Provisioning and growth rates of nestling Fulmars Fulmarus glacialis: stochastic variation or regulation?

The pattern of chick feeding in the Fulmar Fulmarus glacialis at St Kilda, Scotland, was examined by repeated weighing of chicks throughout 14 consecutive days during the first half of the chick-rearing period in 1994. After correcting for metabolic weight losses, the sizes of positive mass increments between weighings were used to assess meal sizes and feeding frequency for each chick. Individual meals fed to chicks averaged 80.8 g (s.d. ± 21.0 g), or approximately 10% of adult mass. Each chick received 0 to 4 meals per day, with an average of 1.9 meals per chick per day, giving an average interval of around 25 h between meals delivered by each parent. The distribution of time intervals between feeds for each chick (whether single or double meals) followed a negative exponential function with a maximum value of 80 h. These results are not compatible with the idea that the purpose of large fat deposits in procellariiform chicks is to guarantee survival over long intervals between feeds. Over 14 days, the chicks' mean daily food requirements for zero-growth increased from 98 g to 160 g. This corresponded with an increase in feeding frequency but not meal size. Chicks with lower scores for body condition after feeding by both parents received more meals during the subsequent 16 h and had shorter intervals to the next feed, indicating that adults regulated feeding frequency in accordance with chick condition at the previous feed. This does not agree with the hypothesis that lipid accumulation by nestling Procellariiformes is a response to stochastic variation in food delivery associated with an absence of regulation. In view of the diversity of growth and feeding patterns present among the Procellariiformes, it is possible that lipid accumulation in this group does not have a unitary explanation.     

Variability and stochasticity of meal size and feeding frequency in the Little Sheanvater Puffinus assimilis

The pattern of chick feeding in Little Shearwater Puffinus assimilis on Selvagem Grande was examined by weighing chicks at 4–h intervals throughout eight successive nights and daily for a further 11 days (19 days in all). Individual meals fed to chicks averaged 23.2 g (s.d. ±4.7) or 13.6% of adult mass. Mass increments over 24 h (NET) were linearly related to the sum of positive mass increments over 4–h intervals during the night (SUM) by the equation NET = 0.36SUM - 5.89 (r²= 0.60). Using this relationship, I estimated that over a period of 18 nights, a mean of 95% of chicks were fed each night, and the mean interval between feeds was 1.05 nights, with a maximum of three nights. There was no significant day-to-day variation in feeding rate. These results were not compatible with the prevalent idea that the purpose of large fat deposits in Procellariiformes is to tide chicks over periodic fasts resulting from poor feeding conditions. On average, chicks required 16 g of food per day to maintain constant mass and converted 33% of their intake of food above this requirement into biomass. Meal size and feeding frequency were independent of chick size and body condition (body-mass corrected for body-size), and the masses of food received by individual chicks each night varied in direct proportion to previous values. These results suggest that the rate of food supply to chicks was not regulated by adjustment according to chicks' nutritional requirements. To some extent, this supports the hypothesis that lipid accumulation among Procellariiformes is related to stochastic variation in food supply rate, resulting from an absence of regulation of feeding. However, feeding was not stochastic, in that adults tended to deliver consistent amounts of food to their chicks, and the pattern of feeding among even the worst-fed chicks was inconsistent with a need for large lipid stores based upon chance variation in food delivery.     

Growth and provisioning strategies of Northern Fulmars Fulmarus glacialis

Growth and foraging strategies in procellariiforms show a great deal of variation, but the fulmarine petrels are notable in that chicks are fed frequently and develop unusually rapidly. This study examined age-related and daily variation in provisioning of the Northern Fulmar Fulmarus glacialis throughout the chick-rearing period at Fair Isle in 1997. In common with many other petrels, meal mass showed an initial rise with age, probably because of a gradual increase in chick gut capacity, but then levelled off. By comparison, feeding frequency showed little age-specific variation until chicks reached the oldest age-class, when the number of meals declined to less than a third of the previous level as chicks underwent mass recession prior to fledging. Compared with the limited day-to-day variation in mean provisioning rates for the whole sample, food delivery to particular chicks was much more variable, suggesting that differences in feeding rates were determined by stochastic factors influencing the feeding success of individual parents. The caloric density of feeds and their size in relation to adult mass were lower in Northern Fulmars than in most other Procellariiformes. This implies that adults are not heavily dependent on stomach oil formation to raise the energy content of the payload, but rely on a high feeding frequency to maintain adequate rates of energy transfer to chicks.     

Experimental evidence for the relationship between food supply, parental effort and chick survival in the Lesser Black-backed Gull Larus fuscus

We studied breeding success, chick growth, parental effort and chick behaviour in two groups of Lesser Black-backed Gulls Larus fuscus whose chicks were provided with additional food until 7 days after hatching or until fledging. These data were compared with those from control pairs which we studied simultaneously to test the hypotheses that food was in short supply during the chick stage at the colony site and that in such circumstances the behaviour of adults and young is mainly responsible for the low success. Pairs whose chicks were fed with additional food until fledging showed a higher fledging success than control pairs (intermediate for pairs of first experimental group). During the first week after hatching, experimental adults of both groups were present together at the territory for longer than control pairs. In adult females of experimental pairs, the length of feeding trips was shorter than in females of control pairs, whilst the rate of chick feeding was more frequent in the experimental broods. After the chicks were 7 days old, differences were significant only for the experimental pairs whose chicks were provided with additional food until fledging. Chicks fed until fledging showed a higher daily mass and wing-length increments and reached a higher fledging mass at an earlier age than both control chicks and chicks which were provided with additional food until day 7. Starvation occurred only in control chicks and in chicks of the first experimental group after we had stopped providing food. When food was in short supply, fledging success of gulls was adversely affected as a result of both starvation (because of the lower feeding rates of chicks) and a higher predation rate (arising from changes in behaviour of both adults and chicks).     

Does the foraging strategy of adult Short‐tailed Shearwaters cause obesity in their chicks?

The chick provisioning behaviour of Short-tailed Shearwaters Puffinus tenuirostris breeding at the northern edge of their distribution on Montague Island, New South Wales, was examined in February and March 1997. The duration of individual foraging trips of parents, weight changes of adults and chicks, and meal sizes delivered to chicks were determined. It was found that individual parents mixed a long foraging trip to Antarctic waters (14.4±2.0 days) with one to three short foraging trips (1.36±0.7 days, mode=1 day). Adults gained body mass on long trips and lost weight on short trips. The size of meals fed to the chicks was significantly greater after a long trip (161±21 g) than after a short trip (135±28 g), although short trips increased the overall chick feeding frequency. The variable number of short trips made by adult Short-tailed Shearwaters and the relationship between short trips and adult body condition were consistent with current life-history theory: adults do not sacrifice their own body condition to increase food delivered to their chicks.
Modelling revealed that this dual foraging strategy inevitable leads to chicks enduring long intervals between meals. These long intervals may have led to the evolution of an over-feeding strategy by parents and the nestling obesity reported in this shearwater. The durations of the long trips from Montague Island were significantly greater than those for Short-tailed Shearwaters breeding at the centre of their distribution in Tasmania, although there was no significant difference in the length of short trips. A commitment to feed regularly in Antarctic waters may explain why the breeding distribution of this species does not extend much further north.     

Plasticity of Noddy Parents and Offspring to Sea-Surface Temperature Anomalies

Behavioral and/or developmental plasticity is crucial for resisting the impacts of environmental stressors. We investigated the plasticity of adult foraging behavior and chick development in an offshore foraging seabird, the black noddy (Anous minutus), during two breeding seasons. The first season had anomalously high sea-surface temperatures and ‘low’ prey availability, while the second was a season of below average sea-surface temperatures and ‘normal’ food availability. During the second season, supplementary feeding of chicks was used to manipulate offspring nutritional status in order to mimic conditions of high prey availability. When sea-surface temperatures were hotter than average, provisioning rates were significantly and negatively impacted at the day-to-day scale. Adults fed chicks during this low-food season smaller meals but at the same rate as chicks in the unfed treatment the following season. Supplementary feeding of chicks during the second season also resulted in delivery of smaller meals by adults, but did not influence feeding rate. Chick begging and parental responses to cessation of food supplementation suggested smaller meals fed to artificially supplemented chicks resulted from a decrease in chick demands associated with satiation, rather than adult behavioral responses to chick condition. During periods of low prey abundance, chicks maintained structural growth while sacrificing body condition and were unable to take advantage of periods of high prey abundance by increasing growth rates. These results suggest that this species expresses limited plasticity in provisioning behavior and offspring development. Consequently, responses to future changes in sea-surface temperature and other environmental variation may be limited.     

Foraging and provisioning strategies of black-browed albatrosses in relation to the requirements of the chick: natural variation and experimental study

We studied regulation of the food supply to black-browed albatrosschicks at Kerguelen by simultaneously recording the provisioningrates achieved by individual parents and satellite trackingforaging birds during two seasons, by studying changes in adultmass, and by experimentally manipulating the food requirementof chicks. In 1994 black-browed albatrosses had a higher breedingsuccess and produced heavier chicks that grew faster than in1995. They spent a similar time foraging but brought heaviermeals to their chick in 1994. Satellite tracking indicated thatin both seasons birds foraged in the same oceanographic area,250 km from the colony. Travel times to and from this area remainedunchanged, and similar times were spent foraging there. In ourstudy area, black-browed albatrosses appear to rely on a foodresource that is predictable in location, but whose availabilityvaries from one year to the next. The principal difference betweenyears of differing food availability was that birds broughtlarger meals when food was more abundant Costs of commutingto nearby feeding areas are probably low and allow the deliveryof energy to the chick at a high rate. A study carried out in1991 indicated that there was no relationship between the changesin adult mass from one trip to the next and the duration offoraging trips or feed mass, suggesting that adult body conditionhad little influence on the provisioning strategy of this species.An experiment whereby some chicks were deprived of food andothers received supplementary food showed that parents of underfedchicks spent the same time foraging and brought slightly largeramounts of food to their chicks as control parents. We suggestthat parents are searching for food to the maximum limits oftheir ability and thus cannot reduce further foraging time,but underfed chicks can swallow more food. Parents of overfedchicks delivered less food and increased the time between feeds.The reduction in provisioning frequency is interpreted as thecapacity of parents to modify their foraging behavior accordingto the nutritional status of the chick, but the reduction offeed mass is probably the result of chicks being close to theirmaximum assimilatory capacity. Comparison between Procellariiformspecies indicates extensive differences in the degree to whichparents can regulate the supply of food to their chicks. Neriticspecies like black-browed albatrosses appear to have a reducedability to regulate, and especially to increase provisioningrates, whereas more pelagic species may have a greater regulationability     

Seasonal changes in the provisioning behaviour and mass of male and female wandering albatrosses in relation to the growth of their chick

Procellariiform seabirds have a number of extreme life-history characteristics in common, in particular low reproductive rates and slow postnatal development, which are generally assumed to reflect the difficulty in acquiring energy in the marine environment. The wandering albatross (Diomedea exulans) is a sexually dimorphic species with the longest postnatal growth found in any bird, suggesting severe constraints on provisioning and possible sex-specific strategies of provisioning. We studied the provisioning behaviour and mass changes of male and female parent wandering albatross throughout the 9-months rearing period to examine how each sex adjusts its foraging effort in relation to the needs of the chicks and the seasonal changes in food availability. The study was carried out on the Crozet Islands, using an automated system recording continuously the attendance pattern of parents between March and December 1994. During the brooding period when energy requirements are highest, parents only perform trips of short duration to sea, and their body condition deteriorates. When the chick is old enough to be left alone, the parents mix short and long foraging trips. The proportion of short trips is very high until July, allowing high rate of food delivery and rapid growth, and at the same time the body condition of adults improves. From August this proportion declines until fledging in December. As a result, the feeding rate decreases from August and adult condition declines, suggesting that feeding conditions at sea are better during the first part of the chick-rearing period, i.e. in autumn and winter. Male parents perform more short trips of shorter duration and provide larger meals than females, delivering an estimated total after brooding of 110 kg of food, compared to 70–80 kg delivered by females. Meal size is inversely related to the body condition of male chicks but not to that of female chicks, suggesting that food delivery is regulated by the adults in response to the condition of the male chick. Male chicks received larger meals and more food every month than female chicks, and overall it was estimated that they receive, after brooding, 195 kg of food compared to 180 kg for the female. As a result, male chicks have a higher growth rate, attain a higher asymptotic mass, and are larger and heavier at fledging than female chicks. However, the differences are relatively small between the chicks of each sex and suggest that energy may be used differently between the sexes to maximise fitness. The results of the study suggest that provisioning effort of wandering albatrosses is adjusted by parents in relation to the availability of food, to the energetic needs of the chick and to the sex of the chick. The adult body mass is likely to play an important role in the long term for the regulation of provisioning, deficits in body mass probably providing the buffer in high power-requirement periods. Accepted: 20 March 2000     

Nestling feeding strategy of the British storm-petrel Hydrobates pelagicus in a Mediterranean colony

The pattern of chick feeding of the British storm-petrel Hydrobates pelagicus in a Mediterranean colony was examined by weighing chicks at 24-h intervals on different days during the nestling period. In order to calibrate daily mass increments (NET) against number of feedings, daily mass changes of chicks were regressed upon the sum of positive mass increments recorded overnight (SUM) during four nights. The average meal size delivered to chicks per night by one parent was 3.5 ß 1.3 g or 12% of adult weight. This was insufficient for sustaining constant chick mass during a day. On average 85% of chicks were fed each night, and the mean interval between feeds was 1.2 ß 0.5 nights. Nightly feeding frequencies differed among days, but this night-to-night variation was not related to meteorological conditions. Both food requirements necessary for chick body maintenance (zero-growth) and meal size were relatively constant for ages up to 59 days. However, feeding frequency decreased throughout the fledging period, accounting for agespecific variation in growth rates until fledging. Food requirement and feeding patterns at Benidorm were different from North Atlantic colonies. None the less, growth patterns were almost identical, suggesting adjustment to maintain chick body mass at a determined level, as food delivery to nestling appears to be regulated to chicks'nutritional requirements.     

Nestling obesity in procellariiform seabirds: temporal and stochastic variation in provisioning and growth of short-tailed shearwaters Puffinus tenuirostris

Procellariiform seabirds such as short-tailed shearwaters Puffinus tenuirostris accumulate large quantities of lipid during the nestling period. The functional significance of this pattern of development remains unclear, but has been related both to temporal variation in feeding conditions around the colony and to stochastic variation in the foraging success of individual parents. This paper examines temporal and age-specific variation in the pattern of food delivery to nestling short-tailed shearwaters, which have one of the lowest provisioning rates of any procellariiforms and are known to experience occasional long intervals between feeds. We assess whether variation in the provisioning rates of chicks was associated primarily with temporal variation in food delivery at the level of the colony or with stochastic variation in food delivery at the level of the individual. We then discuss this variability in the context of nestling obesity. For all but the youngest chick age-classes, individual meals delivered by adults averaged 141 g, which was 25% of adult body mass. The proportion of chicks fed each night was low (49%) and highly variable (coefficient of variation = 82%), which means that occasional long intervals between feeds would be expected to arise simply by chance. In keeping with this, intervals between feeding events for individual chicks followed a negative exponential distribution with a mean of 2 nights and a maximum interval of 13 nights. There was significant temporal variation in food delivery, but deviations from expected values for both feeding frequency and meal size were restricted to a small number of nights, included values both higher and lower than expected and did not persist for more than 2 nights in succession. These data suggest that even among those species with very low feeding frequencies and occasional long intervals between feeds, nestling obesity in Procellariiformes should be regarded as a response to chronic stochastic variability in food delivery at the level of the individual chick rather than as insurance against sporadic temporal variation at the level of the colony. Received: 3 March 1997 / Accepted: 10 May 1997     

Chick provisioning and nest attendance of male and female Wilson’s storm petrels <Emphasis Type="Italic">Oceanites oceanicus</Emphasis>

Seabirds show a range of patterns of sexual size dimorphism and sex-specific parental investment, but the underlying causes remain poorly understood. The aim of the present study was to test two longstanding hypotheses of parental investment in a sexually monomorphic species, Wilson’s storm petrel Oceanites oceanicus, namely that males attend chicks more frequently and females deliver larger meals (Beck and Brown in Br Antarct Surv Sci Rep 69:1–54, 1972). We recorded in eight seasons, both during incubation and chick rearing, which adult was caught first in a nest and found no difference in the probability of catching a male or a female first in any year. Additionally, in five seasons we employed a miniature video camera to record nest attendance during chick rearing and found no significant difference except for 2006, a year with very low krill availability, where females visited the nest less often than males. We then combined video observations with periodic weighing of chicks to estimate mean daily feeding mass (g/day) of males and females and found no difference in the amount of food delivered per day between the sexes. However, in years with low krill availability, males and females tended to use different strategies to achieve the same feeding rates, with females undertaking longer foraging trips and delivering heavier meals. Thus, our results do not support the hypothesis of a general sex-specific parental investment in Wilson’s storm petrels, but a tendency for a context-dependent sex-specific investment in the years of food shortage.     

Chick provisioning rates and growth in Blacklbrowed Albatross Diomedea melanophris and Grey-headed Albatross D. chrysostoma at Bird Island, South Georgia

We compared the parental division of labour and the pattern and rate of parental provisioning by two sympatric species of albatross of similar mass and breeding timetable but differing in diet and in the duration of chick‐rearing. Using electronic weighing platforms inside artificial nests, we recorded chick mass of Black‐browed Albatross and Grey‐headed Albatross at Bird Island, South Georgia every 10 minutes for both species in 1993 and 1994 and for each species in two other years between 1990 and 1996. The chick mass data (nearly one million weighings) were used to calculate meal mass (over 5000 meals) and intervals between meals. Adult birds were fitted with radio‐transmitters which allowed each meal to be allocated to the appropriate parent. The combination of meal mass and foraging trip duration were used to calculate provisioning rates for chicks and individual adults. Overall, Black‐browed Albatrosses delivered significantly lighter meals (569 g) than Grey‐headed Albatrosses (616 g) but more frequently (every 2.07 days and 2.50 days respectively). Thus combining foraging trip data for both parents, Black‐browed Albatross chicks received a meal every 1.22 days compared with 1.26 days for Greyheaded Albatross. These rates did not differ significantly. The contribution of each sex of each species in chick provisioning fluctuated between years, being similar in some years or biased towards males in others. Chicks of both species that failed to fledge received smaller, less frequent meals than successful chicks. In 1990 and 1994, Black‐browed Albatross chick provisioning rates were lower than in 1992 and 1993. In 1990, both meal mass and trip duration were affected, but only in 1994 was trip duration longer. Greyheaded Albatross chick provisioning rate was lower in 1994 than in other years but trip duration was longer. In each species, significant changes in meal mass and trip duration occurred within the chick‐rearing period. Chick provisioning rates invariably declined before chicks attained their peak mass. For both species, chick growth rates and peak and fledging mass, but not fledging age, were affected by differences in provisioning rate.     

Foraging trip decisions by the streaked shearwater Calonectris leucomelas depend on both parental and chick state

Parents of albatross and shearwater species employ a dual foraging strategy, feeding their chicks quickly in repeated short trips and then restoring their own fuel reserves during longer trips. A decline in parental body condition is believed to trigger longer trips, but chick body condition and age may also play a role. To investigate these factors in the little-studied streaked shearwater Calonectris leucomelas, we monitored the nest attendance of 17 pairs on Mikura Island in 2005 using an automated identification system. We also monitored body mass changes and meal masses of 5 of the 17 pairs using an automated weighing system. Although the birds did not show a clear dual foraging pattern, trip duration varied widely from 1 to 15 days. On average, the birds fed chicks 67.6 g during nighttime meals at 2.74-day intervals. Since meal mass did not depend on trip duration, feeding efficiency (meal mass delivered per unit trip duration) decreased as trip duration increased. Parents accumulated more energy reserves when they took longer trips. Parents appeared likely to initiate longer trips when their body condition declined or chick body condition recovered.     

Food provisioning to nestling shearwaters: why parental behaviour should be monitored?

We examined temporal variation in food delivery to nestling Cory's shearwaters Calonectris diomedea, by repeated periodic weighings during the night. We tested whether the magnitude and frequency of meals were influenced by the condition of chicks. In contrast to previous studies of chick provisioning in petrels and shearwaters, the evidence of feeding derived from chick weight gains was complemented by data provided by an electronic system, which logged the entry of each parent to the nest. Estimates of feed size and visiting frequency obtained from chick weighing alone differed from similar estimates obtained using the automatic logging equipment. The data obtained with the logging system combined with chick weighing also showed that, to some extent, food provisioning was regulated, chicks left in poorer condition being more likely to receive food the next night than those left in better condition. The methods based on chick weight gains alone did not detect this regulation effect. Our findings suggest that resolving parental visits to the nest is crucial to obtain accurate parameter estimates, and to address the problem of regulation of provisioning rates in Procellariiformes. Our results do not support the hypothesis that accumulation of fat is just a by-product of chronic overfeeding arising from stochastic variation in foraging success at sea. Copyright 1999 The Association for the Study of Animal Behaviour.     

Influence of adult breeding experience on growth and provisioning of Wandering Albatross Diomedea exulans chicks at South Georgia

For most seabirds, reproductive performance improves with age; in albatrosses this is thought not to be so (experience being acquired before starting breeding) but only one study (of chick growth in a single season at one site) has specifically addressed this. We compared the provisioning performance and growth rates of chicks of Wandering Albatrosses Diomedea exulans breeding for the first (IN), second and third (LE) and fourth or more times (EE) on Bird Island, South Georgia in the austral winters of 1996 and 1997. Eggs from EE adults were significantly heavier than the other two categories and these chicks had a greater mass and longer wings up to 160 days of age and longer culmen and tarsus up to 115 days old. However chicks from all categories fledged at the same average mass, size and age. No significant differences between categories in feeding frequency or meal size were detected but experienced adults made shorter long foraging trips and spent more time at the nest than less experienced birds. Adults that remained at the nest gave chicks smaller meals than those that left immediately after feeding the chick. Although provision of smaller but more frequent meals by experienced adults promotes more rapid chick growth, the resulting differences do not persist into the late chick-rearing period. Our results were very similar to those from Iles Crozet in the Indian Ocean, supporting the hypothesis that when Wandering Albatrosses start to breed they are fully competent foragers but that it takes a while, during early chick-rearing, for birds breeding for the first time to adapt to the additional demands of provisioning a chick.     

Sex differences in provisioning rules: responses of Manx shearwaters to supplementary chick feeding

Sex differences in food provisioning have been found in a numberof socially monogamous birds with biparental care, but the reasonsremain unclear. In Manx shearwaters, males provide 40–50%more food for chicks than do females, and previous empiricaldata have suggested that this difference could arise becausefemales are able to regulate food delivery by reducing the provisioningof well-nourished chicks, whereas males are not (hypothesis1). Alternatively, however, males may be as capable as femalesof assessing and responding to the variation in the nutritionalrequirements of their chick but have a higher threshold forreducing food delivery to well-nourished chicks (hypothesis2). To test these two hypotheses, we used supplementary feedingto manipulate the nutritional status of chicks and then examinedthe responses of male and female parents and their offspring.Supplementary feeding significantly reduced both the beggingbehavior of chicks and the frequency and sizes of meals deliveredby parents. Males and females reduced their overall provisioningrates to a similar extent (males by 38%, females by 42%), somaintaining the same difference in contributions to provisioningin the control group (males 58%, females 42%) and the experimentaltreatment (males 59%, females 41%). These data strongly supporthypothesis 2. Supplementary feeding of chicks resulted in fewervisits by parents and a higher proportion of long trips in bothsexes (4 days for males, 5–7 days for females). However,maximum trip durations were unchanged, suggesting that supplementaryfeeding of chicks had no effect on the foraging ranges or overallfood-provisioning strategies of parents.     

Luxury in freshwater and stress at sea? The foraging of the Common Tern Sterna hirundo

We compared the foraging strategies of Common Terns Sterna hirundo in freshwater (Lake Jeziorsko, Brzeg, Poland) and marine environments (Minsener Oldeoog, German Wadden Sea). Body mass changes, nest relief and duration and number of feeding trips per day were studied by automatically weighing the adults, using electronic balances under the nests. At the freshwater site, adults were lighter both before and after feeding and gained less mass during a trip. in the Wadden Sea, single feeding trips lasted longer than at the freshwater site and the terns made fewer trips per day. To achieve the same mass gain per day as in birds in freshwater, trips at sea had to be longer and food intake per trip was higher. The daily duration of absence for feeding and the daily mass gain were about the same in both areas. The limnetic feeders finished foraging earlier in the evening than the terns foraging at sea. These differences are consistent with the hypothesis that limnetic prey availability was consistent, whereas the tides limited the availability of marine prey. In consequence, foraging over freshwater presents several advantages, such as higher colony attendance, better mate coordination and better parental care.     

Patterns of energy acquisition by a central place forager: benefits of alternating short and long foraging trips

In some seabirds, foraging trips have been defined as eitherlong or short, with the length of time spent traveling to theforaging area apparently a critical feature in determining foragingtrip length. Using logger technology, together with complimentarydata from published studies, we investigated traveling and foragingtimes in 18 free-living Adélie Penguins Pygoscelis adeliae,which were foraging for chicks. Most deep, foraging dives weredistributed around the center of the foraging trip. This centraltendency was particularly apparent if the cumulative amountof undulations in the depth profile (indicative of prey capture)was considered during deep dives; values started to increasebefore 20.9% and ceased after 67.2% of the dives had occurred.This concentration of the feeding activity in the middle ofthe foraging trip indicates that birds traveled to and froma prey patch whose location varied little over the birds' trips.These data form the basis for a simple model that uses travelingand foraging times together with projected rates of prey ingestionand chick and adult gastric emptying to determine that thereare occasions when, to optimize rates of prey ingestion whileat sea for both adults and chicks, birds should conduct foragingtrips of bimodal lengths.     

Mapping pre- and post- fledging foraging locations of thick-billed murres in the North Water polynya

In this study of thick- billed murres in high- arctic Greenland we used electronic data loggers and satellite transmitters (PTTs) to identify the foraging areas of chick-rearing adults, and to map the routes and staging areas of adults accompanying post-fledging chicks during their swimming migration within the North Water (NOW) polynya. During the pre-fledging period the majority of 19 foraging trips performed by 8 birds went to a shelf area north of the colony where 83% of all dives took place. Individual birds headed in different directions during successive trips, and went up to 47 km from the colony. Upon fledging the four PTT-tagged adult/chick pairs initiated swimming migration by heading south-west from the colony. All pairs moved fast until they arrived at a shallow bank area ca 180 km from the colony, where at least two of the pairs remained for more than a week. Speed during the active migration averaged 2.5 km h^-1 with a peak of 6.6 km h^-1. In the pre-fledging period the birds utilised a feeding area outside the normal foraging range of murres from other colonies. Similarly, post-fledging adult/chick pairs may have benefitted from reduced food competition when they moved to a staging area situated at the only shallow area in the polynya without any adjacent murre colony. This initial study suggested that the high-arctic murres did not hasten towards the wintering grounds, and that the NOW remained important even to post-fledging murres.     

Growth and energetics of a small shorebird species in a cold environment: the little stint Calidris minuta on the Taimyr Peninsula, Siberia

The little stint Calidris minuta is one of the smallest shorebird species breeding in the Arctic (weighing 4.3  g on hatching). Their chicks are small and have a high surface area-to-volume ratio. We determined prefledging growth, energy expenditure and time budgets for little stint chicks in northwestern Taimyr, Siberia. A modified power curve was introduced to model the relationship between daily energy expenditure and body mass. Total metabolisable energy, TME, over the 15-d prefledging period was 107% greater than the allometric prediction for a bird the size of a little stint. Their growth rate coefficient was 14% greater than the prediction for a bird their size. The growth of young chicks was reduced in cool weather, possibly due to a reduction in foraging time in order to be brooded and reduced food availability which impact foraging efficiency. We did not detect weather effects on energy expenditure of chicks, but lack of temperature variation during energy expenditure measurements may have prevented this. In sum, both growth rate coefficient and energy expenditure of little stint chicks were greater than predicted and this is similar to that observed in other arctic shorebird species.