Estimating prey capture rates of a planktivorous seabird,the little auk (<Emphasis Type="Italic">Alle alle</Emphasis>), using diet,diving behaviour,and energy consumption

Interpreting the impact of environmental change on food webs requires a clear understanding of predator–prey interactions. Such knowledge is often lacking in the marine environment where the foraging behaviour and prey requirements of some of the major top-predators remains mysterious. For example, very little is known about the underwater foraging behaviour of the little auk, the most numerous seabird in the North Atlantic. In 2004, we used time–depth-recorders at two breeding colonies in East Greenland to examine the diving behaviour of this small, planktivorous seabird during the chick-rearing period. Due to technical difficulties data were only collected for four individuals, but recordings showed that birds dive up to 240 times a day to maximum depths of 27 m (average 10 m), with maximum dive durations of 90 s (average 52 s). In addition, we collected the chick meals from 35 individuals, which were dominated by Calanus copepods (95%), and also determined the field metabolic rates (FMR) of 14 individuals using the doubly labelled water technique, which averaged 609.9 kJ day⁻¹. We integrated information on diving duration with chick diet and FMR to estimate the prey requirements and underwater capture rates of little auks using a Monte Carlo simulation. Chick-rearing little auks needed to catch about 59,800 copepods day⁻¹, which is equivalent to about six copepods caught per second spent underwater. These astonishing results strongly suggest that little auks are, at least partly, filter-feeding, and underline the importance of highly productive, cool marine areas that harbour dense patches of large, energy-rich copepods.     

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.     

The feeding ecology of little auks raises questions about winter zooplankton stocks in North Atlantic surface waters

Copepods are essential components of marine food webs worldwide. In the North Atlantic, they are thought to perform vertical migration and to remain at depths more than 500 m during winter. We challenge this concept through a study of the winter feeding ecology of little auks (Alle alle), a highly abundant planktivorous seabird from the North Atlantic. By combining stable isotope and behavioural analyses, we strongly suggest that swarms of copepods are still available to their predators in water surface layers (less than 50 m) during winter, even during short daylight periods. Using a new bioenergetic model, we estimate that the huge number (20–40 million birds) of little auks wintering off southwest Greenland consume 3600–7200 tonnes of copepods daily, strongly suggesting substantial zooplankton stocks in surface waters of the North Atlantic in the middle of the boreal winter.     

Composition of chick meals from one of the main little auk (Alle alle) breeding colonies in Northwest Greenland

During the last decade, increasing information on little auk (Alle alle) biology, ecology and behaviour has been reported. However, only a few of these studies have focused on the breeding population in the Avanersuaq (Thule) district of Northwest Greenland, where 80 % of the global little auk population is estimated to breed. This study reports on the chick diet composition from one of the largest colonies, the Paakitsoq colony, located on the south-eastern margin of the North Water (NOW) Polynya. Results revealed the highest proportion of Calanus hyperboreus, a large lipid-rich copepod, in chick diet reported for any little auk colony. Results confirmed that the cold, highly productive waters of the NOW Polynya are favourable foraging grounds for the little auks during the breeding season. Species diversity within and between the chick meals was low, which probably reflects a high availability of a few preferred prey species. Individual chick meals were generally low in number of prey items and total energy content compared with other published results. This may be explained by a higher feeding frequency or by the samples being collected late in the breeding period (during late chick rearing), when chicks have a reduced growth rate and may require less energy than at earlier developmental stages.     

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.     

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.     

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.     

Biologging, remotely-sensed oceanography and the continuous plankton recorder reveal the environmental determinants of a seabird wintering hotspot

Marine environments are greatly affected by climate change, and understanding how this perturbation affects marine vertebrates is a major issue. In this context, it is essential to identify the environmental drivers of animal distribution. Here, we focused on the little auk (Alle alle), one of the world's most numerous seabirds and a major component in Arctic food webs. Using a multidisciplinary approach, we show how little auks adopt specific migratory strategies and balance environmental constraints to optimize their energy budgets. Miniature electronic loggers indicate that after breeding, birds from East Greenland migrate >2000 km to overwinter in a restricted area off Newfoundland. Synoptic data available from the Continuous Plankton Recorder (CPR) indicate that this region harbours some of the highest densities of the copepod Calanus finmarchicus found in the North Atlantic during winter. Examination of large-scale climatic and oceanographic data suggests that little auks favour patches of high copepod abundance in areas where air temperature ranges from 0°C to 5°C. These results greatly advance our understanding of animal responses to extreme environmental constraints, and highlight that information on habitat preference is key to identifying critical areas for marine conservation.     

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.     

Climate warming decreases the survival of the little auk (Alle alle), a high Arctic avian predator

Delayed maturity, low fecundity, and high adult survival are traits typical for species with a long‐life expectancy. For such species, even a small change in adult survival can strongly affect the population dynamics and viability. We examined the effects of both regional and local climatic variability on adult survival of the little auk, a long‐lived and numerous Arctic seabird species. We conducted a mark‐resighting study for a period of 8 years (2006‐2013) simultaneously at three little auk breeding sites that are influenced by the West Spitsbergen Current, which is the main carrier of warm, Atlantic water into the Arctic. We found that the survival of adult little auks was negatively correlated with both the North Atlantic Oscillation (NAO) index and local summer sea surface temperature (SST), with a time lag of 2 and 1 year, respectively. The effects of NAO and SST were likely mediated through a change in food quality and/or availability: (1) reproduction, growth, and development of Arctic Calanus copepods, the main prey of little auks, are negatively influenced by a reduction in sea ice, reduced ice algal production, and an earlier but shorter lasting spring bloom, all of which result from an increased NAO; (2) a high sea surface temperature shortens the reproductive period of Arctic Calanus, decreasing the number of eggs produced. A synchronous variation in survival rates at the different colonies indicates that climatic forcing was similar throughout the study area. Our findings suggest that a predicted warmer climate in the Arctic will negatively affect the population dynamics of the little auk, a high Arctic avian predator.     

Breeding density and population of little auks (Alle alle) in a Northwest Greenland colony

The little auk population of the Thule district in Greenland is generally believed to be the largest anywhere and to comprise more than half of the world population, although published numbers have largely been conjectural. In 1996/1997 we estimated breeding density of little auks at Hakluyt Island in this district by colour-marking a number of birds in three study plots and subsequently counting marked and unmarked birds present in the plots. The density estimate considered most representative of the colony was 1.8 birds/m² or 0.73 pairs/m² (±7%). From surveys of the inhabitated area of the scree slopes, this density implies a total little auk population for the island of 130,000 pairs. An extrapolation to the entire Thule district suggests a population in the area of at least 15 million pairs, which is in general agreement with previously published assumptions. Accepted: 3 January 2000     

Temporal variations in the diet of pearly razorfish Xyrichtys novacula (Osteichthyes: Labridae)

The diet of pearly razorfish Xyrichtys novacula, caught monthly along the shores of the Island of Lampedusa, appeared to be mainly composed of crustaceans, followed by colonial ascidians, molluscs and polychaetes. Among prey, sand dwellers and phanerogam‐associated species were recorded. In winter months, the diet was characterized by a small number of prey items, dominated by colonial ascidians, while in spring and summer a wider prey array was recorded. Dietary indices show that X. novacula do not strictly exploit benthic prey but also pelagic organisms, such as copepods. This feeding behaviour reached its peak in March and October, when the abundance of primary consumers was at its highest after phytoplankton blooms. Furthermore, X. novacula caught prey organisms according to their availability and seasonal patterns during their life cycles, irrespective of fish size.     

Oceanographic-related variation in the bone sizes of extinct great auks

Summary We measured the bones of extinct great auks Pinguinis impennis that were killed during recent centuries on Funk Island off the northeast coast of Newfoundland. Comparisons of these measurements with those taken elsewhere suggest that great auks from Funk Island, which is situated in a Low Arctic oceanographic region, were larger than conspecifics from Boreal oceanographic regions. This finding is supported by extant alcid species that inhabit Boreal, Low Arctic or Boreal through High Arctic ocean regions and tend to increase in body size with increasing latitude (generally decreasing sea surface temperature). We suggest that paleoecological sea surface temperatures and food webs may have favored oceanographic-related variation in body sizes of great auks. The variances of the bone sizes of great auks from Funk Island were not less than those of a sample of great auk bones collected from Scandinavian archaeological sites that cover an extensive geographic range and that span seven millenia. This finding is inconsistent with a previously suggested latitudinal cline in body size among great auks in Scandinavia. Research techniques and studies that could address questions of great auk feeding ecology and population genetics are considered.     

Feeding habits of invasive American mink (Neovison vison) in northern Germany—potential implications for fishery and waterfowl

In order to collect ecological data of invasive American mink (Neovison vison) at a fishpond area in northeastern Germany, we conducted a telemetry study in which 14 mink were radio-tracked. During this project, 2,502 scats from radio-tracked individuals were collected in the period from October 2003 to October 2005. Investigated mink principally prey on fish, small mammals and birds (eggs inclusive), whereas amphibians, reptiles and invertebrates were caught infrequently. Analysing mink scats of different seasons, we found significant seasonal variations of diet composition. In spring, fish, mammals and birds were hunted in similar amounts. During summer, birds made up the main part of the diet followed by mammals. In autumn, the proportion of birds in the mink diet decreased, whereas fish gained in importance. This trend continued during the winter period, when mink preyed almost exclusively on fish. Amphibians, crustaceans, insects, molluscs and reptiles were found only occasionally in scat samples. Among birds, the mink preyed mainly on the Eurasian Coot (Fulica atra) followed by the Mallard (Anas platyrhynchos). Mammalian prey was clearly dominated by the water vole (Arvicola terrestris) and among fish, mink hunted especially perch (Perca fluviatilis), roach (Rutilus rutilus) and carp (Cyprinus carpio). Results clearly demonstrate that mink is an opportunistic predator, which hunts its prey according to availability and vulnerability, respectively. Despite the high portions of fish in their autumn and winter diet, the economic damage caused by mink seems to be negligible. However, high predation rates on birds during the breeding season indicate a potential negative impact of mink on waterfowl.     

Weak population genetic differentiation in the most numerous Arctic seabird,the little auk

Quantifying patterns of genetic diversity and differentiation among populations of Arctic birds is fundamental for understanding past and ongoing population processes in the Arctic. However, the genetic differentiation of many important Arctic species remains uninvestigated. Here, phylogeography and population genetics were examined in the worldwide population of a small seabird, the little auk (dovekie, Alle alle)—the most numerous avian species of the Arctic ecosystem. Blood samples or feathers were collected from 328 little auks (325 from the nominate subspecies and 3 from the A. a. polaris) in nine main breeding aggregations in the northern Atlantic and one location from the Pacific Ocean. The mtDNA haplotypes of the two subspecies were not segregated into separate groups. Also, no genetic structure was found within the nominate race based on microsatellite markers. The level of genetic differentiation among populations was low yet significant (mean F _ST = 0.005). Some pairwise F _ST comparisons revealed significant differences, including those involving the most distant Pacific colony as well as among some Atlantic populations. Weak population differentiation following the model of isolation by distance in the little auk is similar to the patterns reported in other high-Arctic bird species, indicating that a lack of distinct genetic structure is a common phenomenon in the Arctic avifauna.     

Diet and prey selectivity of the mesopelagic lanternfish Myctophum asperum from the Perth Canyon,Western Australia

Ichthyological Research - The diet and prey selectivity of Myctophum asperum were examined from the Perth Canyon (Western Australia). Calanoid copepods were consumed by 83% of the M. asperum...     

Diet and fish stock availability as possible factors in the mass death of auks in the North Sea

In February 1983 more than 30 000 birds died on the North Sea coasts of England and Scotland. The birds were unoiled, and there was no immediately obvious cause of death. Stomach contents of auks killed in this incident were examined to ascertain whether they provided any indication of the causal factors. Analysis of food remains showed differences between sample sites and between guillemots, Una aalge (Pont.), and razorbills, Alca tarda L. In all areas gadoid remains comprised a higher, and sandeels a lesser, proportion of the stomach contents in guillemots than in razorbills. The relative importance of clupeid remains varied with area. These differences, combined with current trends in the fisheries biology of major prey species, provided an ecological basis for a hypothesis explaining the incident. It was concluded that starvation was probably a major factor in the death of the auks, and that this may have been related to a combination of adverse weather conditions and changing patterns of abundance in North Sea sprat, Sprattus sprattus (L.), populations.     

Summer feeding strategy of the little auk (Alle alle) from Bjørnøya, Barents Sea

Summer food of the little auks nesting on Bjornoya consisted of large copepods, decapod larvae and pelagic amphipods. Food items of 4–6?mm constituted the most common prey size fraction. This prey size range appears to be optimal (most profitable) with regard to the balance between mean individual mass and density of items in the surface layers of the sea. Little auks from Bjørnøya only sporadically take the abundant copepod Calanus finmarchicus (2–3.5?mm in length) but select the much less abundant and larger Calanus glacialis (3–6?mm).RID="ID=" Acknowledgements The authors would like to thank Dr. Monika Normant from the University of Gdansk for her assistance with caloric measurments, and Lech Illiszko and Adam Wajrak for their help with the fieldwork on Bj&;oslash;rn&;oslash;ya.RID="ID=" Correspondence to J. M. We&;cedil;s&;lstrok;awski     

Seabird species‐ and assemblage‐level isotopic niche shifts associated with changing prey availability during breeding in coastal Newfoundland

Shifting prey availability can lead to altered species interactions, indicated by variation in the dietary niche breadth and position of species within an assemblage. On the Newfoundland coast, annual inshore spawning migration of the dominant forage fish, Capelin Mallotus villosus, provides an excellent opportunity to investigate the influence of varying prey availability on dietary niche breadth and position among species. During June–August 2017, we investigated species‐ and assemblage‐level dietary responses to shifting Capelin availability of three Capelin‐eating, sympatrically breeding auk species, the Atlantic Puffin Fratercula arctica, Razorbill Alca torda and Common Murre Uria aalge. The diet of Leach's Storm Petrels Oceanodroma leucorhoa, which breed alongside the three auk species but are not known to rely on Capelin, was also examined to determine dietary shifts throughout breeding that were unrelated to Capelin availability. We quantified stable isotope ratios (δ¹⁵N, δ¹³C) in seabird blood components (plasma, cellular component) collected both before and after spawning Capelin arrived in the study area and compared isotopic niche breadth within a Bayesian framework. At the species level, auk trophic position increased and isotopic niche breadth narrowed after Capelin arrived, suggesting a more Capelin‐based diet. Simultaneously, trophic diversity of the auk assemblage, reflecting the extent of spacing among niches of species, decreased after spawning Capelin arrived inshore. Contrastingly, increased trophic position but broader isotopic niche breadth during higher relative to lower Capelin availability for Leach's Storm Petrel confirm that this species is probably not affected by the inshore arrival of Capelin, but instead that isotopic changes may be more related to a shift in breeding stage to chick‐rearing. Overall, our findings reiterate the importance of Capelin as a prey resource for breeding auks in coastal Newfoundland, but that the degree of reliance on Capelin varies among species, possibly allowing coexistence of these ecologically similar species. The findings highlight potential changing species interactions, such as increased competition, under declines in Capelin biomass.     

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.