Zooplankton and the oceanography of the eastern tropical Pacific: A review

We review the spatial and temporal patterns of zooplankton in the eastern tropical Pacific Ocean and relationships with oceanographic factors that affect zooplankton distribution, abundance and trophic relationships. Large-scale spatial patterns of some zooplankton groups show broad coincidence with surface water masses, circulation, and upwelling regions, in agreement with an ecological and dynamic partitioning of the pelagic ecosystem. The papers reviewed and a new compilation of zooplankton volume data at large-scale show that abundance patterns of zooplankton biomass have their highest values in the upwelling regions, including the Gulf of Tehuantepec, the Costa Rica Dome, the equatorial cold tongue, and the coast of Peru.Some of the first studies of zooplankton vertical distribution were done in this region, and a general review of the topic is presented. The possible physiological implications of vertical migration in zooplankton and the main hypotheses are described, with remarks on the importance of the oxygen minimum zone (OMZ) as a barrier to both the vertical distribution and migration of zooplankton in the region. Recent results, using multiple-net gear, show that vertical distribution is more complex than previously thought. There are some well-adapted species that do live and migrate within the OMZ.Temporal patterns are reviewed and summarized with historical data. Seasonal variations in zooplankton biomass follow productivity cycles in upwelling areas. No zooplankton time series exist to resolve ENSO effects in oceanic regions, but some El Niño events have had effects in the Peru Current ecosystem. Multidecadal periods of up to 50 years show a shift from a warm sardine regime with a low zooplankton biomass to a cool anchovy regime in the eastern Pacific with higher zooplankton biomasses. However, zooplankton volume off Peru has remained at low values since the 1972 El Niño, a trend opposite to that of anchoveta biomass since 1984.Studies of trophic relations emphasize the difference in the productivity cycle in the eastern tropical Pacific compared to temperate or polar ecosystems, with no particular peaks in the stocks of either zooplankton or phytoplankton. Productivity is more dependent on local events like coastal upwelling or water circulation, especially in the equatorial countercurrent and around the equatorial cool-tongue. Micrograzers are very important in the tropics as are predatory mesozooplankton. Up to 70% of the daily primary productivity is consumed by microzooplankton, which thus regulates the phytoplankton stocks. Micrograzers are an important link between primary producers, including bacteria, and mesozooplankton, constituting up to 80% of mesozooplankton food. Oceanography affects zooplankton trophic relationships through spatial–temporal effects on primary productivity and on the distributions of metabolic factors, food organisms, and predators. This paper is part of a comprehensive review of the oceanography of the eastern tropical Pacific.     

Ecohydrographic constraints on biodiversity and distribution of phytoplankton and zooplankton in coral reefs of the Red Sea,Saudi Arabia

An integral concept of ecological research is the constraint of biodiversity along latitudinal and environmental gradients. The Red Sea features a natural example of a latitudinal gradient of salinity, temperature and nutrient richness. Coral reefs along the Red Sea coasts are supported with allochthonous resources such as oceanic and neritic phytoplankton and zooplankton; however, relatively little is known about how the ecohydrography correlates with plankton biodiversity and abundance. In this article we present the biodiversity of phytoplankton and zooplankton in Red Sea coral reefs. Oceanographic data (temperature, salinity), water samples for nutrient analysis, particulate organic matter, phytoplankton and zooplankton, the latter with special reference to Copepoda (Crustacea), were collected at nine coral reefs over ~1500 km distance along the Red Sea coast of Saudi Arabia. The trophic state of ambient waters [as indicated by chlorophyll a (Chl a)] changed from strong oligotrophy in the north to mesotrophy in the south and was associated with increasing biomasses of Bacillariophyceae, picoeukaryotes and Synechococcus as indicated by pigment fingerprinting (CHEMTAX) and flow cytometry. Net‐phytoplankton microscopy revealed a Trichodesmium erythraeum (Cyanobacteria) bloom north of the Farasan Islands. Several potentially harmful algae, including Dinophysis miles and Gonyaulax spinifera (Dinophyceae), were encountered in larger numbers in the vicinity of the aquaculture facilities at Al Lith. Changes in zooplankton abundance were mainly correlated to the phytoplankton biomass following the latitudinal gradient. The largest zooplankton abundance was observed at the Farasan Archipelago, despite high abundances of copepodites, veligers (Gastropoda larvae) and Chaetognatha at Al Lith. Although the community composition changed over latitude, biodiversity indices of phytoplankton and zooplankton did not exhibit a systematic pattern. As this study constitutes the first current account of the plankton biodiversity in Red Sea coral reefs at a large spatial scale, the results will be informative for ecosystem‐based management along the coastline of Saudi Arabia.     

Brown trout (Salmo trutta) removal by rotenone alters zooplankton and phytoplankton community composition in a shallow mesotrophic reservoir

Brown trout (Salmo trutta) are known to have effects on multiple trophic levels in New Zealand streams, but their impacts on lower trophic levels are less well understood within lentic systems. We examined the effects of brown trout removal using rotenone on zooplankton and phytoplankton community composition in the Upper Karori Reservoir, New Zealand. Significant shifts were observed in zooplankton and phytoplankton composition following removal of brown trout from the reservoir. Shifts in zooplankton community composition did not occur immediately following trout removal (February), but instead followed the likely timing of galaxiid spawning (July). The removal of brown trout likely resulted in reduced predation pressure on galaxiids. A major change occurred in the zooplankton community with the dominance shifting from larger crustaceans to smaller rotifers, indicating an increased predation pressure from the larval native galaxiid. A delayed response in zooplankton community composition change indicates rotenone was not the direct cause of this. A major shift in phytoplankton community composition occurred immediately following trout removal. This was not consistent with the trophic cascade hypothesis of reduced grazing pressure from larger zooplankton due to increased galaxiid predation as a result of brown trout removal.     

The influence of hydrography on the distribution of phytoplankton in the southern Taiwan Strait

During the period August 1985 to May 1986, phytoplankton in the southern Taiwan Strait was collected and studied for distributional variability in relation to hydrography. The results indicated that maximum standing crops of phytoplankton occurred in October and May due to the outgrowth of certain species of diatoms and blue-green algae. The majority of phytoplankton appeared in the water in the top 25 m and occurred in distinct clusters under the influence of water movement. Multivariate analysis indicated that hydrographic parameters, which accounted for the variability of phytoplankton distribution, varied seasonally. Vertical, spatial and temporal variabilities were also apparent. The close relationship between hydrography and algal distribution justifies the use of variations in the phytoplankton population as a useful tracer of water movement.     

Water quality and phytoplankton communities in Lake Pontchartrain during and after the Bonnet Carré Spillway opening,April to October 2008, in Louisiana,USA

The Bonnet Carré Spillway, located 28 miles northwest of New Orleans, was constructed in the early 1930s as part of an integrated flood-control system for the lower Mississippi River system. From 11 April to 8 May 2008, Mississippi River water was diverted through the spillway into the 629-square-mile Lake Pontchartrain, which is hydraulically connected to the Gulf of Mexico. On 8 April, prior to the opening of the spillway, water-quality instruments were deployed and recorded hourly measurements of water temperature, dissolved oxygen, specific conductance, pH, and nitrate. Discrete water-quality and phytoplankton (algae) samples were collected in Lake Pontchartrain from 8 April to 3 October 2008 to assess the water-quality nutrient enrichment effects of the diversion on the lake. The maximum influence of river water in the southern portion of the lake was captured with continuous (hourly) monitoring of nitrate concentrations, and field measurements such as of specific conductance during the critical period in late April to early May. By late May, the deployed instruments had recorded the arrival, peak, and decline of selected constituents associated with the freshwater influx from the Mississippi River/Bonnet Carré Spillway diversion. The continuous monitoring data showed the short-term interactions of high-nitrate, low-specific conductance river water and low-nitrate, high-specific conductance lake water. The phytoplankton community composition, as an indicator of water quality, illustrated an extended response from the river water evident even after the continuous and discrete samples indicated that the lake had returned to pre-diversion conditions. The initial phytoplankton community response to nutrient increases was related to accumulations of diatoms. During periods of low nutrient concentrations, accumulations of blue-greens occurred by July and August. As blue-green algae cell densities and biovolumes increased in the summer, so did the species richness of blue-green algae, particularly the harmful algae bloom taxa. Cell densities and biovolume of the phytoplankton lake indicator taxa Skeletonema costatum, Anabaena sp., and Cylindrospermopsis raciborskii were highest and dominated the diatom and blue-green algae communities during the period of most river water influence on the lake and immediately following the freshwater inflows. The dominance and recession of these indictor taxa reflect the dramatic changes that occurred in the phytoplankton community in response to an increase in nutrient-rich freshwater from the diversion into the lake, and not normal seasonal phytoplankton compositional differences. Water-quality data indicated a gradual reversion to pre-diversion lake conditions by June to July, but shifts in the phytoplankton composition were still evident through August 2008. Observations from this study were similar to results from previous studies of Mississippi River/Bonnet Carré Spillway diversion opening in 1997.     

长江口邻近海域浮游植物粒级结构特征及其与环境的响应关系

浮游植物的分粒级研究是监测浮游植物特征的重要工具,对于深入了解浮游植物动态的作用也不容忽视。本文的研究目的在于揭示长江口邻近海域春秋季不同粒级的浮游植物分布动态,分析浮游植物粒级结构与环境因素以及浮游动物群落结构的关系。通过2010年春季和秋季对粒径分级叶绿素a浓度的现场调查研究发现：春季,浮游植物主要以微型浮游植物占优势;秋季,微微型浮游植物和微型浮游植物共同占优势。相关分析结果表明,温度和富营养化状况是影响微型和微微型浮游植物对总叶绿素a贡献的重要因素。浮游动物的摄食压力可能对小型浮游植物对总叶绿素a的贡献起着重要的作用。     

Trends in the variation of the sea level in the lagoons of the Southeastern Baltic

This paper analyses the global tendency of the sea level rise (SLR) and its long term influence on the sea level upstream drainage cascade based on the example of the level’s variation in the Vistula Lagoon of the Baltic Sea compared to the other lagoons and coastal regions of the southeastern part of the Baltic Sea. A steady positive trend in the water level variations was revealed; its magnitude varies significantly depending on the time period. In general, during the 100–150 year period, the rate of the SLR in the lagoons and coastal areas of the Baltic Sea (1.7–1.8 mm per year) is close to the SLR rate in the World Ocean. In the second half of the 20th century, the increased rate of the SLR in the lagoons and marine areas became stronger (up to 3.6 mm per year in the Vistula Lagoon and in 1959–2006 in the sea and exceeded the rate of global ocean SLR). It dramatically increased at the end of the last century both in the lagoons and in the sea (up to 10.0–15.0 mm per year). This is the response not only to the global climate warming but it is likely that it is also a response to the changes of the climate driving forces that influence the regimes of the local wind and precipitation in the catchment.     

Trophic structure and flows of energy in the Huizache–Caimanero lagoon complex on the Pacific coast of Mexico

The Huizache–Caimanero coastal lagoon complex on the Pacific coast of Mexico supports an important shrimp fishery and is one of the most productive systems in catch per unit area of this resource. Four other less important fish groups are also exploited. In this study, we integrated the available information of the system into a mass-balance trophic model to describe the ecosystem structure and flows of energy using the E approach. The model includes 26 functional groups consisting of 15 fish groups, seven invertebrate groups, macrophytes, phytoplankton, and a detritus group. The resulting model was consistent as indicated by the output parameters. According to the overall pedigree index (0.75), which measures the quality of the input data on a scale from 0 to 1, it is a high quality model. Results indicate that zooplankton, microcrustaceans, and polychaetes are the principal link between trophic level (TL) one (primary producers and detritus) and consumers of higher TLs. Most production from macrophytes flows to detritus, and phytoplankton production is incorporated into the food web by zooplankton. Half of the flow from TL one to the next level come from detritus, which is an important energy source not only for several groups in the ecosystem but also for fisheries, as shown by mixed trophic impacts. The Huizache–Caimanero complex has the typical structure of tropical coastal lagoons and estuaries. The TL of consumers ranges from 2.0 to 3.6 because most groups are composed of juveniles, which use the lagoons as a nursery or protection area. Most energy flows were found in the lower part of the trophic web.     

Role of mysid seasonal migrations in the organic matter transfer in the Curonian Lagoon,south-eastern Baltic Sea

We investigated population dynamics, growth and feeding of an omnivorous mysid, Paramysis lacustris, performing seasonal horizontal migrations in the Curonian Lagoon. In autumn, increased predation pressure and deteriorated feeding conditions, due to zooplankton and phytoplankton decline in the open water, forces mysids to migrate to the shoreline, where large amounts of decaying submersed vegetation occur at the end of plant growth season. Using stable isotope analysis and mixing models, we evaluated relative importance of decaying submersed macrophytes, lagoon phytoplankton and mesozooplankton in the diet of mysids during this period. In September–November, mysids actively grew and produced three cohorts. Specific growth rates of adults (10.9 ± 1.9 mm) and subadults (6 ± 0.5 mm) were 3 and 9% DW d⁻¹, respectively, resulting in population somatic production of 36 g DW m⁻². Both δ¹⁵N and δ¹³C of mysids varied in concert with those in suspended and bottom particulate organic matter (POM) and mesozooplankton. The mixing models estimated that 45% of mysid diets were composed of bottom POM originated from the decaying macrophyte Potamogeton perfoliatus, 45% by suspended POM largely consisting of phytoplankton, while mesozooplankton contributed less than 10% of the diet. This diet composition differs from that in summer, when mysids rely mostly on zooplankton and phytoplankton. Therefore, mysid horizontal migrations and their ability to efficiently utilize littoral detritus improve the efficiency of macrophyte detritus transfer up the food chain and inshore/offshore habitat coupling in the Curonian Lagoon.     

Species Composition and Distribution of Sibling Species of the Genus <Emphasis Type="Italic">Chironomus</Emphasis> Meigen (Diptera: Chironomidae) in the Curonian and Vistula Lagoons,Baltic Sea

The species compositions of the genus Chironomus in the Curonian and Vistula lagoons are very similar, represented by Ch. plumosus and Ch. balatonicus. In the Curonian Lagoon, in addition to these two species, Ch. muratensis and the first-generation interspecific hybrid Ch. muratensis × Ch. plumosus are also found. In this lagoon, Ch. plumosus occurs most frequently (84%); Ch. balatonicus is considered a subdominant species (24%). In the Vistula Lagoon, on the contrary, the high frequency of occurrence is typical of Ch. balatonicus (89%); for Ch. plumosus, it reaches only 11%. The Ch. plumosus population occupies most of the Curonian Lagoon; Ch. balatonicus is present only in its northern part, from the village of Juodkrante to the city of Klaipeda. In the Vistula Lagoon, Ch. balatonicus is distributed over almost its entire area, while Ch. plumosus is found locally, only near the mouth of the Nogat River. The distribution of the sibling species in the lagoons is determined by the salinity gradient.     

印太交汇区浮游植物和浮游动物生态学研究进展

海洋浮游植物和浮游动物是海洋生态系统中的重要组成部分,支撑了整个海洋生态系统的正常运转。因此,海洋浮游植、动物的生态学研究有利于我们全面认识和了解一个海洋生态系统的状况。印太交汇区作为全球最大的海洋生物多样性中心,是国际上生物多样性研究的热点区域,但该区域对浮游生物生态学方面的研究较少,不利于我们深入认识该区域生物多样性中心形成的生态机制。本文针对当前国内、外关于印太交汇区浮游植、动物生态学研究的进展进行综述,介绍了印太交汇区的浮游植物群落结构、生物量、粒级组成和初级生产力水平,以及浮游动物群落结构特征、生物量分布及影响因素等,对未来印太交汇区浮游生态学研究的方向进行了展望,希望可以为该区域相关生态学研究提供借鉴和参考。     

Differences in the structure of the lower trophic levels of pelagic ecosystems in the eastern and western subarctic Pacific

To illustrate areal differences in the structure of lower trophic levels of the pelagic ecosystems in the subarctic Pacific, data collected in the quasi-steady state summer/fall conditions were analysed for five areas, i.e. the Bering Basin, Western Subarctic Gyre, the area south of the Aleutians, the Gulf of Alaska, and the Oyashio Region. Average values of stock size of four components of the lower trophic levels showed a clear difference between areas with ranges of 7.5-fold for nitrate, 3.0 for chlorophyll a, 9.9 for microzooplankton, and 2.4 for mesozooplankton. Such differences were more striking when the structure of the lower trophic levels was expressed as a biomass pyramid. In the Gulf of Alaska, Western Subarctic Gyre, and south of the Aleutians, the relative biomass of microzooplankton to phytoplankton is large and large amounts of nitrate remained unused. In addition to possible iron limitation, grazing control by the microzooplankton on small phytoplankton must be substantial in these areas. Conversely, in the Oyashio Region, the nitrate stock is very small indicating higher efficiency of nitrate consumption by phytoplankton. However, since phytoplankton and zooplankton stocks are not particularly large, their products are likely to be transferred, also efficiently, to the higher trophic levels such as planktivorous pelagic fish. The situation in the Bering Basin is intermediate between the Oyashio Region and the other three areas. Inter-annual fluctuations in stock size of the planktivorous fish which migrate into the Oyashio Region in summer/fall were quite large. However, the inter-annual variation of mesozooplankton biomass was small, suggesting the existence of certain mechanisms to stabilize plankton abundance under increasing predation pressure. As a result, the increasing fish stocks likely keep the transfer efficiency from nitrate through to fish higher, at least in the Oyashio Region.     

基于碳、氮稳定同位素的雅浦海沟底栖生物食物来源和营养级初探

为了分析雅浦海沟中底栖生物群落的食物来源和营养级,本研究分析了雅浦海沟真光层中浮游植物和浮游动物、海底沉积物和巨型底栖生物（海绵、海参、海蛇尾、海星、海葵和钩虾）中的碳、氮稳定同位素组成。研究发现雅浦海沟真光层中的浮游植物和浮游动物δ13C值[（-22.8±0.4）‰和（-21.8±0.8）‰]和δ15N值[（5.4±0.4）‰和（6.8±0.2）‰]与巨型底栖生物的δ13C值（-20.1‰~-16.8‰）和δ15N值（11.9‰~17.9‰）的差异超过了一个营养级,表明作为底栖生物的初始食物来源的浮游植物和浮游动物在向下输送的过程中经历了食物链传递和细菌的降解。巨型底栖生物的δ15N和δ13C值之间无显著的相关性,此外不同物种之间营养级也存在明显差异,表现为海绵的营养级相对较高（3.4~4.7）,海参（3.3~3.6）、海蛇尾（3.4~3.5）和海星（3.2~3.7）的营养级较为接近,钩虾（2.9~3.3）和海葵（3.1）的营养级则相对略低,反映了底栖生物不同物种之间食物来源的多样化。     

Trophic structure and functioning in a eutrophic and poorly flushed lagoon in southwestern Taiwan

Tapong Bay, a eutrophic and poorly flushed tropical lagoon, supports intensive oyster culture. Using the Ecopath approach and network analysis, a mass-balanced trophic model was constructed to analyze the structure and matter flows within the food web. The lagoon model is comprised of 18 compartments with the highest trophic level of 3.2 for piscivorous fish. The high pedigree index (0.82) reveals the model to be of high quality. The most-prominent living compartment in terms of matter flow and biomass in the lagoon is cultured oysters and bivalves, respectively. The mixed trophic impacts indicate that phytoplankton and periphyton are the most-influential living compartments in the lagoon. Comparative analyses with the eutrophic and well-flushed Chiku Lagoon and non-eutrophic tropical lagoons show that high nutrient loadings might stimulate the growth and accumulation of phytoplankton and periphyton and therefore support high fishery yields. However, net primary production, total biomass, fishery yields per unit area, and mean transfer efficiency of Tapong Bay were remarkably lower than those of Chiku Lagoon. The lower transfer efficiency likely results from the low mortality of cultured oysters and invasive bivalves from predation or the lower density of benthic feeders constrained by the hypoxic bottom water as a result of poor flushing. This might therefore result in a great proportion of flows to detritus. However, the hypoxic bottom water might further reduce the recycling of the entering detritus back into the food web. In contrast to many estuaries and tropical lagoons, poor flushing of this eutrophic tropical lagoon might induce a shift from detritivory to herbivory in the food web.     

Structure and function of contemporary food webs on Arctic shelves: A panarctic comparison: The pelagic system of the Kara Sea – Communities and components of carbon flow

After a short introduction to the physical setting and the history of biological research the pelagic ecosystem of the Kara Sea is described. Main emphasis is on regional aspects of the plankton communities and their seasonal dynamics using mostly data collected between 1996 and 2001. In the zooplankton, for which most data were available, four regional aggregations were separated: (1) the rivers and estuaries of the Southern Kara Sea, (2) the south-western and (3) the central Kara Sea, and (4) the northern troughs and slope. The phytoplankton communities had a similar distribution. To provide components for detailed carbon budgets the regional dynamics of bacterial, phytoplankton and zooplankton biomass and production are described and carbon requirements of bacteria and zooplankton are estimated. For completeness a short literature review on higher trophic levels is included. Finally, recent observations of the pelago-benthic coupling are considered. Estimates of the carbon requirements from the plankton and benthos reveal a large underestimation of primary production, which to date, together with seasonal aspects, shows the largest gap in our knowledge.     

Comparative trophodynamics of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela: are species alternations between small pelagic fish trophodynamically mediated?

The results of detailed morphological, experimental, field and modelling studies on various aspects of the trophic ecology of sardine Sardinops sagax and anchovy Engraulis encrasicolus in the Benguela ecosystem are synthesised, and differences in the trophodynamics of these two species are highlighted. Anchovy possess a relatively coarse branchial apparatus; feed predominantly by particulate-feeding and maximise their net energetic gain by using this feeding mode; show higher weight-standardised clearance rates than do sardine for prey >580μm; are most efficient at assimilating nitrogen from zooplankton and excrete <50% of ingested nitrogen; feed inefficiently on phytoplankton and derive the bulk of their dietary input from larger zooplankton; and maximise their scope for growth on mesozooplankton. In contrast, sardine possess a relatively fine branchial apparatus; feed predominantly by filter-feeding and maximise their net energetic gain by using this feeding mode; show higher weightstandardised clearance rates than do anchovy for prey <580μm; are most efficient at assimilating nitrogen from zooplankton but excrete >50% of ingested nitrogen; are able to feed on phytoplankton but derive the bulk of their dietary input from smaller zooplankton; and maximise their scope for growth on microzooplankton. These differences provide compelling evidence that anchovy and sardine are trophically distinct, and indicate that the two species show resource partitioning based on zooplankton size. The implications of these trophic differences for ecosystem functioning are discussed, and support the hypothesis that species alternations between anchovy and sardine, both in the southern Benguela and in other upwelling ecosystems, are likely to be trophodynamically mediated.     

1996年春季副热带环流区浮游植物生态的初步研究

鉴定浮游植物种类54属184种(含变种和变型),其中暖水种为140种(占76.5%).硅藻类和甲藻类等浮游植物的平均总细胞密度为198.71×102个/m3;蓝藻类藻丝体平均密度为44.55×102条/m3.浮游植物各类别的分布与水域的磷酸盐含量、黑潮以及其他水系(涌升水和沿岸水等)等相关;蓝藻类的束毛藻(Trichodesmium)的分布还与50m以浅水体的平均水温关系密切.     

Modeling the seasonal variability of marine ecosystem in the region of the Central-Eastern Atlantic

A 3D eco-hydrodynamical model of high resolution (0.25° × 0.25°, 27 σ-levels) is used to simulate the seasonal variability of the ocean circulation and marine ecosystem in the Central-Eastern Basin of the North Atlantic including the Canary upwelling system. According to the model results, in the winter period, the “patches” of maximal phytoplankton and zooplankton biomass are often located in upwelling zones in the open ocean on the periphery of cyclonic eddies rather than in the coastal upwelling zones. In the summer period, when the phytoplankton biomass reaches maximal (in the annual cycle) values, the maxima of the phytoplankton are located in the coastal upwelling zones. As shown, there is no simple relationship between the nitrate distributions, on the one hand, and the phytoplankton and zooplankton ones, on the other hand.     

山东省大中型水库浮游生物研究──Ⅰ．浮游动物生态

报道１９８９～１９９０年山东省３０多个大中型代表水库浮游动物的种类组成，种类频级、优势类群。主要种和生物量；统计分析掠食性与植食性浮游动物生物量，浮游动物与浮游植物生物量及浮游动物与代表水库营养状态指标的总磷，透明度和深度的关系。提出浮游动物个体越小，其生物量与水库营养水平关系越密切。