Increased production of faecal pellets by the benthic harpacticoid Paramphiascella fulvofasciata: importance of the food source

The re-use of faecal pellets in the water column before sinking to the seafloor is known as an important pathway in marine food webs. Especially planktonic copepods seems to be actively use their faecal pellets. Since benthic copepods (order Harpacticoida) live in the vicinity of their pellets, it remains unclear how important these pellets are for their feeding ecology. In the present study a laboratory experiment was conducted to analyse the importance of faecal pellets for the feeding ecology of the harpacticoid Paramphiascella fulvofasciata and its grazing pressure on two diatom species (Seminavis robusta, Navicula phyllepta). By quantifying the amount and volume of the produced faecel pellets in different treatments, it was tested to what extent the food source and the lack of faecal pellets would influence the production of faecal pellets. We found that the grazing pressure of P. fulvofasciata depended on the diatom density since only a top-down effect could be found on the smaller Navicula cells during its initial exponential growth phase. The grazer had a negative effect on the diatom growth and controlled the cell density to about 4,000 cells/cm². In spite of the fact that the addition of faecal pellets did not show a significant positive effect on the assimilation of diatoms, the removal of faecal pellets strongly promoted the pellet production. Especially when grazing on Navicula the harpacticoid P. fulvofasciata produced significantly more and smaller faecal pellets when the pellets were removed. This outcome illustrates the need for faecal pellets of this harpacticoid copepod when grazing on the diatom Navicula. Apart from its selection for smaller diatom cells, it was suggested that the colonisation of heterotrophic bacteria enriched these pellets. This study is the first to indicate that trophic upgrading occurs on faecal pellets and not only on the initial autotrophic food sources per se.     

Sinking rates of natural copepod fecal pellets

Many pure samples of natural fecal pellets have been collected from mixed small copepods and from the pontellid copepod Anomalocera patersoni in the Ligurian Sea, using a specially designed pellet collection device. Sinking rates of fresh pellets and pellets aged up to 33 days have been determined at 14°C, the mean temperature of the essentially isothermal water column in the Ligurian Sea. Sinking rates of pellets collected during calm sea states increased with increasing pellet volume, but sinking rates of pellets collected during rough sea (Beaufort scale 6) showed little correlation with pellet size. Much of the variability in the sinking rate-pellet size relationships was the result of different pellet composition and compaction, but not pellet age. Pellets produced from laboratory diets of phytoplankton and phytoplankton-sediment mixes showed the expected wide variability in sinking rates, with sediment-ballasted pellets sinking much faster than pellets produced from pure algal diets; thus determination of vertical material fluxes in the sea using laboratory-derived fecal pellet sinking rates is unwarranted. Natural pellet sinking data for small copepods and A. patersoni have been combined with similar data for euphausiids, to yield sinking rates of roughly two orders of magnitude over three orders of magnitude in pellet volume. Pellets from small copepods sank at speeds too slow to be of much consequence to rapid material flux to the deep sea, but they undoubtedly help determine upper water distribution of materials. Recalculation of fecal pellet mass flux estimates from the literature, using our sinking rate data for natural small copepod pellets, yielded estimates about half those of previously published values. Earlier studies had concluded that small fecal pellets were of lesser significance to total material flux than fecal matter; our recalculation strengthens that conclusion. Pellets from large copepods and euphausiids, however, have the capability to transport materials to great depths, and probably do not substantially recycle materials near the surface. The fact that the majority of pellets which had previously been collected in deep traps by other workers were of a size comparable to pellets from our large copepods supports the contention that these larger pellets are the main ones involved in vertical flux.     

Role of bacteria in decomposition of faecal pellets egested by the epiphyte-grazing gastropod Gibbula umbilicaris

The significance of the microheterotrophic utilization of faecal pellets derived from Gibbula umbilicaris — one of the most important gastropod species in the Posidonia oceanica ecosystem around the Isle of Ischia (Italy) — was investigated by means of microcosm-experiments. Initial total organic carbon (TOC) content of faecal pellets was 32 gC (mg faeces dry wt)^-1 and declined continuously over a 2-month incubation period. The low values of TOC coincided with SEM observations of pellets which were found to consist mainly of diatom frustules and other slowly utilizable material. In a long-term experiment freshly egested faecal pellets were rapidly colonized by bacteria, which reached densities of up to 14x10⁵ cells (mg faeces dry wt)^-1 within 12 h. Thereafter bacterial numbers declined and fluctuated in a very narrow range between 2 and 3.5x10⁵ cells (mg faeces dry wt)^-1 during the two months of the investigation period. In short-term experiments (over 5 d) similar trends were observed. Peak densities of attached bacteria were followed by a decline of this population. Concurrently the number of free-living bacteria increased. This observation was confirmed by O₂ consumption measurements of freshly egested faecal pellets over 24 h, where the highest O₂ consumption rates were obtained after 12 h, followed by a rapid decline, thus supporting the view that detachment of bacteria may occur. Moreover, the respiration data indicate that about 4% of faecal TOC are remineralized within the first day.     

Sinking rates of fecal pellets from gelatinous zooplankton (Salps,Pteropods, Doliolids)

Sinking rates were determined for fecal pellets produced by gelatinous zooplankton (salps, Salpa fusiformis and Pegea socia; pteropods, Corolla spectabilis; and doliolids, Dolioletta gegenbaurii) feeding in surface waters of the California Current. Pellets from the salps and pteropods sank at rates up to 2 700 and 1 800 m d^-1, respectively; such speeds exceed any yet recorded for zooplankton fecal pellets. Fecal pellets of salps were rich in organic material, with C:N ratios from 5.4 to 6.2, close to values for living plankton. The relation between volume and sinking rate indicates that salp and pteropod pellets are slightly less dense than those of pelagic Crustacea; moreover, pellet density varied between different collection dates, probably because of differences in composition. In contrast, doliolid pellets sank at rates up to 208 m d^-1, a rate much lower than would be expected from pellet size. Thus, density and sinking rates of pellets are much more variable in zooplankton than would be expected from studies of crustaceans alone. Moreover, the extraordinarily high sinking rates of fecal pellets of salps indicates that these tunicates may be disproportionately important in the flux of biogenic materials during periods when they form dense population blooms.     

Production of DMSP and DMS during a mesocosm study of an Emiliania huxleyi bloom: influence of bacteria and Calanus finmarchicus grazing

We investigated the influence of bacteria and metazooplankton on the production of dimethylsulfoniopropionate (DMSP) and dimethylsulfide (DMS) during blooms of Emiliania huxleyi (Lohmann) Hay and Mohler in seawater mesocosms. The phytoplankton succession was marked by the rapid collapse of an initial Skeletonema costatum (Greville) Cleve bloom followed by a small E. huxleyi bloom. The collapse of the diatom bloom was accompanied by an increase in concentrations of dissolved DMSP (DMSP_d) and bacterial abundance and activity (as determined by the thymidine incorporation technique). The increase in bacterial activity was followed by a rapid decrease in DMSP_d concentrations which remained low for the rest of the experiment, even during the subsequent collapse of the E. huxleyi blooms. The absence of DMSP_d and DMS peaks during the declining phase of the E. huxleyi blooms was attributed to the high bacterial activity prevailing at that time. The influence of metazooplankton grazing on DMSP and DMS production was investigated by adding moderate (24 mg dry weight m^-3) and high (520 mg dry weight m^-3) concentrations of Copepodite Stage V and adults of Calanus finmarchicus to two of four filtered (200 m mesh net) enclosures during the E. huxleyi blooms. The addition of C. finmarchicus, even in high concentrations, had no apparent effect on the dynamics of E. huxleyi, suggesting that the copepods were not grazing significantly on nanophytoplankton. The addition of copepods in high concentrations favored an accumulation of chlorophyll a and particulate DMSP. These results suggest that copepods were preying on the herbivorous microzooplankton which, in turn, was controlling the biomass of nanophytoplankton. DMS production was also enhanced in the enclosure with maximum metazooplankton biomass, suggesting that the grazing of C. finmarchicus on microzooplankton containing DMSP may contribute to DMS production. These results provide strong support to the emerging idea that bacteria and metazooplankton grazing play a dominant role in determining the timing and magnitude of DMS pulses following phytoplankton blooms.     

Characteristic features of body composition and metabolism in some interzonal copepods

Protein, lipid, phosphorus, and organic carbon contents, as well as electron transport system (ETS) activity, lactatedehydrogenase activity, and gut evacuation rate, were measured in four interzonal species of Pacific copepods:Calanus australis, C. pacificus, Eucalanus inermis, andE. elongatus f.hyalinus, collected at the upwelling areas off Peru (8°S) and California (27°N), and in the middle of the North Pacific (30°N), from February to April 1987. The two Eucalanidae species —E. inermis andE. elongatus — have distinctive biochemical and elemental body composition and rates of main physiological processes. Relative protein, lipid, phosphorus, and organic carbon contents (µg mg^–1 wet weight) in these species were, respectively, ca. 1/7 to 1/10, 1/5 to 1/20, 1/5 to 1/10, and 1/5 those inCalanus spp. Likewise, oxygen uptake rate per unit of wet weight (based on ETS activity) inE. inermis andE. elongatus was 5 to 10% of that in calanids; a similar difference was found in phosphorus excretion rate. In addition, gut evacuation rates inE. inermis andE. elongatus were ca. one-fifth of those inCalanus spp. Based on these data, we considered the eucalanids as belonging to a distinctive physiological group, figuratively named jelly-body copepods. In contrast with calanids, active lactatedehydrogenase has been found in the bodies ofE. inermis andE. elongatus, apparently allowing them to survive for a long time in layers of extremely low oxygen content (<0.2 ml l^–1). The adaptive value of physiological features in these eucalanids and typical calanids is compared.     

Cyclical contributions of the digestive epithelium to faecal pellet formation by the copepod Calanus helgolandicus

Faecal pellet formation within the gut of Stage V and adult females of the copepod Calanus helgolandicus Claus involves (1) cyclical processes of digestion and (2) the contribution of parts of the gut epithelium to the pellets. During an experimental regime in which dim lighting was restricted to day-time and feeding to night-time (17.00 to 09.00 hrs), the copepods responded with cyclical changes in both the quantity of pellets they produced and the fine structure of the contents. During the feeding period, the contents showed changes in the relative amounts of materials originating from disintegrated cells of the digestive epithelium and those derived directly from the ingested food. The vacuolar B-cells of the gut contribute to the content of the pellets and the distal, necrotic N-cells appear to be involved in forming the peritrophic membrane which encloses each pellet. Cells of the gut epithelium which are broken down during feeding are all replaced during the non-feeding period. Other individuals were taken directly from the sea and in these, also, the cells of the gut broke down during feeding and contributed to the faecal pellets. The supply of epithelial cells may limit the duration of the feeding period.     

Laboratory and field estimates of the rate of faecal pellet production by Antarctic krill,Euphausia superba

The time course of faecal pellet production (egestion) was monitored in January 1985 for a population of Antarctic krill, Euphausia superba Dana, maintained in flowing seawater aquaria at Palmer Station, Antarctica. Following transfer to filtered seawater, krill produced faecal strings for roughly 40 min, after which time faecal egestion virtually ceased. Similar results were obtained for freshly-trawled krill at sea in February and March 1985. There were wide daily variations in total faecal egestion rate; mean rates varied from 0.54 to 1.66 mg dry wt h^-1 and individual rates from 0.25 to 2.35 mg h^-1 (all data corrected to a standard krill of 600 mg fresh weight). Despite these wide fluctuations in total faecal egestion, the loss of organic matter showed no significant daily variation, with a mean value of 0.13 mgh^-1. The relationship between faecal egestion rate and faecal organic content suggested that feeding rate was governed by food quality; when inorganic load was high, feeding rate increased to ensure sufficient energy intake. The data suggest that superfluous feeding does not occur in krill and that values of gut-clearance time calculated from time intervals greater than about 40 min will not be representative of previous feeding history. the rates of faecal egestion observed in this study indicate that the flux of faecal pellets from krill is substantial. They imply an energy intake in E. superba of 17 to 28% body weight per day, much higher than estimated previously for this species by summing known energy losses, but similar to estimates for other euphausiids.     

Seasonal variations in the densities of fecal pellets produced by Oikopleura vanhoeffeni (C. Larvacea) and Calanus finmarchicus (C. Copepoda)

Two abundant macrozooplankters, Oikopleura vanhoeffeni (Lohmann) and Calanus finmarchicus (Gunnerus) were collected from the coastal waters off Newfoundland in different seasons during 1990–1991 and incubated in natural seawater to collect freshly egested, field produced, fecal pellets. The densities of fecal pellets from O. vanhoeffeni and C. finmarchicus were measured in an isosmotic density gradient. These are the first reported seasonal measurements of fecal pellet densities from two different types of macrozooplankters, O. vanhoeffeni, a larvacean, filter feeder and C. finmarchicus, a crustacean, suspension feeder. Pellet density ranges and medians were significantly different among seasons for both species, depending primarily on the type of phytoplankton ingested and its ability to be compacted. Winter O. vanhoeffeni and fall C. finmarchicus feces filled with nanoplankters and soft bodied organisms had less open space [packing index (% open area) = 3.5 and 4% for O. vanhoeffeni and C. finmarchicus, respectively] and were more dense (1.23 and 1.19 g cm^-3) than spring feces filled with diatoms (packing index = 15 and 23%, density = 1.13 and 1.11 gcm^-3). For copepods, these results contrast with previously published density values and with the predicted copepod fecal pellet density calculated, in the present study, using the conventional mass/volume relationship. Copepod spring and summer diatom-filled feces had a calculated density of 1.12 and 1.24 gcm^-3 vs a measured median density of 1.11 gcm^-3 for both spring and summer feces; the fall feces containing nanoplankters had a calculated density of 1.08 gcm^-3 vs a measured median density of 1.19 gcm^-3. Knowledge of the seasonal variations in fecal pellet densities is important for the development of flux models.     

Gut pigment accumulation and destruction by arctic copepods in vitro and in situ

The results presented here were obtained at six locations during three cruises in 1985 (off the coast of Labrador), 1986 (at the eastern end of Viscount Melbourne Sound) and 1988 (off the coast of Labrador). In situ chlorophyll maximum concentrations were >7 gl^-1 at depths of between 0 and 30 m in all sampling areas. In feeding experiments copepods attained higher gut pigment concentrations the longer they had been previously starved and higher concentrations when fed in the dark than when fed in the light. Community ingestion rates calculated from changes in particulate chlorophyll were higher than estimates derived from gut pigment data except when copepods had been starved for 24 h. Differences between estimates by the two methods suggested pigment destruction. In feeding experiments pigment: biogenic silica ratios in food and faecal pellets suggested that the length of starvation period affected the degree of pigment destruction differently at different stations and that feeding in the light greatly increased pigment destruction. A comparison of pigment: silica ratios in the water column, and in faecal pellets collected from copepods which had fed there, suggested that pigment destruction may occur in situ sometimes and that the degree to which it occurs may be affected by feeding history, light, diel feeding behaviour and species composition.     

Comparison of the chemical composition of particulate material and copepod faecal pellets at stations off the coast of Labrador and in the Gulf of St. Lawrence

Faecal pellets were collected in 1988 from copepods which had fed in situ or in laboratory experiments, using screened natural seawater as food, at two stations off the coast of Labrador and one in the Gulf of St. Lawrence. The chemical composition of the pellets and of particulate material in profiles and in laboratory food were measured in terms of particulate carbon, carbohydrate (soluble and insoluble), protein and lipid. Faecal pellet composition was somewhat similar in all experiments at the first two stations, where the compositions of particulate material in situ and copepod species assemblages were also similar. At the third station the compositions of faecal pellets and particulate material were slightly different from those at the other stations and the copepod species composition varied between sampling times. Faecal pellets at the first two stations had very low levels of soluble carbohydrate, while concentrations in the food were generally high, suggesting that it was efficiently metabolized by copepods, although it might have been absent because of sloppy feeding or release, after passage through the gut, in soluble form or from faecal pellets. Comparisons of POC: biogenic silica ratios in food and faecal pellets, calculated using data presented elsewhere (Head 1992; Mar. Biol. 112: 583–592), suggested that at these stations, where food concentrations were high (chlorophyll concentrations>8 gl^-1), copepods may have been assimilating carbon rather inefficiently.     

Functional responses of copepod nauplii using a high efficiency gut fluorescence technique

To investigate copepod nauplii ingestion rates on phytoplankton, we have adapted the traditional gut fluorescence technique as it can be used with lower gut pigment concentrations. With the improved technique, laboratory experiments were performed to estimate functional responses for nauplii of Calanus helgolandicus and Centropages typicus. Nauplii were raised from eggs to copepodites and the experiments were performed with stages NIV-NV. Gut evacuation rates and ingestion rates were measured on Isochrysis galbana at different concentrations. Specific ingestion rates ranged between 0.038–0.244 μg C μg⁻¹ nauplii C d⁻¹ for C. typicus and 0.041–1.412 μg C μg⁻¹ nauplii C d⁻¹ for C. helgolandicus. Both species showed a type III functional response, reaching a saturation concentration at around 600 μgC l⁻¹ for C. typicus and 800 μgC l⁻¹ for C. helgolandicus. An erratum to this article can be found at     

Morphology and sinking velocities of fecal pellets of copepod, molluscan, euphausiid, and salp taxa in the northeastern tropical Atlantic

Morphological characteristics and sinking velocities of naturally occurring fecal pellets of copepods, euphausiids, salps, and pelagic mollusks collected in the northeastern tropical Atlantic were investigated during the period of May-June 1992. The fecal pellets of copepods and euphausiids were cylindrical and distinguished only by their size. Those of salps were, in general, rectangular, and slight differences were noted according to the species. The fecal pellets of the molluscan pteropod Clio sp. were conical, while those of the molluscan heteropod Carinaria sp. were spiral. The sinking velocities ranged from 26.5 to 159.5 m day^-1 for copepod fecal pellets, from 16.1 to 341.1 m day^-1 for euphausiid pellets, from 43.5 to 1167.6 m day^-1 for salps' pellets (Cyclosalpa affinis, Salpa fusiformis, Iasis zonaria, and two unidentified species), from 65 to 205.7 m day^-1 for Clio sp. pellets, and from 120.3 to 646.4 m day^-1 for Carinaria sp. fecal pellets. The measured sinking velocities were compared with estimates predicted using the equations of Komar et al. (1981; Limnol Oceanogr 26:172-180), Stokes' law, and Newton's second law, using either a constant density of fecal pellets (1.22 g cm^-3) or densities estimated with the three different equations. When a constant density was used, the three equations overestimated the sinking velocities; Stokes' law resulted in the largest overestimation, and Newton's second law, the smallest. At the taxa level, the overestimation was greatest for euphausiid 1 fecal pellets and smallest for copepod fecal pellets. When the three equations were used to estimate fecal pellet density, the density estimated using the equation of Komar et al. was the greatest, and that using Stokes' law, the smallest, resulting in over- and underestimation of sinking velocities, respectively. Newton's second law resulted in an intermediate density and gave the closest estimate of sinking velocities. We propose that measurement of sinking velocities of a portion of the fecal pellets might guide in choosing an appropriate equation to be used for a reasonable interpretation of vertical mass flux.     

Role of salps in the flux of organic matter to the bottom of the Ligurian Sea

Sediment-trap samples were collected during and after the occurrence of a salp (Salpa fusiformis) bloom in the Bay of Villefranche, Mediterranean Sea, in April/May 1985. Large amounts of organic aggregates and faecal pellets were collected during the bloom. The aggregates were rich in carbohydrates and mineral grains and had similar rates of sedimentation (900 to 2 100 m d^-1) to those of the faecal pellets (1 000 to 2 000 m d^-1). The results of mineralogical and organic chemical analyses indicate the potential effect of these mucus-rich aggregates on local biogeochemical fluxes.     

Role of sinking in diatom life-history cycles: ecological,evolutionary and geological significance

Rapid mass sinking of cells following diatom blooms, observed in lakes and the sea, is argued here to represent the transition from a growing to a resting stage in the life histories of these algae. Mass sinking is of survival value in those bloom diatoms that retain viability over long periods in cold, dark water but not in warm, nutrient-depleted surface water. Mechanisms for accelerating sinking speed of populations entering a resting or seeding mode are proposed. Previously unexplained features of diatom form and behaviour take on a new meaning in this context of diatom seeding strategies. Diatoms have physiological control over buoyancy as declining growth is accompanied by increasing sinking rates, where the frustule acts as ballast. Increased mucous secretion in conjunction with the cell protuberances characteristic of bloom diatoms leads to entanglement and aggregate formation during sinking; the sticky aggregates scavenge mineral and other particles during descent which further accelerates the sinking rate. Such diatom flocs will have sinking rates of 100 m d^-1 or more. This is corroborated by recent observations of mass phytoplankton sedimentation to the deep sea. This mechanism would explain the origin of marine snow flocs containing diatoms in high productivity areas and also the well-known presence of a viable deep sea flora. That mortality is high in such a seeding strategy is not surprising. A number of species-specific variables pertaining to size, morphology, physiology, spore formation and frustule dissolution rate will determine the sinking behaviour and thus control positioning of resting stages in the water column or on the bottom. It is argued that sinking behaviour patterns will be environmentally selected and that some baffling aspects of diatom form and distribution can be explained in this light. Rapid diatom sedimentation is currently believed to be mediated by zooplankton faecal pellets, particularly those of copepods. This view is not supported by recently published observations. I speculate that copepod grazing actually retards rather than accelerates vertical flux, because faecal pellets tend to be recycled within the surface layer by the common herbivorous copepods. Egestion of undigested food by copepods during blooms acts as a storage mechanism, as ungrazed cells are likely to initiate mass precipitation and depletion of the surface layer in essential elements. Unique features of diatoms are discussed in the light of their possible evolution from resting spores of other algae. An evolutionary ecology of pelagic bloom diatoms is deduced from behavioural and morphological characteristics of meroplanktonic and tychopelagic forms. Other shell-bearing protistan plankters share common features with diatoms. Similar life-history patterns are likely to be present in species from all these groups. The geological significance of mass diatom sinking in rapidly affecting transfer of biogenic and mineral particles to the sea floor is pointed out.     

Sinking rates of fecal pellets from the marine copepod Pontella meadii

Sinking rates of fecal pellets produced by the marine copepod Pontella meadii, grazing on 4 different phytoplankton diets, ranged from 15 to 153 m/day, with a mean of 66 m/day. Sinking rates, in general, were directly related to fecal pellet volumes, but unrelated to the diets used to produce the fecal pellets. There were two-to-threefold variations in sinking rates between fecal pellets of the same volumes, often produced on the same diets. Twenty repetitions of timed sinking of a single fecal pellet revealed sinking rates varying from 33 to 79 m/day, as well as variations in sinking rates within the course of individual descents. It is suggested that copepod fecal pellets are of such small volumes and densities that their sinking rates are subject to microstructural variations in the most carefully controlled water columns. Scanning electron microscope observations revealed lack of structural damage to some of the diatom frustules in the fecal pellets, suggesting that superfluous feeding may have occurred. Thus, the accelerated sinking rates of copepod fecal pellets may provide a mechanism for nutritional enrichment of the deep-sea ecosystem with organic parcels containing incompletely-assimilated plant material.     

Accumulation of 14C-1-naphthalene by an oceanic and an estuarine copepod during long-term exposure to low-level concentrations

Accumulation of the bi-cyclic aromatic hydrocarbon ¹⁴C-1-naphthalene in adult female Calanus helgolandicus Claus and adult female Eurytemora affinis Poppe in sea water concentrations of hydrocarbon ranging from 0.2 to 992 g/l was studied during exposure periods of up to 15 days as part of an investigation of the possible effects on marine zooplankton of persistent exposure to low levels of petroleum hydrocarbons. With both species the body levels of radioactivity increased rapidly during the first few days of the exposure period, but after exposure for 7 to 8 days to sea water containing 50 g hydrocarbon/l an equilibrium condition was approached; in some experiments where E. affinis was exposed to 1.0 and 10 g hydrocarbon/l for 15 days there was no further increase in body levels of radioactivity after 7 to 8 days. Using a low concentration of hydrocarbon (1 g/l), the quantity of radioactivity accumulated after 10 days was found to be nearly fifty times greater in the smaller species, E. affinis, than in C. helgolandicus, when expressed in terms of body weight. After they had been exposed to the hydrocarbon for several days the copepods contained a considerable proportion of radioactivity that was no longer identifiable as naphthalene and was presumably present as metabolites. Radioactivity accumulated in the copepods after several days was rapidly lost after they were transferred to uncontaminated sea water: e.g. C. helgolandicus lost nearly 90% of its body level of radioactivity in 24 h. Thereafter the rate of loss was greatly reduced, and 5% of the original body level of radioactivity still remained in the copepods at the end of 11 days. Experiments on the breakdown of naphthalene added at low concentrations to sea water samples containing natural microbial populations indicated degradation rates of 0.1 to 0.2 g/l/24 h in oceanic water, and 2.6 g/l/24 h in inshore water samples. The results are discussed in terms of the possible transfer of hydrocarbon to a higher trophic level in areas subjected to constant low-level inputs of petroleum hydrocarbons.     

Copepod grazing during a declining spring phytoplankton bloom in the Northern North Sea

Grazing rates of larger (Calanus finmarchicus) and smaller (Acartia clausii Pseudocalanus elongatus etc.) copepods on naturally occurring phytoplankton populations were measured during a declining spring phytoplankton bloom. During the initial period, dominated by Chaetoceros spp. diatoms, constant ingestion rates were observed in Calanus finmarchicus at suspended particulate concentrations above 300 g carbon l^-1. Average daily intake during this time amounted to 35 to 40% of body carbon and reached a maximum of 50%. The feeding response of the smaller copepods was not so well defined, although a maximum daily intake of 56% body carbon was recorded. In both groups, feeding thresholds were at particulate concentrations around 50 g C l^-1. The feeding response of C. finmarchicus was correlated with both a change in their own population and in the food cell type. Linear regressions describing the concentration-dependent feeding response were: ingestion rate (IR)=1.16 total particulate volume (TPV)-36.15 during the initial part of the period compared with IR=0.41 TPV-12.18 for the latter period. C. finmarchicus filtered out slightly larger (x 1.2 diameter) particles than the small copepods and, in both groups, some filtering adjustment was made to accomodate to modal changes in the phytoplankton population from 20–30 m to 10 m diameter cells. Particle production during feeding was frequently evident in the smallest size ranges of particles and the ratio of particle production to ingestion rate was greater at low feeding rates.     

Production and fate of copepod fecal pellets across the Southern Indian Ocean

The vertical distribution of copepods, fecal pellets and the fecal pellet production of copepods were measured at seven stations across the Southern Indian Ocean from productive areas off South Africa to oligotrophic waters off Northern Australia during October/November 2006. We quantified export of copepod fecal pellet from surface waters and how much was retained. Furthermore, the potential impact of Oncaea spp. and harpacticoid copepods on fecal pellets degradation was evaluated and found to be regional substantial. The highest copepod abundance and fecal pellet production was found in the western nutrient-rich stations close to South Africa and the lowest at the central oligotrophic stations. The in situ copepod fecal pellet production varied between 1 and 1,000 μg C m⁻³ day⁻¹. At all stations, the retention of fecal pellets in the upper 400 m of the water column was more than 99% and the vertical export of fecal pellets was low (<0.02 mg m⁻² day⁻¹).     

Vertical distribution of Calanus finmarchicus and C. helgolandicus in relation to the development of the seasonal thermocline in the Celtic Sea

The geographical distribution and annual mean abundance of Calanus finmarchicus (Gunnerus) and C. helgolandicus (Claus) in the northern North Atlantic Ocean were shown in relation to the seasonal and annual fluctuations of abundance of the species in the Celtic Sea from 1960 to 1981. These congeneric copepods, although showing allopatric distributions over most of their geographical range, have sympatric distributions in the Celtic Sea where they dominate the dry weight biomass of the plankton throughout the year. The two species respond differently to the development of the seasonal thermocline and halocline by taking up different vertical distributions in the water column. C. finmarchicus occurred in the colder, more saline water below the thermocline, while C. helgolandicus occurred in the warmer, less saline water above the thermocline. This behaviour is postulated as a mechanism by which these morphologically similar copepods more fully exploit the resources of their temporally and spatially heterogeneous environment and also minimise interspecific competition. The species have the same foraging techniques and are able to exploit the same size spectrum of particulates. The vertical depth strata in which the populations are found for most of the year in the Celtic Sea means that both species exploit the diatom bloom in early spring but, thereafter, C. helgolandicus grazes on the daily production of the autotrophs in the euphotic zone while C. finmarchicus, below the thermocline, has to rely for its food on sedimenting particulates (whole cells, detritus and faecal material). The isolating mechanisms whereby these two populations partition the habitat in the Celtic Sea are discussed.