ALBERT

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: 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: 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: 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: 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 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: 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: 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: 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: 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: 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: 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: 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: Brown skuas Catharacta antarctica lonnbergi breed across a broad latitudinal range from the Antarctic to temperate regions. While information on the non-breeding distribution and behaviour for Antarctic and subantarctic populations is known, no data exist for populations breeding at temperate latitudes. We combined geolocation sensing and stable isotope analysis of feather tissue to study the non-breeding behaviour of brown skuas from the temperate Chatham Islands, a population that was historically thought to be resident year-round. Analysis of 27 non-breeding tracks across 2 winters revealed that skuas left the colony for a mean duration of 146 d, which is 64% of the duration reported for Antarctic and subantarctic populations from King George Island, South Shetland Islands, and Bird Island, South Georgia. Consistent with populations of brown skuas from Antarctica and the Subantarctic, the distribution was throughout mixed subtropical-subantarctic and shelf waters. Stable isotope analysis of 72 feathers suggests that moulting takes place over mixed subtropical-subantarctic and subtropical shelf waters. We conclude that brown skuas from the Chatham Islands are migratory, but the year-round mild environmental conditions may reduce the necessity to leave their territories for extended periods.

Description: In the northern hemisphere, eelgrass Zostera marina L. is the most important and widespread seagrass species. Despite its ecological importance, baseline data on eelgrass distribution and abundance are mostly absent, particularly in subtidal areas with relatively turbid waters. Here, we report a combined approach of vegetation mapping in the Baltic Sea coupled to a species distribution model (SDM). Eelgrass cover was mapped continuously in the summers of 2010 and 2011 with an underwater towed camera along ~400 km of seafloor. Eelgrass populated 80% of the study region and occurred at water depths between 0.6 and 7.6 m at sheltered to moderately exposed coasts. Mean patch length was 128.6 m but was higher at sheltered locations, with a maximum of 〉2000 m. The video observations (n = 7824) were used as empiric input to the SDMs. Using generalized additive models, 3 predictor variables (depth, wave exposure, and slope), which were selected based on Akaike’s information criterion, were sufficient to predict eelgrass presence/absence. Along with a very good overall discriminative ability (area under the receiver-operating characteristic curve ROC/AUC = 0.82), depth (as a proxy for light), wave exposure, and slope contributed 66, 29, and 5%, respectively, to the final model. The estimated total areal extent of eelgrass in the study region amounts to 140.5 km2 and comprises about 11.5% of all known Baltic seagrass beds. The present work is, to the best of our knowledge, the largest study undertaken to date on vegetation mapping and the first to assess distribution of eelgrass quantitatively in the western Baltic Sea.

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: The nature of many microbe-host interactions is not static, but may shift along a continuum from mutualistic to harmful depending on the environmental conditions. In this study, we assessed the interaction between the foundation plant eelgrass Zostera marina and the frequently associated protist Labyrinthula zosterae. We tested how an important environmental factor, nutrient availability, would modulate their interaction. We experimentally infected naive eelgrass plants in combination with 2 nutrient levels (fertilized and non-fertilized). We followed L. zosterae infection, eelgrass growth parameters and host defense gene expression over 3 wk in large 600 l tanks. Inoculation with L. zosterae and nutrient limitation both reduced eelgrass growth. These effects were additive, whereas no interaction of nutrient treatment and L. zosterae inoculation was detected. Gene expression levels of 15 candidate genes revealed a reduced expression of photosynthesis-related genes but an increased expression of classical stress genes such as Hsp80 in inoculated plants 2 d post-inoculation. However, we found no effects on plant mortality, and plants were able to clear high infection levels within 3 wk to ambient background levels of infection as assessed via specific RT-qPCR designed to quantify endophytic L. zosterae. Thus, we found no evidence that L. zosterae is a facultative mutualist that facilitates eelgrass growth under nutrient-limiting conditions. We suggest that the interaction between contemporary L. zosterae genotypes and Z. marina represents a mild form of parasitism in northern Europe because the damage to the plant is moderate even under nutrient limitation stress.

Description: In fisheries, vulnerability assessments - also commonly known as ecological risk assessments (ERAs) -have been an increasingly popular alternative to stock assessments to evaluate the vulnerability of non-target species in resource- and data-limited settings. The widely-used productivity-susceptibility analysis (PSA) requires detailed species-specific biological information and fishery susceptibility for a large number of parameters to produce a relative vulnerability score. The two major disadvantages of PSA are that each species is assessed against an arbitrary reference point, and PSA cannot quantify cumulative impacts of multiple fisheries. This paper introduces an Ecological Assessment of the Sustainable Impacts of Fisheries (EASI-Fish), a flexible approach that quantifies the cumulative impacts of fisheries on data-limited bycatch species, demonstrated in eastern Pacific Ocean (EPO) tuna fisheries. The method first estimates fishing mortality (F) based on the 'volumetric overlap' of each fishery with the distribution of each species. F is then used in length-structured per-recruit models to assess population vulnerability status using conventional biological reference points. Model results were validated by comparison with stock assessments for bigeye and yellowfin tunas in the EPO for 2016. Application of the model to 24 species of epipelagic and mesopelagic teleosts, sharks, rays, sea turtles and cetaceans and identification of the most vulnerable species is demonstrated. With increasing demands on fisheries to demonstrate ecological sustainability, EASI-Fish allows fishery managers to more confidently identify vulnerable species to which resources can be directed to either implement mitigation measures or collect further data for more formal stock assessment.

Description: Oxygen (O2) deficiency and nutrient concentrations in marine systems are impacting organisms from microbes to higher trophic levels. In coastal and enclosed seas, O2 deficiency is often related to eutrophication and high degradation rates of organic matter. To investigate the impact of O2 concentration on bacterial growth and the turnover of organic matter, we conducted multifactorial batch experiments with natural microbial communities of the central Baltic Sea. Water was collected from suboxic (〈5 µmol L -1) depths in the Gotland Basin during June 2015. Samples were kept for four days under fully oxygenated and low O2 conditions (mean: 34 µmol L-1 O2), with or without nutrient (ammonium, phosphate, nitrate) and labile carbon (glucose) amendments. We measured bacterial abundance, bacterial heterotrophic production, extracellular enzyme rates (leucine-aminopeptidase) and changes in dissolved and particulate organic carbon concentrations. Our results show that the bacterial turnover of organic matter was limited by nutrients under both oxic and low O2 conditions. In nutrient and glucose replete treatments, low O2 concentrations significantly reduced the net uptake of dissolved organic carbon and lead to higher accumulation of more labile dissolved organic matter. Our results therewith suggest that the combined effects of eutrophication and deoxygenation on heterotrophic bacterial activity may potentially favor the accumulation of dissolved organic carbon in the Baltic Sea.

Description: Statoliths of the gonatid squid Gonatus fabricii from Disko Bay, West Greenland, were analysed by laser ablation inductively coupled plasma mass-spectrometry (LA-ICP-MS) to determine the concentrations and spatial distributions of 9 minor and trace elements (Sr, Ba, Mg, U, Zn, Mn, Y, Zr, Na). Element composition was assigned in situ to distinct statolith regions, corresponding to ontogenetic stages of an individual squid’s life. Significant variations in concentrations of all measured elements except Na were found between different regions of the statolith. Variations in Ba/Ca ratios suggest that juveniles inhabit surface waters, while larger specimens move to deeper waters. U/Ca and Sr/Ca ratios increased towards the outer statolith region, suggesting migration of adult squid into colder waters. Mg/Ca ratios decreased progressively from the core to the edge, most likely related to changes in protein concentrations in their microstructure during ontogenesis. This study is one of the first to apply LA-ICP-MS to cephalopod statoliths. Our results emphasise the strong potential of spatially resolved statolith analyses to gather information on life history, migrations and habitat use of cephalopods.

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 North Pacific Ocean population of the neon flying squid Ommastrephes bartramii, which undertakes seasonal north–south migrations, consists of autumn and winter–spring spawning cohorts. We examined life history differences between the 2 cohorts in relation to the oceanographic environment. The differences could be explained by seasonal north–south movements of the following 2 oceanographic zones: (1) the optimum spawning zone defined by sea surface temperatures; and (2) the food-rich zone defined by the position of the transition zone chlorophyll front (TZCF). The 2 cohorts use the food-rich zone in different phases of their life cycles. The spawning grounds for the autumn cohort occur within the subtropical frontal zone (STFZ), characterized by enhanced productivity in winter due to its proximity to the TZCF, whereas the spawning grounds for the winter–spring cohort occur within the subtropical domain, which is less productive. As the TZCF shifts northward in spring, the autumn cohort continues to occur in the productive area north of the TZCF, whereas the winter–spring cohort remains in the less productive area to the south until it migrates into productive waters north of the TZCF in the summer or autumn. Consequently, the autumn cohort grows faster than the winter–spring cohort during the first half of its life cycle, whereas the winter–spring cohort grows faster during the second half. This growth pattern may be responsible for differing migration patterns; males of the autumn cohort do not have to migrate given their early fast growth in the STFZ, whereas those of the winter–spring cohort must migrate to the food-rich subarctic frontal zone to compensate for their slow growth. These biological and ecological differences between the 2 cohorts suggest flexibility of their life history response to oceanographic environments.

Description: The cirrate octopod Stauroteuthis syrtensis is a mesopelagic species commonly collected in the North Atlantic. Individuals were observed at depths 〉600 m and typically within 100 m of the bottom in three ~900 m deep canyons indenting the southern edge of Georges Bank. When first sighted, most octopods were floating passively with their webbed arms gathered into a small ball. When disturbed, they expanded their webs to form a ‘balloon’ shape, swam slowly by sculling their fins, pulsed their webs like medusae and, in some cases, streamlined their arms and webs and moved away smoothly by rapidly sculling their fins. The bodies of 9 octopods comprised 92 to 95% water, with tissue containing 9 to 22% carbon (C) and 2 to 4% nitrogen (N). These values were similar to those reported for medusae and ctenophores. Oxygen (O2) consumption rates of 4.6 to 25.8 µmol O2 g–1 C h–1 were within ranges reported for medusae, ctenophores, and deep-water cephalopods. The stomachs of S. syrtensis, dissected immediately after capture, contained only the calanoid copepod Calanus finmarchicus. Calculations indicated that S. syrtensis need 1.3 to 30.1 ind. d–1 of C. finmarchicus to meet their measured metabolic demand. Excretion rates (0.3 to 12.4 µg NH4+ g–1 C h–1 and 0.06 to 4.83 µg PO43– g–1 C h–1) were at least an order of magnitude lower than rates reported for other octopods or gelatinous zooplankters. O:N ratios (11 to 366) suggested that S. syrtensis catabolized lipids, which may be supplied by C. finmarchicus. Vertical distribution, relatively torpid behavior and low metabolic rates characterized S. syrtensis as a benthopelagic and relatively passive predator on copepods.

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: Mesopelagic fishes occur in all the world’s oceans, but their abundance and consequently their ecological significance remains uncertain. The current global estimate based on net sampling prior to 1980 suggests a global abundance of one gigatonne (109 t) wet weight. Here we report novel evidence of efficient avoidance of such sampling by the most common myctophid fish in the Northern Atlantic, i.e. Benthosema glaciale. We reason that similar avoidance of nets may explain consistently higher acoustic abundance estimates of mesopelagic fish from different parts of the world’s oceans. It appears that mesopelagic fish abundance may be underestimated by one order of magnitude, suggesting that the role of mesopelagic fish in the oceans might need to be revised.

Description: The longfin squid Loligo pealeii is distributed widely in the NW Atlantic and is the target of a major fishery. A previous electrophoretic study of L. pealeii was unable to prove genetic differentiation, and the fishery has been managed as a single unit stock. We tested for population structure using 5 microsatellite loci. In early summer (June), when the squids had migrated inshore to spawn, we distinguished 4 genetically distinct stocks between Delaware and Cape Cod (ca. 490 km); a 5th genetic stock occurred in Nova Scotia and a 6th in the northern Gulf of Mexico. One of the summer inshore stocks did not show genetic differentiation from 2 of the winter offshore populations. We suggest that squids from summer locations overwinter in offshore canyons and that winter offshore fishing may affect multiple stocks of the inshore fishery. In spring, squids may segregate by genetic stock as they undertake their inshore migration, indicating an underlying mechanism of subpopulation recognition.

Description: Pronounced environmental trends across fronts suggest that the otolith chemistry of oceanic fish can resolve zones on either side, promoting application to population questions at similar spatial scales. Trace and minor elements laid down immediately prior to capture - along the edges of otoliths from Patagonian toothfish Dissostichus eleginoides - discriminated frontal zones in the Antarctic Circumpolar Current in the Southwestern Atlantic Ocean. Mean values differentiated sampling areas by up to 2.6 standard deviations, suggesting: (1) otolith Mg/Ca enrichment related to fish activity around the Burdwood Bank; (2) Mn/Ca enrichment associated with South America; (3) Sr/Ca linked to the presence of Circumpolar Deep Water; and (4) Ba/Ca to nutrient production and mixing. In the Polar Frontal Zone, meanders or eddies may account for affinities with neighbouring sampling areas, bringing water from the Subantarctic and Antarctic Zones onto the North Scotia Ridge. Moreover, fish age showed a significant relationship with depth and improved cross-validation by 14%, giving 85% classification rates to South American and Antarctic regions, and 57 to 83% to areas along the Patagonian Shelf. These results indicate that otolith chemistry reflects hydrography, detecting oceanic gradients across the slope of continental shelves and between zones separated by strong trends like fronts.

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

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

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

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

Description: Large Protista of unknown taxonomic affinities are described from 3 species of coleoid squids, and are reported from many other species of cephalopods. The white to yellow-orange, ovoid cyst-like parasites are partially embedded within small pockets on the surface of the gills, often in large numbers. Except for a holdfast region on one side of the large end, the surface of the parasite is elaborated into low triangular plates separated by grooves. The parasites are uninucleate; their cytoplasm bears lipid droplets and presumed paraglycogen granules. Trichocysts, present in a layer beneath the cytoplasmic surface, were found by transmission electron microscopy to be of the dinoflagellate type. Further studies are needed to clarify the taxonomic position of these protists.

Description: A new species of giant unicellular parasite inhabiting the gills of squid hloroteuthis robusta is described from specimens collected from the northeastern Pacific Ocean. Hochbergia moroteuthensis gen. et sp. nov. belongs to a recently reported group of protistans whose taxonomic affinities are as yet unknown. Members of the group parasitize cephalopods and have a characteristic aspect: most of the body wall is raised into close-fitting triangular plates and there is a holdfast region, devoid of plates, by which the parasites attach to the host. H. moroteuthensis is the first species of the group to be named. The general form of the body, the precise number and arrangement of plates, and the morphology of the holdfast reglon are described.

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

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

Description: Global climate change involves an increase in oceanic CO2 concentrations as well as thermal stratification of the water column, thereby reducing nutrient supply from deep to surface waters. Changes in inorganic carbon (C) or nitrogen (N) availability have been shown to affect marine primary production, yet little is known about their interactive effects. To test for these effects, we conducted continuous culture experiments under N limitation and exposed the bloomforming dinoflagellate species Scrippsiella trochoidea and Alexandrium fundyense (formerly A. tamarense) to CO2 partial pressures (pCO(2)) ranging between 250 and 1000 mu atm. Ratios of particulate organic carbon (POC) to organic nitrogen (PON) were elevated under N limitation, but also showed a decreasing trend with increasing pCO(2). PON production rates were highest and affinities for dissolved inorganic N were lowest under elevated pCO(2), and our data thus demonstrate a CO2-dependent trade-off in N assimilation. In A. fundyense, quotas of paralytic shellfish poisoning toxins were lowered under N limitation, but the offset to those obtained under N-replete conditions became smaller with increasing pCO(2). Consequently, cellular toxicity under N limitation was highest under elevated pCO(2). All in all, our observations imply reduced N stress under elevated pCO(2), which we attribute to a reallocation of energy from C to N assimilation as a consequence of lowered costs in C acquisition. Such interactive effects of ocean acidification and nutrient limitation may favor species with adjustable carbon concentrating mechanisms and have consequences for their competitive success in a future ocean.

Description: Heme is the iron-containing prosthetic group of hemoproteins, and is thus required for photosynthesis, respiration and nitrate reduction in marine phytoplankton. Here we report concentrations of heme b in Southern Ocean phytoplankton and contrast our findings with those in coastal species. The concentration of particulate heme b (pmol l-1) observed at the end of the exponential growth phase was related to the concentration of dissolved iron in the culture media. Small Southern Ocean phytoplankton species (〈6 μm in diameter) had heme b quotas 〈1 μmol mol-1 carbon, the lowest yet reported for marine phytoplankton. Heme b was also depleted in these species with respect to chlorophyll a. We calculated the amount of carbon accumulated per mole of heme b per second in our cultures (heme growth efficiency, HGE) and found that small Southern Ocean species can maintain growth rates, even while heme b content is reduced. Small Southern Ocean phytoplankton can thus produce more particulate carbon than larger Southern Ocean or small coastal species at equivalent iron concentrations. Combining primary productivity and heme b concentrations reported for the open ocean, we found that HGE in natural populations was within the range of our laboratory culture results. HGE was also observed to be higher at open ocean stations characterized by low iron concentrations. Our results suggest that low heme b quotas do not necessarily result in reduced growth and that marine phytoplankton can optimize iron use by manipulating the intracellular hemoprotein pool

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

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

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

Description: In the 1930s the wasting disease pathogen Labyrinthula zosterae is believed to have killed 90% of the temperate seagrass Zostera marina in the Atlantic Ocean. Despite the devastating impact of this disease the host–pathogen interaction is still poorly understood, and few field studies have investigated factors correlating with the prevalence and abundance of L. zosterae. This study measures wasting disease in natural populations of Z. marina, showing a strong correlation between the disease and both salinity and water depth. No infection was detected in Z. marina shoots from low salinity (13–25 PSU) meadows, whereas most shoots carried the disease in high salinity (25–29 PSU). Shallow (1 m) living Z. marina shoots were also more infected compared to shoots in deeper (5 m) meadows. In addition, infection and transplantation experiments showed that Z. marina shoots from low salinity meadows with low pathogen pressure were more susceptible to L. zosterae infection. The higher susceptibility could not be explained by lower content of inhibitory defense compounds in the shoots. Instead, extracts from all Z. marina shoots significantly reduced pathogen growth, suggesting that Z. marina contains inhibitory compounds that function as a constitutive defense. Overall, the results show that seagrass wasting disease is common in natural Z. marina populations in the study area and that it increases with salinity and decreases with depth. Our findings also suggest that low salinity areas can act as a refuge against seagrass wasting disease.

Description: The high biodiversity of coral reefs results in complex trophic webs where energy and nutrients are transferred between species through a multitude of pathways. Here, we hypothesize that reef sponges convert the dissolved organic matter released by benthic primary producers (e.g. corals) into particulate detritus that is transferred to sponge-associated detritivores via the sponge loop pathway. To test this hypothesis, we conducted stable isotope (13C and15N) tracer experiments to investigate the uptake and transfer of coral-derived organic matter from the sponges Mycale fistulifera and Negombata magnifica to 2 types of detritivores commonly associated with sponges: ophiuroids (Ophiothrix savignyi and Ophiocoma scolopendrina) and polychaetes (Polydorella smurovi). Findings revealed that the organic matter naturally released by the corals was indeed readily assimilated by both sponges and rapidly released again as sponge detritus. This detritus was subsequently consumed by the detritivores, demonstrating transfer of coral-derived organic matter from sponges to their associated fauna and confirming all steps of the sponge loop. Thus, sponges provide a trophic link between corals and higher trophic levels, thereby acting as key players within reef food webs.

Description: Nitrogen fixation is a key source of nitrogen in the Baltic Sea which counteracts nitrogen loss processes in the deep anoxic basins. Laboratory and field studies have indicated that single-strain nitrogen-fixing (diazotrophic) cyanobacteria from the Baltic Sea are sensitive to ocean acidification and warming, two drivers of marked future change in the marine environment. Here, we enclosed a natural plankton community in twelve indoor mesocosms (volume ~1400 L) and manipulated pCO2 to yield six CO2 treatments with two different temperature treatments (16.6°C and 22.4°C, pCO2 range = 360 – 2030 μatm). We followed the filamentous, heterocystous diazotrophic cyanobacteria community (Nostocales, primarily Nodularia spumigena) over four weeks. Our results indicate that heterocystous diazotrophic cyanobacteria may become less competitive in natural plankton communities under ocean acidification. Elevated CO2 had a negative impact on Nodularia sp. biomass, which was exacerbated by warming. Our results imply that Nodularia sp. may contribute less to new nitrogen inputs in the Baltic Sea in future.

Description: Recent studies on the life history of cephalopods have challenged the paradigm that all coleoid cephalopods have a single reproductive cycle and a short lifespan. Although lifespan has been investigated in several octopod species, few studies have considered their life-history traits in relation to environmental conditions via a comparative approach. We tested the hypothesis that octopod lifespan is correlated with habitat characteristics. For that purpose, life history and environmental data of 25 incirrate octopod species and the vampire squid Vampyroteuthis infernalis were compiled from the literature. Regression analysis showed that the relationship between age at maturity and average habitat temperature was best described by a negative power function (r2 = 0.86). The depth ranges of occurrence (minimum-midpoint-maximum) were positively correlated with time to reach maturity, with maximum depth showing the best fit (r2 = 0.47). Using literature data and our analyses, we estimated that octopods living in polar and deep seas mature after 3 to 5 yr. The reviewed and estimated instantaneous relative growth rates ranged from 0.1% body weight (BW) d-1 in the Antarctic species Pareledone charcoti to nearly 6% BW d-1 in the temperate species Macroctopus maorum. Our analyses suggest that low water temperatures (〈5°C) result in an extended ontogenetic development, potentially as a result of reduced metabolic rates and constraints on protein synthesis, which increases the lifespan of octopods living in cold environments.

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

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

Description: Analysis of stomach contents and predator-prey relations of 168 swordfish from the western North Atlantic Ocean is presented. The predominant food of swordfish is squid (82% by frequency of occurence) followed by fish (53%) consisting primarily of gladis, scombrids, butterfish, bluefish, and sand lance. Average food volume in stomach is 1% of average body weight. Estimates of daily ration range from 0.94% to 1,6% of avergae body weight, with yearly consumption ranging from 3.4 to 5.8 times average body weight per year. The prey biomass consumed by the sowrdfish population in the Georges Bank area is estimated to range from 2,100 to 11,000 mt over a residency time of 5 mo.

Description: Many ecosystems are facing biodiversity loss and environmental change due to anthropogenic activities, with these impacts occurring within the context of natural disturbance. Understanding ecosystem functioning and the response of communities to these impacts is necessary in order to evaluate the effects of future environmental change. The aim of this study was to determine the consequences of the loss of key species on the structure and function of intertidal communities in a context of nutrient enrichment, so as to ascertain the resistance of these communities when disturbance and stresses are compounded. Subarctic rocky intertidal communities in Quebec were subjected to an orthogonal factorial field experiment with 3 stress factors (macroalgae canopy loss, grazer exclusion, and nutrient enrichment), each with 2 disturbance levels. Simple and interactive effects of these factors were followed for 4 mo, and responses in structure (% cover and biomass) and productivity were evaluated. The communities that were not subjected to canopy loss showed greater resistance and very limited effects from enrichment and grazer reduction. The loss of canopy altered the community structure (e.g. reduction in richness and biomass) and functioning (reduced productivity), probably due to increased temperatures and desiccation. This lack of resistance was amplified through the addition of a stress. The application of multiple stresses within field experiments allows for a better understanding of the mechanisms affecting community structure and ecosystem functioning under situations of increased natural and anthropogenic stress.

Description: Ocean acidification has the potential to affect growth and calcification of benthic marine invertebrates, particularly during their early life history. We exposed field-collected juveniles of Asterias rubens from Kiel Fjord (western Baltic Sea) to 3 seawater CO2 partial pressure (pCO2) levels (ranging from around 650 to 3500 µatm) in a long-term (39 wk) and a short-term (6 wk) experiment. In both experiments, survival and calcification were not affected by elevated pCO2. However, feeding rates decreased strongly with increasing pCO2, while aerobic metabolism and NH4+ excretion were not significantly affected by CO2 exposure. Consequently, high pCO2 reduced the scope for growth in A. rubens. Growth rates decreased substantially with increasing pCO2 and were reduced even at pCO2 levels occurring in the habitat today (e.g. during upwelling events). Sea stars were not able to acclimate to higher pCO2, and growth performance did not recover during the long-term experiment. Therefore, the top-down control exerted by this keystone species may be diminished during periods of high environmental pCO2 that already occur occasionally and will be even higher in the future. However, some individuals were able to grow at high rates even at high pCO2, indicating potential for rapid adaption. The selection of adapted specimens of A. rubens in this seasonally acidified habitat may lead to higher CO2 tolerance in adult sea stars of this population compared to the juvenile stage. Future studies need to address the synergistic effects of multiple stressors such as acidification, warming and reduced salinity, which will simultaneously impact the performance of sea stars in this habitat.

Description: Bacterial epiphytes of Gracilaria conferta were quantified. Saprophytic bacteria reached 350 times and agar degraders 25000 times higher numbers g-1 algal wet wt on tissues infected with the 'white tips disease', as compared to healthy tissues. A bacterial inducing agent of the 'white tips disease' was detected. Addition of 10(2) to 10(3) cells of this isolate ml-1 medium led to increased rates of infection. This effect did not occur if the isolate was autoclaved before addition. The virulent bacteria could always be isolated from infected tissues. It frequently, but not always, infected G. conferta and should be regarded as a facultative parasite. Several factors influenced the disease development. Temperatures above 20-degrees-C, in combination with photon flux densities of more than 200 muE m-2 s-1, increased the rate of infection. Relatively low amounts (more than 25 mug ml-1) of certain organic nutrients (peptone and yeast extract) led to strong manifestations of the disease. Addition of agar did not cause any symptoms, while 5 mg l-1 of the antibiotic rifampicin prevented the alga from being infected.

Description: The focus of this study was the 11.55 m long sediment core 303700-7, which was retrieved from the Gdansk Basin during a cruise of RV ‘Poseidon’ within the frame of the Russian–German Project GISEB. The core was analysed for grain size, elemental chemical composition, organic carbon and palynological spectra. The age control was based on palynostratigraphy and 7 radiocarbon datings of bulk sedimentary organic matter. These data provide a high-resolution record of climatic and marine palaeoenvironments in the Gdansk Basin for the last ca. 13 kyr, from the Bølling to late Holocene time. Sedimentation rates were estimated to vary between 0.37 and 1.62 mm yr–1. Major variations in palaeosalinity were estimated from bromine concentrations in the sediment. This method allows the first quantitative reconstruction of palaeosalinity changes in the Baltic Sea, especially profound during the Littorina and Postlittorina periods (middle to late Holocene). In addition, grain size data indicated several Littorina transgressive–regressive stages and a few episodes of increased near-bottom current activity. Our results from the Gdansk Basin are consistent with palaeoceanographic data from other deep basins of the Baltic Sea and provide new insights into the regional Holocene history.

Description: Sprat Sprattus sprattus larvae were used as model organisms to evaluate whether larval lipids reflect in situ feeding conditions and can thus identify match-mismatch situations. In detail, we determined larval lipid content, growth rates based on RNA:DNA ratios, and fatty acid (FA) composition during the spawning season in the Central Baltic Sea, and evaluated these in light of feeding, mortality and recruitment (which were determined in parallel within the project ‘GLOBEC Germany’). Based on the opposing trend of RNA:DNA and lipid content, as well as on previous observations, we hypothesized that lipid content and current feeding conditions are largely uncoupled in the early life stages of sprat due to reduced lipid anabolism. However, lipids still provide information in several ways: (1) segmented generalised linear models proved to be a suitable tool for identifying phases of lipid catabolism during development, with the slope reflecting size-specific environmental starvation pressure. This method detected a previously identified mismatch situation with suitable prey in the early spawning season, which increased mortality of larger larvae. (2) Estimated starvation resistance, a proxy that accounts for temperature- and size-dependent metabolism, reflected the likelihood of near future starvation of individual larvae. (3) Principal component analyses on FAs identified monthly differences in diet composition. Biomarkers indicated a dinoflagellate and/or microbial loop based carbon flux to the larvae. (4) Regression analyses revealed lower docosahexaenoic acid (DHA) levels in spring, but no obvious effect on growth. Food quality was generally high, and its impact on larval survival was less evident than that of prey size suitability.

Description: Bacteria were isolated seasonally from the Mediterranean sponges Chondrilla nucula and Petrosia ficiformis and screened for antibacterial activities. Selected isolates were taxonomically identified by 16S rRNA gene sequencing. A total of 416 different bacterial strains were isolated, 60 (14.4%) of which displayed variable degrees of antimicrobial activity. Of the bioactive strains, 58.3% were able to inhibit Staphylococcus aureus, 6.7% were active against Bacillus subtilis, 11.7% against both Enterococcus faecalis and Escherichia coli, 38.3% against Pseudoalteromonas atlantica and 33.3% against Pseudomonas elongata. 16S rRNA gene sequence analysis showed that 2 isolates, 1 from seawater samples and 1 from P. ficiformis, were most closely related to Bacillus subtilis (99% similarity) and that another isolate from P. ficiformis was most closely related to a previously described sponge-associated Alphaproteobacterium NW001 (98% similarity). Two isolates from C. nucula were most closely related to Brachybacterium paraconglomeratum (99% similarity) and Shewanella algae (89% similarity). The high percentage of bioactive isolates derived from the 2 sponges suggests that marine microorganisms, whether animal-associated or planktonic, are promising sources for drug discovery.

Description: We monitored the dynamics of extracellular organic matter in 3 mesocosms: one dominated by a heterotrophic (microbial) community with negligible autotrophic activity (net heterotrophic system), a second where a small Phaeocystis bloom developed (production and loss almost balanced), and a third harboring a large diatom bloom (net autotrophic system). In all mesocosms, meso- and macroscopic heterotrophic organisms were excluded to primarily study extracellular organic matter production and turnover by specific algae and microbial loop organisms, respectively. Concentration and composition of dissolved organic matter (DOM), i.e. dissolved organic carbon (DOC), monosaccharides and total carbohydrates (MCHO and TCHO), free and combined neutral carbohydrates (DFCHO and DCCHO), as well as free and combined amino acids (DFAA and DCAA) were measured. In addition, net and gross community production rates were determined to calculate C-budgets. Whereas concentrations and composition of MCHO differed very little among the 3 mesocosms, dynamics of TCHO, DFCHO, and DCCHO differed significantly. Concentrations of DFAA were higher in both algal mesocosms compared to the heterotrophic system, and composition of DFAA was significantly different in the Phaeocystis and Diatom tanks. The composition and concentration of DCAA, however, were similar in all 3 mesocosms. Total dissolved carbohydrates and amino acids comprised a substantial fraction of the DOC pool. Dynamics of these DOC fractions, however, could only partly explain those of DOC, implying either that other dissolved compounds were important for overall C-cycling, or that microbial degradation of DOM affects the detection of carbohydrates and protein components.

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

Description: Coccolithophores, a globally distributed group of marine phytoplankton, showed diverse responses to ocean acidification (OA) and to combinations of OA with other environmental factors. While their growth can be enhanced and calcification be hindered by OA under constant indoor light, fluctuation of solar radiation with ultraviolet irradiances might offset such effects. In this study, when a calcifying and a non-calcifying strain of Emiliania huxleyi were grown at 2 CO2 concentrations (low CO2 [LC]: 395 µatm; high CO2 [HC]: 1000 µatm) under different levels of incident solar radiation in the presence of ultraviolet radiation (UVR), HC and increased levels of solar radiation acted synergistically to enhance the growth in the calcifying strain but not in the non-calcifying strain. HC enhanced the particulate organic carbon (POC) and nitrogen (PON) productions in both strains, and this effect was more obvious at high levels of solar radiation. While HC decreased calcification at low solar radiation levels, it did not cause a significant effect at high levels of solar radiation, implying that a sufficient supply of light energy can offset the impact of OA on the calcifying strain. Our data suggest that increased light exposure, which is predicted to happen with shoaling of the upper mixing layer due to progressive warming, could counteract the impact of OA on coccolithophores distributed within this layer.

Description: Nitrification, the step-wise oxidation of ammonium to nitrite and nitrate, is important in the marine environment because it produces nitrate, the most abundant marine dissolved inorganic nitrogen (DIN) component and N-source for phytoplankton and microbes. This study focused on the second step of nitrification, which is carried out by a distinct group of organisms, nitrite-oxidizing bacteria (NOB). The growth of NOB is characterized by nitrite oxidation kinetics, which we investigated for 4 pure cultures of marine NOB (Nitrospina watsonii 347, Nitrospira sp. Ecomares 2.1, Nitrococcus mobilis 231, and Nitrobacter sp. 311). We further compared the kinetics to those of non-marine species because substrate concentrations in marine environments are comparatively low, which likely influences kinetics and highlights the importance of this study. We also determined the isotope effect during nitrite oxidation of a pure culture of Nitrospina (Nitrospina watsonii 347) belonging to one of the most abundant marine NOB genera, and for a Nitrospira strain (Nitrospira sp. Ecomares 2.1). The enzyme kinetics of nitrite oxidation, described by Michaelis-Menten kinetics, of 4 marine genera are rather narrow and fall in the low end of half-saturation constant (Km) values reported so far, which span over 3 orders of magnitude between 9 and 〉1000 µM NO2-. Nitrospina has the lowest Km (19 µM NO2-), followed by Nitrobacter (28 µM NO2-), Nitrospira (54 µM NO2-), and Nitrococcus (120 µM NO2-). The isotope effects during nitrite oxidation by Nitrospina watsonii 347 and Nitrospira sp. Ecomares 2.1 were 9.7 ± 0.8 and 10.2 ± 0.9‰, respectively. This confirms the inverse isotope effect of NOB described in other studies; however, it is at the lower end of reported isotope effects. We speculate that differences in isotope effects reflect distinct nitrite oxidoreductase (NXR) enzyme orientations.

Description: The combined effects of warming and overwintering copepod densities on the spring succession of Baltic Sea plankton were investigated using indoor mesocosms. Three zooplankton (1.5, 4 and 10 copepods L-1) and two temperature levels called ∆0°C and ∆6°C (0°C and 6°C above the present day temperature scenario for Kiel Bight) were chosen. Both, the timing and the duration of the protozooplankton (PZP) bloom were significantly affected by temperature, but not by copepod density. In contrast, the bloom intensity of PZP was highly affected by the factors temperature and copepod density and its interaction. This suggests that at elevated temperature conditions PZP grows faster but, at the same time, are subject to higher top-down control by copepods. At low temperatures and low copepod densities, PZP in turn fully escaped from copepod predation. Further changes in the overwintering copepod densities resulted in a strong ciliate suppression of which small-sized ciliates (〈30 µm) were especially vulnerable to copepod predation while other PZP size classes remained unaffected. In conclusion, the results presented point at a pivotal regulating role of overwintering copepods under future warming condition. Further, warming was shown to cause a distinct match between phytoplankton and PZP thus strengthening trophic pathways through PZP. Our findings are discussed in the context of the ‘trophic link-sink’ debate by considering potential alterations in the flux of matter and energy up the food web.

Description: Within mono-specific meadows of clonal plants, genotypic diversity may functionally replace species diversity. Little is known about the variability in performance and plasticity of different genotypes towards anthropogenically induced stressors. In this field experiment we compared light-limitation stress responses and recovery of different eelgrass Zostera marina genotypes to assess the variability in phenotypic plasticity and gene expression between different genotypes. Replicated monoculture plots of 4 genotypes were subjected to a simulated turbidity period of 4 wk using shading screens, and their performance during light limitation and 4 wk of recovery was compared to non-shaded controls. In addition to growth and biomass, we investigated storage carbohydrates and quantified the expression of genes involved in carbohydrate metabolism, photosynthesis and control of oxidative stress. Plants showed remarkable plasticity in their stress responses and all phenotypic variables recovered to the control level within 4 wk. Depletion and subsequent restoration of sucrose levels differed among genotypes. In terms of gene expression, no consistent patterns were observed. Our study confirms that stress responses and recovery processes can vary substantially between genotypes and the results emphasize the importance of preserving regional genotypic diversity for immediate positive diversity effects and for adaptive evolution in response to global change.

Description: We investigated potential connections over the past 2 decades between mesoscale circulation regimes in the Ionian Sea and newly-observed species and the concurrent rise in sea temperature in the Adriatic Sea. Analyses of plankton samples from 1993 to 2011 in the southern Adriatic revealed marked changes in the non-crustacean zooplankton community. Eleven species were recorded for the first time in the Adriatic, while 3 species reappeared after years of absence. We found that pluriannual changes in the zooplankton community tracked the continuum of circulation regimes in the Northern Ionian Gyre (NIG). The occurrence of Atlantic/Western Mediterranean species coincided with anti-cyclonic circulation in the NIG, probably due to the advection of Modified Atlantic Water into the Adriatic, while the presence of Lessepsian species coincided with the cyclonic pattern, which governs the entry of Eastern Mediterranean waters. The impact has been that newcomers now make a significant contribution to the zooplankton community in the southern Adriatic and, in certain cases, have replaced native species. Our results provide new evidence of the influence of teleconnection processes between the North Atlantic and Eastern Mediterranean on the dynamics of water masses in the southern Adriatic. The synergistic effects of these processes, together with warmer Mediterranean waters, raise concerns over dramatic changes in the marine biodiversity of the Adriatic.

Description: The spatio-temporal origin of surviving juvenile Baltic cod Gadus morhua was investigated by coupling age information from otolith microstructure analysis and hydrodynamic modeling, which allowed backtracking of drift routes in time and space. The suitability of hydrodynamic modeling for drift simulations of early life stages of Baltic cod up to the pelagic juvenile stage was validated by comparing model simulations with the catch distribution from a survey targeting pelagic juveniles, and mortality rates and hatch date distributions of pelagic and demersal juveniles were estimated. Hatch dates and hatch locations of juvenile survivors showed distinct patterns which did not agree well with the abundance and spatial distribution of eggs, suggesting marked spatio-temporal differences in larval survival. The good agreement of the spatio-temporal origin of survivors from this field investigation with previous modeling studies on the survival chances of early-stage larvae and with general spatio-temporal patterns of larval prey availability suggests that differences in survival are related to food availability during the early larval stage. Results are discussed in relation to the recruitment process of Baltic cod, in particular with respect to the critical period and match-mismatch hypotheses, and to possible implications for the placement of a Marine Protected Area which was established to ensure undisturbed spawning of Baltic cod.

Description: Droop’s cell-quota model is the most successful description of phytoplankton growth in laboratory cultures and is increasingly being introduced into the ecosystem components of biogeochemical models. Although the Droop model’s parameters can be easily interpreted in biological terms, it was nevertheless derived empirically and lacks a sound mechanistic foundation. Here we derive Droop’s model from a simple optimality condition which maximises net growth rate. Our approach links the maximum cell quota to the cost of nutrient acquisition and suggests that respiration is influenced more strongly by C fixation than by N assimilation.

Description: The notion that excess phosphorus (P) and high irradiance favour pelagic diazotrophy is difficult to reconcile with diazotroph behaviour in laboratory experiments and also with the observed distribution of N2-fixing Trichodesmium, e.g. in the relatively nitrogen (N)-rich North Atlantic Ocean. Nevertheless, this view currently provides the state-of-the-art framework to understand both past dynamics and future evolution of the oceanic fixed N inventory. In an attempt to provide a consistent theoretical underpinning for marine autotrophic N2 fixation we derive controls of diazotrophy from an optimality-based model that accounts for phytoplankton growth and N2 fixation. Our approach differs from existing work in that conditions favourable for diazotrophy are not prescribed but emerge, indirectly, from trade-offs among energy and cellular resource requirements for the acquisition of P, N, and carbon. Our model reproduces laboratory data for a range of ordinary phytoplankton species and Trichodesmium. The model predicts that (1) the optimal strategy for facultative diazotrophy is switching between N2 fixation and using dissolved inorganic nitrogen (DIN) at a threshold DIN concentration; (2) oligotrophy, especially in P and under high light, favours diazotrophy; (3) diazotrophy is compatible with DIN:DIP supply ratios well above Redfield proportions; and (4) communities of diazotrophs competing with ordinary phytoplankton decouple emerging ambient and supply DIN:DIP ratios. Our model predictions appear in line with major observed patterns of diazotrophy in the ocean. The predicted importance of oligotrophy in P extends the present view of N2 fixation beyond a simple control by excess P in the surface ocean.

Description: Stress often induces metabolically expensive countermeasures. Bivalve shell production is costly and can thus be indirectly impacted by environmental stress. Suboptimal salinity and temperature may constitute stressors that allocate energy away from shell production to cellular processes such as osmoregulation or to the repair of cellular damage. In the course of climate change, water temperatures of the Baltic Sea are predicted to increase, and salinity is predicted to regionally decrease. These shifts may lead to increased stress for temperate marine species adapted to relatively cool water temperatures and high salinity conditions. To better understand the importance of climate change-related stress, we assessed the isolated and interactive effects of salinity and temperature on shell increment (cumulative growth: shell), cellular oxidative stress (accumulation of oxidized lipids and proteins: lipofuscin), instantaneous physiological condition (condition index: CI), and mortality of young Mytilus edulis and Arctica islandica from the western Baltic Sea. Temperature and salinity interactively affected shell increment, lipofuscin accumulation, and mortality of M. edulis as well as shell increment of A. islandica. Shell increment of M. edulis was less affected by hyposalinity than shell increment of A. islandica. In both species the CI decreased and lipofuscin accumulation increased with increasing temperature. Lipofuscin accumulation negatively correlated with shell increment in M. edulis. We conclude that Baltic Sea populations of ecologically relevant bivalve species may experience severe stress by the predicted regional scenario of warming and desalination if evolutionary adaptation does not happen at a similar rate.

Description: Ocean acidification and associated changes in seawater carbonate chemistry negatively influence calcification processes and depress metabolism in many calcifying marine invertebrates. We present data on the cephalopod mollusc Sepia officinalis, an invertebrate that is capable of not only maintaining calcification, but also growth rates and metabolism when exposed to elevated partial pressures of carbon dioxide (pCO2). During a 6 wk period, juvenile S. officinalis maintained calcification under ~4000 and ~6000 ppm CO2, and grew at the same rate with the same gross growth efficiency as did control animals. They gained approximately 4% body mass daily and increased the mass of their calcified cuttlebone by over 500%. We conclude that active cephalopods possess a certain level of pre-adaptation to long-term increments in carbon dioxide levels. Our general understanding of the mechanistic processes that limit calcification must improve before we can begin to predict what effects future ocean acidification will have on calcifying marine invertebrates.

Description: Previous studies have suggested that phytoplankton play an important role in the biogeochemical cycling of iodine, due to the appearance of iodide in the euphotic zone. Changes in the speciation of iodine over the course of the growth cycle were examined in culture media for a variety of phytoplankton taxa (diatoms, dinoflagellates and prymnesiophytes). All species tested showed the apparent ability to reduce iodate to iodide, though production rates varied considerably between species (0.01 to 0.26 nmol l–1 µg–1 chl a d–1), with Eucampia antarctica the least and Pseudo-nitzschia turgiduloides the most efficient iodide producers. Production was found to be species specific and was not related to biomass (indicated by e.g. cell size, cell volume, or chl a content). In all species, except for the mixotrophic dinoflagellate Scrippsiella trochoidea, iodide production commenced in the stationary growth phase and peaked in the senescent phase of the algae, indicating that iodide production is connected to cell senescence. This suggests that iodate reduction results from increased cell permeability, which we hypothesize is due to subsequent reactions of iodate with reduced sulphur species exuded from the cell. A shift from senescence back to the exponential growth phase resulted in a decline in iodide and indicated that phytoplankton-mediated oxidation of iodide to iodate was likely to be occurring. Iodide production could not be observed in healthy cells kept in the dark for short periods. Bacterial processes appeared to play only a minor role in the reduction of iodate to iodide.

Description: Effects of elevated temperature on the formation and subsequent degradation of diatom aggregates were studied in a laboratory experiment with a natural plankton community from the Kiel Fjord (Baltic Sea). Aggregates were derived from diatom blooms that developed in indoor mesocosms at 2.5 and 8.5 degrees C, corresponding to the 1993 to 2002 mean winter in situ temperature of the Western Baltic Sea and the projected sea surface temperature during winter in 2100, respectively. Formation and degradation of diatom aggregates at these 2 temperatures in the dark were promoted with roller tanks over a period of 11 d. Comparison of the 2 temperature settings revealed an enhanced aggregation potential of diatom cells at elevated temperature, which was likely induced by an increased concentration of transparent exopolymer particles (TEP). The enhanced aggregation potential led to a significantly higher proportion of particulate organic matter in aggregates at 8.5 degrees C. Moreover, the elevated temperature favoured the growth of bacteria, bacterial biomass production, and the activities of sugar- and protein-degrading extracellular enzymes in aggregates. Stimulating effects of rising temperature on growth and metabolism of the bacterial community resulted in an earlier onset of aggregate degradation and silica dissolution. Remineralization of carbon in aggregates at elevated temperature was partially compensated by the formation of carbon-rich TEP during dark incubation. Hence, our results suggest that increasing temperature will affect both formation and degradation of diatom aggregates. We conclude that the vertical export of organic matter through aggregates may change in the future, depending on the magnitude and vertical depth penetration of warming in the ocean.

Description: Previous bioassays conducted in the oligotrophic Atlantic Ocean identified availability of inorganic nitrogen (N) as the proximate limiting nutrient control of primary production, but additionally displayed a synergistic growth effect of combined N and phosphorus (P) addition. To classify conditions of nutrient limitation of coastal phytoplankton in the tropical ocean, we performed an 11 d nutrient-enrichment experiment with a natural phytoplankton community from shelf waters off northwest Africa in shipboard mesocosms. We used pigment and gene fingerprinting in combination with flow cytometry for classification and quantification of the taxon-specific photoautotrophic response to differences in nutrient supply. The developing primary bloom was dominated by diatoms and was significantly higher in the treatments receiving initial N addition. The combined supply of N and P did not induce a further increase in phytoplankton abundance compared to high N addition alone. A secondary bloom during the course of the experiment again displayed higher primary producer standing stock in the N-fertilized treatments. Bacterial abundance correlated positively with phytoplankton biomass. Dominance of the photoautotrophic assemblage by N-limited diatoms in conjunction with a probable absence of any P-limited phytoplankton species prevented an additive effect of combined N and P addition on total phytoplankton biomass. Furthermore, after nutrient exhaustion, dinitrogen (N-2)-fixing cyanobacteria succeeded the bloom-forming diatoms. Shelf waters in the tropical eastern Atlantic may thus support growth of diazotrophic cyanobacteria such as Trichodesmium sp. subsequent to upwelling pulses.

Description: Previous studies on trait-mediated trophic interactions in marine ecosystems were restricted to pair-wise interactions between one species of meso-herbivore and plant, though multi-grazer interactions are more common in nature. We investigated whether the feeding of one consumer, either the periwinkle Littorina littorea or the isopod Idotea baltica, affected consumption by the other consumer via anti-herbivory defence induction in the brown seaweed Fucus vesiculosus. To test the generality of our findings, we ran similar experiments with seaweed/grazer populations in the North and Baltic Seas (NE Atlantic). Grazer-specificity in induction strength was assessed by using the same species of grazer for induction and consumption. ‘Indirect’ induction effects were assessed by using different species of grazers for induction and consumption. Palatability assays were run with live algae and with reconstituted food to distinguish between different mechanisms of resistance. Grazing by herbivores induced a chemical defence in F. vesiculosus. In the North Sea population, the induced defences were only effective against I. baltica, regardless of inducer identity. The sensitive responses of I. baltica to the induced defences were also detected in the reconstituted food assays using Baltic Sea organisms. Thus, marine meso-grazers may be affected by previous feeding through the same or a different species of consumer by modified prey traits, such as induced chemical defences. Furthermore, the magnitude of the effect in the induced defences can be determined by species-specific sensitivity.

Description: Ocean acidification and associated changes in seawater carbonate chemistry negatively influence calcification processes and depress metabolism in many calcifying marine invertebrates. We present data on the cephalopod mollusc Sepia officinalis, an invertebrate that is capable of not only maintaining calcification, but also growth rates and metabolism when exposed to elevated partial pressures of carbon dioxide (pCO(2)). During a 6 wk period, juvenile S. officinalis maintained calcification under similar to 4000 and similar to 6000 ppm CO2, and grew at the same rate with the same gross growth efficiency as did control animals. They gained approximately 4%, body mass daily and increased the mass of their calcified cuttlebone by over 500 %. We conclude that active cephalopods possess a certain level of pre-adaptation to long-term increments in carbon dioxide levels. Our general understanding of the mechanistic processes that limit calcification must improve before we can begin to predict what effects future ocean acidification will have on calcifying marine invertebrates.

Description: Effects of global warming on marine ecosystems are far less understood than they are in terrestrial environments. Macrophyte-based coastal ecosystems are particularly vulnerable to global warming, because they often lack species redundancy. We tested whether summer heat waves have negative effects on an ecologically important ecosystem engineer, the eelgrass Zostera marina L., and whether high genotypic diversity may provide resilience in the face of climatic extremes. In a mesocosm experiment, we manipulated genotypic diversity of eelgrass patches fully crossed with water temperature (control vs. temperature stress) over 5 mo. We found a strong negative effect of warming and a positive effect of genotypic diversity on shoot densities of eelgrass. These results suggest that eelgrass meadows and associated ecosystem services will be negatively affected by predicted increases in summer temperature extremes. Genotypic diversity may provide critical response diversity for maintaining seagrass ecosystem functioning, and for adaptation to environmental change.

Description: The diazotrophic cyanobacterium Trichodesmium is a significant contributor to marine nitrogen and carbon cycles and has been incorporated in biogeochemical ocean circulation models. To date, parameterization of light as a controlling factor for nitrogen fixation has been based on field observations, where factors other than light also affect Trichodesmium physiology. Here we present data on light-dependent (15 to 1100 µmol quanta m–2 s–1) diazotrophic growth from controlled laboratory experiments and their implications for modeling approaches. We supply a simple empirical model to describe nitrogen fixation by Trichodesmium in batch cultures. Diazotrophic growth of axenic Trichodesmium IMS-101 was light saturated at 180 µmol quanta m–2 s–1 and did not vary significantly at higher photon irradiances up to 1100 µmol quanta m–2 s–1 (μcarbon based ≈ 0.26 d–1). Chlorophyll a (chl a) normalized N2 fixation rates were significantly affected by light intensity during mid-exponential growth (0.74 to 4.45 mol N fixed mol chl a–1 h–1) over the range of photon irradiances tested. In contrast, nitrogen fixation rates normalized to the cellular carbon content were relatively unaffected by light intensity (0.42 to 0.59, averaging 0.5 mmol N mol particulate organic carbon [POC]–1 h–1). Trichodesmium carbon biomass can be used to estimate the nitrogen input by this diazotroph into the ocean; the maximum input rate is 350 nmol N fixed l–1 h–1.

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: We investigated the nutritional condition of larval fishes caught in daily ichthyoplankton hauls in the ‘Helgoland Roads’ (54°11.18’N and 07°54.00’E) from February to June 2004. We concentrated on larvae of dab Limanda limanda and lesser sandeel Ammodytes marinus in order to compare early life stages of iteroparous and nearly semelparous fish. We analysed length, weight and RNA:DNA ratios as a proxy for condition of the larvae. The relationship between larval nutritional condition and larval size provided an indication of condition-selective mortality due to a loss of larvae in poor condition from the larger size-classes. For the sandeel, larvae in good condition were present in all size-classes, whereas for the dab maximum larval condition increased with increasing size. Variability in both standard length and condition was high in the 2 species during their planktonic stage. Both species showed good nutritional condition in the early to mid portion of the recruitment period and declines in condition in late April. This was more pronounced in larval dab, which showed a higher dependency on feeding conditions than larval sandeel. Together, these findings indicate a more conservative strategy of early life stages of the nearly semelparous sandeel compared with the iteroparous dab.

Description: The conspicuous retreat of the key species Fucus vesiculosus from the deeper parts of its former distribution area in the Baltic Sea has triggered extensive research on the factors that control its growth. Based on recently obtained knowledge on a large number of potential drivers, we developed a numerical model incorporating effects of abiotic factors on the physiological functions of photosynthesis, respiration, and reproduction and the ecological processes of competition, grazing, and epibiosis. For all input combinations, the model delivers the monthly net growth rate near the bladder wrack’s depth limit and the maximum depth of its vertical distribution. The use of data corresponding to conditions presently observed in the western Baltic Sea sets the year’s maximum algal net growth rate in late spring and 2 minima in early spring and autumn. The depth limit of the wrack’s distribution is set at ~9 m. Light and its absorption by phytoplankton represent by far the most important factors controlling the modeled net growth rate and depth penetration, with the role of epibiosis requiring further investigation. Lacking findings on population dynamics and biotic interactions restrict the generated model to an exploratory rather than a predictive tool.

Description: The impact of seawater acidification on calcifying organisms varies at the species level. If the impact differs between predator and prey in strength and/or sign, trophic interactions may be altered. In the present study, we investigated the impact of 3 different seawater pCO2 levels (650, 1250 and 3500 µatm) on the acid–base status or the growth of 2 predatory species, the common sea star Asterias rubens and the shore crab Carcinus maenas, and tested whether the quantity or size of prey consumed is affected. We exposed both the predators and their prey, the blue mussel Mytilus edulis, over a time span of 10 wk and subsequently performed feeding experiments. Intermediate acidification levels had no significant effect on growth or consumption in either predator species. The highest acidification level reduced feeding and growth rates in sea stars by 56%, while in crabs a 41% decrease in consumption rates of mussels could be demonstrated over the 10 wk experimental period but not in the subsequent shorter feeding assays. Because only a few crabs moulted in the experiment, acidification effects on crab growth could not be investigated. Active extracellular pH compensation by means of bicarbonate accumulation was observed in C. maenas, whereas the coelomic fluid pH in A. rubens remained uncompensated. Acidification did not provoke a measurable shift in prey size preferred by either predator. Mussels exposed to elevated pCO2 were preferred by previously untreated A. rubens but not by C. maenas. The observed effects on species interactions were weak even at the high acidification levels expected in the future in marginal marine habitats such as the Baltic Sea. Our results indicate that when stress effects are similar (and weak) on interacting species, biotic interactions may remain unaffected.

Description: Phytoplankton experience strong and abrupt variations in light intensity. How cells cope with these changes influences their competitiveness in a highly dynamical environment. While a considerable amount of work has focused on photoacclimation, it is still unknown whether processes specific of phytoplankton groups (e.g. calcification and silicification) influence their response to changing light. Here we show that the diatom Phaeodactylum tricornutum and the coccolithophore Emiliania huxleyi respond to an abrupt increase in irradiance by increasing carbon fixation rates, decreasing light absorption through the decrease of light-harvesting pigments and increasing energy dissipation through the xanthophyll cycle. In addition, E. huxleyi rapidly increases calcium carbonate precipitation in response to elevated light intensity, thereby providing an additional sink for excess energy. Differences between the 2 species also emerge with regard to the magnitude and timing of their individual responses. While E. huxleyi show a pronounced decrease in chlorophyll a and fucoxanthin cellular contents following increased light intensity, P. tricornutum has a faster increase in diadinoxanthin quota, a slower decrease in Fv/Fm (ratio of variable to maximum fluorescence) and a stronger increase in organic carbon fixation rate during the first 10 min. Our findings provide further evidence of species-specific responses to abrupt changes in light intensity, which may partly depend on the phytoplankton functional groups, with coccolithophores having a supplementary path (calcification) for the rapid dissipation of excess energy produced after an abrupt increase in light intensity. These differences might influence competition between coexisting species and may therefore have consequences at the community level.

Description: Rising atmospheric CO2 concentrations will have profound effects on atmospheric and hydrographic processes, which will ultimately modify the supply and chemistry of trace metals in the ocean. In addition to an increase in sea surface temperatures, higher CO2 also results in a decrease of seawater pH, known as ocean acidification, with implications for inorganic trace metal chemistry. Furthermore, direct or indirect effects of ocean acidification and ocean warming on marine biota will also affect trace metal biogeochemistry via alteration of biological trace metal uptake rates and metal binding to organic ligands. Currently, we still lack a holistic understanding of the impacts of decreasing seawater pH and rinsing temperatures on different trace metals and marine biota, which complicates projections into the future. Here, we outline how ocean acidification and ocean warming will influence the inputs and cycling of Fe and other biologically relevant trace metals globally, and regionally in high and low latitudes of the future ocean, discuss uncertainties, and highlight essential future research fields.

Description: Three bioassay experiments were performed to study the effects of nutrient and Saharan dust additions on natural diazotrophic communities in the tropical North Atlantic Ocean. Samples for nucleic acid analysis were collected at the beginning and end of 48 h incubations. TaqMan probes specific to 7 diazotrophic phylotypes, viz. filamentous cyanobacteria (Trichodesmium spp.), unicellular cyanobacterial (UCYN) Groups A, B, and C, Gamma A and P Proteobacteria, and Cluster III, were used to quantify nifH DNA abundances. N-2 fixation rates were measured in the same experiments using the N-15(2) gas bubble injection method. N-2 fixation was co-limited by P and Fe. Total nifH abundances increased relative to the control with additions of either Fe or P or both in combination. Additions of dissolved N, alone or in combination with phosphate, induced increases in UCYN-A and Gamma A nifH compared with the control. Saharan dust additions significantly stimulated fixation rates. Abundances of all cyanobacterial and Gamma A nifH phylotypes at least doubled after Saharan dust additions where surface water dissolved Fe concentrations were 〈2 nmol l(-1). Laboratory experiments with cultures of T. erythraeum demonstrated that dust addition promoted colony formation and the persistence of T. erythraeum biomass relative to cultures to which no Fe was added. Our results with both field and laboratory experiments indicate that Saharan dust positively affects diazotrophic phylotype abundances and changes T. erythraeum colony morphology.

Description: At 2 modern deep-water cold-seep sites, the North Alex Mud Volcano (eastern Mediterranean Sea, water depth ~500 m) and the Concepción Methane Seep Area (south-east Pacific Ocean, water depth ~700 m), we found abundant catshark (Chondrichthyes: Scylio - rhinidae) and skate (Chondrichthyes: Rajidae) egg capsules, respectively, associated with carbonates and tubeworms. Fossilized catshark egg capsules were found at the 35 million year old Bear River Cold-Seep Deposit (Washington State, USA) closely associated with remains of tubeworms and sponges. We suggest that cold-seep ecosystems have served as nurseries for predatory elasmobranch fishes since at least late Eocene time and therewith possibly play an important role for the functioning of deep-water ecosystems.

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

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

Description: 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: Epibacterial communities on thalli of the algal species Fucus serratus, Fucus vesiculosus, Laminaria saccharina, Ulva compressa, Delesseria sanguinea and Phycodrys rubens were analysed using 16S ribosomal RNA gene-based DGGE. Individuals of all species were collected in the Kiel Fjord (Baltic Sea) and in the rocky intertidal of Helgoland (North Sea). DGGE gels as well as cluster and multidimensional scaling analysis based on the DGGE band patterns of the epibacterial community showed significant differences between the epibacterial communities on the investigated algal species both in the Baltic and North Seas. Epibacterial communities differed less between regions than between host species, and were more similar on closely related host species. Results give the first evidence for lineage-specific bacterial associations to algal thalli. Furthermore, the results suggest that these algal species may control their epibiotic bacterial communities.

Description: The family Solemyidae represents ancestral protobranch bivalves with the shallow-water genus Solemya and the deep-sea genus Acharax. All known members of this family host symbiotic sulfur-oxidizing bacteria in their gill filaments. Analysis of 18S rRNA gene sequences of Acharax specimens from methane-seeps off Makran (Pakistan), Java (Indonesia), the Aleutian Trench and off the Oregon, Costa Rica, and Peru margins revealed that Solemya spp. and Acharax spp. are well-separated genetically. This supports the current systematic distinction based on morphological criteria. We found 2 clearly distinct clusters within the genus Acharax, with specimens from the Makran, Oregon and Peru (MOP) margins in one (MOP–Acharax) cluster, and those from Java, the Aleutian Trench and Costa Rica (JAC) in the other (JAC–Acharax) cluster. The separation of MOP– and JAC–Acharax clusters from each other and from Solemya (S. reidi and S. velum) is well-supported by phylogenetic calculations employing maximum likelihood and maximum parsimony. Compared to genetic distances among other protobranch groups, distances between the MOP– and JAC–Acharax clusters would justify the affiliation of these clusters to separate species. This implies that species differentiation in Acharax based on shell morphology is likely to underestimate true species diversity within this taxon. Furthermore, our results support the hypothesis that genetic separation of Solemya and Acharax is congruent with the phylogeny of their bacterial endosymbionts.

Description: The response of the phytoplankton and bacterial spring succession to the predicted warming of sea surface temperature in temperate climate zones during winter was studied using an indoor-mesocosm approach. The mesocosms were filled with winter water from the Kiel Fjord, Baltic Sea. Two of them were started at ~2°C and the temperature was subsequently increased according to the decadal temperature profile of the fjord (ΔT 0°C, baseline treatment). The other mesocosms were run at 3 elevated temperatures with differences of ΔT +2, +4 and +6°C. All mesocosms were exposed to the same light conditions. Timing of peak phytoplankton primary production (PP) during the experimental spring bloom was not significantly influenced by increasing temperatures, whereas the peak of bacterial secondary production (BSP) was accelerated by about 2 d per °C. This suggests that, in case of warming, the spring peak of bacterial degradation of organic matter (in terms of BSP) would occur earlier in the year. Furthermore, the lag time between the peaks of PP and BSP (about 16 d for ΔT 0°C) would diminish progressively at elevated temperatures. The average ratio between BSP and PP increased significantly from 0.37 in the coldest mesocosms to 0.63 in the warmest ones. Community respiration and the contribution of picoplankton (〈3 µm fraction) to this also increased at elevated temperatures. Our results lead to the prediction that climate warming during the winter/ early spring in temperate climate zones will favor bacterial degradation of organic matter by tightening the coupling between phytoplankton and bacteria. However, if PP is reduced by warming, as in our experiments, this will not necessarily lead to increased recycling of organic matter (and CO2).

Description: The Kasatochi volcanic eruption that occurred in the central Aleutian Islands in Alaska, USA, in August 2008 is thought to have induced a massive diatom bloom in the iron-limited waters of the Gulf of Alaska, which potentially affected the oceanic food web by increasing the abundance of zooplankton and sockeye salmon Oncorhynchus nerka in the northeast Pacific Ocean. We report the first seawater experiments involving volcanic ash ejected from the Kasatochi eruption, showing that the ash released 61 to 83 nmol Fe, 374 to 410 nmol NO3-, 5 to 6 nmol PO43- and 170 to 585 nmol SiO2 when it contacted seawater. Our study suggests that the amount of iron released from Kasatochi ash (an increase of 2.0 to 2.8 nM Fe) was indeed sufficient to cause the observed phytoplankton bloom in the northeastern Pacific Gyre, while the impact of macronutrient release was minimal. We further evaluated the multiple, interdependent processes in the oceanic food web related to the diatom bloom, involving the ocean survival of juvenile salmon that entered the northeast Pacific Ocean in the summer of 2008.

Description: Effects of CO2 concentration on elemental composition of the coccolithophore Emiliania huxleyi were studied in phosphorus-limited, continuous cultures that were acclimated to experimental conditions for 30 d prior to the first sampling. We determined phytoplankton and bacterial cell numbers, nutrients, particulate components like organic carbon (POC), inorganic carbon (PIC), nitrogen (PN), organic phosphorus (POP), transparent exopolymer particles (TEP), as well as dissolved organic carbon (DOC) and nitrogen (DON), in addition to carbonate system parameters at CO2 levels of 180, 380 and 750 µatm. No significant difference between treatments was observed for any of the measured variables during repeated sampling over a 14 d period. We considered several factors that might lead to these results, i.e. light, nutrients, carbon overconsumption and transient versus steady-state growth. We suggest that the absence of a clear CO2 effect during this study does not necessarily imply the absence of an effect in nature. Instead, the sensitivity of the cell towards environmental stressors such as CO2 may vary depending on whether growth conditions are transient or sufficiently stable to allow for optimal allocation of energy and resources. We tested this idea on previously published data sets where PIC and POC divided by the corresponding cell abundance of E. huxleyi at various pCO2 levels and growth rates were available.

Description: The extraordinary molecular phylogenetic diversity of picoeukaryotes and their contributions to marine processes have been highlighted recently. Herein, picoeukaryotic diversity in the coastal Pacific Ocean is characterized for the first time. Close to full length small subunit ribosomal RNA (ssu rRNA) gene amplicons were cloned and sequenced from size fractionated samples (〈 2 μm) taken between September 2000 and October 2001. Sequences belonging to the order Mamiellales were abundant, with Micromonas pusilla-like sequences in all 5 libraries (4 operational taxonomic units, OTUs, at ≥99 sequence similarity), Ostreococcus in 4 (1 OTU) and Bathycoccus in 2 (1 OTU). Phylogenetic reconstruction showed distinct Micromonas clades at this site (although not for other Mamiellales), supporting the idea that the designation M. pusilla harbors cryptic species. In combination with picophytoplankton dynamics previously reported for the same period, the data indicates that picoplanktonic primary production at this site is likely to be dominated by prasinophytes. Furthermore, a Micromonas-clade with no cultured representatives was identified bearing highest identity to Sargasso Sea shotgun clone sequences. Comparison of the Pacific sequences with the shotgun clones showed Ostreococcus and Micromonas at 1 Sargasso site with elevated chlorophyll (chl) levels, but not at other Sargasso sites. Other Pacific clones were primarily Novel Alveolate Group II, which were highly diverse based on OTU analyses. Novel Alveolate Group I, Ciliophora, Cercozoa-like and stramenopile sequences were also retrieved. Although picoeukaryotic diversity has been characterized in only 1 other Pacific Ocean sample (equatorial Pacific), most stramenopile and alveolate sequences corresponded to previously identified phylogenetic clades from studies conducted in other oceans and for which no cultured representatives exist. © Inter-Research 2006.

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: The community composition of denitrifying bacteria was studied in the stratified water column of Lake Kinneret. The nitrite reductase genes nirS and nirK were amplified by PCR from water samples taken at 1, 14, 19 and 22 m depth, which represent the epi-, meta- and hypolimnion of the lake. The PCR products were analyzed with terminal restriction fragment length polymorphism (T-RFLP) and clone libraries. The highest diversity of nirS denitrifying communities was observed at 1 m depth. According to the T-RFLP profiles and clone libraries of nirS products, 2 groups of denitrifiers were common to and dominant in all depths. Deduced protein sequences from one of these groups displayed low identity (77%) with other nirS sequences reported in GenBank. Denitrifying bacterial communities with nirK were most diverse at 22 m and showed highest similarity to those at 19 m depth. Sequences unrelated to nirK dominated the clone libraries from 1 m depth, suggesting that denitrifying bacteria with copper-containing nitrite reductase were less frequent at this depth. The results suggest that microorganisms with nirK and those with nirS respond differently to the environmental conditions in the stratified water column of Lake Kinneret.

Description: Representatives of the phylum Bacteroidetes are repeatedly reported to be abundant members of oceanic bacterioplankton, probably because of their ability to degrade complex organic matter. 16S rDNA analysis was performed in order to address the importance of this phylum in a highly oligotrophic region such as the Eastern Mediterranean Sea and to investigate its distribution patterns and community composition. A new antisense primer was designed for PCR and used in combination with the general eubacterial sense primer 27F to specifically target Bacteroidetes representatives. Data were correlated with water depth and water mass properties. Denaturant gradient gel electrophoresis (DGGE) analysis and sequencing of environmental clone libraries revealed specific distribution patterns. A major fraction of the sequences was assigned to the AGG58 cluster, a branch of yet uncultured members of the Bacteroidetes lineage. Our results demonstrate a considerable diversity within the Bacteroidetes lineage in the oligotrophic Eastern Mediterranean Sea. Differing water mass properties are considered to be of major relevance for the spatial distribution with depth; several environmental clone sequence clusters could be specifically assigned to a defined water mass or to the deep waters of investigated locations. Depth-specific distribution of Bacteroidetes is demonstrated for the first time by the results of this study.

Description: Soft corals of the family Xeniidae are particularly abundant in Red Sea coral reefs. Their success may be partly due to a strong defense mechanism against fish predation. To test this, we conducted field and aquarium experiments in which we assessed the antifeeding effect of secondary metabolites of 2 common xeniid species, Ovabunda crenata and Heteroxenia ghardaqensis. In the field experiment, the metabolites of both investigated species reduced feeding on experimental food pellets in the natural population of Red Sea reef fishes by 86 and 92% for O. crenata and H. ghardaqensis, respectively. In the aquarium experiment, natural concentration of crude extract reduced feeding on experimental food pellets in the common reef fish Thalassoma lunare (moon wrasse) by 83 and 85%, respectively. Moon wrasse feeding was even reduced at extract concentrations as low as 12.5% of the natural concentration in living soft coral tissues. To assess the potential of a structural anti-feeding defence, sclerites of O. crenata were extracted and mixed into food pellets at natural, doubled and reduced concentration without and in combination with crude extract at 25% of natural concentration, and tested in an aquarium experiment. The sclerites did not show any effect on the feeding behavior of the moon wrasse indicating that sclerites provide structural support rather than antifeeding defense. H. ghardaqensis lacks sclerites. We conclude that the conspicuous abundance of xeniid soft coral species in the Red Sea is likely a consequence of a strong chemical defence, rather than physical defences, against potential predators.

Description: Seagrasses worldwide are commonly infected by endophytic protists of the genus Labyrinthula. To date, the nature of interaction of endophyte and host is not well understood. In eelgrass (Zostera marina) Labyrinthula zosterae may become virulent (pathogenic) and lead to the loss of entire sea grass beds. One of the best known examples of any marine epidemic were outbreaks of the ´wasting disease´ on both sides of the Atlantic in the 1930s, but smaller infestations have been reported until recently. Up to now, detection of infection by Labyrinthula was based on the wasting index, i.e. the relative area of leaf lesions or microscopy, while genetic data are virtually absent. We characterized a ~1400 base pair portion of the 18S small subunit rDNA in L. zosterae isolates (N=41) from six northern European sites and one southern location (Adriatic Sea) in order to assess identity and potential diversity of endophytic protists. Because there are indications that low salinity impedes Labyrinthula growth, sampling sites included a wide range of salinities from 5-34 psu. A search against the non-redundant GENBANK data base revealed that most isolates are 99% similar to the only L. zosterae 18S sequence available from the data base at all but the Finish site (salinity values 5-7 psu). At the latter site, a different Labyrinthula species occurred, which was also found in fully marine Wadden Sea cultures. A third species was detected in Skagerrak, south-western Baltic and North Sea samples (20-25 psu). We conclude that L. zosterae is widespread among northern European eelgrass sites across wide ranges of salinity.