Winter diet of the little auk (Alle alle) in the Northwest Atlantic

The little auk (Alle alle) is one of the most numerous seabirds in the world, but their winter prey selection has never been thoroughly studied. In the present study, we analyzed the proventricular contents of 205 little auks caught in coastal areas off southwest Greenland during December–February and off Newfoundland in March. Large Calanoid copepods are known to be the main prey during summer. We found krill to become the crucial winter prey in both areas, followed by Themisto spp. and young capelin (Mallotus villosus). No difference was found between the diet of juvenile and adult birds. Copepods constituted around 1 % of the diet and close to all copepods were in birds caught near Cape Farwell in December. These findings provide new and important insight into the forage ecology of the little auk, and they support other studies showing that large Calanoid copepods in the Arctic decent for winter hibernation at depths that are below the diving range of the little auks. More studies to determine offshore diet and annual variation are, however, needed in order to get a more complete picture.     

Distribution of little auk (Alle alle) breeding colonies in Thule District, northwest Greenland

Large areas with hydrocarbon prospects offshore northwest Greenland necessitate seabird studies in this region. The little auk population breeding in Thule District is the largest seabird population in northwest Greenland, as well as the largest known population of this species. In 1994 and 1995 we mapped the distribution of little auk colonies in Thule District during aerial surveys. Most colonies are situated in sloping screes facing the sea, some as far as 11 km from the coast in valleys or facing glaciers. The total horizontal extent of the colonies is about 400 km. Our surveys generally confirm previous and local knowledge, although the information obtained is more detailed. Received: 11 November 1996 / Accepted: 21 September 1997     

On the importance of fat reserves for the little auk (Alle alle) chicks

Summary The pattern of fat deposition and inter-year variation in the amount of deposited fat, lean dry mass (LDM) and water were studied in the chicks of little auk (Alle alle), an arctic alcid, at Spitsbergen in the 1984, 1986 and 1987 breeding seasons. The lipid index (lipid mass/LDM) rose from 0.29 at hatching to about 0.70 at the age of maximum body mass attained in the nest, the highest yet recorded for seabirds except procellariiforms. Just before fledging the index dropped to 0.38. Fat mass in 1984 chicks and also fresh body mass of older 1984 chicks was significantly lower than in the other two seasons, but there were no statistical year-to-year differences in LDM's of chicks. The amount of fat in chicks was significantly correlated with certain weather parameters. Fasting capability (FC) of the chicks (the time that chicks could survive when using only their lipid stores) was calculated from fat mass and previous data on metabolic rates of chicks. The FC's of all 108 chicks collected in three seasons were greater than maximum interval between two consecutive feeds to chicks reported in the study colony. Also the estimated fat reserves of 99.7% of chicks weighed in 1984 were sufficient to support them for longer than the maximum recorded intervals between feeds. In relation to death by starvation, little auk chicks, at least up to the age of maximum body mass, have a substantial surplus of fat. This is contrary to the hypothesis that fat depots in seabird chicks assist survival during the periodic fasts resulting from unpredictable feeding conditions.     

The finding of a cestode in the little auk Alle alle on Franz Josef Land

One of 14 dovekeys (Alle alle) investigated in August 1993 on Hooker Island (Franz Joseph Land) was found to harbour a single specimen of an early immature dilepidid cestode attributed as Alcataenia sp. The rostellum bears a double crown of 22 hooks, 0.025-0.027 mm in length. Except Threlfall (1971) there was no information concerning dovekey's parasites.     

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.     

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.     

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.     

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.     

Potentially pathogenic yeast isolated from the throat and cloaca of an Arctic colonial seabird: the little auk (Alle alle)

Yeasts are a distinctive group of microfungi, but compared to other microorganisms, their ecological function and biodiversity are poorly known. This is especially so where polar ecosystems are concerned. With climate changes and increasing pollution levels in the Arctic, it can be anticipated that there will be an increase in the prevalence and diversity of fungi colonizing live organisms. With these changes, it is crucial to investigate and monitor species diversity and prevalence of fungi in this fragile environment. In this study, yeasts were examined from throat and cloaca of a small colonial seabird, the little auk (Alle alle), a keystone species in the Arctic ecosystem. Samples were collected from 94 adults and 17 nestlings in breeding colony in Magdalenefjorden (NW Spitsbergen) in 2009. In total, twelve species of yeast from eight genera were found in 12 % of the samples, with the Dipodascus genus being the most prevalent. All yeast species were found in the adults, but only one species, Cryptococcus macerans, was found in a single nestling. In individuals where fungus was isolated, it was only isolated from either the throat or the cloaca, except for two cases, where fungus was found in both throat and cloaca. The presence of yeast was not related to sex but age of the birds, with adults being more prone to colonization by yeasts than the nestlings. The relatively low prevalence and diversity of yeast in little auks suggest that these birds are random carriers of fungi, with minor health impacts.     

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     

Trade‐offs between reproduction and self‐maintenance (immune function and body mass) in a small seabird,the little auk

Breeding season is the most energetically and physiologically demanding phase in the avian annual cycle, challenging adults' physiology and survival. However, the timing and extent that self‐maintenance of breeding adults is compromised during the breeding season is poorly understood. We investigated the trade‐off between reproduction and self‐maintenance in relation to breeding phase (prelaying, incubation, chick rearing) and sex in a small Arctic seabird, the little auk Alle alle. To measure a bird's allocation of time for self‐maintenance, we examined size‐adjusted body mass and immunocompetence expressed by bacteria (Escherichia coli) killing capacity (BKC) of blood plasma, heterophils/lymphocyte ratio (H/L) and their numbers of particular leucocytes per 10 000 red blood cells (RBC). We found that size‐adjusted body mass decreased as the breeding season progressed. BKC, number of heterophils and H/L values were all was significantly higher at prelaying when compared to all other phases. Interestingly, we found that heavier individuals had higher BKC and number heterophils at the prelaying and chick rearing phases than light individuals. There were no differences by sex in any studied variables. Our results indicate that immunocompetence and body mass of breeding adults decreases over the course of breeding season. The efficiency of the immune system appears to be dependent on the bird's body reserves. Our results suggest that little auks allocation of resources into reproduction negatively affects their self‐maintenance.     

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.     

Body size variation of a high-Arctic seabird: the dovekie (Alle alle)

Variation in body size among subpopulations of the same species may reflect phenotypic or genetic responses to environmental gradients or geographical distance. Here, we examine geographical variation in the body size of the dovekie (Alle alle), the most numerous high-Arctic seabird. Locations of dovekie breeding sites are largely restricted to the high-Arctic zone of the Atlantic. We compared wing length, head-bill length, body mass, and a body size index of 1,076 birds from nine main colonies spanning a large part of the breeding range of the species. Results suggest morphological variation across the studied populations of dovekies, with a longitudinal increase in body size from west to east. The smallest birds breed in the western part of the population (Greenland and Jan Mayen), middle-sized individuals on Svalbard, and the largest birds (A. a. polaris subspecies) breed in the eastern part of the studied area, Franz Josef Land. Environmental (air temperature, wind speed, and sea surface temperature) and geographical (intercolonial distance) parameters were analyzed to explore potential mechanisms driving differences in body size. The body size of birds increased significantly with decreasing air temperature, but only when the two subspecies were considered. We did not find a relationship between sea surface temperature and body size of birds. Also, no close relationship was revealed between birds’ body size and the geographical distance between colonies. Whether the body size variation of dovekie can be explained by phenotypic plasticity in response to environmental conditions in wintering areas or a pattern of distance-independent gene flow between colonies remains to be explored.     

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.     

Summer at-sea distribution of little auks Alle alle and harp seals Pagophilus (Phoca) groenlandica in the Fram Strait and the Greenland Sea: impact of small-scale hydrological events

Among the most numerous seabird and pinniped species of the Fram Strait and the Greenland Sea, little auks Alle alle and harp seals Pagophila (Phoca) groenlandica are very abundant in the mixed Polar/Arctic Waters at the front between the two water masses. This must reflect the presence of very high concentrations of their food, Arctic zooplankton and nekton, massively attracting their predators. Such a high biological production seems to be depending on new primary production based on upwelling and high nutrient concentration. This usually takes place at the ice edge (e.g. July 2005), but hydrological conditions such as eddies can modify its position, east of the front in open water as caused by a subsurface eddy (August 2005), or in ice-covered areas if westerly winds push the pack ice to the east, eventually covering an eddy and causing very high concentrations of little auks and harp seals (July 2008). On the other hand, a dramatic decrease of pack ice coverage can move this water mass farther north and west, making it inaccessible to little auks during their breeding season, and apparently causing breeding failure in Jan Mayen in July 2005. In future years, if a much stronger diminution in sea ice coverage will take place, similar to the retreat in 2005 and 2007, the failure might affect the whole Spitsbergen population, as well as other seabird species feeding mainly at the ice edge.     

Variation in faecal corticosterone metabolites in an Arctic seabird,the Little Auk (Alle alle) during the nesting period

Glucocorticoids participate in the control of whole body homoeostasis and an organism’s response to stress. Corticosterone, which is the principal glucocorticoid in birds, has been shown to increase in response to different energetic demands and perturbations that individuals have to cope with. In this study, a non-invasive method to examine the corticosterone secretion by measuring faecal corticosterone metabolites (FCM) has been established for an Arctic seabird, the Little Auk (Alle alle). A group-specific immunoassay was successfully validated for adults and chicks using an adrenocorticotropic challenge test. Then, FCM levels were investigated under different energetic and physiological demands, determined by weather conditions, week of chick rearing in adults, and age in chicks. The amount of rainfall had no effect on FCM levels in adults, whereas it negatively affected FCM levels in chicks. There was no variation in FCM concentrations among weeks of chick rearing in adults. In chicks, the FCM levels increased with age. Moreover, chicks with higher FCM levels had lower body mass and fledged later than chicks with lower FCM levels. This study demonstrates that environmental stress such as poor weather conditions can trigger significant changes in corticosterone levels in seabird chicks. Furthermore, the results indicate that corticosterone may be involved in the physiological and behavioural adjustments necessary for successful fledging and post-fledging survival.     

Fit females and fat polygynous males: seasonal body mass changes in the grey-headed flying fox

When females and males differ in their timing of maximum reproductive effort, this can result in sex-specific seasonal cycles in body mass. Such cycles are undoubtedly under strong selection, particularly in bats, where they affect flying ability. Flying foxes (Old World fruit bats, Pteropus spp.) are the largest mammals that can sustain powered flight and therefore face critical trade-offs in managing body reserves for reproduction, yet little is known about body mass dynamics in this group. I investigated body mass changes in relation to reproductive behaviour in a large colony of grey-headed flying foxes (Pteropus poliocephalus). In this polygynous mammal, females were predicted to maximise reproductive effort during lactation and males during the breeding season. As predicted, female body condition declined during the nursing period, but did not vary in relation to sexual activity. By contrast, males accumulated body reserves prior to the breeding season, but subsequently lost over 20% of their body mass on territory defence and courtship, and lost foraging opportunities as they also defended their day roost territories at night. Males in better condition had larger testes, particularly during territory establishment, prior to maximum sexual activity. Thus, the seasonality of female mass reflected the high metabolic load that lactation imposes on mothers. However, male mass followed a pattern akin to the "fatted male phenomenon", which is commonly observed in large polygynous mammals with seasonal reproduction, but not in bats. This shows the importance of body reserves for reproduction in flying foxes, despite their severe constraints on body mass.     

Geographic patterns in seasonal changes of body mass,skull, and brain size of common shrews

1. Some small mammals exhibit Dehnel''s Phenomenon, a drastic decrease in body mass, braincase, and brain size from summer to winter, followed by a regrowth in spring. This is accompanied by a re‐organization of the brain and changes in other organs. The evolutionary link between these changes and seasonality remains unclear, although the intensity of change varies between locations as the phenomenon is thought to lead to energy savings during winter.
2. Here we explored geographic variation of the intensity of Dehnel''s Phenomenon in Sorex araneus. We compiled literature on seasonal changes in braincase size, brain, and body mass, supplemented by our own data from Poland, Germany, and Czech Republic.
3. We analyzed the effect of geographic and climate variables on the intensity of change and patterns of brain re‐organization.
4. From summer to winter, the braincase height decreased by 13%, followed by 10% regrowth in spring. For body mass, the changes were −21%/+82%, respectively. Changes increased toward northeast. Several climate variables were correlated with these transformations, confirming a link of the intensity of the changes with environmental conditions. This relationship differed for the decrease versus regrowth, suggesting that they may have evolved under different selective pressures.
5. We found no geographic trends explaining variability in the brain mass changes although they were similar (−21%/+10%) to those of the braincase size. Underlying patterns of change in brain organization in northeastern Poland were almost identical to the pattern observed in southern Germany. This indicates that local habitat characteristics may play a more important role in determining brain structure than broad scale geographic conditions.
6. We discuss the techniques and criteria used for studying this phenomenon, as well as its potential presence in other taxa and the importance of distinguishing it from other kinds of seasonal variation.

     

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.     

On the relationship between sweet taste and seasonal body weight changes in a primate (Microcebus murinus)

The relationship between obesity and taste, especially sweettaste, has been and is of interest. From this point of viewof a small primate, the lesser mouse lemur (Microcebus murinus),is of particular interest. It goes through a yearly cycle ofphysiological changes, one of which is an extreme variationin body weight of up to 100%. This occurs concomitantly withsignificant changes of the animal's liking for sucrose; measuredby two-bottle preference tests, the threshold for sucrose changesfrom 28–45 in lean to 77–105 mM in obese animals.It is possible that a change in peripheral taste sensitivitymight be the cause for these preference changes. To test thispossibility we studied the ability of M.murinus to taste sucrosewith electrophysiological and conditioned taste aversion techniques.The electrophysiological recordings were obtained from the chordatympani proper nerve in two heavy and three lean animals. Wedid not record any difference between the two groups in theirneural response to a series of sucrose concentrations. Conditionedtaste aversion experiments with 200 mM sucrose as conditioningstimulus and 50 and 200 mM sucrose as test stimuli gave similarresults. No difference was found between three heavy and fourlean animals; both groups rejected the sucrose concentrations.The results support the notion that the seasonal variationsin preference threshold to sucrose were unrelated to the abilityof M.murinus to taste sucrose.