On the roles of cell size and trophic strategy in North Atlantic diatom and dinoflagellate communities

We have examined the inter- and intra-group seasonal succession of 113 diatom and dinoflagellate taxa, as surveyed by the Continuous Plankton Recorder (CPR) in the North Atlantic, by grouping taxa according to two key functional traits: cell size (mg C cell21) and trophic strategy (photoautotrophy, mixotrophy, or heterotrophy). Mixotrophic dinoflagellates follow photoautotrophic diatoms but precede their obligate heterotrophic counterparts in the succession because of the relative advantages afforded by photosynthesizing when light and nutrients are available in spring. The mean cell size of the sampled diatoms is smallest in the summer, likely because of the higher specific nutrient affinity of smaller relative to larger cells. Contrastingly, we hypothesize that mixotrophy diminishes the size selection based on nutrient limitation and accounts for the lack of a seasonal size shift among surveyed dinoflagellates. Relatively small, heterotrophic dinoflagellates (mg C cell21 , 1023) peak after other, larger dinoflagellates, in part because of the increased abundance of their small prey during nutrientdeplete summer months. The largest surveyed diatoms (mg C cell21 . 1022) bloom later than others, and we hypothesize that this may be because of their relatively slow maximum potential growth rates and high internal nutrient storage, as well as to the slower predation of these larger cells. The new trait database and analysis presented here helps translate the taxonomic information of the CPR survey into metrics that can be directly compared with trait-based models.

Phenological trends and trophic mismatch across multiple levels of a North Sea pelagic food web

Differential phenological responses to climate among species are predicted to disrupt trophic interactions, but datasets to evaluate this are scarce. We compared phenological trends for species from 4 levels of a North Sea food web over 24 yr when sea surface temperature (SST) increased significantly. We found little consistency in phenological trends between adjacent trophic levels, no significant relationships with SST, and no significant pairwise correlations between predator and prey phenologies, suggesting that trophic mismatching is occurring. Finer resolution data on timing of peak energy demand (mid-chick-rearing) for 5 seabird species at a major North Sea colony were compared to modelled daily changes in length of 0-group (young of the year) lesser sandeels Ammodytes marinus. The date at which sandeels reached a given threshold length became significantly later during the study. Although the phenology of all the species except shags also became later, these changes were insufficient to keep pace with sandeel length, and thus mean length (and energy value) of 0-group sandeels at mid-chick-rearing showed net declines. The magnitude of declines in energy value varied among the seabirds, being more marked in species showing no phenological response (shag, 4.80 kJ) and in later breeding species feeding on larger sandeels (kittiwake, 2.46 kJ) where, due to the relationship between sandeel length and energy value being non-linear, small reductions in length result in relatively large reductions in energy. However, despite the decline in energy value of 0-group sandeels during chick-rearing, there was no evidence of any adverse effect on breeding success for any of the seabird species. Trophic mismatch appears to be prevalent within the North Sea pelagic food web, suggesting that ecosystem functioning may be disrupted.

Wasp-Waist Interactions in the North Sea Ecosystem

In a “wasp-waist” ecosystem, an intermediate trophic level is expected to control the abundance of predators through a bottom-up interaction and the abundance of prey through a top-down interaction. Previous studies suggest that the North Sea is mainly governed by bottom-up interactions driven by climate perturbations. However, few studies have investigated the importance of the intermediate trophic level occupied by small pelagic fishes.

Scale-dependent response diversity of seabirds to prey in the North Sea

Functional response diversity is defined as the diversity of responses to environmental change among species that contribute to the same ecosystem function. Because different ecological processes dominate on different spatial and temporal scales, response diversity is likely to be scale dependent. Using three extensive data sets on seabirds, pelagic fish, and zooplankton, we investigate the strength and diversity in the response of seabirds to prey in the North Sea over three scales of ecological organization. Two-stage analyses were used to partition the variance in the abundance of predators and prey among the different scales of investigation: variation from year to year, variation among habitats, and variation on the local patch scale. On the year-to-year scale, we found a strong and synchronous response of seabirds to the abundance of prey, resulting in low response diversity. Conversely, as different seabird species were found in habitats dominated by different prey species, we found a high diversity in the response of seabirds to prey on the habitat scale. Finally, on the local patch scale, seabirds were organized in multispecies patches. These patches were weakly associated with patches of prey, resulting in a weak response strength and a low response diversity. We suggest that ecological similarities among seabird species resulted in low response diversity on the year-to-year scale. On the habitat scale, we suggest that high response diversity was due to interspecific competition and niche segregation among seabird species. On the local patch scale, we suggest that facilitation with respect to the detection and accessibility of prey patches resulted in overlapping distribution of seabirds but weak associations with prey. The observed scale dependencies in response strength and diversity have implications for how the seabird community will respond to different environmental disturbances.

Role of light and photophysiological properties on phytoplankton succession during the spring bloom in the north-western Mediterranean Sea

This study aimed to determine the role of light on the succession of the phytoplankton community during the spring bloom in the northwestern Mediterranean Sea. To this end, three successive Lagrangian experiments were carried out between March and April 2003. The three experiments correspond to distinct phases of the bloom development (pre-bloom, bloom peak and post-bloom, respectively) and therefore to different trophic conditions. Phytoplankton (sampled on a daily scale) was grouped in size-based classes (pico and nano+micro) each of them were characterised in terms of chemotaxonomic composition, primary production and photophysiological properties. The phytoplankton community evolved with time changing in both size-class dominance and specie/group dominance within each size class. The bloom peak was characterised by highly dynamic condition (i.e. vertical mixing) and by the dominance of both small (pico) and large (nano and micro) diatoms, as a result of their capacity to photoacclimate to changing light regimes (‘physiological plasticity’). Concluding, we suggest that the physiological adaptation to light is the main factor driving the succession of the phytoplankton community during the first phases of the bloom (until the onset of thermal stratification) in the western Mediterranean Sea.

Identification of the copepodite developmental stages of twenty-six North Atlantic copepods. Occasional Publication of the Marine Biological Association No. 21 (revised edition)

This identification guide to the copepodite developmental stages of twenty-six North Atlantic copepods has been revised and extended, to include new information, to update the taxonomy and to give additional details on how to determine sex in the later copepodite stages of gymnoplean copepods.

Temporal changes in plankton of the North Sea: community shifts and environmental drivers

This paper analyses long-term and seasonal changes in the North Sea plankton community during the period 1970 to 2008. Based on Continuous Plankton Recorder (CPR) data covering 38 yr, major changes in both phytoplankton and zooplankton abundance and community structure were identified. Regime changes were detected around 1978, 1989 and 1998. The first 2 changes have been discussed in the literature and are defined as a cold episodic event (1978) and a regime shift towards a warm dynamic regime (1989). The effect of these 2 regime changes on plankton indicators was assessed and checked against previous studies. The 1998 change represents a shift in the abundance and seasonal patterns of dinoflagellates and the dominant zooplankton group, the neritic copepods. Furthermore, environmental factors such as air temperature, wind speed and the North Atlantic water inflow were identified as potential drivers of change in seasonal patterns, and the most-likely environmental causes for detected changes were assessed. We suggest that a change in the balance of dissolved nutrients driven by these environmental factors was the cause of the latest change in plankton community structure, which in turn could have affected the North Sea fish community.

The North Pacific Continuous Plankton Recorder Survey

The Continuous Plankton Recorder (CPR) survey of the North Pacific is a PICES project now in its ninth year and facing an uncertain future. CPRs have been towed behind commercial ships along two (north–south and east–west) transects for a total of ~ nine times per year. Samples are collected with a filtering mesh and are then microscopically processed for plankton abundance in the laboratory. The survey, so far, has accumulated 3,648 processed samples (with approximately three times as many archived without processing), each representing 18 km of the transect (Fig. 1) and containing an abundance of data on over 290 phytoplankton and zooplankton taxa. A CTD with a fluorometer has been attached to the CPR sampling at the east–west transect in more recent years to provide supplementary environmental data.

Plankton populations at the bifurcation of the North Pacific Current

As the eastward-flowing North Pacific Current approaches the North American continent it bifurcates into the southward-flowing California Current and the northward-flowing Alaska Current. This bifurcation occurs in the south-eastern Gulf of Alaska and can vary in position. Dynamic height data from Project Argo floats have recently enabled the creation of surface circulation maps which show the likely position of the bifurcation; during 2002 it was relatively far north at 53 degrees N then, during early 2003, it moved southwards to a more normal position at 45 degrees N. Two ship-of-opportunity transects collecting plankton samples with a Continuous Plankton Recorder across the Gulf of Alaska were sampled seasonally during 2002 and 2003. Their position was dependent on the commercial ship's operations; however, most transects sampled across the bifurcation. We show that the oceanic plankton differed in community composition according to the current system they occurred in during spring and fall of 2002 and 2003, although winter populations were more mixed. Displacement of the plankton communities could have impacts on the plankton's reproduction and development if they use cues such as day length, and also on foraging of higher trophic-level organisms that use particular regions of the ocean if the nutritional value of the communities is different. Although we identify some indicator taxa for the Alaska and California currents, functional differences in the plankton communities on either side of the bifurcation need to be better established to determine the impacts of bifurcation movement on the ecosystems of the north-east Pacific.

Changes in development timing and cohort width of Neocalanus plumchrus degree ' flemingeri copepods in the eastern North Pacific

Neocalanus plumchrus/flemingeri copepods make up a large proportion of spring mesozooplankton biomass and are a valuable nutritional source for many higher trophic levels. Copepodites through to sub-adult stage are present in surface waters for a relatively short period of time each spring, and the date of maximum biomass has been calculated as the date when 50% of the population were at the sub-adult, CV stage. This index allows quite a precise date to be calculated from relatively infrequent sampling and interannual comparisons between 1957 and 2004 have demonstrated that the timing of peak abundance is significantly advanced in warmer years. However, recent data from the Continuous Plankton Recorder survey, which samples the surface NE Pacific more frequently during spring, has found that maximum numbers of CV copepodites occur after the 50% point is reached so that maximum biomass occurs some weeks later than predicted by this index (although comparisons between years show that the magnitude of the timing shift is similar). Comparisons with depth-stratified profiles from the BIONESS show that this is not just due to single-depth near-surface sampling by the CPR. We speculate on the cause of this change which could be related to the width of the cohort (which appears to now be narrower, at least in warm years) or the length of time that the CV stage needs to spend in the surface accumulating lipid before beginning diapause. A narrower cohort has implications for predators who will have less time to take advantage of this food source.

Rapid biogeographical plankton shifts in the North Atlantic Ocean

Large-scale biogeographical changes in the biodiversity of a key zooplankton group (calanoid copepods) were detected in the north-eastern part of the North Atlantic Ocean and its adjacent seas over the period 1960–1999. These findings provided key empirical evidence for climate change impacts on marine ecosystems at the regional to oceanic scale. Since 1999, global temperatures have continued to rise in the region. Here, we extend the analysis to the period 1958–2005 using all calanoid copepod species assemblages (nine species assemblages based on an analysis including a total of 108 calanoid species or taxa) and show that this phenomenon has been reinforced in all regions. Our study reveals that the biodiversity of calanoid copepods are responding quickly to sea surface temperature (SST) rise by moving geographically northward at a rapid rate up to about 23.16 km yr−1. Our analysis suggests that nearly half of the increase in sea temperature in the northeast Atlantic and adjacent seas is related to global temperature rises (46.35% of the total variance of temperature) while changes in both natural modes of atmospheric and oceanic circulation explain 26.45% of the total variance of temperature. Although some SST isotherms have moved northwards by an average rate of up to 21.75 km yr−1 (e.g. the North Sea), their movement cannot fully quantify all species assemblage shifts. Furthermore, the observed rates of biogeographical movements are far greater than those observed in the terrestrial realm. Here, we discuss the processes that may explain such a discrepancy and suggest that the differences are mainly explained by the fluid nature of the pelagic domain, the life cycle of the zooplankton and the lesser anthropogenic influence (e.g. exploitation, habitat fragmentation) on these organisms. We also hypothesize that despite changes in the path and intensity of the oceanic currents that may modify quickly and greatly pelagic zooplankton species, these organisms may reflect better the current impact of climate warming on ecosystems as terrestrial organisms are likely to significantly lag the current impact of climate change.

Decadal changes in climate and ecosystems in the North Atlantic Ocean and adjacent seas

Climate change is unambiguous and its effects are clearly detected in all functional units of the Earth system. This study presents new analyses of sea-surface temperature changes and show that climate change is affecting ecosystems of the North Atlantic. Changes are seen from phytoplankton to zooplankton to fish and are modifying the dominance of species and the structure, the diversity and the functioning of marine ecosystems. Changes also range from phenological to biogeographical shifts and have involved in some regions of the Atlantic abrupt ecosystem shifts. These alterations reflect a response of pelagic ecosystems to a warmer temperature regime. Mechanisms are complex because they are nonlinear exhibiting tipping points and varying in space and time. Sensitivity of organisms to temperature changes is high, implicating that a small temperature modification can have sustained ecosystem effects. Implications of these changes for biogeochemical cycles are discussed. Two observed changes detected in the North Sea that could have opposite effects on carbon cycle are discussed. Increase in phytoplankton, as inferred from the phytoplankton colour index derived from the Continuous Plankton Recorder (CPR) survey, has been detected in the North Sea. This pattern has been accompanied by a reduction in the abundance of the herbivorous species Calanus finmarchicus. This might have reduced the grazing pressure and increase diatomaceous ‘fluff’, therefore carbon export in the North Sea. Therefore, it could be argued that the biological carbon pump might increase in this region with sea warming. In the meantime, however, the mean size of organisms (calanoid copepods) has dropped. Such changes have implications for the turnover time of biogenic carbon in plankton organisms and the mean residence time of particulate carbon they produce. The system characterising the warmer period is more based on recycling and less on export. The increase in the minimum turnover time indicates an increase in the ecosystem metabolism, which can be considered as a response of the pelagic ecosystems to climate warming. This phenomenon could reduce carbon export. These two opposite patterns of change are examples of the diversity of mechanisms and pathways the ecosystems may exhibit with climate change. Oversimplification of current biogeochemical models, often due to lack of data and biological understanding, could lead to wrong projection on the direction ecosystems and therefore some biogeochemical cycles might take in a warmer world.