Individual foraging site fidelity in lactating New Zealand fur seals : Continental shelf vs. oceanic habitats

Wide-ranging marine central place foragers often exhibit foraging site fidelity to oceanographic features over differing spatial scales (i.e., localized coastal upwellings and oceanic fronts). Few studies have tested how the degree of site fidelity to foraging areas varies in relation to the type of ocean features used. In order to determine how foraging site fidelity varied between continental shelf and oceanic foraging habitats, 31 lactating New Zealand fur seals (Arctocephalus australis forsteri1) were satellite tracked over consecutive foraging trips (14–108 d). Thirty-seven foraging trips were recorded from 11 females that foraged on the continental shelf, in a region associated with a coastal upwelling, while 65 foraging trips were recorded from 20 females that foraged in oceanic waters. There were no significant differences in the mean bearings (to maximum distance) of individual's consecutive foraging trips, suggesting individual fidelity to foraging areas. However, overlap in area and time spent in area varied considerably between continental shelf and oceanic foragers. Females that foraged on the continental shelf had significantly greater overlap in consecutive foraging trips when compared to females that foraged in oceanic waters (overlap in 5 × 5 km grid cells visited on consecutive trips 55.9%± 20.4% and 13.4%± 7.6%, respectively). Females that foraged on the continental shelf also spent significantly more time within the same grid cell than females that foraged in oceanic waters (maximum time spent in 5 × 5 km grid cells: 14%± 5% and 4%± 2%, respectively). This comparatively high foraging site fidelity may reflect the concentration of productivity associated with a coastal upwelling system, the Bonney Upwelling. Lower foraging site fidelity recorded by seals that foraged in oceanic waters implies a lower density/larger scale habitat, where prey are more dispersed or less predictable at fine scales, when compared to the continental shelf region.

Fur seals at Macquarie Island : post-sealing colonisation, trends in abundance and hybridisation of three species

Commercial sealers exterminated the original fur seal population at Macquarie Island in the early 1800s. The first breeding record since the sealing era was not reported until March 1955. Three species of fur seal now occur at Macquarie Island, the Antarctic (Arctocephalus gazella), subantarctic (A. tropicalis) and New Zealand (A. forsteri) fur seal. Census data from 54 breeding seasons in the period 1954–2007 were used to estimate population status and growth for each species. Between the 1950s and 1970s, annual increases in pup production for the species aggregate were low. Between 1986 and 2007, pup production of Antarctic fur seals increased by about 8.8% per year and subantarctic fur seals by 6.8% per year. The New Zealand fur seal, although the most numerous fur seal species on Macquarie Island, has yet to establish a breeding population, due to the absence of reproductively mature females. Hybridisation among species is significant, but appears to be declining. The slow establishment and growth of fur seal populations on Macquarie Island appears to have been affected by its distance from major population centres and hence low immigration rates, asynchronous colonisation times of males and females of each species, and extensive hybridisation.

Analysis of growth and stable isotopes in teeth of male Australian fur seals reveals interannual variability in prey resources

To detect and monitor long-term ecosystem responses to environmental variability, managers must utilize reliable and quantitative techniques to predict future ecosystem responses. Canine teeth from 67 male Australian fur seals (aged 2-19 yr), collected at Seal Rocks, between 1967 and 1976, were measured for relative growth within the dentine growth layer groups (GLGs), as an index of body growth. Fluctuations in relative growth were apparent during 1956-1971, suggesting interannual variation in prey resources within Bass Strait. These were positively correlated with the Southern Oscillation Index and negatively with the Indian Ocean Subtropical Dipole, both on a 2 yr lag. The observed delay may reflect the time required for the nutrient cascade to filter through to the predominantly benthic prey of Australian fur seals. Stable isotope analysis (?15N/?13C) was also used to investigate whether fluctuations in growth were associated with differences in diet. Relative growth was found to be negatively correlated with ?15N, suggesting years of greater resource availability may be associated with individuals consuming proportionally more prey biomass of lower isotopic value. This study demonstrates that fluctuations in the dentine GLGs of male Australian fur seals are related to environmental parameters, suggesting variation in body growth is mediated by changes in prey resources

Temporal allocation of foraging effort in female Australian fur seals (Arctocephalus pusillus doriferus)

Across an individual's life, foraging decisions will be affected by multiple intrinsic and extrinsic drivers that act at differing timescales. This study aimed to assess how female Australian fur seals allocated foraging effort and the behavioural changes used to achieve this at three temporal scales: within a day, across a foraging trip and across the final six months of the lactation period. Foraging effort peaked during daylight hours (57% of time diving) with lulls in activity just prior to and after daylight. Dive duration reduced across the day (196 s to 168 s) but this was compensated for by an increase in the vertical travel rate (1500–1600 m•h−1) and a reduction in postdive duration (111–90 s). This suggests physiological constraints (digestive costs) or prey availability may be limiting mean dive durations as a day progresses. During short trips (<2.9 d), effort remained steady at 55% of time diving, whereas, on long trips (>2.9 d) effort increased up to 2–3 d and then decreased. Dive duration decreased at the same rate in short and long trips, respectively, before stabilising (long trips) between 4–5 d. Suggesting that the same processes (digestive costs or prey availability) working at the daily scale may also be present across a trip. Across the lactation period, foraging effort, dive duration and vertical travel rate increased until August, before beginning to decrease. This suggests that as the nutritional demands of the suckling pup and developing foetus increase, female effort increases to accommodate this, providing insight into the potential constraints of maternal investment in this species

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.

Variation in body condition during the post-moult foraging trip of southern elephant seals and its consequences on diving behaviour

Mature female southern elephant seals (Mirounga leonina) come ashore only in October to breed and in January to moult, spending the rest of the year foraging at sea. Mature females may lose as much as 50% of their body mass, mostly in lipid stores, during the breeding season due to fasting and lactation. When departing to sea, post-breeding females are negatively buoyant, and the relative change in body condition (i.e. density) during the foraging trip has previously been assessed by monitoring the descent rate during drift dives. However, relatively few drift dives are performed, resulting in low resolution of the temporal reconstruction of body condition change. In this study, six post-breeding females were equipped with time-depth recorders and accelerometers to investigate whether changes in active swimming effort and speed could be used as an alternative method of monitoring density variations throughout the foraging trip. In addition, we assessed the consequences of density change on the swimming efforts of individuals while diving and investigated the effects on dive duration. Both descent swimming speed and ascent swimming effort were found to be strongly correlated to descent rate during drift dives, enabling the fine-scale monitoring of seal density change over the whole trip. Negatively buoyant seals minimized swimming effort during descents, gliding down at slower speeds, and reduced their ascent swimming effort to maintain a nearly constant swimming speed as their buoyancy increased. One per cent of seal density variation over time was found to induce a 20% variation in swimming effort during dives with direct consequences on dive duration.

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.

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.