Interannual measures of nutritional stress during a marine heatwave (the Blob) differ between two North Pacific seabird species.

'The Blob', a mass of anomalously warm water in the Northeast Pacific Ocean peaking from 2014 to 2016, caused a decrease in primary productivity with cascading effects on the marine ecosystem. Among the more obvious manifestations of the event were seabird breeding failures and mass mortality events. Here, we used corticosterone in breast feathers (fCort), grown in the winter period during migration, as an indicator of nutritional stress to investigate the impact of the Blob on two sentinel Pacific auk species (family Alcidae). Feathers were collected from breeding females over 8 years from 2010 to 2017, encompassing the Blob period. Since Pacific auks replace body feathers at sea during the migratory period, measures of fCort provide an accumulated measure of nutritional stress or allostatic load during this time. Changes in diet were also measured using δ15N and δ13C values from feathers. Relative to years prior to the Blob, the primarily zooplanktivorous Cassin's auklets (Ptychoramphus aleuticus) had elevated fCort in 2014-2017, which correlated with the occurrence of the Blob and a recovery period afterwards, with relatively stable feather isotope values. In contrast, generalist rhinoceros auklets (Cerorhinca monocerata) displayed stable fCort values across years and increased δ15N values during the Blob. As marine heatwaves increase in intensity and frequency due to climate change, this study provides insight into the variable response of Pacific auks to such phenomena and suggests a means for monitoring population-level responses to climatological variation.

Exposure to persistent organic pollutants is linked to over-wintering latitude in a Pacific seabird, the rhinoceros auklet, Cerorhinca monocerata.

Seabirds are wide-ranging organisms often used to track marine pollution, yet the effect of migration on exposure over the annual cycle is often unclear. We used solar geolocation loggers and stable isotope analysis to study the effects of post breeding dispersal and diet on persistent organic pollutant (POP) and mercury (Hg) burdens in rhinoceros auklets, Cerorhinca monocerata, breeding on islands along the Pacific Coast of Canada. Hg and four classes of POPs were measured in auklet eggs: organochlorine insecticides (OCs), polychlorinated biphenyls (PCBs), polybrominated diphenyl ethers (PBDEs), and perfluoralkyl substances (PFASs). Stable isotope values of adult breast feathers grown during winter were used in conjunction with geolocation to elucidate adult wintering latitude. Wintering latitude was the most consistent and significant predictor of some POP and of Hg concentrations in eggs. The magnitude and pattern of exposure varied by contaminant, with ∑PCBs, ∑PBDEs and DDE decreasing with wintering latitude, and mirex, perfluoro-n-tridecanoic acid, and Hg increasing with latitude. We suggest that concentrations of these contaminants in rhinoceros auklet eggs are influenced by variation in uptake at adult wintering locations related to anthropogenic inputs and oceanic and atmospheric transport.

Geolocator tagging links distributions in the non-breeding season to population genetic structure in a sentinel North Pacific seabird.

We tested the hypothesis that segregation in wintering areas is associated with population differentiation in a sentinel North Pacific seabird, the rhinoceros auklet (Cerorhinca monocerata). We collected tissue samples for genetic analyses on five breeding colonies in the western Pacific Ocean (Japan) and on 13 colonies in the eastern Pacific Ocean (California to Alaska), and deployed light-level geolocator tags on 12 eastern Pacific colonies to delineate wintering areas. Geolocator tags were deployed previously on one colony in Japan. There was strong genetic differentiation between populations in the eastern vs. western Pacific Ocean, likely due to two factors. First, glaciation over the North Pacific in the late Pleistocene might have forced a southward range shift that historically isolated the eastern and western populations. And second, deep-ocean habitat along the northern continental shelf appears to act as a barrier to movement; abundant on both sides of the North Pacific, the rhinoceros auklet is virtually absent as a breeder in the Aleutian Islands and Bering Sea, and no tagged birds crossed the North Pacific in the non-breeding season. While genetic differentiation was strongest between the eastern vs. western Pacific, there was also extensive differentiation within both regional groups. In pairwise comparisons among the eastern Pacific colonies, the standardized measure of genetic differentiation (FꞌST) was negatively correlated with the extent of spatial overlap in wintering areas. That result supports the hypothesis that segregation in the non-breeding season is linked to genetic structure. Philopatry and a neritic foraging habit probably also contribute to the structuring. Widely distributed, vulnerable to anthropogenic stressors, and exhibiting extensive genetic structure, the rhinoceros auklet is fully indicative of the scope of the conservation challenges posed by seabirds.

Seasonal variability in vulnerability for Cassin's auklets (Ptychoramphus aleuticus) exposed to microplastic pollution in the Canadian Pacific region.

Marine plastic pollution is an emerging global conservation challenge, potentially impacting organisms at all trophic levels. However, currently it is unclear to what extent plastic pollution is impacting marine organisms at the population, species or multispecies level. In this study, we explore seasonal exposure (i.e., vulnerability) of Cassin's Auklet (Ptychoramphus aleuticus) to plastic pollution with exposure models during boreal summer and winter seasons. Based on these models, we infer exposure at the population level for this species, in the Canadian Pacific region where approximately 75% of the global population of this species breeds. The models quantify plastic exposure by determining seasonal core foraging areas and plastic concentrations found in those same areas. Core foraging areas were determined using a Generalized Additive Model based on at-sea observation data (collected year round: 1990-2010) and 50% Home Range Kernels based on aerial telemetry data (May and June 1999-2001). Plastic concentrations within these core areas were interpolated based on seawater microplastic concentrations from the summer of 2012. We found that during the boreal summer, Cassin's Auklets were exposed to relatively low concentrations of plastics. During the winter, auklet distribution shifted towards the coast where plastic concentrations are considerably higher. Model derived seasonal variability in exposure was consistent with necropsy results from bird carcasses recovered during the winter of 2014, and from a multiyear study on chick provisioning during the summer. Local oceanography likely plays a role in determining seasonal shifts in both marine bird as well microplastic concentrations, and hence exposure. As well, individual sensitivity (i.e., dose-dependent effect) may vary with annual cycles. Currently, research is focusing on determining how sensitive individual birds are to microplastic concentrations, and our models will help translate sensitivity found at the individual level to potential impacts at population or species level.

Beach-cast debris surveys on Triangle Island, British Columbia, Canada indicate the timing of arrival of 2011 Tohoku tsunami debris in North America.

We conducted beach-cast debris transect surveys on Triangle Island, British Columbia, Canada in 2012-2017 to (1) establish a baseline against which to track future changes in stranded debris on this small, uninhabited island; and (2) time the arrival in western North America of debris released by the 2011 Tohoku tsunami. Most (90%) of the six-year total of 6784 debris items tallied was composed of Styrofoam or plastic. The number of debris items peaked in 2014 (waste Styrofoam, rope) and 2015 (waste plastic, wood), and cumulative totals for all debris types were ca. 50% higher in 2014-15 than in 2012-13 and 2016-17. The peaks in 2014-15 probably represented the arrival of the bulk of the tsunami debris, based on close correspondence with forecasting models and debris surveys elsewhere. A fuller understanding of the movement of the Tohoku tsunami debris will require information from other beach monitoring programs.

Egg size is independent of variation in pre-breeding feather corticosterone in Cassin's auklets during favorable oceanographic conditions.

The measurement of corticosterone levels in feathers (fCort) is gaining recognition as an effective means for describing links between stages of the annual cycle in birds. Many seabirds are especially good models for exploring these links, or carryover effects, due to their migratory behavior and reproductive investment in a single-egg clutch. Here, we measure fCort in Cassin's auklet (Ptychoramphus aleuticus) breast feathers at two colonies in British Columbia during 2011, a year of favorable oceanographic conditions, and examine its relationship with egg size. These feathers are grown at sea during the late winter period, 1-2 months prior to egg laying. Assuming that fCort provides some measure of nutritional stress and hypothalamic-pituitary-adrenal axis activity during feather growth, we predicted that fCort would be positively correlated with egg size via increased support for foraging activity during the nutritionally demanding molt stage. We also analyzed the δ13C and δ15N stable isotope content of breast feathers, measures commonly used to characterize seabird diet composition. Contrary to prediction, neither fCort nor stable isotope ratios were good predictors of egg size. Our results appear to conflict with two previous studies on alcids in which fCort and stable isotopes showed clear links with egg size; however, both studies were conducted in years when oceanographic and foraging conditions were poor. Under these conditions, upregulation of corticosterone may be needed to mediate the energetic tradeoffs between self-maintenance and reproduction, supporting increased foraging effort and thus increasing both the likelihood of reproduction and large egg size. However, when foraging conditions are favorable, we suggest that such tradeoffs and associated physiological constraints are minimized and regulation of egg size may be effectively independent of circulating corticosterone levels and diet type.

Two forage fishes as potential conduits for the vertical transfer of microfibres in Northeastern Pacific Ocean food webs.

We assessed the potential role played by two vital Northeastern Pacific Ocean forage fishes, the Pacific sand lance (Ammodytes personatus) and Pacific herring (Clupea pallasii), as conduits for the vertical transfer of microfibres in food webs. We quantified the number of microfibres found in the stomachs of 734 sand lance and 205 herring that had been captured by an abundant seabird, the rhinoceros auklet (Cerorhinca monocerata). Sampling took place on six widely-dispersed breeding colonies in British Columbia, Canada, and Washington State, USA, over one to eight years. The North Pacific Ocean is a global hotspot for pollution, yet few sand lance (1.5%) or herring (2.0%) had ingested microfibres. In addition, there was no systematic relationship between the prevalence of microplastics in the fish stomachs vs. in waters around three of our study colonies (measured in an earlier study). Sampling at a single site (Protection Island, WA) in a single year (2016) yielded most (sand lance) or all (herring) of the microfibres recovered over the 30 colony-years of sampling involved in this study, yet no microfibres had been recovered there, in either species, in the previous year. We thus found no evidence that sand lance and herring currently act as major food-web conduits for microfibres along British Columbia's outer coast, nor that the local at-sea density of plastic necessarily determines how much plastic enters marine food webs via zooplanktivores. Extensive urban development around the Salish Sea probably explains the elevated microfibre loads in fishes collected on Protection Island, but we cannot account for the between-year variation. Nonetheless, the existence of such marked interannual variation indicates the importance of measuring year-to-year variation in microfibre pollution both at sea and in marine biota.

Size- and condition-dependent predation: a seabird disproportionately targets substandard individual juvenile salmon.

Selection of prey that are small and in poor body condition is a widespread phenomenon in terrestrial predator-prey systems and may benefit prey populations by removing substandard individuals. Similar selection is widely assumed to operate in aquatic systems. Indeed, size-selective predation is a longstanding and central tenet of aquatic food web theory. However, it is not known if aquatic predators select prey based on their condition or state, compared to their size. Surprisingly, no comparable information is available for marine systems because it is exceedingly difficult to make direct observations in this realm. Thus the role of body condition in regulating susceptibility to predation remains a black box in the marine environment. Here we have exploited an ideal model system to evaluate selective predation on pelagic marine fish: comparing characteristics (fork length, mass corrected for fork length) of fresh, whole, intact juvenile Pacific salmon delivered by a seabird to its single nestling with salmon collected concurrently in coastal trawl surveys. Three species of juvenile salmon (Oncorhynchus spp.) are consumed by provisioning Rhinoceros Auklets (Cerorhinca nionocerata); an abundant, colonial, pursuit-diving seabird. Samples were collected from multiple colonies and fisheries surveys in coastal British Columbia in two years. As predicted, Auklets preyed on small individuals in poor condition and consistently selected them at levels higher than their relative availability. This is the first study to provide direct evidence for both size- and condition-selective predation on marine fish in the wild. We anticipate that our results will be a starting point in evaluating how selective predation may structure or influence marine fish populations and bridges a fundamental incongruity between ecological theory and application; although "bigger is better" is considered a fundamental tenet of marine food webs, marine predators are often assumed to consume indiscriminately.

Leucocyte profiles of Arctic marine birds: correlates of migration and breeding phenology.

Most Arctic marine birds are migratory, wintering south of the limit of annual pack ice and returning north each year for the physiologically stressful breeding season. The Arctic environment is changing rapidly due to global warming and anthropogenic activities, which may influence the timing of breeding in relation to arrival times following migration, as well as providing additional stressors (e.g. disturbance from ships) to which birds may respond. During stressful parts of their annual cycle, such as breeding, birds may reallocate resources so that they have increased heterophil-to-lymphocyte ratios in their white blood cell (leucocyte) profiles. We analysed leucocyte profiles of nine species of marine birds to establish reference ranges for these species in advance of future Arctic change. Leucocyte profiles tended to cluster among taxonomic groups across studies, suggesting that reference values for a particular group can be established, and within species there was evidence that birds from colonies that had to migrate farther had higher heterophil-to-lymphocyte ratios during incubation than those that did not have to travel as far, particularly for species with high wing loading.

Infectious disease, shifting climates, and opportunistic predators: cumulative factors potentially impacting wild salmon declines.

Emerging diseases are impacting animals under high-density culture, yet few studies assess their importance to wild populations. Microparasites selected for enhanced virulence in culture settings should be less successful maintaining infectivity in wild populations, as once the host dies, there are limited opportunities to infect new individuals. Instead, moderately virulent microparasites persisting for long periods across multiple environments are of greatest concern. Evolved resistance to endemic microparasites may reduce susceptibilities, but as barriers to microparasite distributions are weakened, and environments become more stressful, unexposed populations may be impacted and pathogenicity enhanced. We provide an overview of the evolutionary and ecological impacts of infectious diseases in wild salmon and suggest ways in which modern technologies can elucidate the microparasites of greatest potential import. We present four case studies that resolve microparasite impacts on adult salmon migration success, impact of river warming on microparasite replication, and infection status on susceptibility to predation. Future health of wild salmon must be considered in a holistic context that includes the cumulative or synergistic impacts of multiple stressors. These approaches will identify populations at greatest risk, critically needed to manage and potentially ameliorate the shifts in current or future trajectories of wild populations.

Evolution of embryonic developmental period in the marine bird families Alcidae and Spheniscidae: roles for nutrition and predation?

BACKGROUND: Nutrition and predation have been considered two primary agents of selection important in the evolution of avian life history traits. The relative importance of these natural selective forces in the evolution of avian embryonic developmental period (EDP) remain poorly resolved, perhaps in part because research has tended to focus on a single, high taxonomic-level group of birds: Order Passeriformes. The marine bird families Alcidae (auks) and Spheniscidae (penguins) exhibit marked variation in EDP, as well as behavioural and ecological traits ultimately linked to EDP. Therefore, auks and penguins provide a unique opportunity to assess the natural selective basis of variation in a key life-history trait at a low taxonomic-level. We used phylogenetic comparative methods to investigate the relative importance of behavioural and ecological factors related to nutrition and predation in the evolution of avian EDP. RESULTS: Three behavioural and ecological variables related to nutrition and predation risk (i.e., clutch size, activity pattern, and nesting habits) were significant predictors of residual variation in auk and penguin EDP based on models predicting EDP from egg mass. Species with larger clutch sizes, diurnal activity patterns, and open nests had significantly shorter EDPs. Further, EDP was found to be longer among birds which forage in distant offshore waters, relative to those that foraged in near shore waters, in line with our predictions, but not significantly so. CONCLUSION: Current debate has emphasized predation as the primary agent of selection driving avian life history diversification. Our results suggest that both nutrition and predation have been important selective forces in the evolution of auk and penguin EDP, and highlight the importance of considering these questions at lower taxonomic scales. We suggest that further comparative studies on lower taxonomic-level groups will continue to constructively inform the debate on evolutionary determinants of avian EDP, as well as other life history parameters.

Yolk carotenoids and stable isotopes reveal links among environment, foraging behavior and seabird breeding success.

Nutrients that are limited in availability, such as carotenoids, are potentially involved in trade-offs between homeostasis and reproduction. Despite their importance, factors that affect the capacity of female birds to meet their carotenoid requirements are poorly understood. We used delta(15)N stable isotope analysis to relate foraging behavior to yolk carotenoid deposition in two seabirds, Cassin's auklet (Ptychoramphus aleuticus) and rhinoceros auklet (Cerorhinca monocerata), during each of five years. As expected from their narrower trophic range, Cassin's auklets produced yolks with fewer carotenoid types than did rhinoceros auklets (one vs. three). Cassin's auklets also fed on a lower trophic level diet richer in carotenoids, yet had lower total yolk carotenoid levels, which suggests a role for species-specific adaptations for carotenoid uptake and utilization. Within both species, lower trophic-level feeding was linked to higher yolk carotenoid levels, but through different mechanisms. In Cassin's auklets, it was due to a population-wide response to environmental variation: in warm-water years, all females fed at a low trophic level and produced carotenoid-rich yolks. In rhinoceros auklets, it was due to individual differences similarly expressed in all years: females fed across a wide trophic range, and those that fed at a low trophic level produced carotenoid-rich yolks. Rhinoceros auklets bred more successfully in years when their yolks were rich in carotenoids, probably due to a correlated response to stronger marine primary production. Our results are novel because they link avian yolk carotenoid deposition to behavioral and environmental variations.

Stable isotopes link diet to avian yolk carotenoid allocation: a comparative study of five auk species (Charadriiformes: Alcidae).

The allocation of important but environmentally limited nutrients, such as carotenoids, often represents a trade-off between homeostasis and reproduction. However, key questions remain about how diet and species traits influence carotenoid allocation. We studied yolk carotenoid profiles and yolk color in relation to trophic level (based on delta(15)N values) in five species of seabirds belonging to the family Alcidae: common murre (Uria aalge), pigeon guillemot (Cepphus columba), Cassin's auklet (Ptychoramphus aleuticus), rhinoceros auklet (Cerorhinca monocerata), and tufted puffin (Fratercula cirrhata). In three species, which ranged from low (tufted puffin) to high (pigeon guillemot) trophic level, yolks had pink to red hues and contained exclusively astaxanthin, while yolks of species from a high trophic level (common murre) and from a generalist forager (rhinoceros auklet) had yellow to orange hues and contained astaxanthin, lutein, and zeaxanthin. The absence of a systematic relationship between trophic level and carotenoid types suggests that these species possess adaptations for the uptake and use of specific carotenoids. In contrast, total yolk carotenoid concentrations were best explained by the combination of species and trophic level: lower-trophic-level feeding was linked to production of carotenoid-rich yolks, both across species and within the one generalist species. We conclude that both behavioral and physiological traits can play strong roles in the acquisition and allocation of critical nutrients from mothers to their offspring.

Range-wide reproductive consequences of ocean climate variability for the seabird Cassin's Auklet.

We examine how ocean climate variability influences the reproductive phenology and demography of the seabird Cassin's Auklet (Ptychoramphus aleuticus) across approximately 2500 km of its breeding range in the oceanographically dynamic California Current System along the west coast of North America. Specifically, we determine the extent to which ocean climate conditions and Cassin's Auklet timing of breeding and breeding success covary across populations in British Columbia, central California, and northern Mexico over six years (2000-2005) and test whether auklet timing of breeding and breeding success are similarly related to local and large-scale ocean climate indices across populations. Local ocean foraging environments ranged from seasonally variable, high-productivity environments in the north to aseasonal, low-productivity environments to the south, but covaried similarly due to the synchronizing effects of large-scale climate processes. Auklet timing of breeding in the southern population did not covary with populations to the north and was not significantly related to local oceanographic conditions, in contrast to northern populations, where timing of breeding appears to be influenced by oceanographic cues that signal peaks in prey availability. Annual breeding success covaried similarly across populations and was consistently related to local ocean climate conditions across this system. Overall, local ocean climate indices, particularly sea surface height, better explained timing of breeding and breeding success than a large-scale climate index by better representing heterogeneity in physical processes important to auklets and their prey. The significant, consistent relationships we detected between Cassin's Auklet breeding success and ocean climate conditions across widely spaced populations indicate that Cassin's Auklets are susceptible to climate change across the California Current System, especially by the strengthening of climate processes that synchronize oceanographic conditions. Auklet populations in the northern and central regions of this ecosystem may be more sensitive to changes in the timing and variability of ocean climate conditions since they appear to time breeding to take advantage of seasonal productivity peaks.

Carry-over effects in a Pacific seabird: stable isotope evidence that pre-breeding diet quality influences reproductive success.

1. Understanding the interactions between different periods of the annual cycle in migratory animals has been constrained by our inability to track individuals across seasons. In seabirds, virtually nothing is known about how diet quality during the non-breeding period, away from the breeding grounds, might influence subsequent reproductive success. 2. We used stable nitrogen (delta(15)N) and carbon (delta(13)C) isotopes to evaluate the effects of non-breeding diet quality on the timing of breeding and egg size in a population of Cassin's auklets (Ptychoramphus aleuticus) breeding on Triangle Island, British Columbia. Adult feathers are grown during two different periods of the annual cycle, which allowed us to estimate diet quality from the previous fall (October-November) and pre-breeding (February-March) period. 3. We found that the estimated proportion of energetically superior copepods (Neocalanus spp.) in the pre-breeding diet tended to be higher in females that bred earlier and laid larger eggs, whereas energetically poor juvenile rockfish (Sebastes spp.) were dominant in the pre-breeding diets of females that bred later and laid smaller eggs. We detected no effect of fall diet quality on breeding date or egg size, and no effect of pre-breeding diet quality on breeding date in males. 4. Pre-breeding diet quality was not related to body condition measured 1-2 days after laying, which suggests that females may need to attain a threshold condition before they initiate breeding and successfully rear young. 5. Our results suggest that changes in climatic conditions during the pre-breeding period may have severe consequences for reproductive success by influencing breeding date and egg size. Our work emphasizes the importance of determining how events are linked throughout the annual cycle for understanding the fitness and population dynamics of migratory animals.

Are yolk androgens adjusted to environmental conditions? A test in two seabirds that lay single-egg clutches.

It is widely believed that female birds strategically allocate androgens to yolk in the manner that best equips offspring for feeding conditions during their development. Because most avian studies have focused on multi-egg clutch species, and interpreted results within the framework of sibling competition, we still know little about how yolk androgens might be allocated in direct response to environmental conditions. Most oceanic birds are long-lived and lay single-egg clutches, and their breeding success is tightly linked to highly variable marine production. That combination: a variable breeding environment, long lives, and single-egg clutches, makes oceanic birds good subjects to test hypotheses about yolk androgen allocation strategies. We measured concentrations of two yolk androgens, androstenedione (A4) and testosterone (T), in the single-egg clutches laid by early-laying Cassin's (Ptychoramphus aleuticus) and rhinoceros (Cerorhinca monocerata) auklets at Triangle Island, British Columbia, Canada, in 2002-2004. Environmental conditions including sea-surface temperatures and the timing and intensity of marine primary production varied over the 3 years, and in response, both the timing and success of seabird breeding varied. As in other avian species, concentrations of A4 and T varied markedly among individual eggs in both species (by factors of 3-8), yet contrary to expectation, little of the variation could be attributed to year effects. The high interindividual variation and the lack of interannual variation suggest a non-adaptive explanation for yolk androgen deposition relative to environmental conditions in these species.