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.

New insights into the cardiorespiratory physiology of weaned southern elephant seals (Mirounga leonina)

Southern elephant seal (Mirounga leonina) pups must strike a balance between conserving energy during their post-weaning fast and simultaneously developing diving abilities to attain nutritional independence. Little is known about environmental influences on cardiorespiratory patterns, hence energy use, throughout the 6 week fast. Continuous heart rates were recorded for free-ranging, newly weaned southern elephant seals using heart rate time-depth recorders for 5-9 days at Sub-Antarctic Macquarie Island, during October 1994 (n = 1), 1995 (n = 4) and 1996 (n = 1). Daytime observations of respiration and behaviour were made throughout. We present the first instance of synchronous heart rate traces recorded simultaneously for individual weaners. Generalized additive models revealed that a sinusoidal pattern of diurnal heart rate elevation and nocturnal depression was evident in all seals and, on at least one occasion, a conspicuous break in this pattern coincided with an extreme cold weather event. Seals in this study were capable of considerable cardiorespiratory control and regularly demonstrated bradycardia during periods of resting apnoea. Apnoeic duration ranged from 33 to 291 s (mean 134 s). Apnoeic heart rates (mean 67 ± 15 beats min(-1), range 40-114 beats min(-1)) were on average 19.7% lower than those exhibited during periods of eupnoea (mean 83 ± 15 beats min(-1), range 44-124 beats min(-1)). The early development of the cardiorespiratory response is characterized by arrhythmic heart and respiration rates. The strong temporal patterns observed are being driven by the opposing requirements of maximizing time spent fasting in order to develop diving capabilities and of maximizing departure mass. This pilot study has highlighted a potentially large effect of ambient weather conditions on newly weaned southern elephant seal cardiorespiratory activity. Given the increasing westerlies and more erratic and increasing storminess associated with the Southern Annular Mode predicted in the Southern Ocean, the patterns observed here warrant further investigation.

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.

Dive characteristics can predict foraging success in Australian fur seals (arctocephalus pusillus doriferus) as validated by animal-borne video

Dive characteristics and dive shape are often used to infer foraging success in pinnipeds. However, these inferences have not been directly validated in the field with video, and it remains unclear if this method can be applied to benthic foraging animals. This study assessed the ability of dive characteristics from time-depth recorders (TDR) to predict attempted prey capture events (APC) that were directly observed on animal-borne video in Australian fur seals (Arctocephalus pusillus doriferus, n=11). The most parsimonious model predicting the probability of a dive with ≥1 APC on video included only descent rate as a predictor variable. The majority (94%) of the 389 total APC were successful, and the majority of the dives (68%) contained at least one successful APC. The best model predicting these successful dives included descent rate as a predictor. Comparisons of the TDR model predictions to video yielded a maximum accuracy of 77.5% in classifying dives as either APC or non-APC or 77.1% in classifying dives as successful verses unsuccessful. Foraging intensity, measured as either total APC per dive or total successful APC per dive, was best predicted by bottom duration and ascent rate. The accuracy in predicting total APC per dive varied based on the number of APC per dive with maximum accuracy occurring at 1 APC for both total (54%) and only successful APC (52%). Results from this study linking verified foraging dives to dive characteristics potentially opens the door to decades of historical TDR datasets across several otariid species.

Dive behaviour, foraging locations and maternal-attendance patterns in Australian fur seals (Arctocephalus pusillus doriferus)

The dive behaviour, foraging locations, and colony-attendance patterns of female Australian fur seals (Arctocephalus pusillus doriferus) from Kanowna Island (39°10'S, 146°18'E) in Bass Strait, southeastern Australia, were determined throughout lactation during 1997–1999. Foraging-trip durations increased as lactation progressed, being shortest in summer (3.71 ± 0.24 days; mean ± 1 SE) and longest in winter (6.77 ± 0.57 days, P < 0.05), but maternal-attendance periods did not differ in duration (1.70 ± 0.10 days, P > 0.5). Individual mean attendance periods and trip durations were positively correlated (r² = 0.21, P < 0.005). Diving commenced shortly after seals left the colony (2.6 ± 0.4 h), was continuous for long periods (up to 36 h), occurred mostly during daylight hours, and lacked regular diel variation in depth. The majority of dives (78%) were typically U-shaped and reached depths corresponding to the prevailing depths in Bass Strait (65–85 m), indicating that these animals forage mostly on the benthos of the shallow continental shelf in this region. Such behaviour is unusual for fur seals but is reminiscent of that of some sea lion species. Mean dive durations varied between 2.0 and 3.7 min (maximum 8.9 min) and the theoretical aerobic dive limit (3.91–4.26 min) was exceeded on 17.3% of dives. Dive frequency (8.3 ± 0.6/h) and the proportion of time at sea spent diving (40.7 ± 2.1%) were weakly negatively related to the duration of the foraging trip (r²= 0.07, P < 0.004, and r² = 0.13, P < 0.0001, respectively). Data from at-sea locations showed that lactating females forage almost exclusively within Bass Strait during all seasons.

Early-life sexual segregation: ontogeny of isotopic niche differentiation in the Antarctic fur seal

Investigating the ontogeny of niche differentiation enables to determine at which life-stages sexual segregation arises, providing insights into the main factors driving resource partitioning. We investigated the ontogeny of foraging ecology in Antarctic fur seals (Arctocephalus gazella), a highly dimorphic species with contrasting breeding strategies between sexes. Sequential δ(13)C and δ(15)N values of whiskers provided a longitudinal proxy of the foraging niche throughout the whole life of seals, from weaning, when size dimorphism is minimal to the age of 5. Females exhibited an early-life ontogenetic shift, from a total segregation during their first year at-sea, to a similar isotopic niche as breeding females as early as age 2. In contrast, males showed a progressive change in isotopic niche throughout their development such that 5-year-old males did not share the same niche as territorial bulls. Interestingly, males and females segregated straight after weaning with males appearing to feed in more southerly habitats than females. This spatial segregation was of similar amplitude as observed in breeding adults and was maintained throughout development. Such early-life niche differentiation is an unusual pattern and indicates size dimorphism and breeding constraints do not directly drive sexual segregation contrary to what has been assumed in otariid seals.

Flipper strokes can predict energy expenditure and locomotion costs in free-ranging northern and Antarctic fur seals

Flipper strokes have been proposed as proxies to estimate the energy expended by marine vertebrates while foraging at sea, but this has never been validated on free-ranging otariids (fur seals and sea lions). Our goal was to investigate how well flipper strokes correlate with energy expenditure in 33 foraging northern and Antarctic fur seals equipped with accelerometers, GPS, and time-depth recorders. We concomitantly measured field metabolic rates with the doubly-labelled water method and derived activity-specific energy expenditures using fine-scale time-activity budgets for each seal. Flipper strokes were detected while diving or surface transiting using dynamic acceleration. Despite some inter-species differences in flipper stroke dynamics or frequencies, both species of fur seals spent 3.79 ± 0.39 J/kg per stroke and had a cost of transport of ~1.6-1.9 J/kg/m while diving. Also, flipper stroke counts were good predictors of energy spent while diving (R(2) = 0.76) and to a lesser extent while transiting (R(2) = 0.63). However, flipper stroke count was a poor predictor overall of total energy spent during a full foraging trip (R(2) = 0.50). Amplitude of flipper strokes (i.e., acceleration amplitude × number of strokes) predicted total energy expenditure (R(2) = 0.63) better than flipper stroke counts, but was not as accurate as other acceleration-based proxies, i.e. Overall Dynamic Body Acceleration.

Terrestrial apnoeas and the development of cardiac control in Australian fur seal (Arctocephalus pusillus doriferus) pups

The development of cardiac control in association with terrestrial respiration patterns was examined throughout the period of maternal dependence in Australian fur seal pups. Resting eupnoic heart rate and respiration rate were significantly correlated (r² = 0.49) and both decreased with age (P < 0.05 in both cases). From an early age (1 month), pups displayed terrestrial apnoeas (18.1 ± 0.5 s) accompanied by substantial bradycardia (127 beats min^-1, a 13% decrease from eupnoic HR). Terrestrial apnoea duration increased significantly with age reaching a mean of 41 s just prior to weaning, slightly lower than the mean dive duration (52 s) previously recorded for pups of the same age. Correspondingly, mean apnoic heart rate decreased with age to 74 beats min^-1 just prior to weaning, representing a 25% decrease on eupnoic heart rate. Importantly, concomitant with the decrease in mean apnoic heart rate with age, an increase in the control
of bradycardia was evident with the variability in instantaneous apnoic heart decreasing such that older pups were able to maintain a low steady heart rate for the duration of the apnoea. The changes seen in these parameters are similar to those reported during postnatal development in elephant seals (Mirounga spp.) and harbour seals (Phoca vitulina), and are considered indicative of the development of cardiac control. These findings suggest a common strategy for the development of bradycardia control in both otariid and phocid seals.

Accuracy of ARGOS locations of pinnipeds at-sea estimated using Fastloc GPS

Background: ARGOS satellite telemetry is one of the most widely used methods to track the movements of free-ranging marine and terrestrial animals and is fundamental to studies of foraging ecology, migratory behavior and habitat-use. ARGOS location estimates do not include complete error estimations, and for many marine organisms, the most commonly acquired locations (Location Class 0, A, B, or Z) are provided with no declared error estimate.
Methodology/Principal Findings: We compared the accuracy of ARGOS locations to those obtained using Fastloc GPS from the same electronic tags on five species of pinnipeds: 9 California sea lions (Zalophus californianus), 4 Galapagos sea lions (Zalophus wollebaeki), 6 Cape fur seals (Arctocephalus pusillus pusillus), 3 Australian fur seals (A. p. doriferus) and 5 northern elephant seals (Mirounga angustirostris). These species encompass a range of marine habitats (highly pelagic vs coastal), diving behaviors (mean dive durations 2–21 min) and range of latitudes (equator to temperate). A total of 7,318 ARGOS positions and 27,046 GPS positions were collected. Of these, 1,105 ARGOS positions were obtained within five minutes of a GPS position and were used for comparison. The 68^th percentile ARGOS location errors as measured in this study were LC-3
0.49 km, LC-2 1.01 km, LC-1 1.20 km, LC-0 4.18 km, LC-A 6.19 km, LC-B 10.28 km.
Conclusions/Significance: The ARGOS errors measured here are greater than those provided by ARGOS, but within the range of other studies. The error was non-normally distributed with each LC highly right-skewed. Locations of species that make short duration dives and spend extended periods on the surface (sea lions and fur seals) had less error than species like elephant seals that spend more time underwater and have shorter surface intervals. Supplemental data (S1) are provided allowing the creation of density distributions that can be used in a variety of filtering algorithms to improve the quality of ARGOS tracking data.

Morphometric and molecular characterization of the species of Uncinaria Frölich, 1789 (Nematoda) parasitic in the Australian fur seal Arctocephalus pusillus doriferus (Schreber), with notes on hookworms in three other pinniped hosts

This study presents morphological and molecular data on hookworms from the Australian fur seal Arctocephalus pusillus doriferus (Schreber) currently identified in Australian waters as Uncinaria hamiltoni Baylis, 1933. Additional specimens from the Australian sea lion Neophoca cinerea (Péron) and the New Zealand fur seal Arctocephalus forsteri (Lesson) from Australia, and the Southern elephant seal Mirounga leonina (Linnaeus) from Antarctica, were included. Using the internal transcribed spacer (ITS), hookworms from A. p. doriferus, N. cinerea and A. forsteri were found to be genetically similar but distinct from Uncinaria spp. found in M. leonina from Antarctica, as well as from Zalophus californianus (Lesson) and Callorhinus ursinus (Linnaeus) from California. Few morphological differences were detected between these taxa.

Interactions between fur seals and a squid jig fishery in southern Australia

Interactions between Australian fur seals (Arctocephalus pusillus doriferus) and the Southern Squid Jig Fishery (SSJF) were investigated in April–May 2002. Details on the number of seals present, distance from the vessel, age and gender, and their behaviour were recorded using scan sampling techniques over 26 nights from eight vessels operating out of Portland and Port Fairy on the southwest coast of Victoria. Seals were observed on all nights but none were recorded in 30% of all (777) scan observations. Of the seals attending vessels at any one time (1.89 ± 0.24), 67% were involved in activities unrelated to jigging operations with the most common behaviour category being resting/grooming. Only 3.6% of observations involved seals targeting squid caught on jig lures whereas a further 29% were of foraging on squid within 40 m of the vessel. Damage to fishing gear attributable to seals was recorded on only three occasions. There was no evidence of negative impacts on seals from vessel operations. The majority of seals foraging on squid around vessels were adult females (71%) with the remainder being almost exclusively juvenile males. The current level of interactions between Australian fur seals and vessels in the SSJF appears minor.

Peopling of the Northern Territory part 1: a white elephant in a white Australia? The Northern Territory, 1901-1920

As a result of negative net immigration during the 1890s depression, Australians at the time of Federation were preoccupied about the slow rate of growth of the population. The non-Aboriginal population at the end of 1904 was approximately three and three-quarter million. and the publication in March 1904 of the Report of the New South Wales Royal Commission on the Decline of the Birth Rate and the Mortality of Infants did nothing to allay these concerns. Despite the perceived need for more people. the desire for racial unity was paramount. The main goals of policy makers at the time were to preserve a 'white' and essentially British Australia and create an imperial bastion in the Southern Hemisphere.

Effect of proximity to the shelf edge on the diet of female Australian fur seals

The Australian fur seal Arctocephalus pusillus doriferus is a temperate latitude species with a breeding distribution restricted to Bass Strait, Australia. Recent studies of the foraging behaviour of female Australian fur seals indicated that they feed demersally in the shallow continental shelf waters, a behaviour that is in contrast to the epipelagic foraging of females of most other arctocephaline (Southern Hemisphere fur seals) species and akin to that observed in sea lions. These studies, however, were conducted at one colony (Kanowna Island) located in central northern Bass Strait, and it was suggested that the observed foraging behaviour may have been due to the distance of this colony from the continental shelf edge (180 km), making it inefficient to forage beyond it. Here, the diet of lactating Australian fur seals was compared between 2 colonies to test if differing proximity to the continental shelf edge resulted in differences in foraging behaviour. The 2 breeding colonies studied, Kanowna Island and The Skerries, were 180 and 25 km from the nearest shelf edge, respectively. We analysed a total of 917 scat samples collected at the 2 colonies between 1997 and 2001. From faecal analysis, 45 primarily demersal on-shelf species of fishes and cephalopods were identified. Only 4 species had a frequency of occurrence greater than 10%: redbait Emmelichthys nitidus, jack mackerel Trachurus sp., red rock cod Pseudophycis bachus, and Gould’s squid Nototodarus gouldi. No seasonal, annual or spatial differences were found between the 2 colonies, indicating that proximity to the shelf edge does not influence diet.