The thermal physiology of most birds and mammals is characterised by considerable spatial and temporal variation in body temperature. Body temperature is, therefore, a key parameter in physiological, behavioural and ecological research. Temperature measurements on freely moving or free-ranging animals in the wild are challenging but can be undertaken using a range of techniques. Internal temperature may be sampled using thermometry, surgically implanted loggers or transmitters, gastrointestinal or non-surgically placed devices. Less invasive approaches measure peripheral temperature with subcutaneous passive integrated transponder tags or skin surface-mounted radio transmitters and data loggers, or use infrared thermography to record surface temperature. Choice of technique is determined by focal research question and region of interest that reflects appropriate physiological or behavioural causal mechanisms of temperature change, as well as welfare and logistical considerations. Particularly required are further studies that provide opportunities of continuously sampling from multiple sites from within the body. This will increase our understanding of thermoregulation and temperature variation in different parts of the body and how these temperatures may change in response to physiological, behavioural and environmental parameters. Technological advances that continue to reduce the size and remote sensing capability of temperature recorders will greatly benefit field research.
Summary 1.Measures of the field metabolic rate of marine mammals are extremely difficult to make but they are essential for assessing the impacts of marine mammals on prey populations, and for assessing dive performance in relation to aerobic limits. 2. The doubly labelled water (DLW) method is an isotope-based technique for the estimation of the CO 2 production, and hence energy expenditure, of free-living animals. Estimates of field metabolic rate (FMR) from DLW in pinnipeds to date are extremely high and at the upper range for most mammals. DLW has previously been validated in pinnipeds but logistical difficulties meant for these validations were less than ideal, and it has been hypothesised that DLW may overestimate FMR in these animals. 3. To test this hypothesis, we used DLW and simultaneous open-flow respirometry to determine the daily energy expenditures (DEE) of wild grey seals ( Halichoerus grypus ) held temporarily in a captive facility, during 4-5 days of simulated foraging at sea. Comparing DEE from DLW and respirometry, we found that DLW predicted DEE accurately (average difference between the two estimates was 0·7% SD = 17% n = 31), but as with validations of other species there was a large range of individual errors (from -39% to +44%). 4. The results dispel the doubts surrounding the use of DLW as a field method for estimating energy expenditure in grey seals, and by implication other pinnipeds, and simultaneously open a series of questions about their ability to maintain surprisingly high metabolic rates for protracted periods. 5. We make a number of recommendations for future studies of pinniped FMR using DLW. We suggest use of the Speakman two-pool calculation will be most appropriate. Studies should aim for enrichment levels as high as economically feasible but to at least 150 p.p.m. above background for the O 2 isotope. Measurement periods should be extended between one and two half-lives (5-10 days for a typical foraging seal). We also encourage the calculation and presentation of estimates of precision in estimates of FMR.
Southern elephant seals (Mirounga leonina) are known to move and aggregate while molting, but little is known about their behavior on land during this time. In this study, 60 adult females were monitored (23 with GPS tags) during four molting seasons, between 2012 and 2016 at Kerguelen Archipelago, Indian Ocean. Population surveys were recorded each year (N = 230 daily counts), and habitat use was analyzed in relation to the stage of the molt and local weather. Based on stage of molt, habitat use, and movements on land, we classified the molt of elephant seals into three phases: (1) a “search phase” at the initial stage of molt when grass and wallow habitats were used and characterized by greater mean distances travelled on land per day compared with the two other phases; (2) a “resident phase”: during initial and mid‐stage of molt when animals were found in grass and wallow habitats but with less distance moved on land; and (3) a “termination phase” at the final stage of molt where grass and beach habitats were occupied with no change in distances. Windchill and solar radiation influenced individual distances moved per day (mean 590 ± 237.0 m) at the mid‐ and final stage of molt such that animals travelled greater distances on days of low windchill or high solar radiation. Individual variation in distance moved and relative habitat use were also linked to body mass index (BMI) at arrival on the colony, as females with higher BMI moved less and preferred beach habitat. Moreover, the individual rate of molt increased with the use of wallows. Aggregation rate tended to be negatively correlated with distances moved. We therefore suggest that individuals face an energetic trade‐off while molting, balancing energy expenditure between movement and thermoregulation.
SUMMARY The duration of breath-hold dives and the available time for foraging in submerged prey patches is ultimately constrained by oxygen balance. There is a close relationship between swim speed and oxygen utilisation, so it is likely that breath-holding divers optimise their speeds to and from the feeding patch to maximise time spent feeding at depth. Optimal foraging models suggest that transit swim speed should decrease to minimum cost of transport (MCT) speed in deeper and longer duration dives. Observations also suggest that descent and ascent swimming mode and speed may vary in response to changes in buoyancy. We measured the swimming behaviour during simulated foraging of seven captive female grey seals (two adults and five pups). Seals had to swim horizontally underwater from a breathing box to a submerged automatic feeder. The distance to the feeder and the rate of prey food delivery could be varied to simulate different feeding conditions. Diving durations and distances travelled in dives recorded during these experiments were similar to those recorded in the wild. Mean swim speed decreased significantly with increasing distance to the patch, indicating that seals adjusted their speed in response to travel distance, consistent with optimality model predictions. There was, however, no significant relationship between the transit swim speeds and prey density at the patch. Interestingly,all seals swam 10–20% faster on their way to the prey patch compared to the return to the breathing box, despite the fact that any effect of buoyancy on swimming speed should be the same in both directions. These results suggest that the swimming behaviour exhibited by foraging grey seals might be a combination of having to overcome the forces of buoyancy during vertical swimming and also of behavioural choices made by the seals.
Unlike most phocid species (Phocidae), Mirounga leonina (southern elephant seals) experience a catastrophic moult where they not only replace their hair but also their epidermis when ashore for approximately one month. Few studies have investigated behavioural and physiological adaptations of southern elephant seals during the moult fast, a particularly energetically costly life cycle's phase.Recording heart rate is a reliable technique for estimating energy expenditure in the field. For the first time, subcutaneous heart rate data loggers were successfully implanted during the moult in two freeranging southern elephant seals over 3 to 6 days. No substantial postoperative complications were encountered and consistent heart rate data were obtained. This promising surgical technique opens new opportunities for monitoring heart rate in phocid seals.
While endotherms can rely on their insulation to reduce heat loss to adapt to cold environments, renewing of fur during molt impairs insulation while they have to perfuse the periphery to support epidermal tissues. The southern elephant seal Mirounga leonina undertakes an annual catastrophic molt while fasting on land in a wet, windy and cold environment. However, southern elephant seals show characteristic aggregation patterns that are predicted to reduce high metabolic costs during the molt. Between 2012 and 2016, 59 female elephant seals were tracked on land during their molt to study their aggregation behavior in relation to molt stage, habitat type and local weather conditions. Infrared thermography and stomach temperature loggers were used to observe variation in body surface and internal temperature in relation to molt stage and aggregation behavior. We found that thermal constraints varied during the molt, with a peak in surface temperature during the mid-stage of the molt. Wallows (mud pools) appear as favorable habitat to aggregate while molting. Indeed, wallows offered a warmer microclimate with greater ground temperature and lower wind speed. Moreover, there was a greater proportion of aggregated seals and larger group size in wallows. These aggregation patterns in wallows were influenced by local weather such that a greater proportion of seals were located in the center of the aggregation, and larger group size occurred during days of unfavorable meteorological conditions. We also observed a higher proportion of seals at mid-stage of molt amongst aggregated seals compared to isolated individuals. This aggregation behavior may reduce the cost of thermogenesis as surface body temperature and stomach temperature were cooler by 1.0°C and 1.5°C, respectively, in aggregated compared to isolated seals. As a consequence, huddling behavior may be thermally advantageous for female southern elephant seals during the molt.
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