Development of the blood and muscle oxygen stores in gray seals (Halichoerus grypus): implications for juvenile diving capacity and the necessity of a terrestrial postweaning fast

To successfully transition from nursing to foraging, phocid seal pups must develop adequate diving physiology within the limited time between birth and their first independent foraging trip to sea. We studied the postpartum development of oxygen stores in gray seals (Halichoerus grypus, n=40) to better understand the ontogeny of diving capacity in phocids. Hemoglobin (Hb), hematocrit (Hct), blood volume (BV), and myoglobin (Mb) levels in newborn (3 d postpartum [DPP]) and newly weaned (17+/-0.4 DPP) pups were among the lowest measured across age classes. During the pups' terrestrial postweaning fast (PWF), Hb, Hct, mass-specific BV, and Mb increased by 28%, 21%, 13%, and 29%, respectively, resulting in a 35% increase in total body mass-specific oxygen stores and a 23% increase in calculated aerobic dive limit (CADL). Although Hb and Hct levels at the end of the PWF were nearly identical to those of yearlings, total body mass-specific oxygen stores and CADL of weaned pups departing for sea were only 66%-67% and 32%-62%, respectively, of those for yearlings and adult females. The PWF represents an integral component of the physiological development of diving capacity in phocids; however, newly independent phocids still appear to have limited diving capabilities at the onset of foraging.     

Making it through the first year: Ontogeny of movement and diving behavior in harbor seals from Svalbard,Norway

Phocid seal pups must learn successful survival strategies, largely independently, following their abrupt weaning at a relatively young age. To explore the ontogeny of aquatic skills, space use and first‐year habitat choices made by harbor seals, pups (n = 30) were instrumented with satellite relay data loggers (SRDLs) in Svalbard, Norway in 2009 and 2010. Initially, the pups had small home ranges and showed rapid changes in their activity budgets and diving capabilities, displaying steep linear increases in diving depth and duration and in the amount of time spent diving. Most pups underwent an abrupt shift in movement patterns at ca. 50 d of age, which likely marked the end of the postweaning fast. Around this same time, the steep progression in diving performance slowed, though longer, deeper dives gradually became the norm. However, bottom time, ascent and descent rates, and postdive recovery times remained stable after the postweaning fast, suggesting that most aquatic skill acquisition was completed during the first months of life. Few clear effects of environmental variables such as upwelling phenomenon, which are known to influence the diving behavior of adults from the same population, were detected in the diving patterns of pups.     

Maternal attendance behaviour of sympatrically breeding Antarctic and subantarctic fur seals,Arctocephalus spp., at Macquarie Island

Maternal attendance behaviour was studied in Antarctic (Arctocephalus gazella) and subantarctic fur seals (Arctocephalus tropicalis) which breed sympatrically at subantarctic Macquarie Island. Data on attendance were obtained using telemetric methods. Both species undertook two types of foraging trips: overnight foraging trips which were of less than 1 day duration and occurred exclusively overnight, and extended foraging trips which lasted longer than 1 day. The mean duration of overnight foraging trips was 0.43 and 0.39 days, while the duration of extended foraging trips was 3.6 and 3.8 days in A. gazella and A. tropicalis, respectively. The duration of overnight and extended foraging trips did not differ significantly between species. Two types of shore attendance bouts that differed in duration were also observed in these species. Short attendance bouts lasted less than 0.9 days, while long attendance bouts lasted longer than 0.9 days. Short attendance bouts lasted 0.4 and 0.5 days, while long attendance bouts lasted 1.6 and 1.7 days in A. gazella and A. tropicalis, respectively, and did not differ significantly between species. The most significant differences between the attendance behaviour of both species was in the percentage of foraging time allocated to overnight foraging trips (15% and 25% in A. gazella and A. tropicalis, respectively), and the percentage of time spent ashore (30% and 38% in A. gazella and A. tropicalis, respectively). The nearness of pelagic waters to Macquarie Island is considered to be the main reason that lactating females are able to undertake overnight foraging trips. These trips may be used by females as a means of optimising the costs of fasting and nursing ashore. Females may be able to save energy by only nursing pups when milk transfer efficiencies are high, and reduce the time and energy costs of fasting ashore when milk transfer efficiency is low. Of the female A. gazella that still carried transmitters at the end of lactation, 83% continued regular attendance for between 21 and 150 days post-lactation (when data collection ceased). Overwintering of A. gazella females at breeding sites has not been previously reported in other populations. Accepted: 10 November 1998     

Diving Related Changes in the Blood Oxygen Stores of Rehabilitating Harbor Seal Pups (Phoca vitulina)

Harbor seal (Phoca vitulina) pups begin diving within hours of birth, stimulating the development of the blood oxygen (O₂) stores necessary to sustain underwater aerobic metabolism. Since harbor seals experience a brief nursing period, the early-life development of these blood O₂ stores is necessary for successful post-weaning foraging. If mothers and pups become prematurely separated, the pup may be transported to a wildlife rehabilitation center for care. Previous studies suggest that the shallow pools and lack of diving in rehabilitation facilities may lead to under-developed blood O₂ stores, but diving behavior during rehabilitation has not been investigated. This study aimed to simultaneously study the diving behaviors and blood O₂ store development of rehabilitating harbor seal pups. Standard hematology measurements (Hct, Hb, RBC, MCV, MCH, MCHC) were taken to investigate O₂ storage capacity and pups were equipped with time-depth recorders to investigate natural diving behavior while in rehabilitation. Linear mixed models of the data indicate that all measured blood parameters changed with age; however, when compared to literature values for wild harbor seal pups, rehabilitating pups have smaller red blood cells (RBCs) that can store less hemoglobin (Hb) and subsequently, less O₂, potentially limiting their diving capabilities. Wild pups completed longer dives at younger ages (maximum reported <25 days of age: 9 min) in previous studies than the captive pups in this study (maximum <25 days of age: 2.86 min). However, captivity may only affect the rate of development, as long duration dives were observed (maximum during rehabilitation: 13.6 min at 89 days of age). Further, this study suggests that there may be a positive relationship between RBC size and the frequency of long duration dives. Thus, rehabilitating harbor seal pups should be encouraged to make frequent, long duration dives to prepare themselves for post-release foraging.     

Inter-sexual differences in Antarctic fur seal pup growth rates: evidence of environmental regulation?

We investigated the variation of Antarctic fur seal (Arctocephalus gazella) pup growth rates in response to sex, breeding season and duration of both maternal foraging trip and attendance bouts. Data were collected during five consecutive rearing seasons at Cape Shirreff, the most important breeding colony in the South Shetland Islands, Antarctica. Our results showed significant interannual and sexual variations in pup growth rates. Male pups grew significantly faster than female pups during 2000, 2001, 2002 and 2004 seasons, whilst during 2003 no difference was found. The interannual variation in pup growth rates was correlated with the interannual fluctuation in maternal foraging trip and attendance bouts. There was a significant effect of pup sex and maternal foraging trip duration on pup growth rates, which varied between years having foraging trip duration a major effect during 2003, when females spent more time at sea and interestingly on that year there were no sexual differences in pup growth rates. The effect of attendance bout on pup growth rates was not significant. Diet analysis showed that Antarctic krill (Euphausia superba) was the most frequent prey item during the study period. Analysis of krill size distribution showed a significant difference in krill length, during 2003, when A. gazella preyed upon the smallest sizes of krill. In this study, sex was the most important factor on pup growth rates, but when prey availability seemed more limited, there are longer foraging trips and shorter attendance bouts the sex factor became less significant.     

Size and distribution of oxygen stores in harp and hooded seals from birth to maturity

Pinnipeds rely primarily on oxygen stores in blood and muscles to support aerobic diving; therefore rapid development of body oxygen stores (TBO₂) is crucial for pups to transition from nursing to independent foraging. Here, we investigate TBO₂ development in 45 harp (Pagophilus groenlandicus) and 46 hooded (Cystophora cristata) seals ranging in age from neonates to adult females. We found that hooded seal adults have the largest TBO₂ stores yet reported (89.5 ml kg⁻¹), while harp seal adults have values more similar to other phocids (71.6 ml kg⁻¹). In adults, large TBO₂ stores resulted from large blood volume (harp169, hood 194 ml kg⁻¹) and high muscle Mb content (harp 86.0, hood 94.8 mg g⁻¹). In contrast, pups of both species had significantly lower mass-specific TBO₂ stores than adults, and stores declined rather than increased during the nursing period. This decline was due to a reduction in mass-specific blood volume and the absence of an increase in the low Mb levels (harp 21.0, hood 31.5 mg g⁻¹). Comparisons with other phocid species suggests that the pattern of blood and muscle development in the pre- and post-natal periods varies with terrestrial period, and that muscle maturation rates may influence the length of the postweaning fast. However, final maturation of TBO₂ stores does not take place until after foraging begins.     

Biological and environmental drivers of energy allocation in a dependent mammal, the Antarctic fur seal pup

Little is known about how variation in the pattern and magnitude of parental effort influences allocation decisions in offspring. We determined the energy budget of Antarctic fur seal pups and examined the relative importance of timing of provisioning, pup traits (mass, condition, sex), and weather (wind chill and solar radiation) on allocation of energy obtained in milk by measuring milk energy intake, field metabolic rate (FMR), and growth rate in 48 Antarctic fur seal pups over three developmental stages (perinatal, premolt, and molt). The relative amount of milk energy used for growth was 59.1% ± 8.1% during the perinatal period but decreased to 23.4% ± 15.5% and 26.0% ± 13.9% during the premolt and molt. This decrease was associated with a greater amount of time spent fasting, along with an increase in pup activity while the mother was at sea foraging. Average daily milk intake, pup mass, and condition were all important in determining how much energy was available for growth, but the amount of energy obtained as milk was the single most important factor determining pup growth. While mean mass-specific FMR did not change with developmental stage (range = 1.74-1.77 mL O(2)/g/h), the factors that accounted for variation in FMR did. Weather (wind chill and solar radiation) and pup traits (mass and condition) influenced mass-specific FMR, but these impacts varied across development. This study provides information about the factors influencing how offspring allocate energy toward growth and maintenance and improves our predictions about how a changing environment may affect energy allocation in pups.     

Changes in plasma fatty acids indicate change in nutritional status in developing Weddell seal pups

The concentrations and the fatty acid compositions (weight percent) of three plasma lipid fractions (phospholipids, cholesteryl esters, and triacylglycerols) were examined in five Weddell seal pups through suckling, the post-weaning fast and periods of independent foraging. The principal plasma fatty acids in Weddell seal pups during suckling were the same as those previously reported for milk samples collected from lactating Weddell seals. In contrast to the stable milk fatty acid composition suggested by the previous study, significant changes in the fatty acid composition of plasma lipids occurred during suckling. These included an increase in the weight% of 20:4(n-6) and 18:0 in phospholipids, an increase of 20:4(n-6) in cholesteryl esters, and an increase of 20:1(n-9) and a decrease of 20:5(n-3) in triacylglycerols. Weaning and the subsequent fasting period were accompanied by a dramatic drop in the plasma concentration and 14:0 content of triacylglycerols and by marked decrease and increase in the weight% of 20:1(n-9) and 20:5(n-3), respectively, in this lipid fraction. Weight% of 14:0 in triacylglycerols and/or plasma concentration of triacylglycerols clearly distinguish post-weaning from suckling. However, fasting versus foraging pups could not be clearly distinguished. Received: 7 October 1996 / Accepted: 20 May 1997     

Predator‐induced modifications to diving behavior vary with foraging mode

Breath‐hold divers are strongly interacting species whose top–down influence on aquatic communities is shaped by factors governing their diving decisions. Although some of these factors (e.g. physiological constraints, energetic needs) have been scrutinized, the possibility that predation risk influences diving behavior has been largely overlooked, and no study to date has asked if anti‐predator responses by divers depend on foraging mode. We contrasted dive cycle changes by herbivorous dugongs Dugong dugon using two foraging tactics – cropping, which always permits anti‐predator vigilance, and excavation, which limits surveillance at depth – in response to temporal variation in tiger shark Galeocerdo cuvier abundance. Dugongs responded to increasing shark abundance (one component of predation risk) by diving more frequently without changing their surface times and thereby spending a greater proportion of time at the surface, but only while excavating. When threatened, in other words, excavating dugongs sacrificed foraging time at depth to facilitate shark detection. In contrast, cropping dugongs at risk from sharks were able to continue diving and foraging normally. By implication, future studies should consider the influence of predation risk on diving decisions, even by large‐bodied species, and the possibility that behavioral responses by divers to predators may vary with foraging mode.     

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.     

Remarkable development of diving performance and migrations of hooded seals (Cystophora cristata) during their first year of life

Newborn hooded seals (Cystophora cristata) have smaller weight-specific oxygen stores than adults, but nothing is known about how this affects their diving behaviour. Here, we present data on the diving behaviour and migrations of seven weaned hooded seal pups of the Greenland Sea stock during their first year of life, as collected by use of satellite telemetry. The pups started diving 1–2 days after tagging, and during a tracking period of 25–398 days they dispersed over vast areas of the Greenland and Norwegian Seas in a manner similar to adults. The initial development of diving depths and durations in April–May was rapid, and pups reached depths of >100 m and dived for >15 min within 3 weeks of age. During early summer (May–June) this development was temporarily discontinued, to be resumed throughout autumn and winter, during which time maximum depths and durations of >700 m and >30 min, respectively, were reached. Depths and durations were significantly related to age/season, location and time of day. The dive behaviour in early summer, with relatively shallow and short dives without diurnal variations, resembles that of adults and probably reflects the vertical distribution of prey rather than physiological constraints. Dives of pups were nevertheless shallower and shorter than those of adults, but relative to body mass both hooded seal pups and adults display a remarkable diving capacity which makes the species particularly suited for studies of defence mechanisms against hypoxia insult in mammals.     

The contrasting hidden consequences of parasitism: Effects of a hematophagous nematode (Uncinaria sp.) in the development of a marine mammal swimming behavior

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Foraging behavior of redheads (Aythya americana) wintering in Texas and Louisiana

Redheads, Aythya americana, concentrate in large numbers annually in traditional wintering areas along the western and northern rim of the Gulf of Mexico. Two of these areas are the Laguna Madre of Texas and Chandeleur Sound of Louisiana. We collected data on 54,340 activities from 103 redhead flocks in Texas and 51,650 activities from 57 redhead flocks in Louisiana. Males and females fed similarly, differing neither in levels of feeding (percent of all birds in flock that were feeding) (p>0.90) nor in percentages of birds feeding by diving, tipping, dipping, or gleaning from the surface (p>0.10). The foraging level of redheads in the upper Laguna Madre region was relatively constant throughout two winters. Foraging of redheads in early winter in Louisiana was significantly greater than redhead foraging in the upper Laguna Madre, but by late winter, foraging by redheads in Louisiana had declined to the same level as that shown by redheads foraging in the upper Laguna Madre. The overall foraging level of redheads from Chandeleur Sound was greater (41%) than that of redheads in the upper Laguna Madre (26%), yet it was quite similar to the 46% foraging level reported for redheads from the lower Laguna Madre. Redheads in the upper Laguna Madre region of Texas fed more by diving than did those in the Chandeleur Sound and the lower Laguna Madre. Diving increased in frequency in late winter. Greater reliance by redheads on diving in January and February indicates that the birds altered their foraging to feed in deeper water, suggesting that the large concentrations of redheads staging at this time for spring migration may have displaced some birds to alternative foraging sites. Our results imply that the most likely period for food resources to become limiting for wintering redheads is when they are staging in late winter.     

A chronological assessment of foraging activities of the Cotton Pigmy Goose,Nettapus coromandelianus Gmelin, 1789 at Dulahara pond,Ratanpur, Chhattisgarh,India

This study was planned to assess the foraging activity of ecological important avian species the N. coromandelianus at Dulahara pond of Ratanpur, Chhattisgarh. In present work we observed the effect of “time of the day” and “photoperiod” on daytime foraging activities of N. coromandelianus. The most applied foraging skills of N. coromandelianus were documented for two repetitive days each for the period of long days (March and April 2015) and short days (December 2015 and January 2016). One-way ANOVA (by SPSS 16.0) was employed to observe the effects of “photoperiod” and “time of the day” on daytime foraging pattern in order to the frequency of foraging techniques. The rhythms in daytime foraging movement were estimated by means of Cosinor rhythmometry at 24 h and 12 h. The current study clarified that the N. coromandelianus mostly applied three main techniques viz., head and neck dipping, diving and striking on which both the diving and striking were the most employed technique while the head and neck dipping was less employed technique during foraging. Moreover, the N. coromandelianus maximally involved for foraging activities during the short day than long day.     

Fasting metabolism in Antarctic fur seal (Arctocephalus gazella) pups

The metabolism of 52–73-day old Antarctic fur seal pups from Bird Island, South Georgia, was investigated during fasting periods of normal duration while their mothers were at sea foraging. Body mass decreased exponentially with pups losing 3.5–3.8% of body mass per day. Resting metabolic rate also decreased exponentially from 172–197 ml (O₂)·min⁻¹ at the beginning of the fast and scaled to M_b^0.74 at 2.3 times the level predicted for adult terrestrial mammals of similar size. While there was no significant sex difference in RMR, female pups had significantly higher (F_1,18=6.614, P<0.019) mass-specific RMR than male pups throughout the fasting period. Fasting FMR was also significantly (t₁₅=2.37, P<0.035) greater in females (823 kJ·kg⁻¹·d⁻¹) than males (686 kJ·kg⁻¹·d⁻¹). Average protein turnover during the study period was 19.3 g·d⁻¹ and contributed to 5.4% of total energy expenditure, indicating the adoption of a protein-sparing strategy with a reliance on primarily lipid catabolism for metabolic energy. This is supported by observed decreases in plasma BUN, U/C, glucose and triglyceride concentrations, and an increase in β-HBA concentration, indicating that Antarctic fur seals pups adopt this strategy within 2–3 days of fasting. Mean RQ also decreased from 0.77 to 0.72 within 3 days of fasting, further supporting a rapid commencement of protein-sparing. However, RQ gradually increased thereafter to 0.77, suggesting a resumption of protein catabolism which was not substantiated by changes in plasma metabolites. Female pups had higher TBL (%) than males for any given mass, which is consistent with previous findings in this and other fur seal species, and suggests sex differences in metabolic fuel use. The observed changes in plasma metabolites and protein turnover, however, do not support this.     

Brain size in neonatal and adult Weddell seals: Costs and consequences of having a large brain

Little is known about the ontogeny of brain size in pinnipeds despite potential functional implications of brain substrate (glucose, oxygen) requirements for diving, fasting, growth, and lactation strategies. We measured brain mass (brM) and cranial capacity (CC) in newborn and adult Weddell seals. Neonatal Weddell seals had brM that represented ~70% of adult brM. Weddell seals have the largest neonatal brain, proportional to adult brain, reported for any mammal to date, which is remarkable considering the relatively small size of Weddell seal pups at birth (6%–7% of maternal body mass) compared to neonates of other highly precocial mammals. Provision of sufficient glucose to maintain the large, well‐developed brain of the neonatal Weddell seal has a nontrivial metabolic cost to both pup and mother. We therefore hypothesize that this phenomenon must have functional significance, such as allowing pups to acquire complex under‐ice navigation skills during the period of maternal attendance.     

Shifts in specific serum protein concentrations: apolipoprotein-B and albumin in the southern elephant seal during the breeding and molting periods in Antarctica

Serum protein values in Southern elephant seals (Mirounga leonina) were analyzed during the breeding and molting periods at the 25 de Mayo Island (King George Island), Antarctica, during the 1999/2000 and 2000/2001 field seasons, in order to study the reaction of adults to fasting, and of pups to nursing and fasting. The following analyses were carried out: Total Proteins (TP) and Albumin (Alb) were analyzed by a colorimetric technique, and Apolipoprotein B (Apo-B) was determined by the immunodiffusion technique on agarose gel plates. The general ranges of average values for total proteins and specific fractions during the breeding period were: TP (g/dl) = 5.12–9.83; Alb (g/dl) = 1.72–5.71 and Apo-B (mg/dl) = 10–266, and during the post-breeding period: TP = 4.85–9.45; Alb = 2.06–4.20 and Apo-B = 13–232. The essential obtained data were: (a) fasting does not impact adult males, except for a significant decrease of Apo-B during breeding; (b) fasting does not impact adult females, except for a significant decrease in TP during molting; (c) suckling increases significantly TP and Apo-B in pups; (d) post-weaning fast decreases significantly all measured serum components in pups. We can conclude that the adults are adapted to long term fasting, without any metabolic downside they maintain homeostasis during this period, as shown by the serum data. The pups clearly react to both suckling and the post-weaning fast, which are periods driven almost exclusively by lipid chemistry. These impacts may be seen in the chemistry of serum proteins.     

Body condition influences ontogeny of foraging behavior in juvenile southern elephant seals

Ontogeny of diving and foraging behavior in marine top predators is poorly understood despite its importance in population recruitment. This lack of knowledge is partly due to the difficulties of monitoring juveniles in the wild, which is linked to high mortality early in life. Pinnipeds are good models for studying the development of foraging behaviors because juveniles are large enough to robustly carry tracking devices for many months. Moreover, parental assistance is absent after a juvenile departs for its first foraging trip, minimizing confounding effects of parental input on the development of foraging skills. In this study, we tracked 20 newly weaned juvenile southern elephant seals from Kerguelen Islands for up to 338 days during their first trip at sea following weaning. We used a new generation of satellite relay tags, which allow for the transmission of dive, accelerometer, and location data. We also monitored, at the same time, nine adult females from the colony during their post‐breeding trips, in order to compare diving and foraging behaviors. Juveniles showed a gradual improvement through time in their foraging skills. Like adults females, they remarkably adjusted their swimming effort according to temporal changes in buoyancy (i.e., a proxy of their body condition). They also did not appear to exceed their aerobic physiological diving limits, although dives were constrained by their smaller size compared to adults. Changes in buoyancy appeared to also influence their decision to either keep foraging or return to land, alongside the duration of their haul outs and choice of foraging habitat (oceanic vs. plateau). Further studies are thus needed to better understand how patterns in juveniles survival, and therefore elephant seal populations, might be affected by their changes in foraging skills and changes in their environmental conditions.     

Ontogeny of diving behaviour in the Australian sea lion: trials of adolescence in a late bloomer

1. Foraging behaviours of the Australian sea lion (Neophoca cinerea) reflect an animal working hard to exploit benthic habitats. Lactating females demonstrate almost continuous diving, maximize bottom time, exhibit elevated field metabolism and frequently exceed their calculated aerobic dive limit. Given that larger animals have disproportionately greater diving capabilities, we wanted to examine how pups and juveniles forage successfully. 2. Time/depth recorders were deployed on pups, juveniles and adult females at Seal Bay Conservation Park, Kangaroo Island, South Australia. Ten different mother/pup pairs were equipped at three stages of development (6, 15 and 23 months) to record the diving behaviours of 51 (nine instruments failed) animals. 3. Dive depth and duration increased with age. However, development was slow. At 6 months, pups demonstrated minimal diving activity and the mean depth for 23-month-old juveniles was only 44 +/- 4 m, or 62% of adult mean depth. 4. Although pups and juveniles did not reach adult depths or durations, dive records for young sea lions indicate benthic diving with mean bottom times (2.0 +/- 0.2 min) similar to those of females (2.1 +/- 0.2 min). This was accomplished by spending higher proportions of each dive and total time at sea on or near the bottom than adults. Immature sea lions also spent a higher percentage of time at sea diving. 5. Juveniles may have to work harder because they are weaned before reaching full diving capability. For benthic foragers, reduced diving ability limits available foraging habitat. Furthermore, as juveniles appear to operate close to their physiological maximum, they would have a difficult time increasing foraging effort in response to reductions in prey. Although benthic prey are less influenced by seasonal fluctuations and oceanographic perturbations than epipelagic prey, demersal fishery trawls may impact juvenile survival by disrupting habitat and removing larger size classes of prey. These issues may be an important factor as to why the Australian sea lion population is currently at risk.     

Activity‐specific metabolic rates for diving,transiting, and resting at sea can be estimated from time–activity budgets in free‐ranging marine mammals

Time and energy are the two most important currencies in animal bioenergetics. How much time animals spend engaged in different activities with specific energetic costs ultimately defines their likelihood of surviving and successfully reproducing. However, it is extremely difficult to determine the energetic costs of independent activities for free‐ranging animals. In this study, we developed a new method to calculate activity‐specific metabolic rates, and applied it to female fur seals. We attached biologgers (that recorded GPS locations, depth profiles, and triaxial acceleration) to 12 northern (Callorhinus ursinus) and 13 Antarctic fur seals (Arctocephalus gazella), and used a hierarchical decision tree algorithm to determine time allocation between diving, transiting, resting, and performing slow movements at the surface (grooming, etc.). We concomitantly measured the total energy expenditure using the doubly‐labelled water method. We used a general least‐square model to establish the relationship between time–activity budgets and the total energy spent by each individual during their foraging trip to predict activity‐specific metabolic rates. Results show that both species allocated similar time to diving (~29%), transiting to and from their foraging grounds (~26–30%), and resting (~8–11%). However, Antarctic fur seals spent significantly more time grooming and moving slowly at the surface than northern fur seals (36% vs. 29%). Diving was the most expensive activity (~30 MJ/day if done non‐stop for 24 hr), followed by transiting at the surface (~21 MJ/day). Interestingly, metabolic rates were similar between species while on land or while slowly moving at the surface (~13 MJ/day). Overall, the average field metabolic rate was ~20 MJ/day (for all activities combined). The method we developed to calculate activity‐specific metabolic rates can be applied to terrestrial and marine species to determine the energetic costs of daily activities, as well as to predict the energetic consequences for animals forced to change their time allocations in response to environmental shifts.