Calanoid copepod abundances in the northern Benguela Current System from 2009 and 2010

Abundance data of copepods were derived from vertical Multinet hauls at 10 stations, carried out in the northern Benguela upwelling system in December 2009 (FRS Africana) and September/October 2010 (RRS Discovery). Three transects along ~ 17°S, 19°S and 23°S with three stations each (neritic, shelf break, oceanic) and one station at 21°S were analysed for copepod abundance. Maximum sampling depth was either close to the seafloor (neritic and shelf break stations) or 700 m (2009) and 1000 m (2010) for the oceanic stations. Calanoid copepod species and stages were identified and enumerated separately. Adult females, males and copepodite stage 5 (C5) (in case of C. carinatus and N. minor) were included in the abundance calculations. Abundance is expressed as number of individuals per m**3, calculated from the volume of water filtered (calibrated flowmeter, Hydro-Bios) and the maximum sampling depth at each station.

Respiration rates and electron transport system activities of copepod species obtained during Maria S. Merian cruise MSM17/3

Respiration rates and electron transport system (ETS) activities were measured in dominant copepod species from the northern Benguela upwelling system in January-February 2011 to assess the accuracy of the ETS assay in predicting in vivo respiration rates. Individual respiration rates varied from 0.06 to 1.60 µL O2/h/ind, while ETS activities converted to oxygen consumption ranged from 0.14 to 4.46 µL O2/h/ind. ETS activities were significantly correlated with respiration rates (r**2 = 0.79, p = 0.0001). R:ETS ratios were lowest in slow-moving Eucalanidae (0.11) and highest in diapausing Calanoides carinatus copepodids CV (0.76) while fast-moving copepods showed intermediate R:ETS (0.23-0.37). 82% of the variance of respiration rates could be explained by differences in dry mass, temperature and the activity level of different copepod species. Three regression equations were derived to calculate respiration rates for diapausing, slow- and fast-moving copepods, respectively, based on parameters such as body mass and temperature. Thus, knowledge about the activity level and behavioral characteristics of copepod species can significantly increase the predictive accuracy of metabolic models, which will help to better understand and quantify the impact of copepods on nutrient and carbon fluxes in marine ecosystems.

Density and biomass of two copepod size fractions and various species obtained during Almirante Irizar cruises to the Drake Passage and Scotia Sea (2000-2003)

The relative importance of small forms of copepods has been historically underestimated by the traditional use of 200-300-µm mesh nets. This work quantified the distribution and abundance of copepods, considering two size fractions (<300 µm and >300 µm), in superficial waters (9 m deep) of the Drake Passage and contributed to the knowledge of their interannual fluctuations among three summers. Four types of nauplii and eleven species of copepods at copepodite and adult stages were identified, with abundance values of up to 13 ind/L and 28,300 µg C/m**3. The <300-µm fraction, composed of Oithona similis, small cyclopoids and nauplii, dominated the copepod communities in the 3 years; it accounted for more than 77% of the total number and for between 40 and 63% of the total biomass. Changes in density and biomass values among the three cruises differed according to copepod size fraction and water mass; the >300-µm fraction showed no changes among the 3 years, both in Antarctic (density and biomass) and in Subantarctic waters (density), whereas the <300-µm fraction showed higher (density and biomass) values in 2001 both in Subantarctic and in Antarctic waters. Sea surface temperature and its anomaly accounted for the largest proportion of variability in copepod density and biomass, particularly for the <300-µm fraction.

Production, oxygen uptake, and sinking velocity of copepod fecal pellets originated from the central North Sea

Production, oxygen uptake, and sinking velocity of copepod fecal pellets egested by Temora longicornis were measured using a nanoflagellate (Rhodomonas sp.), a diatom (Thalassiosira weissflogii), or a coccolithophorid (Emiliania huxleyi) as food sources. Fecal pellet production varied between 0.8 pellets ind**-1 h**-1 and 3.8 pellets ind**-1 h**-1 and was significantly higher with T. weissflogii than with the other food sources. Average pellet size varied between 2.2 x 10**5 µm**3 and 10.0 x 10**5 µm**3. Using an oxygen microsensor, small-scale oxygen fluxes and microbial respiration rates were measured directly with a spatial resolution of 2 µm at the interface of copepod fecal pellets and the surrounding water. Averaged volume-specific respiration rates were 4.12 fmol O2 µm**-3 d**-1, 2.86 fmol O2 µm**-3 d**-1, and 0.73 fmol O2 µm**-3 d**-1 in pellets produced on Rhodomonas sp., T. weissflogii, and E. huxleyi, respectively. The average carbon-specific respiration rate was 0.15 d**-1 independent on diet (range: 0.08-0.21 d**-1). Because of ballasting of opal and calcite, sinking velocities were significantly higher for pellets produced on T. weissflogii (322 +- 169 m d**-1) and E. huxleyi (200 +- 93 m d**-1) than on Rhodomonas sp. (35 +- 29 m d**-1). Preservation of carbon was estimated to be approximately 10-fold higher in fecal pellets produced when T. longicornis was fed E. huxleyi or T. weissflogii rather than Rhodomonas sp. Our study directly demonstrates that ballast increases the sinking rate of freshly produced copepod fecal pellets but does not protect them from decomposition.

Copepod abundance and fecal pellet production, sinking rate, and degradation in samples obtained during Leg 7 of Galathea 3 cruise 2006

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. 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).

Respiration and ingestion rates of calanoid copepod in the northern Benguela Current System measured ex situ during the RSS Discovery D356 cruise in 2010

Respiration rates of 16 calanoid copepod species from the northern Benguela upwelling system were measured on board RRS Discovery in September/October 2010 to determine their energy requirements and assess their significance in the carbon cycle. Copepod species were sampled by different net types. Immediately after the hauls, samples were sorted to species and stages (16 species; females, males and C5 copepodids) according to Bradford-Grieve et al. (1999). Specimens were kept in temperature-controlled refrigerators for at least 12 h before they were used in experiments. Respiration rates of different copepod species were measured onboard by optode respirometry (for details see Köster et al., 2008) with a 10-channel optode respirometer (PreSens Precision Sensing Oxy-10 Mini, Regensburg, Germany) under simulated in situ conditions in temperature-controlled refrigerators. Experiments were run in gas-tight glass bottles (12-13 ml). For each set of experiments, two controls without animals were measured under exactly the same conditions to compensate for potential bias. The number of animals per bottle depended on the copepods size, stage and metabolic activity. Animals were not fed during the experiments but they showed natural species-specific movements. Immediately after the experiments, all specimens were deep-frozen at - 80 °C for later dry mass determination (after lyophilisation for 48 h) in the home lab. The carbon content (% of dry mass) of each species was measured by mass-spectrometry in association with stable isotope analysis and body dry mass was converted to units of carbon. For species without available carbon data, the mean value of all copepod species (44% dry mass) was applied. For the estimation of carbon requirements of copepod species, individual oxygen consumption rates were converted to carbon units, assuming that the expiration of 1 ml oxygen mobilises 0.44 mg of organic carbon by using a respiratory quotient (RQ) of 0.82 for a mixed diet consisting of proteins (RQ = 0.8-1.0), lipids (RQ = 0.7) and carbohydrates (RQ = 1.0) (Auel and Werner, 2003). The carbon ingestion rates were calculated using the energy budget and the potential maximum ingestion rate approach. To allow for physiological comparisons of respiration rates of deep- and shallow-living copepod species without the effects of ambient temperature and different individual body mass, individual respiration rates were temperature- (15°C, Q10=2) and size-adjusted. The scaling coefficient of 0.76 (R2=0.556) is used for the standardisation of body dry mass to 0.3 mg (mean dry mass of all analysed copepods), applying the allometric equation R= (R15°C/M0.76)×0.30.76, where R is respiration and M is individual dry mass in mg.

Fecal pellet and egg production rates of copepod species at varying suspended sediment concentrations

We investigated the effect of suspended sediments on the vital rates of the copepods Calanus finmarchicus, Pseudocalanus sp. and Metridia longa in a Greenland sub-Arctic fjord. The fjord had a gradient of suspended particulate matter (SPM) with high concentrations (>50 mg/L) in the inner fjord due to glacial melt water runoff. Laboratory experiments showed that when feeding on the diatom Thalassiosira weissflogii specific ingestion rates were low at high concentrations of suspended sediment for C. finmarchicus (>20 mg/L) and Pseudocalanus sp. (>50 mg/L), while no effect was found for M. longa. For C. finmarchicus, a relatively constant fecal pellet production (FPP) and fecal pellet volume suggested ingestion of sediment, which probably led to reduction in egg production rates (EPRs) at high sediment concentrations. For Pseudocalanus sp., FPP decreased with increasing sediment concentrations, while no effect was observed on EPR. No significant difference was observed in FPP for M. longa feeding on the diatom T. weissflogii compared to the ciliate Strombidium sulcatum. The study shows that high sediment concentrations influence the capability of carbon turnover in C. finmarchicus and Pseudocalanus sp., while M. longa appears to be more tolerant to high sediment loads. Therefore, high concentrations of SPM could potentially influence the species composition of glacially influenced fjords.