Colin Southwell

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.

Adelie penguin population diet monitoring by analysis of food DNA in scats.

The Adélie penguin is the most important animal currently used for ecosystem monitoring in the Southern Ocean. The diet of this species is generally studied by visual analysis of stomach contents; or ratios of isotopes of carbon and nitrogen incorporated into the penguin from its food. There are significant limitations to the information that can be gained from these methods. We evaluated population diet assessment by analysis of food DNA in scats as an alternative method for ecosystem monitoring with Adélie penguins as an indicator species. Scats were collected at four locations, three phases of the breeding cycle, and in four different years. A novel molecular diet assay and bioinformatics pipeline based on nuclear small subunit ribosomal RNA gene (SSU rDNA) sequencing was used to identify prey DNA in 389 scats. Analysis of the twelve population sample sets identified spatial and temporal dietary change in Adélie penguin population diet. Prey diversity was found to be greater than previously thought. Krill, fish, copepods and amphipods were the most important food groups, in general agreement with other Adélie penguin dietary studies based on hard part or stable isotope analysis. However, our DNA analysis estimated that a substantial portion of the diet was gelatinous groups such as jellyfish and comb jellies. A range of other prey not previously identified in the diet of this species were also discovered. The diverse prey identified by this DNA-based scat analysis confirms that the generalist feeding of Adélie penguins makes them a useful indicator species for prey community composition in the coastal zone of the Southern Ocean. Scat collection is a simple and non-invasive field sampling method that allows DNA-based estimation of prey community differences at many temporal and spatial scales and provides significant advantages over alternative diet analysis approaches.

SEA ICE COVER AND ITS INFLUENCE ON ADÉLIE PENGUIN REPRODUCTIVE PERFORMANCE

The relationship between Adélie Penguins (Pygoscelis adeliae) and ice is well established, with sea ice influencing penguin populations through a variety of processes operating at different spatial and temporal scales. To further explain the relationship between sea ice and Adélie Penguin reproductive performance, we investigated the relative importance of various measures of sea ice cover on breeding success at Béchervaise Island, East Antarctica. Our results show a clear distinction in the response of penguins to different types of ice, as well as to the timing of the presence of sea ice. Nearshore sea ice, which is composed primarily of fast ice during the guard stage of the breeding season, had an overwhelmingly strong and negative impact on penguin reproductive performance. The influence of winter and offshore guard-stage ice was only evident in conjunction with nearshore ice. Predicting Adélie Penguin population growth in relation to changes in the sea ice environment may be complicated because penguin-ice interactions vary according to the type of sea ice present, the season in which it is present, and the processes contributing to population growth that are influenced by sea ice.

The influence of animal mobility on the assumption of uniform distances in aerial line-transect surveys

Line-transect distance sampling is a widely used method for estimating animal density from aerial surveys. Analysis of line-transect distance data usually relies on a requirement that the statistical distribution of distances of animal groups from the transect line is uniform. We show that this requirement is satisfied by the survey design if all other assumptions of distance sampling hold, but it can be violated by consistent survey problems such as responsive movement of the animals towards or away from the observer. We hypothesise that problems with the uniform requirement are unlikely to be encountered for immobile taxa, but might become substantial for species of high mobility. We test evidence for non-uniformity using double-observer distance data from two aerial surveys of five species with a spectrum of mobility capabilities and tendencies. No clear evidence against uniformity was found for crabeater seals or emperor penguins on the pack-ice in East Antarctica, while minor non-uniformity consistent with responsive movement up to 30 m was found for Adelie penguins. Strong evidence of either non-uniformity or a failure of the capture-recapture validating method was found for eastern grey kangaroos and red kangaroos in Queensland.

Adélie penguin survival: age structure, temporal variability and environmental influences

The driving factors of survival, a key demographic process, have been particularly challenging to study, especially for winter migratory species such as the Adélie penguin (Pygoscelis adeliae). While winter environmental conditions clearly influence Antarctic seabird survival, it has been unclear to which environmental features they are most likely to respond. Here, we examine the influence of environmental fluctuations, broad climatic conditions and the success of the breeding season prior to winter on annual survival of an Adélie penguin population using mark–recapture models based on penguin tag and resight data over a 16-year period. This analysis required an extension to the basic Cormack–Jolly–Seber model by incorporating age structure in recapture and survival sub-models. By including model covariates, we show that survival of older penguins is primarily related to the amount and concentration of ice present in their winter foraging grounds. In contrast, fledgling and yearling survival depended on other factors in addition to the physical marine environment and outcomes of the previous breeding season, but we were unable to determine what these were. The relationship between sea-ice and survival differed with penguin age: extensive ice during the return journey to breeding colonies was detrimental to survival for the younger penguins, whereas either too little or too much ice (between 15 and 80% cover) in the winter foraging grounds was detrimental for adults. Our results demonstrate that predictions of Adélie penguin survival can be improved by taking into account penguin age, prior breeding conditions and environmental features.

Density Can Be Misleading for Low-Density Species: Benefits of Passive Acoustic Monitoring

Climate-induced changes may be more substantial within the marine environment, where following ecological change is logistically difficult, and typically expensive. As marine animals tend to produce stereotyped, long-range signals, they are ideal for repeatable surveying. In this study we illustrate the potential for calling rates to be used as a tool for determining habitat quality by using an Antarctic pack-ice seal, the leopard seal, as a model.With an understanding of the vocal behavior of a species, their seasonal and diurnal patterns, sex and age-related differences, an underwater passive-acoustic survey conducted alongside a visual survey in an arc of 4,225 km across the Davis Sea, Eastern Antarctica, showed that while acoustic and visual surveys identified similar regions as having high densities, the acoustic surveys surprisingly identified the opposite regions as being ‘critical’ habitats. Density surveys of species that cannot be differentiated into population classes may be misleading because overall density can be a negative indicator of habitat quality.Under special circumstances acoustics can offer enormous advantage over traditional techniques and open up monitoring to regions that are remote, difficult and expensive to work within, no longer restricting long-term community assessment to resource-wealthy communities. As climatic change affects a broad range of organisms across geographic boundaries we propose that capitalizing on the significant advances in passive acoustic technology, alongside physical acoustics and population modeling, can help in addressing ecological questions more broadly.

Response behaviour of seals and penguins to helicopter surveys over the pack ice off East Antarctica

Data on the response behaviour of seals (crabeater, leopard and Ross) and penguins (Adélie and emperor) to helicopter surveys over the pack ice off East Antarctica are presented. The surveys involved Sikorsky S76 helicopters flying at altitude 130 m and speed 90 knots along straight-line transects. The relative frequency of alert and movement behaviours by seal and penguin groups decreased with distance out to 800 m from the flight path. Penguin groups were more likely to show a movement response than seals at all distances. The perpendicular distances moved relative to the flight path were small (maximum 20 m, mean ≤ 3 m, for both seal and penguin groups) relative to the width of the area searched (800 m), and there was no evidence that response movement resulted in a spiked detection histogram. Observers were more likely to feel confident in identifying seal and penguin species if the animals responded to the helicopter by changing the body posture or moving. In this application of aerial survey, the response behaviour elicited by the passing helicopter was considered to be beneficial from a technical perspective, and the disturbance caused to the surveyed populations negligible from an ethical perspective.

Optimising the timing of visual surveys of crabeater seal abundance : haulout behaviour as a consideration

The most practical means of estimating pack-ice seal abundance is by conducting visual surveys from ships and aircraft. However, only those seals hauled out on the ice are ‘available’ to such surveys, and additional information on haulout behaviour is required to adjust counts of seals on the ice to estimate the total population size. Consideration of the optimal time to undertake visual surveys with respect to availability is important to ensure that bias and uncertainty in the abundance estimate are minimised for a fixed survey effort. In order to assess the optimal time for conducting visual surveys of crabeater seals (Lobodon carcinophaga), satellite-linked dive recorders were attached to 24 adult seals in the pack-ice off east Antarctica to record haulout behaviour over a 4-month period from mid-September to mid-January. The optimal time for visual surveys within these four months was December to mid-January (after the pupping season) when a high, relatively constant proportion of seals were hauled out over a period of 6–7 h during daylight, and when variation in haulout behaviour between seals was low. Despite the necessity for breeding seals to haul out continuously for extended periods during the pupping season, this was not a preferred time for visual surveys because variability in haulout behaviour between breeding and non-breeding seals was high. The efficiency of surveys before pupping was limited by the relatively short time during daylight when both the proportion of seals hauled out was high and variability in haulout behaviour among seals was small.

New methods and technologies for regional-scale abundance estimation of land-breeding marine animals: application to Adelie penguin populations in East Antarctica

Land-breeding marine animals such as penguins, flying seabirds and pinnipeds are important components of marine ecosystems, and their abundance has been used extensively as an indication of ecosystem status and change. Until recently, many efforts to measure and monitor abundance of these species’ groups have focussed on smaller populations and spatial scales, and efforts to account for perception bias and availability bias have been variable and often ad hoc. We describe a suite of new methods, technologies and estimation procedures for cost-effective, large-scale abundance estimation within a general estimation framework and illustrate their application on large Adélie penguin populations in two regions of East Antarctica. The methods include photographic sample counts, automated cameras for collecting availability data, and bootstrap estimation to adjust counts for the sampling fraction, perception bias, and availability bias, and are applicable for a range of land-breeding marine species. The methods will improve our ability to obtain population data over large spatial and population scales within tight logistic, environmental and time constraints. This first application of the methods has given new insights into the biases and uncertainties in abundance estimation for penguins and other land-breeding marine species. We provide guidelines for applying the methods in future surveys.

Spatially Extensive Standardized Surveys Reveal Widespread, Multi-Decadal Increase in East Antarctic Adélie Penguin Populations

Seabirds are considered to be useful and practical indicators of the state of marine ecosystems because they integrate across changes in the lower trophic levels and the physical environment. Signals from this key group of species can indicate broad scale impacts or response to environmental change. Recent studies of penguin populations, the most commonly abundant Antarctic seabirds in the west Antarctic Peninsula and western Ross Sea, have demonstrated that physical changes in Antarctic marine environments have profound effects on biota at high trophic levels. Large populations of the circumpolar-breeding Adélie penguin occur in East Antarctica, but direct, standardized population data across much of this vast coastline have been more limited than in other Antarctic regions. We combine extensive new population survey data, new population estimation methods, and re-interpreted historical survey data to assess decadal-scale change in East Antarctic Adélie penguin breeding populations. We show that, in contrast to the west Antarctic Peninsula and western Ross Sea where breeding populations have decreased or shown variable trends over the last 30 years, East Antarctic regional populations have almost doubled in abundance since the 1980’s and have been increasing since the earliest counts in the 1960’s. The population changes are associated with five-year lagged changes in the physical environment, suggesting that the changing environment impacts primarily on the pre-breeding age classes. East Antarctic marine ecosystems have been subject to a number of changes over the last 50 years which may have influenced Adélie penguin population growth, including decadal-scale climate variation, an inferred mid-20th century sea-ice contraction, and early-to-mid 20th century exploitation of fish and whale populations.