Continuous moulting by Antarctic krill drives major pulses of carbon export in the north Scotia Sea, Southern Ocean.

Antarctic krill play an important role in biogeochemical cycles and can potentially generate high-particulate organic carbon (POC) fluxes to the deep ocean. They also have an unusual trait of moulting continuously throughout their life-cycle. We determine the krill seasonal contribution to POC flux in terms of faecal pellets (FP), exuviae and carcasses from sediment trap samples collected in the Southern Ocean. We found that krill moulting generated an exuviae flux of similar order to that of FP, together accounting for 87% of an annual POC flux (22.8 g m-2 y-1). Using an inverse modelling approach, we determined the krill population size necessary to generate this flux peaked at 261 g m-2. This study shows the important role of krill exuviae as a vector for POC flux. Since krill moulting cycle depends on temperature, our results highlight the sensitivity of POC flux to rapid regional environmental change.

Author Correction: The importance of Antarctic krill in biogeochemical cycles.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

The importance of Antarctic krill in biogeochemical cycles.

Antarctic krill (Euphausia superba) are swarming, oceanic crustaceans, up to two inches long, and best known as prey for whales and penguins - but they have another important role. With their large size, high biomass and daily vertical migrations they transport and transform essential nutrients, stimulate primary productivity and influence the carbon sink. Antarctic krill are also fished by the Southern Ocean's largest fishery. Yet how krill fishing impacts nutrient fertilisation and the carbon sink in the Southern Ocean is poorly understood. Our synthesis shows fishery management should consider the influential biogeochemical role of both adult and larval Antarctic krill.

Krill faecal pellets drive hidden pulses of particulate organic carbon in the marginal ice zone.

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.

Threatened species drive the strength of the carbonate pump in the northern Scotia Sea.

The efficiency of deep-ocean CO2 sequestration is regulated by the relative balance between inorganic and organic carbon export respectively acting through the biological carbon pump (BCP) and the carbonate counter pump (CCP). The composition and abundance of calcifying species in the prevailing oceanic plankton community plays a major role in driving the CCP. Here we assess the role of these calcifying organisms in regulating the strength of the CCP in a Southern Ocean region (northern Scotia Sea) known to be a major hotspot for the drawdown of atmospheric CO2. We show that, when shelled pteropods dominate the calcifying community, the total annual reduction of CO2 transferred to the deep ocean doubles (17%) compared to when other plankton calcifiers dominate (3-9%). Furthermore, predation enhances their contribution through the removal of organic soft tissue. Pteropods are threatened in polar regions by ocean warming and acidification. We determine that their potential decline would have major implications to the comparative strengths of the BCP and CCP.

The potential role of Antarctic krill faecal pellets in efficient carbon export at the marginal ice zone of the South Orkney Islands in spring.

Antarctic krill (Euphausia superba) play a central role in the food web of the Southern Ocean, forming a link between primary production and large predators. Krill produce large, faecal pellets (FP) which can form a large component of mesopelagic particulate organic carbon (POC) fluxes. However, the patchy distribution of krill swarms, highly variable pellet composition, and variable sinking and attenuation rates means that these episodic, but potentially large, carbon fluxes are difficult to sample or model. We measured particle flux and type using Marine Snow Catchers (MSC) in the marginal ice zone near the South Orkneys, Antarctica. Krill FP were the dominant component of the POC flux in the upper 200 m (typically 60-85%). FP sinking velocities measured onboard were highly variable (15-507 m d-1) but overall high, with mean equivalent velocities of 172, 267, and 161 m d-1 at our three stations. The high numbers of krill FP sinking through the mesopelagic suggest that krill FP can be transferred efficiently and/or that rates of krill FP production are high. We compared our direct MSC-derived estimates of krill FP POC flux (33-154 mg C m-2 d-1) and attenuation to estimates of krill FP production based on previous measurements of krill density and literature FP egestion rates, and estimated net krill FP attenuation rates in the upper mesopelagic. Calculated attenuation rates are sensitive to krill densities in the overlying water column but suggest that krill FP could be transferred efficiently through the upper mesopelagic, and, in agreement with our MSC attenuation estimates, could make large contributions to bathypelagic POC fluxes. Our study contrasts with some others which suggest rapid FP attenuation, highlighting the need for further work to constrain attenuation rates and assess how important the contribution of Antarctic krill FP could be to the Southern Ocean biological carbon pump.

Trophodynamics of Protomyctophum (Myctophidae) in the Scotia Sea (Southern Ocean).

This study investigated spatial and temporal patterns in distribution, population structure and diet of Bolin's lanternfish Protomyctophum bolini, Tenison's lanternfish Protomyctophum tenisoni and gaptooth lanternfish Protomyctophum choriodon in the Scotia Sea using data collected by midwater trawl during spring, summer and autumn. Protomyctophum bolini was the most abundant species of the genus encountered throughout the Scotia Sea with the greatest concentrations occurring around the Antarctic Polar Front (APF). This species had a life cycle of 2+ years, but spatial differences in population structure were apparent as the I-group was absent from all regions south of the APF, suggesting that the species does not recruit in the Scotia Sea. Protomyctophum tenisoni occurred mostly in waters characteristic of the APF and was absent from the southern Scotia Sea. It had a limited size range, but there was clear size-related sexual dimorphism with males significantly larger than females. The species had a life cycle of c. 2 years, but the I-group (c. 1 year old, 1 November to 31 October the next year) occurred only in regions close to the APF suggesting that recruitment is restricted to these waters. A seasonal southward migration for P. choriodon is likely as the species occurred mostly to the south-west of South Georgia in summer, but extended to the sea-ice sectors in autumn. Protomyctophum choriodon had a life cycle of 4+ years in the Scotia Sea and the population was dominated by age classes >3 years old. Larval stages were absent during the surveys for all species. Diurnal variations in vertical distribution were apparent for all three species. Interspecific variations in diet were evident, but all species were primarily copepod feeders, with Metridia spp., Rhincalanus gigas and Calanus simillimus generally dominating their diet. Small euphausiids, principally Thysanoessa spp., were also an important component of their diets, particularly for P. choriodon which had the largest body size. The spatial and temporal variations in diet for both P. bolini and P. tenisoni were broadly consistent with underlying abundance patterns within the mesozooplankton community.

On the comparison of population-level estimates of haplotype and nucleotide diversity: a case study using the gene cox1 in animals.

Estimates of genetic diversity represent a valuable resource for biodiversity assessments and are increasingly used to guide conservation and management programs. The most commonly reported estimates of DNA sequence diversity in animal populations are haplotype diversity (h) and nucleotide diversity (π) for the mitochondrial gene cytochrome c oxidase subunit I (cox1). However, several issues relevant to the comparison of h and π within and between studies remain to be assessed. We used population-level cox1 data from peer-reviewed publications to quantify the extent to which data sets can be re-assembled, to provide a standardized summary of h and π estimates, to explore the relationship between these metrics and to assess their sensitivity to under-sampling. Only 19 out of 42 selected publications had archived data that could be unambiguously re-assembled; this comprised 127 population-level data sets (n ≥ 15) from 23 animal species. Estimates of h and π were calculated using a 456-base region of cox1 that was common to all the data sets (median h=0.70130, median π=0.00356). Non-linear regression methods and Bayesian information criterion analysis revealed that the most parsimonious model describing the relationship between the estimates of h and π was π=0.0081 h(2). Deviations from this model can be used to detect outliers due to biological processes or methodological issues. Subsampling analyses indicated that samples of n>5 were sufficient to discriminate extremes of high from low population-level cox1 diversity, but samples of n ≥ 25 are recommended for greater accuracy.

Swarms of diversity at the gene cox1 in Antarctic krill.

The Antarctic krill, Euphausia superba, is an abundant and key species found in the Southern Ocean that forms dense, discrete swarms. Despite over three decades of research on Antarctic krill, the genetics of individual swarms is yet to be specifically investigated. In this study, we address the genetic diversity, population structure and demographic history of nine Antarctic krill swarms by sequencing 1173 bases of the gene cytochrome c oxidase subunit I (cox1, COI) from 504 individuals. Both haplotype diversity (h=0.9974-1.0000) and nucleotide diversity (pi=0.010275-0.011537) of Antarctic krill swarm samples was consistently high compared with populations of other species reported in the literature. Analysis of molecular variance did not show any significant genetic structure, thus implying that the sampled swarms do not appear to reflect discrete genetic units. Fu's Fs and Bayesian Skyride analyses provided strong evidence for a large increase in the population size of Antarctic krill, or selection favouring a particular mitochondrial lineage, within the last few 100,000 years (Pleistocene). The swarm-level results presented in this study not only further our understanding of Antarctic krill biology but, because of the economical importance of this species, also provide data to consider for future krill stock management.

Spatial and temporal operation of the Scotia Sea ecosystem: a review of large-scale links in a krill centred food web.

The Scotia Sea ecosystem is a major component of the circumpolar Southern Ocean system, where productivity and predator demand for prey are high. The eastward-flowing Antarctic Circumpolar Current (ACC) and waters from the Weddell-Scotia Confluence dominate the physics of the Scotia Sea, leading to a strong advective flow, intense eddy activity and mixing. There is also strong seasonality, manifest by the changing irradiance and sea ice cover, which leads to shorter summers in the south. Summer phytoplankton blooms, which at times can cover an area of more than 0.5 million km2, probably result from the mixing of micronutrients into surface waters through the flow of the ACC over the Scotia Arc. This production is consumed by a range of species including Antarctic krill, which are the major prey item of large seabird and marine mammal populations. The flow of the ACC is steered north by the Scotia Arc, pushing polar water to lower latitudes, carrying with it krill during spring and summer, which subsidize food webs around South Georgia and the northern Scotia Arc. There is also marked interannual variability in winter sea ice distribution and sea surface temperatures that is linked to southern hemisphere-scale climate processes such as the El Niño-Southern Oscillation. This variation affects regional primary and secondary production and influences biogeochemical cycles. It also affects krill population dynamics and dispersal, which in turn impacts higher trophic level predator foraging, breeding performance and population dynamics. The ecosystem has also been highly perturbed as a result of harvesting over the last two centuries and significant ecological changes have also occurred in response to rapid regional warming during the second half of the twentieth century. This combination of historical perturbation and rapid regional change highlights that the Scotia Sea ecosystem is likely to show significant change over the next two to three decades, which may result in major ecological shifts.