Fecal pellets (FP) are a key component of the biological carbon pump, as they can, under some circumstances, efficiently transfer carbon to depth. Like other forms of particulate organic carbon (POC), they can be remineralized in the ocean interior (particularly in the upper 200 m), or alternatively they can be preserved in the sediments. The controls on the attenuation of FP flux with depth are not fully understood, in particular, the relative contributions of zooplankton fragmentation and microbial/zooplankton respiration to FP loss. Collection of sinking particles using Marine Snow Catchers at three ecologically contrasting sites in the Scotia Sea, Antarctica, revealed large differences in POC flux composition (5-96% FP) and flux attenuation despite similar temperatures. To determine the importance of microbial respiration on FP loss in the upper mesopelagic, we made the first ever measurements of small scale oxygen gradients through the boundary layer at the interface of krill FP collected from the Scotia Sea. Estimated carbon-specific respiration rates of microbes within FP (0.010-0.065 d 21 ) were too low to account for the observed large decreases in FP flux over the upper 200 m. Therefore, the observed rapid declines in downward FP flux in the upper mesopelagic are more likely to be caused by zooplankton, through coprophagy, coprorhexy, and coprochaly. Microbial respiration is likely to be more important in regions of higher temperatures, and at times of the year, or in depths of the ocean, where zooplankton abundances are low and therefore grazing and fragmentation processes are reduced.
Abstract. The northern Scotia Sea contains the largest seasonal uptake of atmospheric carbon dioxide yet measured in the Southern Ocean. This study examines one of the main routes by which this carbon fluxes to the deep ocean: through the production of faecal pellets (FPs) by the zooplankton community. Deep sediment traps were deployed at two sites with contrasting ocean productivity regimes (P3, naturally iron-fertilized, and P2, iron-limited) within the same water mass. The magnitude and seasonal pattern of particulate organic carbon (POC) and FPs in the traps was markedly different between the two sites. Maximum fluxes at P3 (22.91 mg C m −2 d −1 ; 2534 FP m −2 d −1 ) were 1 order of magnitude higher than at P2 (4.01 mg C m −2 d −1 ; 915 FP m −2 d −1 ), with flux at P3 exhibiting a double seasonal peak, compared to a single flatter peak at P2. The maximum contribution of FP carbon to the total amount of POC was twice as high at P3 (91 %) compared to P2 (40 %). The dominant FP category at P3 varied between round, ovoidal, cylindrical and tabular over the course of the year, while, at P2, ovoidal FPs were consistently dominant, always making up more than 60 % of the FP assemblage. There was also a difference in the FP state between the two sites, with FPs being relatively intact at P3, while FPs were often fragmented with broken peritrophic membranes at P2. The exception was ovoidal FPs, which were relatively intact at both sites. Our observations suggest that there was a community shift from a herbivorous to an omnivorous diet from spring through to autumn at P3, while detritivores had a higher relative importance over the year at P2. Furthermore, the flux was mainly a product of the vertically migrating zooplankton community at P3, while the FP flux was more likely to be generated by deeper-dwelling zooplankton feeding on recycled material at P2. The results demonstrate that the feeding behaviour and vertical distribution of the zooplankton community plays a critical role in controlling the magnitude of carbon export to the deep ocean in this region.
Scarred shells of polar pteropod Limacina helicina collected from the Greenland Sea inJune 2012 reveal a history of damage, most likely failed predation, in earlier life stages.Evidence of shell fracture and subsequent re-growth is commonly observed in specimens recovered from the sub-Arctic and further afield. However, at one site within sea-ice on the Greenland shelf, shells that had been subject to mechanical damage were also found to exhibit considerable dissolution. It was evident that shell dissolution was localised to areas where the organic, periostracal sheet that covers the outer shell had been damaged at some earlier stage during the animal's life. Where the periostracum remained intact, the shell appeared pristine with no sign of dissolution. Specimens which appeared to be pristine following collection were incubated for four days.Scarring of shells that received periostracal damage during collection only became evident in specimens that were incubated in waters undersaturated with respect to aragonite, Ar≤1. While the waters from which the damaged specimens were collected at the Greenland Sea sea-ice margin were not Ar ≤1, the water column did exhibit the lowest Ar values observed in the Greenland and Barents Seas, and was likely to have approachedAr≤1 during the winter months. We demonstrate that L. helicina shells are only susceptible to dissolution where both the periostracum has been breached and the aragonite beneath the breach is exposed to waters of Ar≤1. Exposure of multiple layers of aragonite in areas of deep dissolution indicate that, as with many molluscs, L. helicina is able to patch up dissolution damage to the shell by secreting additional aragonite 2 internally and maintain their shell. We conclude that, unless breached, the periostracum provides an effective shield for pteropod shells against dissolution in waters Ar≤1, and when dissolution does occur the animal has an effective means of self-repair. We suggest that future studies of pteropod shell condition are undertaken on specimens from which the periostracum has not been removed in preparation.
Downward fluxes of particulate matter were investigated in the polynya of Terra Nova Bay (western Ross Sea) from February 1995 to December 1997. The main biological components were siliceous phytoplankton (diatoms, silicoflagellates and parmales), abundant faecal pellets of several types and zooplankton (mainly shelled pteropods). Vertical fluxes of particles occurred mainly through diatoms and faecal pellets in the first and second part of the summer, respectively. The highest fluxes were recurrently observed in late summer, when faeces contributed up to 100% of organic carbon. Unusually high fluxes were recorded in winter 1995, when faecal pellets accounted for 84.6% of the organic carbon. Peak fluxes were always driven by the sinking of faecal pellets, that hence appear to be the most efficient vector of export in the polynya of Terra Nova Bay. A major flux component was the pteropod Limacina helicina, which repeatedly sank in high amounts after the growing season. In April-June, L. helicina probably transported biogenic carbon to deep layers as a passive sinker. The inclusion of pteropods in flux estimates resulted in values that were up to 20 (for total mass), 25 (for organic matter) and 48 (for carbonate) times higher than the previously measured fluxes. Fluxes are known to be biased by swimmers, but ultimately attention must be paid to a possible erroneous categorization of some zooplankton as swimmers to avoid severe underestimation of fluxes of total mass (up to 95% in our study), organic matter (up to 96%) and carbonate (up to 100%).
The biological carbon pump drives a flux of particulate organic carbon (POC) through the ocean and affects atmospheric levels of carbon dioxide. Short term, episodic flux events are hard to capture with current observational techniques and may thus be underrepresented in POC flux estimates. We model the potential hidden flux of POC originating from Antarctic krill, whose swarming behaviour could result in a major conduit of carbon to depth through their rapid exploitation of phytoplankton blooms and bulk egestion of rapidly sinking faecal pellets (FPs). Our model results suggest a seasonal krill FP export flux of 0.039 GT C across the Southern Ocean marginal ice zone, corresponding to 17–61% (mean 35%) of current satellite-derived export estimates for this zone. The magnitude of our conservatively estimated flux highlights the important role of large, swarming macrozooplankton in POC export and, the need to incorporate such processes more mechanistically to improve model projections.
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