Erling S. Nordøy

Distribution and diving behaviour of crabeater seals (Lobodon carcinophagus) off Queen Maud Land

Eight crabeater seals (Lobodon carcinophagus) (three females, five males), ranging in body mass between 125 and 220 kg, were captured off Queen Maud Land (70–72°S, 7–16°W) during the last week of February, just after moulting, and tagged with Argos satellite-linked dive recorders to provide data on location and diving depth and duration. During the first few weeks of March the seals were moving in the pack ice along the continental shelf edge, close to the coast of Queen Maud Land. In April and May, when the pack ice extended northwards, most of the seals moved north, one reaching 63°S in late May. In the first half of June the two remaining seals turned south and moved back deep into the pack ice. The seals made about 150 dives per day each throughout the study period. Ninety percent of these were made to depths of less than 52 m. Individual maximum diving depths varied between 288 and 528 m. In March the seals were most active at night, when the dive depth was shallower than during the day. In April and May the seals were more active during day-time, with an absence of any diurnal change in divng depth. These results support the notion that crabeater seals predominantly feed on krill in Antarctic pack ice, even when winter returns to the waters off Queen Maud Land.

In vitro digestibility of different prey species of minke whales ( Balaenoptera acutorostrata )

Information on diet composition, daily energy expenditure, energy storage and the utilization of energy in the prey are important factors when evaluating the food consumption of minke whales (Balaenoptera acutorostrata) during their summer stay in northern waters. The purpose of the present study was in this context to obtain information on the digestible energy (DE) of different prey selected by minke whales. An in vitro three-stage digestion technique, simulating the different compartments of the digestive system, has been developed. The initial step simulated the anaerobic microbial fermentation of substrate in the forestomach. The next stage included the addition of pepsin (EC 3.4.23.1)-HCl, simulating ventricle enzymic decomposition, and finally, in the third step, fresh extract from duodenal contents was used to simulate enzymic intestinal degradation of the remaining components of the food. The inoculum was normally obtained from animals which had recently eaten the prey to be tested. In such tests we obtained a dry matter disappearance (DMD) and a DE for herring (Clupea harengus) of 80.4 (SD 5.0)% (n 18) and 92.1 (SD 3.7)% (n 16) respectively, and a DMD of krill (Thysanoessa sp.) of 83.4 (SD 4.9)% (n 6). The DMD of krill was reduced to 73.8 (SD 7.3)% (n 8) while the DE was 70.6 (SD 10.4)% (n 7) when inoculum from whales which had recently eaten cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) was used. These results indicate a high digestibility of the most common species of prey in these animals, and also that the whales have little difficulty in changing from one prey species to another.

Gut length, food transit time and diving habit in phocid seals

Abstract The southern elephant seal (Mirounga leonina) has the ability to dive for 2 h and reach depths of 1200 m. This creature is also exceptional in having a small intestine that is 25 times body length. Krockenberger and Bryden advanced the hypothesis that the long small intestine has developed to compensate for the extended periods with reduced or even abolished intestinal blood perfusion during diving. To test this hypothesis we have measured small-intestinal lengths in crabeater (Lobodon carcinophagus), Weddell (Leptonychotes weddellii), Ross (Ommatophoca rossi), leopard (Hydrurga leptonyx), harp (Phoca groenlandica), ringed (Phoca hispida) and hooded (Cystophora cristata) seals and related them to available data on their maximal dive duration. We found no significant correlation (P > 0.05) between intestinal length relative to body length and diving ability, but we found that small-intestinal internal area was significantly (P < 0.05) related to body length. A crude scanning electron microscopical examination of the small intestines of Weddell, crabeater, hooded and harp seals failed to reveal any gross anatomical differences between small-intestinal surfaces. This suggests that gut dimension in this variety of phocid species with widely differing diving ability is not related to diving habit, but is instead related to body size. The transit time of digesta was determined in two 1-year-old harp seals by use of radiopaque polyethylene rings of 4-mm diameter followed by X-ray examination, as markers for the solid phase passage, and chromium ethylene-diaminetetra acetic acid (Cr-EDTA) as a marker for the liquid phase. The transit time for the Cr-EDTA marker was 6.9 h ± 0.5 SE (range 4.5–8 h, n= 7), while 80% of the polyethylene markers appeared in the colon after 17.6 h ± 1.0 SE (range 14–21.5 h, n= 6) and were sometimes retained in the colon for several hours before defecation. These transit times did not change significantly (P > 0.05) in response to repetitive diving over a period of 8 h. This indicates that the often-used Cr-EDTA is not a good measure for digesta passage time when used alone in seals, and that the hypothesis of Krockenberger and Bryden is most likely wrong.

Digestibility of krill ( Euphausia superba and Thysanoessa sp.) in minke whales ( Balaenoptera acutorostrata ) and crabeater seals ( Lobodon carcinophagus )

Apparent digestible efficiency (% DE) was studied by use of dietary Mn as an inert marker, in minke whales (Balaenoptera acutorostrata) and crabeater seals (Lobodon carcinophagus) which had been eating krill. Median % DE in minke whales (n5) eating krill of the genus Thysanoessa sp. (energy density (ED) 23.8 kJ/g) was 93 (range 87-93). Median % DE in crabeater seals (n6) eating krill of the species Euphausia superba (ED 20.8 kJ/g) was 84 (range 79-85), which is significantly lower than the % DE of krill in minke whales (P = 0.008). Since the chemical composition in E. superba and in Thysanoessa sp. is similar, it is suggested that the complex multi-stomached system of minke whales, which contains both chitinase (EC 3.2.1.14)-producing as well as several other types of bacteria, is superior to the single-stomached system of crabeater seals with regard to krill digestion. It is worth noting, however, that the % DE of krill in the crabeater seal is still very high.

Food requirements of Northeast Atlantic minke whales

Abstract The Northeast Atlantic stock of minke whales (Balaenoptera acutorostrata) numbers ≈87,000 animals. In this report we estimate the food requirements of this stock from the energy requirements of growing and adult animals. The daily energy expenditure of free-swimming whales has been estimated by Blix and Folkow [1] to be 80 kJ/kg per day. This value includes the heat increment of fat deposition, growth and gestation. The additional energy costs of fattening and growth, determined from carcass analysis of whales caught in spring and autumn, was 17 and 8% of their total daily energy requirements, respectively. Based on an age-length relationship it was assumed that 73% of the population was growing, and 100% of all females older than 11 years were pregnant. By use of current estimates of the composition of the diet of Northeast Atlantic minke whales along with information on the seasonal changes in energy density of prey, it was calculated that this population consumes of the order of 1.4 million tonnes of various prey, during an assumed average stay of 6 months in the Northeast Atlantic. The deposited energy (blubber, muscle and visceral fat) does not cover more than about 60 days of the assumed energy requirements, when in other waters.

Application of the two-sample doubly labelled water method alters behaviour and affects estimates of energy expenditure in black-legged kittiwakes.

SUMMARY Despite the widespread use of the doubly labelled water (DLW) method in energetic studies of free-ranging animals, effects of the method on study animals are rarely assessed. We studied behavioural effects of two alternative DLW protocols. During two consecutive breeding seasons, 42 parent black-legged kittiwakes received either the commonly used two-sample (TS) or the less invasive single-sample (SS) DLW treatment. A third group served as a non-treated control. We evaluated the effect of treatment with respect to the time birds took to return to their nest after treatment and recaptures, and the nest attendance during DLW measurement periods. We found that TS kittiwakes took on average 20 times longer to return to their nest than SS kittiwakes after initial treatment, and nest attendance was reduced by about 40% relative to control birds. In contrast, nest attendance did not differ between control and SS kittiwakes. Estimates of energy expenditure of SS kittiwakes exceeded those of TS kittiwakes by 15%. This difference was probably caused by TS birds remaining inactive for extended time periods while at sea. Our results demonstrate that the common assumption that the TS DLW method has little impact on the behaviour of study subjects is in some circumstances fallacious. Estimates of energy expenditure derived by the SS approach may thus more accurately reflect unbiased rates of energy expenditure. However, the choice of protocol may be a trade-off between their impact on behaviour, and hence accuracy, and their differences in precision. Adopting procedures that minimize the impact of TS protocols may be useful.

Estimating population status under conditions of uncertainty: the Ross seal in East Antarctica

Abstract The Ross seal (Ommatophoca rossii) is the least studied of the Antarctic ice-breeding phocids. In particular, estimating the population status of the Ross seal has proved extremely difficult. The Protocol on Environmental Protection to the Antarctic Treaty currently designates the Ross seal as a ‘Specially Protected Species’, contrasting with the IUCNs classification of ‘Least Concern’. As part of a review of the Ross seals classification under the Protocol, a survey was undertaken in 1999/2000 to estimate the status of the Ross seal population in the pack ice off East Antarctica between 64–150°E. Shipboard and aerial sighting surveys were carried out along 9476 km of transect to estimate the density of Ross seals hauled out on the ice, and satellite dive recorders deployed on a sample of Ross seals to estimate the proportion of time spent on the ice. The survey design and analysis addressed the many sources of uncertainty in estimating the abundance of this species in an effort to provide a range of best and plausible estimates. Best estimates of abundance in the survey region ranged from 41 300–55 900 seals. Limits on plausible estimates ranged from 20 500 (lower 2.5 percentile) to 226 600 (upper 97.5 percentile).

Experimental evidence of seawater drinking in juvenile hooded (Cystophora cristata) and harp seals (Phoca groenlandica).

Abstract This study was undertaken to measure whether young harp seals (Phoca groenlandica) and hooded seals (Cystophora cristata) drink seawater and, if so, to investigate how the excess salt load is handled. Blood and urine samples were collected from hooded seal pups (n=3) and harp seal pups (n=3) after 2 weeks of freshwater exposure, at intervals during 3 weeks of seawater exposure and, finally, after 2 weeks of re-exposure to fresh water. Total water turnover, as measured by injection of tritiated water, was 2200 ml · day−1 and 3300 ml · day−1 in hooded seals and harp seals, respectively. The extent of mariposia was taken as the difference between total water turnover and influx of water through food (free and metabolic water) and respiratory water exchange. Seawater drinking amounted to 14% and 27% of total water turnover (rH2O) for the hooded seals and harp seals, respectively. Further evidence of mariposia was obtained from an increase in the excretion rate of the urine osmolytes Na+, Cl− and Mg2+, during the period of seawater exposure. It is concluded that water influx due to seawater drinking can not be excluded as a source of error when estimating food consumption of free-ranging harp seals and hooded seals, by use of labeled water techniques.

Physiological effects of seawater intake in adult harp seals during phase I of fasting.

Previous studies have shown that harp seals may drink considerable amounts of seawater. The current study was undertaken to study the physiological responses to bolus administration of seawater. Adult harp seals (Phoca groenlandica) were fasted without access to water for 48 h and then given 1000 or 1500 ml of seawater by a stomach tube. Changes in urine and plasma parameters were thereafter monitored for another 12-20 h. Urine production and urine excretion rate of Na+ and Cl- increased soon after administration and reached a maximum 3-4 h later. Urine osmolality was kept rather stable and high ( approximately 1500 mOsm x kg(-1)) following seawater administration, due to a drop in urine concentration of urea that was proportional to the simultaneous increase in urine concentration of NaCl. Plasma osmolality remained at approximately 340 mOsm x kg(-1), while plasma concentration of urea decreased some 20-25% due to increased excretion of urea when seawater was ingested. Despite bolus administrations of seawater of up to approximately 2% of body mass, homeostasis was maintained and no ill effects observed. It is concluded that the concentrating abilities of the kidneys of harp seals are sufficient to prevent net loss of body water following seawater ingestion. Seawater ingestion may, moreover, increase urinary osmotic space and thus serve as a mechanism to excrete additional urea produced during phase I of fasting.

The effect of body fat on basal metabolic rate in adult harp seals (Phoca groenlandica)

Three adult harp seals (Phoca groenlandica) were fed different daily amounts of capelin (Mallotus villosus), and their body composition determined by use of the tritiated water method at different levels of fattening. Basal metabolic rate (BMR) was measured after 5 days of fasting by indirect calorimetry, and was on average 1.1 W.kg-1 when 45% of body mass (BM) was fat and 2.3 W.kg-1 when body fat was reduced to 13% of BM. This suggests that body fat contributes little to BMR in these animals. It follows, that predictions of BMR on the basis of BM is questionable in seals, in which body fat may change seasonally between 20 and 60% of BM.