Global Connectivity of Southern Ocean Ecosystems

Southern Ocean ecosystems are globally important. Processes in the Antarctic atmosphere, cryosphere, and the Southern Ocean directly influence global atmospheric and oceanic systems. Southern Ocean biogeochemistry has also been shown to have global importance. In contrast, ocean ecological processes are often seen as largely separate from the rest of the global system. In this paper, we consider the degree of ecological connectivity at different trophic levels, linking Southern Ocean ecosystems with the global ocean, and their importance not only for the regional ecosystem but also the wider Earth system. We also consider the human system connections, including the role of Southern Ocean ecosystems in supporting society, culture, and economy in many nations, influencing public and political views and hence policy. Rather than Southern Ocean ecosystems being defined by barriers at particular oceanic fronts, ecological changes are gradual due to cross-front exchanges involving oceanographic processes and organism movement. Millions of seabirds and hundreds of thousands of cetaceans move north out of polar waters in the austral autumn interacting in food webs across the Southern Hemisphere, and a few species cross the equator. A number of species migrate into the east and west ocean-basin boundary current and continental shelf regions of the major southern continents. Human travel in and out of the Southern Ocean region includes fisheries, tourism, and scientific vessels in all ocean sectors. These operations arise from many nations, particularly in the Northern Hemisphere, and are important in local communities as well as national economic, scientific, and political activities. As a result of the extensive connectivity, future changes in Southern Ocean ecosystems will have consequences throughout the Earth system, affecting ecosystem services with socio-economic impacts throughout the world. The high level of connectivity also means that changes and policy decisions in marine ecosystems outside the Southern Ocean have consequences for ecosystems south of the Antarctic Polar Front. Knowledge of Southern Ocean ecosystems and their global connectivity is critical for interpreting current change, projecting future change impacts, and identifying integrated strategies for conserving and managing both the Southern Ocean and the broader Earth system.

Winter Biogeochemical Cycling of Dissolved and Particulate Cadmium in the Indian Sector of the Southern Ocean (GEOTRACES GIpr07 Transect)

Winter distributions of dissolved cadmium (dCd) and particulate cadmium (pCd) were measured for the first time in the Indian sector of the Southern Ocean thereby contributing a unique spatial and seasonal dataset. Seven depth profiles, between 41°S and 58°S, were collected along the 30°E longitude during the 2017 austral winter to investigate the biogeochemical cycling of cadmium during a period characterized by contrasting upper water column dynamics compared to summer. Our results support an important role for biological uptake during winter months albeit weaker compared to summer. Distinct, biologically driven changes in cadmium cycling across the transect were observed. For example, surface ratios of pCd to phosphorus (P; pCd:P) increased from 0.37 to 1.07 mmol mol–1 between the subtropical zone (STZ) and the Antarctic zone (AAZ) reflecting increased Cd requirements for diatoms at higher latitudes which, in turn, was driven by a complex relationship between the availability of dCd and dissolved iron (dFe), zinc (dZn) and manganese (dMn). Vertical profiles of pCd:P displayed near-surface maxima consistent with (1) P occurring in two phases with different labilities and the lability of Cd being somewhere in-between and (2) increasing dCd to phosphate (PO4; dCd:PO4) ratios with depth at each station. North of the Antarctic Polar Front (APF), a secondary, deeper pCd:P maximum may reflect an advective signal associated with northward subducting Antarctic Intermediate Water (AAIW). The strong southward increase in surface dCd and dCd:PO4, from approximately 10–700 pmol kg–1 and 40–400 μmol mol–1, respectively, reflected the net effect of preferential uptake and regeneration of diatoms with high Cd content and the upwelling of Cd enriched water masses in the AAZ. Furthermore, distinct dCd versus PO4 relationships were observed in each of the intermediate and deep water masses suggesting that dCd and PO4 distributions at depth are largely the result of physical water mass mixing.

Exploring the Microdiversity Within Marine Bacterial Taxa: Toward an Integrated Biogeography in the Southern Ocean

Most of the microbial biogeographic patterns in the oceans have been depicted at the whole community level, leaving out finer taxonomic resolution (i.e., microdiversity) that is crucial to conduct intra-population phylogeographic study, as commonly done for macroorganisms. Here, we present a new approach to unravel the bacterial phylogeographic patterns combining community-wide survey by 16S rRNA gene metabarcoding and intra-species resolution through the oligotyping method, allowing robust estimations of genetic and phylogeographic indices, and migration parameters. As a proof-of-concept, we focused on the bacterial genus Spirochaeta across three distant biogeographic provinces of the Southern Ocean; maritime Antarctica, sub-Antarctic Islands, and Patagonia. Each targeted Spirochaeta operational taxonomic units were characterized by a substantial intrapopulation microdiversity, and significant genetic differentiation and phylogeographic structure among the three provinces. Gene flow estimations among Spirochaeta populations support the role of the Antarctic Polar Front as a biogeographic barrier to bacterial dispersal between Antarctic and sub-Antarctic provinces. Conversely, the Antarctic Circumpolar Current appears as the main driver of gene flow, connecting sub-Antarctic Islands with Patagonia and maritime Antarctica. Additionally, historical processes (drift and dispersal limitation) govern up to 86% of the spatial turnover among Spirochaeta populations. Overall, our approach bridges the gap between microbial and macrobial ecology by revealing strong congruency with macroorganisms distribution patterns at the populational level, shaped by the same oceanographic structures and ecological processes.

Fifty million years of beetle evolution along the Antarctic Polar Front

Global cooling and glacial–interglacial cycles since Antarctica’s isolation have been responsible for the diversification of the region’s marine fauna. By contrast, these same Earth system processes are thought to have played little role terrestrially, other than driving widespread extinctions. Here, we show that on islands along the Antarctic Polar Front, paleoclimatic processes have been key to diversification of one of the world’s most geographically isolated and unique groups of herbivorous beetles—Ectemnorhinini weevils. Combining phylogenomic, phylogenetic, and phylogeographic approaches, we demonstrate that these weevils colonized the sub-Antarctic islands from Africa at least 50 Ma ago and repeatedly dispersed among them. As the climate cooled from the mid-Miocene, diversification of the beetles accelerated, resulting in two species-rich clades. One of these clades specialized to feed on cryptogams, typical of the polar habitats that came to prevail under Miocene conditions yet remarkable as a food source for any beetle. This clade’s most unusual representative is a marine weevil currently undergoing further speciation. The other clade retained the more common weevil habit of feeding on angiosperms, which likely survived glaciation in isolated refugia. Diversification of Ectemnorhinini weevils occurred in synchrony with many other Antarctic radiations, including penguins and notothenioid fishes, and coincided with major environmental changes. Our results thus indicate that geo-climatically driven diversification has progressed similarly for Antarctic marine and terrestrial organisms since the Miocene, potentially constituting a general biodiversity paradigm that should be sought broadly for the region’s taxa.

Exploring the microdiversity within marine bacterial taxa: Towards an integrated biogeography in the Southern Ocean

The phylogeography traditionally correlates the genetic relationships among individuals within a macroorganism species, to their spatial distribution. Most microbial phylogeographic studies so far have been restricted to narrow geographical regions, mainly focusing on isolated strains, either obtained by culture or single-strain natural enrichments. However, the laborious culture-based methodology imposes a low number of studied individuals, leading to poor resolution of haplotype frequency estimation, making difficult a realistic evaluation of the genetic structure of natural microbial populations in the environment.To tackle this limitation, we present a new approach to unravel the phylogeographic patterns of bacteria combining (i) community-wide survey by 16S rRNA gene metabarcoding, (ii) intra-species resolution through the oligotyping method, and (iii) genetic and phylogeographic indices, as well as migration parameters, estimated from populational molecular data as traditionally developed for macroorganisms as models.As a proof-of-concept, we applied this methodology to the bacterial genus Spirochaeta, classically reported as a gut endosymbiont of various invertebrates inhabiting the Southern Ocean (SO), but also described in marine sediment and in open waters. For this purpose, we centered our sampling into three biogeographic provinces of the SO; maritime Antarctica (King George Island), sub-Antarctic Islands (Kerguelen archipelago) and Patagonia in southern South America. Each targeted OTU was chaLRracterized by substantial intrapopulation microdiversity, a significant genetic differentiation and a robust phylogeographic structure among the three distant biogeographic provinces. Patterns of gene flow in Spirochaeta populations support the role of the Antarctic Polar Front (APF) as a biogeographic barrier to bacterial dispersal between Antarctic and sub-Antarctic provinces. Conversely, the Antarctic Circumpolar Current (ACC) appears as the main driver of connectivity between geographically distant sub-Antarctic areas such as Patagonia and Kerguelen archipelago, and between Kerguelen archipelago and maritime Antarctica. Additionnally, we found that historical processes (drift and dispersal limitation) together govern up to 86% of the spatial turnover among Spirochaeta populations. Overall, our approach represents a substantial first attempt to bridge the gap between microbial and macrobial ecology by unifying the way to study phylogeography. We revealed that strong congruency with macroorganisms patterns at the populational level shaped by the same oceanographic structures and ecological processes.

Northward Migration of Antarctic Polar Front during the Quaternary: Planktic Foraminiferal Record from Southeast Indian Ocean

The ODP Hole 763A is influenced by the northward-flowing cold West Australian Current (WAC) and Southward flowing warm Leeuwin Current (LC). LC is a branch of the South Equatorial Current (SEC), which brings relatively warmer waters from the tropical Pacific Ocean into the Indian Ocean via Indonesian Throughflow (ITF). The modern planktic foraminiferal fauna thrives along the western margin of Australia. It consists mainly of warm water assemblages brought by the LC. The present study provides planktic foraminiferal census data from ODP Hole 763A, influenced by the LC and WAC, to document the history of cold water influence at the site during the quaternary. The northward migration of the Antarctic Polar Front (APF) and resultant intensification of the cold West Australian Current have been inferred based on the dramatic increase in the relative abundance of temperate water species group Globoconella at Hole 763A situated in the low latitude region. The Quaternary planktic foraminiferal census data shows several episodes of invasion of Globoconella. These intervals of high abundance of Globoconella group have been attributed to the intensification of WAC, probably due to Antarctic ice volume expansion and resultant northward migration of the APF at 0.05 Ma, 0.2 Ma, 0.45 Ma, 0.7 Ma and 1.2 Ma. We have documented that the amplitude of fluctuations in cold/warm events has increased after the Mid-Pleistocene Transition (MPT). LC is a heat supplier to the higher latitudes, its weakening during such intervals might have contributed to the ice volume expansion over Antarctica. Thus, the study proposes that the Antarctic ice cap formation creates a positive feedback mechanism by lowering sea level, reduced strength of LC due to a decrease in ITF and less heat supply towards the South Pole. All these phenomena add to further cooling.

Wind-driven Advection Across Temperature Gradients Supports Phytoplankton Blooms in the Iron-limited Antarctic Polar Front

We demonstrate how the wind-driven Ekman transport enhances the advection and mixing of cells from the colder waters of the Surface Antarctic Waters from the south to the warmer waters of the northern Polar Front (PF) belt. This mechanism provides cells a mean ambient temperature near optimum levels for species-specific and, ultimately, community growth rates high enough to develop blooms under non-light limiting, macronutrients and iron conditions. A Lagrangian trajectory model was constructed for tracking plankton cells as tracers forced by winds and surface currents. We argue that wind-driven Ekman drift of surface currents can carry phytoplankton cells into warmer waters and thus increase their growth rates to potentially generate blooms, even under iron-limiting conditions. Depending on the region along the circumpolar front, increased winds can enhance this process, and further accelerate such temperature-controlled growth.

Humpback whale abundance south of 60°S from three complete circumpolar sets of surveys

Austral summer estimates of abundance are obtained for humpback whales (Megaptera novaeangliae) in the Southern Ocean from the IWC’s IDCR and SOWER circumpolar programmes. These surveys have encircled the Antarctic three times: 1978/79–1983/84 (CPI), 1985/86–1990/91 (CPII) and 1991/92–2003/04 (CPIII), criss-crossing strata totalling respectively 64.3%, 79.5% and 99.7% of the open-ocean area south of 60°S. Humpback whales were absent from the Ross Sea, but were sighted in all other regions, and in particularly high densities around the Antarctic Peninsula, in Management Area IV and north of the Ross Sea. Abundance estimates are presented for each CP, for Management Areas, and for assumed summer feeding regions of each Breeding Stock. Abundance estimates are negatively biased because some whales on the trackline are missed and because some humpback whales are outside the survey region. Circumpolar estimates with approximate midpoints of 1980/81, 1987/88 and 1997/98 are 7,100 (CV = 0.36), 10,200 (CV = 0.30) and 41,500 (CV = 0.11). When these are adjusted simply for unsurveyed northern areas, the estimated annual rate of increase is 9.6% (95% CI 5.8–13.4%). All Breeding Stocks are estimated to be increasing but increase rates are significantly greater than zero only for those on the eastern and western coasts of Australia. Given the observed rates of increase, the current total Southern Hemisphere abundance is greater than 55,000, which is similar to the summed northern breeding ground estimates (~60,000 from 1999–2008). Some breeding ground abundance estimates are far greater, and others far lower, than the corresponding IDCR/SOWER estimates, in a pattern apparently related to the latitudinal position of the Antarctic Polar Front.