Nadine M. Johnston

ANTARCTIC MARINE BENTHIC DIVERSITY

Species lists have been compiled for all the major groups of Southern Ocean benthic marine invertebrates, eliminating synonymies where possible and providing a subjective estimate of completeness and reliability for each group. Antarctic marine diversity (pelagic and benthic) is relatively high at the phylum and class level, with the gaps mostly comprising minor, meiofaunal or parasitic groups. Most benthic diversity data come from the continental shelves, with relatively few samples from deeper water. Even for the continental shelves, however, sampling is highly patchy with some areas hardly investigated at all. Over 4100 benthic species have been reported from the Southern Ocean, with the most speciose groups being polychaetes, gastropods and amphipods. Comparison with tropical and temperate regions suggest that decapods, bivalves and teleost fishes are poorly represented in the Southern Ocean benthic marine fauna, whereas pycnogonids, echinoderms and many suspension feeding groups are rich and diverse. Some groups that are currently low in diversity were previously well represented in the Antarctic shallow water marine fauna, notably decapods and many fishes. Other groups have undergone marked radiations in the Southern Ocean, including pycnogonids, amphipods, isopods and teleost fishes; in all cases, however, it is only some lineages that have diversified. This indicates that evolutionary questions concerning the origin, diversification or extinction of the Southern Ocean marine fauna will have no single answer; the evolutionary history of each group appears to reflect a different response to the tectonic, climatic and oceanographic changes to which they have been subject through history. The disposition of southern hemisphere continents makes it difficult to assess whether there is a latitudinal dine in shallow-water marine diversity to mirror that known from the northern hemisphere. Within Antarctica, many species appear to have circumpolar distributions, and the long established biogeographical division into continental Antarctic, Antarctic Peninsula and sub-Antarctic regions have not been challenged by recent sampling. For most groups the frequency distribution of species per genus ratios is typical, though none is well described by the predictions from current evolutionary or null models. Where data are available, size spectra indicate that many Southern Ocean taxa are small, a few spectacular examples of gigantism notwithstanding, and species abundance plots are normal. Knowledge of the Southern Ocean benthic marine fauna has reached a stage where we can now ask powerful evolutionary questions, and the development of new molecular techniques provides the mechanism for answering them.

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.

Climatically driven fluctuations in Southern Ocean ecosystems

Determining how climate fluctuations affect ocean ecosystems requires an understanding of how biological and physical processes interact across a wide range of scales. Here we examine the role of physical and biological processes in generating fluctuations in the ecosystem around South Georgia in the South Atlantic sector of the Southern Ocean. Anomalies in sea surface temperature (SST) in the South Pacific sector of the Southern Ocean have previously been shown to be generated through atmospheric teleconnections with El Niño Southern Oscillation (ENSO)-related processes. These SST anomalies are propagated via the Antarctic Circumpolar Current into the South Atlantic (on time scales of more than 1 year), where ENSO and Southern Annular Mode-related atmospheric processes have a direct influence on short (less than six months) time scales. We find that across the South Atlantic sector, these changes in SST, and related fluctuations in winter sea ice extent, affect the recruitment and dispersal of Antarctic krill. This oceanographically driven variation in krill population dynamics and abundance in turn affects the breeding success of seabird and marine mammal predators that depend on krill as food. Such propagating anomalies, mediated through physical and trophic interactions, are likely to be an important component of variation in ocean ecosystems and affect responses to longer term change. Population models derived on the basis of these oceanic fluctuations indicate that plausible rates of regional warming of 1oC over the next 100 years could lead to more than a 95% reduction in the biomass and abundance of krill across the Scotia Sea by the end of the century.

Climate change and Southern Ocean ecosystems I: how changes in physical habitats directly affect marine biota

Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed.

Palaeolimnology of Antarctic fur seal Arctocephalus gazella populations and implications for Antarctic management

A recent rapid increase in the number of Antarctic fur seals on Signy Island has caused extensive changes to terrestrial and freshwater ecosystems. A palaeolimnological study was carried out to determine whether the recent increase in the number of visiting fur seals has exceeded the range of natural variability of past populations and if a management response should be considered. Seal hairs in lake sediments from Sombre Lake were assessed as indicators of visiting fur seals. Results showed that sediments could be used to detect the presence, absence and possibly broad scale changes in the abundance of fur seals and indicated that fur seals have visited the island from the beginning of the sedimentary record (6570 ± 60 14C years B.P.) when the coastline became ice-free. However, the previous maximum number of seals visiting the study site appears to have been no more than a quarter of the number recorded in 1994. The timing of changes in fur seal hair abundance indicates that historical activities of the sealing and whaling industries may be partly responsible for the current increase in fur seal populations. The results of this study have provided a unique historical perspective from which appropriate conservation measures at Signy Island, and possibly at other sites on northern maritime Antarctic islands and on the west coast of the Antarctic Peninsula, can be considered to address and minimise the impact of the Antarctic fur seals.

Antarctic ecosystems: an extreme environment in a changing world.

Since its discovery Antarctica has held a deep fascination for biologists. Extreme environmental conditions, seasonality and isolation have lead to some of the most striking examples of natural selection and adaptation on Earth. Paradoxically, some of these adaptations may pose constraints on the ability of the Antarctic biota to respond to climate change. Parts of Antarctica are showing some of the largest changes in temperature and other environmental conditions in the world. In this volume, published in association with the Royal Society, leading polar scientists present a synthesis of the latest research on the biological systems in Antarctica, covering organisms from microbes to vertebrate higher predators. This book comes at a time when new technologies and approaches allow the implications of climate change and other direct human impacts on Antarctica to be viewed at a range of scales; across entire regions, whole ecosystems and down to the level of species and variation within their genomes. Chapters address both Antarctic terrestrial and marine ecosystems, and the scientific and management challenges of the future are explored.

Introduction. Antarctic ecology from genes to ecosystems: the impact of climate change and the importance of scale

Antarctica offers a unique natural laboratory for undertaking fundamental research on the relationship between climate, evolutionary processes and molecular adaptation. The fragmentation of Gondwana and the development of wide-scale glaciation have resulted in major episodes of extinction and vicariance, as well as driving adaptation to an extreme environment. On shorter time-scales, glacial cycles have resulted in shifts in distribution, range fragmentation and allopatric speciation, and the Antarctic Peninsula is currently experiencing among the most rapid climatic warming on the planet. The recent revolution in molecular techniques has provided a suite of innovative and powerful tools to explore the consequences of these changes, and these are now providing novel insights into evolutionary and ecological processes in Antarctica. In addition, the increasing use of remotely sensed data is providing a large-scale view of the system that allows these processes to be set in a wider spatial context. In these two volumes, we collect a wide range of papers exploring these themes, concentrating on recent advances and emphasizing the importance of spatial and temporal scale in understanding ecological and evolutionary processes in Antarctica.

Introduction: Antarctic ecology from genes to ecosystems. Part 2. Evolution, diversity and functional ecology

The Antarctic biota has evolved over the last 100 million years in increasingly isolated and cold conditions. As a result, Antarctic species, from micro-organisms to vertebrates, have adapted to life at extremely low temperatures, including changes in the genome, physiology and ecological traits such as life history. Coupled with cycles of glaciation that have promoted speciation in the Antarctic, this has led to a unique biota in terms of biogeography, patterns of species distribution and endemism. Specialization in the Antarctic biota has led to trade-offs in many ecologically important functions and Antarctic species may have a limited capacity to adapt to present climate change. These include the direct effects of changes in environmental parameters and indirect effects of increased competition and predation resulting from altered life histories of Antarctic species and the impacts of invasive species. Ultimately, climate change may alter the responses of Antarctic ecosystems to harvesting from humans. The unique adaptations of Antarctic species mean that they provide unique models of molecular evolution in natural populations. The simplicity of Antarctic communities, especially from terrestrial systems, makes them ideal to investigate the ecological implications of climate change, which are difficult to identify in more complex systems.

Understanding the structure and functioning of polar pelagic ecosystems to predict the impacts of change

The determinants of the structure, functioning and resilience of pelagic ecosystems across most of the polar regions are not well known. Improved understanding is essential for assessing the value of biodiversity and predicting the effects of change (including in biodiversity) on these ecosystems and the services they maintain. Here we focus on the trophic interactions that underpin ecosystem structure, developing comparative analyses of how polar pelagic food webs vary in relation to the environment. We highlight that there is not a singular, generic Arctic or Antarctic pelagic food web, and, although there are characteristic pathways of energy flow dominated by a small number of species, alternative routes are important for maintaining energy transfer and resilience. These more complex routes cannot, however, provide the same rate of energy flow to highest trophic-level species. Food-web structure may be similar in different regions, but the individual species that dominate mid-trophic levels vary across polar regions. The characteristics (traits) of these species are also different and these differences influence a range of food-web processes. Low functional redundancy at key trophic levels makes these ecosystems particularly sensitive to change. To develop models for projecting responses of polar ecosystems to future environmental change, we propose a conceptual framework that links the life histories of pelagic species and the structure of polar food webs.

A synergistic approach for evaluating climate model output for ecological applications

Increasing concern about the impacts of climate change on ecosystems is prompting ecologists and ecosystem managers to seek reliable projections of physical drivers of change. The use of global climate models in ecology is growing, although drawing ecologically meaningful conclusions can be problematic. The expertise required to access and interpret output from climate and earth system models is hampering progress in utilizing them most effectively to determine the wider implications of climate change. To address this issue, we present a joint approach between climate scientists and ecologists that explores key challenges and opportunities for progress. As an exemplar, our focus is the Southern Ocean, notable for significant change with global implications, and on sea ice, given its crucial role in this dynamic ecosystem. We combined perspectives to evaluate the representation of sea ice in global climate models. With an emphasis on ecologically-relevant criteria (sea ice extent and seasonality) we selected a subset of eight models that reliably reproduce extant sea ice distributions. While the model subset shows a similar mean change to the full ensemble in sea ice extent (approximately 50% decline in winter and 30% decline in summer), there is a marked reduction in the range. This improved the precision of projected future sea ice distributions by approximately one third, and means they are more amenable to ecological interpretation. We conclude that careful multidisciplinary evaluation of climate models, in conjunction with ongoing modeling advances, should form an integral part of utilizing model output.