The first evidence of alloparental feeding in a crevice-nesting seabird, the little auk.

An alloparent is an individual that cares for a young individual, but it is not its genetic parent. This behaviour is known in many species of animals, but some groups are still underreported. Here, we documented, in camera footage, the alloparental feeding of two chicks of the little auk, a crevice-nesting seabird. This is the first evidence of this behaviour in the little auk despite similar monitoring undertaken between 2016 and 2022 and the second record for a crevice/burrow-nesting seabird. We compared chicks that were fed by an alloparent to other chicks from the same year and explored reasons for the behaviour in the context of seabird breeding biology.

Sex differences in foraging ecology of a zooplanktivorous little auk Alle alle during the pre-laying period: insights from remote sensing and animal-tracking.

BACKGROUND: Energy and time allocation in seabirds differ between consecutive stages of breeding given various requirements of particular phases of the reproductive period. Theses allocations may also be sex-specific considering differential energetic or nutritional requirements of males and females and/or sexual segregation in foraging niches and/or areas. In this study we investigated the foraging ecology of an Arctic, zooplanktivorous seabird, the little auk Alle alle during the pre-laying period using remote sensing of the environment and GPS-TDR loggers deployed on birds. We compared foraging trips range and habitats of birds with other stages of the breeding period and between sexes. RESULTS: We found that little auks during the pre-laying period foraged exclusively in cold sea surface temperature zones (with temperatures < 5 ºC) but in various sea depth zones. They dived to similar depths ranging from -4.0 to -10.9 m, exploring various thermal microhabitats (with mean temperatures values ranging from 2.2 °C in Shelf sea depth zone to 5.9 °C in Deep sea depth zone). The majority of foraging trips and dives characteristics were similar to subsequent phases of breeding. However, home ranges during the pre-laying trips were wider compared to the incubation period. As expected, females exhibited wider foraging niches compared to males (wider range of sea surface temperature and sea depth in foraging locations), which could be explained by sex specific energetic and/or nutritional requirements (females producing an egg). We also delineated local foraging areas important for little auks during their whole breeding season. Protection of these areas is crucial for sustaining the local marine biodiversity. CONCLUSIONS: We found that little auks females during the pre-laying period explored wider foraging niches compared to males. These differences may be attributed to sex-specific nutritional or/and energetical constraints at this stage of breeding. The results of this study also emphasize the importance of shelf Arctic-type water masses as the foraging areas for little auks during successive stages of breeding.

Calls of the little auk (Alle alle) chicks reflect their behavioural contexts.

Animal vocalisations can often inform conspecifics about the behavioural context of production and the underlying affective states, hence revealing whether a situation should be approached or avoided. While this is particularly important for socially complex species, little is known about affective expression in wild colonial animals, and even less to about their young. We studied vocalisations of the little auk (Alle alle) chicks in the Hornsund breeding colony, Svalbard. Little auks are highly colonial seabirds, and adults convey complex behavioural contexts through their calls. We recorded chick calls during two contexts of opposite affective valence: handing by a human, and while they interact with their parents inside the nest. Using permuted discriminant function analysis and a series of linear mixed models, we examined the effect of the production context/associated affective valence on the acoustic parameters of those calls. Calls were reliably classified to their context, with over 97% accuracy. Calls uttered during handling had higher mean entropy, fundamental frequency, as well as lower spectral centre of gravity and a less steep spectral slope compared to calls produced during interactions with a parent inside the nest. The individuality of handling calls, assessed by information content, was lower than the individuality of calls uttered in the nest. These findings suggest that seabird chicks can effectively communicate behavioural/affective contexts through calls, conveying socially important messages early in development. Our results are mostly in line with emotional expression patterns observed across taxa, supporting their evolutionary continuity.

Social calls of the little auk (Alle alle) reflect body size and possibly partnership, but not sex.

Source-filter theory posits that an individual's size and vocal tract length are reflected in the parameters of their calls. In species that mate assortatively, this could result in vocal similarity. In the context of mate selection, this would mean that animals could listen in to find a partner that sounds-and therefore is-similar to them. We investigated the social calls of the little auk (Alle alle), a highly vocal seabird mating assortatively, using vocalizations produced inside 15 nests by known individuals. Source- and filter-related acoustic parameters were used in linear mixed models testing the possible impact of body size. A principal component analysis followed by a permuted discriminant function analysis tested the effect of sex. Additionally, randomization procedures tested whether partners are more vocally similar than random birds. There was a significant effect of size on the mean fundamental frequency of a simple call, but not on parameters of a multisyllable call with apparent formants. Neither sex nor partnership influenced the calls-there was, however, a tendency to match certain parameters between partners. This indicates that vocal cues are at best weak indicators of size, and other factors likely play a role in mate selection.

Mine or my neighbours' offspring: an experimental study on parental discrimination of offspring in a colonial seabird, the little auk Alle alle.

Individual recognition (IR) abilities may result from various ecological and naturally selected features of a species. Complex IR mechanisms should develop when the risk of misidentification of a chick is high. For colonial seabirds, the ability to identify their own brood is crucial to ensure parental fitness. Vocalizations seem to be a key component of most parent-offspring interactions, although few studies have assessed the interindividual differences in seabird chick calls and their potential usage in IR. The little auk (Alle alle), which breeds in dense colonies, constitutes a perfect model for testing IR. In this study, we (1) examined chick calls at different stages of the nesting period, and (2) cross-fostered chicks to examine the rate of acceptance/nonacceptance of chicks by parents. We found significant interindividual differences in chick begging and fledging calls. Surprisingly, all cross-fostered chicks in our experiments were accepted by their foster parents, and male parents were as equally likely to accept cross-fostered chicks as females, even though the sexes would be expected to differ in offspring recognition due to different postfledging interactions with the chick. The revealed individuality of chick calls suggests the potential for chick vocal recognition in the studied species, but parent birds may disregard the individual characteristics enabling chick discrimination. This may take place as long as the chick is found in the nest because of the high likelihood that the chick present there is the focal one. However, IR during and after fledging requires further study. Studying the complexity of IR mechanisms is important for better understanding various avian social relationships and interactions.

Molecular tools prove little auks from Svalbard are extremely selective for Calanus glacialis even when exposed to Atlantification.

Two Calanus species, C. glacialis and C. finmarchicus, due to different life strategies and environmental preferences act as an ecological indicators of Arctic Atlantification. Their high lipid content makes them important food source for higher trophic levels of Arctic ecosystems including the most abundant Northern Hemisphere's seabird, the little auk (Alle alle). Recent studies indicate a critical need for the use of molecular methods to reliably identify these two sympatric Calanus species. We performed genetic and morphology-based identification of 2600 Calanus individuals collected in little auks foraging grounds and diet in summer seasons 2019-2021 in regions of Svalbard with varying levels of Atlantification. Genetic identification proved that 40% of Calanus individuals were wrongly classified as C. finmarchicus according to morphology-based identification in both types of samples. The diet of little auks consisted almost entirely of C. glacialis even in more Atlantified regions. Due to the substantial bias in morphology-based identification, we expect that the scale of the northern expansion of boreal C. finmarchicus may have been largely overestimated and that higher costs for birds exposed to Atlantification could be mostly driven by a decrease in the size of C. glacialis rather than by shift from C. glacialis to C. finmarchicus.

Adoption and feeding of fieldfare nestlings and fledglings by European blackbird.

Interspecific adoption is an intriguing topic in behavioral and evolutionary ecology. As it is a rare phenomenon, seldom documented in the literature, reports of interspecific adoption based on solid data are particularly valuable. A long-term and extensive monitoring programme involving a local population of European blackbirds (Turdus merula, hereafter blackbird) has yielded, among other things, observations of alloparental behavior exhibited by blackbirds toward fieldfare (Turdus pilaris) nestlings (a single nest, the first-ever record) and fledglings (12 cases in all). We discuss the observations in the context of the available literature.

Foraging, Fear and Behavioral Variation in a Traplining Hummingbird.

Traditionally, foraging behavior has been explained as the response to a trade-off between energetic gain from feeding resources and potential costs from concomitant risks. However, an increasing number of studies has shown that this view fails to explain an important fraction of the variation in foraging across a variety of taxa. One potential mechanism that may account for this variation is that various behavioral traits associated with foraging may have different fitness consequences, which may depend on the environmental context. Here, we explored this mechanism by evaluating the foraging efficiency of long-billed hermit hummingbirds (Phaethornis longirostris) with regard to three behavioral traits: (a) exploration (number of feeders used during the foraging visit), (b) risk avoidance (latency to start feeding) and (c) arousal (amount of movements during the foraging visit) in conditions at two different levels of perceived risk (low-control and high-experimental, with a threatening bullet ant model). Foraging efficiency decreased in response to threatening conditions. However, behavioral traits explained additional variation in foraging efficiency in a condition-dependent manner. More exploration was associated with a higher foraging efficiency under control conditions, but this was reversed when exposed to a threat. Regardless of the conditions, arousal was positively associated with foraging efficiency, while risk avoidance was negatively related. Importantly, exploratory behavior and risk avoidance were quite repeatable behaviors, suggesting that they may be related to the intrinsic traits of individuals. Our findings highlight the importance of taking into account additional behavioral dimensions to better understand the foraging strategies of individuals.

Seabird's cry: repertoire and vocal expression of contextual valence in the little auk (Alle alle).

Many seabird species breed in colonies counting up to hundreds of thousands of individuals. Life in such crowded colonies might require special coding-decoding systems to reliably convey information through acoustic cues. This can include, for example, developing complex vocal repertoires and adjusting the properties of their vocal signals to communicate behavioural contexts, and thus regulate social interactions with their conspecifics. We studied vocalisations produced by the little auk (Alle alle)-a highly vocal, colonial seabird-over mating and incubation periods on the SW coast of Svalbard. Using passive acoustic recordings registered in a breeding colony, we extracted eight vocalisation types: single call, clucking, classic call, low trill, short call, short-trill, terror, and handling vocalisation. Calls were grouped by production context (based on the typically associated behaviour), to which a valence (positive vs negative) was later attributed, when possible, according to fitness threats, i.e. predator or human presence (negative) and promoters, i.e. interaction with a partner (positive). The effect of the putative valence on eight selected frequency and duration variables was then investigated. The putative contextual valence significantly affected the acoustic properties of the calls. Calls assigned positive valence had higher fundamental frequency and spectral centre of gravity as well as shorter sound duration than these assigned negative valence. These results indicate that the little auk's vocal communication system may facilitate expression of complex behavioural contexts, and seems to include vocal plasticity within vocalisation types-however, more data are necessary to better understand this effect and possible interplays of other factors.

A keystone avian predator faces elevated energy expenditure in a warming Arctic.

Climate change is transforming bioenergetic landscapes, challenging behavioral and physiological coping mechanisms. A critical question involves whether animals can adjust behavioral patterns and energy expenditure to stabilize fitness given reconfiguration of resource bases, or whether limits to plasticity ultimately compromise energy balance. In the Arctic, rapidly warming temperatures are transforming food webs, making Arctic organisms strong models for understanding biological implications of climate change-related environmental variability. We examined plasticity in the daily energy expenditure (DEE) of an Arctic seabird, the little auk (Alle alle) in response to variability in climate change-sensitive drivers of resource availability, sea surface temperature (SST) and sea ice coverage (SIC), and tested the hypothesis that energetic ceilings and exposure to mercury, an important neurotoxin and endocrine disrupter in marine ecosystems, may limit scope for plasticity. To estimate DEE, we used accelerometer data obtained across years from two colonies exposed to distinct environmental conditions (Ukaleqarteq [UK], East Greenland; Hornsund [HS], Svalbard). We proceeded to model future changes in SST to predict energetic impacts. At UK, high flight costs linked to low SIC and high SST drove DEE from below to above 4 × basal metabolic rate (BMR), a proposed energetic threshold for breeding birds. However, DEE remained below 7 × BMR, an alternative threshold, and did not plateau. Birds at HS experienced higher, relatively invariable SST, and operated above 4 × BMR. Mercury exposure was unrelated to DEE, and fitness remained stable. Thus, plasticity in DEE currently buffers fitness, providing resiliency against climate change. Nevertheless, modeling suggests that continued warming of SST may promote accelerating increases in DEE, which may become unsustainable.

Use of geolocators for investigating breeding ecology of a rock crevice-nesting seabird: Method validation and impact assessment.

Investigating ecology of marine animals imposes a continuous challenge due to their temporal and/or spatial unavailability. Light-based geolocators (GLS) are animal-borne devices that provide relatively cheap and efficient method to track seabird movement and are commonly used to study migration. Here, we explore the potential of GLS data to establish individual behavior during the breeding period in a rock crevice-nesting seabird, the Little Auk, Alle alle. By deploying GLS on 12 breeding pairs, we developed a methodological workflow to extract birds' behavior from GLS data (nest attendance, colony attendance, and foraging activity), and validated its accuracy using behavior extracted from a well-established method based on video recordings. We also compared breeding outcome, as well as behavioral patterns of logged individuals with a control group treated similarly in all aspects except for the deployment of a logger, to assess short-term logger effects on fitness and behavior. We found a high accuracy of GLS-established behavioral patterns, especially during the incubation and early chick rearing period (when birds spend relatively long time in the nest). We observed no apparent effect of logger deployment on breeding outcome of logged pairs, but recorded some behavioral changes in logged individuals (longer incubation bouts and shorter foraging trips). Our study provides a useful framework for establishing behavioral patterns (nest attendance and foraging) of a crevice-nesting seabird from GLS data (light and conductivity), especially during incubation and early chick rearing period. Given that GLS deployment does not seem to affect the breeding outcome of logged individuals but does affect fine-scale behavior, our framework is likely to be applicable to a variety of crevice/burrow nesting seabirds, even though precautions should be taken to reduce deployment effect. Finally, because each species may have its own behavioral and ecological specificity, we recommend performing a pilot study before implementing the method in a new study system.

Mercury Contamination Challenges the Behavioral Response of a Keystone Species to Arctic Climate Change.

Combined effects of multiple, climate change-associated stressors are of mounting concern, especially in Arctic ecosystems. Elevated mercury (Hg) exposure in Arctic animals could affect behavioral responses to changes in foraging landscapes caused by climate change, generating interactive effects on behavior and population resilience. We investigated this hypothesis in little auks (Alle alle), a keystone Arctic seabird. We compiled behavioral data for 44 birds across 5 years using accelerometers while also quantifying blood Hg and environmental conditions. Warm sea surface temperature (SST) and low sea ice coverage reshaped time activity budgets (TABs) and diving patterns, causing decreased resting, increased flight, and longer dives. Mercury contamination was not associated with TABs. However, highly contaminated birds lengthened interdive breaks when making long dives, suggesting Hg-induced physiological limitations. As dive durations increased with warm SST, subtle toxicological effects threaten to increasingly constrain diving and foraging efficiency as climate change progresses, with ecosystem-wide repercussions.

Gone with the wind - Wind speed affects prey accessibility for a High Arctic zooplanktivorous seabird, the little auk Alle alle.

Foraging ecology of chick rearing seabirds is affected mainly by the food availability on feeding grounds, but it can be also modulated by environmental conditions during the foraging trip, in that wind force. Considering predicted strengthening of surface winds over the Arctic Ocean, this factor may have a growing impact on the foraging performance of Arctic seabirds. Here, we studied how wind speed could affect prey accessibility for the High Arctic zooplanktivorous seabird, the little auk Alle alle breeding in Svalbard in 2015-2019. First, we estimated availability of its preferred prey, a cold water copepod Calanus glacialis, based on wider-scale mesozooplankton biomass model and environmental conditions. Then we estimated prey accessibility by including wind speed, the factor affecting the flapping flight performance of little auks commuting from/to the colony. Finally, we compared reproductive performance of the little auks (chick diet, growth rate and survival and duration of foraging flights of adults) between the studied years differing in wind and food availability conditions. We found that wind speed could affect significantly food accessibility for a zooplanktivorous seabird. Despite high spatial and temporal variability in prey availability and accessibility in shelf waters of SW Spitsbergen, interannual differences in duration of foraging flights and chick growth rate, little auks were able to sustain high breeding success confirming their capacity to buffer suboptimal foraging conditions. Our multidisciplinary work, combining multi-year remote sensing of oceanographic conditions, zooplankton availability and accessibility modelling, little auks diet composition and chick growth and survival emphasizes the importance of including wind conditions in the studies of foraging ecology of seabirds.

Post-foraging in-colony behaviour of a central-place foraging seabird.

Studies on time allocation of various activities are crucial to understand which behavioural strategy is the most profitable in a given context, and so why animals behave in a particular way. Such investigations usually focus on a time window when the studied activity is performed, often neglecting how the time devoted to focal activity affects time allocation to following-up behaviours, while that may have its own fitness consequences. In this study, we examined time allocation into three post-foraging activities (entering the nest with food, nest attendance, and colony attendance) in a small seabird species, the little auk (Alle alle). Since little auks alternate foraging trips of different duration (short and long) and purpose (offspring feeding and primarily self-feeding, respectively) we expected that duration of the following up in-colony activities would also vary, being longer after a long absence in the colony (because of greater need of reassessment of the current predation pressure and social interactions in the colony, and re-establishing the bond with the offspring and/or partner and/or neighbours after longer absence). We found that it was not always the case, as time allocation of the post-foraging in-colony activities was primarily year- and sex-specific. It highlights the need to consider year and sex effects in studies of behavioural ecology, as not doing so may lead to spurious conclusions. Interestingly, and despite a great inter-individual variation in time allocation in the post-foraging in-colony activities, little auk individuals were quite repeatable in their behavioural performance, which suggests these activities may reflect birds behavioural profile. Overall, post-foraging in-colony activity of the little auk, although not much dependent on duration/type of the preceding foraging flights, varies with respect to year and sex, and as such may be a proxy of behavioural plasticity of the population.

Eye Region Surface Temperature and Corticosterone Response to Acute Stress in a High-Arctic Seabird, the Little Auk.

Measuring changes in surface body temperature (specifically in eye-region) in vertebrates using infrared thermography is increasingly applied for detection of the stress reaction. Here we investigated the relationship between the eye-region temperature (TEYE; measured with infrared thermography), the corticosterone level in blood (CORT; stress indicator in birds), and some covariates (ambient temperature, humidity, and sex/body size) in a High-Arctic seabird, the Little Auk Alle alle. The birds responded to the capture-restrain protocol (blood sampling at the moment of capturing, and after 30 min of restrain) by a significant TEYE and CORT increase. However, the strength of the TEYE and CORT response to acute stress were not correlated. It confirms the results of a recent study on other species and all together indicates that infrared thermography is a useful, non-invasive measure of hypothalamic-pituitary-adrenal (HPA) axis reactivity under acute activation, but it might not be a suitable proxy for natural variation of circulating glucocorticoid levels.

A quiet extirpation of the breeding little auk Alle alle population in Iceland in the shadow of the famous cousin extermination.

The little auk Alle alle is an Arctic seabird breeding in the North Atlantic. Its southernmost breeding population in the Low Arctic occurred in Iceland but started to decline in numbers at the beginning of the 20th century ending in extirpation at the end of the same century. Climate warming has been blamed for the disappearance of this population. However, it was also exploited by humans (mainly for eggs). Thus, it is unclear what was the main driver for the population collapse. In this study, we reconstruct population dynamics in relation to changes in environmental conditions, and perform stochastic modelling of population viability considering various scenarios including presence/absence of climate warming and/or egg harvest. We found that extirpation of the studied population was attributed to synergistic effects of both climate warming and human harvest. The simulations revealed that climate warming without harvest would cause a 71% decline in population size but not lead to extirpation. Models with climate warming and egg harvesting resulted in population estimates close to the real data (decrease from 400 individuals in 1903 to 2 in 1996). This is one of the few studies documenting synergistic effect of climate warming and human exploitation on extirpation. A strong harvesting component in the explanation for the decline of the study population emphasizes the continuing need to control commercial harvest of animal species in the face of other pressures such as climate warming.

Differences in a Cage Escape Behaviour between Two Migrating Warblers of Different Stop-Over Strategy.

Cognitive abilities play an important role for migratory birds that are briefly visiting a variety of unfamiliar stop-over habitats. Here, we compared cognitive abilities-linked behaviour (escape from an experimental cage) between two long-distant migrants differing in stop-over ecology, Sedge Warbler (Acrocephalus schoenobaenus; not territorial, searching for locally superabundant food) and Reed Warbler (A. scirpaceus; territorial, foraging on a common prey) during the autumn migration. After two minutes of acclimatization in the cage, we remotely opened the cage door and recorded the bird's reaction. We measured latency that individuals needed to escape from a cage. Sedge warblers were 1.61 times more likely to escape from the cage than Reed Warblers. Sedge warblers generally escaped earlier after the door was opened and were 1.79 times more likely to escape at any given time than Reed Warblers. We interpret the prevalence of non-escaped individuals as a general feature of migratory birds. In contrast to resident species, they are more likely to enter an unfamiliar environment, but they are less explorative. We attributed inter-species differences in escape latency to species-specific autumn stop-over refuelling strategies in the context of specialist-generalist foraging. Our study provides ecological insight into the cognitive abilities-linked behaviour of wild animals.

Birds of a feather moult together: Differences in moulting distribution of four species of storm-petrels.

The non-breeding period of pelagic seabirds, and particularly the moulting stage, is an important, but understudied part of their annual cycle as they are hardly accessible outside of the breeding period. Knowledge about the moulting ecology of seabirds is important to understand the challenges they face outside and within the breeding season. Here, we combined stable carbon (δ13C) and oxygen (δ18O) signatures of rectrices grown during the non-breeding period of two pairs of storm-petrel species breeding in the northern (European storm-petrel, Hydrobates pelagicus, ESP; Leach's storm-petrel, Hydrobates leucorhous, LSP) and southern (black-bellied storm-petrel, Fregetta tropica, BBSP; Wilson's storm-petrel, Oceanites oceanicus, WSP) hemispheres to determine differences in moulting ranges within and between species. To understand clustering patterns in δ13C and δ18O moulting signatures, we examined various variables: species, sexes, years, morphologies (feather growth rate, body mass, tarsus length, wing length) and δ15N. We found that different factors could explain the differences within and between the four species. We additionally employed a geographical distribution prediction model based on oceanic δ13C and δ18O isoscapes, combined with chlorophyll-a concentrations and observational data to predict potential moulting areas of the sampled feather type. The northern species were predicted to moult in temperate and tropical Atlantic zones. BBSP was predicted to moult on the southern hemisphere north of the Southern Ocean, while WSP was predicted to moult further North, including in the Arctic and northern Pacific. While moulting distribution can only be estimated on large geographical scales using δ13C and δ18O, validating predictive outcomes with food availability proxies and observational data may provide valuable insights into important moulting grounds. Establishing those, in turn, is important for conservation management of elusive pelagic seabirds.

Seasonal variation of mercury contamination in Arctic seabirds: A pan-Arctic assessment.

Mercury (Hg) is a natural trace element found in high concentrations in top predators, including Arctic seabirds. Most current knowledge about Hg concentrations in Arctic seabirds relates to exposure during the summer breeding period when researchers can easily access seabirds at colonies. However, the few studies focused on winter have shown higher Hg concentrations during the non-breeding period than breeding period in several tissues. Hence, improving knowledge about Hg exposure during the non-breeding period is crucial to understanding the threats and risks encountered by these species year-round. We used feathers of nine migratory alcid species occurring at high latitudes to study bird Hg exposure during both the breeding and non-breeding periods. Overall, Hg concentrations during the non-breeding period were ~3 times higher than during the breeding period. In addition, spatial differences were apparent within and between the Atlantic and Pacific regions. While Hg concentrations during the non-breeding period were ~9 times and ~3 times higher than during the breeding period for the West and East Atlantic respectively, Hg concentrations in the Pacific during the non-breeding period were only ~1.7 times higher than during the breeding period. In addition, individual Hg concentrations during the non-breeding period for most of the seabird colonies were above 5 µg g-1 dry weight (dw), which is considered to be the threshold at which deleterious effects are observed, suggesting that some breeding populations might be vulnerable to non-breeding Hg exposure. Since wintering area locations, and migration routes may influence seasonal Hg concentrations, it is crucial to improve our knowledge about spatial ecotoxicology to fully understand the risks associated with Hg contamination in Arctic seabirds.