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While sexual segregation is expected in highly dimorphic species, the local environment is a major factor driving the degree of resource partitioning within a population. Sexual and individual niche segregation was investigated in the Australian fur seal (Arctocephalus pusillus doriferus), which is a benthic foraging species restricted to the shallow continental shelf region of south-eastern Australia. Tracking data and the isotopic values of plasma, red blood cells and whiskers were combined to document spatial and dietary niche segregation throughout the year. Tracking data indicated that, in winter, males and females overlapped in their foraging habitat. All individuals stayed within central Bass Strait, relatively close (< 220 km) to the breeding colony. Accordingly, both genders exhibited similar plasma and red cell δ13C values. However, males exhibited greater δ13C intra-individual variation along the length of their whisker than females. This suggests that males exploited a greater diversity of foraging habitats throughout the year than their female counterparts, which are restricted in their foraging grounds by the need to regularly return to the breeding colony to suckle their pup. The degree of dietary sexual segregation was also surprisingly low, both sexes exhibiting a great overlap in their δ15N values. Yet, males displayed higher δ15N values than females, suggesting they fed upon a higher proportion of higher trophic level prey. Given that males and females exploit different resources (mainly foraging habitats), the degree of individual specialisation might differ between the sexes. Higher degrees of individual specialisation would be expected in males which exploit a greater range of resources. However, comparable levels of inter-individual variation in δ15N whisker values were found in the sampled males and females, and, surprisingly, all males exhibited similar seasonal and inter-annual variation in their δ13C whisker values, suggesting they all followed the same general dispersion pattern throughout the year.

The foraging strategy of many animals is thought to be determined by their past experiences. However, few empirical studies have investigated whether this is true in diving animals. We recorded three-dimensional movements and mouth-opening events from three Antarctic fur seals during their foraging trips to examine how they adapt their behaviour based on past experience--continuing to search for prey in the same area or moving to search in a different place. Each dive cycle was divided into a transit phase and a feeding phase. The linear horizontal distance travelled after feeding phases in each dive was affected by the mouth-opening rate during the previous 244 s, which typically covered two to three dive cycles. The linear distance travelled tended to be shorter when the mouth-opening rate in the previous 244 s was higher, i.e. seals tended to stay in the same areas with high prey-encounter rates. These results indicate that Antarctic fur seals follow decision-making strategies based on the past foraging experience over time periods longer than the immediately preceding dive., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)

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., (© 2015 The Authors. Journal of Animal Ecology © 2015 British Ecological Society.)

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.

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.

We developed models to predict foraging habitat of adult female northern fur seals (Callorhinus ursinus) using stable carbon (δ13C) and nitrogen (δ15N) isotope values from plasma and red blood cells. Binomial generalized linear mixed models were developed using blood isotope samples collected from 35 adult female fur seals on three breeding colonies in Alaska during July-October 2006. Satellite location and dive data were used to define habitat use in terms of the proportion of time spent or dives made in different oceanographic/bathymetric domains. For both plasma and red blood cells, the models accurately predicted habitat use for animals that foraged exclusively off or on the continental shelf. The models did not perform as well in predicting habitat use for animals that foraged in both on- and off-shelf habitat; however, sample sizes for these animals were small. Concurrently collected scat, fatty acid, and dive data confirmed that the foraging differences predicted by isotopes were associated with diet differences. Stable isotope samples, dive data, and GPS location data collected from an additional 15 females during August-October 2008 validated the effective use of the models across years. Little within year variation in habitat use was indicated from the comparison between stable isotope values from plasma (representing 1-2 weeks) and red blood cells (representing the prior few months). Constructing predictive models using stable isotopes provides an effective means to assess habitat use at the population level, is inexpensive, and can be applied to other marine predators.

During wild foraging, Australian fur seals (Arctocephalus pusillus doriferus) encounter many different types of prey in a wide range of scenarios, yet in captive environments they are typically provided with a narrower range of opportunities to display their full repertoire of behaviours. This study aimed to quantitatively explore the effect of foraging-based enrichment on the behaviour and activity patterns displayed by two captive Australian fur seals at Melbourne Zoo, Australia. Food was presented as a scatter in open water, in a free-floating ball device, or in a static box device, with each treatment separated by control trials with no enrichment. Both subjects spent more time interacting with the ball and static box devices than the scatter feed. The total time spent pattern swimming was reduced in the enrichment treatments compared to the controls, while the time spent performing random swimming behaviours increased. There was also a significant increase in the total number of bouts of behaviour performed in all three enrichment treatments compared to controls. Each enrichment method also promoted a different suit of foraging behaviours. Hence, rather than choosing one method, the most effective way to increase the diversity of foraging behaviours, while also increasing variation in general activity patterns, is to provide seals with a wide range of foraging scenarios where food is encountered in different ways.

We tagged 82 lactating northern fur seals (Callorhinus ursinus) with tri-axial accelerometers and magnetometers on two eastern Bering Sea islands (Bogoslof and St. Paul) with contrasting population trajectories. Using depth data, accelerometer data and spectral analysis we classified time spent diving (30%), resting (~7%), shaking and grooming their pelage (9%), swimming in the prone position (~10%) and two types of previously undocumented rolling behavior (29%), with the remaining time (~15%) unspecified. The reason for the extensive rolling behavior is not known. We ground-truthed the accelerometry signals for shaking and grooming and rolling behaviors--and identified the acceleration signal for porpoising--by filming tagged northern fur seals in captivity. Speeds from GPS interpolated data indicated that animals traveled fastest while in the prone position, suggesting that this behavior is indicative of destination-based swimming. Very little difference was found in the percentages of time spent in the categorical behaviors with respect to breeding islands (Bogoslof or St. Paul Island), forager type (cathemeral or nocturnal), and the region where the animals foraged (primarily on-shelf <200 m, or off-shelf > 200 m). The lack of significant differences between islands, regions and forager type may indicate that behaviors summarized over a trip are somewhat hardwired even though foraging trip length and when and where animals dive are known to vary with island, forager type and region.

Strategies employed by wide-ranging foraging animals involve consideration of habitat quality and predictability and should maximise net energy gain. Fidelity to foraging sites is common in areas of high resource availability or where predictable changes in resource availability occur. However, if resource availability is heterogeneous or unpredictable, as it often is in marine environments, then habitat familiarity may also present ecological benefits to individuals. We examined the winter foraging distribution of female Antarctic fur seals, Arctocephalus gazelle, over four years to assess the degree of foraging site fidelity at two scales; within and between years. On average, between-year fidelity was strong, with most individuals utilising more than half of their annual foraging home range over multiple years. However, fidelity was a bimodal strategy among individuals, with five out of eight animals recording between-year overlap values of greater than 50%, while three animals recorded values of less than 5%. High long-term variance in sea surface temperature, a potential proxy for elevated long-term productivity and prey availability, typified areas of overlap. Within-year foraging site fidelity was weak, indicating that successive trips over the winter target different geographic areas. We suggest that over a season, changes in prey availability are predictable enough for individuals to shift foraging area in response, with limited associated energetic costs. Conversely, over multiple years, the availability of prey resources is less spatially and temporally predictable, increasing the potential costs of shifting foraging area and favouring long-term site fidelity. In a dynamic and patchy environment, multi-year foraging site fidelity may confer a long-term energetic advantage to the individual. Such behaviours that operate at the individual level have evolutionary and ecological implications and are potential drivers of niche specialization and modifiers of intra-specific competition.

Predatory interaction of marine mammals with longline fisheries is observed globally, leading to partial or complete loss of the catch and in some parts of the world to considerable financial loss. Depredation can also create additional unrecorded fishing mortality of a stock and has the potential to introduce bias to stock assessments. Here we aim to characterise depredation in the Patagonian toothfish (Dissostichus eleginoides) fishery around South Georgia focusing on the spatio-temporal component of these interactions. Antarctic fur seals (Arctocephalus gazella), sperm whales (Physeter macrocephalus), and orcas (Orcinus orca) frequently feed on fish hooked on longlines around South Georgia. A third of longlines encounter sperm whales, but loss of catch due to sperm whales is insignificant when compared to that due to orcas, which interact with only 5% of longlines but can take more than half of the catch in some cases. Orca depredation around South Georgia is spatially limited and focused in areas of putative migration routes, and the impact is compounded as a result of the fishery also concentrating in those areas at those times. Understanding the seasonal behaviour of orcas and the spatial and temporal distribution of "depredation hot spots" can reduce marine mammal interactions, will improve assessment and management of the stock and contribute to increased operational efficiency of the fishery. Such information is valuable in the effort to resolve the human-mammal conflict for resources.

This is the first study estimating the ability of sea mammals to perceive and analyze information under the conditions of an almost complete sleep deprivation.

Rationale: The feeding success of predators is associated with the competition level for resources, and, thus, sympatric species are exposed to a potential trophic overlap. Isotopic Bayesian mixing models should provide a better understanding of the contribution of preys to the diet of predators and the feeding behavior of a species over time., Methods: The carbon and nitrogen isotopic signatures from pup hair samples of 93 Galapagos sea lions and 48 Galapagos fur seals collected between 2003 and 2009 in different regions (east and west) of the archipelago were analyzed. A PDZ Europa ANCA-GSL elemental analyzer interfaced with a PDZ Europa 20-20 continuous flow gas source mass spectrometer was employed. Bayesian models, SIAR and SIBER, were used to estimate the contribution of prey to the diet of predators, the niche breadth, and the trophic overlap level between the populations., Results: Statistical differences in the isotopic values of both predators were observed over the time. The mixing model determined that Galapagos fur seals had a primarily teutophagous diet, whereas the Galapagos sea lions fed exclusively on fish in both regions of the archipelago. The SIBER analysis showed differences in the trophic niche between the two sea lion populations, with the western rookery of the Galapagos sea lion being the population with the largest trophic niche area., Conclusions: A trophic niche partitioning between Galapagos fur seals and Galapagos sea lions in the west of the archipelago is suggested by our results. At intraspecific level, the western population of the Galapagos sea lion (ZwW) showed higher trophic breadth than the eastern population, a strategy adopted by the ZwW to decrease the interspecific competition levels in the western region., (Copyright © 2014 John Wiley & Sons, Ltd.)

Foraging behaviours used by two female Australian fur seals (Arctocephalus pusillus doriferus) were documented during controlled feeding trials. During these trials the seals were presented with prey either free-floating in open water or concealed within a mobile ball or a static box feeding device. When targeting free-floating prey both subjects primarily used raptorial biting in combination with suction, which was used to draw prey to within range of the teeth. When targeting prey concealed within either the mobile or static feeding device, the seals were able to use suction to draw out prey items that could not be reached by biting. Suction was followed by lateral water expulsion, where water drawn into the mouth along with the prey item was purged via the sides of the mouth. Vibrissae were used to explore the surface of the feeding devices, especially when locating the openings in which the prey items had been hidden. The mobile ball device was also manipulated by pushing it with the muzzle to knock out concealed prey, which was not possible when using the static feeding device. To knock prey out of this static device one seal used targeted bubble blowing, where a focused stream of bubbles was blown out of the nose into the openings in the device. Once captured in the jaws, prey items were manipulated and re-oriented using further mouth movements or chews so that they could be swallowed head first. While most items were swallowed whole underwater, some were instead taken to the surface and held in the teeth, while being vigorously shaken to break them into smaller pieces before swallowing. The behavioural flexibility displayed by Australian fur seals likely assists in capturing and consuming the extremely wide range of prey types that are targeted in the wild, during both benthic and epipelagic foraging.

For central place foragers, forming colonies can lead to extensive competition for prey around breeding areas and a zone of local prey depletion. As populations grow, this area of reduced prey can expand impacting foraging success and forcing animals to alter foraging behaviour. Here, we examine a population of marine predators, the northern fur seal (Callorhinus ursinus), which colonized a recently formed volcanic island, and assess changes in foraging behaviour associated with increasing population density. Specifically, we measured pup production and adult foraging behaviour over a 15-year period, during which the population increased 4-fold. Using measures of at-sea movements and dive behaviour, we found clear evidence that as the population expanded, animals were required to allot increasing effort to obtain resources. These changes in behaviour included longer duration foraging trips, farther distances travelled, a larger foraging range surrounding the island and deeper maximum dives. Our results suggest that as the northern fur seal population increased, local prey resources were depleted as a result of increased intraspecific competition. In addition, the recent slowing of population growth indicates that this population may be approaching carrying capacity just 31 years after a natural colonization event. Our study offers insight into the dynamics of population growth and impacts of increasing population density on a large marine predator. Such data could be vital for understanding future population fluctuations that occur in response to the dynamic environment, as natural and anthropogenic factors continue to modify marine habitats., (Published 2014. This article is a U.S. Government work and is in the public domain in the USA.)

Behavioral responses by top marine predators to oceanographic features such as eddies, river plumes, storms, and coastal topography suggest that biophysical interactions in these zones affect predators' prey, foraging behaviors, and potentially fitness. However, examining these pathways is challenged by the obstacles inherent in obtaining simultaneous observations of surface and subsurface environmental fields and predator behavior. In this study, migratory movements and, in some cases, diving behavior of 40 adult female northern fur seals (NFS; Callorhinus ursinus) were quantified across their range and compared to remotely-sensed environmental data in the Gulf of Alaska and California Current ecosystems, with a particular focus off the coast of Washington State (USA)--a known foraging ground for adult female NFS and where autonomous glider sampling allowed opportunistic comparison of seal behavior to subsurface biophysical measurements. The results show that in these ecosystems, adult female habitat utilization was concentrated near prominent coastal topographic, riverine, or inlet features and within 200 km of the continental shelf break. Seal dive depths, in most ecosystems, were moderated by surface light level (solar or lunar), mirroring known behaviors of diel vertically-migrating prey. However, seal dives differed in the California Current ecosystem due to a shift to more daytime diving concentrated at or below the surface mixed layer base. Seal movement models indicate behavioral responses to season, ecosystem, and surface wind speeds; individuals also responded to mesoscale eddies, jets, and the Columbia River plume. Foraging within small scale surface features is consistent with utilization of the inner coastal transition zone and habitats near coastal capes, which are known eddy and filament generation sites. These results contribute to our knowledge of NFS migratory patterns by demonstrating surface and subsurface behavioral responses to a spatially and temporally dynamic ocean environment, thus reflecting its influence on associated NFS prey species.

Adult male and female northern fur seals (Callorhinus ursinus) are sexually segregated in different regions of the North Pacific Ocean and Bering Sea during their winter migration. Explanations for this involve interplay between physiology, predator-prey dynamics, and ecosystem characteristics, however possible mechanisms lack empirical support. To investigate factors influencing the winter ecology of both sexes, we deployed five satellite-linked conductivity, temperature, and depth data loggers on adult males, and six satellite-linked depth data loggers and four satellite transmitters on adult females from St. Paul Island (Bering Sea, Alaska, USA) in October 2009. Males and females migrated to different regions of the North Pacific Ocean: males wintered in the Bering Sea and northern North Pacific Ocean, while females migrated to the Gulf of Alaska and California Current. Horizontal and vertical movement behaviors of both sexes were influenced by wind speed, season, light (sun and moon), and the ecosystem they occupied, although the expression of the behaviors differed between sexes. Male dive depths were aligned with the depth of the mixed layer during daylight periods and we suspect this was the case for females upon their arrival to the California Current. We suggest that females, because of their smaller size and physiological limitations, must avoid severe winters typical of the northern North Pacific Ocean and Bering Sea and migrate long distances to areas of more benign environmental conditions and where prey is shallower and more accessible. In contrast, males can better tolerate often extreme winter ocean conditions and exploit prey at depth because of their greater size and physiological capabilities. We believe these contrasting winter behaviors 1) are a consequence of evolutionary selection for large size in males, important to the acquisition and defense of territories against rivals during the breeding season, and 2) ease environmental/physiological constraints imposed on smaller females.

Predicting how climatic variations will affect marine predator populations relies on our ability to assess foraging success, but evaluating foraging success in a marine predator at sea is particularly difficult. Dive metrics are commonly available for marine mammals, diving birds and some species of fish. Bottom duration or dive duration are usually used as proxies for foraging success. However, few studies have tried to validate these assumptions and identify the set of behavioral variables that best predict foraging success at a given time scale. The objective of this study was to assess if foraging success in Antarctic fur seals could be accurately predicted from dive parameters only, at different temporal scales. For this study, 11 individuals were equipped with either Hall sensors or accelerometers to record dive profiles and detect mouth-opening events, which were considered prey capture attempts. The number of prey capture attempts was best predicted by descent and ascent rates at the dive scale; bottom duration and descent rates at 30-min, 1-h, and 2-h scales; and ascent rates and maximum dive depths at the all-night scale. Model performances increased with temporal scales, but rank and sign of the factors varied according to the time scale considered, suggesting that behavioral adjustment in response to prey distribution could occur at certain scales only. The models predicted the foraging intensity of new individuals with good accuracy despite high inter-individual differences. Dive metrics that predict foraging success depend on the species and the scale considered, as verified by the literature and this study. The methodology used in our study is easy to implement, enables an assessment of model performance, and could be applied to any other marine predator.

Bio-energetic models used to characterize an animal's energy budget require the accurate estimate of different variables such as the resting metabolic rate (RMR) and the heat increment of feeding (HIF). In this study, we estimated the in air RMR of wild juvenile South American fur seals (SAFS; Arctocephalus australis) temporarily held in captivity by measuring oxygen consumption while at rest in a postabsorptive condition. HIF, which is an increase in metabolic rate associated with digestion, assimilation and nutrient interconversion, was estimated as the difference in resting metabolic rate between the postabsorptive condition and the first 3.5h postprandial. As data were hierarchically structured, linear mixed effect models were used to compare RMR measures under both physiological conditions. Results indicated a significant increase (61%) for the postprandial RMR compared to the postabsorptive condition, estimated at 17.93±1.84 and 11.15±1.91mL O2 min(-1)kg(-1), respectively. These values constitute the first estimation of RMR and HIF in this species, and should be considered in the energy budgets for juvenile SAFS foraging at-sea., (Copyright © 2013 Elsevier Inc. All rights reserved.)

We propose a method to model the physiological link between somatic survival and reproductive output that reduces the number of parameters that need to be estimated by models designed to determine combinations of birth and death rates that produce historic counts of animal populations. We applied our Reproduction and Somatic Survival Linked (RSSL) method to the population counts of three species of North Pacific pinnipeds (harbor seals, Phoca vitulina richardii (Gray, 1864); northern fur seals, Callorhinus ursinus (L., 1758); and Steller sea lions, Eumetopias jubatus (Schreber, 1776))--and found our model outperformed traditional models when fitting vital rates to common types of limited datasets, such as those from counts of pups and adults. However, our model did not perform as well when these basic counts of animals were augmented with additional observations of ratios of juveniles to total non-pups. In this case, the failure of the ratios to improve model performance may indicate that the relationship between survival and reproduction is redefined or disassociated as populations change over time or that the ratio of juveniles to total non-pups is not a meaningful index of vital rates. Overall, our RSSL models show advantages to linking survival and reproduction within models to estimate the vital rates of pinnipeds and other species that have limited time-series of counts.

1. Analyses of animal resource selection functions (RSF) using data collected from relocations of individuals via remote telemetry devices have become commonplace. Increasing technological advances, however, have produced statistical challenges in analysing such highly autocorrelated data. Weighted distribution methods have been proposed for analysing RSFs with telemetry data. However, they can be computationally challenging due to an intractable normalizing constant and cannot be aggregated (i.e. collapsed) over time to make space-only inference. 2. In this study, we take a conceptually different approach to modelling animal telemetry data for making RSF inference. We consider the telemetry data to be a realization of a space-time point process. Under the point process paradigm, the times of the relocations are also considered to be random rather than fixed. 3. We show the point process models we propose are a generalization of the weighted distribution telemetry models. By generalizing the weighted model, we can access several numerical techniques for evaluating point process likelihoods that make use of common statistical software. Thus, the analysis methods can be readily implemented by animal ecologists. 4. In addition to ease of computation, the point process models can be aggregated over time by marginalizing over the temporal component of the model. This allows a full range of models to be constructed for RSF analysis at the individual movement level up to the study area level. 5. To demonstrate the analysis of telemetry data with the point process approach, we analysed a data set of telemetry locations from northern fur seals (Callorhinus ursinus) in the Pribilof Islands, Alaska. Both a space-time and an aggregated space-only model were fitted. At the individual level, the space-time analysis showed little selection relative to the habitat covariates. However, at the study area level, the space-only model showed strong selection relative to the covariates., (© 2013 The Authors. Journal of Animal Ecology © 2013 British Ecological Society.)

Cape fur seal (Arctocephalus pusillus) pups spend the first weeks of life exclusively or mainly ashore. They are exposed to intense solar radiation and high temperatures for long time periods, which results in temperatures up to at least 80°C on their black natal coat. To test the hypothesis that the natal coat has a crucial function in coping with these extreme conditions, we investigated the insulating properties of the natal coat in six captive newborn Cape fur seals during the first 50 days after birth. The natal fur differs from the adult fur not only in colour, but also in density, structure, and water repellence. We measured temperature on the fur surface and within the fur, as well as skin and rectal temperature under varying environmental conditions, comparable to the species' habitat. Experiments were designed to not influence the spontaneous behaviour of the pups. Rectal temperature was constant as long as the pups stayed dry, even during long-lasting intense solar radiation for up to 3 h. Skin temperature remained close to rectal temperature as long as the fur was dry, while with wet fur, skin temperature was significantly reduced as well. Our results show that the natal coat provides an effective insulation against overheating. The severely reduced insulation of wet natal fur against cold supports the assumption that the natal fur is an adaptation to the pups' terrestrial phase of life.

In 2010, Coxiella burnetii was identified in 75% of northern fur seal placentas from a single rookery in Alaska, but nothing was known about the significance of this organism in the population. Although many infectious organisms cause increased cell death, C. burnetii has been shown to suppress apoptosis of the host macrophages as an intracellular survival mechanism. To determine if infection induces a similar functional change in the placenta, immunohistochemistry for antibodies to cleaved caspase-3 (activated caspase-3) and the (TDT)-mediated dUTP-digoxigenin nick end labeling (TUNEL) technique were used to compare the amount of placental apoptosis in infected and noninfected placentas. There was a statistically significant difference in the frequency of apoptotic cells between infected and uninfected placentas, with more apoptosis identified in the uninfected placentas. This finding suggests that the survival mechanism of C. burnetii in host macrophages to reduce apoptosis may also be utilized in trophoblasts. The significance of decreased trophoblastic apoptosis for the northern fur seal fetus requires further investigation.

Two major drivers in population dynamics are bottom-up processes, such as environmental factors that affect foraging success, and the top-down impacts of predation. Many populations of marine mammal and seabird species appear to be declining in response to reductions in prey associated with the bottom-up effects of climate change. However, predation, which usually occurs at sea and is difficult to observe, may also play a key role. We analysed drivers of population dynamics of Antarctic fur seals Arctocephalus gazella at Cape Shirreff from 1997 to 2009, including a predator that targets pre-weaned pups and bottom-up environmental effects in an ecosystem particularly sensitive to small changes in temperature. We use Bayesian mark-recapture analysis to demonstrate that although large-scale environmental variability affects annual adult survival and reproduction, first year survival appears to be driving the current decline in this population (as defined by a decline in the annual number of pups born). Although the number of pups increased during the first third of the study, first year survival and recruitment of those pups in later years was very low. Such low survival may be driven by leopard seal Hydrurga leptonyx predation, particularly prior to weaning. Our results suggest that without leopard seal predation, this population would most likely increase in size, despite the observed bottom-up effects of climate changes on adult vital rates. More broadly, our results show how age-targeted predation could be a major factor in population decline of K-selected colonial breeders., (© 2013 The Authors. Journal of Animal Ecology © 2013 British Ecological Society.)

Pinnipeds, that is true seals (Phocidae), eared seals (Otariidae), and walruses (Odobenidae), possess highly developed vibrissal systems for mechanoreception. They can use their vibrissae to detect and discriminate objects by direct touch. At least in Phocidae and Otariidae, the vibrissae can also be used to detect and analyse water movements. Here, we review what is known about this ability, known as hydrodynamic perception, in pinnipeds. Hydrodynamic perception in pinnipeds developed convergently to the hydrodynamic perception with the lateral line system in fish and the sensory hairs in crustaceans. So far two species of pinnipeds, the harbour seal (Phoca vitulina) representing the Phocidae and the California sea lion (Zalophus californianus) representing the Otariidae, have been studied for their ability to detect local water movements (dipole stimuli) and to follow hydrodynamic trails, that is the water movements left behind by objects that have passed by at an earlier point in time. Both species are highly sensitive to dipole stimuli and can follow hydrodynamic trails accurately. In the individuals tested, California sea lions were clearly more sensitive to dipole stimuli than harbour seals, and harbour seals showed a superior trail following ability as compared to California sea lions. Harbour seals have also been shown to derive additional information from hydrodynamic trails, such as motion direction, size and shape of the object that caused the trail (California sea lions have not yet been tested). The peculiar undulated shape of the harbour seals' vibrissae appears to play a crucial role in trail following, as it suppresses self-generated noise while the animal is swimming.

Using a food-rewarded two-choice instrumental conditioning paradigm we assessed the ability of South African fur seals, Arctocephalus pusillus, to discriminate between 12 enantiomeric odor pairs. The results demonstrate that the fur seals as a group were able to discriminate between the optical isomers of carvone, dihydrocarvone, dihydrocarveol, menthol, limonene oxide, α-pinene, fenchone (all p < 0.01), and β-citronellol (p < 0.05), whereas they failed to distinguish between the (+)- and (-)-forms of limonene, isopulegol, rose oxide, and camphor (all p > 0.05). An analysis of odor structure-activity relationships suggests that a combination of molecular structural properties rather than a single molecular feature may be responsible for the discriminability of enantiomeric odor pairs. A comparison between the discrimination performance of the fur seals and that of other species tested previously on the same set of enantiomers (or subsets thereof) suggests that the olfactory discrimination capabilities of this marine mammal are surprisingly well developed and not generally inferior to that of terrestrial mammals such as human subjects and non-human primates. Further, comparisons suggest that neither the relative nor the absolute size of the olfactory bulbs appear to be reliable predictors of between-species differences in olfactory discrimination capabilities. Taken together, the results of the present study support the notion that the sense of smell may play an important and hitherto underestimated role in regulating the behavior of fur seals.

Underwater, the contrast between object and background is much larger reduced with increasing distance between object and observer than in air. For marine predators, such as pinnipeds, it would therefore be advantageous to possess a high sensitivity for brightness differences, since this would increase the distance at which prey can be detected visually. Few studies have examined the brightness discrimination thresholds of pinnipeds. Two studies with phocid seals have confirmed low brightness discrimination thresholds in pinnipeds whereas the threshold obtained for the South African fur seal seems to be twice as high as that of the phocids. However, the experiments with the South African fur seal have been conducted under inadequate conditions which likely resulted in an underestimation of the brightness discrimination ability of this species. The study at hand reinvestigated the brightness discrimination threshold of the South African fur seal under well controlled conditions. In a two alternative forced choice task, one fur seal was trained to indicate the position of the brighter of two gray discs presented on a black background on a monitor. The thresholds were determined for 11 standard intensities each tested against 8 lower comparison intensities. It was found that the fur seal was able to perceive brightness differences of 8-10%, which is better than the phocid species tested so far. For low standard intensities, however, the threshold increased which could to be due to a relative slow dark adaptation rate of the fur seal. The results are discussed in terms of the relevance of visual information for pinnipeds during foraging dives and are directly compared to the results obtained for the harbor seal which has been tested under the same conditions as the fur seal in a previous study., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

1. In vertebrates, patterns of resource utilization change throughout development according to age- and or size-specific abilities and requirements. Thus, interspecific competition affects different age classes differently. 2. Adults of sympatric species often show distinct foraging niche segregation, but juvenile resource use might overlap with adult competitors of similar body size. Resultant negative effects on juveniles can have important consequences for population dynamics, yet such interactions have received little attention in studies of mammalian communities. 3. Using GPS tracking devices, time-depth recorders and stable isotope data, we compared diving depth, activity time, trophic position and foraging habitat characteristics to investigate foraging niche overlap between similar-sized sympatric juvenile Galapagos sea lions (Zalophus wollebaeki) and adult Galapagos fur seals (Arctocephalus galapagoensis) and compared each group with much larger-bodied adult Galapagos sea lions. 4. We found little indication for direct competition but a complex pattern of foraging niche segregation: juvenile sea lions and adult fur seals dived to shallow depths at night, but foraged in different habitats with limited spatial overlap. Conversely, juvenile and adult sea lions employed different foraging patterns, but their foraging areas overlapped almost completely. 5. Consistency of foraging habitat characteristics between juvenile and adult sea lions suggests that avoidance of competition may be important in shaping foraging habitat utilization. Resultant specialization on a limited habitat could contribute to low sea lion numbers that contrast with high fur seal abundance. Our data suggest that exploitation by multiple predators within spatially restricted foraging ranges of juveniles might negatively impact juvenile foraging success and ultimately influence population dynamics., (© 2013 The Authors. Journal of Animal Ecology © 2013 British Ecological Society.)

Spatial coherence between predators and prey has rarely been observed in pelagic marine ecosystems. We used measures of the environment, prey abundance, prey quality, and prey distribution to explain the observed distributions of three co-occurring predator species breeding on islands in the southeastern Bering Sea: black-legged kittiwakes (Rissa tridactyla), thick-billed murres (Uria lomvia), and northern fur seals (Callorhinus ursinus). Predictions of statistical models were tested using movement patterns obtained from satellite-tracked individual animals. With the most commonly used measures to quantify prey distributions--areal biomass, density, and numerical abundance--we were unable to find a spatial relationship between predators and their prey. We instead found that habitat use by all three predators was predicted most strongly by prey patch characteristics such as depth and local density within spatial aggregations. Additional prey patch characteristics and physical habitat also contributed significantly to characterizing predator patterns. Our results indicate that the small-scale prey patch characteristics are critical to how predators perceive the quality of their food supply and the mechanisms they use to exploit it, regardless of time of day, sampling year, or source colony. The three focal predator species had different constraints and employed different foraging strategies--a shallow diver that makes trips of moderate distance (kittiwakes), a deep diver that makes trip of short distances (murres), and a deep diver that makes extensive trips (fur seals). However, all three were similarly linked by patchiness of prey rather than by the distribution of overall biomass. This supports the hypothesis that patchiness may be critical for understanding predator-prey relationships in pelagic marine systems more generally.

Unihemispheric sleep has been observed in numerous species, including birds and aquatic mammals. While knowledge of its functional role has been improved in recent years, the physiological mechanisms that generate this behavior remain poorly understood. Here, unihemispheric sleep is simulated using a physiologically based quantitative model of the mammalian ascending arousal system. The model includes mutual inhibition between wake-promoting monoaminergic nuclei (MA) and sleep-promoting ventrolateral preoptic nuclei (VLPO), driven by circadian and homeostatic drives as well as cholinergic and orexinergic input to MA. The model is extended here to incorporate two distinct hemispheres and their interconnections. It is postulated that inhibitory connections between VLPO nuclei in opposite hemispheres are responsible for unihemispheric sleep, and it is shown that contralateral inhibitory connections promote unihemispheric sleep while ipsilateral inhibitory connections promote bihemispheric sleep. The frequency of alternating unihemispheric sleep bouts is chiefly determined by sleep homeostasis and its corresponding time constant. It is shown that the model reproduces dolphin sleep, and that the sleep regimes of humans, cetaceans, and fur seals, the latter both terrestrially and in a marine environment, require only modest changes in contralateral connection strength and homeostatic time constant. It is further demonstrated that fur seals can potentially switch between their terrestrial bihemispheric and aquatic unihemispheric sleep patterns by varying just the contralateral connection strength. These results provide experimentally testable predictions regarding the differences between species that sleep bihemispherically and unihemispherically., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Slow wave sleep (SWS) in the northern fur seal (Callorhinus ursinus) is characterized by a highly expressed interhemispheric electroencephalogram (EEG) asymmetry, called 'unihemispheric' or 'asymmetrical' SWS. The aim of this study was to examine the regional differences in slow wave activity (SWA; power in the range of 1.2-4.0 Hz) within one hemisphere and differences in the degree of interhemispheric EEG asymmetry within this species. Three seals were implanted with 10 EEG electrodes, positioned bilaterally (five in each hemisphere) over the frontal, occipital and parietal cortex. The expression of interhemispheric SWA asymmetry between symmetrical monopolar recordings was estimated based on the asymmetry index [AI = (L-R)/(L+R), where L and R are the power in the left and right hemispheres, respectively]. Our findings indicate an anterior-posterior gradient in SWA during asymmetrical SWS in fur seals, which is opposite to that described for other mammals, including humans, with a larger SWA recorded in the parietal and occipital cortex. Interhemispheric EEG asymmetry in fur seals was recorded across the entire dorsal cerebral cortex, including sensory (visual and somatosensory), motor and associative (parietal or suprasylvian) cortical areas. The expression of asymmetry was greatest in occipital-lateral and parietal derivations and smallest in frontal-medial derivations. Regardless of regional differences in SWA, the majority (90%) of SWS episodes with interhemispheric EEG asymmetry meet the criteria for 'unihemispheric SWS' (one hemisphere is asleep while the other is awake). The remaining episodes can be described as episodes of bilateral SWS with a local activation in one cerebral hemisphere., (© 2012 European Sleep Research Society.)

The objective of this study was to assess whether the freezing and thawing of pinniped carcasses prior to post-mortem examination could create artefacts that resembled lesions caused by trauma. Necropsy findings in New Zealand fur seals (Arctocephalus forsteri), captured incidental to commercial fishing, and either chilled (n=5) or frozen (n=5), were compared. Changes in frozen, but not in chilled, carcasses included: pseudo-bruising of subcutis; the accumulation of thick dark red fluid (resembling haemorrhage) in the thoracic cavity, pericardial sac and abdominal cavity; apparent subcapsular renal haemorrhage; pseudo-contusions of the brain; apparent haemorrhage from the nares; and blood-staining of the anterior ocular chamber. The processes of freezing and thawing were strongly associated with subcutaneous pseudo-bruises, the presence of thick, dark red abdominal fluid and renal subcapsular 'haemorrhage' (P=0.004). These artefacts probably develop due to a combination of autolysis and 'freeze-thaw' effects including lysis of cell membranes, fluid shifts into the extracellular space, and disruption of blood vessel walls. The results of the study demonstrate that artefacts resembling traumatic lesions are created during freezing and thawing of pinniped bodies. Such changes must be taken into consideration at post-mortem examination of previously frozen carcasses., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Thoracic auscultation is an important diagnostic method used in cases of suspected pulmonary disease in many species, as respiratory sounds contain significant information on the physiology and pathology of the lungs and upper airways. Respiratory diseases are frequent in marine mammals and are often listed as one of their main causes of death. The aim of this study was to investigate and report baseline parameters for the electronic-mediated thoracic auscultation of one cetacean species and two pinniped species in captivity. Respiratory sounds from 20 captive bottlenose dolphins (Tursiops truncatus), 6 California sea lions (Zalophus californianus), and 5 South African fur seals (Arctocephalus pusillus) were recorded with an electronic stethoscope. The sounds were analyzed for duration of the respiratory cycle, adventitious sounds, and peak frequencies of recorded sounds during expiration and inspiration as well as for sound intensity as reflected by waveform amplitude during the respiratory cycle. In respiratory cycles of the bottlenose dolphins' expiring "on command," the duration of the expiration was significantly shorter than the duration of the inspiration. In the examined pinnipeds of this study, there was no clear pattern concerning the duration of one breathing phase: Adventitious sounds were detected most often in bottlenose dolphins that were expiring on command and could be compared with "forced expiratory wheezes" in humans. This is the first report of forced expiratory wheezes in bottlenose dolphins; they can easily be misinterpreted as pathologic respiratory sounds. The peak frequencies of the respiratory sounds reached over 2,000 Hz in bottlenose dolphins and over 1,000 Hz in California sea lions and South African fur seals, but the variation of the frequency spectra was very high in all animals. To the authors' knowledge, this is the first systematic analysis of respiratory sounds of bottlenose dolphins and two species of pinnipeds.

Life-history theory predicts that selection will favor optimal levels of parental effort that balance benefits of current reproduction with costs to survival and future reproduction. The optimal level of effort depends on parental traits, offspring traits, and provisioning strategy. Additionally, how these factors influence effort may differ depending on the stage of reproduction. The relative importance of maternal and offspring traits on energy allocation to offspring was investigated in known-age Antarctic fur seals Arctocephalus gazella across four stages of reproduction, using birth mass and milk-consumption measurements. Maternal traits were important during three of the four stages investigated, with larger females giving birth to larger pups and investing more in pups during perinatal and molt stages. Pup mass influenced maternal effort during the premolt stage, and provisioning strategy influenced postnatal maternal effort at all stages. Energy provided to the offspring during an attendance visit was positively related to the duration of the foraging-trip/visit cycle; however, when investment was controlled for trip/visit cycle duration, the overall rate of energy transfer was similar across trip durations. In addition to strong effects of maternal mass, pup traits affected energy allocation, suggesting that pup demand is important in determining maternal care. These findings emphasize the importance of considering state variables in life-history studies and suggest that timing of measurements of effort in species with long provisioning periods may influence conclusions and our ability to make comparisons of reproductive effort among species.

Northern fur seal (Callorhinus ursinus; NFS) populations have been declining, perhaps due to limited foraging ability of pups. Because a marine mammal's proficiency at exploiting underwater prey resources is based on the ability to store large amounts of oxygen (O(2)) and to utilize these reserves efficiently, this study was designed to determine if NFS pups had lower blood, muscle, and total body O(2) stores than adults. Pups (<1-month old) had a calculated aerobic dive limit only ~40% of adult females due to lower blood and, to a much greater extent, muscle O(2) stores. Development of the Pectoralis (Pec) and Longissimus dorsi (LD) skeletal muscles was further examined by determining their myosin heavy chain (MHC) composition and enzyme activities. In all animals, the slow MHC I and fast-twitch IIA proteins typical of oxidative fiber types were dominant, but adult muscles contained more (Pec ~50%; LD ~250% higher) fast-twitch MHC IID/X protein characteristic of glycolytic muscle fibers, than pup muscles. This suggests that adults have greater ability to generate muscle power rapidly and/or under anaerobic conditions. Pup muscles also had lower aerobic and anaerobic ATP production potential, as indicated by lower metabolically scaled citrate synthase, β-hydroxyacyl CoA dehydrogenase, and lactate dehydrogenase activities (all P values ≤0.001). In combination, these findings indicate that pups are biochemically and physiologically limited in their diving capabilities relative to adults. This may contribute to lower NFS first year survival.

We created a Bayesian hierarchical model (BHM) to investigate ecosystem relationships between the physical ecosystem (sea ice extent), a prey measure (krill density), predator behaviors (diving and foraging effort of female Antarctic fur seals, Arctocephalus gazella, with pups) and predator characteristics (mass of maternal fur seals and pups). We collected data on Antarctic fur seals from 1987/1988 to 1994/1995 at Seal Island, Antarctica. The BHM allowed us to link together predators and prey into a model that uses all the data efficiently and accounts for major sources of uncertainty. Based on the literature, we made hypotheses about the relationships in the model, which we compared with the model outcome after fitting the BHM. For each BHM parameter, we calculated the mean of the posterior density and the 95% credible interval. Our model confirmed others' findings that increased sea ice was related to increased krill density. Higher krill density led to reduced dive intensity of maternal fur seals, as measured by dive depth and duration, and to less time spent foraging by maternal fur seals. Heavier maternal fur seals and lower maternal foraging effort resulted in heavier pups at 22 d. No relationship was found between krill density and maternal mass, or between maternal mass and foraging effort on pup growth rates between 22 and 85 days of age. Maternal mass may have reflected environmental conditions prior to the pup provisioning season, rather than summer prey densities. Maternal mass and foraging effort were not related to pup growth rates between 22 and 85 d, possibly indicating that food was not limiting, food sources other than krill were being used, or differences occurred before pups reached age 22 d.

Australian fur seals breed on thirteen islands located in the Bass Strait, Australia. Land access to these islands is restricted, minimising human presence but boat access is still permissible with limitations on approach distances. Thirty-two controlled noise exposure experiments were conducted on breeding Australian fur seals to determine their behavioural response to controlled in-air motor boat noise on Kanowna Island (39°10'S, 146°18'E). Our results show there were significant differences in the seals' behaviour at low (64-70 dB) versus high (75-85 dB) sound levels, with seals orientating themselves towards or physically moving away from the louder boat noise at three different sound levels. Furthermore, seals responded more aggressively with one another and were more alert when they heard louder boat noise. Australian fur seals demonstrated plasticity in their vocal responses to boat noise with calls being significantly different between the various sound intensities and barks tending to get faster as the boat noise got louder. These results suggest that Australian fur seals on Kanowna Island show behavioural disturbance to high level boat noise. Consequently, it is recommended that an appropriate level of received boat sound emissions at breeding fur seal colonies be below 74 dB and that these findings be taken into account when evaluating appropriate approach distances and speed limits for boats.

Background: Individual variations in the use of the species niche are an important component of diversity in trophic interactions. A challenge in testing consistency of individual foraging strategy is the repeated collection of information on the same individuals., Methodology/principal Findings: The foraging strategies of sympatric fur seals (Arctocephalus gazella and A. tropicalis) were examined using the stable isotope signature of serially sampled whiskers. Most whiskers exhibited synchronous δ(13)C and δ(15)N oscillations that correspond to the seal annual movements over the long term (up to 8 years). δ(13)C and δ(15)N values were spread over large ranges, with differences between species, sexes and individuals. The main segregating mechanism operates at the spatial scale. Most seals favored foraging in subantarctic waters (where the Crozet Islands are located) where they fed on myctophids. However, A. gazella dispersed in the Antarctic Zone and A. tropicalis more in the subtropics. Gender differences in annual time budget shape the seal movements. Males that do not perform any parental care exhibited large isotopic oscillations reflecting broad annual migrations, while isotopic values of females confined to a limited foraging range during lactation exhibited smaller changes. Limited inter-individual isotopic variations occurred in female seals and in male A. tropicalis. In contrast, male A. gazella showed large inter-individual variations, with some males migrating repeatedly to high-Antarctic waters where they fed on krill, thus meaning that individual specialization occurred over years., Conclusions/significance: Whisker isotopic signature yields unique long-term information on individual behaviour that integrates the spatial, trophic and temporal dimensions of the ecological niche. The method allows depicting the entire realized niche of the species, including some of its less well-known components such as age-, sex-, individual- and migration-related changes. It highlights intrapopulation heterogeneity in foraging strategies that could have important implications for likely demographic responses to environmental variability.

Emerging Bayesian analytical approaches offer increasingly sophisticated means of reconstructing historical population dynamics from genetic data, but have been little applied to scenarios involving demographic bottlenecks. Consequently, we analysed a large mitochondrial and microsatellite dataset from the Antarctic fur seal Arctocephalus gazella, a species subjected to one of the most extreme examples of uncontrolled exploitation in history when it was reduced to the brink of extinction by the sealing industry during the late eighteenth and nineteenth centuries. Classical bottleneck tests, which exploit the fact that rare alleles are rapidly lost during demographic reduction, yielded ambiguous results. In contrast, a strong signal of recent demographic decline was detected using both Bayesian skyline plots and Approximate Bayesian Computation, the latter also allowing derivation of posterior parameter estimates that were remarkably consistent with historical observations. This was achieved using only contemporary samples, further emphasizing the potential of Bayesian approaches to address important problems in conservation and evolutionary biology., (© 2011 Blackwell Publishing Ltd.)

Locomotion is one of the major energetic costs faced by animals and various strategies have evolved to reduce its cost. Birds use interspersed periods of flapping and gliding to reduce the mechanical requirements of level flight while undergoing cyclical changes in flight altitude, known as undulating flight. Here we equipped free-ranging marine vertebrates with accelerometers and demonstrate that gait patterns resembling undulating flight occur in four marine vertebrate species comprising sharks and pinnipeds. Both sharks and pinnipeds display intermittent gliding interspersed with powered locomotion. We suggest, that the convergent use of similar gait patterns by distinct groups of animals points to universal physical and physiological principles that operate beyond taxonomic limits and shape common solutions to increase energetic efficiency. Energetically expensive large-scale migrations performed by many vertebrates provide common selection pressure for efficient locomotion, with potential for the convergence of locomotory strategies by a wide variety of species.

The hormonal factors that influence development from birth to weaning in otariid seals is still largely unknown. In the present study, a suite of thyroid hormones and cortisol were measured in Australian fur seal pups in order to determine baseline concentrations as well as to describe their endocrinology over this critical developmental period. A cross-section of newborn pups from a breeding colony located on Kanowna Island, Australia were sampled at six different times over the course of the 10 month lactation period. Sample times were designed to correspond to periods of heightened physiological change during pre-weaning development: post-natal, pre-molt, the initiation of molt, mid-molt, period of peak milk intake and weaning. Results indicate that the greatest hormonal changes were associated with the post-natal stage and molt, with molt showing the greatest changes, as has been reported for several species of pinnipeds. Two forms of thyroid hormones analyzed (Total T(4), and Free T(3)), increased with the initiation of the molt, and Free T(3) exhibited a second increase that was associated with the period of peak milk intake. The T(3):T(4) ratio was significantly lower during the initiation of molt than either pre- or mid-molt. The study was able to describe physiological change during the first year of life in Australian fur seals as well as document basal concentrations of thyroid hormones and cortisol in pups of this species., (Copyright © 2011 Elsevier Inc. All rights reserved.)

Data on fine-scale animal movement are being collected worldwide, with the number of species being tagged and the resolution of data rapidly increasing. In this study, a general methodology is proposed to understand the patterns in these high-resolution movement time series that relate to marine animal behavior. The approach is illustrated with dive data from a northern fur seal (Callorhinus ursinus) tagged on the Pribilof Islands, Alaska, USA. We apply a state-space model composed of a movement model and corresponding high-resolution vertical movement data. The central goal is to estimate parameters of this movement model, particularly their variation on appropriate time scales, thereby providing a direct link to behavior. A particle filter with state augmentation is used to jointly estimate the movement parameters and the state. A multiple iterated filter using overlapping data segments is implemented to match the parameter time scale with the behavioral inference. The time variation in the auto-covariance function facilitates identification of a movement model, allows separation of observation and process noise, and provides for validation of results. The analysis yields fitted parameters that show distinct time-evolving changes in fur seal behavior over time, matching well what is observed in the original data set.

Zero offset correction of diving depth measured by time-depth recorders is required to remove artifacts arising from temporal changes in accuracy of pressure transducers. Currently used methods for this procedure are in the proprietary software domain, where researchers cannot study it in sufficient detail, so they have little or no control over how their data were changed. GNU R package diveMove implements a procedure in the Free Software domain that consists of recursively smoothing and filtering the input time series using moving quantiles. This paper describes, demonstrates, and evaluates the proposed method by using a "perfect" data set, which is subsequently corrupted to provide input for the proposed procedure. The method is evaluated by comparing the corrected time series to the original, uncorrupted, data set from an Antarctic fur seal (Arctocephalus gazella Peters, 1875). The Root Mean Square Error of the corrected data set, relative to the "perfect" data set, was nearly identical to the magnitude of noise introduced into the latter. The method, thus, provides a flexible, reliable, and efficient mechanism to perform zero offset correction for analyses of diving behaviour. We illustrate applications of the method to data sets from four species with large differences in diving behaviour, measured using different sampling protocols and instrument characteristics.

Understanding animal movement and resource selection provides important information about the ecology of the animal, but an animal's movement and behavior are not typically constant in time. We present a velocity-based approach for modeling animal movement in space and time that allows for temporal heterogeneity in an animal's response to the environment, allows for temporal irregularity in telemetry data, and accounts for the uncertainty in the location information. Population-level inference on movement patterns and resource selection can then be made through cluster analysis of the parameters related to movement and behavior. We illustrate this approach through a study of northern fur seal (Callorhinus ursinus) movement in the Bering Sea, Alaska, USA. Results show sex differentiation, with female northern fur seals exhibiting stronger response to environmental variables.

Thirteen standard hematology values were determined for a healthy and growing population of free-ranging, lactating northern fur seals (Callorhinus ursinus) from Lovushki Island in the Kuril Islands of far-east Russia. Results are presented from 24 females sampled between June and August during the 3-yr period of 2006-08. Hematologic values have been made available for future comparisons with the declining population of northern fur seals on the Pribilof Islands, Alaska, and are compared with published values for other otariid species.

The parallelisms in communicative behavior of domestic sheep and Northern fur seals within a herd are accompanied by parallelisms in parameters of sound signal, the calling scream. This signal ensures ties between babies and their mothers at a long distance. The basis of parallelisms is formed by amplitude modulation at two levels: the one being a direct amplitude modulation of the carrier frequency and the other--modulation of the carrier frequency oscillation. Parallelisms in the signal oscillatory process result in corresponding parallelisms in the structure of its frequency spectrum.

Behavioral thermoregulation represents an important strategy for reducing energetic costs in thermally challenging environments, particularly among terrestrial vertebrates. Because of the cryptic lifestyle of aquatic species, the energetic benefits of such behaviors in marine endotherms have been much more difficult to demonstrate. In this study, I examined the importance of behavioral thermoregulation in the northern fur seal (Callorhinus ursinus) pup, a small-bodied endotherm that spends prolonged periods at sea. The thermal neutral zones of three weaned male northern fur seal pups (body mass range = 11.8-12.8 kg) were determined by measuring resting metabolic rate using open-flow respirometry at water temperatures ranging from 2.5° to 25.0°C. Metabolic rate averaged 10.03 ± 2.26 mL O₂kg⁻¹ min⁻¹ for pups resting within their thermal neutral zone; lower critical temperature was 8.3° ± 2.5°C , approximately 8°C higher than the coldest sea surface temperatures encountered in northern Pacific waters. To determine whether behavioral strategies could mitigate this potential thermal limitation, I measured metabolic rate during grooming activities and the unique jughandling behavior of fur seals. Both sedentary grooming and active grooming resulted in significant increases in metabolic rate relative to rest (P = 0.001), and percent time spent grooming increased significantly at colder water temperatures (P < 0.001). Jughandling metabolic rate (12.71 ± 2.73 mL O₂kg⁻¹ min ⁻¹) was significantly greater than resting rates at water temperatures within the thermal neutral zone (P < 0.05) but less than resting metabolism at colder water temperatures. These data indicate that behavioral strategies may help to mitigate thermal challenges faced by northern fur seal pups while resting at sea.

Life history trade-offs have often been assumed to be the consequence of restrictions in the availability of critical resources such as energy and nutrients, which necessitate the differential allocation of resources to costly traits. Here, we examined endocrine (testosterone) and health (parasite burdens) parameters in territorial and non-territorial New Zealand fur seal males. We documented intra-sexual differences in sexual behaviours, testosterone levels, and parasitism that suggest a trade-off exists between reproductive success and physical health, particularly susceptibility to helminths and acanthocephalans, in males displaying different mating tactics (i.e., territorial and non-territorial tactics). Levels of testosterone were higher in territorial males and correlated positively with reproductive effort (i.e., intra- and inter-sexual interactions). However, these territorial males also exhibited high levels of parasitic infection, which may impair survival in the long-term. Our study, while limited in sample size, provides preliminary evidence for a link between male mating tactics, testosterone levels and parasite loads, and potential effects on reproductive success and life history that should be explored further.

Background: In systems where two or more species experience secondary contact, behavioural factors that regulate interspecific gene flow may be important for maintaining species boundaries and reducing the incidence of hybridisation. At subantarctic Macquarie Island, two species of fur seal breed in close proximity to one another, hybridise at very high levels (up to 21% of hybrid pups are born annually), yet retain discrete gene pools. Using spatial and genetic information collected for pups and adults over twelve years, we assessed two behavioural traits - inter-annual site fidelity and differences in habitat use between the species - as possible contributors to the maintenance of this species segregation. Further, we explored the breakdown of these traits in pure-species individuals and hybrids., Results: We found virtually complete spatial segregation of the parental species, with only one exception; a single territory that contained adults of both species and also the highest concentration of hybrid pups. The spatial distribution of each species was closely linked to habitat type (pebbled vs boulder beaches), with members of each species breeding almost exclusively on one type or the other but hybrids breeding on both or at the junction between habitats. Inter-annual site fidelity was high for both sexes of pure-species adults, with 66% of females and all males returning to the same territory or a neighbouring one in different years. An important consequence for pure females of breeding on the 'wrong' habitat type, and thus in a heterospecific aggregation, was the production of hybrid pups. Low habitat fidelity of hybrid females facilitated bi-directional backcrossing, resulting in more diverse hybrid offspring., Conclusion: In a disturbed system where two sympatric fur seal species breed in close proximity, discrete gene pools are retained by extremely fine-scale and strong spatial segregation of the species. Two behavioural traits were found to be important in maintaining this stable population structure, and habitat type was a strong indicator of where species locate and a potentially powerful predictor of future directions of hybridisation. A direct consequence of the breakdown of this trait was the production of hybrid offspring, which may have severe implications if hybrids have reduced fitness.