Geology topics

Sarah K. Schoen

Publications and source records attributed to Sarah K. Schoen.

16 recordsLinked to original sources

USGS2025

Paralytic shellfish toxins and seabirds: Evaluating sublethal effects, behavioral responses, and ecological implications of saxitoxin ingestion by common murres (Uria aalge)

Paralytic shellfish toxins (PSTs), including saxitoxin (STX) and its congeners, are neurotoxins that can be produced during harmful algal blooms and cause illness or death in humans, fish, seabirds, and marine mammals. Since 2014, multiple large-scale seabird mortality events have occurred in Alaska waters, with STXs detected in some carcasses. To investigate the sublethal behavioral and ecological effects of STX on seabirds, we conducted captive dosing trials with common murres ( Uria aalge ). We gavaged purified STX (dehydrated STX dihydrocholoride, STX-diHCl) or an Alexandrium catenella culture extract into murres, monitored behavioral responses and recovery times, and assessed tissue concentrations in individuals that died or were euthanized. Using a modified up-and-down dose-finding scheme, we estimated a median effective dose (ED 50 ) of 89 µg STX-equivalents (eq) kg -1 for STX-diHCl and 366 µg STX-eq kg -1 for the A. catenella extract based on ecologically relevant behavior. Differences between the ED 50 estimates could reflect uncertainties in toxin equivalency factors for PST congeners, which are based on studies using purified toxins in mice and may vary across taxa or toxin matrices. Post-dosing concentrations of STX varied by tissue type across individuals, with quantifiable levels ranging from 3 to 379 µg STX-eq 100g -1 . Evidence of biotransformation of STX in A. catenella extract-dosed birds was observed. We also measured the chronic effects of dosing with sublethal levels of STX-diHCl over seven-days, which resulted in lower fish intake among treatment birds compared to controls (-187 g day -1 ). This investigation improves our understanding of the ecological effects of PSTs on seabird health.

Alaska↗

USGS2024

Lingering impacts of the 2014-2016 northeast Pacific marine heatwave on seabird demography in Cook Inlet, Alaska (USA)

A protracted period (2014-2016) of anomalously warm water in the northeast Pacific Ocean precipitated an extensive die-off of common murres Uria aalge (hereafter ‘murres’) during 2015-2016, accompanied by reduced colony attendance and reproductive success of murres and black-legged kittiwakes Rissa tridactyla (‘kittiwakes’) starting in 2015. Most murres died of starvation following a large-scale reduction in abundance and quality of forage fish. To assess murre and kittiwake recovery following the marine heatwave, we monitored their demographics at 2 colonies (Chisik and Gull Islands) in Cook Inlet, Alaska (USA), from 2016 to 2019. Compared to historic data (1995-1999), we observed declines and increased variability in colony attendance and productivity across species and colonies, and predation was widespread. At Chisik, where food limitations were common during historic studies, both species experienced substantial population declines and reproductive failures in all 4 years (2016-2019) following the heatwave. At Gull, a typically productive colony during historic studies, murres failed to fledge chicks for 3 years (2016-2018) following the heatwave. By 2019, murre productivity recovered to about half that observed during historic studies (0.28 vs. 0.54 chicks per pair), but populations had declined by half. Kittiwake population size at Gull declined a quarter from historic counts, and reproduction alternated between complete breeding failures (2016/2018) and high productivity (2017/2019). These multi-year demographic impacts indicate lingering effects of the heatwave on kittiwakes and murres through forage fish depletion and increased predator disturbance, and possibly other stressors. It remains unknown whether populations can rebound to historic levels. If so, recovery would likely take decades.

Alaska↗

USGS2024

Predator disturbance contributed to Common Murre Uria aalge breeding failures in Cook Inlet, Alaska following the 2014–2016 Pacific marine heatwave

The 2014-2016 Pacific marine heatwave caused unprecedented die-offs and multi-year reproductive failures for Common Murres Uria aalge along the west coast of North America. Lingering impacts, such as declines in colony attendance and productivity, have persisted at some colonies following the heatwave and are attributed largely to changes in prey availability and quality. Here, we present evidence of an additional, top-down mechanism contributing to Common Murre breeding failures on Gull Island (Alaska): disturbance of nesting birds by aerial predators and associated egg depredation. We collected time-lapse images over five murre breeding seasons (2016-2020) on Gull Island to document the frequency, duration, and intensity of disturbances caused by aerial predators, as well as to quantify disturbance-associated egg depredation. To identify seasonal and inter-annual variability of disturbances, we calculated a daily disturbance index and compared years using generalized additive models. In all years, Bald Eagles Haliaeetus leucocephalus were the primary cause of disturbance, which led to periods of prolonged colony abandonment by murres and facilitated high levels of murre egg depredation by Glaucous-winged Gulls Larus glaucescens and Herring Gulls L. argentatus . We found that the seasonality of disturbance was an important factor in determining egg depredation rates. In years when disturbance levels were high and persisted later in the season, the colony experienced complete breeding failures due to disturbance-associated egg depredation. Our study revealed that the response of nesting murres to a strong environmental perturbation, such as the Pacific marine heatwave, can be complex and involve multiple stressors from both bottom-up and top-down factors.

Alaska↗

USGS2023

Joint spatiotemporal models to predict seabird densities at sea

Introduction: Seabirds are abundant, conspicuous members of marine ecosystems worldwide. Synthesis of distribution data compiled over time is required to address regional management issues and understand ecosystem change. Major challenges when estimating seabird densities at sea arise from variability in dispersion of the birds, sampling effort over time and space, and differences in bird detection rates associated with survey vessel type. Methods: Using a novel approach for modeling seabirds at sea, we applied joint dynamic species distribution models (JDSDM) with a vector-autoregressive spatiotemporal framework to survey data collected over nearly five decades and archived in the North Pacific Pelagic Seabird Database. We produced monthly gridded density predictions and abundance estimates for 8 species groups (77% of all birds observed) within Cook Inlet, Alaska. JDSDMs included habitat covariates to inform density predictions in unsampled areas and accounted for changes in observed densities due to differing survey methods and decadal-scale variation in ocean conditions. Results: The best fit model provided a high level of explanatory power (86% of deviance explained). Abundance estimates were reasonably precise, and consistent with limited historical studies. Modeled densities identified seasonal variability in abundance with peak numbers of all species groups in July or August. Seabirds were largely absent from the study region in either fall (e.g., murrelets) or spring (e.g., puffins) months, or both periods (shearwaters). Discussion: Our results indicated that pelagic shearwaters ( Ardenna spp.) and tufted puffin ( Fratercula cirrhata ) have declined over the past four decades and these taxa warrant further investigation into underlying mechanisms explaining these trends. JDSDMs provide a useful tool to estimate seabird distribution and seasonal trends that will facilitate risk assessments and planning in areas affected by human activities such as oil and gas development, shipping, and offshore wind and renewable energy.

Alaska↗

USGS2022

Kittlitz’s murrelet seasonal distribution and post-breeding migration from the Gulf of Alaska to the Arctic Ocean

Kittlitz’s Murrelets ( Brachyramphus brevirostris ) nest during summer in glaciated or recently deglaciated (post-Wisconsin) landscapes. They forage in adjacent marine waters, especially those influenced by glacial meltwater. Little is known of their movements and distribution outside the breeding season. To identify post-breeding migrations of murrelets, we attached satellite transmitters to birds (n = 47) captured at sea in the Gulf of Alaska and Aleutian Islands during May – July 2009 – 15 and tracked 27 birds that migrated from capture areas. Post-breeding murrelets migrated toward the Bering Sea, with short periods of movement (median 2 d) separated by short stopovers (median 1 d). Travel speeds averaged 79.4 km d-1 (83.5 SD, 449.1 maximum). Five Kittlitz’s Murrelets tagged in Prince William Sound in May migrated to the Bering Sea by August and four continued north to the Arctic Ocean, logging 2500 – 4000 km of travel. Many birds spent 2‒3 weeks with little movement along coasts of the Alaska Peninsula or eastern Bering Sea during late August through September, also the pre-basic molt period. Ship-based surveys, many of which were conducted concurrently with our telemetry studies, confirmed that substantial numbers of Kittlitz’s Murrelets migrate into the Arctic Ocean during autumn. They also revealed that some birds spend winter and spring in the Bering Sea in association with ice-edge, polynya, or marginal ice zone habitats before returning to summer breeding grounds. We conclude that this species is best characterized as a sub-Arctic and Arctic species, which has implications for future risk assessments and threat mitigation.

Alaska↗

USGS2022

Paralytic shellfish toxins associated with Arctic Tern mortalities in Alaska

Harmful algal blooms produce biotoxins that can injure or kill fish, wildlife, and humans. These blooms occur naturally but have intensified in many locations globally due to recent climatic changes, including ocean warming. Such changes are especially pronounced in northern regions, where the effects of paralytic shellfish toxins (PSTs) on marine wildlife are of growing concern. In Alaska, seabird mortality events have increased in frequency, magnitude, and duration since 2015 alongside anomalously high ocean temperatures. Although starvation has been implicated as the apparent cause of death in many of these die-offs, saxitoxin (STX) and other PSTs have been identified as possible contributing factors. Here, we describe a mortality event at a nesting colony of Arctic Terns ( Sterna paradisaea ) near Juneau, Alaska in 2019 and report elevated concentrations of PSTs in bird, forage fish, and mussel samples. Concentrations of STX and other PSTs in tern tissues (2.5–51.2 µg 100g −1 STX-equivalents [STX-eq]) were of similar magnitude to those reported from other PST-induced bird die-offs. We documented high PST concentrations in blue mussels (>11,000 µg 100g −1 STX-eq; Mytilus edulis spp.) collected from nearby beaches, as well as in forage fish (up to 494 µg 100g −1 STX-eq) retrieved from Arctic Tern nests, thereby providing direct evidence of PST exposure via the terns’ prey. At maximum concentrations measured in this study, a single 5 g Pacific Sand Lance ( Ammodytes personatus ) could exceed the median lethal STX dose (LD 50 ) currently estimated for birds, offering strong support for PSTs as a likely source of tern mortality. In addition to describing this localized bird mortality event, we used existing energetics data from adult and nestling Arctic Terns to calculate estimated cumulative daily PST exposure based on ecologically relevant concentrations in forage fish. Our estimates revealed potentially lethal levels of PST exposure even at relatively low (≤30 ug 100g −1 STX-eq) toxin concentrations in prey. These findings suggest that PSTs present a significant hazard to Arctic Terns and other northern seabirds and should be included in future investigations of avian mortality events as well as assessments of population health.

Alaska↗

USGS2022

Melanism in a Common Murre Uria aalge in Kachemak Bay, Alaska

In accord with melanism being uncommon in birds, we could find only six published records of completely melanistic Common Murres Uria aalge , one of the most widely and intensively studied of all seabirds. We added to the record by observing a Common Murre in completely dark, melanistic alternate plumage every summer from 2017 to 2021 at Gull Island in Kachemak Bay, Alaska, USA. In 2017, the bird frequented the colony periphery, indicating that it could have been a subadult. Subsequently, it occupied the same narrow rock ledge within the colony every summer from 2018 to 2021, an indication that it may have been attempting to breed. Because we have been conducting long-term monitoring on Gull Island, we are in the unique position to be able to monitor the attendance and reproductive performance of this distinctively marked murre into the future.

Alaska↗

USGS2021

Heatwave-induced synchrony within forage fish portfolio disrupts energy flow to top pelagic predators

During the Pacific marine heatwave of 2014–2016, abundance and quality of several key forage fish species in the Gulf of Alaska were simultaneously reduced throughout the system. Capelin ( Mallotus catervarius ), sand lance ( Ammodytes personatus ), and herring ( Clupea pallasii ) populations were at historically low levels, and within this community abrupt declines in portfolio effects identify trophic instability at the onset of the heatwave. Although compensatory changes in age‐structure, size, growth or energy content of forage fish were observed to varying degrees among all these forage fish, none were able to fully mitigate adverse impacts of the heatwave, which likely included both top‐down and bottom‐up forcing. Notably, changes to the demographic structure of forage fish suggested size‐selective removals typical of top‐down regulation. At the same time, zooplankton community structure may have driven bottom‐up regulation as copepod community structure shifted towards smaller, warm‐water species, and euphausiid biomass was reduced owing to the loss of cold‐water species. Mediated by these impacts on the forage fish community, an unprecedented disruption of the normal pelagic food web was signaled by higher trophic level disruptions during 2015–2016, when seabirds, marine mammals, and groundfish experienced shifts in distribution, mass mortalities, and reproductive failures. Unlike decadal‐scale variability underlying ecosystem regime shifts, the heatwave appeared to temporarily overwhelm the ability of the forage fish community to buffer against changes imposed by warm water anomalies, thereby eliminating any ecological advantages that may have accrued from having a suite of coexisting forage species with differing life history compensations.

Alaska↗

USGS2021

Reduced quality and synchronous collapse of forage species disrupts trophic transfer during a prolonged marine heatwave

The Gulf of Alaska forage fish community includes a few key species that differ markedly in their timing of spawning, somatic growth and lipid storage, and in their migration behavior. This diversity in life history strategies facilitates resilience in marine food webs because it buffers predators against the naturally high variance in abundance of pelagic forage fish populations by decreasing the likelihood that all species will be scarce at the same time. During the prolonged North Pacific marine heatwave of 2014-2016, the availability and quality of at least three key forage species with different life history strategies were reduced simultaneously in the system. Capelin and sand lance occurrence in predator diets declined abruptly, and Prince William Sound herring spawning biomass declined to historically low levels. Biomass of euphausiids was also reduced, in part due to the loss of a cold-water species. Changes in age structure, growth, and energy content of capelin, sand lance, and herring were also associated with warming during the heatwave, but not all species responded in the same way. For example, spawning capelin grew faster and matured at a younger age but were shorter in length than usual, while sand lance in Prince William Sound experienced anomalously low growth rates and lipid storage in 2015-2016. Changes in forage fish populations had immediate impacts on predator populations in 2015-2016, when seabirds and marine mammals experienced shifts in distribution, mass mortality, and reproductive failures in the Gulf of Alaska. In contrast, copepod abundance increased on the shelf and in some coastal regions during the heatwave, suggesting that food availability was not a primary factor limiting forage fish populations at this time. The reduced quality and collapse of multiple forage fish populations reduced the efficiency of energy transfer through the middle trophic level of pelagic food webs, disrupting energy flow to piscivorous pelagic predators and causing abrupt and extreme reductions in their numbers and productivity.

Alaska↗

USGS2021

Monitoring the recovery of seabirds and forage fish following a major ecosystem disruption in Lower Cook Inlet

Following the North Pacific marine heatwave of 2014-2016 and associated extreme die-off of seabirds in the winter of 2015-2016, we repeated historical marine bird and forage fish surveys around two seabird colonies (Gull Island, Chisik Island) in lower Cook Inlet during 2016-2019 in order to document immediate and lingering impacts of the heatwave on resident fish and seabird populations. At sea, we conducted acoustic/trawl surveys for fish in near-colony areas, while simultaneously counting seabirds on survey transects. At colonies, we monitored bird numbers on whole colony censuses and population plot counts and assessed annual reproductive success of Black-legged Kittiwakes ( Rissa tridactyla ) and Common Murres ( Uria aalge ). Our studies suggest that impacts of the heatwave on seabirds and their forage base were severe and extended for several years beyond the heatwave itself. Although fish biomass indices were greatest in 2016-2017, the forage community was largely dominated by juvenile fish with low nutritional value following a collapse of key species of forage fish during the marine heatwave. Lowest fish biomass was observed in 2018, coincident with a marked decline in the at-sea abundance of many seabirds around colonies. By 2019 there were signs of improvement in prey abundance and quality, and significant aggregations of pre-spawning capelin and large sand lance were observed in and around Kachemak Bay. In comparison with historical (1996-1999) bird colony surveys, kittiwake population counts decreased by more than 70% at Chisik Island, and by 8-29% at Gull Island. Similarly, murre counts decreased at Chisik Island by 72% and by more than 20% at Gull Island. Kittiwakes had complete reproductive failure at Chisik Island from 2016-2018, with minimal success in 2019. At Gull Island, kittiwakes also failed in 2016 and 2018 but had unusually high productivity in 2017 and 2019. Murres also had complete reproductive failures at Chisik Island in all four years and at Gull Island during 2016, 2017 and 2018. Murres finally fledged chicks in 2019, albeit at about half the normal rate. Finally, we also observed unusually high levels of predator disturbance (egg predation, flushing adults from plots) and unprecedented numbers of adult birds in poor body condition (“skinny murres”), particularly in 2018. Together, these results indicate that an extreme disruption of marine food webs occurred during and after the heatwave. Given the prolonged reproductive failures, continued monitoring is required to assess continuing impacts and recovery from the 2014-2016 heatwave. For example, murre chicks that would have been produced in 2016 should, after 4-5 years away, start recruiting to the colony in 2021, and thereby replace aged adults that would have died naturally in the past year(s). Without such natal recruitment for at least the next 3-4 years, we should document a continuing decline in bird numbers at the colonies.

Alaska↗

USGS2021

Another bad year for seabirds on Gull Island

The USGS monitored Black-legged Kittiwake and Common Murre populations on Gull Island from 1995-1999, and from 2016-2021, following the 2014-2016 northeast Pacific marine heatwave. USGS research, which is supported by the USGS and BOEM, includes population monitoring, and quantifying breeding success and predator disturbances. This information is preliminary or provisional and is subject to revision. Learn more: https://www.usgs.gov/centers/alaska-science-center/science/cook-inlet-seabird-and-forage-fish-study

Alaska↗

USGS2020

Extreme mortality and reproductive failure of common murres resulting from the northeast Pacific marine heatwave of 2014-2016

About 62,000 dead or dying common murres ( Uria aalge ), the trophically dominant fish-eating seabird of the North Pacific, washed ashore between summer 2015 and spring 2016 on beaches from California to Alaska. Most birds were severely emaciated and, so far, no evidence for anything other than starvation was found to explain this mass mortality. Three-quarters of murres were found in the Gulf of Alaska and the remainder along the West Coast. Studies show that only a fraction of birds that die at sea typically wash ashore, and we estimate that total mortality approached 1 million birds. About two-thirds of murres killed were adults, a substantial blow to breeding populations. Additionally, 22 complete reproductive failures were observed at multiple colonies region-wide during (2015) and after (2016–2017) the mass mortality event. Die-offs and breeding failures occur sporadically in murres, but the magnitude, duration and spatial extent of this die-off, associated with multi-colony and multi-year reproductive failures, is unprecedented and astonishing. These events co-occurred with the most powerful marine heatwave on record that persisted through 2014–2016 and created an enormous volume of ocean water (the “Blob”) from California to Alaska with temperatures that exceeded average by 2–3 standard deviations. Other studies indicate that this prolonged heatwave reduced phytoplankton biomass and restructured zooplankton communities in favor of lower-calorie species, while it simultaneously increased metabolically driven food demands of ectothermic forage fish. In response, forage fish quality and quantity diminished. Similarly, large ectothermic groundfish were thought to have increased their demand for forage fish, resulting in greater top-predator demands for diminished forage fish resources. We hypothesize that these bottom-up and top-down forces created an “ectothermic vise” on forage species leading to their system-wide scarcity and resulting in mass mortality of murres and many other fish, bird and mammal species in the region during 2014–2017.

PLoS ONE↗

USGS2020

Algal toxins in Alaskan seabirds: Evaluating the role of saxitoxin and domoic acid in a large-scale die-off of Common Murres

Elevated seawater temperatures are linked to the development of harmful algal blooms (HABs), which pose a growing threat to marine birds and other wildlife. During late 2015 and early 2016, a massive die-off of Common Murres (Uria algae; hereafter, murres) was observed in the Gulf of Alaska coincident with a strong marine heat wave. Previous studies have documented illness and death among seabirds resulting from exposure to the HAB neurotoxins saxitoxin (STX) and domoic acid (DA). Given the unusual mortality event, corresponding warm water anomalies, and recent detection of STX and DA throughout coastal Alaskan waters, HABs were identified as a possible factor of concern. To evaluate whether algal toxins may have contributed to murre deaths, we tested for STX and DA in a suite of tissues obtained from beach-cast murre carcasses associated with the die-off as well as from apparently healthy murres and Black-legged Kittiwakes (Rissa tridactyla; hereafter, kittiwakes) in the preceding and following summers. We also tested forage fish and marine invertebrates collected in the Gulf of Alaska in 2015–2017 to evaluate potential sources of HAB toxin exposure for seabirds. Saxitoxin was present in multiple tissue types of both die-off (36.4%) and healthy (41.7%) murres and healthy kittiwakes (54.2%). Among birds, we detected the highest concentrations of STX in liver tissues (range 1.4 –10.8 µg 100 g-1) of die-off murres. Saxitoxin was relatively common in forage fish (20.3%) and marine invertebrates (53.8%). No established toxicity limits currently exist for seabirds, but concentrations of STX in birds and forage fish in our study were lower than values reported from most other bird die-offs in which STX intoxication was causally linked. We detected low concentrations of DA in a single bird sample and in 33.3% of marine invertebrates and 4.0% of forage fish samples. Although these results do not support the hypothesis that acute exposure to STX or DA was a primary factor in the 2015–2016 die-off event, additional information about the sensitivity of murres to these toxins is needed before we can discount their potential role in the die-off. The widespread occurrence of STX in seabirds, forage fish, and marine invertebrates in the Gulf of Alaska indicates that algal toxins should be considered in future assessments of seabird health, especially given the potential for greater occurrence of HABs in the future.

Alaska↗

USGS2020

Seabird synthesis

Overall, the status of seabirds was fair to good in the WGOA in 2020, with limited data available from Middleton Island, Cook Inlet, and the Kodiak Archipelago (Figure 63). Colony attendance remains low in some populations compared to historic levels, and some colonies were newly abandoned. However, when birds did arrive to breed, reproductive success generally appeared fair to good for fish-eating, surface-feeding birds and fish-eating, diving birds. There was spatial variability in colony attendance and reproductive success, with Middleton Island birds performing more strongly than Kodiak Island or Cook Inlet. Middleton Island populations from both these groups experienced their strongest breeding seasons since the marine heatwave began in 2014, suggesting an increase in the availability of small schooling fish in that region of WGOA. No large-scale mortality event was recorded based on monthly beach surveys in the WGOA. This year’s integrated approach to reporting seabird status is less comparable to previous Ecosystem Status Reports, as the Alaska Maritime National Wildlife Refuge’s seabird reproductive success time series were not updated, due to COVID-19 related survey cancellations.

Alaska↗

USGS2019

Extreme reduction in nutritional value of a key forage fish during the Pacific marine heatwave of 2014–2016

Pacific sand lance Ammodytes personatus are a key forage fish in the North Pacific for many species of salmon, groundfish, seabirds, and marine mammals and have historically been important to predators in relatively warm years. However, extreme declines in the nutritional value of sand lance in Prince William Sound, Alaska, USA, during 2012-2016 indicate that energy transfer from lower trophic levels to predators via sand lance may have been disrupted during the North Pacific marine heatwave in 2015 and 2016. Nutritional value (length, energy density, and whole-body energy) was measured in age-0 and age-1 sand lance collected during July in cool (2012-2013) and increasingly warm (2014-2016) years. The value of age-0 fish was relatively stable, with only minor differences among years for length and whole-body energy. By contrast, the value of age-1 fish significantly declined in 2015, and by 2016 they were 38% shorter and 13% lower in energy density compared to cooler years. This contributed to significant declines in whole-body energy of 44% in 2015 and 89% in 2016 compared to cooler years (2012-2014). The 2015 sand lance cohort experienced little growth or lipid accumulation from July 2015 at age-0 to July 2016 at age-1. This effective disruption of energy flow through pelagic food webs probably contributed to population declines and/or breeding failures observed among several predators in the Gulf of Alaska and suggests that tipping points were reached during the heatwave.

Marine Ecology Progress Series↗

USGS2018

Avian predator buffers against variability in marine habitats with flexible foraging behavior

How well seabirds compensate for variability in prey abundance and composition near their breeding colonies influences their distribution and reproductive success. We used tufted puffins ( Fratercula cirrhata ) as forage fish samplers to study marine food webs from the western Aleutian Islands (53°N, 173°E) to Kodiak Island (57°N, 153°W), Alaska, during August 2012–2014. Around each colony we obtained data on: environmental characteristics (sea surface temperature and salinity, seafloor depth and slope, tidal range, and chlorophyll- a ), relative forage fish biomass (hydroacoustic backscatter), and seabird community composition and density at-sea. On colonies, we collected puffin chick-meals to characterize forage communities and determine meal energy density, and measured chicks to obtain a body condition index. There were distinct environmental gradients from west to east, and environmental variables differed by ecoregions: the (1) Western-Central Aleutians, (2) Eastern Aleutians, and, (3) Alaska Peninsula. Forage fish biomass, species richness, and community composition all differed markedly between ecoregions. Forage biomass was strongly correlated with environmental gradients, and environmental gradients and forage biomass accounted for ~ 50% of the variability in at-sea density of tufted puffins and all seabird taxa combined. Despite the local and regional variability in marine environments and forage, the mean biomass of prey delivered to puffin chicks did not differ significantly between ecoregions, nor did chick condition or puffin density at-sea. We conclude that puffins can adjust their foraging behavior to produce healthy chicks across a wide range of environmental conditions. This extraordinary flexibility enables their overall success and wide distribution across the North Pacific Ocean.

Aleutian Islands, Gulf of Alaska↗