Macroecological study of Centropages typicus in the North Atlantic Ocean

Centropages typicus is a temperate neritic-coastal species of the North Atlantic Oceans, generally found between the latitudes of the Mediterranean and the Norwegian Sea. Therefore, the species experiences a large number of environments and adjusts its life cycle in response to changes in key abiotic parameters such as temperature. Using data from the Continuous Plankton Recorder (CPR) Survey, we review the macroecology of C. typicus and factors that influence its spatial distribution, phenology and year-to-year to decadal variability. The ecological preferences are identified and quantified. Mechanisms that allow the species to occur in such different environments are discussed and hypotheses are proposed as to how the species adapts to its environment. We show that temperature and both quantity and quality of phytoplankton are important factors explaining the space and time variability of C. typicus. These results show that C. typicus will not respond only to temperature increase in the region but also to changes in phytoplankton abundance, structure and composition and timing of occurrence. Methods such as a decision tree can help to forecast expected changes in the distribution of this species with hydro-climatic forcing. (C) 2007 Elsevier Ltd. All rights reserved.

Long-term responses of North Atlantic calcifying plankton to climate change

The global increase in atmospheric carbon dioxide concentration is potentially threatening marine biodiversity in two ways. First, carbon dioxide and other greenhouse gases accumulating in the atmosphere are causing global warming1. Second, carbon dioxide is altering sea water chemistry, making the ocean more acidic2. Although temperature has a cardinal influence on all biological processes from the molecular to the ecosystem level3, acidification might impair the process of calcification or exacerbate dissolution of calcifying organisms4. Here, we show however that North Atlantic calcifying plankton primarily responded to climate-induced changes in temperatures during the period 1960–2009, overriding the signal from the effects of ocean acidification. We provide evidence that foraminifers, coccolithophores, both pteropod and nonpteropod molluscs and echinoderms exhibited an abrupt shift circa 1996 at a time of a substantial increase in temperature5 and that some taxa exhibited a poleward movement in agreement with expected biogeographical changes under sea temperature warming6,7. Although acidification may become a serious threat to marine calcifying organisms, our results suggest that over the study period the primary driver of North Atlantic calcifying planktonwas oceanic temperature.

Relationships between North Atlantic salmon, plankton and hydroclimatic change in the Northeast Atlantic

The abundance of wild salmon (Salmo salar) in the North Atlantic has declined markedly since the late 1980s as a result of increased marine mortality that coincided with a marked rise in sea temperature in oceanic foraging areas. There is substantial evidence to show that temperature governs the growth, survival, and maturation of salmon during their marine migrations through either direct or indirect effects. In an earlier study (2003), long-term changes in three trophic levels (salmon, zooplankton, and phytoplankton) were shown to be correlated significantly with sea surface temperature (SST) and northern hemisphere temperature (NHT). A sequence of trophic changes ending with a stepwise decline in the total nominal catch of North Atlantic salmon (regime shift in ∼1986/1987) was superimposed on a trend to a warmer dynamic regime. Here, the earlier study is updated with catch and abundance data to 2010, confirming earlier results and detecting a new abrupt shift in ∼1996/1997. Although correlations between changes in salmon, plankton, and temperature are reinforced, the significance of the correlations is reduced because the temporal autocorrelation of time-series substantially increased due to a monotonic trend in the time-series, probably related to global warming. This effect may complicate future detection of effects of climate change on natural systems.

Comparing copepod time-series in the north of Spain: Spatial autocorrelation of community composition

Four time-series of copepod species biomass in the north of Spain were contrasted to demonstrate spatial autocorrelation of local communities and their responses to short-term local and regional variability in oceanographic conditions. The series represented coastal and oceanic environments along a marked gradient of influence of seasonal upwelling from Galicia to the Mar Cantábrico (S Bay of Biscay), and each one included at least 10 years of continuous data collected at monthly frequency. Community composition (i.e. species number and diversity) was very consistent through the region, but local variations in the presence of new species and the relative proportions of common species allowed for the characterisation of the response to the environment at each site. Small-sized species were more frequent near the coast. A few species, however, captured the main patterns of variability in all series. Calanus helgolandicus and Acartia (mainly Acartia clausi) were generally the main contributors to total biomass, while other species as Paracalanus parvus and Clausocalanus spp. were important only at some locations. Most copepod indices were positively correlated with upwelling, either considering the whole community (biomass, species richness and diversity) or individual species, but only in the coastal series analysed since 1991. Copepods in the nearby ocean, however, showed negative correlations with upwelling in the period 1960–1986. The effects of upwelling may have been modulated by local factors, as showed by the increases in biomass, number of species and diversity in associations with increases in sea surface temperature in Galicia, while in the Mar Cantábrico only the warming-tolerant species increased and those typical of upwelling decreased. Density stratification of the water column was associated with decreases in total copepod biomass in Galicia, while it favoured the increase in species richness in the Mar Cantábrico. Nearly all significant responses of copepods to environmental variability were delayed by up to 5 months, showing the importance of considering time-lags in the analysis of temporal responses of zooplankton.

Comparative seasonal dynamics of Centropages typicus at seven coastal monitoring stations in the North Sea, English Channel and Bay of Biscay

We review current knowledge and understanding of the biology and ecology of Centropages typicus in the European shelf-seas (e.g. North Sea, English Channel and Bay of Biscay). Our study is based on observations at seven coastal time-series stations as well as on the Continuous Plankton Recorder dataset. This paper focuses on the influence of the environmental parameters (e.g. temperature and Chla) on the life cycle and distribution of C typicus and provides a comparison with its congeneric species C. hamatus and C. chierchiae in the study area. Data on abundance, seasonality and egg production have been used to define the temperature and chlorophyll optima for occurrence and reproduction of Centropages spp. within this region of the European shelf-seas. (C) 2007 Elsevier Ltd. All rights reserved.

Feeding of Calanus finmarchicus and Oithona similis on the microplankton assemblage in the Irminger Sea, North Atlantic

The present investigation reviews published data on the feeding rates and prey selection of Oithona similis females, Calanus finmarchicus nauplii and females in the Irminger Sea in April/May and July/August 2002. Our aim was to examine how the feeding rates and prey selection of these three copepod stages respond to concomitant changes in microplankton community composition and prey abundance. Copepods typically ingested prey overall according to its ambient concentration although significant species and stage-specific differences in prey-type ingestion and selection were apparent. Despite being of comparable weight, the ingestion rates of C. finmarchicus nauplii were always higher than those of the O. similis females. Moreover, C. finmarchicus nauplii and O. similis females fed preferentially on diatoms and ciliates respectively, whereas adult female C. finmarchicus showed limited prey selectivity. Copepod grazing impact on total and on ciliates/dinoflagellates standing stock was <0.5 and <2%, respectively. We attribute this result to a combination of low grazing rates, low copepod abundance and low microplankton biomass, all of which are indicative of the non-bloom conditions under which these experiments were conducted. The differences in copepod feeding rates and prey selection we report reflect species and stage-specific eco-physiological adaptations, which may act as important driving forces for marine ecosystem structuring and functioning.

Morphological and genetic variation in the north Atlantic copepod, Centropages typicus

This study describes phenotypic and genotypic variations in the planktonic copepod, Centropages typicus (Copepoda: Calanoida) that indicate differentiation between geographical samples. We found consistent differences in the morphology of the chela of the sexually modified fifth pereiopod (P5) of male C. typicus between samples from the Mediterranean, western North Atlantic and eastern North Atlantic. A 560 base pairs (bp) region of the C. typicus mitochondrial cytochrome c oxidase subunit I (COI) and a 462 bp fragment of the nuclear rDNA internal transcribed spacer (ITS) tandem array were analysed to determine whether these morphological variations reflect population genetic differentiation. Mitochondrial haplotype diversity was found to be high with 100 unique COI haplotypes among 116 individuals. Analysis of mtCOI variation suggested differentiation between the Mediterranean and Atlantic populations but no separation was detected within the Atlantic. Intragenomic variation in the ITS array suggested genetic differentiation between samples from the western North Atlantic and those from the eastern North Atlantic and Mediterranean. Breeding experiments would be required to elucidate the extent of genetic isolation between C. typicus from the different population centres.

Temporal and spatial variability of Oithona spp. abundance and phenology in the North Atlantic, North Pacific and Southern Ocean using the CPR data

The genus Oithona is considered the most ubiquitous and abundant copepod group in the world oceans. Although they generally make-up a lower proportion of the total copepod biomass, because of their high numerical abundance, preferential feeding for microzooplankton and motile preys, Oithona spp. plays an important role in microbial food webs and can provide a food source for other planktonic organisms. Thus, changes in Oithona spp. overall abundance and the timing of their annual maximum (i.e. phenology) can have important consequences for both energy flow within marine food webs and secondary production. Using the long term data (1954-2005) collected by the Continuous Plankton Recorder (CPR), the present study, investigates whether global climate warming my have affected the long term trends in Oithona spp. population abundance and phenology in relation to biotic and abiotic variables and over a wide latitudinal range and diverse oceanographic regions in the Atlantic, Pacific and Southern Ocean.

Influence of the Pacific Decadal Oscillation on phytoplankton phenology and community structure in the western North Pacific

Phytoplankton phenology and community structure in the western North Pacific were investigated for 2001–2009, based on satellite ocean colour data and the Continuous Plankton Recorder survey. We estimated the timing of the spring bloom based on the cumulative sum satellite chlorophyll adata, and found that the Pacific Decadal Oscillation (PDO)-related interannual SST anomaly in spring significantly affected phytoplankton phenology. The bloom occurred either later or earlier in years of positive or negative PDO (indicating cold and warm conditions, respectively). Phytoplankton composition in the early summer varied depending on the magnitude of seasonal SST increases, rather than the SST value itself. Interannual variations in diatom abundance and the relative abundance of non-diatoms were positively correlated with SST increases for March–April and May–July, respectively, suggesting that mixed layer environmental factors, such as light availability and nutrient stoichiometry, determine shifts in phytoplankton community structure. Our study emphasised the importance of the interannual variation in climate-induced warm–cool cycles as one of the key mechanisms linking climatic forcing and lower trophic level ecosystems.

Spatial variability of marine climate change impacts in the North Atlantic

There is an accumulating body of evidence to suggest that many marine ecosystems in the North Atlantic, both physically and biologically are responding to changes in regional climate caused predominately by the warming of air and sea surface temperatures (SST) and to a varying degree by the modification of oceanic currents, precipitation regimes and wind patterns. The biological manifestations of rising SST and oceanographic changes have variously taken the form of biogeographical, phenological, physiological and community changes. For example, during the last 40 years there has been a northerly movement of warmer water plankton by 10 degree latitude in the north-east Atlantic and a similar retreat of colder water plankton to the north. This geographical movement is much more pronounced than any documented terrestrial study, presumably due to advective processes playing an important role. Other research has shown that the plankton community in the North Sea has responded to changes in SST by adjusting their seasonality (in some cases a shift in seasonal cycles of over six weeks has been detected), but more importantly the response to climate warming varied between different functional groups and trophic levels, leading to mismatch. Therefore, while it has been documented that marine ecosystems in certain regions of the Atlantic have undergone some conspicuous changes over the last few decades it is not known whether this is a pan-oceanic homogenous response. Using these two most prominent responses and/or indicative signals of pelagic ecosystems to hydro-climatic change, changes in species phenology and the biogeographical movement of populations, we attempt to identify vulnerable regional areas in terms of particularly rapid and marked change.

Predator–prey reversal: A possible mechanism for ecosystem hysteresis in the North Sea?

Removal of large predatory fishes from marine ecosystems has resulted in persistent ecosystem shifts, with collapsed predator populations and super-abundant prey populations. One explanation for these shifts is reversals of predator–prey roles that generate internal feedbacks in the ecosystems. Pelagic forage fish are often predators and competitors to the young life stages of their larger fish predators. I show that cod recruitment in the North Sea has been negatively related to the spawning-stock biomass of herring for the last 44 years. Herring, together with the abundance of Calanus finmarchicus, the major food for cod larvae, were the main predictors of cod recruitment. These predictors were of equivalent importance, worked additively, and explained different parts of the dynamics in cod recruitment. I suggest that intensive harvesting of cod has released herring from predator control, and that a large population of herring suppresses cod recruitment through predation on eggs and larvae. This feedback mechanism can promote alternative stable states and therefore cause hysteresis to occur under changing conditions; however, harvesting of herring might at present prevent a shift in the ecosystem to a herring-dominated state.

Transregional linkages in the North-Eastern Atlantic - An ‘end-to-end ’ analysis of pelagic ecosystems

This review examines interregional linkages and gives an overview perspective on marine ecosystem functioning in the north-eastern Atlantic. It is based on three of the 'systems' considered by the European Network of Excellence for Ocean Ecosystems Analysis (EUR-OC EANS was established in 2004 under the European Framework VI funding programme to promote integration of marine ecological research within Europe), the Arctic and Nordic Seas, North Atlantic shelf seas and North Atlantic. The three systems share common open boundaries and the transport of water, heat, nutrients and particulates across these boundaries modifies local processes. Consistent with the EUR-OC EANS concept of 'end-to-end' analyses of marine food webs, the review takes an integrated approach linking ocean physics, lower trophic levels and working up the food web to top predators such as marine mammals. We begin with an overview of the regions focusing on the major physical patterns and their implications for the microbial community, phytoplankton, zooplankton, fish and top predators. Human-induced links between the regional systems are then considered and finally possible changes in the regional linkages over the next century are discussed. Because of the scale of potential impacts of climate change, this issue is considered in a separate section. The review demonstrates that the functioning of the ecosystems in each of the regions cannot be considered in isolation and the role of the atmosphere and ocean currents in linking the North Atlantic Ocean, North Atlantic shelf seas and the Arctic and Nordic Seas must be taken into account. Studying the North Atlantic and associated shelf seas as an integrated 'basin-scale' system will be a key challenge for the early twenty-first century. This requires a multinational approach that should lead to improved ecosystem-based approaches to conservation of natural resources, the maintenance of biodiversity, and a better understanding of the key role of the north-eastern Atlantic in the global carbon cycle.