Food sources, behaviour, and distribution of hydrothermal vent shrimps at the Mid-Atlantic Ridge

Five species of bresilioid shrimp were investigated at seven hydrothermal sites on the Mid-Atlantic Ridge: Menez Gwen, Lucky Strike, Rainbow, Broken Spur, TAG, Snake Pit and Logatchev. Samples were prepared for analysis of stable isotopes, elemental composition and lipids. Shrimp behaviour was observed from the submersible ‘Alvin’ and in the laboratory aboard RV ‘Atlantis’. The distribution and zonation of the shrimp species was recorded. Juvenile shrimp of all species arrive at the vents carrying reserves of photosynthetic origin, built-up in the pelagic larval stages. These reserves are used while the shrimp metamorphose to the adult form and, in Rimicaris exoculata and Chorocaris chacei, while they develop epibiotic bacteria supporting structures, the modified mouthparts and the inside of the carapace. The main food of adult R. exoculata is filamentous bacteria that grow on these structures. The intermediate sizes of C. chacei also feed on such bacteria, but the final stage gets some food by scavenging or predation. Mirocaris species scavenge diverse sources; they are not trophically dependent on either R. exoculata or mussels. Adults of Alvinocaris markensis are predators of other vent animals, including R. exoculata. The dense swarms of R. exoculata, with their exosymbionts, can be compared to endosymbiont-containing animals such as Bathymodiolus and the vestimentiferan tube-worms of the Pacific vents. Such associations, whether endo- or ectosymbiotic, may be necessary for the development of flourishing communities at hydrothermal vents.

Impact of climate change on Antarctic krill

Antarctic krill Euphausia superba (hereafter ‘krill’) occur in regions undergoing rapid environmental change, particularly loss of winter sea ice. During recent years, harvesting of krill has increased, possibly enhancing stress on krill and Antarctic ecosystems. Here we review the overall impact of climate change on krill and Antarctic ecosystems, discuss implications for an ecosystem-based fisheries management approach and identify critical knowledge gaps. Sea ice decline, ocean warming and other environmental stressors act in concert to modify the abundance, distribution and life cycle of krill. Although some of these changes can have positive effects on krill, their cumulative impact is most likely negative. Recruitment, driven largely by the winter survival of larval krill, is probably the population parameter most susceptible to climate change. Predicting changes to krill populations is urgent, because they will seriously impact Antarctic ecosystems. Such predictions, however, are complicated by an intense inter-annual variability in recruitment success and krill abundance. To improve the responsiveness of the ecosystem-based management approach adopted by the Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR), critical knowledge gaps need to be filled. In addition to a better understanding of the factors influencing recruitment, management will require a better understanding of the resilience and the genetic plasticity of krill life stages, and a quantitative understanding of under-ice and benthic habitat use. Current precautionary management measures of CCAMLR should be maintained until a better understanding of these processes has been achieved.

Severe tissue damage in Atlantic cod larvae under increasing ocean acidification

Ocean acidification, caused by increasing atmospheric concentrations of CO2 (refs 1-3), is one of the most critical anthropogenic threats to marine life. Changes in seawater carbonate chemistry have the potential to disturb calcification, acid-base regulation, blood circulation and respiration, as well as the nervous system of marine organisms, leading to long-term effects such as reduced growth rates and reproduction(4,5). In teleost fishes, early life-history stages are particularly vulnerable as they lack specialized internal pH regulatory mechanisms(6,7). So far, impacts of relevant CO2 concentrations on larval fish have been found in behaviour(8,9) and otolith size(10,11), mainly in tropical, non-commercial species. Here we show detrimental effects of ocean acidification on the development of a mass-spawning fish species of high. commercial importance. We reared Atlantic cod larvae at three levels of CO2, (1) present day, (2) end of next century and (3) an extreme, coastal upwelling scenario, in a long-term (2; months) mesocosm experiment. Exposure to CO2 resulted in severe to lethal tissue damage in many internal organs, with the degree of damage increasing with CO2 concentration. As larval survival is the bottleneck to recruitment, ocean acidification has the potential to act as an additional source of natural mortality, affecting populations of already exploited fish stocks.

Climate, plankton and cod

While a few North Atlantic cod stocks are stable, none have increased and many have declined in recent years. Although overfishing is the main cause of most observed declines, this study shows that in some regions, climate by its influence on plankton may exert a strong control on cod stocks, complicating the management of this species that often assumes a constant carrying capacity. First, we investigate the likely drivers of changes in the cod stock in the North Sea by evaluating the potential relationships between climate, plankton and cod. We do this by deriving a Plankton Index that reflects the quality and quantity of plankton food available for larval cod. We show that this Plankton Index explains 46.24% of the total variance in cod recruitment and 68.89% of the variance in total cod biomass. Because the effects of climate act predominantly through plankton during the larval stage of cod development, our results indicate a pronounced sensitivity of cod stocks to climate at the warmer, southern edge of their distribution, for example in the North Sea. Our analyses also reveal for the first time, that at a large basin scale, the abundance of Calanus finmarchicus is associated with a high probability of cod occurrence, whereas the genus Pseudocalanus appears less important. Ecosystem-based fisheries management (EBFM) generally considers the effect of fishing on the ecosystem and not the effect of climate-induced changes in the ecosystem state for the living resources. These results suggest that EBFM must consider the position of a stock within its ecological niche, the direct effects of climate and the influence of climate on the trophodynamics of the ecosystem.

Diel vertical migration of decapod larvae in the Portuguese coastal upwelling ecosystem: implications for offshore transport

The vertical distribution of decapod larvae off the northwest Portuguese coast was analysed in relation to associated environmental conditions from sampling during a 69 h period around a current meter mooring located on the shelf, approximately 21 km off the coast. Plankton samples were collected every 2 h at the surface with a neuston net and through the water column with a Longhurst Hardy Plankton Recorder (Pro-LHPR), allowing a very detailed resolution of larval vertical distribution. Environmental data (temperature, salinity, and chlorophyll a) were obtained every hour. To investigate the horizontal distribution of decapod larvae in relation to the coast, a plankton-sampling grid was carried out before the 69 h fixed station. Larvae of shelf decapod species were widely distributed over the shelf, while those of inshore species were found much closer to the coast. Decapod larvae (zoeae and megalopae) showed clear diel vertical migrations, only appearing in the upper 20 m at night, a migration that did not appear to be affected by physical conditions in the water column. Larval densities were highly variable, 0.01 to 215 ind. m super(-3) for zoeae and 0 to 93 ind. m super(-3) for megalopae, the zoeae being generally more abundant. The results indicated that during the day larvae accumulate very close to the bottom. The diel vertical migration behaviour is discussed as one of the contributing mechanisms for larval retention over the shelf, even with offshore transport conditions promoted by coastal upwelling, and is hence of major relevance for the recruitment success of decapod species that inhabit inshore and shelf zones of coastal upwelling systems.

Long-Term Retrospective Analysis of Mackerel Spawning in the North Sea: A New Time Series and Modeling Approach to CPR Data

We present a unique view of mackerel (Scomber scombrus) in the North Sea based on a new time series of larvae caught by the Continuous Plankton Recorder (CPR) survey from 1948-2005, covering the period both before and after the collapse of the North Sea stock. Hydrographic backtrack modelling suggested that the effect of advection is very limited between spawning and larvae capture in the CPR survey. Using a statistical technique not previously applied to CPR data, we then generated a larval index that accounts for both catchability as well as spatial and temporal autocorrelation. The resulting time series documents the significant decrease of spawning from before 1970 to recent depleted levels. Spatial distributions of the larvae, and thus the spawning area, showed a shift from early to recent decades, suggesting that the central North Sea is no longer as important as the areas further west and south. These results provide a consistent and unique perspective on the dynamics of mackerel in this region and can potentially resolve many of the unresolved questions about this stock

Climate-induced effects on the meroplankton and the benthic-pelagic ecology of the North Sea

Analyses of long-term time series of North Sea plankton and sea surface temperature (SST) data reveal that the annual planktonic larval abundance of three benthic phyla, Echinodermata, Arthropoda, and Mollusca, responds positively and immediately to SST. Long-term outcomes for the planktonic abundance of these three phyla are different, however. The planktonic larvae of echinoderms and decapod crustaceans have increased in abundance from 1958 to 2005, and especially since the mid-1980s, as North Sea SST has increased. In contrast, the abundance of bivalve mollusc larvae has declined, despite the positive year-to-year relationship between temperature and bivalve larval abundance continuing to hold. We argue that the changes in meroplankton abundance, coincident with increased phytoplankton and declining holoplankton, reflect the synchronous effect of rising SST and related changes in the pelagic community on the reproduction and recruitment of many benthic marine invertebrates. Under this scenario, the long-term decline in bivalve mollusc larvae will reflect increased predation on the settled larvae and adults by benthic decapods. These alterations in the zooplankton may therefore describe an ecosystem-wide restructuring of North Sea trophic interactions.

Distribution and Seasonal Cycles of Decapod Crustacean Larvae in Continuous Plankton Records from the North Pacific Ocean

Decapoda taken in Continuous Plankton Recorder (CPR) samples from the Pacific in 1997 and 2000-2003 have been identified and measured. Some previously un-described larval stages were referred to species and characteristics of these are described. Distributions and seasonal occurrence of decapod taxa in the samples are described and discussed with particular emphasis on the dendrobranchiate shrimp Sergestes similis and the brachyurans Cancer spp. And Chionoecetes spp. There is a prolonged larval season at low levels of abundance off the Californian coast but in the more northern waters there is a shorter productive period but numbers of larvae per sample are high, particularly in June. Larvae of Chionoecetes and other Oregoninae were found only from May to July.

Fish data from the Continuous Plankton Recorder survey

The Continuous Plankton Recorder (CPR) survey has been sampling plankton in the North Sea since 1931. However the identification of the larval and juvenile fish taken in the survey has not been a part of the routine analysis of the samples. Specialist analysis of the fish was carried out between 1948 and the early 1980s but the data were available as hard copy only. As part of MarBEF, data on >60 taxa from 1948 to 1972 have been entered on a database which is now linked to EUROBIS to show the biogeographical information and the data are available for general research. Examples of the data are shown. These data provide a background on the variability of fish stocks before the recent period of rapid warming and in some cases before significant fisheries developed. Data for subsequent years will be made available as possible and work is now underway to bring the analysis up to date. This will provide time series over six decades.

Recruitment in a changing environment: the 2000s North Sea herring recruitment failure

Environmentally induced change appears to be impacting the recruitment of North Sea herring (Clupea harengus). Despite simultaneously having a large adult population, historically low exploitation, and Marine Stewardship Council accreditation (implying sustainability), there have been an unprecedented 6 sequential years of poor juvenile production (recruitment). Analysis suggests that the poor recruitment arises during the larval overwintering phase, with recent survival rates greatly reduced. Contemporary warming of the North Sea has caused significant changes in the plankton community, and a recently identified regime shift around 2000 shows close temporal agreement with the reduced larval survival. It is, therefore, possible that we are observing the first consequences of this planktonic change for higher trophic levels. There is no indication of a recovery in recruitment in the short term. Fishing mortality is currently outside the agreed management plan, and forecasts show a high risk of the stock moving outside safe biological limits soon, potentially precipitating another collapse of the stock. However, bringing the realized fishing mortality back in line with the management plan would likely alleviate the problem. This illustrates again that recruitment is influenced by more than just spawning-stock biomass, and that changes in other factors can be of equal, or even greater, importance. In such dynamically changing environments, recent management success does not necessarily guarantee future sustainability.

Empirically modelling the potential effects of changes in temperature and prey availability on the growth of cod larvae in UK shelf seas

It has been hypothesized that changes in zooplankton community structure over the past four decades led to reduced growth and survival of prerecruit Atlantic cod (Gadus morhua) and that this was a key factor underlying poor year classes, contributing to stock collapse, and inhibiting the recovery of stocks around the UK. To evaluate whether observed changes in plankton abundance, species composition and temperature could have led to periods of poorer growth of cod larvae, we explored the effect of prey availability and temperature on early larval growth using an empirical trophodynamic model. Prey availability was parameterized using species abundance data from the Continuous Plankton Recorder. Our model suggests that the observed changes in plankton community structure in the North Sea may have had less impact on cod larval growth, at least for the first 40 days following hatching, than previously suggested. At least in the short term, environmental and prey conditions should be able to sustain growth of cod larvae and environmental changes acting on this early life stage should not limit stock recovery.

Black-legged kittiwakes as indicators of environmental change in the North Sea; evidence from long-term studies

Top predators, particularly seabirds, have repeatedly been suggested as indicators of marine ecosystem status. One region currently under pressure from human fisheries and climate change is the North Sea. Standardized seabird monitoring data have been collected on the Isle of May, an important seabird colony in the northwestern North Sea, over the last 10–20 years. Over this period oceanographic conditions have varied markedly, and between 1990 and 1999 a major industrial fishery for sandlance (Ammodytes marinus), the main prey of most seabird species, was prosecuted nearby. Sandlance fishing grounds close to seabird colonies down the east coast of the UK were closed in 2000 in an attempt to improve foraging opportunities for breeding seabirds, particularly black-legged kittiwakes (Rissa tridactyla). Initially this closure seemed to be beneficial for kittiwakes with breeding success recovering to pre-fishery levels. However, despite the ban continuing, kittiwakes and many other seabird species in the North Sea suffered severe breeding failures in 2004. In this paper, we test the predictive power of four previously established correlations between kittiwake breeding success and climatic/trophic variables to explain the observed breeding success at the Isle of May in 2004. During the breeding season, kittiwakes at this colony switch from feeding on 1+ group to 0 group sandlance, and results up until 2003 indicated that availability of both age classes had a positive effect on kittiwake breeding success. The low breeding success of kittiwakes in 2004 was consistent with the late appearance and small body size of 0 group sandlance, but at odds with the two variables likely to operate via 1 group availability (lagged winter sea surface temperature and larval sandlance cohort strength in 2003). The reason for the discrepancy is currently unknown, but analysis of 1 group sandlance body composition indicated that lipid content in 2004 was extremely low, and thus fish eaten by kittiwakes during pre-breeding and early incubation were likely to be of poor quality. Monitoring of reproductive success of kittiwakes, although useful, was clearly not sufficient to tease apart the complex causation underlying the 2004 event. Monitoring programs such as this, therefore, need to be complemented by detailed research to identify the mechanisms involved, and to attribute and predict the effects of natural and human-induced environmental change.

Feeding selectivity of bivalve larvae on natural plankton assemblages in the Western English Channel

Meroplankton, including bivalve larvae, are an important and yet understudied component of coastal marine food webs. Understanding the baseline of meroplankton ecology is imperative to establish and predict their sensitivity to local and global marine stressors. Over an annual cycle (October 2009–September 2010), bivalve larvae were collected from the Western Channel Observatory time series station L4 (50°15.00′N, 4°13.02′W). The morphologically similar larvae were identified by analysis of the 18S nuclear small subunit ribosomal RNA gene, and a series of incubation experiments were conducted to determine larval ingestion rates on natural plankton assemblages. Complementary gut content analysis was performed using a PCR-based method for detecting prey DNA both from field-collected larvae and those from the feeding experiments. Molecular identification of bivalve larvae showed the community composition to change over the course of the sampling period with domination by Phaxas in winter and higher diversity in autumn. The larvae selected for nanoeukaryotes (2–20 µm) including coccolithophores (<20 µm) which together comprised >75 % of the bivalve larvae diet. Additionally, a small percentage of carbon ingested originated from heterotrophic ciliates (<30 µm). The molecular analysis of bivalve larvae gut content provided increased resolution of identification of prey consumed and demonstrated that the composition of prey consumed established through bottle incubations conferred with that established from in situ larvae. Despite changes in bivalve larvae community structure, clearance rates of each prey type did not change significantly over the course of the experiment, suggesting different bivalve larvae species may consume similar prey.

Sub-Arctic populations of European lobster (Homarus gammarus) in Northern Norway.

The European lobster is distributed throughout the south and western regions of the Norwegian coast. A previous lobster allozyme investigation (1993) in the Tysfjord region, north of the Arctic Circle demonstrated that the lobster population from this region was genetically different from lobster samples collected in other parts of Norway. More detailed investigation including supplementary extensive sampling and additional allozyme, microsatellite and mtDNA analyses are reported here. This investigation supports the genetic distinctness of the Tysfjord population and shows that this is mainly due to a reduction (60�70%) in gene diversity (observed heterozygosities and number of alleles) compared with lobsters from more southern regions. In addition to the Tysfjord region, the comprehensive sampling also included lobsters found in the adjacent Nordfolda fjord system. Genetic analyses provided evidence for significant differences between the lobster populations of Tysfjord and Nordfolda, even though they are separated by a coastal distance of only 142 km. The two populations were also different with regards to several biological characteristics such as body size. The genetic difference between these two geographically close populations is likely to be due to the local hydrological conditions, preventing larval dispersal between the fjord systems. Assessment of lobster abundance in the north-west region suggests that the sub-arctic lobster populations are geographically isolated.

Commercial oyster stocks as a potential source of larvae in the regeneration of Ostrea edulis in Strangford Lough, Northern Ireland.

Ostrea edulis was extremely rare in the wild in Strangford Lough from the early 1900s until renewed spatfall was observed at a number of sites in the 1990s. A monitoring programme was undertaken to investigate the presence and distribution of planktonic oyster larvae at nine sites around the lough between June and September in 1997 and 1998 as a precursor to studies of spatfall patterns. Larval densities at sites in the northern basin of the lough were significantly higher than those in the southern basin where larvae were lacking or in low numbers. Densities and sizes of oyster larvae showed significant temporal variation suggesting pulsed larval release. Larval densities also showed significant spatial variation with higher densities at sites closer to commercial stocks pointing to these as the main source of oyster larvae. This hypothesis was supported during a larval flux study over a complete tidal cycle which indicated a 90% net tidal movement of O. edulis larvae from the entrance of the bay where commercial stocks were held to the main body of the lough. Thus the maintenance of dense commercial stocks of flat oysters may provide the key to the redevelopment of native oyster beds in Strangford Lough and elsewhere by providing an initial broodstock nucleus from which larvae can be exported.