ALBERT

Description: The exploitation of marine resources has caused drastic declines of many large predatory fishes. Amongst these, sharks are of major conservation concern due to their high vulnerability to overfishing and their ecological role as top predators. The 2 protected and endangered shark species tope Galeorhinus galeus and smooth hammerhead Sphyrna zygaena use overlapping coastal areas around the globe as essential fish habitats, but data to assess their trophic ecology and niche partitioning are scarce. We provide the first comparative assessment of the trophic ecology, ontogenetic shifts, and niche partitioning of the co-occurring tope and juvenile smooth hammerhead around the Azores Islands, mid-north Atlantic, based on delta 13C, delta 15N, and delta 34S (CNS) stable isotope analysis of muscle tissue of the sharks and their putative prey species. Overall, isotopic niches of both species indicated a reliance on similar resources throughout the sampled sizes (tope: 35-190; smooth hammerhead 54-159 cm total length), with significant ontogenetic shifts. Topes displayed a gradual shift to higher trophic levels and a more generalist diet with increasing size (increasing delta 15N values and isotopic niche volumes, respectively), whereas smooth hammerhead diet shifted towards prey with lower delta 34S at a constant trophic level and a more specialized diet than tope of comparable body size (decreasing delta 34S and constant delta 15N and delta 13C values, respectively). Our results indicate contrasting ontogenetic shifts in delta 13C and delta 34S along with pronounced differences between niche overlap of life stages pointing to intra- and interspecific niche partitioning of habitat and prey.

Description: Spatiotemporal observations are data rich and offer insights into links between ecological patterns and underlying processes. We present fine-scale autonomous observations from repeated ferry transects in the Strait of Georgia (British Columbia, Canada) during the 2020 spring bloom period using a FerryBox system (temperature, salinity, chlorophyll a fluorescence) and a digital inline holographic microscope. Despite instrument cleaning interruptions related to COVID-19 restrictions, 3 periods from late winter (February) to springtime (March and April) contained 14 days of high-quality holograms (〉70 000) capturing 〉10 500 identifiable micro- to mesoplankton using automatic object detection. The ferry set-up provided automatic data storage through Ocean Networks Canada, which also automatized data flagging and guaranteed remote access. The highest-quality holograms repeatedly covered the central and eastern Strait and showed aspects of bloom succession. Fast-growing diatoms (Skeletonema sp.) emerged first, followed by a diverse assemblage including Chaetoceros spp., Ditylum spp., and Eucampia spp., and by April, larger centric cells prevailed. The combined approach captured local suppression of chlorophyll a fluorescence and diatom concentrations in Fraser River plume waters during the freshet, suggesting fine-scale spatial patterns in seasonal planktonic community composition. This work is among the first of its kind to autonomously generate in situ imaging and physicochemical data with spatiotemporal resolution.

Description: Abundance, biomass and respiration rates of dominant medium- to larger-sized copepod species (ML class) from the upwelling system off Peru (8.5-16°S) were determined along with their carbon ingestion and egestion rates. Small copepods (S class) were included for comparisons of community rates. Overall, abundance/biomass was highest in the upper 50 m and decreased with depth and thus also community ingestion and egestion. Ingestion of the ML class (0-50 m) in shelf regions (14-515 mg C m -2 d -1 ) was lower in the south compared to the north and central study areas, while their offshore ingestion (11-502 mg C m -2 d -1 ) was comparable across regions (8.5-16°S). Ingestion rates (0-50 m) of the S class were in a range similar to those of the ML class in shelf regions (100-417 mg C m -2 d -1 ) but were higher offshore (177-932 mg C m -2 d -1 ). Calanus chilensis and the S class contributed most to total ingestion in the north, while in the south, Centropages brachiatus had the highest community ingestion aside from the S class. Egestion varied from 3-155 mg C m -2 d -1 for the ML class and 30-280 mg C m -2 d -1 for the S class. The high community rates highlight the crucial role of both size classes for carbon budgets in the northern Humboldt Current System off Peru and indicate that the ML class may enhance passive vertical carbon flux, whereas the S class may support carbon remineralization rates in surface waters.

Description: The Humboldt Upwelling System (HUS) supports high levels of primary production and has the largest single-stock fishery worldwide. The high fish production is suggested to be related to high trophic transfer efficiency in the HUS. Mucous-mesh grazers (pelagic tunicates and gastropods) are mostly of low nutritious value and might reduce trophic transfer efficiency when they are locally abundant. Unfortunately, little is known about the spatial dynamics of mucous-mesh grazers from Peruvian waters, limiting our understanding of their potential ecological role(s). We provide a spatial assessment of mucous-mesh grazer abundance from the Peruvian shelf in austral summer 2018/2019 along six cross-shelf transects spanning from 8.5 to 16° S latitude. The community was dominated by appendicularians and doliolids. Salps occurred in high abundance but infrequently and pelagic gastropods were mostly restricted to the North. At low latitudes, the abundance of mucous-mesh grazers was higher than some key species of crustacean mesozooplankton. Transects in this region had stronger Ekman-transport, higher temperature, lower surface turbidity and a broader oxygenated upper water layer compared to higher-latitude transects. Small-scale lateral intrusions of upwelled water were potentially associated with high abundances of doliolids at specific stations. The high abundance and estimated ingestion rates of mucous-mesh grazers in the northern HUS suggest that a large flux of carbon from lower trophic levels is shunted to tunicates in recently upwelled water masses. The data provide important information on the ecology of mucous mesh grazers and stress the relevance to increase research effort on investigating their functioning in upwelling systems.

Description: The blue mussel (Mytilus species complex) is an important ecosystem engineer, and salinity can be a major abiotic driver of mussel functioning in coastal ecosystems. However, little is known about the interactive effects of abiotic drivers and trematode infection. This study investigated the combined effects of salinity and Himasthla elongata and Renicola roscovita metacercarial infections on the filtration capacity, growth, and condition of M. edulis from the Baltic Sea. In a laboratory experiment, groups of infected and uninfected mussels were exposed to a wide range of salinities (6−30, in steps of 3) for 1 mo. Shell growth was found to be positively correlated with salinity and optimal at 18−24 at the end of the experiment, imposed by constraints in shell calcification under lower salinities. Mussel shell growth was not affected by H. elongata infection. While salinity had only a minor effect on tissue dry weight, infected mussels had a significantly lower tissue dry weight than uninfected mussels. Most interestingly, the combination of salinity and trematode infections negatively affected the mussels’ condition indices at lower salinity levels (6 and 9), suggesting that trematode infections are more detrimental to mussels when combined with freshening. A significant positive effect of salinity on mussel filtration was found, with an initial optimum at salinity 18 shifting to 18−24 by the end of the experiment. These findings indicate that salinity and parasite infections act as synergistic stressors for mussels, and enhance the understanding of potential future ecosystem shifts under climate change-induced freshening in coastal waters.

Description: In the deep sea, benthic communities largely depend on organic material from the overlying water column for food. The remains of organisms on the seafloor (food falls) create areas of organic enrichment that attract scavengers. The scavenging rates and communities of food falls of medium-sized squid, fish and jellyfish (1-100 cm) are poorly known. To test our hypothesis that scavenging responses are specific for different food falls, we deployed camera landers baited with squid, jellyfish and fish for 9 to 25 h at 1360 to 1440 m in the southern Norwegian Sea. Image analysis of 8 deployments showed rapid food fall consumption (20.3 +/- 1.4 [SD] to 31.6 +/- 3.7 g h(-1)) by an amphipod-dominated scavenging community that was significantly different between the food fall types. Fish and squid carcasses were mostly attended by amphipods of the genus Eurythenes. Smaller unidentified amphipods dominated the jellyfish experiments together with brittle stars (cf. Ophiocten gracilis) and decapod shrimps (cf. Bythocaris spp.); the latter only occurred on jellyfish carcasses. The removal time for jellyfish (similar to 17 h) was almost twice as long as that for squid and fish (9-10 h). The maximum scavenger abundance was significantly higher on fish carcasses than on jellyfish and squid. The times at which abundances peaked were similar for jellyfish and fish (after 8-9 h) but significantly sooner for squid (3.00 +/- 0.35 h). Our results, although based on a small number of experiments, demonstrate differences in scavenging responses between food fall species, suggesting tight coupling between the diversity and ecology of benthic scavenging communities in the Norwegian Sea.

Description: In this work, we focused on the functional characterization of unicellular eukaryotic assemblages that had previously been taxonomically characterized by 18S rRNA gene amplicon sequencing in a eutrophic coastal site with marked plankton blooms. Biological traits of different functional groups were assigned to the retrieved operational taxonomic units (OTUs). The traits included size, trophic strategy, the presence of spines, mucilage production, colony formation, motility, spore formation, and potential harmfulness. Functional diversity indices were calculated and compared to analogous taxonomic diversity indices, indicating a strong positive coupling of richness and dominance and a negative coupling of evenness, even at a low taxonomic resolution (at the family/genus/species level). Biological trait trade-offs and co-occurrences of specific traits were evident during the succession of plankton blooms. The trophic strategy dominating in the assemblages frequently alternated between autotrophy, mixotrophy, and a few recorded cases of parasitism. Given that there was no indication of nutrient limitation, we suggest that biotic pressures force marine eukaryotes to exploit narrow niches by adopting specific strategies/traits that favour their survival. These traits act by increasing resource acquisition potential and via predator avoidance. This leads to a unique succession of blooms in the system, characterized by adaptations of the bloom taxa that are a direct response to the preceding assemblage.

Description: Recent advancements in telemetry have redefined our ability to quantify the fine-scale movements of aquatic animals and derive a mechanistic understanding of movement behaviours. The VEMCO Positioning System (VPS) is a fine-scale commercial positioning system used to generate highly accurate semi-continuous animal tracks. To date, VPS has been used to study 86 species, spanning 25 taxonomic orders. It has provided fine-scale movement data for critical life stages, from tracking day-old turtle hatchlings on their first foray into the sea to adult fish returning to natal rivers to spawn. These high-resolution tracking data have improved our understanding of the movements of species across environmental gradients within rivers, estuaries and oceans, including species of conservation concern and commercial value. Existing VPS applications range from quantifying spatio-temporal aspects of animal space use and key aspects of ecology, such as rate of movement and resource use, to higher-order processes such as interactions among individuals and species. Analytical approaches have seen a move towards techniques that incorporate error frameworks such as autocorrelated kernel density estimators for home range calculations. VPS technology has the potential to bridge gaps in our fundamental understanding of fine-scale ecological and physiological processes for single and multi-species studies under natural conditions. Through a systematic review of the VPS literature, we focus on 4 principle topics: the diversity of species studied, current ecological and ecophysiological applications and data analysis techniques, and we highlight future frontiers of exploration.

Description: Juvenile sea turtles can disperse thousands of kilometers from nesting beaches to oceanic development habitats, aided by ocean currents. In the North Atlantic, turtles dispersing from American beaches risk being advected out of warm nursery grounds in the North Atlantic Gyre into lethally cold Northern European waters (e.g. around the United Kingdom). We used an ocean model simulation to compare simulated numbers of turtles that were advected to cold waters around the UK with observed numbers of turtles reported in the same area over ~5 decades. Rates of virtual turtles predicted to encounter lethal temperatures (≤10 and 15°C, mean 19% ± 2.7) and reach the UK were consistently low (median 0.83%, lower quartile 0.67%, upper quartile 1.02%), whereas there was high inter-annual variability in the numbers of dead or critically ill turtles reported in the UK. Generalized additive models suggest inter-annual variability in the North Atlantic Oscillation (NAO) index to be a good indicator of annual numbers of turtle strandings reported in the UK. We demonstrate that NAO variability drives variability in the dispersion scenarios of juvenile turtles from key nesting regions into the North Atlantic. Coastal effects, such as the number of storms and mean sea surface temperatures in the UK were significant but weak predictors, with a weak effect on turtle strandings. Further understanding how changing environmental conditions such as NAO variability and storms affect the fate of juvenile turtles is vital for understanding the distribution and population dynamics of sea turtles.

Description: Mothers impact the survival and performance of their offspring through the resources they provision, and the degree of maternal investment in an individual offspring can be broadly estimated by egg size for organisms that lack parental care. Animals may also actively maintain symbiotic partnerships with microorganisms through the germ line, but whether microbes are a fundamental component of maternal provisioning is an untested hypothesis in evolutionary symbiosis. We present a preliminary test of this by comparing the egg-associated microbiota of ten sea urchin species with ecological factors known to influence egg size. We found that the microbiota associated with sea urchin eggs had a phylogenetic signal in both composition and richness, which varied between years but not between individuals or within a clutch. Moreover, we found a negative correlation between microbiome richness and taxonomic dominance, and that community diversity covaried with egg size and energetic content but not with pelagic larval duration or latitude. These data suggest that there are multiple parallels between the ecological factors that govern changes in egg size and microbiome composition and diversity, implying that microbial symbionts may be another constituent potentially provided by the mother.

Description: A large-volume mesocosm-based nutrient perturbation experiment was conducted off the island of Hawai‘I, USA, to investigate the response of surface ocean phytoplankton communities to nutrient addition of macronutrients, trace metals, and vitamins and to assess the feasibility of using mesocosms in the open ocean. Three free-drifting mesocosms (~60 m3) were deployed: one mesocosm served as a control (no nutrient amendments), a second (termed +P) was amended with nitrate (N), silicate (Si), phosphate (P) and a trace metal + vitamin mixture, and a third (termed -P) was amended with N, Si, and a trace metal + vitamin mixture but no P. These mesocosms were unreplicated due to logistical constraints and hence differences between treatments are qualitative. After 6 d, the largest response of the phytoplankton community was observed in the +P mesocosm where chlorophyll a (chl a) and 14C-based primary production were 2–3× greater than the -P mesocosm and 4–6× greater than the control. Comparison between mesocosm and ‘microcosm’ incubations (20 l) revealed differences in the magnitude and timing of production and marked differences in community structure with a reduced response of diatoms in microcosm treatments. Notably, we also observed pronounced declines in Prochlorococcus populations in all treatments: although these were greater in microcosms (up to 99%). Overall, this study confirmed the feasibility of deploying free-drifting mesocosms in the open ocean as a potentially powerful tool to investigate ecological impacts of nutrient perturbations and constitutes a valuable first step towards scaling plankton manipulation experiments.

Description: The trophic ecology of mixotrophic, zooxanthellate jellyfishes potentially spans a wide spectrum between autotrophy and heterotrophy. However, their degree of trophic plasticity along this spectrum is not well known. To better characterize their trophic ecology, we sampled the zooxanthellate medusa Mastigias papua in contrasting environments and sizes in Palau (Micronesia). We characterized their trophic ecology using isotopic (bulk δ13C and δ15N), elemental (C:N ratios), and fatty acid compositions. The different trophic indicators were correlated or anti-correlated as expected (Pearson’s correlation coefficient, rP 〉 0.5 or 〈 -0.5 in 91.1% of cases, p 〈 0.05), indicating good agreement. The sampled M. papua were ordered in a trophic spectrum between autotrophy and heterotrophy (supported by decreasing δ13C, C:N, proportion of neutral lipid fatty acids (NLFA:TLFA), n-3:n-6 and increasing δ15N, eicosapentaenoic acid to docosahexaenoic acid ratio (EPA:DHA)). This trophic spectrum was mostly driven by sampling location with little influence of medusa size. Moreover, previous observations have shown that in a given location, the trophic ecology of M. papua can change over time. Thus, the positions on the trophic spectrum of the populations sampled here are not fixed, suggesting high trophic plasticity in M. papua. The heterotrophic end of the trophic spectrum was occupied by non-symbiotic M. papua, whereas the literature indicates that the autotrophic end of the spectrum corresponds to dominant autotrophy, where more than 100% of the carbon requirement is obtained by photosynthesis. Such high trophic plasticity has critical implications for the trophic ecology and blooming ability of zooxanthellate jellyfishes

Description: The Asian shore crab Hemigrapsus takanoi, native to the northwest Pacific Ocean, was recently discovered in Kiel Fjord (southwestern Baltic Sea). In laboratory experiments, we tested the salinity tolerance of H. takanoi across 8 levels (0 to 35) and across 3 life history stages (larvae, juveniles and adults) to assess its potential to invade the brackish Baltic Sea. Larval development at different salinities was monitored from hatching to the megalopa stage, while survival and feeding of juveniles and adults were assessed over 17 d. Larvae of H. takanoi were able to complete their development to megalopa at salinities 〉= 20 and the time needed after hatch to reach this stage did not differ between salinities of 20, 25, 30 and 35. At a salinity of 15, larvae still reached the last zoea stage (zoea V), but development to the megalopa stage was not completed. All juveniles and adults survived at salinities from 5 to 35. Feeding rates of juveniles increased with increasing salinity across the entire salinity range. However, feeding rates of adults reached their maximum between salinities of 15 and 35. Our results indicate that both juveniles and adults of H. takanoi are euryhaline and can tolerate a wide range of salinities, at least for the time period tested (2 wk). However, larval development was impaired at salinities lower than 20, which may prevent the spread of H. takanoi into the Baltic Proper.

Description: Understanding the underlying ecological factors that affect the distribution patterns of organisms is vital for their conservation. Cephalopods such as giant warty squids Moroteuthopsis longimana are important in the diets of marine predators, including grey-headed albatrosses Thalassarche chrysostoma, yet our understanding of their habitat and trophic ecology remains limited. We investigated the habitat and trophic niche utilised by M. longimana through the delta C-13 and delta N-15 profiles captured in their beaks. M. longimana beaks were collected around grey-headed albatross nests at the Prince Edward Islands during 2004 and 2013 (n = 40 beaks). The results showed distinctly Antarctic distributions (delta C-13 = -24.0 +/- 1.0 parts per thousand, mean +/- SD) for M. longimana, consistent with albatrosses foraging at the Southwest Indian Ridge, as opposed to broader foraging zones utilised by albatrosses from Iles Crozet and Iles Kerguelen. Slightly lower delta N-15 values (5.4 +/- 0.7 parts per thousand) were found compared to other islands in the Indian Sector of the Southern Ocean, which may indicate more crustaceans in the squid diets. Sequential sampling along the lateral walls of individual beaks (n = 4) revealed ontogenetic shifts in delta C-13 and delta N-15 values, but individual variation in these shifts requires further investigation.

Description: The maraena whitefish Coregonus maraena is a threatened anadromous species in the North Sea, which in the past was decimated to near extinction. Since the late 1980s, several re-establishment programs have been implemented in rivers draining into the North Sea, but the scientific basis for sustainable conservation measures is often lacking, since little is known about the biology of this species. In this study, otolith microchemistry of fish ranging from 24.6 to 58.4 cm in total length (median 31.3 cm, SD 8.4 cm) was used to characterize the migration behavior of a reintroduced population of maraena whitefish from the River Elbe, Germany. Our analyses revealed the presence of 3 different migration patterns: (1) one-time migration into high-salinity habitat (North Sea) within the first year of life (29.6%), (2) multiple migrations between lowland high-salinity habitats starting in the first year of life (14.8%) and (3) permanent residency within low-salinity habitats, a pattern displayed by the majority (55.6%) of sampled individuals. Not only do these results reveal differential migration behavior, but they also indicate that permanent river residency is common in the River Elbe population of C. maraena. The role of the Elbe as both a feeding and a spawning habitat should thus be considered more explicitly in current conservation measures to support recovery of this species.

Description: Cold‐water coral (CWC) reefs and mounds are and have been biodiversity hotspots of the deep sea. As their occurrence depends on specific environmental parameters, gaining hindsight on changing ocean conditions under on‐going climate change is the key to a better understanding of CWC mound development through time. A convenient technique for reconstructing the palaeoenvironment during periods of CWC mound growth is by extracting geochemical proxies from biologically mediated carbonates. Here, the focus is on probably the two most abundant calcareous archives, that are, cold‐water Scleractinia and Foraminifera, with an overview of the geochemical proxies (selection) used in these aragonitic and calcitic skeletons from CWC mounds. A particular emphasis is set on constraining proxies for temperature, salinity, seawater density, seawater carbonate systems parameters (pH, CO 3 2− ), nutrients, oxygen and water mass tracers.

Description: Understanding the behavioural ecology of endangered taxa can inform conservation strategies. The activity budgets of the loggerhead turtle Caretta caretta are still poorly understood because many tracking methods show only horizontal displacement and ignore dives and associated behaviours. However, time-depth recorders have enabled researchers to identify flat, U-shaped dives (or type 1a dives) and these are conventionally labelled as resting dives on the seabed because they involve no vertical displacement of the animal. Video- and acceleration-based studies have demonstrated this is not always true. Focusing on sea turtles nesting on the Cabo Verde archipelago, we describe a new metric derived from magnetometer data, absolute angular velocity, that integrates indices of angular rotation in the horizontal plane to infer activity. Using this metric, we evaluated the variation in putative resting behaviours during the bottom phase of type 1a dives for 5 individuals over 13 to 17 d at sea during a single inter-nesting interval (over 75 turtle d in total). We defined absolute resting within the bottom phase of type 1a dives as periods with no discernible acceleration or angular movement. Whilst absolute resting constituted a significant proportion of each turtle’s time budget for this 1a dive type, turtles allocated 16−38% of their bottom time to activity, with many dives being episodic, comprised of intermittent bouts of rest and rotational activity. This implies that previously considered resting behaviours are complex and need to be accounted for in energy budgets, particularly since energy budgets may impact conservation strategies. © The authors 2021. Open Access under Creative Commons by Attribution Licence. Use, distribution and reproduction are unrestricted. Authors and original publication must be credited

Description: Cape anchovy Engraulis encrasicolus is an ecologically and economically important pelagic fish species occurring along the coast of South Africa. A recent eastward shift in Cape anchovy distribution indicates that environmental conditions are becoming more favorable for the species on the east coast. This shift is particularly important in the sheltered Algoa Bay region, a nursery area for fish larvae. However, the relatively low productivity of the Agulhas Current Large Marine Ecosystem on the eastern coast of South Africa may result in an anchovy population in poorer nutritional condition and with slower growth rates than the west coast population. Using otolith and nucleic acid analyses, the growth rates of anchovy larvae from the western and southeastern coasts of South Africa were compared. The otolith analysis results indicated that, at any given age, individual growth rates for anchovy larvae were higher on the southeast coast than on the west coast. The RNA:DNA values also indicated that instantaneous growth rates of anchovy larvae were higher in Algoa Bay than on the west coast. At the time of sampling, chlorophyll and zooplankton productivity were higher at sampling sites in Algoa Bay than sites on the west coast, potentially due to favorable oceanographic features in the bay. As such, the results suggest that Algoa Bay is a suitable and potentially favorable nursery area for the early stages of anchovy, highlighting the importance of separate management of the southeast coast region in a changing world.

Description: A mesocosm approach was used to investigate the effects of ocean acidification (OA) on a natural plankton community in coastal waters off Norway by manipulating CO2 partial pressure ( pCO2). Eight enclosures were deployed in the Raunefjord near Bergen. Treatment levels were ambient (~320 µatm) and elevated pCO2 (~2000 µatm), each in 4 replicate enclosures. The experiment lasted for 53 d in May-June 2015. To assess impacts of OA on the plankton community, phytoplankton and protozooplankton biomass and total seston fatty acid content were analyzed. In both treatments, the plankton community was dominated by the dinoflagellate Ceratium longipes. In the elevated pCO2 treatment, however, biomass of this species as well as that of other dinoflagellates was strongly negatively affected. At the end of the experiment, total dinoflagellate biomass was 4-fold higher in the control group than under elevated pCO2 conditions. In a size comparison of C. longipes, cell size in the high pCO2 treatment was significantly larger. The ratio of polyunsaturated fatty acids to saturated fatty acids of seston decreased at high pCO2. In particular, the concentration of docosahexaenoic acid (C 22:6n3c), essential for development and reproduction of metazoans, was less than half at high pCO2 compared to ambient pCO2. Thus, elevated pCO2 led to a deterioration in the quality and quantity of food in a natural plankton community, with potential consequences for the transfer of matter and energy to higher trophic levels

Description: We investigated trace element stoichiometries of the nitrogen-fixing marine cyanobacterium Crocosphaera subtropica ATCC51142 under steady-state growth conditions. We utilized exponentially fed batch cultures and varied iron (Fe) concentrations to establish nutrient limitation in C. subtropica growing at a constant growth rate (0.11 d -1 ). No statistical difference in cell density, chlorophyll a , particulate organic carbon (C), nitrogen (N) and phosphorus (P) were observed between consecutive days after Day 14, and cultures were assumed to be at steady state with respect to growth for the remaining 11 d of the experiment. Cultures were limited by P in the highest Fe treatment (41 nmol l -1 ) and by Fe in the 2 lower-concentration Fe treatments (1 and 5 nmol l -1 ). Cell size and in vivo fluorescence changed throughout the experiment in the 1 nmol l -1 Fe treatment, suggesting ongoing acclimation of C. subtropica to our lowest Fe supply. Nevertheless, Fe:C ratios were not significantly different between the Fe treatments, and we calculated an average (±SD) Fe:C ratio of 32 ± 14 µmol mol -1 for growth at 0.11 d -1 . Steady-state P-limited cells had lower P quotas, whilst Fe-limited cells had higher manganese (Mn) and cobalt (Co) quotas. We attribute the increase in Mn and Co quotas at low Fe to a competitive effect resulting from changes in the supply ratio of trace elements. Such an effect has implications for variability in elemental stoichiometry in marine phytoplankton, and potential consequences for trace metal uptake and cycling in marine systems.

Description: Precipitation extremes with devastating socioeconomic consequences within the South American Monsoon System (SAMS) are expected to become more frequent in the near future. The complexity in SAMS behavior, however, poses severe challenges for reliable future projections. Thus, robust paleomonsoon records are needed to constrain the high spatiotemporal variability in the response of SAMS rainfall to different climatic drivers. This study uses Ti/Ca ratios from X‐ray fluorescence scanning of a sediment core retrieved off eastern Brazilian to trace precipitation changes over the past 322 Kyr. The results indicate that despite the spatiotemporal complexity of the SAMS, insolation forcing is the primary pacemaker of variations in the monsoonal system. Additional modulation by atmospheric p CO 2 suggests that SAMS intensity over eastern Brazil will be suppressed by rising CO 2 emissions in the future. Lastly, our record reveals an unprecedented strong and persistent wet period during Marine Isotope Stage 6 driven by anomalously strong trade winds.

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.