Foraging strategy of the little auk Alle alle throughout breeding season – switch from unimodal to bimodal pattern

Energy and time allocation differs between incubation and chick‐rearing periods, which may lead to an adjustment in the foraging behaviour of parent birds. Here, we investigated the foraging behaviour of a small alcid, the little auk Alle alle during incubation and compared it with the chick‐rearing period in West Spitsbergen, using the miniature GPS (in Hornsund) and temperature loggers (in Magdalenefjorden). GPS‐tracking of 11 individuals revealed that during incubation little auks foraged 8–55 (median 46) km from the colony covering 19–239 (median 120) km during one foraging trip. Distance from the colony to foraging areas was similar during incubation and chick‐rearing period. During incubation 89% of foraging positions were located in the zone over shallower parts of the shelf (isobaths up to 200–300 m) with sea surface temperature below 2.5°C. Those environmental conditions are preferred by Arctic zooplankton community. Thus, little auks in the Hornsund area restrict their foraging (both during the incubation and chick‐rearing period) to the area under influence of cold, Arctic‐origin water masses where its most preferred prey, copepod Calanus glacialis is most abundant. The temperature logger data (from 4 individuals) indicate that in contrast to the chick‐rearing period, when parent birds alternated short and long trips, during the incubation they performed only long trips. Adopting such a flexible foraging strategy allows little auks to alter their foraging strategy to meet different energy and time demands during the two main stages of the breeding.     

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.     

A fine-scale time budget of Cape gannets provides insights into the foraging strategies of coastal seabirds

Central-place foragers organize their feeding trips both to feed themselves and to provide their offspring with food. In seabirds, several long-range foragers have been shown to alternate long and short trips to balance these dual needs. However, the strategies of short-range foragers remain poorly understood. We used a precise, miniaturized motion sensor to examine the time budget of 20 breeding Cape gannets, Morus capensis, foraging off the coast of South Africa. Birds stayed at sea for 5.5-25.3 h, occasionally spending the night at sea. The large number of isolated dives and extended flight time observed during these overnight trips suggested that birds either experienced poor foraging conditions or exploited more distant, yet more profitable prey patches. Conversely, birds that stayed at sea for less than 1 day had relatively consistent activity patterns. Most of these birds (88%) foraged actively at the beginning and at the end of the foraging trip. These feeding bouts were separated by protracted periods of sitting on the sea surface. Such resting periods probably allow birds to digest the food ingested during the first part of the foraging trip, so they initially feed themselves, and then obtain food for their chick on the way back to the breeding site.     

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.     

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.     

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.     

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.     

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.     

Visual prey availability and distribution of foraging little auks (Alle alle) in the shelf waters of West Spitsbergen

As diving seabirds use vision underwater, it is presumed they should preferentially select sites where their preferred food items are not only abundant but also clearly visible. To test this, we studied the optical properties of the seawater in the West Spitsbergen Shelf, in combination with zooplankton abundance in the feeding grounds of the planktivorous little auks from the nearby colonies in Hornsund. We estimated the relative attractiveness of the foraging sites using a novel parameter—visual prey availability (VPAv), which relates density and proportion of the preferred food item (Calanus glacialis) of the little auk, in total zooplankton, to the optical properties of the seawater. We found a significant positive correlation between the density of foraging little auks and VPAv values. Birds chose areas where C. glacialis was both abundant and clearly visible, because of the clarity of the water and low proportion of other zooplankton species. The birds avoided foraging over the warmer Atlantic-type waters, characterised by a high abundance of zooplankton taxa mostly ignored by birds and where VPAv values were low. VPAv values could potentially also be applied to other visual planktivores for which prey preference and visual acuity are known.     

Foraging patterns of breeding Dolphin Gulls Larus scoresbii at Punta Tombo, Argentina

The Dolphin Gull Larus scoresbii is a little-known, rare species endemic to southern South America. Knowledge of its feeding ecology is essential for development of management and conservation strategies. To obtain information on their use of food resources and on the frequency and duration of their foraging trips, we followed seven individuals by radiotelemetry during the breeding season of 2002 at the Punta Tombo Reserve, Argentina. In 99% of mapped locations ( n  = 4069) the Gulls were within 2.4 km of their colony. Dolphin Gulls foraged mostly at the colonies of three other species, feeding on Magellanic Penguin Spheniscus magellanicus and Imperial Cormorant Phalacrocorax atriceps regurgitates and on Southern Sea Lion Otaria flavescens faeces. The sea lion colony was the most frequently visited foraging area, accounting for 64% of the total ( n  = 260 trips). A similar trend was observed during both the incubation (60%, n  = 38 trips) and the chick (64%, n  = 40 trips) stages. The mean number of trips per day was 3.8 (range = 3–5), with a mean duration of 123 min (sd = ±27). Trip duration differed between foraging areas, being significantly longer at the sea lion than at the penguin and cormorant colonies. The small foraging range given by its particular feeding strategy suggests that Dolphin Gulls, in contrast to most other seabirds that range over large expanses of ocean in search of food, may be effectively conserved within protected areas during the breeding season.     

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.     

Satellite tracking of male rockhopper penguins Eudyptes chrysocome during the incubation period at the Falkland Islands

The foraging patterns of ten male rockhopper penguins Eudyptes chrysocome from the Falkland Islands were recorded during the incubation period by using satellite telemetry. Irrespective of study site and year, two different foraging areas could be identified. Three foraging trips were directed towards the slope of the Patagonian Shelf c. 140 km to the northeast of the breeding colony and these trips had a duration of 11–15 days. The seven other rockhopper penguins travelled c. 400 km towards the edge of the Falkland Islands' waters; all these foraging trips followed an anti-clockwise direction and lasted 16–27 days. The calculated mean daily travelling speed, based on the time spent underwater and the distance covered between two positions, was significantly higher in birds travelling to the edge of the Falkland Islands' waters compared with those foraging at the shelf slope (4.4±1.6 km/h vs. 3.4±2.0 km/h, respectively). The consistent foraging patterns exhibited during the long trips may be linked to the Falklands current, allowing the penguins to reach remote areas while reducing their energy expenditure. Potential interactions between commercial fisheries and hydrocarbon exploration are discussed.     

Foraging behaviour and habitat partitioning of two sympatric cormorants in Patagonia, Argentina

Radiotelemetry was used to assess the distribution and diving behaviour of Rock Shags Phalacrocorax magellanicus and Red-legged Cormorants Phalacrocorax gaimardi breeding in sympatry, and Rock Shags breeding in isolation. When breeding in sympatry there was little overlap in the foraging locations of the two species, with the highest densities of each species separated by 10 km. Red-legged Cormorants fed significantly closer to the breeding colony than did Rock Shags and undertook shorter foraging trips, making almost twice as many foraging trips per day as Rock Shags. Rock Shags breeding in isolation had a shorter foraging range than the birds breeding in sympatry with Red-legged Cormorants and foraging trip duration was significantly shorter. However, the number of feeding trips per day was similar between areas of sympatry and allopatry. Differences in the foraging ecology of Rock Shags in areas of sympatry and allopatry may be due to interspecific competition, which forces niche differentiation. The distance between foraging sites, the speed of movement of the prey, a species tendency to move into prey-depleted areas and the length of the breeding season (during which the birds are constrained to be in the same area) may play critical roles in determining the extent to which differential area use by competitors is a strategy that benefits both parties.     

Diet and diving behaviour of European Storm Petrels Hydrobates pelagicus in the Mediterranean (ssp. melitensis)

Capsule Unlike Atlantic populations, which feed on krill, Mediterranean populations feed mainly on pelagic fish Gymnammodites cicerellus.

Aims To determine the diet and dive depth of the Mediterranean subspecies of European Storm Petrels Hydrobates pelagicus melitensis.

Methods Analysis of regurgitates of adults arriving at the colony for chick feeding and by determination of dives depth using the capillary tube method.

Results The main prey is Gymnammodites cicerellus, a pelagic fish. Storm Petrels dive for their prey and can reach up to 5 m in depth. They also make short foraging trips just outside the colony where they capture Opossum Shrimps Misydacea.

Conclusions European Storm Petrels in the Mediterranean exploit pelagic fish which are taken by diving. This contrasts with the Atlantic populations which feed mainly on krill. Mediterranean birds also feed on Opossum Shrimps Mysidacea during short foraging trips made at night just outside the colony. Differences in diet between long and short foraging trips may be because adults have to forage for both themselves and their chicks.     

Low incubation investment in the burrow‐nesting Crab Plover Dromas ardeola permits extended foraging on a tidal food resource

We used GPS data‐loggers, video‐recordings and dummy eggs to assess whether foraging needs may force the low incubation attentiveness (< 55%) of the Crab Plover Dromas ardeola, a crab‐eating wader of the Indian Ocean that nests colonially in burrows. The tidal cycle was the major determinant of the time budget and some foraging trips were more distant from the colony than previously known (up to 26 km away and lasting up to 45 h). The longest trips were mostly made by off‐duty parents, but on‐duty parents also frequently left the nest unattended while foraging for 1–7 h. However, the time spent at the colony area (47%) and the time spent roosting on the foraging grounds (16%) would have allowed almost continuous incubation, as in other species with shared incubation. Therefore, the low incubation attentiveness is not explained by the need for long foraging trips but is largely dependent on a high intermittent rhythm of incubation with many short recesses (5.8 ± 2.6 recesses/h) that were not spent foraging but just outside the burrow or thermoregulating at the seashore. As a result, the eggs were warmed on average only 1.7 °C above burrow temperature, slightly more during high tide periods and when burrow temperature was lower between 20:00 and 10:00 h, only partly counteracting the temperature fluctuations of the incubation chamber. These results suggest that low incubation attentiveness is due to the favourable thermal conditions provided by safe nesting burrows and by the hot tropical breeding season, a combination that allows simultaneous foraging by parents and the exploitation of distant foraging grounds. Why Crab Plovers engage in many short recesses from incubation still remains to be clarified but the need to thermoregulate at the seashore and to watch for predators may play a role.     

Water temperature sampling by foraging Brünnich's Guillemots with bird-borne data loggers

We describe the features of waters where seabirds were feeding by sampling vertical water temperature profiles with data loggers mounted on five Brünnich's Guillemots in Svalbard, Norway. The guillemots foraged in a cold water (−0.5–0.5°C SST (sea surface temperature)) by making 1.8 dive bouts in short trips (32–257 min duration) as well as in moderate (0.5–2.0°C SST) and warm waters (2.5–4.0°C SST) by making 6.0 dive bouts during long trips (411–688 min duration). Judging from outbound flying time (15.7–24.4 min), time between dive bouts (23.9–43.3 min) and water types, the birds probably fed in fjord or coastal waters during short trips and in both coastal and offshore waters during long trips. Water temperature and diving behaviour can be simultaneously recorded by small data loggers, which therefore will provide useful information on marine features and foraging activity of top predators.     

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.     

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.     

The effects of food abundance on breeding performance and adult time budgets of Guillemots Uria aalge

The breeding performance, food fed to chicks and adult time budgets of Guillemots Uria 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 insufficient to compensate for 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 available for foraging.     

How area restricted search of a pelagic seabird changes while performing a dual foraging strategy

Movements and foraging strategies of marine predators should cope with the hierarchical spatial distribution of resources. Therefore, in order to predict the at‐sea distribution of aerial predators, it is crucial to understand the factors governing trajectory decisions at different scales. Using first passage time (FPT) analysis on precision tracking information (GPS‐loggers data) we were able to examine the foraging strategy of Cory's shearwaters Calonectris diomedea and to detect the adoption of area‐restricted search (ARS), measuring the scale and duration of this behaviour. Data were collected from three different populations foraging in different oceanographic conditions. During long excursions birds only commuted between their colony and prey patches, while on their short movements birds increased the amount of looping movements. On short trips, birds addopted ARS behaviour at an average scale of 18 km and at a second nested scale of around 2 km. When engaging in long trips, first scale of ARS occurred on average at about 67 km of radii and than a second nested scale at a radii of 24 km. Overall, the different populations showed foraging patterns matching the habitats exploited: a) at smaller scales of ARS, sea‐surface temperature, chlorophyll‐a concentration and depth influenced the time of residence (i.e. FPT) of birds (with variations at a population level); b) at larger scales of ARS, FPT increased within regions of higher gradients of sea‐surface temperature, chlorophyl‐a concentration and depth. This study demonstrates that Cory's shearwaters adopt scale‐dependent adjustments of movement in relation to the hierarchical distribution of the environment they exploit, matching the scale and duration of ARS with the hierarchical distribution of the environmental features.