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.

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.

Contrasting Spatial and Seasonal Trends of Methylmercury Exposure Pathways of Arctic Seabirds: Combination of Large-Scale Tracking and Stable Isotopic Approaches.

Despite the limited direct anthropogenic mercury (Hg) inputs in the circumpolar Arctic, elevated concentrations of methylmercury (MeHg) are accumulated in Arctic marine biota. However, the MeHg production and bioaccumulation pathways in these ecosystems have not been completely unraveled. We measured Hg concentrations and stable isotope ratios of Hg, carbon, and nitrogen in the feathers and blood of geolocator-tracked little auk Alle alle from five Arctic breeding colonies. The wide-range spatial mobility and tissue-specific Hg integration times of this planktivorous seabird allowed the exploration of their spatial (wintering quarters/breeding grounds) and seasonal (nonbreeding/breeding periods) MeHg exposures. An east-to-west increase of head feather Hg concentrations (1.74-3.48 µg·g-1) was accompanied by significant spatial trends of Hg isotope (particularly Δ199Hg: 0.96-1.13‰) and carbon isotope (δ13C: -20.6 to -19.4‰) ratios. These trends suggest a distinct mixing/proportion of MeHg sources between western North Atlantic and eastern Arctic regions. Higher Δ199Hg values (+0.4‰) in northern colonies indicate an accumulation of more photochemically impacted MeHg, supporting shallow MeHg production and bioaccumulation in high Arctic waters. The combination of seabird tissue isotopic analysis and spatial tracking helps in tracing the MeHg sources at various spatio-temporal scales.

Sex- and breeding stage-specific hormonal stress response of seabird parents.

Changes in corticosterone (CORT) and prolactin (PRL) levels are thought to provide complementary information on parental decisions in birds in the context of stressful situations. However, these endocrine mechanisms have yet to be fully elucidated, appearing to vary among avian species without any clear pattern. Here, we examined CORT and PRL stress responses in a small Arctic seabird, the little auk (Alle alle). We analysed the levels of these hormones (baseline, and stress response, i.e. the change in the baseline in response to stress) with respect to the breeding phase (mid incubation and mid chick rearing) and the sex of the birds. Baseline CORT concentrations were similar during both breeding phases but baseline PRL levels were higher during incubation than chick rearing. The CORT and PRL stress responses were stronger during incubation than chick rearing (although with respect to CORT the effect was only marginally significant). There were also some sex-specific baseline levels and stress responses for both hormones (during the incubation period males compared to females exhibited higher CORT stress response and lower baseline PRL; during the chick rearing period males exhibited higher PRL stress response). Our results suggest that in the case of the little auk, both the incubation and the chick rearing periods may represent similar levels of physiological stress. However, the birds may be more sensitive to stress during incubation than during chick rearing, possibly because of inter-phase differences in predation pressure. The sex differences suggest differential exposure of males and females to stressors.

Differential timing and latitudinal variation in sex ratio of Aquatic Warblers during the autumn migration.

Differential migration has been extensively reported in spring, but less so in autumn, particularly in relation to sex in monomorphic bird species. Here, we analysed the autumn passage of a monomorphic, globally threatened passerine, the Aquatic Warbler Acrocephalus paludicola throughout Western Europe, with regard to age and sex. We showed that, overall, adults migrated earlier than first-year birds, and males migrated earlier than females during the autumn migration. This may be caused by an overall social dominance of adults over immatures, and differentiated migration strategy of males and females. In addition, we found male-skewed sex proportions, with a tendency to an equalised ratio in more southern stopover sites. This may indicate a male bias in the global population or different migration strategies of the sexes. Differential migration may cause the age and sex classes to be exposed differently to various threats affecting demographic structure of the species.

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.

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.

Coordination of parental performance is breeding phase-dependent in the Dovekie (Alle alle), a pelagic Arctic seabird.

Currently, parental care is becoming increasingly perceived as male and female cooperation, instead of being primarily shaped by sexual conflict. Most studies examining cooperating performance consider coordination of parental activities, and doing so focuses on a short time-window including only one stage of breeding (i.e., incubation or chick rearing period). Here, we considered the cooperation of breeding partners, investigating the coordination of parental care in a long-lived seabird species with long and extensive biparental care, the Dovekie (or Little Auk), Alle alle, and looked at the issue throughout the breeding season. Previous studies on this species revealed coordinated chick provisioning, but parental coordination during incubation remains unstudied. Using video recordings collected over the course of two breeding seasons, we tested whether coordination was subject to small-scale changes within each stage and whether there was a relationship between coordination levels across the two stages. We found that the level of parental coordination is overall high and increases during the incubation period but decreases through the chick rearing phases. There were some inter-annual differences in the coordination level both at the incubation and chick rearing stages. We also found some dependency between the coordination during the incubation and chick rearing periods. All these results suggest that coordination is not a fixed behavior but breeding-phase dependent. The present study thus provides insights into how parental care and parents' cooperation is shaped by brood needs and conditions. It also highlights a relationship between coordination levels during chick rearing and incubation periods, suggesting some extent of temporal dependence in coordination of parental performance within the breeding season.

Decline in the West Greenland population of a zooplanktivorous seabird, the little auk Alle alle.

The warming of the Arctic had lead to a diverse range of impacts on local biota, including northward shifts of some species range. Here, we report past and present distribution and abundance of an Arctic zooplanktivorous seabird, the little auk Alle alle in West Greenland south of 74° N, and examine the changes in sea surface temperature (SST) and sea ice concentration (SIC) in the birds foraging areas in 1850-2007. We estimated the little auk population in the studied region to be 5,200 pairs in the 1930s, 6,000-6,500 pairs in the 1940-1970s and 70-80 pairs by the 2000s. We found that periods with increased SST and reduced SIC, especially in the last few decades, coincided with little auk population declines. Besides, years with little auk presence in breeding sites were characterized by either low SST and low to moderate SIC or higher SST but moderate to high SIC. Observed contraction of the breeding range and a decrease in abundance of the little auk may be attributed to more complex climate-driven changes in the marine ecosystem at finer spatial and temporal scales and/or cannot be easily detected given the coarseness of data used. It is possible that the population in this region has never been very numerous being subjected to local impacts such as disease, bycatch, predation, etc. The climate warming that is currently being observed, along with corresponding shifts in zooplankton communities, may lead to extirpation of the studied little auk populations.

Diversity of major histocompatibility complex of II B gene and mate choice in a monogamous and long-lived seabird, the Little Auk (Alle alle).

The major histocompatibility complex (MHC) plays a key role in the adaptive immune system of vertebrates, and is known to influence mate choice in many species. In birds, the MHC has been extensively examined but mainly in galliforms and passerines while other taxa that represent specific ecological and evolutionary life-histories, like seabirds, are underexamined. Here, we characterized diversity of MHC Class II B exon 2 in a colonial pelagic seabird, the Little Auk (or Dovekie Alle alle). We further examined whether MHC variation could be maintained through balancing selection and disassortative mating. We found high polymorphism at the genotyped MHC fragment, characterizing 99 distinct alleles across 140 individuals from three populations. The alleles frequencies exhibited a similar skewed distribution in both sexes, with the four most commonly occurring alleles representing approximately 35% of allelic variation. The results of a Bayesian site-by-site selection analysis suggest evidence of balancing selection and no direct evidence for MHC-dependent disassortative mating preferences in the Little Auk. The latter result might be attributed to the high overall polymorphism of the examined fragment, which itself may be maintained by the large population size of the species.

Behavioural and hormonal stress responses during chick rearing do not predict brood desertion by female in a small Arctic seabird.

We examined behavioural and hormonal stress responses in a small seabird (little auk, Alle alle), which exhibits a transition from biparental to male-only care towards the end of the nesting period, in order to understand the mechanisms underlying this parental strategy. We hypothesized that the male staying with the chick should be less sensitive to stressors. As such the male might offer the offspring more efficient protection during the fledging period than the female. We tested this hypothesis by observing male and female behaviour in a neophobia test. We also measured the birds' baseline and stress-induced levels of corticosterone and prolactin using the standardized capture-and-restraint protocol. Both sexes respond rapidly to foreign objects, delaying the entry time to the nest with food, consuming the food load, and/or temporarily abandoning feeding. However, we did not find any differences between the sexes in the frequency of each behaviour or in the time of the first reaction to the experimental treatment. Level of both corticosterone and prolactin increased after the experimental treatment. However, we did not find sex differences in baseline and stress-induced hormone levels. The results indicate that the males are as much sensitive to the stress situation as the females. Thus, the pattern of male and female behavioural and hormonal responses to stress does not predict their behaviour at the final breeding stage.

Differential autumn migration of the aquatic warbler Acrocephalus paludicola.

Relatively little attention has been paid to sex differences in the migration of birds in autumn. We studied the autumn migration strategy of molecularly sexed males and females in the globally threatened aquatic warbler Acrocephalus paludicola. We captured 176 birds at a stopover site in the Loire estuary at Donges, France. The median date for the passage of adults was 8 days earlier in males than females, although the timing of migration in first-year males and females was similar. This indicates that males, who are without parental duties, can start their migration earlier than females and first-year birds. Adults were significantly heavier than immature birds but did not have higher fat scores. In both age categories, more males (two to three times more) were captured. However, various factors (including tape-luring) can affect observed sex ratio.

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.

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.

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.

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.

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.

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.

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.

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.