Research Publications

Penguins: Natural History and Conservation
Edited by Dr. Pablo Garcia-Borboroglu and Dr. Dee Boersma

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Selected Publications

Changing course: Relocating commercial tanker lanes significantly reduces threat of chronic oiling for a top marine predator (2023)

Eric L. Wagner, Esteban Frere, P. Dee Boersma

A goal for conservation biologists is to show that policies enacted on behalf of an imperiled species results in direct benefits for it. In Argentina, tens of thousands of Magellanic penguins (Spheniscus magellanicus) were estimated to have died from chronic oil pollution each year through the early 1980s. From 1982 to 1990, surveys
at sites along approximately 900 km of Chubut Province coastline found that >60 % of penguin carcasses had evidence of oiling in some years. In response to these findings, as well as pressure from non-governmental organizations and the public, provincial and federal authorities in Chubut moved the commercial tanker lanes 20 nautical miles farther offshore in 1997 and required oil tankers to have double hulls. During a second round of surveys in 2001, using most of the same sites as the first survey period, the number of dead and oiled penguins dropped effectively to zero. A policy change not only led to fewer oiled penguins, but also likely increased the survival of adult Magellanic penguins near some of their most significant breeding colonies in Argentina.

Climate change as a global amplifier of human–wildlife conflict (2023)

Briana Abrahms, Neil H. Carter, T. J. Clark-Wolf, Kaitlyn M. Gaynor, Erik Johansson, Alex McInturff, Anna C. Nisi, Kasim Rafiq & Leigh West

Climate change and human–wildlife conflict are both pressing challenges for biodiversity conservation and human well-being in the Anthropocene. Climate change is a critical yet underappreciated amplifier of human– wildlife conflict, as it exacerbates resource scarcity, alters human and animal behaviours and distributions, and increases human–wildlife encounters. We synthesize evidence of climate-driven conflicts occurring among ten taxonomic orders, on six continents and in all five oceans. Such conflicts disrupt both subsistence livelihoods and industrial economies and may accelerate the rate at which human–wildlife conflict drives wildlife declines. We introduce a framework describing distinct environmental, ecological and sociopolitical pathways through which climate variability and change percolate via complex social–ecological systems to influence patterns and outcomes of human–wildlife interactions. Identifying these pathways allows for developing mitigation strategies and proactive policies to limit the impacts of human–wildlife conflict on biodiversity conservation and human well-being in a changing climate.

Divergent foraging strategies between populations of sympatric matrilineal killer whales (2023)

Jennifer B Tennessen, Marla M Holt, Brianna M Wright, M Bradley Hanson, Candice K Emmons, Deborah A Giles, Jeffrey T Hogan, Sheila J Thornton, Volker B Deecke

In cooperative species, human-induced rapid environmental change may threaten cost–benefit tradeoffs of group behavioral strategies that evolved in past environments. Capacity for behavioral flexibility can increase population viability in novel environments. Whether the partitioning of individual responsibilities within social groups is fixed or flexible across populations is poorly understood, despite its relevance for predicting responses to global change at the population and species levels and designing successful conservation programs. We leveraged bio-logging data from two populations of fish-eating killer whales (Orcinus orca) to quantify patterns of fine-scale foraging movements and their relationships with demography. We reveal striking interpopulation differences in patterns of individual foraging behavior. Females from the endangered Southern Resident (SRKW) population captured less prey and spent less time pursuing prey than SRKW males or Northern Resident (NRKW) females, whereas NRKW females captured more prey than NRKW males. The presence of a calf (≤3 years) reduced the number of prey captured by adult females from both populations, but disproportionately so for SRKW. SRKW adult males with a living mother captured more prey than those whose mother had died, whereas the opposite was true for NRKW adult males. Across populations, males foraged in deeper areas than females, and SRKW captured prey deeper than NRKW. These population-level differences in patterns of individual foraging behavior challenge the existing paradigm that females are the disproportionate foragers in gregarious resident killer whales, and demonstrate considerable variation in the foraging strategies across populations of an apex marine predator experiencing different environmental stressors.

Additional Publications


  • Changing course: Relocating commercial tanker lanes significantly reduces threat of chronic oiling for a top marine predator. Marine Pollution Bulletin.
  • Chasing inter-species communication: what marine mammals are telling us about our oceans. ICES Journal of Marine Science, 2023. Open Access.
  • Climate Change as a Global Amplifier of Human-Wildlife Conflict. Nature Climate Change, 2023. Open Access.
    Coverage: KUOW, NPR, The Guardian, Newsweek, Scientific American
  • Climate Presses and Pulses Mediate the Decline of a Migratory Predator. Proceedings of the National Academy of Sciences. Open Access.
  • Divergent foraging strategies between populations of sympatric matrilineal killer whales. Behavioral Ecology, 2023. Open Access.
    Coverage: The Seattle Times, Skagit Valley Herald, Seattle King 5 News, KUOW Public Radio, Oregon Public Broadcasting’s “Think Out Loud”, Victoria Times Colonist, North Shore News
  • Resilience to a severe marine heatwave at two Pacific seabird colonies. Marine Ecology Progress Series, 2023. Open Access.
  • Retrospective analysis of conservation measures to reduce the risk of large whale entanglements in a highly lucrative fishery. Biological Conservation, 2023. Open Access.


  • Acoustic signature reveals blue whales tune life-history transitions to oceanographic conditions. Functional Ecology, 2022.
  • Long-term, climate-driven phenological shift in a tropical large carnivore. Proceedings of the National Academy of Sciences, 2022. Open Access.
  • Should I stay or should I go: factors influencing mate retention and divorce in a colonial seabird. Animal Behaviour, 2022.
    Coverage: New Scientist
  • Site fidelity as a maladaptive behavior in the Anthropocene. Frontiers in Ecology and the Environment, 2022. Open Access.
  • Site fidelity increases reproductive success by increasing foraging efficiency in a marine predator. Behavioral Eoclogy, 2022.
  • Unprecedented heat mortality of Magellanic Penguins. Ornithological Applications, 2022. Open Access.
  • Variability in Foraging Range and Direction Among Colonies in a Widespread Seabird, the Magellanic Penguin. Frontiers in Marine Science, 2022. Open Access.


  • Dynamic strategies offer potential to reduce lethal ship collisions with large whales under changing climate conditions. Marine Policy, 2021. Open Access.


  • Consequences of phenological shifts and a compressed breeding period in Magellanic penguins. Ecology, 2021. Open Access.
  • The power of penguins: Where tourists travel to see penguins in the wild. Ocean & Coastal Management, 2020. Open Access.


  • Applying Science to Pressing Conservation Needs for Penguins. Conservation Biology, 2019. Open Access.
  • Important At-Sea Areas of Colonial Breeding Marine Predators on the Southern Patagonian Shelf. Nature, 2019.
  • Applying Science to Pressing Conservation Needs for Penguins. Conservation Biology, 2019.
  • Happy Feet in a Hostile World? The Future of Penguins Depends on Proactive Management of Current and Expected Threats. Frontiers, 2019.
  • Lateralization (handedness) in Magellanic penguins. PeerJ, 2019.
  • Coverage: Hakai Magazine
  • Food allocation and feeding behaviours of Magellanic penguin, Spheniscus magellanicus, adults and chicks. Animal Behaviour, 2019.
  • Coverage: UW News
  • Sex-biased survival contributes to population decline in long-lived seabird, the Magellanic Penguin. Ecological Applications, 2019.
  • Coverage:UW News


  • Oceanographic conditions in wintering grounds affect date and body condition in breeding female Magellanic penguins. Marine Ecology Progress Series, 2018.
  • Coverage: UW News
  • Effects of El Nino and La Nina Southern Oscillation events on the adrenocortical responses to stress in birds of the Galapagos Islands. General and Comparitive Endocrinology, 2018.
  • Framework for mapping key areas for marine megafauna to inform Marine spatial Planning: The Falkland Islands case study. Marine Policy, 2018.
  • Sexing Galapos penguins Spheniscus mendiculus by morphological measurements. Endangered Species Research, 2018.
  • Coverage: UW News
  • Spatial scales of marine conservation management for breeding seabirds. Marine Policy, 2018.


  • Comparing reproductive success of a colonial seabird , the Magellanic penguin, estimated by course- and fine-scale temporal sampling. The Condor, 2017.
  • Telomeres shorten and then lengthen before fledging in Magellanic penguins (Spheniscus magellanicus). Aging, 2017.
  • First observations of post-fledging care in Galápagos penguins (Spheniscus mendiculus). The Wilson Journal of Ornithology, 2017.
  • The strong connection between forage fish and their predators: A response to Hillborn et al. (2017). Fisheries Research, 2017.


  • Natural Selection on morphology varies among years and by sex in Magellanic Penguins (Spheniscus magellanicus). The Auk, 2016.


  • Population regulation on Magellanic penguins: What determines changes in colony size? PLOS One, 2015.
  • Marine protection is needed for Magellanic penguins in Argentina based on long-term data. Biological Conservation, 2015.
  • Coverage: Argos
  • Changes in habitat use and nesting density in a declining seabird colony. Population Ecology, 2015.


  • Pollution, habitat loss, fishing, and climate change as a critical threats to penguins. Conservation Biology, 2014.
  • Winter migration of Magellanic penguins (Spheniscus magellanicus) along Southeastern Pacific. Waterbirds, 2014.
  • Predator Responses to the Exploitation of Forage Fish: The PREP Equation. Conservation Letters, 2014.
  • Conservation of Migratory Magellanic Penguins Requires Marine Zoning. Biological Conservation, 2014.
  • Climate Change Increases Reproductive Failure in Magellanic Penguins. PLOS One, 2014.
  • The Glucocorticoid stress response in Magellanic penguins: comparing within and between breeding seasons, by age and colony, after fighting, and with other penguin species. Canadian Journal of Zoology, 2014.


  • Beach attendance in Magellanic penguins (Spheniscus magellanicus). Ardeola, 2013.
  • Parental behavior controls incubation period and asynchrony of hatching in Magellanic Penguins: Reply to Demongin, Poisbleau, and Eens (2013). The Condor, 2013.


  • Research priorities for seabirds: improving conservation and management in the 21st century. Endangered Species Research, 2012.
  • The global contribution of forage fish to marine fisheries and ecosystems. Fish and Fisheries, 2012.
  • Occurrence of Magellenic penguins along the Brazilian Northeast coast during the 2008 austral winter. The Scientific World Journal, 2012.
  • Avian pox in Magellanic penguins (Spheniscus magellanicus). Journal of Wildlife Disease, 2012.
  • Mercury levels in feathers of Magellanic penguins. Marine Pollution Bulletin, 2012.
  • Patterns of acceptance of artificial eggs and chicks by Magellanic penguins (Spheniscus magellanicus). Journal of Ornithology, 2012.
  • MHC Diversity and Mate Choice in the Magellanic Penguin, Spheniscus magellanicus. Journal of Heredity, 2012.
  • Penguins and Petroleum: Lessons in Conservation Ecology. Frontiers in Ecology and the Environment, 2012.


  • Parental behavior controls incubation period and asynchrony of hatching in Magellanic penguins. The Condor, 2011.
  • Effects of predation risk on the nocturnal activity budgets of thin-billed prions Pachyptila belcheri on New Island, Falkland Islands. Polar Biology, 2011.
  • Effects of fisheries on seabird community ecology. Reviews in Fisheries Science, 2011.


  • Calculating egg volume when shape differs: when are equations appropriate. Journal of Field Ornithology, 2010.
  • Effects of double bands on Magellanic penguins. Journal of Field Ornithology, 2010.
  • Feather-loss disorder in the African and Magellanic penguins. Waterbirds, 2010.
  • Magellanic penguin mortality in 2008 along the SW Atlantic coast. Marine Pollution Bulletin, 2010.


  • Flipper bands do not affect foraging-trip duration of Magellanic penguins. Journal of Field Ornithology, 2009.
  • Foraging distance affects reproductive success in Magellanic penguins. Marine Ecology Progress Series, 2009.
  • Intraclutch egg-size dimorphism in Magellanic penguins (Spheniscus magellanicus): adaptation, constraint, or noise? The Auk, 2009.
  • Following the fish: penguins and productivity in the South Atlantic. Ecological Monographs, 2009.
  • Magellanic penguin eggshell pores: does number matter? British Ornithologists’ Union, 2009.
  • Marine reserve in Chile would benefit penguins and ecotourism. Ocean and Coastal Management, 2009.
  • Regional genetic structure in the Magellanic penguin (Spheniscus magellanicus) suggests metapopulation dynamics. The Auk, 2009.


  • Penguins as Marine Sentinels. BioScience, 2008.
  • Tourism and Magellanic Penguins (Spheniscus magellanicus): An Example of Applying Field Endocrinology to Conservation Problems. Neotropical, 2008.


  • Oceans apart: conservation models for two temperate penguin species shaped by the marine environment. Marine Ecological Progress Series, 2007.
  • Anchovy fisheries threat to Patagonia ecosystem. Science, 2007.
  • Humboldt penguins (Spheniscus Humboldti) in the Northern Hemisphere. The Wilson Journal of Ornithology, 2007.
  • Egg size and parental quality in thin-billed prions, Pachyptila belcheri: effects on offspring fitness. Animal Behaviour, 2007.
  • Modeling the effect of El Niño on the persistence of small populations: the Galápagos penguin as a case study. Biological Conservation, 2007.


  • Chronic oil pollution harms Magellanic penguins in the Southwest Atlantic. Marine Pollution Bulletin, 2006.
  • Habituation of adult Magellanic penguins to human visitation as expressed through behavior and corticosterone secretion. Conservation Biology, 2006.
  • Agonistic interaction in wild male Magellanic penguins: when and how do they interact? Journal of Ethology, 2006.
  • Southern elephant seal (Mirounga leonina) kills Magellanic penguins (Spheniscus magellanicus) on land. Marine Mammal Science, 2006.
  • Flipper Banding of Penguins a Problem? Marine Ornithology, 2006.
  • Name that Tune: Call Discrimination and Individual Recognition in Magellanic Penguins. Animal Behaviour, 2006.


  • Living laboratory in peril. Science. May, 2005.
  • Los pingüinos de las costas Argentinas: estado poblacional y conservación. Hornero, 2005.
  • Age and food deprivation affects expression of glucocorticosteriod stress response in Magellanic penguin (Spheniscus magellanicus). Chicks Physiological and Biochemical Zoology, 2005.
  • Intraclutch egg-size variation in Magellanic penguins. The Condor, 2005.
  • Field endocrinology and conservation biology. Integrative and Comparative Biology, 2005.
  • Physiological and behavioral differences in Magellanic penguin chicks in undisturbed and tourist-visited locations of a colony. Conservation Biology, 2005.


  • Physiological condition in Magellanic penguins: does it matter if you have to walk a long way to your nest? The Condor, 2004.
  • Why penguin eggshells are thick. The Auk, 2004.


  • Fighting in female Magellanic penguins: when, why, and who wins? Wilson Bulletin, 2003.
  • Marine birds and harmful algal blooms: sporadic victims or under-reported events? Harmful Algae, 2003.
  • Diving behavior of Magellanic penguins (Spheniscus magellanicus) at Punta Tombo. Canadian Journal of Zoology, 2003.


  • Applying ecology to conservation: tracking breeding penguins at New Island South Reserve, Falkland Islands. Aquatic Conservation: Marine and Freshwater Ecosystems, 2002.
  • A comprehensive review of Endangered Species Act recovery plans. Ecological Applications, 2002.
  • Winning and losing: causes for variability in outcome of fights in male Magellanic penguins (Spheniscus magellanicus). Behavioral Ecology, 2002.
  • A comparison of genetic diversity between the Galápagos penguin and the Magellanic penguin. Conservation Genetics, 2002.
  • Improving U.S. Endangered Species Act recovery plans. Conservation Biology, 2002.
  • Habitat use and breeding distribution of Magellanic penguins in northern San Jorge Gulf, Patagonia, Argentina. Auk, 2002.


  • Response from Boersma and Colleagues. BioScience, 2001.
  • How good are endangered species recovery plans? BioScience, 2001.


  • Three Southern right whales dead along Argentine coast. Marine Mammal Science, 2000.
  • Nesting density and reproductive success in a colonial seabird, the Magellanic penguin. Ecology, 2000.


  • Nest concealment and its relationship to predation and reproductive success in the magellanic penguin at its southern-most continental colony. Ornithologia Neotropical, 1999.
  • Limiting abuse: marine protected areas, a limited solution. Ecological Economics, 1999.
  • Where breeding Magellanic penguins (Spheniscus magellanicus) forage: satellite telemetry results and their implications for penguin conservation. Marine Ornithology, 1999.


  • Population trends of the Galápagos penguin: impacts on El Niño and La Niña. Condor, 1998.
  • Satellite tracking of Magellanic penguin (Spheniscus magellanicus) migration. Condor, 1998.
  • Nest-site characteristics and reproductive success in Magellanic penguins (Spheniscus magellanicus). Auk, 1998.
  • Independent scientific review in natural resource management. Conservation Biology, 1998.
  • Principles for sustainable governance of the oceans. Science, 1998.
  • The breeding ecology of Magellanic penguins at Cabo Virgenes, Argentina: what factors determine reproductive success? Colonial Waterbirds, 1998.
  • The adrenocortical response to stress in incubating Magellanic penguins (Spheniscus magellanicus). Auk, 1998.
  • Flexible growth rates in fork-tailed storm-petrels: a response to environmental variability. Auk, 1998.


  • Magellanic Penguins Decline in South Atlantic. Penguin Conservation, 1997.
  • Feeding chases and food allocation in Adélie penguins (Pygoscelis adeliae). Animal Behaviour, 1997.
  • Efectos de la calidad de habitat sobre el éxito reproductivo del pinguino de magellanes (Spheniscus magellanicus) en Cabo Virgenes Santa Cruz, Argentina. Ornithologia Neotropical, 1997.


  • Breeding biology of the dolphin gull at Punta Tombo, Argentina. Condor, 1996.
  • Status and conservation of Magellanic penguins Spheniscus magellanicus in Patagonia, Argentina. Bird Conservation International, 1996.
  • Aspectos particulares de la biologia de reproduccion y tendencia poblactional del pinguino de magellanes (Spheniscus magellanicus) en la colonia de Cabo Virgenes, Santa Cruz, Argentina. Hornero, 1996.
  • Satellite telemetry of the winter migration of Adélie penguins (Pygoscelis adeliae).Polar Biology, 1996.
  • Feeding behavior, size asymmetries, and food distribution in Magellanic penguin (Spheniscus magellanicus) chicks. Auk, 1996.


  • Un delicado equilibrio en Chubut. America, 1995.
  • Hormonal and reproductive effects of low levels of petroleum fouling in the Magellanic penguin (Spheniscus magellanicus). Auk, 1995.
  • Water over the bridge. American Scientists, 1995.
  • Muddy waters: seabird mortality following the Exxon Valdez oil spill. American Scientists, 1995.


  • Consequences of nest desertion and inattendance for Magellanic penguin hatching success. Auk, 1994.
  • Causes of nest desertion during incubation in the Magellanic penguin (Spheniscus magellanicus). Condor, 1994.
  • Magellanic penguins (Spheniscus magellanicus) are affected by chronic petroleum pollution along the coast of Chubut, Argentina. Auk, 1994.
  • Reproductive endocrinology and weight change in relation to reproductive success in the Magellanic penguin (Spheniscus magellanicus). General and Comparative Endocrinology, 1994.


  • The effects of human disturbance on Magellanic penguin breeding success. Bird Conservation International, 1992.
  • Effects of nest type and location on reproductive success of the Magellanic penguins Spheniscus magellanicus. Marine Ornithology, 1992.


  • Status of wild and captive penguin population. Tree, 1991.
  • Effects of substrate on the distribution of Magellanic penguins (Spheniscus magellanicus). Auk, 1991.


  • Direct and indirect effects of peregrine falcon predation on seabird abundance. Auk, 1990.
  • Parental quality and selection on egg size in the Magellanic penguin. Evolution, 1990.


  • Penguins oiled in Argentina. Science, 1998.


  • Body temperature, torpor and growth in chicks of fork-tailed storm-petrels (Oceanodroma furcata). Physiological Zoology, 1986.
  • Seabirds reflect petroleum pollution. Science, 1986.


  • Why do some birds take so long to hatch? American Naturalists, 1982.


  • Egg neglect in the Procellariiformes: reproductive adaptations in the fork-tailed storm-petrel. Condor, 1979.
  • Egg chilling and the thermal environment of the fork-tailed storm-petrel (Oceanodroma furcata) nest. Physiological Zoology, 1979.


  • Breeding patterns of Galapagos penguins as indicators of oceanographic conditions. Science, 1978.


  • An ecological and behavioral study of the Galápagos penguin. Living Bird, 1977.
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