Variability and predictability of Antarctic krill swarm structure

Swarming is a fundamental part of the life of Euphausia superba, yet we still know very little about what drives the considerable variability in swarm shape, size and biomass. We examined swarms across the Scotia Sea in January and February 2003 using a Simrad EK60 (38 and 120 kHz) echosounder, concurrent with net sampling. The acoustic data were analysed through applying a swarm-identification algorithm and then filtering out all non-krill targets. The area, length, height, depth, packing-concentration and inter-swarm distance of 4525 swarms was derived by this method. Hierarchical clustering revealed 2 principal swarm types, which differed in both their dimensions and packing-concentrations. Type 1 swarms were generally small (<50 m long) and were not very tightly packed (<10 ind. m⁻³), whereas type 2 swarms were an order of magnitude larger and had packing concentrations up to 10 times greater. Further sub-divisions of these types identified small and standard swarms within the type 1 group and large and superswarms within the type 2 group. A minor group (swarm type 3) was also found, containing swarms that were isolated (>100 km away from the next swarm). The distribution of swarm types over the survey grid was examined with respect to a number of potential explanatory variables describing both the environment and the internal-state of krill (namely maturity, body length, body condition). Most variables were spatially averaged over scales of 100 km and so mainly had a mesoscale perspective. The exception was the level of light (photosynthetically active radiation (PAR)) for which measurements were specific to each swarm. A binary logistic model was constructed from four variables found to have significant explanatory power (P<0.05): surface fluorescence, PAR, krill maturity and krill body length. Larger (type 2) swarms were more commonly found during nighttime or when it was overcast during the day, when surface fluorescence was low, and when the krill were small and immature. A strong pattern of diel vertical migration was not observed although the larger and denser swarms tended to occur more often at night than during the day. The vast majority of krill were contained within a minor fraction of the total number of swarms. These krill-rich swarms were more common in areas dominated by small and immature krill. We propose that, at the mesoscale level, the structure of swarms switches from being predominantly large and tightly packed to smaller and more diffuse as krill grow and mature. This pattern is further modulated according to feeding conditions and then level of light.     

Ecological niches and areas of overlap of the squat lobster ‘munida’ (Pleuroncodes monodon) and anchoveta (Engraulis ringens) off Peru

The world’s largest mono-specific fishery, the Peruvian anchovy or anchoveta (Engraulis ringens) fishery, has been the subject of many studies since the 1960s. Details of its relationship with other species have mainly focused on alternations with sardine, Sardinops sagax, and little effort has so far been paid to interactions with other species sharing the same ecosystem. This is the case for Pleuroncodes monodon, the crustacean squat lobster or ’munida’, which has become highly abundant along the Peruvian coast since the mid-1990s. Munida is now an important prey for seabirds, mammals and coastal predatory fish. Knowledge of patterns of distribution and ecological niche of munida is scarce however off Peru. Here we describe and compare spatial patterns of distribution of anchoveta and munida and their ecological niches based on data from 26 acoustic surveys performed along the Peruvian coast between 1998 and 2006. The results indicate that munida and anchoveta share ecological niches but that munida is restricted to the coldest part of the productive cold coastal waters whereas anchoveta do not present any temperature preference over a large range (14–23 °C). The recent increase in munida abundance off Peru is concomitant with colder conditions; with their onset munida extended its range from central Chile northwards. Off Peru the very shallow oxycline keeps munida from its usual bottom habitat and has forced it to adopt pelagic behaviour.     

Physical dynamics and biological implications of Cyclone Noah in the lee of Hawaii during E-Flux I

Quasi-synoptic observations of the horizontal and vertical structure of a cold-core cyclonic mesoscale eddy feature (Cyclone Noah) were conducted in the lee of Hawai’i from November 4–22, 2004 as part of the E-Flux interdisciplinary collaborative research program. Cyclone Noah appears to have spun up to the southwest of the ‘Alenuihaha Channel (between Maui and Hawai’i) as a result of strong and persistent northeasterly trade winds through the channel. Shipboard hydrographic surveys 2.5 months later suggest that Noah weakened and was in a hypothesized spin-down phase of its life cycle. Although the initial surface expression of Noah was limited in scale to 40 km in diameter and, as evidenced by surface temperatures, 2–3 °C cooler than the surrounding waters, depth profiles revealed a fully developed semi-elliptical shallow feature (200 m), 144 km long and 90 km wide (based on sigma-t=23 kg m⁻³) with tangential speeds of 40–80 cm s⁻¹, and substantial isopycnal doming. Potential vorticity distribution of Noah suggests that radial horizontal flow of the core water was inhibited from the surface to depths of 75 m, with high vorticity confined above the sigma-t=23.5 kg m⁻³ isopycnal surface. Upward displacements of isopycnal surfaces in the eddy's center (50 m) were congruent with enhanced pigment concentrations (0.50 mg m⁻³). Comparisons of the results obtained for E-Flux I (Noah) and E-Flux III (Opal) suggest that translation characteristics of cyclonic Hawaiian lee eddies may be important in establishing the biogeochemical and biological responses of the oligotrophic ocean to cyclonic eddies.     

Short-term under-ice variability of prokaryotic plankton communities in coastal Antarctic waters (Cape Hallett, Ross Sea)

During the 2006 Italian Antarctic expedition a diel sampling was performed close to Cape Hallett (Ross Sea) during the Austral summer. Under-ice seawater samples (4 m) were collected every 2 h for 28 h in order to estimate prokaryotic processes' variability and community structure dynamics. Prokaryotic and viral abundances, exoenzymatic activities (β-glucosidase, chitinase, lipase, alkaline phosphatase and leucine aminopeptidase), prokaryotic carbon production (³H-leucine incorporation) and community structure (Denaturing Gradient Gel Electrophoresis – DGGE fingerprints) were analysed. Results showed that the diel variability of the prokaryotic activity followed a variation in salinity, probably as a consequence of the periodical thawing of sea ice (driven by solar radiation and air temperature cycles), while negligible variation in viral and prokaryotic abundances occurred. The Bacterial and Archaeal community structures underwent an Operational Taxonomic Units (OTUs) temporal shift from the beginning to the end of the sampling, while Flavobacteria-specific primers highlighted high variations in this group possibly related to sea ice melting and substrate release.     

Accumulation of Cu and Zn in discarded antifouling paint particles by the marine gastropod, Littorina littorea

The short-term (5 day) accumulation of Cu and Zn in different tissues of the marine gastropod, Littorina littorea, has been studied in the presence of 10 mg l⁻¹ of antifouling paint particles and pre- or simultaneously contaminated algal food (Ulva lactuca). Accumulation of Cu was observed in the head–foot, digestive gland–gonad complex and gills to extents dependent on how and when food was contaminated and administered. However, retention of Zn was only observed in the gills and only when L. littorea and U. lactuca were simultaneously exposed to paint particles. Relative to the alga, faecal material was highly enriched in Zn, suggesting that the animal is able to rapidly eliminate this metal, most likely through the formation and egestion of insoluble phosphate granules. Thus, L. littorea is a useful biomonitor of marine contamination by antifouling applications in respect of Cu but not Zn.     

Uptake and metabolism of arsenate by anexic cultures of the microalgae Dunaliella tertiolecta and Phaeodactylum tricornutum

Axenic cultures of the microalgae species, Dunaliella tertiolecta and Phaeodactylum tricornutum were grown at arsenic (As) concentrations typically found in uncontaminated marine environments ( 2 µg L^− 1) under different phosphorus concentrations. D. tertiolecta accumulated higher arsenic concentrations (mean: 13.7 ± 0.7 µg g^− 1 dry mass) than P. tricornutum (mean: 1.9 ± 0.2 µg g−1 dry mass). Media phosphorus concentrations (0.6–3 mg/L) had little influence on microalgae growth rates or arsenic accumulation. Arsenic was present as lipid bound (29–38%; 4.2–9.5%), water-soluble (20–29%; 26–34%) and residue bound (41–45%; 57–69%) arsenic species in D. tertiolecta and P. tricornutum respectively. Hydrolysed lipids contained mostly glycerol arsenoribose (OH- ribose), dimethylarsinate (DMA) and inorganic arsenic (As(V)) moieties. Water-soluble species of microalgae were very different. D. tertiolecta contained inorganic arsenic (54–86%) with variable amounts of DMA (7.4–20%), arsenoriboses (5–25%) and traces of methylarsonate (MA) ( 1%). P. tricornutum contained mostly DMA (32–56%) and phosphate arsenoribose (PO₄-ribose, 23–49%) and small amounts of OH-ribose (3.8–6.5%) and As(V) (9–16%). Both microalgae contained an unknown cationic arsenic species. The residue fractions of both microalgae contained predominately inorganic arsenic (99–100%). These results show that at natural seawater arsenic concentrations, both algae take up substantial amounts of inorganic arsenic that is complexed with structural elements or sequestered in vacuoles as stable complexes. A significant portion is also incorporated into lipids. Arsenic is metabolised to simple methylated species and arsenoriboses.     

Alterations in seawater pH and CO2 affect calcification and photosynthesis in the tropical coralline alga, Hydrolithon sp. (Rhodophyta)

Calcification in the marine environment is the basis for the accretion of carbonate in structures such as coral reefs, algal ridges and carbonate sands. Among the organisms responsible for such calcification are the Corallinaceae (Rhodophyta), recognised as major contributors to the process world-wide. Hydrolithon sp. is a coralline alga that often forms rhodoliths in the Western Indian Ocean. In Zanzibar, it is commonly found in shallow lagoons, where it often grows within seagrass beds and/or surrounded by green algae such as Ulva sp. Since seagrasses in Zanzibar have recently been shown to raise the pH of the surrounding seawater during the day, and since calcification rates are sensitive to pH, which changes the saturation state of calcium carbonate, we measured the effects of pH on photosynthetic and calcification rates of this alga. It was found that pH had significant effects on both calcification and photosynthesis. While increased pH enhanced calcification rates both in the light and in the dark at pH >8.6, photosynthetic rates decreased. On the other hand, an increase in dissolved CO₂ concentration to 26 μmol kg⁻¹ (by bubbling with air containing 0.9 mbar CO₂) caused a decrease in seawater pH which resulted in 20% less calcification after 5 days of exposure, while enhancing photosynthetic rates by 13%. The ecological implications of these findings is that photosynthetically driven changes in water chemistry by surrounding plants can affect calcification rates of coralline algae, as may future ocean acidification resulting from elevated atmospheric CO₂.     

Revisiting Peruvian anchovy (Engraulis ringens) trophodynamics provides a new vision of the Humboldt Current system

The Peruvian anchovy or anchoveta (Engraulis ringens) forages on plankton and is a main prey for marine mammals, seabirds, fish, and fishers, and is therefore a key element of the food web in the Humboldt Current system (HCS). Here, we present results from the analysis of 21,203 anchoveta stomach contents sampled during 23 acoustic surveys over the period 1996–2003. Prey items were identified to the genus level, and the relative dietary importance of different prey was assessed by determination of their carbon content. Variability in stomach fullness was examined relative to the diel cycle, the distance from the coast, sea surface temperature, and latitude, using generalized additive models (GAMs). Whereas phytoplankton largely dominated anchoveta diets in terms of numerical abundance and comprised >99% of ingested prey items, the carbon content of prey items indicated that zooplankton was by far the most important dietary component, with euphausiids contributing 67.5% of dietary carbon followed by copepods (26.3%). Stomach fullness data showed that anchoveta feed mainly during daytime between 07h00 and 18h00, although night-time feeding also made a substantial contribution to total food consumption. Stomach fullness also varied with latitude, distance from the coast, and temperature, but with substantial variability indicating a high degree of plasticity in anchoveta feeding behaviour. The results suggest an ecological role for anchoveta that challenges current understanding of its position in the foodweb, the functioning of the HCS, and trophic models of the HCS.     

东南太平洋秘鲁鳀资源量预报模型的构建

秘鲁鳀（Engraulis ringens）是栖息于东南太平洋沿岸的一种小型中上层鱼类。有效地对秘鲁鳀资源量进行预报将有助于为我国鱼粉进口企业提供决策支撑。为此,本研究结合秘鲁鳀生物（上一个渔汛季度的资源量、渔获物中的幼鱼比例）和环境（渔场水温和nino1+2区的温度距平）因素及捕捞量为预报因子,利用主成分分析和多元线性回归的方法对17个渔汛季度（2006年至2014年第一渔汛季度）秘鲁鳀的资源量建立预报模型并利用主成分分析的结果对影响秘鲁鳀资源变动的因子进行初步评价。研究表明,随着时间的推移样本量的增加,模型拟合资源量与真实资源量的平均相对误差逐渐下降,拟合资源量序列与真实资源量序列的相关系数逐渐增加。最终模型5（建模数据为2006-2013年第二渔汛季度的数据,验证数据为2014年第一渔汛季度的数据）能够很好地拟合出秘鲁鳀资源量的大小及变动趋势拟合资源量序列与真实资源量序列的相关系数为0.861;拟合资源量序列与真实资源量序列的平均相对误差为12%;预报得到的2014年第一渔汛季度的数据与真实值相比,相对误差为1%。结合主成分分析的结果对影响秘鲁鳀资源变动的因素进行评价结果表明,第一主成分的方差贡献率为46%,其中环境因素占据了最大的载荷;第二主成分的方差贡献率为23%,同样环境因素占据了最大载荷,但是,排名第二和第三的因素分别是上一个渔汛季度的资源量和捕捞量,其载荷相当;第三和第四主成分的方差贡献率分别为9%和7%,其中幼鱼比例占的载荷最大。根据各主成分得分序列与资源量序列的相关系数结果,环境因子对秘鲁鳀资源变动有重要影响。     

Retention of crab larvae in a coastal null zone

Alongshelf transport in the southern Middle Atlantic Bight is forced by buoyancy-driven currents originating in three large estuaries along the bight. These currents are strongest in the coastal ocean near the southern terminus of each estuary, while the analogous region on the northern side is characterized by weak subtidal flow. We used a combination of field observations and numerical modeling to test the hypothesis that these regions of weak subtidal flow are coastal null zones that serve as retention areas for larvae. The field study consisted of a four-day, shipboard investigation of the distribution of blue crab larvae (Callinectes sapidus) near the mouth of Delaware Bay (39°N, 75°W) in late summer, 2004. Hydrographic surveys of the study site were conducted with a hull-mounted, surface-measuring system. Results showed a sharp boundary between the null zone and the buoyancy-driven current to the south. Blue crab larvae were collected in surface plankton tows along a 30-km transect that encompassed these two areas. Stations with higher densities of larvae were clustered in the null zone during both ebb and flood tides. A numerical model was used to examine the physical mechanisms responsible for the observed distribution. Model results agreed with the field survey and showed that simulated larvae are aggregated in the null zone. The simulations also demonstrated that larvae spawned within the null zone have a much greater probability of settling in juvenile nursery habitat within the bay. The close agreement between field and model results provides consistent support for the hypothesis that coastal null zones associated with the buoyancy-driven circulation of large estuaries may allow retention of larvae in the vicinity of the natal spawning population.     

基于复合模型的西南大西洋阿根廷滑柔鱼（Illex argentinus）时空分布研究

本文利用2003-2011年西南大西洋阿根廷滑柔鱼渔业数据和海洋环境数据,包括海表温度（sea surface temperature, SST）,海面高度（sea surface height, SSH）和叶绿素浓度（chlorophyll a, Chl a）,开发基于广义加性模型（GAM）和神经网络模型（NNM）的复合模型研究滑柔鱼资源时空分布。GAM用于选择关键影响因子,并分析与单位捕捞努力量渔获量（catch per unit effort, CPUE）的关系,NNM用于建立关键影响因子与CPUE之间的预报模型。结果表明：GAM选择的影响因子的偏差解释率为53.8%,空间变量（经度和纬度）,环境变量（SST、SSH、Chl a）均匀CPUE之间存在显著相关性。CPUE与SST和SSH之间为非线性关系,与Chl a之间为线性关系。NNM模型的MSE和ARV较低,其精度高且稳定。此复合模型也能够解释解释西南大西洋阿根廷滑柔鱼时空变化趋势和迁徙模式。     

Habitat selection and quality for multiple cohorts of young-of-the-year bluefish (Pomatomus saltatrix): Comparisons between estuarine and ocean beaches in southern New Jersey

In this study, seasonal and annual variability in the use of estuarine and ocean beaches by young-of-the-year bluefish, Pomatomus saltatrix, was evaluated by indices of abundance in coastal areas of southern New Jersey (1998–2000). Biological and physical factors measured at specific sites were correlated with bluefish abundance to determine the mechanisms underlying habitat selection. In addition, integrative and discrete indicators of bluefish growth were used to examine spatio-temporal dynamics in habitat quality and its effect on habitat selection by multiple cohorts of bluefish. Intra-annual recruitment to coastal areas of southern New Jersey was episodic, and resulted from the ingress of spring-spawned bluefish (hatch-date April) to estuarine beaches in late May to early June, followed by the recruitment of summer-spawned fish (hatch-date early July) to ocean beaches from July to October. Bluefish utilized estuarine and ocean beaches in a facultative manner that was responsive to dynamics in prey composition and temperature conditions. The recruitment and residency of bluefish in the estuary (1998–1999) and ocean beaches (1998), for example, was coincidental with the presence of the Atlantic silverside Menidia menidia and bay anchovy Anchoa mitchilli, the principal prey species for bluefish occupying these respective habitat-types. Bluefish abundance in the estuary (2000) and ocean beaches (1999–2000) was also correlated with water temperature, with the greatest catches of juveniles coinciding with their optimal growth temperature (24 °C). Bluefish growth, estimated as the slope of age–length relationships and daily specific growth rates, equaled 1.27–2.63 mm fork length (FL) d⁻¹ and 3.8–8.7% body length increase d⁻¹, respectively. The growth of sagittal otoliths was also used as a proxy for changes in bluefish size during and shortly before their time of capture. Accordingly, otolith growth rates of summer-spawned bluefish were greater at ocean beaches relative to the estuary and were explained by the more suitable temperature conditions found at ocean beaches during the mid- to late summer. Notwithstanding the fast growth of oceanic summer-spawned bluefish, individuals spawned in the spring were still larger in absolute body size at the end of the summer growing season (240 and 50–200 mm FL for spring- and summer-spawned bluefish, respectively). The size discrepancy between spring- and summer-spawned bluefish at the onset of autumn migrations and during overwintering periods may account for the differential recruitment success of the respective cohorts.     

Depth-stratified phytoplankton dynamics in Cyclone Opal, a subtropical mesoscale eddy

As part of E-Flux III cruise studies in March 2005, we investigated phytoplankton community dynamics in a cyclonic cold-core eddy (Cyclone Opal) in the lee of the Hawaiian Islands. Experimental incubations were conducted under in situ temperature and light conditions on a drift array using a two-treatment dilution technique. Taxon-specific estimates of growth, grazing and production rates were obtained from analyses of incubation results based on phytoplankton pigments, flow cytometry and microscopy. Cyclone Opal was sampled at a biologically and physically mature state, with an 80–100 m doming of isopycnal surfaces in its central region and a deep biomass maximum of large diatoms. Depth-profile experimentation defined three main zones. The upper (mixed) zone (0–40 m), showed little compositional or biomass response to eddy nutrient enrichment, but growth, grazing and production rates were significantly enhanced in this layer relative to the ambient community outside of the eddy. Prochlorococcus spp. dominated the upper mixed layer, accounting for 50–60% of its estimated primary production both inside and outside of Opal. In contrast, the deep zone of 70–90 m showed little evidence of growth rate enhancement and was principally defined by a 100-fold increase of large (>20-μm) diatoms and a shift from Prochlorococcus to diatom dominance (80%) of production. The intermediate layer of 50–60 m marked the transition between the upper and lower extremes but also contained an elevated biomass of physiologically unhealthy diatoms with significantly depressed growth rates and proportionately greater grazing losses relative to diatoms above or below. Microzooplankton grazers consumed 58%, 65% and 55%, respectively, of the production of diatoms, Prochlorococcus and the total phytoplankton community in Cyclone Opal. The substantial grazing impact on diatoms suggests that efficient recycling was the major primary fate of diatom organic production, consistent with the low export fluxes and selective export of biogenic silica, as empty diatom frustules, in Cyclone Opal.     

Zooplankton communities and oceanographic structures in a high-resolution grid in the south-eastern corner of the Bay of Biscay

A one year long monthly sampling series at a fixed station was combined with a high-spatial resolution survey (May 2003) to investigate factors controlling zooplankton distribution in the south-eastern corner of the Bay of Biscay. Species abundance and gonad maturity for Calanoides carinatus and Calanus helgolandicus were estimated to clarify whether the observed spatial patterns were attributable to increased population growth or to mechanical accumulation. The two studies included distinct oceanographic regimes. The eastern corner of the grid was characterized by Adour river plume waters and the Cap Breton canyon, and the rest of the area by an alternating pattern of physical convergences and divergences. In the river plume, the zooplankton community diversity and abundances were highest in response to the nutrient enriched river plume waters and the submarine canyon. In the rest of the area, distribution and abundance patterns were the result of a combination of behaviour and physical transport.     

Dissolved manganese and silicic acid in the Columbia River plume: A major source to the California current and coastal waters off Washington and Oregon

The spatial distributions of dissolved manganese and nutrients were examined in the Columbia River plume off Oregon and Washington during the summer of 2004 and 2005 as part of the River Influence on Shelf Ecosystems (RISE) program. Factors influencing the hydrochemical characteristics of the freshly formed and aged Columbia River plume were investigated. Hydrographic data and nutrient concentrations were used to delineate three distinct water sources for the Columbia River Plume: California Current surface water, coastal upwelled water, and Columbia River water. The warm, intermediate salinity, nutrient poor California Current water contains low levels of dissolved manganese (< 5 nM) and silicic acid (< 5 μM), and is depleted in nitrate. The cold, high salinity, nutrient rich, freshly upwelled water is highly variable (2–20 nM) in dissolved manganese and can be as high as  45 μM in silicic acid and  30 μM nitrate. The variable Columbia River has summer temperatures ranging from  13 to 24 °C, high silicic acid concentrations (ranging from  120 to 200 μM), and lower nitrate concentrations (ranging from  2 to 20 μM). During the summer, the concentrations of silicic acid and dissolved manganese can exceed 100 μM and 200 nM, respectively, in near-field Columbia River plumes. These values are markedly greater than those of surface coastal waters (even during upwelling conditions). As the plume advects and mixes, the concentrations of these two constituents remain relatively high within plume waters. The concentrations of dissolved manganese in the near-field plume vary with tidal amplitude, exhibiting much higher concentrations for a given salinity during spring tides than during neap tides. For example, the Columbia River plume at a salinity of 20 has a concentration of dissolved manganese of  240 nM during spring tides, as compared to only  60 nM during low amplitude tides. Silicic acid concentrations in the near-field plume remain relatively constant throughout the tidal month. Calculations indicate there is roughly an equivalent yearly delivery of dissolved manganese and silicic acid to the coastal waters off Oregon and Washington by upwelled waters and by the Columbia River plume.     

A multiple-scale analysis of metazoan meiofaunal distribution in the deep Mediterranean Sea

In order to identify environmental factors driving the distribution and functioning of deep-sea fauna and the spatial scales of interactions, we carried out a multiple-scale investigation in the Mediterranean basin in which we compared two bathyal plains, located at the same depth (ca. 3000 m), but characterised by contrasting trophic conditions. We investigated meiofaunal abundance, biomass, community structure and biodiversity (expressed as richness of taxa) in relation to sediment characteristics, downward fluxes and food availability in the sediment. Samples were collected at all spatial scales (from small to macroscale) in two seasons. Our results indicated that deep-sea systems with different trophic conditions displayed different responses to the distribution of available energy and its spatio-temporal variability in the sediment. The analysis at a macroscale (>1000 km) indicated that meiofauna were controlled primarily by the trophic inputs to the deep-sea system. Spatial variability of meiofaunal parameters at a mesoscale (>50 km) was highest in the eastern Mediterranean and lowest in the western Mediterranean. Such differences are the consequence of the unpredictable inputs of organic matter in the oligotrophic eastern Mediterranean versus a more homogeneous distribution of food inputs in the mesotrophic western Mediterranean. At a smaller scale (local scale 7 km), in the western Mediterranean, the distribution of meiofaunal parameters was highly homogeneous, reflecting the homogeneous distribution of the food availability in the sediment. Our results indicated that the highly variable input and distribution of food sources in the deep eastern Mediterranean did not provide any “insurance” for the sustainability of the deep-sea faunal assemblages in the long term, thus leading to an uncoupling between resource availability and distribution of organisms. We conclude that the influence of energy availability on the deep-sea faunal distributions change at different spatial scales and that the analysis of spatial variability at mesoscales is crucial for understanding the relationships between deep-sea benthic fauna and environmental drivers.     

Zooplankton and forage fish species off Peru: Large-scale bottom-up forcing and local-scale depletion

The Humboldt Current System, like all upwelling systems, has dramatic quantities of plankton-feeding fish, which suggested that their population dynamics may ‘drive’ or ‘control’ ecosystem dynamics. With this in mind we analysed the relationship between forage fish populations and their main prey, zooplankton populations. Our study combined a zooplankton sampling program (1961–2005) with simultaneous acoustic observations on fish from 40 pelagic surveys (1983–2005) conducted by the Peruvian Marine Research Institute (IMARPE) and landing statistics for anchoveta (Engraulis ringens) and sardine (Sardinops sagax) along the Peruvian coast from 1961 to 2005. The multi-year trend of anchoveta population abundance varied consistently with zooplankton biovolume trend, suggesting bottom-up control on anchovy at the population scale (since oceanographic conditions and phytoplankton production support the changes in zooplankton abundance). For a finer-scale analysis (km) we statistically modelled zooplankton biovolume as a function of geographical (latitude and distance from the 200-m isobath), environmental (sea surface temperature), temporal (year, month and time-of-day) and biological (acoustic anchovy and sardine biomass within 5 km of each zooplankton sample) covariates over all survey using both classification and regression trees (CART) and generalized additive models (GAM). CART showed local anchoveta density to have the strongest effect on zooplankton biovolume, with significantly reduced levels of biovolume for higher neighbourhood anchoveta biomass. Additionally, zooplankton biovolume was higher offshore than on the shelf. GAM results corroborated the CART findings, also showing a clear diel effect on zooplankton biovolume, probably due to diel migration or daytime net avoidance. Apparently, the observed multi-year population scale bottom-up control is not inconsistent with local depletion of zooplankton when anchoveta are locally abundant, since the latter effect was observed over a wide range of overall anchoveta abundance.     

Phytoplankton patchiness: the role of lateral stirring and mixing

Explanations for the patchy distribution of marine phytoplankton are critically reviewed with the focus on the role played by lateral advection and mixing. Generating mechanisms for what is increasingly seen as a ubiquitous feature of the oceans have been sought in biology, behaviour, population dynamics, physics and various combinations thereof. As the mesoscale and sub-mesoscale (1-500 km) contain the “weather” of the ocean—eddies and fronts capable of strong stirring and mixing of any tracers advected by them—the physical circulation in this regime may have a strong role in the generation of patchiness. As a consequence, the focus here is on the ways the turbulent currents at these scales can interact with other processes to produce the complex spatial structures in phytoplankton distributions seen in countless satellite images and cruise data. The mechanisms reviewed include patch formation (KiSS theory), filamentation, Turing and related instabilities, diffusion waves, shear effects and turbulent stirring. Parallel to this is a discussion of observations and of the techniques that have been used to analyse them, particularly spectral analysis. It will be seen that many of the limitations in extant theories stem from a paucity of data and ambiguity in its interpretation. A synthesis of recent developments in sampling, instrumentation, image analysis and turbulence theory is used to suggest alternatives to conventional approaches, to highlight important work in related fields and to motivate new lines of research.