Centennial relationships between ocean temperature and Atlantic puffin production reveal shifting decennial trends.

The current warming of the oceans has been shown to have detrimental effects for a number of species. An understanding of the underlying mechanisms may be hampered by the non-linearity and non-stationarity of the relationships between temperature and demography, and by the insufficient length of available time series. Most demographic time series are too short to study the effects of climate on wildlife in the classical sense of meteorological patterns over at least 30 years. Here we present a harvest time series of Atlantic puffins (Fratercula arctica) that goes back as far as 1880. It originates in the world's largest puffin colony, in southwest Iceland, which has recently experienced a strong decline. By estimating an annual chick production index for 128 years, we found prolonged periods of strong correlations between local sea surface temperature (SST) and chick production. The sign of decennial correlations switches three times during this period, where the phases of strong negative correlations between puffin productivity and SST correspond to the early 20th century Arctic warming period and to the most recent decades. Most of the variation (72%) in chick production is explained by a model in which productivity peaks at an SST of 7.1°C, clearly rejecting the assumption of a linear relationship. There is also evidence supporting non-stationarity: The SST at which puffins production peaked has increased by 0.24°C during the 20th century, although the increase in average SST during the same period has been more than three times faster. The best supported models indicate that the population's decline is at least partially caused by the increasing SST around Iceland.

Local prey shortages drive foraging costs and breeding success in a declining seabird, the Atlantic puffin.

As more and more species face anthropogenic threats, understanding the causes of population declines in vulnerable taxa is essential. However, long-term datasets, ideal to identify lasting or indirect effects on fitness measures such as those caused by environmental factors, are not always available. Here we use a single year but multi-population approach on populations with contrasting demographic trends to identify possible drivers and mechanisms of seabird population changes in the north-east Atlantic, using the Atlantic puffin, a declining species, as a model system. We combine miniature GPS trackers with camera traps and DNA metabarcoding techniques on four populations across the puffins' main breeding range to provide the most comprehensive study of the species' foraging ecology to date. We find that puffins use a dual foraging tactic combining short and long foraging trips in all four populations, but declining populations in southern Iceland and north-west Norway have much greater foraging ranges, which require more (costly) flight, as well as lower chick-provisioning frequencies, and a more diverse but likely less energy-dense diet, than stable populations in northern Iceland and Wales. Together, our findings suggest that the poor productivity of declining puffin populations in the north-east Atlantic is driven by breeding adults being forced to forage far from the colony, presumably because of low prey availability near colonies, possibly amplified by intraspecific competition. Our results provide valuable information for the conservation of this and other important North-Atlantic species and highlight the potential of multi-population approaches to answer important questions about the ecological drivers of population trends.

Mercury contamination and potential health risks to Arctic seabirds and shorebirds.

Since the last Arctic Monitoring and Assessment Programme (AMAP) effort to review biological effects of mercury (Hg) on Arctic biota in 2011 and 2018, there has been a considerable number of new Arctic bird studies. This review article provides contemporary Hg exposure and potential health risk for 36 Arctic seabird and shorebird species, representing a larger portion of the Arctic than during previous AMAP assessments now also including parts of the Russian Arctic. To assess risk to birds, we used Hg toxicity benchmarks established for blood and converted to egg, liver, and feather tissues. Several Arctic seabird populations showed Hg concentrations that exceeded toxicity benchmarks, with 50 % of individual birds exceeding the "no adverse health effect" level. In particular, 5 % of all studied birds were considered to be at moderate or higher risk to Hg toxicity. However, most seabirds (95 %) were generally at lower risk to Hg toxicity. The highest Hg contamination was observed in seabirds breeding in the western Atlantic and Pacific Oceans. Most Arctic shorebirds exhibited low Hg concentrations, with approximately 45 % of individuals categorized at no risk, 2.5 % at high risk category, and no individual at severe risk. Although the majority Arctic-breeding seabirds and shorebirds appeared at lower risk to Hg toxicity, recent studies have reported deleterious effects of Hg on some pituitary hormones, genotoxicity, and reproductive performance. Adult survival appeared unaffected by Hg exposure, although long-term banding studies incorporating Hg are still limited. Although Hg contamination across the Arctic is considered low for most bird species, Hg in combination with other stressors, including other contaminants, diseases, parasites, and climate change, may still cause adverse effects. Future investigations on the global impact of Hg on Arctic birds should be conducted within a multi-stressor framework. This information helps to address Article 22 (Effectiveness Evaluation) of the Minamata Convention on Mercury as a global pollutant.

Multicomponent shell traits are consistent with an individual recognition function of the appearance of common murre (Uria aalge) eggs: A biological replication study.

In dense breeding colonies, and despite having no nest structure, common murres (or guillemots: Uria aalge) are still able to identify their own eggs. Each female murre's egg is thought to be recognized individually by the shell's avian-perceivable traits. This is because the eggshells' visible traits conform to expectations of the identity-signaling hypothesis in that they show both high intraindividual repeatability and high interindividual variability. Identity signaling also predicts a lack of correlation between each of the putative multicomponent recognition traits, yielding no significant relationships between those eggshell traits that are generated by mutually exclusive physiological factors. Using a multivariate analysis across eggshell size and shape, avian-perceivable background coloration, spot (maculation) shape, and spot density, we detected no unexpected statistical correlations between Icelandic common murre egg traits lacking known physiological or mathematical relationships with one another. These results biologically replicate the conclusions of a recent eggshell trait study of Canadian common murres using similar methodology. We also demonstrate the use of static correlations to infer identity signaling function without direct behavioral observations, which in turn may also be applied to rare or extinct species and provide valuable insight into otherwise unknown communicative and behavioral functions.

Complex population structure of the Atlantic puffin revealed by whole genome analyses.

The factors underlying gene flow and genomic population structure in vagile seabirds are notoriously difficult to understand due to their complex ecology with diverse dispersal barriers and extensive periods at sea. Yet, such understanding is vital for conservation management of seabirds that are globally declining at alarming rates. Here, we elucidate the population structure of the Atlantic puffin (Fratercula arctica) by assembling its reference genome and analyzing genome-wide resequencing data of 72 individuals from 12 colonies. We identify four large, genetically distinct clusters, observe isolation-by-distance between colonies within these clusters, and obtain evidence for a secondary contact zone. These observations disagree with the current taxonomy, and show that a complex set of contemporary biotic factors impede gene flow over different spatial scales. Our results highlight the power of whole genome data to reveal unexpected population structure in vagile marine seabirds and its value for seabird taxonomy, evolution and conservation.

The chemical basis of a signal of individual identity: shell pigment concentrations track the unique appearance of Common Murre eggs.

In group-living species with parental care, the accurate recognition of one's own young is critical to fitness. Because discriminating offspring within a large colonial group may be challenging, progeny of colonial breeders often display familial or individual identity signals to elicit and receive parental provisions from their own parents. For instance, the common murre (or common guillemot: Uria aalge) is a colonially breeding seabird that does not build a nest and lays and incubates an egg with an individually unique appearance. How the shell's physical and chemical properties generate this individual variability in coloration and maculation has not been studied in detail. Here, we quantified two characteristics of the avian-visible appearance of murre eggshells collected from the wild: background coloration spectra and maculation density. As predicted by the individual identity hypothesis, there was no statistical relationship between avian-perceivable shell background coloration and maculation density within the same eggs. In turn, variation in both sets of traits was statistically related to some of their physico-chemical properties, including shell thickness and concentrations of the eggshell pigments biliverdin and protoporphyrin IX. These results illustrate how individually unique eggshell appearances, suitable for identity signalling, can be generated by a small number of structural mechanisms.

Ocean-wide Drivers of Migration Strategies and Their Influence on Population Breeding Performance in a Declining Seabird.

Which factors shape animals' migration movements across large geographical scales, how different migratory strategies emerge between populations, and how these may affect population dynamics are central questions in the field of animal migration [1] that only large-scale studies of migration patterns across a species' range can answer [2]. To address these questions, we track the migration of 270 Atlantic puffins Fratercula arctica, a red-listed, declining seabird, across their entire breeding range. We investigate the role of demographic, geographical, and environmental variables in driving spatial and behavioral differences on an ocean-basin scale by measuring puffins' among-colony differences in migratory routes and day-to-day behavior (estimated with individual daily activity budgets and energy expenditure). We show that competition and local winter resource availability are important drivers of migratory movements, with birds from larger colonies or with poorer local winter conditions migrating further and visiting less-productive waters; this in turn led to differences in flight activity and energy expenditure. Other behavioral differences emerge with latitude, with foraging effort and energy expenditure increasing when birds winter further north in colder waters. Importantly, these ocean-wide migration patterns can ultimately be linked with breeding performance: colony productivity is negatively associated with wintering latitude, population size, and migration distance, which demonstrates the cost of competition and migration on future breeding and the link between non-breeding and breeding periods. Our results help us to understand the drivers of animal migration and have important implications for population dynamics and the conservation of migratory species.