Foraging by little auks in the distant marginal sea ice zone during the chick-rearing period

During the chick-rearing period, little auks Alle alle adopt a bimodal foraging strategy, alternating long trips with several short ones. It has been postulated that they reach more remote areas during long feeding trips than during short ones. However, the range of their foraging flights has never actually been measured. The aims of this study were to find the exact location of the little auk feeding grounds and to investigate whether they reach remote areas during long foraging trips using miniature GPS and temperature loggers. The study was conducted in 2009 in Magdalenefjorden (79°34′N, 11°04′E), one of the main breeding grounds of little auks on Spitsbergen. The temperature logger records indicated that during short trips, little auks visit warmer waters (situated close to the colony) than during long ones. The tracks of two GPS-equipped birds indicated that during long trips little auks foraged in the distant, food-abundant marginal sea ice zone, at least 100 km away from the colony. During long trips, birds make several stops at sea, perhaps sampling the foraging area with respect to prey distribution. Since food conditions near the studied colony are usually suboptimal, little auks may be exploiting distant feeding areas to compensate for the poorer-quality food available at nearby foraging grounds. The extended duration of long foraging trips may enable birds to collect food for chicks on food-abundant, remote foraging grounds as well as acquire, process and excrete food needed for self-maintenance, reducing the costs of flight to the colony.     

Divergent diving behavior during short and long trips of a bimodal forager,the little auk Alle alle

The purpose of this study was to characterize for the first time seabird diving behavior during bimodal foraging. Little auks Alle alle, small zooplanktivorous Alcids of the High Arctic, have recently been shown to make foraging trips of short and long duration. Because short (ST) and long trips (LT) are thought to occur in different locations and serve different purposes (chick‐ and self‐feeding, respectively) we hypothesized that foraging differences would be apparent, both in terms of water temperature and diving characteristics. Using Time Depth Recorders (TDRs), we tested this hypothesis at three colonies along the Greenland Sea with contrasting oceanographic conditions. We found that diving behavior generally differed between ST and LT. However, the magnitude of the disparity in diving characteristics depended on local foraging conditions. At the study site where conditions were favorable, diving behavior differed only to a small degree between LT and ST. Together with a lack of difference in diving depth and ocean temperature, this indicates that these birds did not increase their foraging effort during ST nor did they travel long distances to seek out more profitable prey. In contrast, where local foraging conditions were poor, birds increased their diving effort substantially to collect a chick meal during ST as indicated by longer, more U‐shaped dives with slower ascent rates and shorter resting times (post‐dive intervals and extended surface pauses). In addition, large differences in diving depth and ocean temperature indicate that birds forage on different prey species and utilize different foraging areas during LT, which may be up to 200 km away from the colony. Continued warming and deteriorating near‐colony foraging conditions may have energetic consequences for little auks breeding in the eastern Greenland Sea.     

The effects of loggers on the foraging effort and chick-rearing ability of parent little auks

We studied the effects of loggers attached to chick-rearing little auks (Alle alle) on their daily time budget (proportion of time spent in the colony and at sea), foraging activity (duration and proportion of long and short foraging flights), chick provisioning rate and their growth and development on Spitsbergen. We found that experimental parent birds performed shorter but more frequent long foraging flights and reduced the frequency of short foraging flights. They spent more time at the colony and reduced chick provisioning rate compared to control birds. Nestlings reared by experimental parents weighed significantly less at their middle, peak and fledging age and departed colony later than chicks of control parents. Little auks depend on energy-rich copepods associated with cold Arctic waters and are expected to face the climate-induced worsening of the foraging conditions, which may have negative impact on their time/energy budget and survival. The study may help to determine the level of extra effort little auks need to invest to breed successfully.     

Stage-dependent foraging in breeding black-legged kittiwakes Rissa tridactyla: distinguishing behavioural responses to intrinsic and extrinsic factors

Different phases of the annual cycle in birds and mammals are often associated with characteristic and recurrent foraging behaviours. The extent to which stage‐dependent changes in foraging behaviour are caused by intrinsic or extrinsic factors is unclear. We controlled for the effects of extrinsic factors by synchronising groups of incubating and chick‐rearing black‐legged kittiwakes Rissa tridactyla. Synchrony amongst incubators and rearers was achieved experimentally by switching eggs between nests. Behavioural responses to the treatment varied between the sexes. Male kittiwakes with prolonged‐incubation made fewer foraging trips but of greater duration compared to those rearing chicks resulting in no change in the time spent on trips between the two groups. Females with prolonged‐incubation carried out fewer trips than those rearing chicks but trip duration did not differ between the two stages which resulted in prolonged‐incubating birds spending a lower percentage time on trips. In contrast, foraging ranges did not differ between prolonged‐incubation and chick‐rearing birds for either sex. This suggests that extrinsic factors, such as food availability and distribution determine kittiwake foraging locations and ranges, whereas intrinsic factors, reflected in parental duties, constrain nest attendance. Female prolonged‐incubators invested lower levels of parental effort, in terms of daily energy expenditure, compared to chick‐rearers whereas males did not show stage‐related differences in energy expenditure. This provides evidence that incubation could be an energetically cheaper stage although under normal conditions this difference may be masked by temporal variation in environmental factors. We conclude that while conditions differ between the incubation and chick rearing stages for kittiwakes at this colony, they are not the main factors prompting changes in stage‐related foraging patterns. Intrinsic factors such as sex differences, or behaviours required for each stage of the annual cycle, rather than extrinsic factors related to seasonal environments, are likely to be the main proximate cause of recurring changes in behaviour between breeding stages.     

Arctic warming: nonlinear impacts of sea‐ice and glacier melt on seabird foraging

Arctic climate change has profound impacts on the cryosphere, notably via shrinking sea‐ice cover and retreating glaciers, and it is essential to evaluate and forecast the ecological consequences of such changes. We studied zooplankton‐feeding little auks (Alle alle), a key sentinel species of the Arctic, at their northernmost breeding site in Franz‐Josef Land (80°N), Russian Arctic. We tested the hypothesis that little auks still benefit from pristine arctic environmental conditions in this remote area. To this end, we analysed remote sensing data on sea‐ice and coastal glacier dynamics collected in our study area across 1979–2013. Further, we recorded little auk foraging behaviour using miniature electronic tags attached to the birds in the summer of 2013, and compared it with similar data collected at three localities across the Atlantic Arctic. We also compared current and historical data on Franz‐Josef Land little auk diet, morphometrics and chick growth curves. Our analyses reveal that summer sea‐ice retreated markedly during the last decade, leaving the Franz‐Josef Land archipelago virtually sea‐ice free each summer since 2005. This had a profound impact on little auk foraging, which lost their sea‐ice‐associated prey. Concomitantly, large coastal glaciers retreated rapidly, releasing large volumes of melt water. Zooplankton is stunned by cold and osmotic shock at the boundary between glacier melt and coastal waters, creating new foraging hotspots for little auks. Birds therefore switched from foraging at distant ice‐edge localities, to highly profitable feeding at glacier melt‐water fronts within <5 km of their breeding site. Through this behavioural plasticity, little auks maintained their chick growth rates, but showed a 4% decrease in adult body mass. Our study demonstrates that arctic cryosphere changes may have antagonistic ecological consequences on coastal trophic flow. Such nonlinear responses complicate modelling exercises of current and future polar ecosystem dynamics.     

Foraging location and range of White-chinned Petrels Procellaria aequinoctialis breeding in the South Atlantic

The foraging range and principal feeding areas of White‐chinned Petrels breeding at South Georgia were determined using satellite telemetry. Foraging trips during incubation lasted 12–15 days and covered 3000–8000 km and 2–11 days and 1100–5900 km during chick‐rearing. Adults covered less distance per day during chick‐rearing (71 km) than during incubation (91 km) but the proportion covered at night (47%) was the same. Mean (31–34 km/h) and maximum (80 km/h) flight velocities were similar during both periods of the breeding season and during day and night. Between incubation shifts, White‐chinned Petrels travelled to the Patagonian shelf; during chick‐rearing they foraged more extensively. Most locations were between 30° to 55°W and 52° to 60°W around South Georgia/Shag Rocks and south to the South Orkney Islands. Diet samples from known foraging locations suggested birds fed mainly on krill and squid. They caught the squid Brachioteuthis? picta and Galiteuthis glacialis around Shag Rocks/South Georgia and also at sites close to the South Orkney Islands; Illex argentinus on the Patagonian shelf. Dispersal of adults after breeding failure was south to the South Orkney Islands then west to the Falkland Islands. This study confirms that breeding White‐chinned Petrels are amongst the widest‐ranging of seabirds; they may minimise competition with other Procellariiformes in the South Atlantic by their more extensive foraging range. The nature and extent of their range also brings substantial risk of high mortality rate in South Atlantic long‐line fisheries.     

Time and travelling costs during chick‐rearing in relation to habitat quality in Little Owls Athene noctua

In many bird species, parents adjust their home‐ranges during chick‐rearing to the availability and distribution of food resources, balancing the benefits of energy intake against the costs of travelling. Over recent decades, European agricultural landscapes have changed radically, resulting in the degradation of habitats and reductions in food resources for farmland birds. Lower foraging success and longer foraging trip distances that result from these changes are often assumed to reduce the reproductive performance of parents, although the mechanisms are not well understood. We tested the behavioural response of chick‐rearing Little Owls Athene noctua to variation in habitat diversity in an agricultural landscape. We equipped females with GPS loggers and received adequate range‐use data for 19 individuals (6063–14 439 locations per bird). In habitats dominated by homogeneous cropland habitats, home‐ranges were over 12 ha in size, whereas in highly diverse habitats they were below 2 ha. Large home‐ranges were associated with increased flight activity (117% of that of birds in small home‐ranges) and distances travelled per night (152%), increased duration of foraging trips (169%) covering larger distances (246%), and reduced nest visiting rates (81%). The study therefore provides strong correlative evidence that Little Owls breeding in monotonous farmland habitats expend more time and energy for a lower benefit in terms of feeding rates than do birds in more heterogeneous landscapes. As nestling food supply is the main determinant of chick survival, these results suggest a strong impact of farmland characteristics on local demographic rates. We suggest that preserving and creating islands of high habitat diversity within uniform open agricultural landscapes should be a key target in the conservation of Little Owl populations.     

Exploitation of distant Antarctic waters and close neritic waters by short‐tailed shearwaters breeding in South Australia

Identifying the primary foraging grounds of abundant top predators is of importance in marine management to identify areas of high biological significance, and to assess the extent of competition with fisheries. We studied the search effort and habitat selection of the highly abundant short‐tailed shearwater Puffinus tenuirostris to assess the search strategies employed by this wide‐ranging seabird. During the chick‐rearing period 52 individuals were tracked performing 39 short foraging trips (1–2 days), and 13 long trips (11–32 days). First‐passage time analysis revealed that 46% of birds performing short trips employed area‐restricted searches, concentrating search effort at an average scale of 14 ± 5 km. Foraging searches were more continuous for the other 54%, who travelled faster to cover greater distances, with little evidence of area‐restricted searches. The prey returned indicated that continuous searchers consumed similar prey mass, but greater prey diversity than area‐restricted search birds. On long trips 23% of birds travelled 500–1000 km to neritic (continental shelf) habitats, showing weak evidence of preference for areas of higher chlorophyll a concentration, and foraged at a similar spatial scale to short trips. The other 76% performed rapid outbound flights of 1000–3600 km across oceanic habitats commuting to regions with higher chlorophyll a. The spatial scale of search effort in oceanic habitat varied widely with some performing broad‐scale searches (260–560 km) followed by finer‐scale nested searches (16–170 km). This study demonstrates that a range of search strategies are employed when exploiting prey across ocean basins. The trade‐offs between different search strategies are discussed to identify the value of these contrasting behaviours to wide‐ranging seabirds.     

Nest attendance and foraging movements of northern fulmars rearing chicks at Bjørnøya Barents Sea

We studied several aspects of the foraging ecology of fulmars rearing young chicks on Bjørnøya. To determine precisely the duration of foraging trips during the brooding period, we used an automated logging system that recorded the presence of fulmars fitted with transponders. We also tracked, with satellite transmitters, four parent fulmars during the brooding period, and two after the chick had been left alone. When brooding the chick, fulmars appeared to alternate very rapidly on the nest, with foraging trips lasting on average 8?h. This period appeared constraining for the birds since parents lost mass. The growth of chicks was dependent on the ability of the female (and not the male) to do short foraging trips. At this time birds are foraging at an average distance of 60?km from the colony, with birds concentrating on the shelf around Bjørnøya. They did not return from one trip to the next to the same foraging area. As the season progressed and the chicks were left alone on the nest, parents increased the duration and maximum range of foraging trips as well as the distance covered. However, they still perform a succession of relatively short foraging trips to the east of the Bjørnøya shelf but they interspersed these short trips with longer foraging trips. One bird returned twice to the same site along the Norwegian coast 570?km from Bjørnøya, the other foraged at 580?km in the mid-Barents Sea. Average flight speed including time spent on the water was 28?km/h and reached 70?km/h during bouts of more than 1?h when the bird was probably continuously in flight.     

Overnight foraging trips by chick‐rearing Nazca Boobies Sula granti and the risk of attack by predatory fish

Most tropical booby species complete breeding foraging trips within daylight hours, thus avoiding nights at sea. Nazca Boobies Sula granti are unusual in this respect, frequently spending one or more nights away from the nest. We used GPS dataloggers, time‐depth recorders, and changes in body weight to characterize foraging trips and to evaluate potential influences on the decisions of 64 adult Nazca Boobies to spend a night at sea, or to return to their chicks on Isla Española, Galápagos, in daylight hours. The tagged birds foraged east of Isla Española, undertaking both single‐day (2–15 h, 67% of trips) and overnight trips (28 h–7.2 days, 33%), and executing 1–19 foraging plunge‐dives per single‐day trip. Birds might forage longer if they are in nutritional stress when they depart, but body weight at departure was not correlated with trip length. Birds might be expected to return from longer trips with more prey for young, but they returned from single‐day and overnight trips with similar body weights, consistent with previous indications that Nazca Boobies forage until accumulating a target value of prey weight. Birds with a lower dive frequency during the first 5 h of a trip were more likely to spend the night at sea, suggesting that they might choose to spend the night at sea if prey capture success was low. At night, birds almost never dived and spent most of their time resting on the water’s surface (11.8–12.1 h, > 99% of the time between civil sunset and civil dawn). Thus, the night is an unproductive time spent among subsurface predators under low illumination. The birds’ webbed feet provided evidence of this risk: 24% of birds were missing > 25% of their foot tissue, probably due to attacks by predatory fish, and the amount of foot tissue lost increased with age, consistent with a cumulative risk across the lifespan. In contrast, other tropical boobies (Blue‐footed Sula nebouxii and Brown Boobies Sula leucogaster), which do not spend the night on the water, showed no such damage. These results suggest that chick‐rearing Nazca Boobies accept nocturnal predation risk on occasions of low prey encounter during a foraging trip’s first day.     

Strong temporal consistency in the individual foraging behaviour of Imperial Shags Phalacrocorax atriceps

Individual consistency in foraging behaviour can generate behavioural variability within populations and may, ultimately, lead to species diversification. However, individual‐based long‐term behavioural studies are particularly scarce in seabird species. Between 2008 and 2011, breeding Imperial Shags Phalacrocorax atriceps at the Punta León colony, Argentina, were tracked with GPS devices to evaluate behavioural consistency during their foraging trips. Within a breeding season, individuals were highly consistent in the maximum distances they reached from the shore and the colony, as well as in the time invested in flight and diving across consecutive days during early chick rearing. In addition, each individual had its specific foraging area distinct from the foraging area of other individuals. Comparing between early and late chick rearing in the same season, individuals were consistent, to a lesser degree, in the maximum distance they reached from the colony and the shore, increasing in consistency later on in the season. Within the season, females were more consistent than males in the maximum distance they moved from the colony and the shore, the sexes segregated in their foraging areas and individual females were segregated from one another. Twenty‐eight individuals tracked in different breeding seasons were marginally consistent in their trip durations and maximum distance reached from shore across seasons. Among seasons, foraging locations differed between sexes and among individual females. Individuals from this colony exhibited consistency over time in several aspects of foraging behaviour, which may be due to a combination of individual characteristics such as learning abilities, breeding experience or health, as well as targeted prey type and stability of the environment at this location.     

High foraging site fidelity and spatial segregation among individual great black‐backed gulls

Individual foraging site fidelity, whereby individuals repeatedly visit the same foraging areas, is widespread in nature, and likely benefits individuals through higher foraging efficiency and potentially, higher breeding success. It may arise as a consequence of habitat or resource specialisation, or alternatively, where resources are abundant or predictable, the partitioning of space might guarantee individuals exclusive foraging opportunities. We tracked seven adult great black‐backed gulls Larus marinus at a North Sea colony from early incubation to the end of the breeding season in 2016, providing a total of 1170 foraging trips over a mean ± SD tracking period of 67 ± 16 days. There was clear spatial segregation between individuals, with almost no overlap of their core areas (50% utilisation distribution) during incubation and chick‐rearing. Core areas were relatively small and there was high repeatability (R ± SE) in foraging parameters, including initial departure direction (0.73 ± 0.11), foraging range (0.41 ± 0.14) and cumulative distance travelled (0.19 ± 0.1) throughout the breeding season. Despite the low spatial overlap, there was little evidence of differential habitat use by individuals. The near‐exclusive individual foraging areas of this species, usually considered to be a generalist, indicate that where there is high resource availability throughout the breeding season and a small local population, individuals appear to adopt a territorial strategy which likely reduces intraspecific competition.     

Fledging success of little auks in the high Arctic: do provisioning rates and the quality of foraging grounds matter?

Long-lived birds often face a dilemma between self-maintenance and reproduction. In order to maximize fitness, some seabird parents alternate short trips to collect food for offspring with long trips for self-feeding (bimodal foraging strategy). In this study, we examined whether temporal and spatial variation in the quality of foraging grounds affect provisioning and fledging success of a long-lived, bimodal forager, the little auk (Alle alle), the most abundant seabird species in the Arctic ecosystem. We predicted that an increase in sea surface temperature (SST), with an associated decrease in the preferred Arctic zooplankton prey, would increase foraging trip durations, decrease chick provisioning rates and decrease chick fledging success. Chick provisioning and survival were observed during three consecutive years (2008–2010) at two colonies with variable foraging conditions in Spitsbergen: Isfjorden and Magdalenefjorden. We found that a change in SST (range 1.6–5.4 °C) did not influence trip durations or provisioning rates. SST was, however, negatively correlated with the number of prey items delivered to a chick. Furthermore, provisioning rates did not influence chick’s probability to fledge; instead, SST was also negatively correlated with fledging probability. This was likely related to the prey availability and quality in the little auk’s foraging grounds. Our findings suggest that predicted warmer climate in the Arctic will negatively influence the ability of parents to provide their chicks, and consequently, the fledging prospects of little auk chicks.     

Prey capture and selection throughout the breeding season in a deep‐diving generalist seabird,the thick‐billed murre

Generalist seabirds forage on a variety of prey items providing the opportunity to monitor diverse aquatic fauna simultaneously. For example, the coupling of prey consumption rates and movement patterns of generalist seabirds might be used to create three‐dimensional prey distribution maps (‘preyscapes’) for multiple prey species in the same region. However, the complex interaction between generalist seabird foraging behaviour and the various prey types clouds the interpretation of such preyscapes, and the mechanisms underlying prey selection need to be understood before such an application can be realized. Central place foraging theory provides a theoretical model for understanding such selectivity by predicting that larger prey items should be 1) selected farther from the colony and 2) for chick‐feeding compared with self‐feeding, but these predictions remain untested on most seabird species. Furthermore, rarely do we know how foraging features such as handling time, capture methods or choice of foraging location varies among prey types. We used three types of animal‐borne biologgers (camera loggers, GPS and depth‐loggers) to examine how a generalist Arctic seabird, the thick‐billed murre Uria lomvia, selects and captures their prey throughout the breeding season. Murres captured small prey at all phases of a dive, including while descending and ascending, but captured large fish mostly while ascending, with considerably longer handling times. Birds captured larger prey and dove deeper during chick‐rearing. As central place foraging theory predicted, birds travelling further also brought bigger prey items for their chick. The location of a dive (distance from colony and distance to shore) best explained which prey type was the most likely to get caught in a dive, and we created a preyscape surrounding our study colony. We discuss how these findings might aid the use of generalist seabirds as bioindicators.     

Inter‐colony foraging area segregation quantified in small colonies of Adélie Penguins

Theoretical models on the movement of colonial animals predict that neighbouring colonies may segregate their foraging areas, and many seabird studies have reported the presence of such segregations. However, these studies have often lacked the appropriate null model to test the effect of neighbouring colonies on foraging areas, especially in small colonies or in short‐ranging species. Here, we examined the foraging areas of Adélie Penguins Pygoscelis adeliae from two neighbouring (2 km apart) colonies by using bird‐borne GPS loggers. The field study was conducted at Hukuro Cove colony (104 pairs) and Mizukuguri Cove colony (338 pairs) in Lützow‐Holm Bay, East Antarctica. We obtained GPS tracks for 504 foraging trips from 48 chick‐rearing Adélie Penguins and quantified the degree of overlap in the foraging areas between two colonies. We also produced simulated movement tracks by using correlated random‐walks assuming no inter‐colony competition and quantified the degree of overlap in the simulated foraging areas. Finally, we compared the results from real GPS tracks with those from simulated tracks to examine the effect of neighbouring colonies on Adélie Penguin movement. The results indicate that the degree of overlap was significantly smaller in real tracks than in simulated tracks. In real tracks, the foraging area of the smaller Hukuro Cove colony extended to the other side of the larger Mizukuguri Cove colony, unlike in simulated tracks. Consequently, we suggest that Adélie Penguins from two neighbouring colonies segregated their foraging areas and that the larger colony appeared to affect the foraging area of the smaller colony.     

The individual counts: within sex differences in foraging strategies are as important as sex‐specific differences in masked boobies Sula dactylatra

Understanding how animals allocate their foraging time is a central question in behavioural ecology. Intrinsic factors, such as body mass and size differences between sexes or species, influence animals’ foraging behaviour, but studies investigating the effects of individual differences in body mass and size within the same sex are scarce. We investigated this in chick‐rearing masked boobies Sula dactylatra, a species with reversed sexual dimorphism, through the simultaneous deployment of GPS and depth‐acceleration loggers to obtain information on foraging movements and activity patterns. Heavier females performed shorter trips closer to the colony than lighter females. During these shorter trips, heavier females spent higher proportions of their flight time flapping and less time resting on the water than lighter females did during longer trips. In contrast, body mass did not affect trip duration of males, however heavier males spent less time flapping and more time resting on the water than lighter males. This may occur as a result of higher flight costs associated with body mass and allow conservation of energy during locomotion. Body size (i.e. wing length) had no effect on any of the foraging parameters. Dive depths and dive rates (dives h^?1) were not affected by body mass, but females dived significantly deeper than males, suggesting that other factors are important. Other studies demonstrated that females are the parent in charge of provisioning the chick, and maintain a flexible investment under regulation of their own body mass. Variation in trip length therefore seems to be triggered by body condition in females, but not in males. Consequently, shorter trips are presumably used to provision the chick, while longer trips are for self‐maintenance. Our findings underline the importance of accounting for the effects of body mass differences within the same sex, if sex‐specific foraging parameters in dimorphic species are being investigated.     

Sex differences in Adélie penguin foraging strategies

Consistent sex differences in foraging trip duration, feeding locality and diet of breeding Adélie penguins (Pygoscelis adeliae) were demonstrated at two widely separated locations over several breeding seasons. Differences in foraging behaviour were most pronounced during the guard stage of chick rearing. Female penguins made on average longer foraging trips than males, ranged greater distances more frequently and consumed larger quantities of krill. In contrast, males made shorter journeys to closer foraging grounds during the guard period and fed more extensively on fish throughout chick rearing. Mean guard stage foraging trip durations over four seasons at Béchervaise Island, Eastern Antarctica and over two seasons at Edmonson Point, Ross Sea ranged between 31 and 73 h for females and 25 and 36 h for males. Ninety percent of males tracked from Béchervaise Island by satellite during the first 3 weeks post-hatch foraged within 20 km of the colony, while the majority (60%) of females travelled to the edge of the continental shelf (80–120 km from the colony) to feed during this period. Received: 10 December 1997 / Accepted: 10 April 1998     

Intra-seasonal variation in zooplankton availability,chick diet and breeding performance of a high Arctic planktivorous seabird

Different phenological responses to climate changes by species representing preys and predators may lead to mismatch between functionally dependent components of an ecosystem, with important effects on its structure and functioning. Here, we investigate within-season variation in zooplankton availability, chick diet composition and breeding performance of a small planktivorous seabird, the little auk (Alle alle) in two large colonies in Hornsund and Magdalenefjorden, Spitsbergen, differing in synchrony of breeding (11-day vs. 22-day hatching period, respectively). Assuming similar zooplankton phenology and existing differences in duration of the little auk breeding period, we expected lower availability of the preferred food in the less synchronized colony in Magdalenefjorden and in consequence a negative effect on nestling body mass and survival. We found that in both colonies Calanus glacialis (copepodite stage CV) was the most important prey item in the chick diet making up 68–87 % of the biomass and energy of all prey items. The only exception was the end of the chick-rearing period in Magdalenefjorden, when contribution of this prey item was significantly lower (24–26 %). Thus, late breeders in Magdalenefjorden were apparently mismatched regarding C. glacialis CV availability. However, the hatching date did not affect birds fitness (reproductive output and chick pre-fledging mass) significantly. Results of our study indicate that little auks breeding on Spitsbergen can respond to a wide range of environmental conditions and prey availabilities through the plasticity of their foraging behaviour, which may help them to maintain their optimum fitness level in changing and unpredictable environments.     

Flexibility in the bimodal foraging strategy of a high Arctic alcid, the little auk Alle alle

A bimodal foraging strategy has previously been described for procellariiform seabird species and is thought to have evolved in response to local resource availability being too low for adult birds to meet chick requirements and simultaneously maintain their own body condition. Here, we examine the dual foraging trip pattern of an alcid, the little auk Alle alle , at five colonies with contrasting oceanographic conditions. In spite of large variation in local conditions, little auks at all colonies showed the same general pattern of alternating a single long-trip with several consecutive short-trips. However, we found that the foraging pattern was flexible and could be adjusted at three levels: (1) the length of long-trips, (2) the frequency of short-trips, and (3) the total time spent foraging. Birds facing unfavorable conditions increased the duration of long-trips and reduced the number of short-trips. These adjustments resulted in reduced provisioning rates of chicks despite the fact that birds also increased the time allocated to foraging. Travel times during foraging trips were positively correlated to the total duration of the trip suggesting that differences in trip length among colonies were partly driven by variation in the distance to foraging areas. Most birds spent substantially more time traveling during long compared to short-trips, indicating that they accessed distant foraging areas during long-trips but remained close to the colony during short-trips. However, the difference in travel times was small at the site with the most favorable conditions suggesting that bimodal foraging in the little auk may be independent of the existence of high-quality areas at distance from the breeding ground.     

Breeding phased dependent oxidative balance in a small High Arctic seabird,the little auk

Reproduction is a demanding phase of bird's life cycle and imposes various physiological challenges. Increasing oxidative stress (OS) has been proposed as one of the costs associated with reproduction. In this study, we investigate the level of OS in a small Arctic seabird, the little auk Alle alle in relation to sex and phase of breeding (incubation, chick‐rearing). We also examine whether OS is related to the birds leucocyte profile. We expected increase in OS with the progress of breeding period (due to increasing energetic demands) and higher values in females (due to high initial investments in production of a large egg). Surprisingly, we found higher OS during incubation compared to chick‐rearing period, suggesting that incubation is a highly demanding reproductive phase in terms of oxidative balance. We suggests that those changes may be attributed to changes in hormone levels affecting oxidative status. Also, in contrasts to our expectations, we did not find sex differences in OS throughout the studied periods of breeding. Finally, we found positive relationships between OS level and haematological parameters: heterophils/lymphocytes ratio, number of leucocytes and lymphocytes per 10 000 erythrocytes, suggesting the effect of the oxidative stress on the immunological system.