Effects of capture stress on free-ranging, reproductively active male Weddell seals

Physiological stress responses to capture may be an indicator of welfare challenges induced by animal handling. Simultaneously, blood chemistry changes induced by stress responses may confound experimental design by interacting with the biological parameters being measured. Cortisol elevation is a common indicator of stress responses in mammals and reproductive condition can profoundly influence endocrine response. We measured changes in blood cortisol and testosterone induced by handling reproductively active male Weddell seals (Leptonychotes weddellii) early and late in the breeding season. Weddell seals have the highest resting cortisol levels of all mammals yet showed a clear, prolonged elevation in cortisol in response to capture. Responses were similar when first caught and when caught a second time, later in the breeding season. Baseline testosterone levels declined over the breeding season but were not altered by capture. Administering a light dose of diazepam significantly ameliorated the cortisol response of handled animals without affecting testosterone levels. This may be an effective way of reducing acute capture stress responses. Male breeding success in years males were handled was no different to the years they were not, despite the acute capture response, suggesting no long-term impact of handling on male reproductive output.     

A method for reconstructing three-dimensional dive profiles of marine mammals using geomagnetic intensity data: results from two lactating Weddell seals

The under-ice behavior of two free-ranging female Weddell seals (Leptonychotes weddellii) was studied using geomagnetic, acceleration and velocity sensors at Big Razorback Island in McMurdo Sound, Antarctica. The seals' body angle and posture were calculated from the acceleration data and the heading from the geomagnetic intensity data. Together with swim speed, the seals' three-dimensional underwater dive path, heading and even posture were reconstructed for each dive. Each instrument was deployed for 2 days, during which time these females made multiple, deep (₞ m) dives, with average maximum depths of 236ᆯ m (n=4) and 244끁 m (n=40). Each seal appeared to choose a particular heading on which to descend. These headings were significantly different between seals and bouts (Watson's U² test, P<0.05). These new instruments and methodologies are shown to provide valuable information on the fine-scale and complex movements of diving animals.     

Hydrographic influences on the summer dive behaviour of Weddell seals (Leptonychotes weddellii) in Atka Bay, Antarctica

In order to gain insights into species-level behavioural responses to the physical environment, it is necessary to obtain information from various populations and at all times of year. We analysed the influences of physical environmental parameters on the mid-summer dive behaviour of Weddell seals (Leptonychotes weddellii) from a little-known population at Atka Bay, Antarctica. Dive depth distributions followed a typical bimodal pattern also exhibited by seals from other populations and seals targeted both shallow water layers of <50 m and depths near the seafloor. Increased stratification of temperature layers within the water column resulted in increased forage efforts by the seals through relatively high numbers of dives to the seafloor, as well as forage effort associated with shallow dives. We interpret these behavioural responses to be due to increased water temperature stratification resulting in the concentration of prey species in particular depth layers.     

The tide as zeitgeber for Weddell seals

The haul-out and underwater activity patterns of five Weddell seals were investigated in relation to the tide. Electronic recorders measured the seals' dive depth and internal body temperature simultaneously. Diving and temperature data were analysed with the help of time series analyses tools. We identified a half-day (semicircadian) rhythm in underwater activity and body temperature. A close correlation was found between the rhythmicity of the tide and the timing of the seals' underwater activity and oscillations of their body temperature. We propose that the tide is the principal zeitgeber for the Weddell seals' ultradian cycle of behaviour during the austral summer. Received: 25 August 1997 / Accepted: 17 May 1998     

Blood rheology of Weddell seals and bowhead whales

Red blood cell (RBC) aggregation and blood viscosity are important determinants of in vivo blood flow dynamics and, in marine mammals, these parameters may impact diving physiology by altering blood oxygen delivery during the diving response. Weddell seals are superb divers and exhibit age-related patterns in blood oxygen chemistry and diving ability. By contrast, bowhead whales are not long duration divers, and little is known of their blood properties relative to diving. The present study was designed to compare rheological characteristics of blood from Weddell seal pups, Weddell seal adults, and from adult bowhead whales: blood viscosity and RBC aggregation in plasma and in polymer solutions (i.e., RBC "aggregability") were measured. Salient findings included: (1) significant 4- to 8-fold greater aggregation in blood from adult seals compared with pups and human subjects; (2) 2-to 8-fold greater aggregation in bowhead whale blood compared with human blood; (3) compared to human red cells, enhanced RBC aggregability of RBC from adult seals and whales as determined by their greater aggregation in polymer solutions; (4) increasing RBC aggregation and aggregability of seal pup blood over a seven day period following birth; (5) significantly greater blood viscosity for adult seals compared with pups at both native and standardized hematocrits. These results indicate that, for both species, hemorheological parameters differ markedly from those of humans, and suggest progressive changes with seal age; the physiological implications of these differences have yet to be fully defined.     

Variation in probability of first reproduction of Weddell seals

1. For many species, when to begin reproduction is an important life-history decision that varies by individual and can have substantial implications for lifetime reproductive success and fitness. 2. We estimated age-specific probabilities of first-time breeding and modelled variation in these rates to determine age at first reproduction and understand why it varies in a population of Weddell seals in Erebus Bay, Antarctica. We used multistate mark-recapture modelling methods and encounter histories of 4965 known-age female seals to test predictions about age-related variation in probability of first reproduction and the effects of annual variation, cohort and population density. 3. Mean age at first reproduction in this southerly located study population (7.62 years of age, SD=1.71) was greater than age at first reproduction for a Weddell seal population at a more northerly and typical latitude for breeding Weddell seals (mean=4-5 years of age). This difference suggests that age at first reproduction may be influenced by whether a population inhabits the core or periphery of its range. 4. Age at first reproduction varied from 4 to 14 years, but there was no age by which all seals recruited to the breeding population, suggesting that individual heterogeneity exists among females in this population. 5. In the best model, the probability of breeding for the first time varied by age and year, and the amount of annual variation varied with age (average variance ratio for age-specific rates=4.3%). 6. Our results affirmed the predictions of life-history theory that age at first reproduction in long-lived mammals will be sensitive to environmental variation. In terms of life-history evolution, this variability suggests that Weddell seals display flexibility in age at first reproduction in order to maximize reproductive output under varying environmental conditions. Future analyses will attempt to test predictions regarding relationships between environmental covariates and annual variation in age at first reproduction and evaluate the relationship between age at first reproduction and lifetime reproductive success.     

Serum chemistry and antibodies against pathogens in antarctic fur seals, Weddell seals, crabeater seals, and Ross seals

Information on health parameters, such as antibody prevalences and serum chemistry that can reveal exposure to pathogens, disease, and abnormal physiologic conditions, is scarce for Antarctic seal species. Serum samples from Antarctic fur seals (Arctocephalus gazella, n=88) from Bouvet?ya (2000-2001 and 2001-2002), and from Weddell seals (Leptonychotes weddellii, n=20), Ross seals (Ommatophoca rossii, n=20), and crabeater seals (Lobodon carcinophagus, n=9) from the pack-ice off Queen Maud Land, Antarctica (2001) were analyzed for enzyme activity, and concentrations of protein, metabolites, minerals, and cortisol. Adult Antarctic fur seal males had elevated levels of total protein (range 64-99 g/l) compared to adult females and pups (range 52-79 g/l). Antarctic fur seals had higher enzyme activities of creatine kinase, lactate dehydrogenase, and amylase, compared to Weddell, Ross, and crabeater seals. Antibodies against Brucella spp. were detected in Weddell seals (37%), Ross seals (5%), and crabeater seals (11%), but not in Antarctic fur seals. Antibodies against phocine herpesvirus 1 were detected in all species examined (Antarctic fur seals, 58%; Weddell seals, 100%; Ross seals, 15%; and crabeater seals, 44%). No antibodies against Trichinella spp., Toxoplasma, or phocine distemper virus (PDV) were detected (Antarctic fur seals were not tested for PDV antibodies). Antarctic seals are challenged by reduced sea ice and increasing temperatures due to climate change, and increased anthropogenic activity can introduce new pathogens to these vulnerable ecosystems and represent a threat for these animals. Our data provide a baseline for future monitoring of health parameters of these Antarctic seal species, for tracking the impact of environmental, climatic, and anthropogenic changes in Antarctica over time.     

A safe and practical inhalation anaesthesia for Weddell seals

Nine adult female Weddell seals were anaesthetized during the breeding season using the inhalation anaesthetic sevoflurane. The seals were captured with a head bag. Sevoflurane was administered and vaporized into the bag. A dose of 5 ml sevoflurane induced narcosis 1 min after application. Anaesthesia was maintained by multiple follow-up doses of 2 ml at an interval of 2 min. The average duration for complete recovery was 11Dž min after the terminal application. Sevoflurane is a reliable volatile anaesthetic that requires a minimum of equipment even under the low temperature regime of Antarctica.     

Through-ice communication by Weddell seals may not be practicable

Possible communication between territorial male Weddell seals (Leptonychotes weddellii) under the ice with females on the ice was investigated. In-air and underwater recordings of underwater calls were made at three locations near Davis, Antarctica. Most underwater calls were not detectable in air, often because of wind noise. In-air call amplitudes of detectable calls ranged from 32–74 dB re. 20 Pa at 86 Hz down to 4–38 dB re. 20 Pa at 3.6 kHz. Most of these would be audible to humans. Only 26 of 582 amplitude measurements (from 230 calls) ranged from 5 dB to a maximum of 15 dB above the minimum harbour-seal (Phoca vitulina) in-air detection threshold. Seals on the ice could likely hear a few very loud underwater calls but only if the caller was nearby and there were no wind noises. The low detectability of underwater calls in air likely precludes effective communication between underwater seals and those on the ice.     

Movements of adult female Weddell seals during the winter months

The focus of this study was the distribution of adult female Weddell seals during winter at the Vestfold Hills. Satellite tracking of Weddell seals had never been done before at this location. Hence, this was a pilot study to evaluate the following methods. We attached satellite transmitters to the lower back, where there was least potential to change the seals’ behaviour or to damage instruments on the ice. Location data were obtained only where the seals hauled out, not necessarily where they were feeding. All locations were within the area of fast-ice that was associated with the Vestfold Hills. There were gaps of up to 30 days in the location data sets. Each instrument (n=3) remained attached and functioning for ca. 6 months. During that time, two of the three seals hauled out within small areas adjacent to, or nearby, open water. The same seals hauled out sporadically. We inferred that these seals foraged offshore whilst returning to fast-ice to rest. If Weddell seals forage beneath dynamic ice but return to stable ice as their preferred resting substrate, then evidence of haulout sites will always be a biased measure of foraging range. Tracking seals in the water may be possible using alternative placement of transmitters. However, there is potential for instruments to interfere with movement (breathing and prey capture). For this reason, we recommend a combination of sensors, diet and tracking haulout sites to research winter foraging.     

Immobilisation of free-living Weddell seals Leptonychotes weddellii using midazolam and isoflurane

Eleven lactating female Weddell seals were immobilised using inhaled isoflurane and oxygen, having initially been sedated using an intramuscular injection of midazolam. The seals were selected from colonies in Long Fjord, East Antarctica. Isoflurane was delivered using a precision, out-of-circle vaporiser in a portable, heated, semi-closed circle system anaesthetic machine. Induction time (time from injection of midazolam to detected maximal effect of midazolam) ranged from 12 min to 29 min. The maximal effect of midazolam was assessed as being either moderate sedation (n=9) or heavy sedation (n=2), and the maximal effect of inhaled isoflurane and oxygen was assessed as being light anaesthesia (n=11). The level of chemical restraint achieved using this combination allowed attachment of heart rate monitoring units and collection of biological samples. Recovery time ranged from 1 min to 11 min. The anaesthetic regime proved a practical, safe and reliable method for the immobilisation of lactating Weddell seals under conditions of low environmental temperature.     

Annual and diurnal variations in the underwater vocalizations of Weddell seals

Summary Weddell seal vocalizations were recorded once a fortnight throughout 1984 at a major pupping site near Davis, Antarctica. Few vocalizations were heard between January and June. The number of vocalizations recorded increased from July to a peak in November and early December during the breeding period and then fell off rapidly through December. The increase in the number of seal vocalizations was correlated with a build up in seal numbers but some of the increase may be explained by an increased rate of vocalization by males.     

Spatial utilisation of fast-ice by Weddell seals Leptonychotes weddelli during winter

This study describes the distribution of Weddell seals Leptonychotes weddelli in winter (May–September 1999) at the Vestfold Hills, in Prydz Bay, East Antarctica. Specifically, we describe the spatial extent of haul-out sites in shore–fast sea-ice, commonly referred to as fast-ice. As winter progressed, and the fast-ice grew thick (ca 2 m), most of the inshore holes closed over, and the seals' distribution became restricted to ocean areas beyond land and islands. Using observations from the end of winter only, we fitted Generalised Additive Models (GAMs) to generate resource selection functions, which are models that yield values proportional to the probability of use. The models showed that seal distribution was defined mainly by distance to ice-edge and distance to land. Distance to ice-bergs was also selected for models of some regions. We present the results as maps of the fitted probability of seal presence, predicted by the binomial GAM for offshore regions, both with and without autocorrelation terms. The maps illustrate the expected distribution encompassing most of the observed distribution. On this basis, we hypothesise that propensity for the fast-ice to crack is the major determinant of Weddell seal distribution in winter. Proximity to open water and pack-ice habitats could also influence the distribution of haul-out sites in fast-ice areas. This is the first quantitative study of Weddell seal distribution in winter. Potential for regional variation is discussed.     

Spatial variation in age-specific probabilities of first reproduction for Weddell seals

Spatial variation in vital rates can affect the dynamics and persistence of a population. We evaluated the prediction that age-specific probabilities of survival and first reproduction for Weddell seals would vary as a function of birth location in Erebus Bay, Antarctica. We used multi-state mark–resight models and 25 years of data to estimate demographic rates for female seals. We predicted that probabilities of survival and first reproduction would be higher for seals born at near-shore colonies or more southerly-located colonies with consistent ice conditions. Contrary to predictions, results revealed higher age-specific probabilities of first reproduction at offshore colonies relative to near-shore colonies and no spatial variation in survival rates. For 7-year old females (average age at 1st reproduction=7.6 years old) born at offshore colonies to mothers aged 10.8 years (average maternal age), probability of first reproduction was 0.43 (SE=0.07), whereas probability of first reproduction for females born at near-shore colonies was 0.30 (SE=0.05) based on estimates from our top-ranked model. Breeding probabilities following first reproduction were also higher at offshore colonies. Thus, our results (1) provide evidence of spatial variation in breeding probabilities, (2) reveal the importance of birth location on a female's vital rates, and (3) suggest that the effect persisted for many years. Birth-colony effects may be attributed to spatial variation in prey availability, or to heterogeneity in female quality in this population. If females who are superior competitors consistently chose offshore colonies for pupping, pups born at these locations may have inherited those superior qualities and displayed higher probabilities of first reproduction, relative to seals born at other colonies. Further research into physical or food-related differences among colonies may offer insight into spatial variation in breeding probabilities documented in this paper.     

Accuracy of satellite positions from free-ranging grey seals using ARGOS

We have undertaken an analysis of the locations derived from an ARGOS system using satellite relay data loggers mounted on free-ranging grey seals (Halichoerus grypus Fab.) in the Gulf of Bothnia, between Sweden and Finland. The accuracy of the classes reported within this study is similar to that reported by Service ARGOS and we suggest that it would be beneficial to receive the A and B location classes. Locations-derived location classes 0 and A can be used for studies involving long-range movements, 10 km or more, but would not be suitable for fine-scale analyses.     

Influence of maternal mass and condition on energy transfer in Weddell seals

1. Environmental variation influences food abundance and availability, which is reflected in the reproductive success of top predators. We examined maternal expenditure, offspring mass and condition for Weddell seals in 2 years when individuals exhibited marked differences in these traits. 2. For females weighing > or = 355 kg there was a positive relationship between maternal post-partum mass (MPPM) and lactation length, but below this there was no relationship, suggesting that heavier females were able to increase lactation length but lighter females were restricted to a minimum lactation period of 33 days. 3. Overall, females were heavier in 2002, but in 2003 shorter females were lighter than similar-sized females in 2002 suggesting that the effects of environmental variability on foraging success and condition are more pronounced in smaller individuals. 4. There was no relationship between MPPM and pup birth mass, indicating pre-partum investment did not differ between years. However, there was a positive relationship between MPPM and pup mass gain. Mass and energy transfer efficiency were 10.2 and 5.4% higher in 2002 than 2003, which suggests costs associated with a putatively poor-resource year were delayed until lactation. 5. Heavier females lost a higher proportion of mass during lactation in both years, so smaller females may not have been able to provide more to their offspring to wean a pup of similar size to larger females. 6. MPPM had only a small influence on total body lipid; therefore, regardless of mass, females had the same relative body composition. Females with male pups lost a higher percentage of lipid than those with female pups, but by the end of lactation female pups had 4.5% higher lipid content than males. 7. It appears that for Weddell seals the consequences of environmentally induced variation in food availability are manifested in differences in maternal mass and expenditure during lactation. These differences translate to changes in pup mass and condition at weaning with potential consequences for future survival and recruitment.     

Evaluation of reproductive costs for Weddell seals in Erebus Bay, Antarctica

1. Organisms balance current reproduction against future survival and reproduction, which results in life-history trade-offs. These trade-offs are also known as reproductive costs and may represent significant factors shaping life-history strategy for many species. 2. Using multistate mark-resight models and 26 years of mark-resight data (1979-2004), we estimated the costs of reproduction to survival and reproductive probabilities for Weddell seals in Erebus Bay, Antarctica and evaluated whether this species either conformed to the 'prudent parent' reproductive strategy predicted by life-history theory for long-lived mammals or alternatively, incurred costs to survival in order to reproduce in a variable environment (flexible-strategy hypothesis). 3. Results strongly supported the presence of reproductive costs to survival (mean annual survival probability was 0.91 for breeders vs. 0.94 for nonbreeders), a notable difference for a long-lived mammal, demonstrating that investment in reproduction does result in a cost to survival for Weddell seals, contrary to the prudent parent hypothesis. 4. Reproductive costs to subsequent reproductive probabilities were also present for first-time breeders (mean probability of breeding the next year was 31.3% lower for first-time breeders than for experienced breeders), thus supporting our prediction of the influence of breeding experience. 5. We detected substantial annual variation in survival and breeding probabilities. Breeding probabilities were negatively influenced by summer sea-ice extent, whereas weak evidence suggested that survival probabilities were affected more by winter sea-ice extent, and the direction of this effect was negative. However, a model with annual variation unrelated to any of our climate or sea-ice covariates performed best, indicating that further study will be needed to determine the appropriate mechanism or combination of mechanisms underlying this annual variation.     

Viral and bacterial serology of six free-ranging bearded seals Erignathus barbatus

Serum or heparinized plasma samples were obtained from 3 male (2 adult and 1 weaned calf) and 3 adult female free-ranging bearded seals Erignathus barbatus in May of 1994, 1995, or 1996. Blood samples were obtained from animals taken in subsistence hunts near St. Lawrence Island, Alaska and screened for antibodies to a suite of bacteria and viruses potentially pathogenic for pinnipeds and/or humans. No samples had detectable antibodies to Brucella spp., Phocine distemper virus, influenza A virus or caliciviruses (San Miguel sea lion virus strains 1, 2, and 4 to 13, vesicular exanthema of swine serotypes A48, B51, C52, D53, E54, F55, G55, H54, 155, J56, K54, 1934B, and Tillamook and Walrus calicivirus). One seal had a low titer of 100 to Leptospira interrogans serovar