Michael D. Sumner

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.

Bayesian Estimation of Animal Movement from Archival and Satellite Tags

The reliable estimation of animal location, and its associated error is fundamental to animal ecology. There are many existing techniques for handling location error, but these are often ad hoc or are used in isolation from each other. In this study we present a Bayesian framework for determining location that uses all the data available, is flexible to all tagging techniques, and provides location estimates with built-in measures of uncertainty. Bayesian methods allow the contributions of multiple data sources to be decomposed into manageable components. We illustrate with two examples for two different location methods: satellite tracking and light level geo-location. We show that many of the problems with uncertainty involved are reduced and quantified by our approach. This approach can use any available information, such as existing knowledge of the animals potential range, light levels or direct location estimates, auxiliary data, and movement models. The approach provides a substantial contribution to the handling uncertainty in archival tag and satellite tracking data using readily available tools.

How many seabirds do we need to track to define home-range area?

1. In recent years, marine predator and seabird tracking studies have become ever more popular. However, they are often conducted without first considering how many individuals should be tracked and for how long they should be tracked in order to make reliable predictions of a population’s home-range area. 2. Home-range area analysis of two seabird-tracking data sets was used to define the area of active use (where birds spent 100% of their time) and the core foraging area (where birds spent 50% of their time). Analysis was conducted on the first foraging trip undertaken by the birds and then the first two, three and four foraging trips combined. Appropriate asymptotic models were applied to the data, and the calculated home-range areas were plotted as a function of an increasing number of individuals and trips included in the sample. Data were extrapolated from these models to predict the area of active use and the core foraging area of the colonies sampled. 3. Significant variability was found in the home-range area predictions made by analysis of the first foraging trip and the first four foraging trips combined. For shags, the first foraging trip predicted a 56% smaller area of active use when compared to the predictions made by combining the first four foraging trips. For kittiwakes, a 43% smaller area was predicted when comparing the first foraging trip with the four combined trips. 4. The number of individuals that would be required to predict the home range area of the colony depends greatly on the number of trips included in the analysis. This analysis predicted that 39 (confidence interval 29–73) shags and 83 (CI: 109–161) kittiwakes would be required to predict 95% of the area of active use when the first four foraging trips are included in the sample compared with 135 (CI 96–156) shags and 248 (164–484) kittiwakes when only the first trip is included in the analysis. 5. Synthesis and applications. Seabird and marine mammal tracking studies are increasingly being used to aid the designation of marine conservation zones and to predict important foraging areas. We suggest that many studies may be underestimating the size of these foraging areas and that better estimates could be made by considering both the duration and number of data logger deployments. Researchers intending to draw conclusions from tracking data should conduct a similar analysis of their data as used in this study to determine the reliability of their home-range area predictions.

Migrations and foraging of juvenile southern elephant seals from Macquarie Island within CCAMLR managed areas

Twenty-three juvenile (8–14 months of age) southern elephant seals (Mirounga leonina L.) from Macquarie Island were tracked during 1993 and 1995. Migratory tracks and ocean areas with concentrated activity, presumed to be foraging grounds, were established from location data gathered by attached geolocation-time depth recorders. The seals ranged widely (811–3258 km) and foraging activity centred on oceanographic frontal systems, especially the Antarctic Polar Front and bathymetric features such as the Campbell Plateau region. The seals spent 58.6% of their sea time within managed fishery areas while the remainder was spent on the high seas, an area of unregulated fishing. The Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR) areas 58.4.1, 88.2 and especially 88.1 were important and distant foraging areas for these juvenile elephant seals. From fisheries records, diet and the foraging ecology studies of the seals there appears to be little, if any, overlap or conflict between the seals and commercial fishing operations within the regulated commercial areas. However, attention is drawn to the possibility of future interactions if Southern Ocean fisheries expand or new ones commence.

Depletion of deep marine food patches forces divers to give up early

Many optimal foraging models for diving animals examine strategies that maximize time spent in the foraging zone, assuming that prey acquisition increases linearly with search time. Other models have considered the effect of patch quality and predict a net energetic benefit if dives where no prey is encountered early in the dive are abandoned. For deep divers, however, the energetic benefit of giving up is reduced owing to the elevated energy costs associated with descending to physiologically hostile depths, so patch residence time should be invariant. Others consider an asymptotic gain function where the decision to leave a patch is driven by patch-depletion effects - the marginal value theorem. As predator behaviour is increasingly being used as an index of marine resource density and distribution, it is important to understand the nature of this gain function. We investigated the dive behaviour of the worlds deepest-diving seal, the southern elephant seal Mirounga leonina, in response to patch quality. Testing these models has largely been limited to controlled experiments on captive animals. By integrating in situ measurements of the seals relative lipid content obtained from drift rate data (a measure of foraging success) with area-restricted search behaviour identified from first-passage time analysis, we identified regions of high- and low-quality patches. Dive durations and bottom times were not invariant and did not increase in regions of high quality; rather, both were longer when patches were of relatively low quality. This is consistent with the predictions of the marginal value theorem and provides support for a nonlinear relationship between search time and prey acquisition. We also found higher descent and ascent rates in high-quality patches suggesting that seals minimized travel time to the foraging patch when quality was high; however, this was not achieved by increasing speed or dive angle. Relative body lipid content was an important predictor of dive behaviour. Seals did not schedule their diving to maximize time spent in the foraging zone in higher-quality patches, challenging the widely held view that maximizing time in the foraging zone translates to greater foraging success.

Temporal changes in the quality of hot-iron brands on elephant seal (Mirounga leonina) pups

Hot-iron brands were used to mark permanently 14 000 six-week-old southern elephant seal (Mirounga leonina L.) pups at Macquarie Island between 1993 and 2000. We assessed temporal changes in the quality of 4932 brands applied in 1998 and 1999 to determine the duration of the brand wound, and the relationships between brand healing, brand readability and the amount of skin and hair damage peripheral to the brand characters. Most (98%) brand wounds were healed within one year. Brand-mark healing, peripheral skin damage and brand readability were significantly (P 95%) for the marked population. The mean number of brand characters with peripheral skin damage decreased significantly over the same period. The seal’s annual hair and skin moult is the process that contributed most to the healing of brand wounds. We also assessed our branding technique to determine whether any of the features we measured contributed to a poor-quality brand. Excessive pressure used during brand-iron application is the most probable cause of unsightly peripheral skin damage, but this damage is short lived.

Remote sensing of Southern Ocean sea surface temperature: implications for marine biophysical models

Nineteen years of Advanced Very High Resolution Radiometer Multi-Channel Sea Surface Temperature (AVHRR MCSST) data were used to calculate monthly averages of sea surface temperatures (SSTs) for a large region of the Southern Ocean centred on Macquarie Island. Between October and February, the MCSST data were a reliable source of SSTs north of 60°S, but their quality (i.e. spatial and temporal density) was degraded severely at higher latitudes. Between April and August, the interpolated MCSST data were found to be unreliable in the study area, even at latitudes as low as 45°S. A specific problem has been identified to the west of Tasmania in the winter interpolated climatology which is likely an artefact of interpolation of sparse data. The poor coverage of valid MCSSTs during this time limits the datas application for many uses. Although the limitations of the MCSST data set are well recognised, the possibilities for ecological applications of these data remain largely unexplored and unquantified. The potential exists to maximize the information available from this and other similar data sets by determining the appropriate spatial and temporal scales at which they are best applied to biophysical models.

Under ice habitats for Antarctic krill larvae: could less mean more under climate warming?

Overwintering of larvae underneath Antarctic pack ice is a critical stage in the life cycle of Antarctic krill. However, there are no circumpolar assessments of available habitat for larval krill, making it difficult to evaluate how climate change may impact this life stage. We use outputs from a circumpolar sea ice model, together with a set of simple assumptions regarding key habitat features, to identify possible regions of larval krill habitat around Antarctica during winter. We assume that the location and suitability of habitat is determined by both food availability and three-dimensional complexity of the sea ice. A comparison of the combined area of these regions under current conditions with a warm climate scenario indicates that while total areal sea ice extent decreases, there is a consistently larger area of potential larval krill habitat under warm conditions. These findings suggest that decreases in sea ice extent may not necessarily be detrimental for krill populations.

The winter pack-ice zone provides a sheltered but food-poor habitat for larval Antarctic krill

A dominant Antarctic ecological paradigm suggests that winter sea ice is generally the main feeding ground for krill larvae. Observations from our winter cruise to the southwest Atlantic sector of the Southern Ocean contradict this view and present the first evidence that the pack-ice zone is a food-poor habitat for larval development. In contrast, the more open marginal ice zone provides a more favourable food environment for high larval krill growth rates. We found that complex under-ice habitats are, however, vital for larval krill when water column productivity is limited by light, by providing structures that offer protection from predators and to collect organic material released from the ice. The larvae feed on this sparse ice-associated food during the day. After sunset, they migrate into the water below the ice (upper 20 m) and drift away from the ice areas where they have previously fed. Model analyses indicate that this behaviour increases both food uptake in a patchy food environment and the likelihood of overwinter transport to areas where feeding conditions are more favourable in spring.Winter sea ice is thought to provide critical grazing habitat for overwintering Antarctic krill. In contrast, here the authors show that the pack-ice zone is a food-poor habitat, but does serve as an important sheltering ground for developing larvae.

Abundance and richness of key Antarctic seafloor fauna correlates with modelled food availability

Most seafloor communities at depths below the photosynthesis zone rely on food that sinks through the water column. However, the nature and strength of this pelagic–benthic coupling and its influence on the structure and diversity of seafloor communities is unclear, especially around Antarctica where ecological data are sparse. Here we show that the strength of pelagic–benthic coupling along the East Antarctic shelf depends on both physical processes and the types of benthic organisms considered. In an approach based on modelling food availability, we combine remotely sensed sea-surface chlorophyll-a, a regional ocean model and diatom abundances from sediment grabs with particle tracking and show that fluctuating seabed currents are crucial in the redistribution of surface productivity at the seafloor. The estimated availability of suspended food near the seafloor correlates strongly with the abundance of benthic suspension feeders, while the deposition of food particles correlates with decreasing suspension feeder richness and more abundant deposit feeders. The modelling framework, which can be modified for other regions, has broad applications in conservation and management, as it enables spatial predictions of key components of seafloor biodiversity over vast regions around Antarctica.Combining data on sea-surface chlorophyll-a with a regional ocean model and diatom abundance from sediment grabs, the authors determine the strength of pelagic–benthic coupling across the George V region in East Antarctica.