Error quantification of a high resolution coupled hydrodynamic-ecosystem coastal-ocean model: Part3, validation with Continuous Plankton Recorder data

Data collected during the Continuous Plankton Recorder (CPR) survey has been used to validate a three-dimensional hydrodynamic ecosystem model simulation of the North-west European Shelf for the years 1988–89. The CPR time series is unique to the North Atlantic region as a validation tool. Data were extracted from the model to correspond with those collected by the CPR survey, and both the model and survey plankton data were standardised to allow the comparison of model biomass with survey counts. Simple linear regression and absolute error maps provide a qualitative evaluation of spatio-temporal model performance of simulated diatoms, flagellates, total phytoplankton and omnivorous mesozooplankton. Comparisons of z-scores indicate that the model reproduces the main pelagic seasonal features, and there is good correlation between magnitudes of these features with respect to standard deviations from a long-term mean. The model is replicating up to 62% of the mesozooplankton seasonality across the domain, with variable results for the phytoplankton. There are, however, differences in the timing of patterns in plankton seasonality. The validation exercise has highlighted that the spring diatom bloom in the model is too early, suggesting the need to reparameterise the response of phytoplankton to changing light levels in the model. Errors in the north and west of the domain imply that model turbulence and vertical density structure need to be improved to more accurately capture plankton dynamics.     

Evidence of episodic changes in deep-sea mesozooplankton abundance and composition in the Levantine Sea (Eastern Mediterranean)

Mesozooplankton samples from the 4270-m deep Ierapetra Basin in the oligotrophic Levantine Sea have revealed a strong temporal variability in abundance and composition hitherto unknown for the deep sea pelagic zone. The phenomenon reflected by a survey in June 1993 as compared to January 1987 is assessed by (i) the increase of the mesozooplankton standing crop throughout the water column, (ii) the absence of a decline in mesozooplankton abundance with increasing depth below 1000 m, (iii) the outstanding dominance of two interzonal calanoid copepods at bathypelagic and abyssopelagic depths and (iv) a substantial faunal change due to the codominance of Calanus helgolandicus, which was found for the first time in the area. The underlying hydrographical factors are addressed in the light of the Eastern Mediterranean Transient (EMT). Apart from a possible long-term faunal change, the EMT may exaggerate episodic plankton blooms and surface-abyssopelagic coupling in space and time.     

A model study with light-dependent mortality rates of copepod stages

This paper is based on an advanced ecosystem model of the Baltic Sea (ERGOM [J. Mar. Sys. 25 (3–4) (2005) 405]), but with an increased resolution of the zooplankton stage variable [J. Plankton Res. 23 (2001) 1217; ICES Marine Science 219 (2003) 208]. The model copepods are represented by five stages: eggs, an aggregated variable of nauplii, two aggregated groups of copepodites and adults. The transfer among the stages, i.e., hatching, molting and reproduction, is controlled by food availability and temperature.As usual, the model food web is truncated at the level of zooplankton. The study explores the effects of different parametrization of zooplankton mortality and looks in particular on light-dependent rates. The light climate may serve a proxy for the effects of visual feeding of fish larvae and fish. Different choices of the mortality parameters can result in remarkable differences in abundances and biomass of the model zooplankton and in the timing of its development.It is found that the different choices of mortality affect the development of populations in several ways: Relative small initial differences of abundances at the beginning of the spring bloom are important for the development of the model populations. Higher mortality rates are less important at food rich conditions than at scarce resources. At low phytoplankton levels, the individual development of the copepods through the stages can be faster for elevated mortality rates because then less animals have to share the available food.     

Sea ice and the onshore–offshore gradient in pre-winter zooplankton assemblages in southeastern Beaufort Sea

Zooplankton communities were studied in southeastern Beaufort Sea (Arctic Ocean) in September–October 2002. Cluster analysis and non-metric multidimensional scaling revealed three distinct mesozooplankton assemblages. A neritic assemblage occurred on the Mackenzie Shelf and in Franklin Bay, while distinct off-shelf assemblages prevailed in the Cape Bathurst Polynya and on the Beaufort Slope respectively. Over 95% of the mesozooplankton was comprised of eight copepod taxa. Pseudocalanus spp. contributed predominantly to the discrimination of the three assemblages and was the only significant indicator of the Shelf assemblage. Oithona similis, Oncaea borealis, Metridia longa and Calanus hyperboreus were indicators of the Polynya assemblage. Cyclopina sp. and Microcalanus pygmaeus were indicative of the overall off-shelf community (Polynya and Slope assemblages). The importance of omnivores and carnivores increased from the shelf to the polynya and the slope. Station depth and duration of reduced ice conditions during summer (< 50% ice concentration) underpinned the distribution of the assemblages (r² = 0.71 and 0.45 respectively). The abundance of Pseudocalanus spp. was independent of depth and increased with the duration of reduced ice conditions (r_s = 0.438). The abundance of Cyclopina sp., M. pygmaeus and other indicators of the offshore assemblages followed the opposite trend (r_s = − 0.467 and − 0.5 respectively). Under continued climate warming, a reduction of the ice cover will affect the biogeography of mesozooplankton on and around the Mackenzie Shelf, to the potential advantage of Pseudocalanus spp. and other calanoid herbivores.     

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.     

Top-down effects and carbon flux in the ocean: A hypothesis

The close relationships between mesozooplankton, chlorophyll, and primary production normally observed across latitudes may be regarded as a bottom-up effect of increased productivity or, conversely, as a top-down effect of mesozooplankton on microzooplankton, releasing phytoplankton growth. The time-course of primary producers and mesozooplankton observed during the Late Winter Bloom in subtropical waters suggests a top-down effect, similar to the results found in lakes and mesocosm experiments. Two scenarios were observed during the development of the bloom in the Canary Island waters. The first was the increase in mesozooplankton as the effect of higher primary production due to mixing. The second scenario was the decrease in mesozooplankton due to the top-down effect of predation by diel vertical migrants upon epipelagic mesozooplankton. The structure of the food web in the euphotic zone was different depending on the presence or not of mesozooplankton, suggesting important consequences for the transport of carbon in those large areas of the ocean.     

Hydrographic structure and zooplankton abundance and diversity off Paita, northern Peru (1994 to 2004) — ENSO effects, trends and changes

The objective of the present study was to verify possible spatial, seasonal, and inter-annual changes in the zooplankton off Paita (northern Peru), an upwelling area located closely to the limits of cold Humboldt Current and warm Equatorial Surface Waters. Zooplankton was sampled at subsurface on 53 occasions from August 1994 to December 2004 at four stations located 2 to 30 km offshore with a WP-2 net (300 µm). Extremely high surface water temperatures combined with low salinities were observed during the 1997/98 El Niño up to 29.0 °C) and in April 2002 (up to 25.0 °C). Temperatures more than 2 °C above monthly average were also observed in October 1994, in April 2000, and in November 2004. Significant trends were observed for oxygen concentration (increase) and several horizontal and vertical gradients. Among the copepods (72% of all individuals), the most abundant species were Paracalanus parvus (28%), Acartia tonsa (26%), and Calanus sp. (10%). The strong 1997–98 El Niño (EN) event led to drastic changes in species composition that were reversed during the 1998–99 La Niña (LN) event. Community parameters such as total abundance, diversity, equitability and species richness displayed marked variations associated with the 1997–98 EN and long-term trends. Long-term trends were significant for several vertical and horizontal temperature and oxygen gradients, indicating an increase in upwelling intensity at the shelf during the study period. 10-year-trends were also significant for total zooplankton abundance (increase) and community evenness (J, decline). Our data confirmed the importance of the weak EN in 2002/03 for the study region. Within the trend of increasing zooplankton abundance, a sharp step or shift was observed from 1999 to 2000. When using sequential t-tests to detect shifts in (x + 1) transformed abundance data, a significant rupture was found between the last sampling in 1999 and the first sampling in 2000. Also, a substantial decrease in diel variability occurred after 1999, probably due to changes in vertical migration patterns. The considerable increase in zooplankton abundance over the study period, the ENSO effect, and the 1999–2000 transition are discussed with regard to synchronicity with other zooplankton time series. The present study contributes with the first evidence from an important area located in the Humboldt Current for synchronous trends and changes that were previously observed elsewhere in the Pacific. Our results demonstrated the importance of long-term zooplankton monitoring studies in upwelling areas, and confirms the idea of dramatic changes in pelagic ecosystem structure occurring in the East Pacific.     

A “horizontal Sverdrup mechanism” may control the spring bloom around small oceanic islands and over banks

One of the most basic paradigms in marine ecology is the “Sverdrup mechanism,” where the spring bloom is retarded if the surface mixed layer is too deep, due to the algae being mixed vertically out of the euphotic layer. A similar mechanism may operate in vertically homogeneous water over a small shallow area, surrounded by deep waters, if the horizontal exchange is sufficiently intense. In systems with inter-annual variations in the horizontal exchange rate, this may induce inter-annual variations in the timing and intensity of the spring bloom. A numerical primary production model with circular symmetry and prescribed horizontal exchange rate is developed. Using the Faroe Shelf as an example, the model and observations show that the timing and intensity of the spring bloom in the shallow parts of the system may be critically dependent upon the rate of the horizontal exchange. Numerical experiments confirm that the effect of horizontal exchange on the spring bloom is less pronounced for banks than for shelf systems around islands of similar scales, and that the effect increases in importance as the horizontal scale is reduced.