This study investigated the foraging behaviour, diving behaviour, movements and diet of lactating southern sea lions in the Falkland Islands. They appeared to be primarily nocturnal, benthic foragers hunting at depths ranging from the surf line down to 250 m, and from just off their breeding sites out to 45 km offshore. Occasional longer trips may extend foraging ranges to over 150 km from breeding sites. Attendance patterns suggest that Falklands' sea lions do not haulout simply to feed their pups. Other factors such as predation and thermoregulation may cause them to come ashore. When at sea they travelled at approximately 1 m s 71 , slower than previous studies of otariids indicated. Absence from the breeding site did not imply continuous foraging; all study animals used remote haulout sites away from their pups. Repeated use of the same areas and the short duration of foraging trips suggest that they were able to catch enough food within a fairly small foraging range. Data on diet support the suggestion that lactating sea lions are mainly benthic foragers. They take a wide range of prey species. The most common cephalopods were Loligo gahi, similar in size to those taken by the ®shery. There is therefore potential for competition between the sea lion population and the Loligo trawl ®shery. There was considerable overlap between the diet of sea lions and those of Gentoo and Magellanic penguins, which suggests that sea lions may compete with both species.

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Seals may delay costly physiological processes (e.g. digestion) that are incompatible with the physiological adjustments to diving until after periods of active foraging. We present unusual profiles of metabolic rate (MR) in grey seals measured during long-term simulation of foraging trips (4-5 days) that provide evidence for this. We measured extremely high MRs (up to almost seven times the baseline levels) and high heart rates during extended surface intervals, where the seals were motionless at the surface. These occurred most often during the night and occurred frequently many hours after the end of feeding bouts. The duration and amount of oxygen consumed above baseline levels during these events was correlated with the amount of food eaten, confirming that these metabolic peaks were related to the processing of food eaten during foraging periods earlier in the day. We suggest that these periods of high MR represent a payback of costs deferred during foraging.

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.

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.