Relationship between long-term environmental fluctuations and diving effort of female Australian fur seals

For predators foraging within spatially and temporally heterogeneous marine ecosystems, environmental fluctuations can alter prey availability. Using the proportion of time spent diving and foraging trip duration as proxies of foraging effort, a multi-year dataset was used to assess the response of 58 female Australian fur seals Arctocephalus pusillus doriferus to interannual environmental fluctuations. Multiple environmental indices (remotely sensed ocean colour data and numerical weather predictions) were assessed for their influence on inter-annual variations in the proportion of time spent diving and trip duration. Model averaging revealed strong evidence for relationships between 4 indices and the proportion of time spent diving. There was a positive relationship with effort and 2 yr-lagged spring sea-surface temperature, current winter zonal wind and southern oscillation index, while a negative relationship was found with 2 yr-lagged spring zonal wind. Additionally, a positive relationship was found between foraging trip duration and 1 yr-lagged spring surface chlorophyll a. These results suggest that environmental fluctuations may influence prey availability by affecting the survival and recruitment of prey at the larval and post-larval phases while also affecting current distribution of adult prey.

Identification of prey captures in Australian Fur Seals (Arctocephalus pusillus doriferus) using head-mounted accelerometers: field validation with animal-borne video cameras

This study investigated prey captures in free-ranging adult female Australian fur seals (Arctocephalus pusillus doriferus) using head-mounted 3-axis accelerometers and animal-borne video cameras. Acceleration data was used to identify individual attempted prey captures (APC), and video data were used to independently verify APC and prey types. Results demonstrated that head-mounted accelerometers could detect individual APC but were unable to distinguish among prey types (fish, cephalopod, stingray) or between successful captures and unsuccessful capture attempts. Mean detection rate (true positive rate) on individual animals in the testing subset ranged from 67-100%, and mean detection on the testing subset averaged across 4 animals ranged from 82-97%. Mean False positive (FP) rate ranged from 15-67% individually in the testing subset, and 26-59% averaged across 4 animals. Surge and sway had significantly greater detection rates, but also conversely greater FP rates compared to heave. Video data also indicated that some head movements recorded by the accelerometers were unrelated to APC and that a peak in acceleration variance did not always equate to an individual prey item. The results of the present study indicate that head-mounted accelerometers provide a complementary tool for investigating foraging behaviour in pinnipeds, but that detection and FP correction factors need to be applied for reliable field application.

Use of anthropogenic sea floor structures by Australian fur seals: potential positive ecological impacts of marine industrial development?

Human-induced changes to habitats can have deleterious effects on many species that occupy them. However, some species can adapt and even benefit from such modifications. Artificial reefs have long been used to provide habitat for invertebrate communities and promote local fish populations. With the increasing demand for energy resources within ocean systems, there has been an expansion of infrastructure in near-shore benthic environments which function as de facto artificial reefs. Little is known of their use by marine mammals. In this study, the influence of anthropogenic sea floor structures (pipelines, cable routes, wells and shipwrecks) on the foraging locations of 36 adult female Australian fur seals (Arctocephalus pusillus doriferus) was investigated. For 9 (25%) of the individuals, distance to anthropogenic sea floor structures was the most important factor in determining the location of intensive foraging activity. Whereas the influence of anthropogenic sea floor structures on foraging locations was not related to age and mass, it was positively related to flipper length/standard length (a factor which can affect manoeuvrability). A total of 26 (72%) individuals tracked with GPS were recorded spending time in the vicinity of structures (from <1% to >75% of the foraging trip duration) with pipelines and cable routes being the most frequented. No relationships were found between the amount of time spent frequenting anthropogenic structures and individual characteristics. More than a third (35%) of animals foraging near anthropogenic sea floor structures visited more than one type of structure. These results further highlight potentially beneficial ecological outcomes of marine industrial development.

Influence of intrinsic variation on foraging behaviour of adult female Australian fur seals

Phenotypic variation and individual experience can create behavioural and/or dietary variation within a population. This may reduce intra-specific competition, creating a buffer to environmental change. This study examined how intrinsic variation affects foraging behaviour of Australian fur seals. Foraging movements of 29 female Australian fur seals were recorded using FastLoc GPS and dive behaviour recorders. For each individual, body mass, flipper length and axis length were recorded, a tooth was sampled to determine age, and milk was collected for diet analysis. Clustering of fatty acid dietary analysis revealed 5 distinct groups in the population. Behaviour was described using 19 indices, which were then reduced to 7 principal components (>80% of the behavioural variation). Bayesian mixed effect models were developed to describe the relationship between these components and intrinsic variation. No association was found between diet and age or body shape; however, age had a negative relationship with component 1 (27% of variation). Older females spent less time at-sea and foraged nearer to the colony. Age had an effect on component 5 (7% of variation), which represented haul-outs and dive depth; older females made fewer visits to haul-out sites and dived deeper to the benthos. This suggests that as animals age they are able to utilise prior knowledge to exploit nearby foraging sites that younger animals are either unaware of, or have yet to gain the experience required to efficiently utilise. Mass had a negative effect on components representing the directedness of a foraging trip, suggesting heavier individuals were more likely to travel directly to a foraging site.

Determinants of individual foraging specialization in large marine vertebrates, the Antarctic and subantarctic fur seals

The degree of individual specialization in resource use differs widely among wild populations where individuals range from fully generalized to highly specialized. This interindividual variation has profound implications in many ecological and evolutionary processes. A recent review proposed four main ecological causes of individual specialization: interspecific and intraspecific competition, ecological opportunity and predation. Using the isotopic signature of subsampled whiskers, we investigated to what degree three of these factors (interspecific and intraspecific competition and ecological opportunity) affect the population niche width and the level of individual foraging specialization in two fur seal species, the Antarctic and subantarctic fur seals (Arctocephalus gazella and Arctocephalus tropicalis), over several years. Population niche width was greater when the two seal species bred in allopatry (low interspecific competition) than in sympatry or when seals bred in high-density stabilized colonies (high intraspecific competition). In agreement with the niche variation hypothesis (NVH), higher population niche width was associated with higher interindividual niche variation. However, in contrast to the NVH, all Antarctic females increased their niche width during the interbreeding period when they had potential access to a wider diversity of foraging grounds and associated prey (high ecological opportunities), suggesting they all dispersed to a similar productive area. The degree of individual specialization varied among populations and within the annual cycle. Highest levels of interindividual variation were found in a context of lower interspecific or higher intraspecific competition. Contrasted results were found concerning the effect of ecological opportunity. Depending on seal species, females exhibited either a greater or lower degree of individual specialization during the interbreeding period, reflecting species-specific biological constraints during that period. These results suggest a significant impact of ecological interactions on the population niche width and degree of individual specialization. Such variation at the individual level may be an important factor in the species plasticity with significant consequences on how it may respond to environmental variability.

From video recordings to whisker stable isotopes: a critical evaluation of timescale in assessing individual foraging specialisation in Australian fur seals

Estimating the degree of individual specialisation is likely to be sensitive to the methods used, as they record individuals' resource use over different time-periods. We combined animal-borne video cameras, GPS/TDR loggers and stable isotope values of plasma, red cells and sub-sampled whiskers to investigate individual foraging specialisation in female Australian fur seals (Arctocephalus pusillus doriferus) over various timescales. Combining these methods enabled us to (1) provide quantitative information on individuals' diet, allowing the identification of prey, (2) infer the temporal consistency of individual specialisation, and (3) assess how different methods and timescales affect our estimation of the degree of specialisation. Short-term inter-individual variation in diet was observed in the video data (mean pairwise overlap = 0.60), with the sampled population being composed of both generalist and specialist individuals (nested network). However, the brevity of the temporal window is likely to artificially increase the level of specialisation by not recording the entire diet of seals. Indeed, the correlation in isotopic values was tighter between the red cells and whiskers (mid- to long-term foraging ecology) than between plasma and red cells (short- to mid-term) (R (2) = 0.93-0.73 vs. 0.55-0.41). δ(13)C and δ(15)N values of whiskers confirmed the temporal consistency of individual specialisation. Variation in isotopic niche was consistent across seasons and years, indicating long-term habitat (WIC/TNW = 0.28) and dietary (WIC/TNW = 0.39) specialisation. The results also highlight time-averaging issues (under-estimation of the degree of specialisation) when calculating individual specialisation indices over long time-periods, so that no single timescale may provide a complete and accurate picture, emphasising the benefits of using complementary methods.

Utilisation of intensive foraging zones by female Australian fur seals

Within a heterogeneous environment, animals must efficiently locate and utilise foraging patches. One way animals can achieve this is by increasing residency times in areas where foraging success is highest (area-restricted search). For air-breathing diving predators, increased patch residency times can be achieved by altering both surface movements and diving patterns. The current study aimed to spatially identify the areas where female Australian fur seals allocated the most foraging effort, while simultaneously determining the behavioural changes that occur when they increase their foraging intensity. To achieve this, foraging behaviour was successfully recorded with a FastLoc GPS logger and dive behaviour recorder from 29 individual females provisioning pups. Females travelled an average of 118 ± 50 km from their colony during foraging trips that lasted 7.3 ± 3.4 days. Comparison of two methods for calculating foraging intensity (first-passage time and first-passage time modified to include diving behaviour) determined that, due to extended surface intervals where individuals did not travel, inclusion of diving behaviour into foraging analyses was important for this species. Foraging intensity 'hot spots' were found to exist in a mosaic of patches within the Bass Basin, primarily to the south-west of the colony. However, the composition of benthic habitat being targeted remains unclear. When increasing their foraging intensity, individuals tended to perform dives around 148 s or greater, with descent/ascent rates of approximately 1.9 m•s-1 or greater and reduced postdive durations. This suggests individuals were maximising their time within the benthic foraging zone. Furthermore, individuals increased tortuosity and decreased travel speeds while at the surface to maximise their time within a foraging location. These results suggest Australian fur seals will modify both surface movements and diving behaviour to maximise their time within a foraging patch.

A colostrum trypsin inhibitor gene expressed in the Cape fur seal mammary gland during lactation

The colostrum trypsin inhibitor (CTI) gene and transcript were cloned from the Cape fur seal mammary gland and CTI identified by in silico analysis of the Pacific walrus and polar bear genomes (Order Carnivora), and in marine and terrestrial mammals of the Orders Cetartiodactyla (yak, whales, camel) and Perissodactyla (white rhinoceros). Unexpectedly, Weddell seal CTI was predicted to be a pseudogene. Cape fur seal CTI was expressed in the mammary gland of a pregnant multiparous seal, but not in a seal in its first pregnancy. While bovine CTI is expressed for 24-48h postpartum (pp) and secreted in colostrum only, Cape fur seal CTI was detected for at least 2-3months pp while the mother was suckling its young on-shore. Furthermore, CTI was expressed in the mammary gland of only one of the lactating seals that was foraging at-sea. The expression of β-casein (CSN2) and β-lactoglobulin II (LGB2), but not CTI in the second lactating seal foraging at-sea suggested that CTI may be intermittently expressed during lactation. Cape fur seal and walrus CTI encode putative small, secreted, N-glycosylated proteins with a single Kunitz/bovine pancreatic trypsin inhibitor (BPTI) domain indicative of serine protease inhibition. Mature Cape fur seal CTI shares 92% sequence identity with Pacific walrus CTI, but only 35% identity with BPTI. Structural homology modelling of Cape fur seal CTI and Pacific walrus trypsin based on the model of the second Kunitz domain of human tissue factor pathway inhibitor (TFPI) and porcine trypsin (Protein Data Bank: 1TFX) confirmed that CTI inhibits trypsin in a canonical fashion. Therefore, pinniped CTI may be critical for preventing the proteolytic degradation of immunoglobulins that are passively transferred from mother to young via colostrum and milk.