we calculated that approximately 10 and 8.5%, respectively, of the carbon ingested by total mesozooplankton was due to veliger and appendicularian grazing. The importance of bivalve veligers might be seen in their grazing on seston particles that escape predation by copepods and on the amount of energy that is therefore directed from the water column to the benthos when larvae settle.

Description: Destructive macroalgal mass blooms threaten estuarine and coastal ecosystems worldwide. We asked which factors regulate macroalgal bloom intensity, distribution and species composition. In field experiments in the Baltic Sea, we analyzed the relative effects of nutrients, herbivores and algal propagule banks on population development and dominance patterns in two co-occurring bloom-forming macroalgae, Enteromorpha intestinalis and Pilayella littoralis. Both species were highly affected by the combined effects of a propagule bank, herbivory and nutrients. The magnitude of effects varied with season. The propagule bank was an important overwintering mechanism for both algae, and allowed for recruitment two months earlier than recruitment via freshly dispersed propagules. This provided a seasonal escape from intense herbivory and nutrient limitation later in the year. Favored by massive recruitment from the propagule bank, Enteromorpha was the superior space occupier in early spring, thereby reducing recruitment of Pilayella. Elimination of the propagule bank and recruitment via freshly dispersed propagules favored Pilayella. Strong and selective herbivory on Enteromorpha supported Pilayella in the presence, but not in the absence of the propagule bank. Nutrient enrichment in summer counteracted herbivore pressure on Enteromorpha, thereby negatively affecting Pilayella. Herbivore and nutrient effects were more pronounced for early life stages than adult algae. These results show that recruitment processes and forces affecting early life stages at the beginning of the vegetation period determine development and dominance patterns of macroalgal blooms. Herbivores naturally suppress blooms but increasing nutrient enrichment can override this important control mechanism. The propagule bank plays a previously unrecognized role for population and community dynamics.

Description: Spatial and temporal variability in environmental factors can exert major influences on survival and growth of living organisms. However, in many key areas of fisheries science (e.g. growth, survival and recruitment determination), environmental heterogeneity is usually ignored because of insufficient environmental or fisheries data or lack of evidence that such heterogeneity impacts response variables. For the eastern Baltic Sea (ICES Subdivisions 25 to 32), we evaluated spatial and temporal differences in conditions affecting the survival of cod Gadus morhua L. eggs at survival on four distinct spawning sites within the assessment area. We intercalibrated ways of quantifying the volume of water ('reproductive volume') at each site where salinity, oxygen and temperature conditions permitted successful egg development. We have developed and compared a time series (1952 to 1996) of reproductive volumes among the areas to identify spatial differences. The results of 2 independent volume-estimation methods are comparable, indicating that highly significant differences exist among the sites, and that the westernmost spawning ground, Bornholm Basin, has on average the highest reproductive volume and the lowest variability among the 4 sites. These findings may be useful in evaluating how spatial and temporal variability in environmental conditions affect egg hatching success and possibly recruitment in the Baltic stock.

Description: Throughout the last 2 decades a shift from a cod- to a sprat-dominated system occurred in the upper trophic levels of the Central Baltic Sea. This was caused by a decline in the cod stock, due to recruitment failure and high fishing intensity, resulting in a decrease in predation pressure on sprat. Concurrently with the lowest cod stock size on record, sprat reached biomass values of above 2 × 106 t in 1992, being relatively stable afterwards. Besides predation mortality through cod and in recent years also an increasing fishing pressure, cannibalism on eggs may be a compensatory process limiting the reproductive success of sprat and hence contributing to the population regulation in the Central Baltic. Based on sprat stomach sampling on 21 cruises between March 1988 and July 1996 cannibalism on sprat eggs was investigated in the Bornholm Basin, one of the main spawning areas of Central Baltic sprat. Using a model of gastric evacuation to estimate daily food intake rates and a Virtual Population/Extended Survivor Analysis for computing predator population sizes, egg cannibalism rates were estimated. These were compared to egg abundance data from ichthyoplankton surveys and to preliminary estimates of seasonal egg productions. The study revealed significant interannual differences in the intensity of sprat egg cannibalism with considerable predation in 1990 to 1992 (〉15% of the egg abundance during peak spawning and 〉60% of the seasonal production) and a reduction in most recent years (〈16% of the corresponding abundance and production). As a possible reason for these differences a combination of changes in the vertical overlap of predator/prey and variability in the food environment were identified. Shortcomings of the applied methods and the possible impact of cannibalism on the reproductive success and population development of sprat in the Central Baltic Sea are discussed.

Description: Nanoplankton and picoplankton abundance and community grazing on picoplankton were determined in summer and autumn at several stations in a productive coastal environment (Georges Bank, NW Atlantic Ocean) and in an oligotrophic oceanic ecosystem (Sargasso Sea). Ranges of heterotrophic nanoplankton (HNAN) abundance were 1.2 to 3.6 x 103 cells ml-1 on Georges Bank, and 2.2 to 6.8 x 102 cells ml-1 in the Sargasso Sea. Ranges of phototrophic nanoplankton (PNAN) abundance in these ecosystems were 1.9 to 6.0 x 103 and 1.3 to 4.7 x 102, respectively. Mixotrophic nanoplankton (MNAN), operationally defined here as chloroplast-bearing nanoplankton that ingested fluorescent tracers, comprised an average of 12 to 17% of PNAN in surface waters in both environments during August and October. Mixotrophs at specific stations constituted as much as 38% of total PNAN abundance on Georges Bank and 30% in the Sargasso Sea. Mixotrophs represented up to 39% of the total phagotrophic nanoplankton abundance (MNAN/[MNAN + HNAN]). Community grazing impact was estimated from the disappearance of fluorescent prey surrogates (fluorescently labeled bacteria, FLB; cyanobacteria, FLC; and 〈\3 µm algae, FLA). Absolute grazing rates (total picoplankton cells removed d-1) on Georges Bank exceeded those in the Sargasso Sea due to the greater abundances of predators and prey. However, there was overlap in the specific grazing losses at the 2 sites (ranges = 0.08 to 0.38 d-1 in the coastal ocean and 0.05 to 0.24 d-1 in the oligotrophic ocean). Rates of bacterivory were in approximate balance with rates of bacterial production (3H-thymidine uptake), but production exceeded bacterivory on Georges Bank during the summer cruise. These data are among the first documenting the impact of grazing on picoplankton in these environments, and they are consistent with the prediction that nanoplanktonic protists are major predators of picoplankton. While the proportion of phototrophs that are phagotrophic was highly variable, our study indicates that algal mixotrophy is widespread in the marine environment, occurring in both coastal and oligotrophic sites, and should be considered quantitatively in microbial food web investigations.

Description: During the large-scale deep-sea programme BIGSET in situ measurements of sediment community oxygen consumption (SCOC) were carried out during three cruises between 1995–1998 at five abyssal sites (3190–4450 m water depth) in the deep Arabian Sea in order to elucidate the regional and temporal variation of benthic carbon remineralisation. SCOC ranged from 0.9–6.3 mmol O2 m−2 d−1, with highest values in the western and northern Arabian Sea and lowest values in the southern Arabian Sea. For the central Arabian Sea intermediate oxygen uptake rates were detected. This regional pattern mirrors the overall regional pattern of primary productivity in surface waters and vertical particulate organic carbon (POC) flux at 1000 mab. Primary productivity in Arabian Sea surface waters and particulate flux into the deep-sea are controlled by the monsoon system and the flux maxima during the SW and NE monsoon are among the highest particle fluxes recorded in the deep open ocean. Highest flux rates were recorded in the western and northern Arabian Sea and decreased towards the central and southern Arabian Sea. SCOC at our western, northern and eastern Arabian Sea stations WAST, NAST and EAST were considerably higher than so far detected in other abyssal areas of the global oceans, and vertical POC flux can account for only 20–50% of benthic carbon remineralisation (BCR). Possible explanations for the high rates of BCR at these stations that are situated close to the continental margins are discussed: the accelerated deposition of very labile organic matter due to eolian dust particles, enhanced rain efficiencies, and lateral advection. A significant temporal variability in SCOC only could be detected at the eastern and western Arabian Sea stations WAST and EAST.

Description: This study tested whether the extreme scarcity of larger nanophytoplankton and microphytoplankton in the Gulf of Aqaba and in the open northern Red Sea is caused by nutrient limitation or by selective removal by grazers. Samples of near surface phytoplankton were incubated on board under a fully factorial combination of release from grazing pressure and release from nutrient stress. Release from grazing pressure by different size classes was obtained by sieving through 100, 20, and 10 µm size mesh screens. Release from nutrient stress was obtained by enrichment of Si alone and a full enrichment by N, P, Si and trace elements. Growth rates of most phytoplankton taxa showed a strong, positive response to the full nutrient enrichment and a weaker, but significant response to grazer exclusion. Several diatom taxa showed a weak positive response to Si enrichment. Thus, bottom-up control of medium-sized algae appears to be more important than top-down control.

Description: Increased nutrient loading favors macroalgal blooms in eutrophied coastal ecosystems. The main counteracting factor on this bottom-up support is top-down control by consumers. We asked (1) whether herbivore control on 2 bloom-forming macroalgae in the Baltic Sea varies between different algal life stages, (2) whether herbivores selectively feed on Enteromorpha spp. (Chlorophyceae) thereby supporting dominance of Pilayella littoralis (Phaeophyceae), and (3) whether various herbivore species differ in their effects. In comparative field and laboratory experiments, we analyzed herbivore pressure and selectivity on germling density and adult thalli of Enteromorpha spp. and P. littoralis. In the field, herbivores reduced macroalgal recruitment by 80% within 14 d indicating strong herbivore control at early life stages. Recruits of Enteromorpha spp. were significantly preferred over P. littoralis Adult thalli of both algae showed similar growth rates, but grazing rates were significantly higher on Enteromorpha spp. In laboratory experiments, Idotea chelipes (Isopoda), Littorina saxatilis (Gastropoda) and Gammarus locusta (Amphipoda) strongly reduced germling density, whereas effects of L. littorea were weak. I. chelipes and L. saxatilis significantly preferred germlings of Enteromorpha spp. over P. littoralis. I. chelipes had strong effects on adult Enteromorpha spp. but not on P. littoralis. The effects of G. locusta and L. littorea on both adult algae were smaller and not selective, and L. saxatilis did not feed on adults at all. Different herbivore feeding modes are discussed. We conclude that strong and selective herbivory on juvenile and adult Enteromorpha spp. favors dominance of P. littoralis in the Baltic. An effective control of macroalgal blooms in eutrophied areas can be supported by sustaining high herbivore density and species richness considering the variable and complementary effects of different herbivore species on different algal life stages.

Description: Experiments were carried out on Georges Bank, a productive coastal region in the northwestern sector of the North Atlantic Ocean, and in the oligotrophic western Sargasso Sea to examine the effects of nutrient (inorganic nitrogen and phosphorus) and organic carbon (glucose) additions on bacterial and phytoplankton growth. Four experiments were conducted in each environment. Phytoplankton growth was monitored over a 36 h period by following changes in the concentration of chlorophyll in unfiltered seawater and in seawater prefiltered through 5 μm screening to reduce grazing pressure. Bacterial production was estimated initially and after 24 h using the 3H-thymidine (TdR) method in unfiltered seawater and in 1 μm filtrate. Phytoplankton biomass increased significantly in response to nutrient additions in all but 1 experiment, whereas chlorophyll concentrations remained unchanged or decreased in all of the unamended (control) treatments or treatments supplemented with glucose. Responses of the phytoplankton community were similar for the 〈5 μm and unfiltered treatments. Bacterial production increased after 24 h in all of the treatments on Georges Bank, and there was little effect of nutrient or glucose addition in unfiltered seawater relative to unamended controls. However, glucose addition to the 〈1 μm filtrate caused substantial increases in bacterial production relative to controls and N/P-amended treatments in 2 of the experiments from this environment. Glucose had no stimulatory effect (relative to unamended treatments) in 3 of the 4 Sargasso Sea experiments, and only a marginal effect in the fourth. However, the addition of inorganic nitrogen and phosphorus in the latter ecosystem resulted in higher bacterial production (relative to unamended treatments or glucose addition) in 2 of the experiments with unfiltered seawater, and very large increases in 3 of the experiments with 1 μm filtrate. The magnitude of the changes in bacterial production differed greatly between unfiltered and filtered seawater in both ecosystems, indicating an important role for bacterial grazers in controlling bacterial population growth. The results of this study indicate different nutritional restraints on bacterial production in these contrasting environments.

Description: As one of the elements for a model on the food requirements of Humboldt penguins Spheniscus humboldti we determined, via gas respirometry, metabolic rates while swimming and resting in water. During rest in water at 19°C Humboldt penguins (mean body mass 3.6 kg) required 5.95 W kg-1. This corresponds to a thermal conductance in water of 0.2975 W (kg °C)-1 (at Ta 19°C and assuming a Tb of 39°C). When swimming in a 20 m long channel, metabolism rose from 8 W kg-1 at a speed of 0.6 ms-1 to 23.1 W kg-1 at 2.2 m s-1. Transport costs (the cost to move 1 kg of body mass over a distance of 1 m) reached a minimum at 1.4 ms-1 with 8.1 J (kg m)-1, which corresponds to 0.89 J (Nm)-1. We corrected for acceleration and deceleration in the channel to determine transport costs of free-ranging Humboldt penguins travelling at sea, which were calculated as 7 J (kg m)-1 (0.71 J [Nm]-1), at 1.7 m s-1. Birds feeding chicks need to balance the costs of either (1) returning to the breeding island for the night and travelling back to the feeding grounds in the morning or (2) incurring increased thermoregulatory costs associated with resting at sea overnight. Simple calculations show that at water temperatures of 19°C we expect Humboldt penguins to show a tendency to remain at sea overnight if foraging areas are 〉4 km from their island. In colder waters (12°C), this distance increases to 〉9 km. Using previously published data on at-sea activity of Humboldt penguins, we found that foraging costs during chick rearing amount to 340 g anchovies d-1. Finally, we present a general model to convert Humboldt penguin activity data at sea to food requirements.

Description: The large temporal and spatial variability in carbon isotope fractionation of marine phytoplankton (ε p) is thought to reflect differences in environmental conditions. Meaningful interpretation of this variability requires an understanding of the processes responsible for phytoplankton isotope fractionation. While numerous factors have been suggested to potentially influence ε p, recent theoretical and experimental evidence has emphasized the primary role of phytoplankton growth rate (µ) and CO2 concentration ([CO2aq]) in controlling ε p. Experimental examination of the relationship of ε p with µ and [CO2aq] in studies using different experimental approaches, however, has yielded inconsistent results. Here we directly compare new and previously published data on ε p as a function of CO2 concentration and growth rate for the marine diatom Phaeodactylum tricornutum. When grown under nitrogen-deficient conditions (nitrate-limited chemostat), ε p of P. tricornutum decreases with increasing growth rate. In contrast, under N-replete conditions ε p values are considerably lower at comparable growth rates and CO2 concentrations and are largely insensitive to a 3-fold increase in growth rate due to increasing photon flux density. In both experimental approaches, ε p shows a relatively small CO2 sensitivity in the range of CO2 concentrations naturally occurring in the ocean (8 to 25 µmol kg-1). Below ca 5 µmol CO2 kg-1, a strong decline in ε p with decreasing [CO2aq] is observed. The apparent difference in ε p responses between nitrate-limited and light-controlled cultures of P. tricornutum suggests a principal difference in carbon acquisition for different growth-rate-limiting resources. A mechanistic explanation is proposed and potential implications for the interpretation of phytoplankton carbon isotope fractionation are discussed.

Description: Benthic fluxes and pore-water compositions of silicic acid, nitrate and phosphate were investigated for surface sediments of the abyssal Arabian Sea during four cruises (1995-1998). Five sites located in the northern (NAST), western (WAST), central (CAST), eastern (EAST), and southern (SAST) Arabian Sea were revisited during intermonsoonal periods after the NE- and SW-Monsoon. At these sites, benthic fluxes of remineralized nutrients from the sediment to the bottom water of 36-106, 102-350 and 4-16 mmol m-2 yr-1 were measured for nitrate, silicic acid and phosphate, respectively. The benthic fluxes and pore-water compositions showed a distinct regional pattern. Highest fluxes were observed in the western and northern region of the Arabian Sea, whereas decreasing fluxes were derived towards the southeast. At WAST, the general temporal pattern of primary production, related to the NE- and SW-Monsoon, is reflected by benthic fluxes. In contrast, at sites NAST, SAST, CAST, and EAST a temporal pattern of fluxes in response to the monsoon is not obvious. Our results reveal a clear coupling between the general regional pattern of production in surface waters and the response of the benthic environment, as indicated by the flux of remineralized nutrients, though a spatially differing degree of decoupling during transport and remineralization of particulate organic matter and biogenic opal was observed. This has to be taken into account regarding budget calculations and paleoceanographic topics.

Description: In the Baltic Sea, rye tested how short nutrient pulses of different lengths and frequencies affect macroalgae, epiphytes, grazers and their interactions. We hypothesized that even small-scale variations in nutrient supply may have significant impacts by favoring fast-growing epiphytes which can cause large-scale declines of canopy-forming macroalgae. In a factorial field experiment single plants of the canopy-forming macroalga Fucus vesiculosus with and without epiphytes were exposed to pulses of elevated nutrients (N and P) over 25 d. Five 1 h pulses given every 5 d had no significant effects. A single 5 h pulse increased the epiphyte load but not F. vesiculosus growth rate. In contrast, increasing epiphyte load caused F. vesiculosus growth rate to decline and attracted higher densities of gastropod grazers. These results indicate that a single nutrient pulse can have rapid direct and indirect effects on macroalgae and their associated epiphytes and grazers. Temporal variability of nutrient supply (five 1 h vs one 5 h pulse) plays a significant role in determining the response of primary producers and consumers to elevated nutrients.

Description: Effects of monsoon-induced enhanced depositional regimes of particulate organic carbon (POC) on regional variability and distribution patterns and size spectra of metazoan meiofauna, particularly of nematodes, were investigated at five sites 3158–4414 m deep in the Arabian Sea. The sampling sites were subjected to different flux rates of POC. Total meiofaunal abundance ranged from 109 to 320 ind./10 cm2. Nematodes were the numerically most abundant taxon, with a relative abundance of 82.5–88.7%, followed by copepods and ostracods. Mean individual nematode biomass ranged from 0.0272 to 0.1033 μgC, and Mean nematode population biomass varied between 0.0026 and 0.0133 mgC/10 cm2. Mean nematode lengths ranged from 614.2 to 832.6 μm. The length distributions of nematodes at the different sites were typically skewed with the distributions extending into the longer size classes. At the sites with higher POC deposition rates, nematodes displayed deeper distributions in the sediment column (47.4–58.5% of nematodes in the top 1 cm layer of the sediment) in contrast to very shallow distributions at a site of low POC flux (75.1% of nematodes in the top 1 cm of the sediment). Regional variability of nematode biomass, size and vertical distribution was related to monsoon-driven gradients of POC- and chlorophyll a (chl. a) flux rates and bacterial biomass i.e. bioavailable organic carbon. This was in contrast to nematode abundance which did not correlate significantly with any of these environmental parameters. The differential pattern between biomass and abundance, distribution might be related to POC-dependent alterations in the species composition of the nematode assemblages at the different sites. The hypothesis of increased meiobenthic stocks due to monsoon-induced enhanced sedimentation could not be confirmed compared to data from other less productive oceanic regions. Nematode abundance and biomass in the Arabian Sea were similar to values obtained from the abyssal temperate NE-Atlantic.

Description: Generalisations on the combined effects of consumers and resources on autotrophs in aquatic food webs largely rely on freshwater studies. In this study, we tested these general concepts with marine benthic microalgae, which are important components of coastal food webs. We manipulated nitrogen availability and herbivore presence in a factorial field experiment in the Western Baltic Sea. Moreover, we investigated how herbivore control varied among 3 sites and 2 seasons and tested for trophic cascades by enhancing demersal fish density at 2 sites. Nitrogen availability and herbivore presence had strong and antagonistic effects on microalgal biomass, species composition and diversity. Herbivores significantly reduced algal biomass, whereas nutrient enrichment led to an increase in biomass. Herbivore effects on microalgal biomass increased with increasing nitrogen availability, indicating a functional response of herbivores to nutrient enrichment. The response of microalgae at the species level suggested a trade-off between nutrient use and grazing resistance which appeared to be linked to algal growth form. Compared to other growth forms, large erect species were most responsive to both nitrogen loading and herbivory. Grazing reduced microalgal diversity at low nutrient supply, but enhanced it at high nutrient supply. Herbivore effects varied considerably among different sites and were stronger in spring than in summer. Manipulations of fish density during summer did not have any effects on microalgal community structure. In conclusion, our results demonstrate that herbivores and nutrients have strong and balancing effects on marine microbenthic community structure.

Description: Squids typically demonstrate considerable plasticity in individual growth rates. However, it is not known if individuals growing at different rates also differ at lower levels of organisation. We wished to determine if Sepioteuthis lessoniana individuals that were larger than predicted for their age differed in their digestive gland and mantle muscle tissue proximal composition or mantle muscle structure, compared with individuals that were smaller for their age than predicted. The residual, the difference between the observed size-at-age and that predicted by the growth equation, was used as a measure of the difference in an individual's lifetime growth from the population average. Individual squid varied considerably in their size-at-age, with juveniles showing less variation than adults. Juveniles had greater concentrations of lipid in their muscle tissue, perhaps due to an emphasis on storing energy reserves in this critical period of their life. Differences in biochemical constituents in both the digestive gland and muscle tissue were not related to the size-at-age of individuals, despite biochemical make-up being the lowest organisational level of growth. This may be due to whole animal growth and changes in biochemical composition occurring on different time scales. There was no relationship between the size-at-age of individuals and average mantle muscle fibre size. A strong relationship, however, existed between the size of mantle muscle blocks and the size-at-age of individuals for both juvenile and adult individuals, suggesting that larger muscle blocks are related to both body size and faster individual growth rates. This study demonstrates a clear relationship between mantle muscle structure and growth and the size-at-age of S. lessoniana individuals.

Description: Multiple-choice feeding experiments were performed with the isopod Idotea granulosa and the amphipod Gammarus locusta as consumers. In a first experiment, 2 different types of tissues of the brown seaweed Fucus vesiculosus and its main macroepiphytes, Ulva lactuca and Elachista fucicola, were offered. I. granulosa rejected apices of F. vesiculosus and preferred E. fucicola, while G. locusta clearly preferred F. vesiculosus tissue, especially the meristematic apices. In a second experiment, F. vesiculosus tissue with and without E. fucicola was offered together. For I. granulosa, the consumption of F. vesiculosus was enhanced by the presence of the epiphyte, while for G. locusta there was no difference in consumed F. vesiculosus mass. G. locusta, however, showed behavioural rejection of E. fucicola, and thus, the epiphyte acted as Œprotective coating¹. We conclude that host (F. vesiculosus) tissue could be Œco-consumed¹ by mesograzers (I. granulosa) that were attracted by the presence of epiphytes and that these epiphytes therefore may have a 2-fold negative effect on the host (i.e. competion for light, nutrients etc. and attraction of consumers). ŒCo-consumption¹ and Œprotective coating¹ add 2 more facets to the very variable and case-dependent interrelationships of mesograzer-epiphyte-host systems; their relevance in nature, however, remains to be demonstrated.

Description: As part of the large-scale, interdisciplinary deep-sea study “BIGSET”, the relationship between the monsoon-induced regional and temporal variability of POC deposition and the small-sized benthic community was investigated at several sites 2316–4420 m deep in the Arabian Sea during four cruises between 1995 and 1998. Vertical and horizontal distribution patterns of chloroplastic pigments (a measure of phytodetritus deposition), readily soluble protein and activity, and biomass parameters of the small-sized benthic community (Electron Transport System Activity (ETSA); bacterial ectoenzymatic activity (FDA turnover) and DNA concentrations) were measured concurrently with the vertical fluxes of POC and chloroplastic pigments. Sediment chlorophyll a (chl. a) profiles were used to calculate chl. a flux rates and to estimate POC flux across the sediment water interface using two different transport reaction models. These estimates were compared with corresponding flux rates determined in sediment traps. Regional variability of primary productivity and POC deposition at the deep-sea floor creates a trophic gradient in the Arabian Basin from the NW to the SE, which is primarily related to the activity of monsoon winds and processes associated with the topography of the Arabian Basin and the vicinity of land masses. Inventories of sediment chloroplastic pigments closely corresponded to this trophic gradient. For ETSA, FDA and DNA, however, no clear coupling was found, although stations WAST (western Arabian Sea) and NAST (northern Arabian Sea) were characterised by high concentrations and activities. These parameters exhibited high spatial and temporal variability, making it impossible to recognise clear mechanisms controlling temporal and spatial community patterns of the small-sized benthic biota. Nevertheless, the entire Arabian Basin was recognised as being affected by monsoonal activity. Comparison of two different transport reaction models indicates that labile chl. a buried in deeper sediment layers may escape rapid degradation in Arabian deep-sea sediments.

Description: Albatrosses have among the most remarkable travelling capacities of any extant animal. However, previous studies regarding their movements at sea have mainly focused on breeding birds commuting between the nest site and offshore feeding grounds. In this study, we compare the movement patterns and at-sea activity of breeding and inter-breeding black-browed albatrosses Diomedea melanophris from the Falkland Islands. Data were recorded via global location and activity sensors for 26 incubating birds [during single foraging trips lasting 6.8 d on average) and 6 inter-breeding individuals (during non-stop offshore journeys of 127.5 d on average). Our results showed that foraging black-browed albatrosses utilise vast offshore areas (the average foraging area was 102000 +_ 132 000 km2 by incubating birds and 1552 000 * 970 000 km2 by inter-breeding birds). However, mean forag~ngr ange was similar in both groups (691 * 330 km and 680 t 192 km by incubating and interbreeding birds, respectively) as were their core foraging areas and their at-sea activity patterns. Our results thus indicate that black-browed albatrosses from the Falkland Islands, which represent the largest albatross population world-wide (ca 800 000 individuals), mainly rely on marine resources available within the Patagonian Shelf area. Although this hghly productive continental shelf is the largest of the Southern Hemisphere, rapid development of industrial fisheries currently results in the removal of over 1.4 million tonnes of fish and squid per year in this zone. As our data also show significant spatio-temporal overlap between human and albatross fishing activities within the Patagonian Shelf, we anticipate major detrimental effects on the albatross population in terms of competition for food and additional mortality caused by bird bycatch.