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.     

The feeding ecology of the copepod Centropages typicus (Kröyer)

Here we report on the current knowledge on the feeding ecology of the planktonic copepod Centropages typicus. We describe the foraging process of C. typicus from the detection of prey to their digestion, considering also the effects of several physical variables on the feeding activity of the species. C. typicus is an omnivorous copepod that feeds on a wide spectrum of prey, from small algae (3–4 μm equivalent spherical diameter, ESD) to yolk-sac fish larvae (3.2–3.6 mm length). It uses both suspensivorous and ambush feeding strategies, depending on the characteristics of the prey. In general, C. typicus exhibits selection for large motile prey, such as ciliates or dinoflagellates, both in nature and laboratory, and this selective pattern is enhanced under moderate intensities of turbulence. Daily rations in the field are somewhat lower than those found in the laboratory, which indicates food limitation. This fact, together with the relatively modest capacity of the species to adapt to fluctuations in food availability may explain the geographical distribution of C. typicus, being restricted to near-shelf waters. In general, it does not seem that C. typicus feeding severely impacts planktonic populations. However, the occasional importance of the species in certain ecosystems is also apparent.     

Reproduction,hatching success,and early naupliar survival in Centropages typicus

The broadcast spawner, Centropages typicus, is a very successful copepod species in many coastal areas of the Mediterranean Sea and North Atlantic Ocean. This review assembles the large amount of information on the reproduction and early life history of C. typicus that has emerged since the 1970s and has made this species one of the best-studied copepods, similar in that regard to species of Acartia and Calanus. Observations on mating behavior and the female gametogenic and oogenic cycles are presented, together with information on seasonal cycles of egg production rates in Mediterranean and Atlantic populations from various regions. These studies indicate a strong latitudinal gradient, with continuous reproduction and the main spawning season occurring earlier (late winter/spring) in warmer waters such as the Mediterranean Sea, compared to northern areas such as the North Sea and in the Kattegat, where C. typicus actively reproduces mainly in late summer and fall with reproduction ceasing altogether in winter in the German Bight. These observations strongly suggest that temperature is the controlling factor for reproductive activity in this species. Egg development times are also temperature dependent but do not vary with latitude, and there is as yet no conclusive evidence that diapause egg production occurs in C. typicus. Laboratory experiments have shown that food quantity and quality both affect fecundity and offspring fitness, but most of these studies have focused on diatom and dinoflagellate diets and non-algal prey have been strongly underrepresented, despite their importance for this omnivorous copepod. Large fluctuations in hatching success and naupliar survival have been reported in field surveys and have subsequently been related to maternal feeding history and food quality or toxicity in laboratory experiments. We identify future lines of research that will help to explain the interannual variability in breeding intensity and recruitment of C. typicus in the field based on better understanding of the species’ reproductive biology.     

Temporal variability of Centropages typicus in the Mediterranean Sea over seasonal-to-decadal scales

Centropages typicus is one of the most common, abundant and best studied calanoid copepods in neritic waters of the Mediterranean Sea, which means it can provide useful information about the long-term dynamics of the Mediterranean epipelagic ecosystem. This paper presents the first comparative overview of the seasonal and long-term variability of C. typicus in different Mediterranean regions. This review is based on quantitative information from the published literature and novel data from five ongoing zooplankton time-series carried out in the Mallorca Island (Balearic Sea), the Bay of Villefranche (Ligurian Sea), the Gulf of Naples (Tyrrhenian Sea), the Gulf of Trieste (North Adriatic Sea), and the Saronikos Gulf (Aegean Sea). In most Mediterranean regions, C. typicus has a perennial occurrence, with peaks of abundance that reflect the succession of different generations. Throughout the Mediterranean, the annual cycle of C. typicus is characterized by minima in winter and major peaks in April–June, which is earlier than those observed in European Atlantic waters, where the peaks are more frequently recorded in summer and fall. In the regions investigated, the annual cycle shows remarkable similarities in terms of timing, but notable differences in the peak height; populations are far more abundant in coastal north-western regions and less abundant in the eastern basin. In the long-term, changes in C. typicus phenology observed in the Bay of Villefranche and in the Gulf of Naples are related to the North Atlantic Oscillation (NAO) index. In these two regions, the species responds to climate forcing similarly in terms of average seasonal patterns (bi-modal patterns in years of positive NAO, unimodal patterns in years of negative NAO) but oppositely in terms of quantity, indicating different influence of the NAO on the two regions. At decadal scales, C. typicus populations show high interannual variability with marked geographical differences. In some areas, the patterns are clearly characterized by alternate phases of higher and lower annual abundances, at higher frequency (mainly 1–2 years) in the Gulf of Naples, and lower frequency in the Saronikos Gulf (mainly 4–5 years) and in the Gulf of Trieste (mainly 5–6 years). Synchronous phases of increasing or decreasing abundance are discernable only for a few sites and short periods, for example from 1998 to 2000 in the Gulf of Naples, Gulf of Trieste and Saronikos Gulf. The regional differences observed in the long-term patterns of C. typicus populations suggest that the temporal dynamics of this species are significantly more affected by local conditions than by any possible common driving force acting at basin scale through teleconnections.     

Metabolism of Centropages species in the Mediterranean Sea and the North Atlantic Ocean

Information on the metabolism rates of Centropages typicus and congeneric species (C. hamatus, C. furcatus, C. brachiatus and C. abdominalis) in neritic areas of the Mediterranean Sea, the North Atlantic Ocean and the Pacific Ocean are reported here. Respiration rates and excretion rates are strongly influenced by abiotic (i.e. temperature, salinity) and biotic factors (i.e. food availability and composition). Differences in the response of respiratory rates to temperature of acclimated, acclimatized and adapted individuals are clearly observed among regions of the Mediterranean Sea and the West and East shores of the Atlantic Ocean. Food supply also strongly affects respiration and excretion rates, as well as the size, sex and stage development of the individuals. The co-measurement of these two rates allows confirmation of the omnivory or carnivory oriented feeding habits of these species. The role of this neritic genus in coastal environment is also discussed.     

Development and growth rates of Centropages typicus

This paper presents an updated review of our knowledge about Centropages typicus development and growth based on a synthesis of all data found in the literature on stage durations, development rates, size, weight, and C:N ratios. Developmental and growth characteristics have been recorded during field time series and cruises, as well as in controlled laboratory experiments. A few laboratory studies have contributed to an understanding of the impact of food quantity, food quality and temperature on growth and development.C. typicus has a sigmoidal development pattern when the contribution of the different stages to total development time is considered, CIV and CV having much longer durations than preceding stages. The development times of this species are closely related to temperature with a nearly equiproportional development. In turn, food conditions, both in quantity and quality, have less effect on development than on growth rate, leading to large variations in weight within stage, even larger than those predicted from changes in length and biovolume. Actual measurements of weight and biochemical composition are required to better understand the growth of C. typicus in relation to food composition. As a consequence, our review shows that weight–length relationships obtained in particular environmental conditions may be inadequate for others.The combination of a temperature-dependent development rate and a food-dependent growth rate may explain why shelf regions are a more favourable environment for C. typicus as compared to offshore regions, as they support large females, production of more and better quality eggs, and probably better survival of the offspring.     

Seasonal and spatial dynamics of Centropages typicus and C. hamatus in the western North Atlantic

Centropages typicus and Centropages hamatus are two of the most abundant copepods on the continental shelf in the mid-Atlantic region of the western North Atlantic. Their range extends from the Scotian Shelf (C. typicus) and the Grand Banks (C. hamatus) in the north to Cape Hatteras in the south. South of Cape Hatteras they have only been observed in inshore waters of North and South Carolina and not offshore on the continental shelf or in coastal waters of eastern Florida. However, C. hamatus has been observed in western Florida. Abundances of both species are greatest in inshore regions in the mid-Atlantic Bight with C. hamatus tending to have a more coastal distribution. In this region seasonal variability is low with high abundances from late fall through mid-summer. In the north year-round presence of both species is confined to inshore areas and offshore banks such as Georges Bank, and Browns Bank, Emerald Bank and Western Bank on the Scotian Shelf. In this northern region there is a pronounced seasonal cycle in abundance with high abundances during late summer and fall. Periods of high reproductive rates are closely linked to blooms of large phytoplankton and food availability rather than temperature appears to be controlling population abundances.     

The biology and ecology of Centropages typicus: An introduction

We briefly introduce the different chapters of this special issue which review the current knowledge and understanding of the biology and ecology of Centropages typicus obtained over the last 30 years. The papers are grouped into two major theme areas: the first five papers review behaviour, feeding, metabolism, growth and development, and reproduction, and the four following papers have a regional focus dealing with temporal and spatial distribution in the Mediterranean Sea and in the North Atlantic shelf areas. The findings reviewed in the two groups of papers converge to explain why C. typicus is such a very successful copepod species in these coastal areas. In the different chapters, similarities and differences with congeneric species (C. velificatus, C. hamatus, C. brachiatus, C. chierchiae, C. furcatus, and C. abdominalis) have been explored where these are known.     

Food-web structure and trophodynamics of mesopelagic–suprabenthic bathyal macrofauna of the Algerian Basin based on stable isotopes of carbon and nitrogen

The trophodynamics of mesopelagic (macrozooplankton/micronekton) and benthic boundary layer (suprabenthos=hyperbenthos) faunas from the Algerian Basin were characterized on a seasonal scale through stable carbon and nitrogen isotopic analyses of a total of 34 species and two broad taxa (Copepoda and Cumacea). This is the first study simultaneously focused on trophodynamics of deep-sea zooplankton and suprabenthos. Samples were collected southeast of Mallorca (Algerian Basin, Western Mediterranean), on the continental slope close to Cabrera Archipelago, at 650–780 m depths, ca. bi-monthly between August 2003 and June 2004. Mean δ¹³C values of suprabenthos ranged from ?21.1‰ (Munnopsurus atlanticus) to ?16.7‰ (Cyclaspis longicaudata). Values of δ¹⁵N ranged from 2.8‰ (Lepechinella manco) to 9.9‰ (larvae of Gnathia sp.). The stable isotope ratios of suprabenthic fauna displayed a continuum of values, confirming a wide spectrum of feeding guilds (from filter feeders/surface deposit feeders to predators). According to the available information on diets for suprabenthic species, the highest annual mean δ¹⁵N values were found for the hematophagous isopod Gnathia sp. parasite on fish (represented by Praniza larvae) and carnivorous amphipods (e.g. Rhachotropis spp., Nicippe tumida) consuming copepods, and the lowest δ¹⁵N values were found for two cumaceans (Cyclaspis longicaudata and Platysympus typicus) feeding on detritus. Assuming a ¹⁵N-enrichment factor of 2.5‰ and deposit feeders as baseline, we found three trophic levels in suprabenthic food webs. δ¹³C ranges were particularly wide among deposit feeders (ranging from ?21.8% to ?17.3‰) and omnivores (from ?20.5% to ?18.8‰), suggesting exploitation of particulate organic matter (POM) of different characteristics. Our isotopic analyses revealed lower ranges of δ¹³C and δ¹⁵N for macrozooplankton/micronekton, compared with suprabenthos. δ¹³C values of zooplankton taxa ranged from ?21.1‰ (the hyperiid Phrosina semilunata) to ?19.9‰ (the decapod Pasiphaea multidentata), while δ¹⁵N values ranged from 3.9‰ (P. semilunata) to 7.5‰ (P. multidentata). Among zooplankton, more enriched δ¹⁵N values were found among carnivores (e.g. the fish Cyclothone spp. and Pasiphaea multidentata) preying on copepods, hyperiids, euphausiids and small fish. The lowest δ¹⁵N values were found for hyperiids that feed on the mucus nets of salps (e.g. Vibilia armata). After contrasting isotope analysis with dietary data, we conclude there were two trophic levels among zooplankton/micronekton. Strong correlation between the mean annual δ¹⁵N and δ¹³C values was found for zooplankton (R²=0.7), but not for suprabenthos, which suggests a single source of carbon for plankton. We found a general seasonal trend for δ¹³C enrichment from late autumn (November) to late winter–spring (February–April) for both suprabenthos and zooplankton. The δ¹³C enrichment in February–April was correlated in zooplankton with higher surface chlorophyll a concentration 1 month before sampling. As evidenced by δ¹³C–δ¹⁵N correlations, the response of zooplankton to the peak of surface primary production was almost immediate (an increase of δ¹³C–δ¹⁵N correlations in February), and stronger than for suprabenthos. The response among suprabenthos was weak, with slight increase in δ¹³C–δ¹⁵N relationships in April–June.     

Food preferences and rhythms of feeding activity of two co‐existing demersal fish,the longspine snipefish,Macroramphosus scolopax (Linnaeus, 1758), and the boarfish Capros aper (Linnaeus, 1758), on the Mediterranean deep shelf

The feeding behaviour of two potentially competing species, the longspine snipefish, Macroramphosus scolopax, and the boarfish Capros aper was examined. While both species are very abundant along the Mediterranean coast and are regularly caught by demersal trawlers, they are of no commercial value. The diets of boarfish and longspine snipefish were investigated from samples collected between January 2001 and May 2002. Variations in the diet with fish size and season, as well as diet overlap and diversity were explored. Mysid shrimps, amphipods and gastropods were the most important food items in the diet of longspine snipefish. During ontogenetic development, M. scolopax occupies different trophic levels: the diet shifts from being predominantly composed of mysids (Anchialina agilis, Lophogaster typicus, Erythrops sp., Leptomysis spp.) in the smaller longspine snipefish [<6.5 cm total length (TL)] towards decapods (Anapagurus laevis) and amphipods (Leucothoe incisa, Eusirus longipes, Hyperidea) in the larger individuals (>6.5 cm TL). Crustacean decapods and copepods were the most important prey in the stomachs of boarfish. Mysids (Lo. typicus), euphausiids and nematodes were present in the larger individuals (>8 cm TL). A more generalist diet, still containing copepods, crustacean decapods, gastropods (Limacina retroversa) and a large variety of amphipods (e.g. Phtysica marina, Stenotoe bosphorana) and mysids (e.g. A. agilis, Leptomysis spp., Erythrops sp.), dominated the diet of C. aper between 2 and 8 cm TL. Diet overlap between longspine snipefish and boarfish was very low and the differences in stomach species diversity were explained by season and fish size.     

Note on the CO2 air–sea gas exchange at high temperatures

The temperature dependency of ocean–atmosphere gas transfer velocities is commonly estimated in terms of Schmidt numbers, i.e. the ratio of kinematic viscosity to diffusivity. In numerical models least square regressions are used to fit the limited number of experimentally derived Schmidt numbers to a function of temperature. For CO₂ a well established fit can be found in the literature. This fit constitutes an integral part in standardized carbon cycle simulation projects (e.g. C4MIP, OC4MIP, Friedlingstein et al., 2006). However, the fit is valid only in the range where diffusivity measurements exist, i.e., from 0 to about 30 °C. In many climate warming simulations like e.g. the MPI contribution to the fourth Intergovernmental Panel on Climate Change Assessment Report (IPCC AR 4), sea surface temperatures largely exceed the validated range and approach or even reach the range, where the standard fits leave the physically meaningful range. Thus, this paper underlines the demand for new measurements of seawater diffusivities for CO₂ and other trace gases especially for the temperature range >30 °C.In this paper we provide improved fits for the temperature dependence of the Schmidt number. For carbon dioxide our fit is compared to the established fit under identical climate change simulations carried out with the 3D-carbon cycle model HAMOCC. We find that in many tropical and subtropical high temperature regions the established fit leads to unrealistically short adaption times of the surface water pCO₂ to altered atmospheric pCO₂. In regions where the local oceanic pCO₂ is not primarily controlled by the atmospheric boundary pCO₂ but by other processes such as biological activity, the atmosphere ocean pCO₂ gradient is clearly underestimated when using the established fit. The effect on global oceanic carbon uptake in a greenhouse world is rather small and the potential climate feedback introduced by this bias seems to be negligible. However, the bias will clearly gain in significance the more regions warm up to over 30 °C. On a regional scale, especially in coastal regions at low latitudes, the effect is not negligible and a different steady state is approached.     

Trophic ecology of European sardine Sardina pilchardus and European anchovy Engraulis encrasicolus in the Bay of Biscay (north-east Atlantic) inferred from δ13C and δ15N values of fish and identified mesozooplanktonic organisms

European sardine (Sardina pilchardus) and European anchovy (Engraulis encrasicolus) are two species of economical and ecological significance in the Bay of Biscay (north-east Atlantic). However, the trophic ecology of both species is still poorly known in the area, and more generally, few studies have considered the potential trophic overlap between sardines and anchovies worldwide. This study aims to highlight the trophic links between the mesozooplankton and adults of these two pelagic fish in the Bay of Biscay, through carbon and nitrogen stable isotope analysis (SIA). Mesozooplankton and individuals of sardines and anchovies were collected during one season (spring 2010), over spatially contrasted stations within the study area. First, the potential effect of preservation (ethanol vs. freezing) and of delipidation (by cyclohexane) on mesozooplankton δ¹³C and δ¹⁵N values was assessed. Results demonstrated the necessity to correct for the preservation effect and for lipid contents in mesozooplankton for further analyses of sardines' and anchovies' diet through SIA. Next, this study highlighted the interest of working on identified mesozooplanktonic organisms instead of undetermined assemblages when unravelling food sources of planktivorous fish using stable isotopes. The inter-specific variability of isotope values within a planktonic assemblage was effectively high, probably depending on the various feeding behaviours that can occur among mesozooplankton species. Intra-specific variability was also significant and related to the spatial variations of baseline signatures in the area. To investigate the foraging areas and potential diet overlap of S. pilchardus and E. encrasicolus, mixing models (SIAR) were applied. Both fish species appeared to feed mainly in the neritic waters of the Bay of Biscay in spring and to select mainly small- to medium-sized copepods (e.g. Acartia sp., Temora sp.). However, E. encrasicolus showed a greater trophic plasticity by foraging more offshore and on a wider range of prey sizes, while S. pilchardus seemed more limited to coastal areas and the mesozooplanktonic species of these waters for feeding.     

A communicative turnaround: Shifting the burden of proof in European fisheries governance

Current and prospective changes in European fisheries governance suggest not only a “communicative turn” but a complete turnaround in the relationships between government, science, and the fishing industry. At the heart of these changes are the so-called Regional Advisory Councils (RACs) and the idea of partially replacing the burden of proof on the resource users (fishing industry). This change entails new forms of interaction between fishers’ representatives, other stakeholders (e.g. NGOs), policy-makers and scientists. By drawing on experiences from the Baltic Sea RAC, the analysis focuses on two aspects of fisheries governance: institutional design and the process of negotiation and decision-making. It is concluded that to allow for a partial shift in the burden of proof, stakeholder organisations such as RACs need to adapt both institutionally as well as process-wise to enable a more constructive and responsible fisheries governance system.     

Inter-decadal variability in zooplankton and phytoplankton abundance on the Newfoundland and Scotian shelves

Continuous Plankton Recorder (CPR) sampling on the Newfoundland and Scotian shelves covers three multi-year periods characterised by negative (1962–1971), positive (1992–2000) and negative/neutral (2001–2003) values of the NAO index. Water temperatures respond differently to changes in the NAO in different regions: a positive NAO index tends to lead to reduced temperatures on the Newfoundland shelf and to increased temperatures on the central/western Scotian shelf, and a negative NAO index to the reverse. Since the 1960s, the hydrographic changes due to the NAO have been superimposed on a freshening of the water column throughout the region, which is attributed to increased contribution of Arctic water outflow. Changes in plankton abundance measured by the CPR for the three time periods were generally, but not always, similar on the Newfoundland and eastern and western regions of the Scotian shelf, although Arctic species (e.g. Calanus glacialis, Calanus hyperboreus) were notably more abundant and warm water species (e.g. Metridia lucens, euphausiids) less abundant on the Newfoundland shelf than on the Scotian shelf. Three categories of phytoplankton (colour, diatoms, dinoflagellates) increased in abundance in the 1990s, and these increases generally persisted into 2001–2003. This is believed to be a response to the persistent freshening of the water column, probably due to increased stratification. The Arctic species C. glacialis and C. hyperboreus also showed persistent increases in abundance after 1992, perhaps due to increased transport from the Arctic, although the abundance of the Arctic slope water species Metridia longa decreased. Two groups, Calanus 1–4 and euphausiids, both thought to play important roles in the food chain, showed persistent decreases in abundance after 1992, especially on the Newfoundland shelf. In all regions, Calanus finmarchicus 5–6, Oithona spp. and Centropages hamatus abundance changed in association with variations in the NAO, although no common mechanism could be identified. C. finmarchicus 5–6 abundance decreased in the 1990s and increased after 2001, while the other two species showed the opposite pattern. Centropages typicus and M. lucens abundance on the Scotian shelf increased with rising temperature. This is attributed to increased production rates for the former and an increased influx of warm, M. lucens-rich, slope water on to the shelf for the latter. A comparison between ring net and CPR sampling on the Newfoundland shelf suggests that the Calanus 1–4 category is dominated by C. finmarchicus and that late stage C. glacialis and C. hyperboreus are grossly under-sampled compared to late stage C. finmarchicus.     

Growth dynamics of the deep-water Asian eelgrass, <Emphasis Type="Italic">Zostera asiatica</Emphasis>, in the eastern coastal waters of Korea

Among the seagrasses that occur along the coast of Korea, Zostera asiatica inhabits the deepest depth; however, to date, there is limited information on its ecology. This study presents the first quantitative data on the seasonal growth dynamics of Z. asiatica in Korea. We measured seasonal growth and morphological characteristics, as well as environmental factors, including underwater irradiance, water temperature, salinity and nutrient concentrations of the water column and sediment pore water, bimonthly from July 2012 to May 2015. Underwater irradiance showed clear seasonal trends, increasing in the spring and summer and decreasing in the fall and winter, ranging from 2.4 ± 0.2 mol photons m^-2 d^-1 in November 2012 to 12.8 ± 1.3 mol photons m^-2 d^-1 in July 2014. Water temperature also followed a strong seasonal trend similar to underwater irradiance, ranging from 9.8 ± 0.1°C in January 2013 to 20.5 ± 0.2°C in September 2013. Nutrient availability fluctuated substantially, but there was no evidence of distinct seasonal variations. Shoot density, biomass, leaf productivity, and morphological characteristics of Z. asiatica exhibited significant seasonal variations: maximum values of these variables occurred in summer, and the minima were recorded in winter. Total shoot density was highest (218.8 ± 18.8 shoots m^-2) in July 2012 and lowest (106.3 ± 6.3 shoots m^-2) in January 2013. Total biomass ranged from 182.6 ± 16.9 g dry weight (DW) m^-2 in January 2015 to 310.9 ± 6.4 g DW m^-2 in July 2014.Areal leaf production was highest (4.9 ± 0.0 g DW m^-2 d^-1) in July 2012 and lowest (1.4 ± 0.2 g DW m^-2 d^-1) in January 2013. The optimum water temperature for the growth of Z. asiatica was between 16-19°C. Growth of Z. asiatica was more strongly correlated with underwater irradiance than water temperature, suggesting that light is the most important factor determining seasonality of Z. asiatica at the study site.     

Use of formation pressure test results over a hydrate interval for long-term production forecasting at the Mount Elbert Gas Hydrate Stratigraphic Test Well, Alaska North Slope: Implications of uncertainties

In the first part of this paper, the pressure and temperature measurements obtained using Schlumberger's Modular Formation Dynamics Tester (MDT) conducted on the C2 interval in the BPXA-DOE-USGS Mount Elbert Gas Hydrate Stratigraphic Test Well (Mount Elbert Well) are history matched, with the following three objectives: (i) to obtain a better understanding of hydrate decomposition and its reformation as conditions cross the p/T stability, (ii) to obtain formation properties (e.g., permeability) that are consistent with the measurements, and (iii) to explore the non-uniqueness in the history match; i.e., to explore the ranges of parameters that allow a reasonable match of the measured quantities. In the second part of this paper, long-term production performance is predicted, and the effect of the uncertain parameters on the predictions is demonstrated. The results are used to demonstrate the range of long-term production that may be expected, when a model is calibrated using the MDT data. Usefulness of short-term tests for long-term forecast prediction is then discussed.     

Centropages behaviour: Swimming and vertical migration

The evolutionary success of any species living in a variable environment depends on its capacity to enhance the probability of finding food and mates, and escaping predators. In the case of copepods of the genus Centropages, as in all planktonic copepods, their swimming behaviour is closely tied to these vital aspects, and shows a high degree of plasticity and adaptive capacity. Swimming mechanisms of Centropages change radically during development, mainly in the transition between naupliar stages to the 1st copepodite; nauplii do not produce feeding currents, whereas copepodites do. Adults and late developmental stages of C. typicus, C. hamatus and C. velificatus spend most of the time in slow swimming and resting breaks, with occasional and brief fast swimming (escape reactions) and grooming events. Slow swimming is closely related to the creation of feeding currents, and results from the beating of the cephalic appendages in a “fling and clap” manner. The proportion of time allocated to the different swimming activities depends on sensory cues like type and concentration of food, presence of potential mates, light intensity, hydrodynamic flow, etc.The responses of Centropages to changes in flow velocity fluctuations (small-scale turbulence) are similar to the escape responses (fast swimming) triggered by the presence of potential predators. Centropages generally have standard nocturnal vertical migration patterns involving considerable vertical displacements. This behaviour is closely related to the narrow spectral sensitivity and the low intensity threshold of the genus, and has important consequences for the active vertical transport of matter and energy.The variety of responses of Centropages to environmental changes, and in general all the aspects related to its swimming behaviour seem to be controlled by the trade-off between energetic gains (food intake), losses (swimming energy expenditure), and predation risk. Behavioural plasticity and adaptation appear to be the most relevant characteristics for the success of the genus in a wide range of marine environments.     

An overview of Calanus helgolandicus ecology in European waters

We review current knowledge and understanding of the biology and ecology of the calanoid copepod Calanus helgolandicus in European waters, as well as provide a collaborative synthesis of data from 18 laboratories and 26 sampling stations in areas distributed from the northern North Sea to the Aegean and Levantine Seas. This network of zooplankton time-series stations has enabled us to collect and synthesise seasonal and multi-annual data on abundance, body size, fecundity, hatching success and vertical distribution of C. helgolandicus. An aim was to enable comparison with its congener Calanus finmarchicus, which has been studied intensively as a key component of European and north east Atlantic marine ecosystems. C. finmarchicus is known to over-winter at depth, whereas the life-cycle of C. helgolandicus is less well understood. Overwintering populations of C. helgolandicus have been observed off the Atlantic coast between 400 and 800 m, while in the Mediterranean there is evidence of significant deep-water populations at depths as great as 4200 m. The biogeographical distribution of C. helgolandicus in European coastal waters covers a wide range of habitats, from open ocean to coastal environments, and its contribution to mesozooplankton biomass ranges from 6% to 93%. Highest abundances were recorded in the Adriatic and off the west coast of Spain. C. helgolandicus is generally found in 9-20 °C water, with maximum abundances from 13-17 °C. In contrast, C. finmarchicus is found in cooler water between 0 and 15 °C, with peak abundances from 0 to 9 °C. As water has warmed in the North Atlantic over recent decades, the range of C. helgolandicus and its abundance on the fringes of its expanding range have increased. This review will facilitate development of population models of C. helgolandicus. This will not only help answer remaining questions but will improve our ability to forecast future changes, in response to a warming climate, in the abundance and distribution of this important species.