A numerical study of stochastic larval settlement in the California Current system

Key to the predictive understanding of many nearshore marine ecosystems is the transport of larvae by ocean circulation processes. Many species release thousands to billions of larvae to develop in pelagic waters, but only a few lucky ones successfully settle to suitable habitat and recruit to adult life stages. Methodologies for predicting the larval dispersal are still primitive, and simple diffusive analyses are still used for many important applications. In this study, we investigate mechanisms of larval dispersal using idealized simulations of time-evolving coastal circulations in the California Current system with Lagrangian particles as models for planktonic larvae. Connectivity matrices, which describe the source-to-destination relationships for larval dispersal for a given larval development time course, are used to diagnose the time–space dynamics of larval settlement. The resulting connectivity matrices are shown to be a function of several important time scales, such as the planktonic larval duration, the frequency and duration of larval release events and inherent time scales for the coastal circulations. Many important fishery management applications require knowledge of fish stocks on a year-to-year or generation-to-generation basis. For these short time scales (typically less than 1 year), larval dispersal is generally far from a simple diffusive process and the consideration of the stochastic and episodic nature of larval dispersal is required. This work provides new insights into the spatial–temporal dynamics of nearshore fish stocks.     

Primary productivity of the Palmer Long Term Ecological Research Area and the Southern Ocean

A major objective of the Palmer Long Term Ecological Research (Palmer LTER) project is to obtain a comprehensive understanding of the various components of the Antarctic marine ecosystem. Phytoplankton production plays a key role in this so-called high nutrient, low chlorophyll environment, and factors that regulate production include those that control cell growth (light, temperature, and nutrients) and those that control cell accumulation rate and hence population growth (water column stability, grazing, and sinking). Sea ice mediates several of these factors and frequently conditions the water column for a spring bloom which is characterized by a pulse of production restricted in both time and space. This study models the spatial and temporal variability of primary production within the Palmer LTER area west of the Antarctic Peninsula and discusses this production in the context of historical data for the Southern Ocean. Primary production for the Southern Ocean and the Palmer LTER area have been computed using both light-pigment production models [Smith, R.C., Bidigare, R.R., Prézelin, B.B., Baker, K.S., Brooks, J.M., 1987. Optical characterization of primary productivity across a coastal front. Mar. Biol. (96), 575–591; Bidigare, R.R., Smith, R.C., Baker, K.S., Marra, J., 1987. Oceanic primary production estimates from measurements of spectral irradiance and pigment concentrations. Global Biogeochem. Cycles (1), 171–186; Morel, A., Berthon, J.F., 1989. Surface pigments, algal biomass profiles and potential production of the euphotic layer—relationships reinvestigated in view of remote-sensing applications. Limnol. Oceanogr. (34), 1545–1562] and an ice edge production model [Nelson, D.M., Smith, W.O., 1986. Phytoplankton bloom dynamics of the western Ross Sea ice edge: II. Mesoscale cycling of nitrogen and silicon. Deep-Sea Res. (33), 1389–1412; Wilson, D.L., Smith, W.O., Nelson, D.M., 1986. Phytoplankton bloom dynamics of the Western Ross Sea ice edge: I. primary productivity and species-specific production. Deep-Sea Res., 33, 1375–1387; Smith, W.O., Nelson, D.M., 1986. Importance of ice edge phytoplankton production in the Southern Ocean. BioScience (36), 251–257]. Chlorophyll concentrations, total photosynthetically available radiation (PAR) and sea ice concentrations were derived from satellite data. These same parameters, in addition to hydrodynamic conditions, have also been determined from shipboard and Palmer Station observations during the LTER program. Model results are compared, sensitivity studies evaluated, and productivity of the Palmer LTER region is discussed in terms of its space time distribution, seasonal and interannual variability, and overall contribution to the marine ecology of the Southern Ocean.     

Spatial fisheries ecology: Recent progress and future prospects

We review recent progresses made in the study of fish distribution and survival over space — i.e., fisheries spatial ecology. This is achieved by first surveying the most common statistical approaches and relative challenges associated with the analysis of fisheries spatial data, loosely grouped in geostatistical and regression approaches. Then we review a selected number of case-studies implementing the discussed techniques. We conclude by proposing new areas of statistical and ecological research to further our understanding of how fish distribute and survive in space. This review serves a dual purpose by emphasizing the scientific importance of studying spatial interactions to better understand the temporal dynamics of fish abundance, and by promoting the development of new analytical and ecological approaches for the analysis of spatial data. Through our survey we cover different statistical techniques, marine ecosystems and life stages. This analytical, geographic and ontogenetic variety is also purposely selected to highlight the importance of comparative and multidisciplinary studies across diverging ecological disciplines, ecosystems and life stages. Besides having a general ecological relevance this review also bears a more applied significance, owing to the increasing need for protecting renewable marine resources along with their primary habitat.     

Mesoscale high-frequency variability in the Alboran Sea and its influence on fish larvae distributions

This work analyses a multidisciplinary data set including hydrological and meteorological data, satellite images, and fish larvae abundance from a high-frequency experiment conducted along a north–south transect across the Western Alboran Sea anticyclonic gyre. Four consecutive occupations of the transect, crossing the frontal area, evidenced the high-frequency variability of hydrological structures and its influence on the latitudinal and vertical distribution of fish larvae in a period of a few days. The influence of dynamical processes on fish larvae depends on the location of the spawning as well as on the larval fish position in the water column. Wind induced upwelling and/or the southward drift of the Atlantic current transport larvae of neritic species, such as Sardina pilchardus and Engraulis encrasicolus to open sea areas. At the same time, these events bring about alterations in the latitudinal and vertical distributions of mesopelagic fish larvae. The species with a relatively surface distribution, as Ceratoscopelus maderensis, were transported and accumulated to the south of the Atlantic Jet (AJ), while those with a deeper distribution in the water column, as Maurolicus muelleri or Benthosema glaciale, would be upwelled and concentrated to the north. This study shows that it is difficult to extract conclusions about the mean or preferential distributions of fish larvae when one single/synoptic survey is considered and that time evolution of hydrological structures has to be considered in order to conclude.     

Nutrient cycling and primary production in the marine systems of the Arctic and Antarctic

Primary production events in both the Arctic and the Antarctic are highly localized. Carbon-14 incubations that did not account for this caused antarctic primary production estimates to be revised too far downwards from the historic view of high productivity. The primary production regime in the Arctic is even more heterogeneous than in the Antarctic. Arctic primary production rates are in the process of being revised upwards because of a better spatial and temporal distribution of incubation experiments and a re-awakening of interest in estimating new production from the distribution of chemical variables. Similarly, recent examination of temporal changes in nitrate concentrations and recognition of the importance of ice-edge blooms has caused antarctic primary productivity to be revised upwards. In both the Arctic and the Antarctic, the ratio of “new” to total primary production is high, and neglect of this fact can lead to an underestimation of the potential that these regions have for influencing global cycles of bioactive chemicals. Some recent data on temporal changes in nitrate from Fram Strait emphasize the poor state of our knowledge by suggesting an unexpectedly high “new” production rate of 1 g C m⁻² d⁻¹ for a 35 day experiment that encountered an early Phaeocystis bloom. Chemical distributions suggest that new production over the shelf seas that border the Polar Basin is about 50 g Cm⁻² yr⁻¹.The shelves in the Arctic Ocean's marginal and adjacent seas comprise 25% of the total global continental shelf. These extensive shallow regions have much higher rates of primary production than the Polar Basin and may be globally significant sites of denitrification. Globally significant silica deposition could occur on these shelves or on the adjacent slopes.Because of the differences in geomorphology and stratification, global warming is likely to increase primary production in the Arctic and will probably decrease antarctic primary production.In addition to sharing high ratios of “new” to total primary production, high ammonium concentrations occur in the Arctic and Antarctic. It is possible that these accumulations arise from a strong repression of nitrification at low temperatures.     

Analysis of the parasitic copepod species richness among Mediterranean fish

The Mediterranean ichthyofauna is composed of 652 species belonging to 405 genera and 117 families. Among these, 182 were studied for their parasitic copepods. The analysis of all the works conducted on these crustacea yielded 226 species distributed in 88 genera and 20 families. For each fish species we have established a file providing the species name of the fish, its family, its geographical distribution within the Mediterranean and some of its bio-ecological characteristics. Within each file, all the parasitic copepod species reported on each host species were listed. This allowed to know the species richness (SR) of these hosts. We thus produced 182 files within which 226 copepod species are distributed. A program was created under the Hypercard software, in order to analyse our data. Two parameters were studied. The first one is the mean species richness (MSR), which corresponds to the mean of the different SR found on the different host species. The second is the parasite–host ratio (P/H), which is the ratio of the number of copepod species by the number of host species. These parameters are calculated by our program for all the 182 species of Mediterranean fishes retained in our investigation, on the first hand, and, on the second hand, for one particular group of fish species. We used the following variables to investigate their correlations with copepod species richness: taxonomy—fish families, genera and species; biometry—maximal size of the adult fish; eco-ethology—mode of life (benthic, pelagic or nectonic), displacements (sedentary, migratory with environmental change, or migratory without environmental change), behaviour (solitary or gregarious). Other variables (colour, food, reproduction, abundance, distribution area) were also analysed but did not reveal any clear correlation. Providing that our study does not rely on quantitative (prevalence, intensity) but qualitative basis our aim was only to reveal some tendencies. These tendencies are as follows: (1) In many cases, parasite and host phylogeny seem to play an important role. There are fish families with copepods and families with few species of these parasites. The phyletic constraints could be due to the morphological characteristics of the habitat (e.g. structure of the gills) or biological/ecological characteristics that we were unable to identify. (2) It appears that the presence in a same environment of related fish species (e.g. several species of the same genus, or numerous genera of the same family) is correlated with high parasite richness. A likely explanation is that such situations favours alternated processes of lateral transfers and speciation. (3) Some eco-ethological criteria seem to favour the establishment of a large parasite species richness. It should be noted for instance that Mediterranean fishes the most often infected with copepods are generally nectonic or pelagic, migratory, and gregarious species.     

Decadal-scale variations of trophic levels at high trophic levels in the Yellow Sea and the Bohai Sea ecosystem

A total of 2759 stomachs collected from a bottom trawl survey carried out by R/V “Bei Dou” in the Yellow Sea between 32°00 and 36°30N in autumn 2000 and spring 2001 were examined. The trophic levels (TL) of eight dominant fish species were calculated based on stomach contents, and trophic levels of 17 dominant species in the Yellow Sea and the Bohai Sea reported in later 1950s and mid-1980s were estimated so as to be comparable. The results indicated that the mean trophic level at high trophic levels declined from 4.06 in 1959–1960 to 3.41 in 1998–1999, or 0.16–0.19·decade^− 1 (mean 0.17·decade^− 1) in the Bohai Sea, and from 3.61 in 1985–1986 to 3.40 in 2000–2001, or 0.14·decade^− 1 in the Yellow Sea; all higher than global trend. The dominant species composition in the Yellow Sea and the Bohai Sea changed, with the percentage of planktivorous species increases and piscivorous or omnivorous species decreases, and this was one of the main reasons for the decline in mean trophic level at high tropic levels. Another main reason was intraspecific changes in TL. Similarly, many factors caused decline of trophic levels in the dominant fish species in the Yellow Sea and the Bohai Sea. Firstly, TL of the same prey got lower, and anchovy (Engraulis japonicus) as prey was most representative. Secondly, TLs of diet composition getting lower resulted in not only decline of trophic levels but also changed feeding habits of some species, such as spotted velvetfish (Erisphex pottii) and Trichiurus muticus in the Yellow Sea. Thirdly, species size getting smaller also resulted in not only decline of trophic levels but also changed feeding habits of some species, such as Bambay duck (Harpodon nehereus) and largehead hairtail (Trichiurus haumela). Furthermore, fishing pressure and climate change may be interfering to cause fishing down the food web in the China coastal ocean.     

The importance of oceanographic fronts to marine birds and mammals of the southern oceans

During the last 30 years, at-sea studies of seabirds and marine mammals in the oceans south of the Subtropical Front have described an association with major frontal areas. More recently, the advancement in microtechnology has allowed the tracking of individuals and investigations into how these marine predators actually use the frontal zones. In this review, we examine 1) the relative importance to apex predators of the different frontal zones in terms of spatial distribution and carbon flux; 2) the processes that determine their preferential use; and 3) how the mesoscale dynamics of frontal structures drive at-sea foraging strategies of these predators. We review published results from southern waters and place them in a broader context with respect to what has been learned about the importance of fronts in oceans farther north.Some fronts constitute important boundaries for seabird communities in southern waters. At a mesoscale the maximum values of seabird diversity and abundance correspond to the location of the main fronts. At-sea surveys show a strong curvilinear correlation between seabird abundance and sea surface temperatures. High mean species richness and diversity for whales and seabirds are consistently associated with the southern water mass boundary of the Antarctic Circumpolar Current, the Subtropical Front and the Subantarctic Front; in the case of the Polar Front mean seabird densities are more variable. At small-scales, variation in seabird occurrence has been directly related to the processes at fronts in a limited number of cases. A significant positive relation was found between some plankton feeding species and frontal temperature gradient–phytoplankton variables.Telemetric studies have revealed that several apex predators (penguins, albatrosses, seals) perform long, directed foraging trips either to the Subtropical front or Polar Front, depending on locality. Seabirds with low flight costs, such as albatrosses, are able to reach fronts at long distances from colonies, showing variable foraging strategies as a function of the distances involved. Diving birds such as King penguins, that travel at a higher cost and lower speed, rely on the predictable spatial distribution of mesopelagic fish found close to the Polar Front. They may use the currents associated with eddies as oceanographic cues in the active search for frontal zones. Once in these areas they dive preferentially in and below the depth of the thermocline where catches per unit effort are high. Elephant seals concentrate foraging activity principally inside or at the boundary of cyclonic eddies. These mesoscale features appear to offer exceptional productivity favourable for foraging by various diving top predators.The connection between biophysical parameters at fronts and predators is likely to be made through biological enhancement. Top predators appear to forage at locations where prey are advected by physical processes and others where prey are produced locally. Long-term research on at-sea distributions and demographic parameters of top predators are essential to assess the consequences of potential shift in front distributions in relation to global warming. Such environmental changes would add to the impact of fish extraction by the industrial fisheries on the southern food webs.     

Morphological and biochemical differentiation in Antarctic krill

During the February 1981 cruise FIBEX MD-25 between 30–50°E and 61–64°S, hydrography showed the presence of two gyres, confirmed by the geostrophic circulation relative to 1000 m from Levitus climatology, at the borders of these gyres concentrations of highly morphologically differentiated krill were found. Gaussian component analysis of krill samples, pooled by sectors, showed three cohorts of Euphausia superba in the western sector and one in the eastern sector. Across the sampling area, Thysanoessa macrura and E. superba occurred at separate stations. Analysis of cohorts in T. macrura separated two size groups in both the western and the eastern sectors. The use of a Differentiation Index (D.I.) [Färber-Lorda, J., 1990. Somatic length relationships and ontogenetic morphometric differentiation of Euphausia superba and Thysanoessa macrura of the southwest Indian Ocean during summer (February 1981). Deep-Sea Res. 37, 1135–1143.], based on somatic lengths, allows studying certain morphological differences within the populations sampled. Morphologically different and bigger males II (D.I. from 2.8 to 3.5) were present only in the southern transect while smaller males I (D.I. from 3.5 to 5.0) were present over the entire area. Biochemical composition of both species showed significant differences among stations for protein, lipids, and carbohydrates. A significant difference in lipid content was found between males I, and males II. For T. macrura, percentage of lipid content in mature animals was much higher than that in E. superba. The D.I. size distribution showed that when populations of E. superba were highly differentiated (corresponding to mature animals) in morphology, lipid content was high, and they were located near a gyre. Differences in morphometry can influence distribution of the species, because different developing stages have different swimming capacities. It is shown that, together with hydrography and trophic conditions, lipid content and morphometry of krill populations, are different but complementary aspects that help to understand krill ecology and distribution.     

Biochemical composition of marine sediment from the eastern Weddell Sea (Antarctica): High nutritive value in a high benthic-biomass environment

Within the SCAR's international EASIZ programme, as part of the benthic–pelagic coupling experiment, grain size and organic matter contents in marine surface sediment were measured. Samples were taken during the austral autumn of 2000 from 3 regions in the eastern Weddell Sea: Kapp Norvegia, Four Seasons Bank, and Austasen.In general, sediments were fine sand with a grain size fraction < 200 μm representing more than 40% of the total weight. The sediments from Four Seasons Bank (64 to 107 m depth) were coarser than those from Austasen and Kapp Norvegia (209 to 480 m depth), presumably due to winnowing of fine sediment at shallow depths. Organic carbon (OC) content ranged from 0.25% to 1.2% and constituted 10% to 97% of the total carbon. The samples from Kapp Norvegia presented the highest OC values. Overall, protein (PRT), lipid (LPD), and carbohydrate (CHO) contents were similar to those in sediment from cold regions (e.g., the North Atlantic and the Ross Sea) but higher than those in sediment from other Antarctic and more septentrional regions (e.g., the Ross Sea and the Mediterranean). The difference within the Antarctic is explained through the local conditions in Terra Nova Bay and Kapp Norvegia. In the Antarctic, PRT and LPD carbon were the main contributors to the biopolymeric carbon (BPC). In the eastern Weddell Sea shelf, the BPC accounted for more than 90% of the OC in most of the samples. More than 82% of the total PRT, LPD, and CHO were present in the fraction < 200 μm. This work remarks the existence of sediments with a high nutritional value persistent several weeks after the spring–summer pulse of fresh organic matter. It is also highlighted the high potential availability of these sediments (due to its grain size) for the benthic communities inhabiting this high-latitude continental shelf.     

Topological constraints on the dynamics of wasp-waist ecosystems

Small pelagic fish species like anchovy or sardines are of high ecological and economical importance. As marine food webs are fished down, these small pelagics tend to be more exploited and overfished. It is not yet very well known what the possible effects of their collapse can be, therefore there is an urgent need to outline a theoretical framework for understanding their dynamics. These fish occupy very special position in food webs, ensuring energy transfer between species lower and higher levels, while forming narrow “wasp-waists” poor in number of species (but very abundant). Our purpose was to quantify the interaction structure of model food webs of equal complexity but different levels of “wasp-waistedness”. We analysed the topological properties of the webs by characterising every direct and indirect interactions between individual species, as well as by assessing the relative positional importance of each species in each web. We found that (1) the shorter the interaction pathways considered, the weaker the predictive power of node degree for positional importance, (2) the importance of species varies more in wasp-waist food webs, (3) if longer indirect chain effects are considered, indirect effects can well be stronger than direct ones, (4) interactions between coexisting wasp-waist species are stronger than the average, and (5) the “self-regulatory” looping effects are also stronger for wasp-waist species. Based on the topological properties of the networks, our results describe constraints acting on the dynamical behaviour of wasp-waist ecosystems. We give explanations, from this viewpoint, for regime shifts in which one WW species replaces another, and for the unpredictable dynamics of these fish stocks. From a marine conservation viewpoint, we illustrate that as the abundance of wasp-waist species decreases, the architecture of energy flows becomes highly vulnerable and unreliable. We provide an approach for quantifying these structural changes.     

Application of new parameterizations of gas transfer velocity and their impact on regional and global marine CO2 budgets

One of the dominant sources of uncertainty in the calculation of air–sea flux of carbon dioxide on a global scale originates from the various parameterizations of the gas transfer velocity, k, that are in use. Whilst it is undisputed that most of these parameterizations have shortcomings and neglect processes which influence air–sea gas exchange and do not scale with wind speed alone, there is no general agreement about their relative accuracy.The most widely used parameterizations are based on non-linear functions of wind speed and, to a lesser extent, on sea surface temperature and salinity. Processes such as surface film damping and whitecapping are known to have an effect on air–sea exchange. More recently published parameterizations use friction velocity, sea surface roughness, and significant wave height. These new parameters can account to some extent for processes such as film damping and whitecapping and could potentially explain the spread of wind-speed based transfer velocities published in the literature.We combine some of the principles of two recently published k parameterizations [Glover, D.M., Frew, N.M., McCue, S.J. and Bock, E.J., 2002. A multiyear time series of global gas transfer velocity from the TOPEX dual frequency, normalized radar backscatter algorithm. In: Donelan, M.A., Drennan, W.M., Saltzman, E.S., and Wanninkhof, R. (Eds.), Gas Transfer at Water Surfaces, Geophys. Monograph 127. AGU,Washington, DC, 325–331; Woolf, D.K., 2005. Parameterization of gas transfer velocities and sea-state dependent wave breaking. Tellus, 57B: 87–94] to calculate k as the sum of a linear function of total mean square slope of the sea surface and a wave breaking parameter. This separates contributions from direct and bubble-mediated gas transfer as suggested by Woolf [Woolf, D.K., 2005. Parameterization of gas transfer velocities and sea-state dependent wave breaking. Tellus, 57B: 87–94] and allows us to quantify contributions from these two processes independently.We then apply our parameterization to a monthly TOPEX altimeter gridded 1.5° × 1.5° data set and compare our results to transfer velocities calculated using the popular wind-based k parameterizations by Wanninkhof [Wanninkhof, R., 1992. Relationship between wind speed and gas exchange over the ocean. J. Geophys. Res., 97: 7373–7382.] and Wanninkhof and McGillis [Wanninkhof, R. and McGillis, W., 1999. A cubic relationship between air−sea CO2 exchange and wind speed. Geophys. Res. Lett., 26(13): 1889–1892]. We show that despite good agreement of the globally averaged transfer velocities, global and regional fluxes differ by up to 100%. These discrepancies are a result of different spatio-temporal distributions of the processes involved in the parameterizations of k, indicating the importance of wave field parameters and a need for further validation.     

Community structure of mesopelagic copepods (>500 μm) in the Ligurian Sea (Western Mediterranean)

The distribution of mesopelagic copepods between 250 and 1000 m depth was studied along a transect off the coast of Villefranche-sur-Mer in June 1991 by means of a BIONESS multiple-net sampler. Among the most abundant species at all stations were Pleuromamma gracilis, Paraeuchaeta acuta and overwintering Calanus helgolandicus CV copepodites, each species inhabiting different depth layers with maximum abundances in the 250–350 m, 450–550 m and the 700–850 m layer, respectively. Cluster analysis of the vertical distribution of all species caught revealed three distinct strata (‘TOP', ‘INTERMEDIATE', ‘BOTTOM') consisting of characteristic species that occurred in the same stratum at all stations. Differences in total abundances of the species assemblage occupying each layer are discussed with regard to predation pressure by mesopelagic macroplanktonic crustaceans and fish. We show that each stratum is characterized by distinct trophic interactions and life strategies, which play an important role in the structuring of mesopelagic zooplankton communities.     

The Mercator global ocean operational analysis system: Assessment and validation of an 11-year reanalysis

This paper presents Prototype Système 2 Global (PSY2G), the first Mercator global Ocean General Circulation Model (OGCM) to assimilate along-track sea level anomaly (SLA) satellite data. Based on a coarse resolution ocean model, this system was developed mainly for climatic purposes and will provide, for example, initial oceanic states for coupled ocean-atmosphere seasonal predictions. It has been operational since 3 September 2003 and produces an analysis and a two-week forecast for the global ocean every week. The PSY2G system uses an incremental assimilation scheme based on the Cooper and Haines [Cooper, M., Haines, K., 1996. Data assimilation with water property conservation. J. Geophys. Res., 101, 1059-1077.] lifting–lowering of isopycnals. The SLA increment is obtained using an optimal interpolation method then the correction is partitioned into baroclinic and barotropic contributions. The baroclinic ocean state correction consists of temperature, salinity and geostrophic velocity increments and the barotropic correction is a barotropic velocity increment. A reanalysis (1993–2003) was carried out that enabled the PSY2G system to perform its first operational cycle. All available SLA data sets (TOPEX/Poséïdon, ERS2, Geosat-Follow-On, Jason1 and Envisat) were assimilated for the 1993–2003 period. The major objective of this study is to assess the reanalysis from both an assimilation and a thermodynamic point of view in order to evaluate its realism, especially in the tropical band which is a key region for climatic studies. Although the system is also able to deliver forecasts, we have mainly focused on analysis. These results are useful because they give an a priori estimation of the qualities and capabilities of the operational ocean analysis system that has been implemented. In particular, the reanalysis identifies some regional biases in sea level variability such as near the Antarctic Circumpolar Current, in the eastern Equatorial Pacific and in the Norwegian Sea (generally less than 1 cm) with a small seasonal cycle. This is attributed to changes in mean circulation and vertical stratification caused by the assimilation methodology. But the model's low resolution, inaccurate physical parameterisations (especially for ocean–ice interactions) and surface atmospheric forcing also contribute to the occurrence of the SLA biases. A detailed analysis of the thermohaline structure of the ocean reveals that the isopycnal lifting–lowering tends to diffuse vertically the main thermocline. The impact on temperature is that the surface layer (0–200 m) becomes cooler whereas in deeper waters (from 500 to 1500 m), the ocean becomes slightly warmer. This is particularly true in the tropics, between 30°N and 30°S. However it can be demonstrated that the assimilation improves the variability in both surface currents and sub-surface temperature in the Equatorial Pacific Ocean.     

Modelling the effects of physical–biological interactions and spatial variability in spawning and nursery areas on transport and retention of sardine Sardinops sagax eggs and larvae in the southern Benguela ecosystem

A particle-tracking individual-based model (IBM) coupled with a 3-D hydrodynamic model was used to investigate how spatial variability in spawning and nursery grounds may influence transport and retention of sardine (Sardinops sagax) eggs and larvae in the southern Benguela ecosystem. A temperature-dependent Bělehrádek development model based on laboratory growth data was used to determine the duration of one egg and two larval stages. Successful transport/retention depended on each particle's stage of development rather than its age. Results show that recruitment could occur in two distinct nursery areas on the west and south coasts of South Africa. Three viable recruitment ‘systems’ were identified: two are retention-based (spawned and retained on the west coast (WC–WC) or the south coast (SC–SC)) and one is transport-based (spawned on the south coast and transported to the west coast (SC–WC)). In the WC–WC system, the vertical distribution of eggs influenced retention; at intermediate depths of spawning (25–50 m) eggs avoided both offshore Ekman drift and deep cold water. In the SC–SC system, the area of spawning was important; > 50% of eggs from the eastern Agulhas Bank (EAB) were retained in the south coast nursery grounds, whereas very few eggs were transported there from west of Cape Agulhas. In the SC–WC system, area of spawning was also important; 40% of the eggs spawned on the western Agulhas Bank (WAB) were transported to the west coast nursery ground. Sardine life history strategy could be divided between two main systems: the west coast system (spawning on the WAB and WC, and recruiting to the WC) and the Agulhas Bank system (spawning on the central and eastern Agulhas Bank, and recruiting to the SC).     

Population dynamics of the red shrimp Aristeus antennatus in the Balearic Islands (western Mediterranean): Short spatio-temporal differences and influence of environmental factors

The red shrimp Aristeus antennatus is one of the target species of the bottom trawl fishery of the Balearic Islands. The objective of the present paper is to study the short spatial and temporal differences of this important economic resource between two different locations off Mallorca (Cabrera: CA; Sóller: SO), where a fleet mobility pattern has been detected, and to study the influence of environmental conditions on this species. Six simultaneous bottom-trawl and oceanographic surveys were carried out at these two locations in order to collect data from the demersal species, hydrography (temperature and salinity), trophic resources and sediment characteristics. The commercial fleet from both locations was monitored by monthly on-board sampling, log-books and daily landings obtained from sales slips. Additional data was obtained from other fishing surveys. Short spatial and temporal differences have been detected between both locations. The population at CA was more demographically homogeneous, while that at SO showed important variations, like high abundance of juveniles recruiting to fishing grounds in autumn–winter and high abundance of large females during summer. Several differences have also been found in the biology of the species between locations, such as males were more abundant in SO than in CA. Also, the reproductive period started sooner in SO than in CA, and the condition of pre-spawning females was better in SO. The percentage of total lipids in the hepatopancreas was minimal during the spawning period, showing their importance as a reserve of energy for the ovary ripening. Water masses could play an important role in these differences, the characteristics of water masses being more stable in CA than in SO. Red shrimp adult females seemed to be more correlated with the warmer and more saline Levantine Intermediate Waters, while juveniles (males and females) and adult males were more correlated with the colder Western Mediterranean Deep Waters, detected only in SO during autumn–winter. Two different hypothesis of mobility patterns for the species are discussed in relation to these observed differences.     

Dynamics of suprabenthos-zooplankton communities around the Balearic Islands (western Mediterranean): Influence of environmental variables and effects on the biological cycle of Aristeus antennatus

Dynamics of suprabenthos and zooplankton were analyzed in two areas located in the NW (off Sóller harbour) and S (off Cabrera Archipelago) of Mallorca (Balearic Islands, western Mediterranean) at depths ranging between 135–780 m. Four stations situated respectively at 150 m (shelf-slope break), and at bathyal depths of 350, 650 and 750 m were sampled at bi-monthly intervals during six cruises performed between August 2003 and June 2004. Suprabenthos showed maximum biomass in both areas from late spring to summer (April to August), while minimum biomass was found in autumn (September–November). Though variable, temporal dynamics of zooplankton showed peaks of biomass in late winter and summer (February and June), while minimals occurred in autumn (August–September) and, at bathyal depths, in April. Suprabenthos (abundance; MDS analyses) showed a sample aggregation as a function of depth (3 groups corresponding to the shelf-slope break, upper slope — over 350 m; and the middle, deeper part of the slope — over 650–750 m), without any separation of hauls by season. By contrast, zooplankton samples were separated by season and not by depth. There was evidence of three seasonal groups corresponding to summer (June 2004–August 2003), autumn–winter (September and November 2003, February 2004), and spring (April 2004), being especially well established off Sóller. In general, suprabenthos was significantly correlated with the sediment variables (e.g. total organic matter content (% OM), potential REDOX), whereas zooplankton was almost exclusively dependent on Chl a at the surface, which suggests two different food sources for suprabenthos and zooplankton. The increase of suprabenthos abundance in April–June was paralleled by a sharp increase (ca. 2.8 times) in the %OM on sediment during the same period, coupled ca. 1–2 months of delay with the peak of surface Chl a recorded in February–March (from satellite imagery data). Suprabenthos biomass was also correlated with salinity close to the bottom, suggesting a link between suprabenthos abundance and changes in the oceanographic condition of water masses close to the bottom. It is suggested that a higher suprabenthos biomass recorded off Sóller in comparison to that off Cabrera in June could, in turn, be related to a seasonal inflow of Levantine Intermediate Water (LIW) in April–June in this area at mid bathyal depths (350–650 m). This trend would be based on: 1) it was evident only at mid-slope depths between 350–750 m, coinciding with the LIW distribution, and 2) it was not recorded among zooplankton (collected throughout the water column). The possible effect of the fluctuations of suprabenthos and zooplankton on higher trophic levels has been explored studying the diet and food consumption rates of the red shrimp Aristeus antennatus, as indicator species by its dominance in bathyal communities. A. antennatus increased its food consumption from February to April–June 2004 off Sóller, which in the case of large (CL > 40 mm) specimens was found in both areas. In addition, there was a shift of diet from winter to spring–early summer. In this last period, A. antennatus preyed upon euphausiids and mesopelagic decapods and fish, while benthos (e.g. polychaetes and bivalves) decreased in the diet. This indicates an increase in the food consumption and probably in the caloric content of the diet in pre-spawning females in April–June 2004, which is synchronized with the period when gonad development begins in A. antennatus females (May–June). Anyway, macrozooplankton, and not suprabenthos, is crucial as a high energetic food source in the coupling between food intake and reproduction in the red shrimp.     

The influence of different turbulence schemes on modelling primary production in a 1D coupled physical–biological model

A one-dimensional (1D) coupled physical–microbiological model has been applied to a site in the central North Sea. The impact of the choice of the turbulence closure scheme on the modelling the primary production has been investigated.The model was run with four different parameterisations of vertical mixing of heat, momentum and dissolved and suspended matters, using M2 tidal forcing and the hourly mean meteorological forcing of 1989 to reproduce the annual thermal structure and primary production. The four mixing parameterisations are: Level 2 turbulence closure scheme [Mellor, G.L., Yamada, T., 1974. A hierarchy of turbulence closure models for planetary boundary layers. J. Atmos. Sci. 31, 1791–1806; Mellor, G.L., Yamada, T., 1982. Development of a turbulence closure model for geophysical Fluid problems. Rev. Geophys. Space Phys. 20 (4) 851–875] using an explicit numerical scheme [Sharples, J., Tett, P., 1994. Modelling the effect of physical variability on the midwater chlorophyll maximum. J. Mar. Res. 52, 219–238]; a version of the Level 2.5 turbulence closure scheme [Galperin, B., Kantha, L.H., Hassid, S., Rosati, A., 1988. A quasi-equilibrium turbulent energy model for geophysical flows. J. Atmos. Sci. 45, 55–62; Ruddick, K.G., Deleersnijder, E., Luyten, P.J., Ozer, J., 1995. Haline stratification in the rhine/meuse freshwater plume: a 3D model sensitivity analysis. Cont. Shelf Res. 15 (13) 1597–1630] simplified to use an algebraic mixing length by Sharples and Simpson [Sharples, J., Simpson, J.H., 1995. Semidiurnal and longer period stability cycles in the Liverpool Bay region of freshwater influence. Cont. Shelf Res. 15, 295–313], also solved explicitly; the same simplified L2.5 scheme with an implicit numerical solution and modified vertical discretisation scheme [Annan, J.D., 1999. Numerical methods for the solution of the turbulence energy equations in the shelf seas. Int. J. Numer. Methods Fluids 29, 193–206]; and another version of the same scheme (but using a different algebraic mixing length) as described by Xing and Davies [Xing, J., Davies, A.M., 1996a. Application of turbulence energy models to the computation of tidal currents and mixing intensities in the shelf edge regions. J. Phys. Oceanogr. 26, 417–447; Xing, J., Davies, A.M., 1996b. Application of a range of turbulence models to the computation of tidal currents and mixing intensities in shelf edge regions. Cont. Shelf. Res. 16, 517–547; Xing, J., Davies, A.M., 1998. Application of a range of turbulence energy models to the computation of the internal tide. Int. J. Numer. Methods Fluids 26, 1055–1084]. Various model outputs at the sea surface and in depth profiles have been compared with data collected in 1989 as part of the North Sea Project [Huthnance, J.M., 1990. Progress on North Sea Project. NERC News, vol. 12, pp. 25–29, UK]. It is shown that the biological results are extremely sensitive to the small changes in the physical conditions, which arise due to the different turbulence schemes tested. The timing of the spring bloom and the maintenance of the midwater chlorophyll maximum all differ greatly between model runs, and the gross primary production varies by a factor of two from the highest to lowest results. The simplified Level 2.5 scheme, implemented using the numerical methods of Annan [Annan, J.D., 1999. Numerical methods for the solution of the turbulence energy equations in the shelf seas. Int. J. Numer. Methods Fluids 29, 193–206], produces results, which give the best agreement with the available data.     

Enrichment, concentration and retention processes in relation to anchovy (Engraulis ringens) eggs and larvae distributions in the northern Humboldt upwelling ecosystem

A Lagrangian model is used to simulate and quantify in the northern Humboldt upwelling ecosystem the processes of enrichment, concentration and retention, identified by Bakun [Bakun, A., 1996. Patterns in the ocean. Ocean processes and marine population dynamics. University of California Sea Grant, California, USA, in cooperation with Centro de Investigaciones Biologicas de Noroeste, La Paz, Baja California Sur, Mexico, 323 pp.] as being important for the survival and recruitment of early life stages of pelagic fish. The method relies on tracking the positions of particles within water velocity fields generated by a three-dimensional hydrodynamic model. Simple criteria for considering particles as participating to enrichment, concentration or retention are used to derive indices of the three processes. We analyse the spatial distribution of and seasonal variability in these indices. The results are discussed in relation to anchovy (Engraulis ringens) eggs and larvae distributions off Peru, and to a comparable study conducted in the southern Benguela upwelling ecosystem.