Advances in remote sensing of emperor penguins: first multi-year time series documenting trends in the global population.

Like many polar animals, emperor penguin populations are challenging to monitor because of the species' life history and remoteness. Consequently, it has been difficult to establish its global status, a subject important to resolve as polar environments change. To advance our understanding of emperor penguins, we combined remote sensing, validation surveys and using Bayesian modelling, we estimated a comprehensive population trajectory over a recent 10-year period, encompassing the entirety of the species' range. Reported as indices of abundance, our study indicates with 81% probability that there were fewer adult emperor penguins in 2018 than in 2009, with a posterior median decrease of 9.6% (95% credible interval (CI) -26.4% to +9.4%). The global population trend was -1.3% per year over this period (95% CI = -3.3% to +1.0%) and declines probably occurred in four of eight fast ice regions, irrespective of habitat conditions. Thus far, explanations have yet to be identified regarding trends, especially as we observed an apparent population uptick toward the end of time series. Our work potentially establishes a framework for monitoring other Antarctic coastal species detectable by satellite, while promoting a need for research to better understand factors driving biotic changes in the Southern Ocean ecosystem.

Human infrastructures correspond to higher Adélie penguin breeding success and growth rate.

Anthropogenic activities generate increasing disturbance in wildlife especially in extreme environments where species have to cope with rapid environmental changes. In Antarctica, while studies on human disturbance have mostly focused on stress response through physiological and behavioral changes, local variability in population dynamics has been addressed more scarcely. In addition, the mechanisms by which breeding communities are affected around research stations remain unclear. Our study aims at pointing out the fine-scale impact of human infrastructures on the spatial variability in Adélie penguin (Pygoscelis adeliae) colonies dynamics. Taking 24 years of population monitoring, we modeled colony breeding success and growth rate in response to both anthropic and land-based environmental variables. Building density around colonies was the second most important variable explaining spatial variability in breeding success after distance from skua nests, the main predators of penguins on land. Building density was positively associated with penguins breeding success. We discuss how buildings may protect penguins from avian predation and environmental conditions. The drivers of colony growth rate included topographical variables and the distance to human infrastructures. A strong correlation between 1-year lagged growth rate and colony breeding success was coherent with the use of public information by penguins to select their initial breeding site. Overall, our study brings new insights about the relative contribution and ecological implications of human presence on the local population dynamics of a sentinel species in Antarctica.

The impact of boldness on demographic rates and life-history outcomes in the wandering albatross.

Differences among individuals within a population are ubiquitous. Those differences are known to affect the entire life cycle with important consequences for all demographic rates and outcomes. One source of among-individual phenotypic variation that has received little attention from a demographic perspective is animal personality, which is defined as consistent and heritable behavioural differences between individuals. While many studies have shown that individual variation in individual personality can generate individual differences in survival and reproductive rates, the impact of personality on all demographic rates and outcomes remains to be assessed empirically. Here, we used a unique, long-term, dataset coupling demography and personality of wandering albatross (Diomedea exulans) in the Crozet Archipelago and a comprehensive analysis based on a suite of approaches (capture-mark-recapture statistical models, Markov chains models and structured matrix population models). We assessed the effect of boldness on annual demographic rates (survival, breeding probability, breeding success), life-history outcomes (life expectancy, lifetime reproductive outcome, occupancy times), and an integrative demographic outcome (population growth rate). We found that boldness had little impact on female demographic rates, but was very likely associated with lower breeding probabilities in males. By integrating the effects of boldness over the entire life cycle, we found that bolder males had slightly lower lifetime reproductive success compared to shyer males. Indeed, bolder males spent a greater proportion of their lifetime as non-breeders, which suggests longer inter-breeding intervals due to higher reproductive allocation. Our results reveal that the link between boldness and demography is more complex than anticipated by the pace-of-life literature and highlight the importance of considering the entire life cycle with a comprehensive approach when assessing the role of personality on individual performance and demography.

How does maternal age influence reproductive performance and offspring phenotype in the snow petrel (Pagodroma nivea)?

In wild vertebrates, the increase of breeding success with advancing age has been extensively studied through laying date, clutch size, hatching success, and fledging success. However, to better evaluate the influence of age on reproductive performance in species with high reproductive success, assessing not only reproductive success but also other proxies of reproductive performance appear crucial. For example, the quality of developmental conditions and offspring phenotype can provide robust and complementary information on reproductive performance. In long-lived vertebrate species, several proxies of developmental conditions can be used to estimate the quality of the produced offspring (i.e., body size, body condition, corticosterone levels, and telomere length), and therefore, their probability to survive. By sampling chicks reared by known-aged mothers, we investigated the influence of maternal age on reproductive performance and offspring quality in a long-lived bird species, the snow petrel (Pagodroma nivea). Older females bred and left their chick alone earlier. Moreover, older females had larger chicks that grew faster, and ultimately, those chicks had a higher survival probability at the nest. In addition, older mothers produced chicks with a higher sensitivity to stress, as shown by moderately higher stress-induced corticosterone levels. Overall, our study demonstrated that maternal age is correlated to reproductive performance (hatching date, duration of the guarding period and survival) and offspring quality (body size, growth rate and sensitivity to stress), suggesting that older individuals provide better parental cares to their offspring. These results also demonstrate that maternal age can affect the offspring phenotype with potential long-term consequences.

Machine and deep learning approaches to understand and predict habitat suitability for seabird breeding.

The way animals select their breeding habitat may have great impacts on individual fitness. This complex process depends on the integration of information on various environmental factors, over a wide range of spatiotemporal scales. For seabirds, breeding habitat selection integrates both land and sea features over several spatial scales. Seabirds explore these features prior to breeding, assessing habitats' quality. However, the information-gathering and decision-making process by seabirds when choosing a breeding habitat remains poorly understood. We compiled 49 historical records of larids colonies in Cuba from 1980 to 2020. Then, we predicted potentially suitable breeding sites for larids and assessed their breeding macrohabitat selection, using deep and machine learning algorithms respectively. Using a convolutional neural network and Landsat satellite images we predicted the suitability for nesting of non-monitored sites of this archipelago. Furthermore, we assessed the relative contribution of 18 land- and marine-based environmental covariates describing macrohabitats at three spatial scales (i.e. 10, 50 and 100 km) using random forests. Convolutional neural network exhibited good performance at training, validation and test (F1-scores >85%). Sites with higher habitat suitability (p > .75) covered 20.3% of the predicting area. Larids breeding macrohabitats were sites relatively close to main islands, featuring sparse vegetation cover and high chlorophyll-a concentration at sea in 50 and 100 km around colonies. Lower sea surface temperature at larger spatial scales was determinant to distinguish the breeding from non-breeding sites. A more comprehensive understanding of the seabird breeding macrohabitats selection can be reached from the complementary use of convolutional neural networks and random forest models. Our analysis provides crucial knowledge in tropical regions that lack complete and regular monitoring of seabirds' breeding sites.

Investigating two consecutive catastrophic breeding seasons in a large king penguin colony.

Large-scale breeding failures, such as offspring die-offs, can disproportionately impact wildlife populations that are characterized by a few large colonies. However, breeding monitoring-and thus investigations of such die-offs-is especially challenging in species with long reproductive cycles. We investigate two unresolved dramatic breeding failures that occurred in consecutive years (2009 and 2010) in a large king penguin Aptenodytes patagonicus colony, a long-lived species with a breeding cycle lasting over a year. Here we found that a single period, winter 2009, was likely responsible for the occurrence of breeding anomalies during both breeding seasons, suggesting that adults experienced poor foraging conditions at sea at that time. Following that unfavorable winter, the 2009 breeding cohort-who were entering the late stage of chick-rearing-immediately experienced high chick mortality. Meanwhile, the 2010 breeding cohort greatly delayed their arrival and egg laying, which would have otherwise started not long after the winter. The 2010 breeding season continued to display anomalies during the incubation and chick-rearing period, such as high abandonment rate, long foraging trips and eventually the death of all chicks in winter 2010. These anomalies could have resulted from either a domino-effect caused by the delayed laying, the continuation of poor foraging conditions, or both. This study provides an example of a large-scale catastrophic breeding failure and highlights the importance of the winter period on phenology and reproduction success for wildlife that breed in few large colonies.

Individual life histories: neither slow nor fast, just diverse.

The slow-fast continuum is a commonly used framework to describe variation in life-history strategies across species. Individual life histories have also been assumed to follow a similar pattern, especially in the pace-of-life syndrome literature. However, whether a slow-fast continuum commonly explains life-history variation among individuals within a population remains unclear. Here, we formally tested for the presence of a slow-fast continuum of life histories both within populations and across species using detailed long-term individual-based demographic data for 17 bird and mammal species with markedly different life histories. We estimated adult lifespan, age at first reproduction, annual breeding frequency, and annual fecundity, and identified the main axes of life-history variation using principal component analyses. Across species, we retrieved the slow-fast continuum as the main axis of life-history variation. However, within populations, the patterns of individual life-history variation did not align with a slow-fast continuum in any species. Thus, a continuum ranking individuals from slow to fast living is unlikely to shape individual differences in life histories within populations. Rather, individual life-history variation is likely idiosyncratic across species, potentially because of processes such as stochasticity, density dependence, and individual differences in resource acquisition that affect species differently and generate non-generalizable patterns across species.

Global assessment of marine plastic exposure risk for oceanic birds.

Plastic pollution is distributed patchily around the world's oceans. Likewise, marine organisms that are vulnerable to plastic ingestion or entanglement have uneven distributions. Understanding where wildlife encounters plastic is crucial for targeting research and mitigation. Oceanic seabirds, particularly petrels, frequently ingest plastic, are highly threatened, and cover vast distances during foraging and migration. However, the spatial overlap between petrels and plastics is poorly understood. Here we combine marine plastic density estimates with individual movement data for 7137 birds of 77 petrel species to estimate relative exposure risk. We identify high exposure risk areas in the Mediterranean and Black seas, and the northeast Pacific, northwest Pacific, South Atlantic and southwest Indian oceans. Plastic exposure risk varies greatly among species and populations, and between breeding and non-breeding seasons. Exposure risk is disproportionately high for Threatened species. Outside the Mediterranean and Black seas, exposure risk is highest in the high seas and Exclusive Economic Zones (EEZs) of the USA, Japan, and the UK. Birds generally had higher plastic exposure risk outside the EEZ of the country where they breed. We identify conservation and research priorities, and highlight that international collaboration is key to addressing the impacts of marine plastic on wide-ranging species.

Causes and consequences of pair-bond disruption in a sex-skewed population of a long-lived monogamous seabird.

Many animals form long-term monogamous pair bonds, and the disruption of a pair bond (through either divorce or widowhood) can have significant consequences for individual vital rates (survival, breeding, and breeding success probabilities) and life-history outcomes (lifetime reproductive success [LRS], life expectancy). Here, we investigated the causes and consequences of pair-bond disruption in wandering albatross (Diomedea exulans). State-of-the-art statistical and mathematical approaches were developed to estimate divorce and widowhood rates and their impacts on vital rates and life-history outcomes. In this population, females incur a higher mortality rate due to incidental fishery bycatch, so the population is male-skewed. Therefore, we first posited that males would show higher widowhood rates negatively correlated with fishing effort and females would have higher divorce rates because they have more mating opportunities. Furthermore, we expected that divorce could be an adaptive strategy, whereby individuals improved breeding success by breeding with a new partner of better quality. Finally, we posited that pair-bond disruptions could reduce survival and breeding probabilities owing to the cost of remating processes, with important consequences for life-history outcomes. As expected, we showed that males had higher widowhood rates than females and females had higher divorce rates in this male-skewed population. However, no correlation was found between fishing effort and male widowhood. Secondly, contrary to our expectation, we found that divorce was likely nonadaptive in this population. We propose that divorce in this population is caused by an intruder who outcompetes the original partner in line with the so-called forced divorce hypothesis. Furthermore, we found a 16.7% and 18.0% reduction in LRS only for divorced and widowed males, respectively, owing to missing breeding seasons after a pair-bond disruption. Finally, we found that divorced individuals were more likely to divorce again, but whether this is related to specific individual characteristics remains an important area of investigation.

Boldness predicts divorce rates in wandering albatrosses (Diomedea exulans).

Personality predicts divorce rates in humans, yet how personality traits affect divorce in wild animals remains largely unknown. In a male-skewed population of wandering albatross (Diomedea exulans), we showed that personality predicts divorce; shyer males exhibited higher divorce rates than bolder males but no such relationship was found in females. We propose that divorce may be caused by the intrusion of male competitors and shyer males divorce more often because of their avoidance of territorial aggression, while females have easier access to mates regardless of their personality. Thus, personality may have important implications for the dynamics of social relationships.

When the going gets tough, the tough get going: Effect of extreme climate on an Antarctic seabird's life history.

Individuals differ in many ways. Most produce few offspring; a handful produce many. Some die early; others live to old age. It is tempting to attribute these differences in outcomes to differences in individual traits, and thus in the demographic rates experienced. However, there is more to individual variation than meets the eye of the biologist. Even among individuals sharing identical traits, life history outcomes (life expectancy and lifetime reproduction) will vary due to individual stochasticity, that is to chance. Quantifying the contributions of heterogeneity and chance is essential to understand natural variability. Interindividual differences vary across environmental conditions, hence heterogeneity and stochasticity depend on environmental conditions. We show that favourable conditions increase the contributions of individual stochasticity, and reduce the contributions of heterogeneity, to variance in demographic outcomes in a seabird population. The opposite is true under poor conditions. This result has important consequence for understanding the ecology and evolution of life history strategies.

Temporal correlations among demographic parameters are ubiquitous but highly variable across species.

Temporal correlations among demographic parameters can strongly influence population dynamics. Our empirical knowledge, however, is very limited regarding the direction and the magnitude of these correlations and how they vary among demographic parameters and species' life histories. Here, we use long-term demographic data from 15 bird and mammal species with contrasting pace of life to quantify correlation patterns among five key demographic parameters: juvenile and adult survival, reproductive probability, reproductive success and productivity. Correlations among demographic parameters were ubiquitous, more frequently positive than negative, but strongly differed across species. Correlations did not markedly change along the slow-fast continuum of life histories, suggesting that they were more strongly driven by ecological than evolutionary factors. As positive temporal demographic correlations decrease the mean of the long-run population growth rate, the common practice of ignoring temporal correlations in population models could lead to the underestimation of extinction risks in most species.

A U-Turn for Mercury Concentrations over 20 Years: How Do Environmental Conditions Affect Exposure in Arctic Seabirds?

Mercury (Hg) is highly toxic in its methylated form (MeHg), and global change is likely to modify its bioavailability in the environment. However, it is unclear how top predators will be impacted. We studied blood Hg concentrations of chick-rearing black-legged kittiwakes Rissa tridactyla (2000-2019) in Svalbard (Norway). From 2000 to 2019, Hg concentrations followed a U-shaped trend. The trophic level, inferred from nitrogen stable isotopes, and chlorophyll a (Chl a) concentrations better predicted Hg concentrations, with positive and U-shaped associations, respectively. As strong indicators of primary productivity, Chl a concentrations can influence production of upper trophic levels and, thus, fish community assemblage. In the early 2000s, the high Hg concentrations were likely related to a higher proportion of Arctic prey in kittiwake's diet. The gradual input of Atlantic prey in kittiwake diet could have resulted in a decrease in Hg concentrations until 2013. Then, a new shift in the prey community, added to the shrinking sea ice-associated release of MeHg in the ocean, could explain the increasing trend of Hg observed since 2014. The present monitoring provides critical insights about the exposure of a toxic contaminant in Arctic wildlife, and the reported increase since 2014 raises concern for Arctic seabirds.

The call of the emperor penguin: Legal responses to species threatened by climate change.

Species extinction risk is accelerating due to anthropogenic climate change, making it urgent to protect vulnerable species through legal frameworks in order to facilitate conservation actions that help mitigate risk. Here, we discuss fundamental concepts for assessing climate change risks to species using the example of the emperor penguin (Aptenodytes forsteri), currently being considered for protection under the US Endangered Species Act (ESA). This species forms colonies on Antarctic sea ice, which is projected to significantly decline due to ongoing greenhouse gas (GHG) emissions. We project the dynamics of all known emperor penguin colonies under different GHG emission scenarios using a climate-dependent meta-population model including the effects of extreme climate events based on the observational satellite record of colonies. Assessments for listing species under the ESA require information about how species resiliency, redundancy and representation (3Rs) will be affected by threats within the foreseeable future. Our results show that if sea ice declines at the rate projected by climate models under current energy system trends and policies, the 3Rs would be dramatically reduced and almost all colonies would become quasi-extinct by 2100. We conclude that the species should be listed as threatened under the ESA.

Landfast ice: a major driver of reproductive success in a polar seabird.

In a fast-changing world, polar ecosystems are threatened by climate variability. Understanding the roles of fine-scale processes, and linear and nonlinear effects of climate factors on the demography of polar species is crucial for anticipating the future state of these fragile ecosystems. While the effects of sea ice on polar marine top predators are increasingly being studied, little is known about the impacts of landfast ice (LFI) on this species community. Based on a unique 39-year time series of satellite imagery and in situ meteorological conditions and on the world's longest dataset of emperor penguin (Aptenodytes forsteri) breeding parameters, we studied the effects of fine-scale variability of LFI and weather conditions on this species' reproductive success. We found that longer distances to the LFI edge (i.e. foraging areas) negatively affected the overall breeding success but also the fledging success. Climate window analyses suggested that chick mortality was particularly sensitive to LFI variability between August and November. Snowfall in May also affected hatching success. Given the sensitivity of LFI to storms and changes in wind direction, important future repercussions on the breeding habitat of emperor penguins are to be expected in the context of climate change.

The Indian Ocean 'garbage patch': Empirical evidence from floating macro-litter.

Marine litter has become a global issue with 'garbage patches' documented in all ocean gyres. The Pacific and Atlantic garbage patches have been well described, but there are few empirical data for the Indian Ocean. In the austral summer 2019-2020, we conducted an at-sea survey of macro-litter in the rarely investigated south-west Indian Ocean. Over 24 days, 1623 man-made items were observed including plastic fragments, packaging and fishing-related items during 216 h of observations covering 5464 km. More than 99% of the litter items were plastics of which almost 60% were white. Floating litter was patchily distributed with only five items (0.2%) recorded south of 40°S (0.1 items·km-2). Half of the items were encountered over a two-day period south-east of Madagascar (30°S; 59-67°E; 75.2 items·km-2). Our survey detected an accumulation of litter in the southern Indian Ocean and demonstrated that this area warrants more research.

"Home alone!" influence of nest parental attendance on offspring behavioral and hormonal stress responses in an Antarctic seabird, the snow petrel (Pagodroma nivea).

In altricial species, parents brood their chicks constantly before leaving them unattended sometimes for extended periods when they become thermally independent. During this second phase, there is sometimes important inter-individual differences in parental attendance and the fitness costs and benefits of parental strategies have previously been extensively investigated. However, the impact of parental presence on offspring behaviors and stress physiology has been overlooked. Here, we examined the influence of parental presence on offspring hormonal and behavioral stress sensitivities in snow petrel chicks. We demonstrated for the first time in a wild bird species that attended chicks had lower stress-induced corticosterone levels and a lower probability to show defensive behavior compared to the alone chicks. This reduced stress sensitivity is certainly explained by the well-known link between corticosterone and nutritional status, and by the recent delivery of meals to the attended chicks and the improvement of their nutritional status. It may also be explained by the parental protection against predators or inclement weather, or/and by the psychosocial comfort of parental presence for the offspring. Overall, these results suggest that the presence of a parent in the nest reduces offspring stress sensitivity in wild birds. Further studies would now be required to disentangle the impact of nutritional status and parental presence on stress sensitivity and to better understand the potential impact of parental presence and circulating corticosterone levels on growth and cognitive development in wild birds.

Selection against immigrants in wild seabird populations.

Immigration is a major demographic parameter shaping population dynamics and is an important driver of eco-evolutionary patterns, but the fitness consequences for individuals following their settlement to a new population (immigrants) remain poorly tested in wild animal populations, particularly among long-lived species. Here we show that immigrants have a lower fitness than residents in three wild seabird populations (wandering albatross Diomedea exulans, southern fulmar Fulmarus glacialoides, snow petrel Pagodroma nivea). Across all species and during a 32-year period, immigrants made on average -9 to 29% fewer breeding attempts, had 5-31% fewer fledglings, had 2-16% lower breeding success and produced 6-46% fewer recruits. Female immigration and male residency were also favored through differences in breeding performance. We provide evidence for selection against immigrants in wild populations of long-lived species and our results are consistent with female-biased dispersal in birds being driven by asymmetric limiting resources and the competitive ability of dispersers vs. non-dispersers.

Linking 19th century European settlement to the disruption of a seabird's natural population dynamics.

Recent estimates indicate that ∼70% of the world's seabird populations have declined since the 1950s due to human activities. However, for almost all bird populations, there is insufficient long-term monitoring to understand baseline (i.e., preindustrial) conditions, which are required to distinguish natural versus anthropogenically driven changes. Here, we address this lack of long-term monitoring data with multiproxy paleolimnological approaches to examine the long-term population dynamics of a major colony of Leach's Storm-petrel (Hydrobates leucorhous) on Grand Colombier Island in the St. Pierre and Miquelon archipelago-an overseas French territory in the northwest Atlantic Ocean. By reconstructing the last ∼5,800 y of storm-petrel dynamics, we demonstrate that this colony underwent substantial natural fluctuations until the start of the 19th century, when population cycles were disrupted, coinciding with the establishment and expansion of a European settlement. Our paleoenvironmental data, coupled with on-the-ground population surveys, indicate that the current colony is only ∼16% of the potential carrying capacity, reinforcing concerning trends of globally declining seabird populations. As seabirds are sentinel species of marine ecosystem health, such declines provide a call to action for global conservation. In response, we emphasize the need for enlarged protected areas and the rehabilitation of disturbed islands to protect ecologically critical seabird populations. Furthermore, long-term data, such as those provided by paleoecological approaches, are required to better understand shifting baselines in conservation to truly recognize current rates of ecological loss.