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Climate warming can reduce food resources for animal populations. In species exhibiting parental care, parental effort is a 'barometer' of changes in environmental conditions. A key issue is the extent to which variation in parental effort can buffer demographic rates against environmental change. Seabirds breed in large, dense colonies and globally are major predators of small fish that are often sensitive to ocean warming. We explored the causes and consequences of annual variation in parental effort as indicated by standardised checks of the proportions of chicks attended by both, one or neither parent, in a population of common guillemots Uria aalge over four decades during which there was marked variation in marine climate and chick diet. We predicted that, for parental effort to be an effective buffer, there would be a link between environmental conditions and parental effort, but not between parental effort and demographic rates. Environmental conditions influenced multiple aspects of the prey delivered by parents to their chicks with prey species, length and energy density all influenced by spring sea surface temperature (sSST) in the current and/or previous year. Overall, the mean annual daily energy intake of chicks declined significantly when sSST in the current year was higher. In accordance with our first prediction, we found that parental effort increased with sSST in the current and previous year. However, the increase was insufficient to maintain chick daily energy intake. In contrast to our second prediction, we found that increased parental effort had major demographic consequences such that growth rate and fledging success of chicks, and body mass and overwinter survival of breeding adults all decreased significantly. Common guillemot parents were unable to compensate effectively for temperature-mediated variation in feeding conditions through behavioural flexibility, resulting in immediate consequences for breeding population size because of lower adult survival and potentially longer-term impacts on recruitment because of lower productivity. These findings highlight that a critical issue for species' responses to future climate change will be the extent to which behavioural buffering can offer resilience to deteriorating environmental conditions.

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Density-dependent regulation can offer resilience to wild populations experiencing fluctuations in environmental conditions because, at lower population sizes, the average quality of habitats or resources is predicted to increase. Site-dependent regulation is a mechanism whereby individuals breed at the highest quality, most successful, sites, leaving poorer quality, less successful sites vacant. As population size increases, higher quality sites become limiting but when populations decline, lower quality sites are vacated first, offering resilience. This process is known as the 'buffer effect'. However, few studies have tested whether such regulation operates in populations experiencing changes in size and trend. We used data from a population of common guillemots Uria aalge, a colonially breeding seabird, to investigate the relationship between site occupancy probability, site quality and population size and trend. These data were collected at five sub-colonies spanning a 38-year period (1981-2018) comprising phases of population increase, decrease and recovery. We first tested whether site quality and population size in sub-colonies explained which sites were occupied for breeding, and if this was robust to changes in sub-colony trend. We then investigated whether disproportionate use of higher quality sites drove average site quality and breeding success across sub-colony sizes and trends. Finally, we tested whether individuals consistently occupied higher quality sites during periods of decline and recovery. Higher quality sites were disproportionality used when sub-colony size was smaller, resulting in higher average site quality and breeding success at lower population sizes. This relationship was unaffected by changes in sub-colony trend. However, contrary to the predictions of the buffer effect, new sites were established at a similar rate to historically occupied sites during sub-colony decline and recovery despite being of lower quality. Our results provide support for the buffer effect conferring resilience to populations, such that average breeding success was consistently higher at lower population size during all phases of population change. However, this process was tempered by the continued establishment of new, lower quality, sites which could act to slow population recovery after periods when colony size was low.

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The predictable oscillation between the light of day and the dark of night across the diel cycle is a powerful selective force that has resulted in anticipatory mechanisms in nearly all taxa. At polar latitude, however, this oscillation becomes highly attenuated during the continuous light of polar day during summer. A general understanding of how animals keep time under these conditions is poorly understood. We tested the hypothesis that the common murre (a seabird, Uria aalge) can use melatonin and corticosterone, hormones associated with timekeeping, to track the diel cycle despite continuous light. We also tested the assumption that common murres breeding during polar summer schedule their colony attendance by time of day and sex, as they do at subpolar latitude. In the Atlantic population, common murres have a plumage color dimorphism associated with fitness-related traits, and we investigated the relationship of this dimorphism with colony attendance, melatonin, and corticosterone. The common murres did not schedule their attendance behavior by time of day or sex, yet they had higher concentrations of melatonin and, to a more limited extent, corticosterone during "night" than "day". Melatonin also linked to behavioral state. The two color morphs tended to have different colony-attendance behavior and melatonin concentrations, lending support for balancing selection maintaining the plumage dimorphism. In common murres, melatonin can signal time of day despite continuous light, and the limited diel variation of corticosterone contributes to the mounting evidence that polar-adapted birds and mammals require little or no diel variation in circulating glucocorticoids during polar day.

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A global analysis recently showed that seabird breeding phenology (as the timing of egg-laying and hatching) does not, on average, respond to temperature changes or advance with time (Keogan et al. 2018 Nat. Clim. Change8, 313-318). This group, the most threatened of all birds, is therefore prone to spatio-temporal mismatches with their food resources. Yet, other aspects of the breeding phenology may also have a marked influence on breeding success, such as the arrival date of adults at the breeding site following winter migration. Here, we used a large tracking dataset of two congeneric seabirds breeding in 14 colonies across 18° latitudes, to show that arrival date at the colony was highly variable between colonies and species (ranging 80 days) and advanced 1.4 days/year while timing of egg-laying remained unchanged, resulting in an increasing pre-laying duration between 2009 and 2018. Thus, we demonstrate that potentially not all components of seabird breeding phenology are insensitive to changing environmental conditions.

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Very little is known about how morphology affects the motion, spatial stability and resulting viability of avian eggs. The limited existing research focuses on the uniquely pyriform egg shapes found in the Alcidae bird family. This unusual shell shape was originally thought to suppress displacement and prevent egg loss on the cliffside nesting habitat of the Uria genus. Unfortunately, these early studies never isolated or quantified the specific morphological features (elongation, asymmetry and conicality) of these pyriform eggs, which limits their applicability to other taxa and has hampered a robust proof of concept. We isolated each feature as an enumerated variable, produced model 3D printed eggs with incremental expressions of a single variable and then with all three features co-varying simultaneously. Recorded motion (egg-rolling) trials were conducted to test the individual and combined effects of each morphological characteristic on displacement over a range of inclines representative of the conditions found in natural habitats. Increasing elongation and asymmetry significantly increased displacement, whereas increased conicality decreased displacement in the single-variable egg models. In the multivariable egg models, only conicality consistently suppressed displacement, while lower levels of asymmetry significantly increased displacement. Our findings broadly support previous studies' assertions of the adaptive value of the pyriform eggs while also providing methodology and comparative data for future analyses of the interactions between nesting habitat, behavior and egg shape, beyond the confines of a handful of focal species.

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Two Guillemot carcases were dissected, each providing 12 discrete tissue samples and 3 samples of partially digested food. One hundred and five PCBs from the 209 PCBs determined by GCxGC-ToFMS were detected. The relative proportions of individual PCBs did not vary greatly within tissue types, although the PCB profile from undigested food could be distinguished. Enantiomer fractions (EFs) were determined for CB-95, CB-136 and CB-149 by GC-HRqToFMS. EFs in the partially digested food were near racemic, with high levels of enrichment for E1 CB-95 in the kidneys and liver (EF of 0.80 and 0.84 respectively). This provides some of the clearest evidence to date that fractionation takes place in the organs where metabolic biotransformation and elimination of PCBs occurs. Our findings also confirm the ability of non-lethal sampling techniques, such as collection of small (<1 g) blood samples, to provide PCB signatures that are representative of an individual organism.

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Nutritional support is a primary therapy administered to oiled animals during responses to oil spills, but data informing nutritional decision-making during events are limited. In this study, 44 common murres ( Uria aalge ) and 6 Western grebes ( Aechmophorus occidentalis ), naturally oiled by oceanic seeps off the coast of Ventura and Santa Barbara Counties, CA, USA, were assigned to 1 of 4 groups fed diets with varying levels (6.8% [no added oil], 11%, and 20%) and types (salmon, corn) of oil added to a partially purified basal diet. Birds used in the study ranged from extremely emaciated to thin body condition (62%-80% wild bird mean body mass). Acid-insoluble ash was used as an indigestible dietary marker to quantify nitrogen retention, apparent nitrogen digestibility, nitrogen-corrected apparent metabolizable energy, energy digestibility, fat retention, fat digestibility, and estimated fat excretion. Fat excretion is important in these species because once birds have been cleaned they are at risk of plumage recontamination from excreted fat during care. Lower fat diets resulted in lower fat excretion but higher nitrogen retention, higher apparent nitrogen digestibility, and higher apparent metabolizable energy. Decreases in nitrogen retention were significantly related to increases in fat excretion. Regardless of diet, energy digestibility significantly declined with declines in body mass, suggesting severity of emaciation reduced a birds' ability to extract energy from food. Energy digestibility was highest in the 11% (low) salmon oil diet; hence, this diet had the highest effective energy content despite a lower gross kcal/kg diet. Diets fed during oil spills historically have had high fat concentrations to provide maximum caloric support. Results of this study suggest that lower fat diets may be more efficacious for nutritionally depleted seabirds. This study provides valuable data to guide clinical decision making regarding nutritional support during oil spills and other mass stranding events.

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The duration of parental care in animals varies widely, from none to lifelong. Such variation is typically thought to represent a trade-off between growth and safety. Seabirds show wide variation in the age at which offspring leave the nest, making them ideal to test the idea that a trade-off between high energy gain at sea and high safety at the nest drives variation in departure age (Ydenberg's model). To directly test the model assumptions, we attached time-depth recorders to murre parents (fathers [which do all parental care at sea] and mothers; [Formula: see text] of each). Except for the initial mortality experienced by chicks departing from the colony, the mortality rate at sea was similar to the mortality rate at the colony. However, energy gained by the chick per day was ∼2.1 times as high at sea compared with at the colony because the father spent more time foraging, since he no longer needed to spend time commuting to and from the colony. Compared with the mother, the father spent ∼2.6 times as much time diving per day and dived in lower-quality foraging patches. We provide a simple model for optimal departure date based on only (1) the difference in growth rate at sea relative to the colony and (2) the assumption that transition mortality from one life-history stage to the other is size dependent. Apparently, large variation in the duration of parental care can arise simply as a result of variation in energy gain without any trade-off with safety.

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After major oil spills, hundreds to thousands of live stranded birds enter rehabilitative care. To target aspects of rehabilitative efforts for improvement and to evaluate which initial physical examination and biomedical parameters most effectively predict survival to release, medical records were examined from 913 Common Murres ( Uria aalge ; COMUs) oiled during the November 2001-January 2003 oil spill associated with the sunken S.S. Jacob Luckenbach off San Francisco, California, US. Results showed that 52% of all deaths occurred during the first 2 days of treatment. Birds stranding closest to the wreck had greater amounts of oil on their bodies than birds stranding farther away. More heavily oiled birds were in better clinical condition than birds with lesser amounts of oil, as shown by higher body mass (BM), packed cell volumes (PCV), total plasma protein (TP), and higher survival proportions. Additionally, BM, PCV, TP, and body temperature were positively correlated. For comparison, medical records from all nonoiled COMUs admitted for rehabilitation at the same facility during 2007-09 (n=468) were examined, and these variables were also found to be positively correlated. Oiled birds with BM under 750 g had approximately 5% lower PCV than BM-matched nonoiled COMUs. More heavily oiled COMUs may be in better condition than less oiled birds because heavily oiled birds must beach themselves immediately to avoid drowning and hypothermia, whereas lightly oiled birds may postpone beaching until exhausted due to extreme body catabolism. The strong relationship of PCV to BM regardless of oiling provides evidence that anemia commonly encountered in oiled seabirds may be a sequela to overall loss of body condition rather than solely due to toxic effects of oiling. Clinical information garnered in this study provides guidance for triage decisions during oil spills.

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Dietary salt intake in domestic fowl affects epithelial transport and morphology of the lower intestine (colon and coprodeum). This study investigated lower intestinal morphology and transport activity in two wild bird species with natural diets containing either low or high salt. Tissues from rock ptarmigan (Lagopus mutus) and common murres (Uria aalge) were sampled for histology and electrophysiological analyses. The ptarmigan exists on a low salt diet, while the murre lives on a high protein and high salt diet. The ptarmigan colon and coprodeum had villi/folds and crypts and the epithelium contained absorptive epithelial cells, mitochondria-rich cells and goblet cells. The colon had significant amiloride-inhibitable Isc, 5-15 µA/cm(2), with no glucose-stimulated Isc, and no significant phloridzin inhibition. The coprodeum also had high amiloride-inhibitable Isc. This transport pattern corresponded to that of chickens on low-salt diets. However, the ptarmigan colon also had a significant lysine/leucine-stimulated Isc of 3±1.0 µA/cm(2). The short U. aalge colon was similar to that of ptarmigans, but with no villi. It demonstrated a significant lysine/leucine-stimulated Isc (11±3.5 µA/cm(2)) with no amiloride-inhibitable Isc, similar to the high-salt chicken colon, but with no Na(+)-glucose cotransport. The murre coprodeum was inert to all substances and showed high resistance (1000 Ω·cm(2)), with a multilayered squamous epithelium. Despite some variations possibly associated with dietary protein intake, we conclude that natural high and low salt diets in different avian species are associated with different lower intestinal transport patterns, providing for post-renal adjustments in ion and water excretion.

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In many seabird studies, single annual proxies of prey abundance have been used to explain variability in breeding performance, but much more important is probably the timing of prey availability relative to the breeding season when energy demand is at a maximum. Until now, intraseasonal variation in prey availability has been difficult to quantify in seabirds. Using a state-of-the-art ocean drift model of larval cod Gadus morhua, an important constituent of the diet of common guillemots Uria aalge in the southwestern Barents Sea, we were able to show clear, short-term correlations between food availability and measurements of the stress hormone corticosterone (CORT) in parental guillemots over a 3-year period (2009-2011). The model allowed the extraction of abundance and size of cod larvae with very high spatial (4 km) and temporal resolutions (1 day) and showed that cod larvae from adjacent northern spawning grounds in Norway were always available near the guillemot breeding colony while those from more distant southerly spawning grounds were less frequent, but larger. The latter arrived in waves whose magnitude and timing, and thus overlap with the guillemot breeding season, varied between years. CORT levels in adult guillemots were lower in birds caught after a week with high frequencies of southern cod larvae. This pattern was restricted to the two years (2009 and 2010) in which southern larvae arrived before the end of the guillemot breeding season. Any such pattern was masked in 2011 by already exceptionally high numbers of cod larvae in the region throughout chick-rearing period. The findings suggest that CORT levels in breeding birds increase when the arrival of southern sizable larvae does not match the period of peak energy requirements during breeding.

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Within Europe, the Marine Strategy Framework Directive (MSFD) is aimed at addressing the chemical status and quality of the marine environment. One of the main goals is to achieve Good Environmental Status (GES) in the marine environment. Environmental monitoring of biota e.g., Baltic herring and guillemot eggs, is conducted annually in Sweden to follow temporal changes in environmental contaminants. To determine the suitability of guillemot eggs as a sentinel species for investigating GES, we compared temporal trends of polychlorinated dibenzo-p-dioxins and dibenzofurans (PCDD/Fs) and dioxin-like polychlorinated biphenyls (dl-PCBs) in these two species from single sampling sites within Sweden. Lipid content from guillemot eggs was consistently high and stable (yearly mean for >40 years, ∼12%) compared to that of herring (yearly mean for >20 years, ∼3%). A significant decreasing trend of ΣPCDD/F in TEQ WHO1998 was observed in guillemot eggs, but no trend was seen in herring. CB118 significantly decreased in both species, but in the last 10 years this decrease was not significant in herring. A number of advantages, such as high lipid content in the egg and a low coefficient of variation make guillemot suitable as a sentinel species. The advantages and disadvantages of using either guillemot eggs or Baltic herring are compared.

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Plastic ingestion by seabirds is a growing conservation issue, but there are few time series of plastic ingestion with large sample sizes for which one can assess temporal trends. Common and Thick-billed Murres (Uria aalge and U. lomvia) are pursuit-diving auks that are legally harvested in Newfoundland and Labrador, Canada. Here, we combined previously unpublished data on plastic ingestion (from the 1980s to the 1990s) with contemporary samples (2011-2012) to evaluate changes in murres' plastic ingestion. Approximately 7% of murres had ingested plastic, with no significant change in the frequency of ingestion among species or periods. The number of pieces of plastic/bird, and mass of plastic/bird were highest in the 1980s, lowest in the late 1990s, and intermediate in contemporary samples. Studying plastic ingestion in harvested seabird populations links harvesters to conservation and health-related issues and is a useful source of large samples for diet and plastic ingestion studies.

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