Pattern of benthic biomass size spectra from shallow waters in the East China Seas

Benthic biomass size spectra (BSS) and normalized biomass size spectra were constructed, and benthic secondary production was estimated by a size spectrum equation in the shallow waters in the East China Sea, ranging latitudinally from 40A degrees N to 29A degrees N. The BSS patterns were bimodal, two biomass peaks corresponding to meiofauna and macrofauna, respectively, separated by a trough of low biomass at 8-256 mu g individual dry weight which varied in position with median sediment particle size. The BSS also displayed bimodality within meiofauna size ranges, which in most stations was due to the relative proportions of nematodes and other meiofauna taxa. Re-analysis of data from sites in the UK, South Africa, and Antarctic showed a similar bimodality in the adult species body size distribution within the meiofauna size range. Macrofaunal production estimated by the size spectrum equation was very similar to the results of Brey90 empirical equation. However, these production values were much lower than those calculated by Brey01. Different individual dry-to-wet conversion ratios, temperature deviation, and macrofauna taxonomic composition might be responsible for the between-model differences. The macrofaunal P/B ratios calculated by this equation ranged from 0.3 to 3.4 which were in accordance with values from Northern Hemisphere mid-latitudes. Meiofaunal production estimates will need further empirical support.

Shortened duration of the annual Neocalanus plumchrus biomass peak in the Northeast Pacific

The calanoid copepod Neocalan us plumchrus (Marukawa) is a dominant member of the spring mesozooplankton in the subarctic North Pacific and Bering Sea. Previous studies have shown interdecadal and latitudinal variation in seasonal developmental timing, with peak biomass occurring earlier in years and places with warmer upper ocean temperatures. Because N. plumchrus normally has a single dominant annual cohort, its seasonal timing can be indexed from measurements of total population biomass or by following progressive changes in stage composition. Early studies empirically found that peak upper ocean biomass occurred when about half of the pre-dormant population had reached copepodite stage 5 (C5). However, more recent comparisons derived from recent Continuous Plankton Recorder (CPR) data now show peak biomass when a larger fraction (> 80%) of the population is at C5. CPR samples the surface 10 to 15 m, but comparisons to depth-resolved BIONESS data show that this discrepancy is not an artefact of sampling depth. Other causes are either a prolongation of duration of pre-dormant C5 or a narrowing of the age range making up the annual cohort. We assessed changes in cohort width using a modification of Greve's cumulative percentile method, and found that average cohort widths in the Alaska Gyre were significantly narrower in 2000-2007 than in 1957-1965 (1968-1980 were intermediate). Net tow sampling of Strait of Georgia populations showed a similar significant narrowing of cohorts in the 2003-2005 sampling period. This study provides evidence that in addition to the shift to an earlier occurrence of peak biomass reported previously, the duration of the peak has also decreased in the last decade.

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.

Diet of auklet chicks in the Aleutian Islands, Alaska: similarity among islands, interspecies overlap, and relationships to ocean climate

Seabirds are effective samplers of the marine environment, and can be used to measure resource partitioning among species and sites via food loads destined for chicks. We examined the composition, overlap, and relationships to changing climate and oceanography of 3,216 food loads from Least, Crested, and Whiskered Auklets (Aethia pusilla, A. cristatella, A. pygmaea) breeding in Alaska during 1994–2006. Meals comprised calanoid copepods (Neocalanus spp.) and euphausiids (Thysanoessa spp.) that reflect secondary marine productivity, with no difference among Buldir, Kiska, and Kasatochi islands across 585 km of the Aleutian Islands. Meals were very similar among species (mean Least–Crested Auklet overlap C = 0.68; Least–Whiskered Auklet overlap C = 0.96) and among sites, indicating limited partitioning of prey resources for auklets feeding chicks. The biomass of copepods and euphausiids in Least and Crested Auklet food loads was related negatively to the summer (June–July–August) North Pacific Gyre Oscillation, while in Whiskered Auklet food loads, this was negatively related to the winter (December–January–February) Pacific Decadal Oscillation, both of which track basin-wide sea-surface temperature (SST) anomalies. We found a significant quadratic relationship between the biomass of calanoid copepods in Least Auklet food loads at all three study sites and summer (June–July) SST, with maximal copepod biomass between 3–6°C (r 2 = 0.71). Outside this temperature range, zooplankton becomes less available to auklets through delayed development. Overall, our results suggest that auklets are able to buffer climate-mediated bottom-up forcing of demographic parameters like productivity, as the composition of chick meals has remained constant over the course of our study.

Long-term changes in North Atlantic zooplankton biomass

New measures of zooplankton biomass have been derived from CPR samples in the North Atlantic from 1958 to 2005. The final aim was to investigate how the zooplankton standing stock had varied throughout the last decades, knowing that in different areas of the North Atlantic significant changes in the distribution of the dominant zooplankton species as well as the plankton assemblage have been observed. During the forty-five years of monitoring the contribution of the different groups (e.g. copepods, euphausiids, meroplankton larvae) to the total zooplankton biomass has been evaluated. The changes in the phenology of the biomass were also considered. The relationship between quantity, quality and seasonal timing of plankton and the poor fish recruitment seen in recent years in the North Sea are also discussed.

Effects of environmental conditions on the seasonal distribution of phytoplankton biomass in the North Sea

We study the spatial and seasonal variability of phytoplankton biomass (as phytoplankton color) in relation to the environmental conditions in the North Sea using data from the Continuous Plankton Recorder survey. By using only environmental fields and location as predictor variables we developed a nonparametric model (generalized additive model) to empirically explore how key environmental factors modulate the spatio-temporal patterns of the seasonal cycle of algal biomass as well as how these relate to the ,1988 North Sea regime shift. Solar radiation, as manifest through changes of sea surface temperature (SST), was a key factor not only in the seasonal cycle but also as a driver of the shift. The pronounced increase in SST and in wind speed after the 1980s resulted in an extension of the season favorable for phytoplankton growth. Nutrients appeared to be unimportant as explanatory variables for the observed spatio-temporal pattern, implying that they were not generally limiting factors. Under the new climatic regime the carrying capacity of the whole system has been increased and the southern North Sea, where the environmental changes have been more pronounced, reached a new maximum.

A long-term chlorophyll dataset reveals regime shift in North Sea phytoplankton biomass unconnected to nutrient levels

During the 1980s, a rapid increase in the Phytoplankton Colour Index (PCI), a semiquantitative visual estimate of algal biomass, was observed in the North Sea as part of a regionwide regime shift. Two new data sets created from the relationship between the PCI and SeaWiFS chlorophyll a (Chl a) quantify differences in the previous and current regimes for both the anthropogenically affected coastal North Sea and the comparatively unaffected open North Sea. The new regime maintains a 13% higher Chl a concentration in the open North Sea and a 21% higher concentration in coastal North Sea waters. However, the current regime has lower total nitrogen and total phosphorus concentrations than the previous regime, although the molar N: P ratio in coastal waters is now well above the Redfield ratio and continually increasing. Besides becoming warmer, North Sea waters are also becoming clearer (i.e., less turbid), thereby allowing the normally light-limited coastal phytoplankton to more effectively utilize lower concentrations of nutrients. Linear regression analyses indicate that winter Secchi depth and sea surface temperature are the most important predictors of coastal Chl a, while Atlantic inflow is the best predictor of open Chl a; nutrient concentrations are not a significant predictor in either model. Thus, despite decreasing nutrient concentrations, Chl a continues to increase, suggesting that climatic variability and water transparency may be more important than nutrient concentrations to phytoplankton production at the scale of this study.

Long-term declining trend of zooplankton biomass in the Tropical Atlantic

We report evidences that the zooplankton biomass in the tropical Atlantic has declined with an almost 10-fold drop from the 1950s to 2000. The results of the multiple regression analysis showed that the decline in zooplankton biomass was positively related to the NAO-index and to phosphate concentration. We also found that the depth of the thermocline has decreased over the period of our investigation. Thus, the decline we report in zooplankton biomass may be related to the combined effect of two phenomena driven by global temperature increase: (1) the widening of the distributional range of tropical species due to the expansion of the ‘tropical belt’ and (2) a decrease in primary production resulting from the thinning of the thermocline. The decline of zooplankton biomass we report suggests that global warming of the ocean may be altering tropical food webs, and through them, it may also indirectly impact tropical oceans biogeochemical cycles.

Temporal and spatial variation in the Nazaré Canyon (Western Iberian margin): Inter-annual and canyon heterogeneity effects on meiofauna biomass and diversity

The Nazaré Canyon on the Portuguese Margin (NE Atlantic) was sampled during spring-summer for three consecutive years (2005–2007), permitting the first inter-annual study of the meiofaunal communities at the Iberian Margin at two abyssal depths (~3500 m and ~4400 m). Using new and already published data, the meiofauna standing stocks (abundance and biomass) and nematode structural and functional diversity were investigated in relation to the sediment biogeochemistry (e.g. organic carbon, nitrogen, chlorophyll a, phaeopigments) and grain size. A conspicuous increase in sand content from 2005 to 2006 and decrease of phytodetritus at both sites, suggested the occurrence of one or more physical disturbance events. Nematode standing stocks and trophic diversity decreased after these events, seemingly followed by a recovery/recolonisation period in 2007, which was strongly correlated with an increase in the quantity and bioavailability of phytodetrital organic matter supplied. Changes in meiofauna assemblages, however, also differed between stations, likely because of the contrasting hydrodynamic and food supply conditions. Higher meiofauna and nematode abundances, biomass and trophic complexity were found at the shallowest canyon station, where the quantity, quality and bioavailability of food material were higher than at the deeper site. The present results suggest that even though inter-annual variations in the sedimentary environment can regulate the meiofauna in the abyssal Nazaré Canyon, heterogeneity between sampling locations in the canyon were more pronounced.

Interactions between multiple large macrofauna species and nematode communities — Mechanisms for indirect impacts of trawling disturbance

Highlights •We exposed meiofauna to 7 different large macrofauna species at high and low densities. •Macrofauna presence altered nematode community structure and reduced their abundance. •Macrofauna species had similar effects by reducing the few dominant nematode species. •Meio–macrofauna resource competition and spatial segregation are the main drivers. •Trawling effects on macrofauna affect nematode communities indirectly. Diverse assemblages of infauna in sediments provide important physical and biogeochemical services, but are under increasing pressure by anthropogenic activities, such as benthic trawling. It is known that trawling disturbance has a substantial effect on the larger benthic fauna, with reductions in density and diversity, and changes in community structure, benthic biomass, production, and bioturbation and biogeochemical processes. Largely unknown, however, are the mechanisms by which the trawling impacts on the large benthic macro- and megafauna may influence the smaller meiofauna. To investigate this, a mesocosm experiment was conducted whereby benthic nematode communities from a non-trawled area were exposed to three different densities (absent, low, normal) of 7 large (> 10 mm) naturally co-occurring, bioturbating species which are potentially vulnerable to trawling disturbance. The results showed that total abundances of nematodes were lower if these large macrofauna species were present, but no clear nematode abundance effects could be assigned to the macrofauna density differences. Nematode community structure changed in response to macrofauna presence and density, mainly as a result of the reduced abundance of a few dominant nematode species. Any detectable effects seemed similar for nearly all macrofauna species treatments, supporting the idea that there may be a general indirect, macrofauna-mediated trawling impact on nematode communities. Explanations for these results may be, firstly, competition for food resources, resulting in spatial segregation of the meio- and macrobenthic components. Secondly, different densities of large macrofauna organisms may affect the nematode community structure through different intensities of bioturbatory disturbance or resource competition. These results suggest that removal or reduced densities of larger macrofauna species as a result of trawling disturbance may lead to increased nematode abundance and hints at the validity of interference competition between large macrofauna organisms and the smaller meiofauna, and the energy equivalence hypothesis, where a trade-off is observed between groups of organisms that are dependent on a common source of energy.

Biomass changes and trophic amplification of plankton in a warmer ocean

Ocean warming can modify the ecophysiology and distribution of marine organisms, and relationships between species, with nonlinear interactions between ecosystem components potentially resulting in trophic amplification. Trophic amplification (or attenuation) describe the propagation of a hydroclimatic signal up the food web, causing magnification (or depression) of biomass values along one or more trophic pathways. We have employed 3-D coupled physical-biogeochemical models to explore ecosystem responses to climate change with a focus on trophic amplification. The response of phytoplankton and zooplankton to global climate-change projections, carried out with the IPSL Earth System Model by the end of the century, is analysed at global and regional basis, including European seas (NE Atlantic, Barents Sea, Baltic Sea, Black Sea, Bay of Biscay, Adriatic Sea, Aegean Sea) and the Eastern Boundary Upwelling System (Benguela). Results indicate that globally and in Atlantic Margin and North Sea, increased ocean stratification causes primary production and zooplankton biomass to decrease in response to a warming climate, whilst in the Barents, Baltic and Black Seas, primary production and zooplankton biomass increase. Projected warming characterized by an increase in sea surface temperature of 2.29 ± 0.05 °C leads to a reduction in zooplankton and phytoplankton biomasses of 11% and 6%, respectively. This suggests negative amplification of climate driven modifications of trophic level biomass through bottom-up control, leading to a reduced capacity of oceans to regulate climate through the biological carbon pump. Simulations suggest negative amplification is the dominant response across 47% of the ocean surface and prevails in the tropical oceans; whilst positive trophic amplification prevails in the Arctic and Antarctic oceans. Trophic attenuation is projected in temperate seas. Uncertainties in ocean plankton projections, associated to the use of single global and regional models, imply the need for caution when extending these considerations into higher trophic levels.

Plankton indicators and ocean observing systems: support to the marine ecosystem state assessment

Ecological indicators are used extensively as tools to manage environmental resources. In the oceans, indicators of plankton can be measured using a variety of observing systems including: mooring stations, ships, autonomous floats and ocean colour remote sensing. Given the broad range of temporal and spatial sampling resolutions of these different observing systems, as well as discrepancies in measurements obtained from different sensors, the estimation and interpretation of plankton indicators can present significant challenges. To provide support to the assessment of the state of the marine ecosystem, we propose a suite of plankton indicators and subsequently classify them in an ecological framework that characterizes key attributes of the ecosystem. We present two case studies dealing with plankton indicators of biomass, size structure and phenology, estimated using the most spatially extensive and longest in situ and remote-sensing observations. Discussion of these studies illustrates how some of the challenges in estimating and interpreting plankton indicators may be addressed by using for example relative measurement thresholds, interpolation procedures and delineation of biogeochemical provinces. We demonstrate that one of the benefits attained, when analyzing a suite of plankton indicators classified in an ecological framework, is the elucidation of non-trivial changes in composition, structure and functioning of the marine ecosystem.