Simultaneous enumeration of diatom, protozoa and meiobenthos from marine sediments using Ludox-QPS method

The Ludox-QPS method is a newly developed technique, which combines the Ludox HS 40 density centrifugation and quantitative protargol stain, to enumerate marine ciliates with good taxonomic resolution. We tested the method for simultaneous enumeration of diatoms, protozoa and meiobenthos and compared its extraction efficiency for meiobenthos with that of the routine Ludox-TM centrifugation and a modified protocol using Ludox HS 40. We conducted the evaluation with a sample size of 8.3 ml each from sandy, muddy-sand and muddy sediments collected from the intertidal area of the Yellow Sea in summer 2006 and spring 2007. The Ludox-QPS method not only produced high extraction efficiencies of 97 +/- 1.3% for diatoms and 97.6 +/- 0.8% for ciliates, indicating a reliable enumeration for eukaryotic microbenthos, but also produced excellent extraction efficiencies of on average 97.3% for total meiobenthos, 97.9% for nematodes and 97.8% for copepods from sands, muddy sands and mud. By contrast, the routine Ludox-TM centrifugation obtained only about 74% of total meiobenthos abundance with one extraction cycle, and the modified Ludox HS 40 centrifugation yielded on average 93% of total meiobenthos: 89.4 +/- 2.0% from sands, 93 +/- 4.1% from muddy sands and 97.1 +/- 3.0% from mud. Apart from the sediment type, sample volume was another important factor affecting the extraction efficiency for meiobenthos. The extraction rate was increased to about 96.4% when using the same modified Ludox centrifugation for a 4 ml sediment sample. Besides the excellent extraction efficiency, the Ludox-QPS method obtained higher abundances of meiobenthos, in particular nematodes, than the routine Ludox centrifugation, which frequently resulted in an uncertain loss of small meiobenthos during the sieving process. Statistical analyses demonstrated that there were no significant differences between the meiobenthos communities revealed by the Ludox-QPS method and the modified Ludox HS 40 centrifugation, showing the high efficiency of the Ludox-QPS method for simultaneous enumeration of diatom, protozoa and meiobenthos. Moreover, the comparatively high taxonomic resolution of the method, especially for diatoms and ciliates, makes it feasible to investigate microbial ecology at community level.

A new species of Tortanus (Atortus) (Copepoda: Calanoida) from the Seychelles, Mauritius and Madagascar

A new species of the planktonic copepod Tortanus (Atortus) (Calanoida: Tortanidae), T. (A.) magnonyx is described from Seychelles, Mauritius and Madagascar. This is the sixth species of the Indian Ocean recticauda species group, of the Indo-West Pacific recticauda species complex, that has been described from the western Indian Ocean. The inshore areas where these copepods are found have been poorly surveyed, so the number of species found implies a high diversity.

Epipelagic mesozooplankton distribution and abundance over the Mascarene Plateau and Basin, south-western Indian Ocean

The crescent shaped Mascarene Plateau (southwestern Indian Ocean), some 2200 km in length, forms a partial barrier to the (predominantly westward) flow of the South Equatorial Current. Shallow areas of the Mascarene Plateau effectively form a large shelf sea without an associated coastline. Zooplankton sampling transects were made across the plateau and also the basin to the west, to investigate the role the partial interruption of flow has on zooplankton biomass and community structure over the region. Biomass data from Optical Plankton Counter (OPC) analysis, and variability in community structure from taxonomic analysis, appear to indicate that the obstruction by the plateau causes upwelling, nutrient enrichment and enhanced chlorophyll and secondary production levels downstream. The Mascarene Basin is clearly distinguishable from the ridge itself, and from the waters to the south and north, both in terms of size-distributed zooplankton biomass and community structure. Satellite remote sensing data, particularly remotely-sensed ocean colour imagery and the sea surface height anomaly (SSHA), indicate support for this hypothesis. A correlation was found between OPC biovolume and SSHA and sea surface temperature (SST), which may indicate the physical processes driving mesozooplankton variability in this area. Biomass values away from the influence of the ridge averaged 24 mg m-3, but downstream if the ridge biomass averaged 263 mg m-3. Copepods comprised 60% of the mean total organisms. Calanoid copepods varied considerably between regions, being lowest away from the influence of the plateau, where higher numbers of the cyclopoid copepods Oithona spp., Corycaeus spp. and Oncaea spp., and the harpacticoid Microsetella spp. were found.

How well does the Continuous Plankton Recorder (CPR) sample zooplankton? A comparison with the Longhurst Hardy Plankton Recorder (LHPR) in the northeast Atlantic

The Continuous Plankton Recorder (CPR) survey has collected data on basin- scale zooplankton abundance in the North Atlantic since the 1930s. These data have been used in many studies to elucidate seasonal patterns and long-term change in plankton populations, as well as more recently to validate ecosystem models. There has, however, been relatively little comparison of the data from the CPR with that from other samplers. In this study we compare zooplankton abundance estimated from the CPR in the northeast Atlantic with near-surface samples collected by a Longhurst-Hardy Plankton Recorder (LHPR) at Ocean Weather Station India (59 degree N, 19 degree W) between 1971 and 1975. Comparisons were made for six common copepods in the region: Acartia clausi, Calanus finmarchicus, Euchaeta norvegica, Metridia lucens, Oithona sp. and Pleuromamma robusta. Seasonal cycles based on CPR data were similar to those recorded by the LHPR. Differences in absolute abundances were apparent, however, with the CPR underestimating abundances by a factor of between 5 and 40, with the exception of A. clausi. Active avoidance by zooplankton is thought to be responsible. This avoidance is species specific, so that care must be taken describing communities, as the CPR emphasises those species that are preferentially caught, a problem common to many plankton samplers.

Long-term changes in phytoplankton, zooplankton and salmon related to climate

Recently, large-scale changes in the biogeography of calanoid copepod crustaceans have been detected in the northeastern North Atlantic Ocean and adjacent seas. Strong biogeographical shifts in all copepod assemblages were found with a northward extension of more than ° in latitude of warm-water species associated with a decrease in the number of colder-water species. These changes were attributed to regional increase in sea surface temperature. Here, we have extended these studies to examine long-term changes in phytoplankton, zooplankton and salmon in relation to hydro-meteorological forcing in the northeast Atlantic Ocean and adjacent seas. We found highly significant relationships between (1) long-term changes in all three trophic levels, (2) sea surface temperature in the northeastern Atlantic, (3) Northern Hemisphere temperature and (4) the North Atlantic Oscillation. The similarities detected between plankton, salmon, temperature and hydro-climatic parameters are also seen in their cyclical variability and in a stepwise shift that started after a pronounced increase in Northern Hemisphere Temperature anomalies at the end of the 1970s. All biological variables show a pronounced change which started after circa 1982 for euphausiids (decline), 1984 for the total abundance of small copepods (increase), 1986 for phytoplankton biomass (increase) and Calanus finmarchicus (decrease) and 1988 for salmon (decrease). This cascade of biological events led to an exceptional period, which is identified after 1986 to present and followed another shift in large-scale hydro-climatic variables and sea surface temperature. This regional temperature increase therefore appears to be an important parameter that is at present governing the dynamic equilibrium of northeast Atlantic pelagic ecosystems with possible consequences for biogeochemical processes and fisheries.

Congruent trends in long-term zooplankton decline in the north-east Atlantic and basking shark (Cetorhinus maximus ) fishery catches off west Ireland

Trends in basking shark (Cetorhinus maximus) fishery catches off Achill Island, west Ireland between 1949 and 1975 were examined in relation to zooplankton (total copepod) abundance in four adjacent sea areas over a 27-year period. The numbers of basking sharks caught and copepod abundance showed downward trends and were positively correlated (r-value range, 0.44–0.74). A possible explanation for the downward trend in shark catches was that progressively fewer basking sharks occurred there between 1956 and 1975 because fewer copepods, their food resource, occurred near the surface off west Ireland over the same period. We suggest that the decline in basking sharks may have been due to a distributional shift of sharks to more productive areas, rather than a highly philopatric, localized stock that was over-exploited.

Spatial, seasonal and long-term fluctuations of plankton in relation to hydroclimatic features in the English Channel, Celtic Sea and Bay of Biscay

Plankton collected by the Continuous Plankton Recorder (CPR) survey were investigated for the English Channel, Celtic Sea and Bay of Biscay from 1979 to 1995. The main goal was to study the relationship between climate and plankton and to understand the factors influencing it. In order to take into account the spatial and temporal structure of biological data, a three-mode principal component analysis (PCA) was developed. It not only identified 5 zones characterised by their similar biological composition and by the seasonal and inter-annual evolution of the plankton, it also made species associations based on their location and year-to-year change. The studied species have stronger year-to-year fluctuations in abundance over the English Channel and Celtic Sea than the species offshore in the Bay of Biscay. The changes in abundance of plankton in the English Channel are negatively related to inter-annual changes of climatic conditions from December to March (North Atlantic Oscillation [NAO] index and air temperature). Thus, the negative relationship shown by Fromentin and Planque (1996; Mar Ecol Prog Ser 134:111-118) between year-to-year changes of Calanus finmarchicus abundance in the northern North Atlantic and North Sea and NAO was also found for the most abundant copepods in the Channel. However, the hypothesis proposed to explain the plankton/NAO relationship is different for this region and a new hypothesis is proposed. In the Celtic Sea, a relationship between the planktonic assemblage and the air temperature was detected, but it is weaker than for the English Channel. No relationship was found for the Bay of Biscay. Thus, the local physical environment and the biological composition of these zones appear to modify the relationship between winter climatic conditions and the year-to-year fluctuations of the studied planktonic species. This shows, therefore, that the relationship between climate and plankton is difficult to generalise.

Feeding of anchovy Engraulis encrasicolus larvae in the northwestern Adriatic Sea in response to changing hydrobiological conditions

Results from depth integrated and vertically stratified plankton sampling in the northwestern Adriatic Sea were used for comparison of gut contents of larvae of European anchovy Engraulis encrasicolus with composition and concentration of potential prey in the plankton. Sampling was carried out over a grid of stations both before and after a period of increased wind mixing to investigate changes in food availability and larval feeding success. All larvae had empty guts soon after dusk, indicating daytime feeding and rapid gut clearance. With increasing larval length there was a greater percentage of specimens with empty guts, despite suitable food being available in the plankton for these larger larvae; this suggests differential gut evacuation during sampling-possibly related to the degree of gut development. Larval diet was principally the various developmental stages of copepods, especially calanoid and cyclopoid nauplii, which were preferentially selected by larvae, whereas selection was against harpacticoid nauplii. Lamellibranch larvae and Peridinium were generally abundant in the plankton, but were only present in the gut contents in any number when the preferred dietary organisms were present in the plankton at low concentrations. The number of food organisms in the gut contents increased with concentration of the preferred food organisms in the plankton up to a limit of similar to 50 organisms/l. Within the upper 18 m of the water column, there was a reduction in the proportion of larvae with food in their guts with increasing depth, irrespective of the vertical profile of food concentration. Following a period of wind mixing the composition of the plankton changed. This was reflected in the diet of anchovy larvae, which altered in parallel. There was also an overall 41% decrease in concentration of the preferred food particles of larvae in the plankton following the period of wind mixing, but larvae were still able to maintain their food intake. These results show that anchovy larvae can successfully adapt their diet to a changing prey field and suggest that in the conditions observed in the northern Adriatic, quite radical changes in the feeding environment were probably insufficient to affect overall larval mortality.

Digestion of natural food by larval and post-larval turbot Scophthalmus maximus

The digestion of natural, mainly crustacean zooplankton, by different age groups of turbot Scophthalmus maximus larvae was evaluated by comparisons of visual appearance, dry weight and carbon and nitrogen content of fresh food organisms with material recovered from faeces. Visually, the degree of digestion of food particles ranged from no discernible change of lamellibranch larvae, copepod eggs, intact copepod faecal pellets and some phytoplankton species, to varying degrees of removal of body constituents in copepods, cladocerans and decapod zoea. For crustaceans, the proportion of body constituents removed was related to the size and construction of their apparently indigestible exoskeleton. Uppon defaecation larger organisms showed the greatest percentage loss in dry weight and carbon. A high percentage of nitrogen was extracted from all organisms.

Digestion of copepod eggs by larval turbot Scophthalmus maximus and egg viability following gut passage

Between 20.5 and 93.6 % of the subitaneous eggs of 6 species of egg-carrying copepods passed undigested through the digestive tracts of larval and early postlarval turbot Scophthalmus maximus. Viability of the eggs of Eurytemora affinis, E. velox and Euterpina acutifrons remained high on egestion (67.0 to 91.7 %), Pseudocalanus elongatus and Oncaea venusta eggs had low viability (1.1 to 1.5 %), while all Corycaeus anglicus eggs were rendered inviable. The indigestibility of the eggs denies the turbot larvae a potentially valuable food resource, while retention of high egg viability in certain species reduces the effect of predation.

Effect of Grazing-Mediated Dimethyl Sulfide (DMS) Production on the Swimming Behavior of the Copepod Calanus helgolandicus

Chemical interactions play a fundamental role in the ecology of marine foodwebs. Dimethyl sulfide (DMS) is a ubiquitous marine trace gas that acts as a bioactive compound by eliciting foraging behavior in a range of marine taxa including the copepod Temora longicornis. Production of DMS can rapidly increase following microzooplankton grazing on phytoplankton. Here, we investigated whether grazing-induced DMS elicits an increase in foraging behavior in the copepod Calanus helgolandicus. We developed a semi-automated method to quantify the effect of grazing-mediated DMS on the proportion of the time budget tethered females allocate towards slow swimming, typically associated with feeding. The pooled data showed no differences in the proportion of the 25 min time budget allocated towards slow swimming between high (23.6 +/- 9.74%) and low (29.1 +/- 18.33%) DMS treatments. However, there was a high degree of variability between behavioral responses of individual copepods. We discuss the need for more detailed species-specific studies of individual level responses of copepods to chemical signals at different spatial scales to improve our understanding of chemical interactions between copepods and their prey.

Effects Of Preservation On Wet Weight Dry-Weight Nitrogen And Carbon Contents Of Calanus-Helgolandicus (Crustacea Copepoda)

The lengths, wet and dry weights, nitrogen and carbon contents of fresh, frozen and formaldehyde-fixed specimens of Calanus helgolandicus (Claus) were determined. Samples were collected during May 1980 in the Celtic Sea. Individual Copepodite Stages 3, 4, 5, and Adult Male and Female Stage 6 were measured and analysed, and 36 linear regression equations derived for these variables together with mean values, standard deviations and 95% confidence limits. The range of nitrogen values in the fresh material, expressed as a percentage of dry weight, ranged from 8.08%±0.80 (Copepodite Stage 3) to 10.89%±0.27 (adult female); carbon values changed from 41.6%±3.05 (mean ±95% confidence limits) for Copepodite Stage 3 to 50.97%±2.63 in Copepodite Stage 5. The adult females had a high nitrogen and relatively low carbon content, while the converse was true for Stage 5 copepodites. There was a loss of dry weight from the frozen samples (57%) and the fixed samples (38%) compared with the mean of the fresh dry weight of all stages. The material lost from the copepods was rich in nitrogen, thus, artificially high percentage carbon values were determined from the frozen and fixed samples (52.0 to 60.3% and 44.7 to 58.5%, respectively).

Continuous Plankton Records - Persistence In Time-Series And The Population-Dynamics Of Pseudocalanus-Elongatus And Acartia-Clausi

Rates of population increase in early spring and the sizes of overwintering stocks were calculated for the planktonic copepods Pseudocalanus elongatus and Acartia clausi for a set of areas covering the open waters of the north-east Atlantic Ocean and the North Sea for the period 1948 to 1979. For both species, the rates of population increase were higher in the open ocean than in the North Sea and appear to be related to temperature. The overwintering stocks in the North Sea were larger than those in the open ocean and are probably related to phytoplanton concentration. P. elongatus shows higher overwintering stocks and lower rates of population increase than A. clausi, resulting in different levels of persistence in the stocks of the two species. It is suggested that this difference in persistence is responsible for differences between the two species with respect to geographical distribution in summer and different patterns of year-to-year fluctuations in abundance.

The Nematode-Copepod Ratio And Its Use In Pollution Ecology

The nematode/copepod ratio is critically examined with a view to adding some precision to its proposed use in pollution ecology. At two unpolluted intertidal sites, differing markedly in sediment grade, the metabolic requirements of copepods are shown to be equivalent to the requirements of that fraction of the nematode population which feeds in the same way. The partitioning of this total energy requirement among individuals depends on the distribution of individual metabolic body sizes and the relative rates of metabolism. The distribution of body sizes is constrained by the sediment granulometry, which affects nematodes and copepods differently. These considerations enable precise predictions of the nematode/copepod ratios expected in unpolluted situations, against which observed ratios can be compared.

North Atlantic climatic signals and the plankton of the European Continental Shelf

Climatic variability on the European Continental Shelf is dominated by events over the North Atlantic Ocean, and in particular by the North Atlantic Oscillation (NAO). The NAO is essentially a winter phenomenon, and its effects will be felt most strongly by populations for which winter conditions are critical. One example is the copepod Calanus finmarchicus, whose northern North Sea populations overwinter at depth in the North Atlantic. Its annual abundance in this region is strongly dependent on water transports at the end of the winter, and hence on the NAO index. Variations in the NAO give rise to changes in the circulation of the North Atlantic Ocean, with additional perturbations arising from El Ni (n) over tildeo - Southern Oscillation (ENSO) events in the Pacific, and these changes can be delayed by several years because of the adjustment time of the ocean circulation. One measure of the circulation is the latitude of the north wall of the Gulf Stream (GSNW index). Interannual variations in the plankton of the Shelf Seas show strong correlations with the fluctuations of the GSNW index, which are the result of Atlantic-wide atmospheric processes. These associations imply that the interannual variations are climatically induced rather than due to natural fluctuations of the marine ecosystem, and that the zooplankton populations have not been significantly affected by anthropogenic processes such as nutrient enrichment or fishing pressure. While the GSNW index represents a response to atmospheric changes over two or more years, the zooplankton populations correlated with it have generation times of a few weeks. The simplest explanation for the associations between the zooplankton and the GSNW index is that the plankton are responding to weather patterns propagating downstream from the Gulf Stream system. It seems that these meteorological processes operate in the spring. Although it has been suggested that there was a regime shift in the North Sea in the late 1980s, examination of the time-series by the cumulative sum (CUSUM) technique shows that any changes in the zooplankton of the central and northern North Sea are consistent with the background climatic variability. The abundance of total copepods increased during this period but this change does not represent a dramatic change in ecosystem processes. It is possible some change may have occurred at the end of the time-series in the years 1997 and 1998.