We measured the breeding performance, body condition, time budgets and foraging ranges of Kittiwakes Rissa tridactyla at Sumburgh Head, Shetland, in two years of contrasting food availability. Kittiwakes in Shetland generally feed their young almost entirely on sandeels, and fisheries data indicated that stocks of sandeels in Shetland waters were at least ten times higher in 1991 than in 1990. Fledging success of Kittiwakes was nil in 1990 and 68% of eggs laid in 1991, although clutch-size and hatching success were no different between years. Post-hatching foraging trips in 1991 were of comparable duration to those recorded at other colonies in conditions of good food supply (2-3 h), while trips recorded during incubation or post-hatching in 1990 were approximately three times longer on average than at corresponding stages of the breeding season in 1991. Radio-tracking data indicated that adults generally stayed within 5 km of the colony in 1991 but flew more than 40 km from the colony on each trip in 1990. Eggs were apparently not left unattended in either year, despite the fact that this required adults to incubate for periods in excess of 4. 4 h in 1990. The extent to which adults were able to increase trip durations, foraging ranges and incubation shift lengths between years, while maintaining hatching success, indicates the degree to which Kittiwakes are normally buffered against adverse feeding conditions during incubation. Reduced nest attendance and lower body-condition of adults post-hatching in 1990. in conjunction with complete post-hatching breeding failure, indicate that adults were beyond the limits of their buffering capacity during chick-rearing in 1990.
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The foraging behaviour of Guillemots Uria aalge at sea was compared between 2 years of radically different food abundance. Radio telemetry was used to determine foraging locations and diving patterns. In the poor compared with the good food year, foraging trips were much longer, the birds foraged more than six times further from their breeding sites, they spent over five times as much time diving when at sea and their estimated energy expenditure was twice as great. Time spent foraging in the poor food year was at the expense of time spent sitting at the colony. The duration of a foraging trip was a poor indicator of distance travelled but a good indicator of the amount of time spent diving. Mean dive durations, surface pause durations and interbout periods did not differ between years, but individuals made more than four times as many dives per diving bout in the poor food year. Surface pause lengths did not vary with water depth in either year. In the poor food year, birds made shorter surface pauses for a dive of a given duration than in the good food year, possibly accepting a lactic acid debt in order to maximize searching time, The duration of the interbout period was positively related to the number of dives in the previous bout, and dives tended to get shorter in long diving sequences, suggesting possible exhaustion effects. These data demonstrate that breeding Guillemots have the capacity to adjust their foraging behaviour and time budgets in response to changes in food abundance, but this flexibility was not sufficient to compensate fully for the very low food abundance experienced by birds in this study.
The breeding performance, food fed to chicks and adult time budgets of Guillemots Urin aalge were examined in a year of high and a year of low food availabiIity. There was no difference between the 2 years in reproductive success, although the rate of chick feeding, chick weight and fledging success were greater in the year of high food availability. On average, chick prey items were larger in the poor food year, but this was i n s u~~i e n t to compensate €or the lower feeding frequency. Chick feeding frequency did not differ between days in the good year but did increase later in the season in the poor food year. Compared with the high food availability year, adult Guillemots in the year of low food availability spent much less time resting at the breeding colony. and their foraging trips were twice as long. Foraging birds tended to make several successive trips before resuming brooding duties from their mates when food supplies were good, but in the low food availability year single trips were the norm. These results demonstrate that predators experiencing reduced food supply may mitigate the effects on their reproductive output by shifting their time allocation such that more time is availab~e for foraging.
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