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.     

Seasonal variations of water system distribution and flow patterns in the southern sea area of Hokkaido,Japan

Hydrographic data and composite current velocity data (ADCP and GEK) were used to examine the seasonal variations of upper-ocean flow in the southern sea area of Hokkaido, which includes the “off-Doto” and “Hidaka Bay” areas separated by Cape Erimo. During the heating season (April–September), the outflow of the Tsugaru Warm Current (TWC) from the Tsugaru Strait first extends north-eastward, and then one branch of TWC turns to the west along the shelf slope after it approaches the Hidaka Shelf. The main flow of TWC evolves continuously, extending eastward as far as the area off Cape Erimo. In the late cooling season (January–March), part of the Oyashio enters Hidaka Bay along the shallower part of the shelf slope through the area off Cape Erimo, replacing almost all of the TWC water, and hence the TWC devolves. It is suggested that the bottom-controlled barotropic flow of the Oyashio, which may be caused by the small density difference between the Oyashio and the TWC waters and the southward migration of main front of TWC, permits the Oyashio water to intrude along the Hidaka shelf slope.     

Winter-spring phytoplankton blooms in Dabob Bay, Washington

Scientific investigations in Dabob Bay, Washington State, USA, have been extensive since the early 1960s, but phytoplankton blooms have been studied mostly with regard to chlorophyll concentrations and little is known about the phytoplankton species themselves. Here we provide information on the species present, their abundances during blooms, their contribution to organic carbon concentrations and the ability of some phytoplankton species to produce toxic aldehydes that may impact metazoan grazers.Multiple blooms of phytoplankton, dominated by diatoms, occurred in the late winter-early spring period, with depth-integrated chlorophyll levels ranging from <20 to 230 mg m⁻² and peaks in February and April. The major bloom species included Skeletonema costatum, Thalassiosira spp. and Chaetoceros spp; Phaeocystis cf. pouchetii occurred in 2002 and 2004. Other taxa or groups of organisms that were sometimes abundant included unidentified small flagellates <10 μm in size and unidentified heterotrophic dinoflagellates. Large diatoms usually comprised most of the cell carbon, but a large, heterotrophic dinoflagellate, identified only as Gyrodinium “tear” because of its shape, was a major contributor to the microplankton carbon when present even in small numbers. Five Thalassiosira species and S. costatum were found to produce polyunsaturated aldehydes (PUA) that are known to affect copepod reproduction and hatching success. Our findings are similar to the few previous studies in the last four decades that included phytoplankton species and suggest long-term similarities and relative stability in the phytoplankton species present and their timing in Dabob Bay.     

The fate of dumped sediments monitored by a high-resolution multibeam echosounder system,Weser Estuary,German Bight

With the development of high-resolution multibeam echosounder systems (MBES) for surveying shallow-water areas a new tool is available to monitor rapid changes in seabed morphology as, e.g., caused by the dumping of dredge spoil in coastal waters. In this study, four data sets of repeated bathymetric surveys with a MBES were processed and analyzed. The data were collected in a 1.94-km² dumping site in the outer Weser Estuary (German Bight). Between June and December 1998, 2.6 million m³ of dredged sediment were deposited there. The bathymetric maps generated in the course of this study reveal features such as subaqueous dunes, scour holes, and mounds of dumped dredge spoil. The mean water depth decreased by about 1 m during the dumping period. Furthermore, difference grids showing changes in sediment volume allowed a calculation of the sediment budget for the monitored area. After a time period of only 5 months, 0.5 million m³ of the originally dumped 2.6 million m³ of dredge spoil had already been removed from the dumping site.     

Effects of water-borne copper on metallothionein and lipid peroxidation in the marine amphipod Gammarus locusta

The aim of the current work was to determine over 10 days the effects of water-borne exposure of the marine amphipod Gammarus locusta to 4 microgCu l(-1) on the metallothionein (MT; measured by differential pulse polarography) protection system and lipid peroxidation (LP: thiobarbituric acid-reactive malondialdehyde equivalents) as a measure of oxidative damage. MT levels in exposed animals increased significantly at day 2 (36% > control; P < 0.001) and remained high at days 6 and 10 (55 and 38%, respectively, P < 0.001). The maximum level of MT at day 6 coincided with the highest Cu body-burden. LP increased within I day of exposure, indicative of Cu as an oxidative stressor. However, in contrast to MT, the highest LP level was seen at day 4 (68% > control, P < 0.001) before returning to control values by day 6, indicating a protective role of MT against the pro-oxidant effects of Cu.     

Ecology of Quaternary benthic foraminiferal assemblages on the Amazon shelf,northern Brazil

The distribution of benthic foraminiferal microfauna recovered from two offshore transects on the Amazon shelf was studied. Eight box-core samples were collected along the inner, middle, and outer parts of the continental shelf. Each box-core sample was divided into 10 subsamples, of 1- to 5-cm intervals, in order to investigate the foraminiferal microhabitat preferences. The variations of salinity, oxygen, lithology, and sediment accumulation rates were used to evaluate ecological patterns. Four benthic foraminiferal assemblages identified in the study area characterize a record of oceanographic changes from early Holocene to the present. An abundant assemblage, with opportunistic species, was found near the river mouth. Two assemblages were identified northwest of the river mouth, with dominant epifaunal and infaunal species, respectively. A relict assemblage was encountered northwest of the river mouth.     

Metridia pacifica in Dabob Bay, Washington: The diatom effect and the discrepancy between high abundance and low egg production rates

Information on life cycle strategies and reproductive parameters of Metridia pacifica is scarce, despite its importance in the zooplankton of the subarctic Pacific. In many regions it occurs in high abundance, but reproductive rates, when reported, are usually low. This discrepancy was studied in Dabob Bay, Washington, USA, in the context of an investigation of the effect of diatom blooms on the reproductive success of copepod grazers.In situ egg production rates of M. pacifica were measured in spring and mid-summer with standard methods (multi-wells) and a new incubation chamber (spawning towers) that separates the spawning female from its eggs and allows the eggs to develop undisturbed. Many females did not produce eggs, possibly due to a high fraction of immature individuals. Egg production rates were variable, but clutch sizes were higher in spawning towers, and estimates of female egg cannibalism revealed that females consume many eggs shortly after their release. Thus, a separation of females and eggs is mandatory for accurate measurements of M. pacifica egg production rates. The maximum clutch sizes recorded in our study were comparable to measurements for other calanoids. However, unviable eggs were a large fraction of those spawned, independent of incubation method, especially in late winter and early spring. In order to assess whether the diatom effect may be responsible for low viability of embryos and nauplii, we also measured in situ grazing. Adult females were omnivorous, but they ingested some diatoms that rank among the strongest anti-mitotic toxin producers known so far. Although M. pacifica’s vertical migration behavior suggests opportunistic feeding on abundant food during their short stay in the phytoplankton-rich surface, they often ignored the food items that contributed most to microplankton carbon concentrations. Thus, their feeding strategy remains ambiguous. Due to severe reproductive failure early in the season, recruitment was impaired in spring, while the population increased, reaching high abundance in mid-summer when reproductive output was low. While advection and interannual variability in bloom conditions might compensate for the losses described here, the paradox of high Metridia abundance versus low reproductive success still requires further investigation, and methodological constraints need to be ruled out in future studies.     

Longshore drift estimation using fluorescent tracers: New insights from an experiment at Comporta Beach, Portugal

A fluorescent sand-tracer experiment was performed at Comporta Beach (Portugal) with the aim of acquiring longshore sediment transport data on a reflective beach, the optimization of field and laboratory tracer procedures and the improvement of the conceptual model used to support tracer data interpretation.

The field experiment was performed on a mesotidal reflective beach face in low energetic conditions (significant wave height between 0.4 and 0.5 m). Two different colour tracers (orange and blue) were injected at low tide and sampled in the two subsequent low tides using a high resolution 3D grid extending 450 m alongshore and 30 m cross-shore. Marked sand was detected using an automatic digital image processing system developed in the scope of the present experiment.

Results for the two colour tracers show a remarkable coherence, with high recovery rates attesting data validity. Sand tracer displayed a high advection velocity, but with distinct vertical distribution patterns in the two tides: in the first tide there was a clear decrease in tracer advection velocity with depth while in the second tide, the tracer exhibited an almost uniform vertical velocity distribution. This differing behaviour suggests that, in the first tide, the tracer had not reached equilibrium within the transport system, pointing to a considerable time lag between injection and complete mixing. This issue has important implications for the interpretation of tracer data, indicating that short term tracer experiments tend to overestimate transport rates. In this work, therefore, longshore estimates were based on tracer results obtained during the second tide.

The estimated total longshore transport rate at Comporta Beach was 2 × 10^− 3 m³/s, more than four times larger than predicted using standard empirical longshore formulas. This discrepancy, which results from the unusually large active moving layer observed during the experiment, confirms the idea that most common longshore transport equations under-estimate total sediment transport in plunging/surging waves.     

Vertical distribution of the macrobenthic community and its relationships to trace metals and natural sediment characteristics in the lower Douro estuary, Portugal

Vertical distribution (0–15 cm) of the macrobenthic community and its relationships to natural sediment characteristics and trace metal contents and bioavailability were studied at five locations in the lower Douro estuary, Portugal. An analysis of vertical metal distribution, for the interpretation of anthropogenic impact on the estuarine sediments, was also investigated. Sediment characterisation included organic matter, grain size, metals (Al, Fe, Cu, Pb, Cr, Ni, Cd, Zn and Mn), acid volatile sulphide (AVS) and simultaneously extracted metals (SEM). The macrobenthic community had low diversity (14 species), was dominated by small size opportunists and seemed to be controlled mainly by natural factors such as grain size distribution, Al and Fe contents and sediment depth. The vertically heterogeneous distribution of macrobenthic community appears to affect redox status of the sediments and consequently metal bioavailability. Despite anthropogenic contamination in terms of Zn, Cu, Pb, Cr and Ni having already been detected in the north bank, the analysis of vertical distribution was essential for the identification of current anthropogenic contamination in terms of Zn, Pb and Cd in the south bank.