ALBERT

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

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

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

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

Description: 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: 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: 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.

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

Description: The 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: 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: 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.