Biodiversity and ecosystems: Change at the community level

Some commercial fish species of the northeast Atlantic Ocean have relocated in response to warming. The impact of warming on marine assemblages in the region may already be much greater than appreciated, however, with over 70% of common demersal fish species responding through changes in abundance, rather than range. The northeast Atlantic Ocean is one of the most productive marine ecoregions in the world with a substantial commercial fishery. It is also a region that has undergone particularly rapid warming over the past 50 years, up to four times faster than the global average1. Compared with other marine regions worldwide, the biological response in the northeast Atlantic Ocean has been particularly dramatic, reflecting this rapid warming. Studies have documented biogeographical movements in marine plankton of over 1,000 km northwards2 and advances in the onset of key life-history events by six to eight weeks3. In addition, there has been limited evidence of distributional shifts in some fish species along latitudinal and depth gradients in response to warming4, 5. Writing in Current Biology, Stephen Simpson and colleagues6 present the most comprehensive analysis so far of the impact of warming on commercially important European continental-shelf fish species in the region, and in doing so show that there has been a profound reorganization of local communities.

Is the Russell Cycle a true cycle? Multidecadal zooplankton and climate trends in the western English Channel

The Russell Cycle is one of the classical examples of climate influence on biological oceanography, represented as shifts in the marine plankton over several decades with warm and cool conditions. While the time-series data associated with the phenomenon indicate cyclical patterns, the question remains whether or not the Russell Cycle should be considered a “true cycle”. Zooplankton time-series data from 1924 to 2011 from the western English Channel were analysed with principal component (PC), correlation and spectral analyses to determine the dominant trends, and cyclic frequencies of the Russell Cycle indicators in relation to long-term hydroclimatic indices. PC1 accounted for 37.4% of the variability in the zooplankton data with the main contributions from non-clupeid fish larvae, southwestern zooplankton, and overall zooplankton biovolume. For PC2 (14.6% of data variance), the dominant groups were northern fish larvae, non-sardine eggs, and southern fish larvae. Sardine eggs were the major contributors to PC3 (representing 12.1% of data variance). No significant correlations were observed between the above three components and climate indices: Atlantic Multidecadal Oscillation, North Atlantic Oscillation, and local seawater temperature. Significant 44- and 29-year frequencies were observed for PC3, but the physical mechanisms driving the cycles are unclear. Harmonic analysis did not reveal any significant frequencies in the physical variables or in PCs 1 and 2. To a large extent, this is due to the dominant cycles in all datasets generally being long term (>50 years or so) and not readily resolved in the examined time frame of 88 years, hence restricting the ability to draw firm conclusions on the multidecadal relationship between zooplankton community dynamics in the western English Channel and environmental indices. Thus, the zooplankton time-series often associated and represented as the Russell Cycle cannot be concluded as being truly cyclical.

Is the Russell Cycle a true cycle? Multidecadal zooplankton and climate trends in the western English Channel

The Russell Cycle is one of the classical examples of climate influence on biological oceanography, represented as shifts in the marine plankton over several decades with warm and cool conditions. While the time-series data associated with the phenomenon indicate cyclical patterns, the question remains whether or not the Russell Cycle should be considered a “true cycle”. Zooplankton time-series data from 1924 to 2011 from the western English Channel were analysed with principal component (PC), correlation and spectral analyses to determine the dominant trends, and cyclic frequencies of the Russell Cycle indicators in relation to long-term hydroclimatic indices. PC1 accounted for 37.4% of the variability in the zooplankton data with the main contributions from non-clupeid fish larvae, southwestern zooplankton, and overall zooplankton biovolume. For PC2 (14.6% of data variance), the dominant groups were northern fish larvae, non-sardine eggs, and southern fish larvae. Sardine eggs were the major contributors to PC3 (representing 12.1% of data variance). No significant correlations were observed between the above three components and climate indices: Atlantic Multidecadal Oscillation, North Atlantic Oscillation, and local seawater temperature. Significant 44- and 29-year frequencies were observed for PC3, but the physical mechanisms driving the cycles are unclear. Harmonic analysis did not reveal any significant frequencies in the physical variables or in PCs 1 and 2. To a large extent, this is due to the dominant cycles in all datasets generally being long term (>50 years or so) and not readily resolved in the examined time frame of 88 years, hence restricting the ability to draw firm conclusions on the multidecadal relationship between zooplankton community dynamics in the western English Channel and environmental indices. Thus, the zooplankton time-series often associated and represented as the Russell Cycle cannot be concluded as being truly cyclical.

Use of Continuous Plankton Recorder information in support of marine management: applications in fisheries, environmental protection, and in the study of ecosystem response to environmental change

The Continuous Plankton Recorder (CPR) survey was conceived from the outset as a programme of applied research designed to assist the fishing industry. Its survival and continuing vigour after 70 years is a testament to its utility, which has been achieved in spite of great changes in our understanding of the marine environment and in our concerns over how to manage it. The CPR has been superseded in several respects by other technologies, such as acoustics and remote sensing, but it continues to provide unrivalled seasonal and geographic information about a wide range of zooplankton and phytoplankton taxa. The value of this coverage increases with time and provides the basis for placing recent observations into the context of long-term, large-scale variability and thus suggesting what the causes are likely to be. Information from the CPR is used extensively in judging environmental impacts and producing quality status reports (QSR); it has shown the distributions of fish stocks, which had not previously been exploited; it has pointed to the extent of ungrazed phytoplankton production in the North Atlantic, which was a vital element in establishing the importance of carbon sequestration by phytoplankton. The CPR continues to be the principal source of large-scale, long-term information about the plankton ecosystem of the North Atlantic. It has recently provided extensive information about the biodiversity of the plankton and about the distribution of introduced species. It serves as a valuable example for the design of future monitoring of the marine environment and it has been essential to the design and implementation of most North Atlantic plankton research.

From plankton to top predators: bottom-up control of a marine food web across four trophic levels

1. Abundant mid-trophic pelagic fish often play a central role in marine ecosystems, both as links between zooplankton and top predators and as important fishery targets. In the North Sea, the lesser sandeel occupies this position, being the main prey of many bird, mammal and fish predators and the target of a major industrial fishery. However, since 2003, sandeel landings have decreased by > 50%, and many sandeel-dependent seabirds experienced breeding failures in 2004. 2. Despite the major economic implications, current understanding of the regulation of key constituents of this ecosystem is poor. Sandeel abundance may be regulated 'bottom-up' by food abundance, often thought to be under climatic control, or 'top-down' by natural or fishery predation. We tested predictions from these two hypotheses by combining unique long-term data sets (1973–2003) on seabird breeding productivity from the Isle of May, SE Scotland, and plankton and fish larvae from the Continuous Plankton Recorder survey. We also tested whether seabird breeding productivity was more tightly linked to sandeel biomass or quality (size) of individual fish. 3. The biomass of larval sandeels increased two- to threefold over the study period and was positively associated with proxies of the abundance of their plankton prey. Breeding productivity of four seabirds bringing multiple prey items to their offspring was positively related to sandeel larval biomass with a 1-year lag, indicating dependence on 1-year-old fish, but in one species bringing individual fish it was strongly associated with the size of adult sandeels. 4. These links are consistent with bottom-up ecosystem regulation and, with evidence from previous studies, indicate how climate-driven changes in plankton communities can affect top predators and potentially human fisheries through the dynamics of key mid-trophic fish. However, the failing recruitment to adult sandeel stocks and the exceptionally low seabird breeding productivity in 2004 were not associated with low sandeel larval biomass in 2003, so other mechanisms (e.g. predation, lack of suitable food after metamorphosis) must have been important in this case. Understanding ecosystem regulation is extremely important for predicting the fate of keystone species, such as sandeels, and their predators.

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.

Marine snow studies in the Northeast Atlantic Ocean: distribution, composition and role as a food source for migrating plankton

During a 25 d Lagrangian study in May and June 1990 in the Northeast Atlantic Ocean, marine snow aggregates were collected using a novel water bottle, and the composition was determined microscopically. The aggregates contained a characteristic signature of a matrix of bacteria, cyanobacteria and autotrophic picoplankton with inter alia inclusions of the tintiniid Dictyocysta elegans and large pennate diatoms. The concentration of bacteria and cyanobacteria was much greater on the aggregates than when free-living by factors of 100 to 6000 and 3000 to 2 500 000, respectively, depending on depth. Various species of crustacean plankton and micronekton were collected, and the faecal pellets produced after capture were examined. These often contained the marine snow signature, indicating that these organisms had been consuming marine snow. In some cases, marine snow material appeared to dominate the diet. This implies a food-chain short cut wherby material, normally too small to be consumed by the mesozooplankton, and considered to constitute the diet of the microplankton can become part of the diet of organisms higher in the food-chain. The micronekton was dominated by the amphipod Themisto compressa, whose pellets also contained the marine snow signature. Shipboard incubation experiments with this species indicated that (1) it does consume marine snow, and (2) its gut-passage time is sufficiently long for material it has eaten in the upper water to be defecated at its day-time depth of several hundred meters. Plankton and micronekton were collected with nets to examine their vertical distribution and diel migration and to put into context the significance of the flux of material in the guts of migrants. “Gut flux” for the T. compressa population was calculated to be up to 2% of the flux measured simultaneously by drifting sediment traps and <5% when all migrants are considered. The in situ abundance and distribution of marine snow aggregates (>0.6 mm) was examined photographically. A sharp concentration peak was usually encountered in the depth range 40 to 80 m which was not associated with peaks of in situ fluorescence or attenuation but was just below or at the base of the upper mixed layer. The feeding behaviour of zooplankton and nekton may influence these concentration gradients to a considerable extent, and hence affect the flux due to passive settling of marine snow aggregates.