Dietary divergence is associated with increased intra-specific competition in a marine predator

Optimal foraging theory predicts that when food is plentiful all individuals should take a small range of preferred prey types, but as competition increases less preferred prey will be included in the diet. This dietary switching may not be uniform among individuals, which produces discrete dietary clusters. We tested this hypothesis for gentoo penguins at Bird Island, South Georgia, using stable isotope analysis and biologging. Competition, in the form of the density of foraging dives, increased markedly from incubation to chick-rearing owing to increased foraging effort. Birds responded behaviourally by exploiting a greater portion of the available foraging radius and increasing dive depths. Dietary niche width doubled and two discrete dietary clusters appeared; one comprising birds that consumed mostly krill and another that ate a greater proportion of demersal fish. There were no differences in morphology between the dietary classes, but birds in the fish class had a tendency to dive deeper, which suggests a behavioural basis for specialization. Our findings are consistent with the hypothesis that intra-specific competition expands the population’s dietary niche width and drives divergence in diets among individuals

The contribution of zooplankton faecal pellets to deep carbon transport in the Scotia Sea (Southern Ocean)

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 in 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 × 10 FP m−2 d−1) were an order of magnitude higher than at P2 (4.01 mg C m−2 d−1; 915 × 10 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 community shift from an 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 behavior 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

Experimental determination of reflectance spectra of Antarctic krill (Euphausia superba) in the Scotia Sea

Antarctic krill are the dominant metazoan in the Southern Ocean in terms of biomass; however, their wide and patchy distribution means that estimates of their biomass are still uncertain. Most currently employed methods do not sample the upper surface layers, yet historical records indicate that large surface swarms can change the water colour. Ocean colour satellites are able to measure the surface ocean synoptically and should theoretically provide a means for detecting and measuring surface krill swarms. Before we can assess the feasibility of remote detection, more must be known about the reflectance spectra of krill. Here, we measure the reflectance spectral signature of Antarctic krill collected in situ from the Scotia Sea and compare it to that of in situ water. Using a spectroradiometer, we measure a strong absorption feature between 500 and 550 nm, which corresponds to the pigment astaxanthin, and high reflectance in the 600–700 nm range due to the krill's red colouration. We find that the spectra of seawater containing krill is significantly different from seawater only. We conclude that it is tractable to detect high-density swarms of krill remotely using platforms such as optical satellites and unmanned aerial vehicles, and further steps to carry out ground-truthing campaigns are now warranted

Predatory impact of the myctophid fish community on zooplankton in the Scotia Sea (Southern Ocean)

Myctophids are the most abundant mesopelagic fishes in the Southern Ocean, although their trophic role within the predominantly krill-based food web in regions south of the Antarctic Polar Front (APF) is poorly resolved. This study therefore examined the diets of 10 species of myctophid fishes: Electrona antarctica, E. carlsbergi, Gymnoscopelus braueri, G. fraseri, G. nicholsi, Krefftichthys anderssoni, Protomyctophum bolini, P. tenisoni, P. choriodon and Nannobrachium achirus, in the Scotia Sea, together with their predatory impact on the underlying zooplankton community. Myctophids and their prey were sampled in different seasons by scientific nets deployed across the Scotia Sea from the sea-ice zone to the APF. Based on the percentage index of relative importance, myctophids had high overlap in their diets, although the data indicate dietary specialisation in some species. There was also a distinct switch in diet, from copepods to euphausiids and amphipods, with increasing myctophid size. Myctophid predation impacted daily copepod production by between 0.01 and 5%, with Calanus simillimus being most impacted. Total annual consumption of copepods was around 1.5 million t (Mt) per year. All myctophids preyed upon the euphausiid Thysanoessa spp., consuming ~12% of its daily productivity and around 4 Mt per year. However, only larger myctophid species preyed upon Antarctic krill Euphausia superba, consuming 2% of its daily productivity, which could amount to as much as 17 Mt per year. Themisto gaudichaudii was also an important dietary component, with 4% of its daily productivity being consumed, amounting to around 2 Mt per year. This study demonstrates that myctophids link secondary productivity to higher predators both through krill-dependent and krill-independent trophic pathways

Intra-specific niche partitioning in Antarctic fur seals, Arctocephalus gazella

Competition for resources within a population can lead to niche partitioning between sexes, throughout ontogeny and among individuals, allowing con-specifics to co-exist. We aimed to quantify such partitioning in Antarctic fur seals, Arctocephalus gazella, breeding at South Georgia, which hosts ~95% of the world’s population. Whiskers were collected from 20 adult males and 20 adult females and stable isotope ratios were quantified every 5 mm along the length of each whisker. Nitrogen isotope ratios (δ15N) were used as proxies for trophic position and carbon isotope ratios (δ13C) indicated foraging habitat. Sexual segregation was evident: δ13C values were significantly lower in males than females, indicating males spent more time foraging south of the Polar Front in maritime Antarctica. In males δ13C values declined with age, suggesting males spent more time foraging south throughout ontogeny. In females δ13C values revealed two main foraging strategies: 70% of females spent most time foraging south of the Polar Front and had similar δ15N values to males, while 30% of females spent most time foraging north of the Polar Front and had significantly higher δ15N values. This niche partitioning may relax competition and ultimately elevate population carrying capacity with implications for ecology, evolution and conservation

Respiration of mesopelagic fish: a comparison of respiratory electron transport system (ETS) measurements and allometrically calculated rates in the Southern Ocean and Benguela Current

Mesopelagic fish are an important component of marine ecosystems, and their contribution to marine biogeochemical cycles is becoming increasingly recognized. However, major uncertainties remain in the rates at which they remineralize organic matter. We present respiration rate estimates of mesopelagic fish from two oceanographically contrasting regions: the Scotia Sea and the Benguela Current. Respiration rates were estimated by measuring the enzyme activities of the electron transport system. Regression analysis of respiration with wet mass highlights regional and inter-specific differences. The mean respiration rates of all mesopelagic fish sampled were 593.6 and 354.9 µl O2 individual−1 h−1 in the Scotia Sea and Benguela Current, respectively. Global allometric models performed poorly in colder regions compared with our observations, underestimating respiratory flux in the Scotia Sea by 67–88%. This may reflect that most data used to fit such models are derived from temperate and subtropical regions. We recommend caution when applying globally derived allometric models to regional data, particularly in cold (<5°C) temperature environments where empirical data are limited. More mesopelagic fish respiration rate measurements are required, particularly in polar regions, to increase the accuracy with which we can assess their importance in marine biogeochemical cycles

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

Mesozooplankton community composition controls faecal pellet flux and remineralisation depth in the Southern Ocean

Zooplankton faecal pellets (FPs) are important conduits of carbon from the surface to the deep ocean, as shown by their presence in deep-sea sediment traps. Zooplankton themselves are thought to play an important role in the breakdown and reworking of FPs as they sink, whilst processes such as diel vertical migration (DVM) may enhance the supply of carbon to the mesopelagic. However, comparatively little is known about the processes or variability of FP sinking/ transport within the upper mesopelagic and how this relates to deeper ocean export. Profiles of FP type and size, and the contribution made by FPs to mesopelagic carbon flux to a depth of 400 m, were considered. Three contrasting locations in the Scotia Sea were compared, which together reflect the variability in physical regime and productivity encountered across the Southern Ocean. Comparing observed FPs with predictions from the mesozooplankton community, we show that, even at shallow depths, the smallest fraction of FP is under-represented, suggesting rapid remineralisation, incorporation into larger aggregates or reworking into larger FPs, and that the flux is dominated by FPs from larger zooplankton. In contrast to models where POC attenuation rates are set to increase with temperature, we find that FP carbon flux attenuates rapidly in low productivity, colder regions dominated by krill, while remineralisation is deeper in warmer areas where productivity is high and copepods dominate. This emphasises the strong modulation of the zooplankton community on the supply and transfer of FP carbon between the epi- and mesopelagic. Evidence was found to suggest that DVM enhances FP flux across the upper mesopelagic, producing a pulse of fresh, dense material that may support secondary production and heterotrophic respiration in the mesopelagic. This illustrates that variability in flux at short (daily) as well as longer (seasonal) timescales may have important implications for the supply of FP carbon to deeper waters

Mercury levels in Southern Ocean squid : variability over the last decade

Authors acknowledge the Portuguese Foundation for the Science and Technology (FCT) through a PhD grant to José Seco (SRFH/PD/BD/113487). Acknowledgments are due also to the Integrated Program of SR&TD ‘Smart Valorization of Endogenous Marine Biological Resources Under a Changing Climate’ (reference Centro-01-0145-FEDER-000018), co-funded by Centro (2020) program, Portugal 2020, European Union, through the European Regional Development Fund, for personal funding to J.P.Coelho. The IUF (Institut Universitaire de France) is acknowledged for its support to P. Bustamante as a Senior Member. This research was also within José Xavier strategic program of MARE (MARE - UID/MAR/04292/2013). GAT, GS and SF were supported by the Ecosystems programme at the British Antarctic Survey.The concentrations of total and proportions of organic mercury were measured in tissues of 355 individuals of 8 species of Southern Ocean squid (Alluroteuthis antarcticus, Bathyteuthis abyssicola, Filippovia knipovitchi, Galiteuthis glacialis, Gonatus antarcticus, Kondakovia longimana, Psychroteuthis glacialis and Slosarczykovia circumantarctica). Squid were caught around South Georgia (Scotia Sea) during 5 cruises, between the austral summers of 2006/07 to 2016/17 to evaluate temporal changes in bioaccumulation and tissue partitioning. Total mercury concentrations varied between 4 ng g−1 and 804 ng g−1 among all tissues. Net accumulation of mercury in muscle with size was observed in A. antarcticus, B. abyssicola and P. glacialis, but no relationship was found for S. circumantarctica and lower concentrations were observed in larger individuals of G. glacialis. Muscle tissues had the highest mercury concentrations in the majority of species, except for F. knipovitchi for which the digestive gland contained highest concentrations. In terms of the percentage of organic mercury in the tissues, muscle always contained the highest values (67%–97%), followed by the digestive gland (22%–38%). Lowest organic mercury percentages were found consistently in the gills (9%–19%), suggesting only low levels of incorporation through the dissolved pathway and/or a limited redistribution of dietary organic mercury towards this tissue. Overall, results are indicative of a decreasing trend of mercury concentrations in the majority of analysed species over the last decade. As cephalopods are an important Southern Ocean trophic link between primary consumers and top predators, these changes suggest decreasing mercury levels in lower trophic levels and an alleviation of the mercury burden on higher predators that consume squid.PostprintPeer reviewe

Seasonal variation in the predatory impact of myctophids on zooplankton in the Scotia Sea (Southern Ocean)

Myctophids are the biomass-dominant mesopelagic fishes in the Southern Ocean, but their trophic role within the pelagic food web south of the Antarctic Polar Front is poorly resolved from a seasonal perspective at the ocean-basin scale. In this study, the predatory impact of the predominant Southern Ocean myctophid community (Electrona antarctica, Electrona carlsbergi, Gymnoscopelus braueri, Gymnoscopelus fraseri, Gymnoscopelus nicholsi, Protomyctophum bolini, Protomyctophum tenisoni, Protomyctophum choriodon, Krefftichthys anderssoni and Nannobrachium achirus) on their zooplankton prey was examined during austral spring, summer and autumn in the Scotia Sea, one of the most productive regions of the Southern Ocean. Seasonal variations in diet and predation rates were apparent for all species. Based on the percentage index of relative importance, myctophids had high overlap in their diets, with all species mostly consuming copepods, small euphausiids and amphipods. Myctophid size was a key determinant of diet in the region, with larger species and intra-specific size classes consuming larger prey. Cluster analyses revealed myctophid feeding guilds that appeared to change seasonally, although there was little evidence of dietary specialisation. Myctophid predation on the daily productivity of most copepod species was relatively low across seasons (<7%), except for Calanus simillimus that was predated upon highly in summer (∼26%). From the macrozooplankton component of the prey field, the myctophid community consumed substantial proportions of the euphausiid Thysanoessa spp. in each season (∼7 to 76% daily productivity), particularly in summer. Relatively high proportions of the daily Antarctic krill (Euphausia superba) productivity (∼8–58%) were also consumed by the larger myctophid species, particularly in summer by Electrona antarctica, suggesting increased competition for krill resources during the higher predator breeding season and possible reductions in food web stability during periods of reduced krill availability at this time. The amphipod Themisto gaudichaudii formed an important part of the larger myctophid species’ diet in all seasons, with between 10 and 38% of its daily productivity being consumed. Myctophid predation on the daily productivity of salps was up to 4%, whilst their impact on ostracods and pteropods was negligible (<0.1% of daily productivity) in all seasons. This study demonstrates that Southern Ocean myctophids link secondary productivity to higher predators through both krill-independent and krill-dependent trophic pathways across seasons, with myctophids comprising a more krill-dependent pathway during austral summer