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


Carroll, G., Abrahms, B., Cimino, M., and Brodie, S. Spatial match-mismatch between predators and prey under climate change. Nature Ecology & Evolution (accepted, in press).

Clark-Wolf, T. J., Holt, K. A., Johansson, E., Nisi, A. C., Rafiq, K., West, L., Boersma, P. D., Hazen, E. L., Moore, S. E., & Abrahms, B. (2024). The capacity of sentinel species to detect changes in environmental conditions and ecosystem structure. Journal of Applied Ecology, 00, 1–11.

Gaynor, K.M., Abrahms, B., Manlove, K.R., Oestreich, W.K., Smith, J.A. Anthropogenic impacts at the interface of animal spatial and social behaviour. Philosophical Transactions of the Royal Society B (accepted, in press). 

Gaynor, K.,M. McInturff, A.M., Abrahms, B., Smith, A., and Brashares, J. Hunting mode and habitat selection mediate the success of human hunters. Movement Ecology, 12: 29 (2024).

Hazen, E.L., Savoca, M.S., Clark-Wolf, T.J., Czapansky, M., Rabinowitz, P.M., and Abrahms, B. Ecosystem Sentinels as Early Warning Indicators in the Anthropocene. Annual Review of Environment and Resources (accepted, in press).

Picardi, S., Abrahms, B., and Merkle, J. Scale at the interface of spatial and social ecology. Philosophical Transactions of the Royal Society B (accepted, in press). 

Rafiq, K., Jordan, N.R., McNutt, J.W., Neelo, J., Attias, N., Boersma, P.D., Palmer, M., Ruesink, J., and Abrahms, B. (2024). The value of field research in academia. Science 384, 855-856.

Wagner, E. L., Rebstock, G. A., & Boersma, P. D. (2024). A fearful scourge to the penguin colonies: Southern giant petrel (Macronectes giganteus) predation on living Magellanic penguins (Spheniscus magellanicus) may be more common than assumed. Ecology and Evolution, 14, e11258.


Abrahms, B., Carter, N., Clark-Wolf, T.J., Gaynor, K., Johansson, E., McInturff, A., Nisi, A., Rafiq, K., and West, L. 2023. Climate change as a global amplifier of human-wildlife conflict. Nature Climate Change, 13: 224-234. 
Coverage: KUOW, NPR, The Guardian, Newsweek, Scientific American

Clark-Wolf, T.J., Boersma, P.D., Rebstock, G.A., and Abrahms, B. 2023. Climate presses and pulses mediate the decline of a migratory predator. Proceedings of the National Academy of Sciences, 120 (3): e2209821120.

Liu, O.R., Fisher, M., Feist, B.E., Abrahms, B., Richerson, K., and Samhouri, J.F. 2023. Mobility and flexibility enable resilience of human harvesters to environmental perturbation. Global Environmental Change, 78 (2023): 102629.

Moore, S.E. 2023. Chasing inter-species communication: what marine mammals are telling us about our oceans. ICES Journal of Marine Science, 80(4), 690–697.

Nisi, Anna C., Benson, J.F., King, R., Wilmers, C.C. 2023. Habitat fragmentation reduces survival and drives source–sink dynamics for a large carnivore. Ecological Applications, 33(4):e2822.

Rafiq, K., Jordan, N., Golabek, K., McNutt, J.W., Wilson, A., and Abrahms, B. 2023. Increasing ambient temperatures trigger shifts in activity patterns and temporal partitioning in a large carnivore guild. Proceedings of the Royal Society Biological Sciences, 290: 20231938. 

Rafiq, K., Appleby, R., Davies, A., and Abrahms, B. 2023. SensorDrop: a system to remotely detach individual sensors from wildlife tracking collars. Ecology and Evolution, 13 (7): e10220.

Rebstock GA, Boersma PD. 2023. Sex-specific migratory behavior in a marine predator results in higher risks to females. Mar Ecol Prog Ser 725:141-156.

Riekkola, L., Liu, O.R., Feist, B.E., Forney, K.A., Abrahms, B., Hazen, E., and Samhouri, J.F. 2023. Retrospective analysis of conservation measures to reduce the risk of large whale entanglements in a highly lucrative fishery. Biological Conservation, 278: 109880.

Stafford, K.M., J.C. George, Q. Harcharek, S.E. Moore. 2023. Humpback whale sightings in northern Arctic Alaska. Marine Mammal Science. https://doi/org/10.1111/mms13051 

Tennessen, J., Holt, M.M., Wright, B.M., Hanson, M.B., Emmons, C.K., Giles, D.A., Hogan, J.T., Thornton, S.J., Deecke, V.B.. 2023. Divergent foraging strategies between populations of sympatric matrilineal killer whales. Behavioral Ecology, 34(3), 373–386.
Coverage: The Seattle Times, Skagit Valley Herald, Seattle King 5 News, KUOW Public Radio, Oregon 

Wagner, E., Frere, E., Boersma, P.D. 2023. Changing course: Relocating commercial tanker lanes significantly reduces threat of chronic oiling for a top marine predator. Marine Pollution Bulletin, 193: 115-195.

Wagner, E.L., Pearson, S.F., Good, T.P., Hodum P.J., Buhle E.R., Schrimpf M.B. 2023. Resilience to a severe marine heatwave at two Pacific seabird colonies. Mar Ecol Prog Ser :HEATav4. 

Welch, H., Liu, O., Riekkola, L., Abrahms, B., Hazen, E., and Samhouri, J. 2023. Designing dynamic management strategies to reduce human-wildlife conflict. Conservation Biology,

Woodroffe, R., Abrahms, B., English, H. Jumbam, K., Linden, J., Ngatia, D., Rabaiotti, D., and McNutt, J.W. 2023. African wild dogs are hot and hungry: Response to Creel et al. (2023). Biological Conservation, 284: 110198.


Abrahms, B., Rafiq, K., Jordan, N.R., and McNutt, J.W. 2022. Long-term, climate-driven phenological shift in a tropical large carnivore. Proceedings of the National Academy of Sciences , 119 (27): e2121667119.
Coverage: Scientific American and New Scientist

Emmet, R., Augustine, B., Abrahms, B., Rich, L., and Gardner, B. 2022. A spatial capture-recapture model for group-living species. Ecology, 103(10):e3576.

Fahlbusch, J., Czapanskiy, M., Calambokidis, J., Cade, D.E., Abrahms, B., Hazen, E.L., Goldbogen, J.A. 2022. Blue whales increase feeding rates at fine-scale ocean features. Proceedings of the Royal Society Biological Sciences , 289: 20221180.

Gulland, F.M.D., J.D. Baker, M. Howe, E. LaBrecque, L. Leach, S.E. Moore, R.R. Reeves, P.O. Thomas. 2022. A review of climate change effects on marine mammals in United States waters: Past predictions, observed impacts, current research and conservation imperatives. Climate Change Ecology 3, 100054. 

Holt, K.A. and Boersma, P.D. 2022. Unprecedented heat mortality of Magellanic penguins. Ornithological Applications, 124: 1-12.  

McClintock, B., Abrahms, B., Chandler, R., Conn, P., Converse, S., Emmet, R., Gardner, B., Hostetter, N., and Johnson, D.  2022. An integrated path for spatial capture-recapture and animal movement modeling. Ecology, 103(10):e3473.

Merkle, J., Abrahms, B., Armstrong, J., Sawyer, H., Costa, D., and Chalfoun, A. 2022. Site fidelity as a maladaptive behavior in the Anthropocene. Frontiers in Ecology and the Environment, 20(3): 187-194. 

Moore, S.E., J.T. Clarke, S.R. Okkonen, J.M. Grebmeier, C.L. Berchok, K.M. Stafford. 2022. Changes in gray whale phenology and distribution related to prey variability and ocean biophysics in the northern Bering and eastern Chukchi seas. PLoS ONE. 

Oestreich, W.K., Abrahms, B., McKenna, M.F.,  Goldbogen, J.A., Crowder, L.B., and Ryan, J.P. 2022. Acoustic signature reveals blue whales tune life history transitions to oceanographic conditions. Functional Ecology, 36(4): 882-895.

Oestreich, W.K., Aiu, K.M.,  Crowder, L.B., McKenna, M.F.,  Berdahl, A.M. , and Abrahms, B. 2022. The influence of social cues on timing of animal migrations. Nature Ecology & Evolution, .

Picardi, S., Abrahms, B., Morrison, T.A., Verzuh, T., and Merkle, J.A. 2022. Defining Null Expectations for Animal Site Fidelity. Ecology Letters, 10.1111/ele.14148.

Rebstock, G., Abrahms, B., and Boersma, D. 2022. Site fidelity increases reproductive success by increasing foraging efficiency in a marine predator. Behavioral Ecology, 33(4): 868-875.

Rebstock, G., Borboroglu P.G., and Boersma P.D. 2022. Effective marine protection depends on colony and year for a widespread seabird, the Magellanic penguin. Frontiers in Marine Science, 9:815706.

Stafford, K.M., H. Melling, S.E. Moore, C.L. Berchok. E.K. Braen, A.M. Brewer, B.M. Kimber. 2022. Marine mammal detections on the Chukchi Plateau 2009-2020. Journal of the Acoustical Society of America 151(4): 2521-2529.

Wagner, E.L., Cappello, C.D., and Boersma, P.D. 2022. Should I stay or should I go: Factors in- fluencing mate retention and divorce rates in a colonial seabird. Animal Behaviour, 192: 133-144.


Abrahms, B. Human-wildlife conflict under climate change. 2021. Science, 373(6554): 484-485.
Coverage: Popular Science and The Independent

Abrahms, B., Aikens, E.O.,  Armstrong, J.B., Deacy, W.W., Kauffman, M.J., and Merkle, J.A. 2021. Emerging perspectives on resource tracking and animal movement ecology. Trends in Ecology and Evolution, 36(4): 308-320.

Abrahms, B., Teitelbaum, C., Mueller, T.M., and Converse, S.J. 2021. Ontogenetic shifts from social to experiential learning drive avian migration timing. Nature Communications, 12:7326. 

Ahonen, H., K.M. Stafford, C. Lyderson, C.L. Berchok, S.E. Moore and K.M. Kovacs. 2021. Interannual variability in acoustic detection of blue and fin whale calls in the Northeast Atlantic High Arctic between 2008 and 2019. Endangered Species Research 45: 209-224. 

Baylis, A.M.M., de Lecea, A.M., Tierney, M., Orben, R.A., Ratcliffe, N., Wakefield, E., Catry, P., Campioni, L., Costa, M., Boersma, P.D., GalimberN, F., Granadeiro, J.P., Masello, J.F., Pütz, K., Quillfeldt, P., Rebstock, G.A., et al. 202., Overlap between marine predators and proposed Marine Managed Areas on the Patagonian Shelf. Ecological Applications, 00( 00):e02426.

Brodie, S., Abrahms, B., Bograd, S., Carroll, G., Hazen, E., Muhling, B., Pozo Buil, M., Smith, J., Welch, H., and Jacox, M. 2021. Exploring timescales of predictability in species distributions. Ecography, 44 (6): 832-844.

Cappello, C.D. and Boersma, P.D. 2021. Consequences of phenological shifts and a compressed breeding period in Magellanic penguins. Ecology, e03443. doi:10.1002/ecy.3443

Cole, T., Zhou, C., Fang, M., Pan, H., Ksepka, D.T., Fiddaman, S., Emerling C., Thomas, D.B., Bi, X., Fang, Q., Ellegaard, M., Feng, S., Smith, A.L., Heath, T.A., Tennyson, A.J.D., Borboroglu, P.G., Wood, J.R., Hadden, P., Grosser, S., Bost, C, Cherel, Y, Mattern, T., Hart, T., Sinding, M.S., Shepherd, L.D., Phillips, R.A, Quillfeldt, P., Masello, J.F., Bouzat, J.L., Ryan, P.G., Thompson, D.R., Ellenberg, U., Dann, P., Miller, G., Boersma, P.D., Zhao, R., Gilbert, M.T.P, Yang, H., Zhang, D., and Zhang G. Genomic insights into the secondary aquatic transition of penguins. Nature Genetics.

Esmaeili, S., Jesmer, B.R., Albeke, S.E.,  Aikens, E.O., Schoenecker, K.A., King, S.R.B., Abrahms, B., et al. 2021. Body size and digestive system shape resource selection by ungulates: a cross-taxa test of the Forage Maturation Hypothesis. Ecology Letters, 24(10): 2178-2191.

Gownaris, N.J., and Boersma, P.D. 2021. Feet first: Adaptive growth in magellanic penguin chicks. Ecol Evol, 11: 4339–

Hazen, E., Abrahms, B., Brodie, S., Carroll, G., Welch, H., and Bograd, S. 2021. Where did they not go? Considerations for generating pseudo-absences for telemetry-based habitat models. Movement Ecology, 9(5): 1-13. 

Hausner, A., Samhouri, J., Hazen, E., Delgerjargal, D., and Abrahms, B. 2021. Dynamic strategies offer potential to reduce lethal ship collisions with large whales under changing climate conditions. Marine Policy, 130: 104565.

Moore, S. E., J. C. George, R. R. Reeves. 2021. Bowhead whale ecology in changing high-latitude ecosystems. In: George, J.C. and J. G. M. Thewissen, Eds. The Bowhead Whale, Balaena mysticetus, Biology and Human Interactions. Academic Press. 

Samhouri, J., Feist, B., Fisher, M., Liu, O., Woodman, S., Abrahms, B., Forney, K., Hazen, E., Lawson, D., Redfern, J., and Saez, L. 2021. Marine heatwave challenges solutions to human-wildlife conflict. Proceedings of the Royal Society Biological Sciences, 288(1964): 20211607.

Sydeman, W., Hunt, G., Pikitch, E., Parrish, J., Piatt, J., Boersma, P.D., Kaufman, L., Anderson, D., Thompson, S., Sherley, R.. 2021. South Africa’s experimental fisheries closures and recovery of the endangered African penguin. ICES Journal of Marine Science.

Sydeman, W., Schoeman D. S., Thompson A, Hoover B.A., García-Reyes M, Daunt, F, Agnew, P., Anker-Nilssen T., Barbraud C, BarreI R., Becker, P.H., Bell, E., Boersma, P.D., et al. 2021. Hemispheric asymmetry in ocean change and the productivity of ecosystem sentinels. Science, 372 (6545): 980-983.


Blondin, H., Abrahms, B., Crowder, L., Hazen, E.L. 2020. Combining high temporal resolution whale distribution and vessel tracking data improves estimates of ship strike risk. Biological Conservation, 250: 108757.

Dodson, S., Abrahms, B., Hazen, E.L, Bograd, S.J., Feitcher, J. 2020. Disentangling the biotic and abiotic drivers of emergent migratory behavior using individual-based models. Ecological Modeling, 432: 109225.

George, J.C., Moore, S.E., J.G.M. Thewissen. 2020. Bowhead whales: recent insights into their biology, status and resilience. Arctic Report Card 2020, R. L. Thoman, J. Richter-Menge, and M. L. Druckenmiller, Eds. 

Noonan, M., Fleming, C., Tucker, M., Kays, R., Harrison, A.-L., Crofoot, M., Abrahms, B., et al. 2020. Effects of body size on estimation of mammalian area requirements. Conservation Biology, 34(4): 1017-1028. H., Cole TL. Xupeng B., Fang M., Zhou C., Yang Z., Ksepka D.T., Hart T., Bouzat J.L., Argilla L.S., Bertelsen M.F., Boersma P.D., et al. 2020. Correction to: High-coverage genomes to elucidate the evolution of penguins, GigaScience, 9(3), giaa031.


Baylis, A., Tierney M., Orben R., Warwick-Evans V., Wakefield E., Grecian J., Trathan P., Reisin- ger R., Ratcliffe N., Croxall J., Campioni L., Catry P., Crofts S., Boersma P.D, Galimberti F.,Hedd A., Granadeiro J.P., Handley J., Hayes S., Masello J., Montevecchi W., Pütz K., Quillfeldt P., Rebstock G., Sanvito S., Staniland I., Brickle P. Important At-Sea Areas of Colonial Breeding Marine Predators on the Southern Patagonian Shelf. Sci Rep. 9: 8517. 

Boersma, P. D., Garcia Borboroglu, P., Gownaris, N.J., Bost, CA., Chiaradia, A., Ellis, S., Schneider, T., Seddor, P.J., Simeone, A., Trathanm P.N., Waller, L.J., Wienecke, B. 2019. Applying Science to Pressing Conservation Needs for Penguins. Conservation Biology 34: 103-112.

Gownaris, N. and Boersma, P.D. 2019. Sex-Biased Survival Contributes to Population Decline in a Long-Lived Seabird, the Magellanic Penguin. Ecological Applications 29(1): e01826.
Coverage:UW News

Moore, S.E., Haug, T., Vikingsson, G.A., Stenson, G.B. 2019. Baleen whale ecology in Arctic and Subarctic Seas in an era of rapid habitat alteration. Progress in Oceanography 176: 102-118. 

Moore, S.E. and Hauser, D.D.W. 2019. Marine mammals as indicators of Arctic ecosystem variability: finding common ground between Conventional Science and Indigenous Knowledge. Environmental Research Letters 14. 

Moore, S.E. and KuletzK.J. 2019. Marine birds and mammals as ecosystem sentinels in and near Distributed Biological Observatory regions: an abbreviated review of published accounts. Deep-Sea Research II, DBO Special Issue 162: 211-217. 

Ropert-Coudert Y., Chiaradia, A, Ainley, David G., Barbosa, A., Boersma, PD., Brasso, R., Dewar, M., Ellebnurg, U., Garcia-Borborglu, P., Emmerson, L., Hickox, R., Jenouvrier, S., Kato, A., McIntosh, R., Lewis, P., Ramirez, F., Ruoppolo, V., Ryan, P.G., Seddor P.J., Sherley, R.B., Vanstreels, R.E.T., Waller, L., Woehler, E. Trathan, P.N. 2019. Happy Feet in a Hostile World? The Future of Penguins Depends on Proactive Management of Current and Expected Threats. 2019. Frontiers 6: 248. 

Stor, T., Rebstock, G., Garcia-Borboroglu, P., and Boersma, P.D. 2019. Lateralization (handed- ness) in Magellanic penguins. PeerJ 7:e06936.
Coverage: Hakai Magazine

Wagner, E.L. and Boersma, P.D. 2019. Food allocation and feeding behaviours of Magellanic penguins, Spheniscus magellanicus, adults and chicks. Animal Behaviour 148: 161-168.
Coverage: UW News


Auge A.A, M.P. Dias, B. Lascelles, A.M.M. Baylis, A. Black, P.D. Boersma, P. Catry, S. Crofts, F. Galimberti, J.P. Granadeiro, A. Hedd, K. Ludynia, J.F. Masello, W. Montevecchi, R.A. Phillips, K. Putz, P. Quillfeldt, G.A. Rebstock, S. Sanvito, I.J. Staniland, A. Stanworth, D. Thompson, M. Tierney, P.N. Trathan and J.P. Croxall. 2018. Framework for mapping key areas for marine mega-fauna to inform Marine Spatial Planning: the Falkland Islands case study. Marine Policy. 92: 61-72. 

Cappello, C., and P.D. Boersma. 2018. Sexing Galápagos penguins (Spheniscus mendiculus) by morphological measurement. Endangered Species Research. 35: 169-173.
Coverage: UW News

Moore, S.E., Grebmeier, J.M. 2018. The Distributed Biological Observatory: Linking Physics to Biology in the Pacific Arctic Region. The Arctic Institute of North America, 71(5).

Moore, S.E. and Reeves, R.R. 2018. Tracking arctic marine mammal resilience in an era of rapid ecosystem alteration. PLOS Biology 16(10). 

Moore, S.E., Stabeno, P.J., Van Pelt, T.I. 2018. The Synthesis of Arctic Research (SOAR) Project Deep-Sea Research II. 

Moore, S.E., Stabeno, P.J., Grebmeier, J.M., Okkonen S. 2018. The Arctic Marine Pulses (AMP) Model: linking temporal processes to contiguous ecological domains in the Pacific Arctic. Deep-Sea Research II.

Oppel, S., Bolton, M., Carneiro, A.P.B., Dias, M.P., Green, J.A., Masello, J.F., Phillips, R.A., Owen, E., Quillfeldt, P., Beard, A., Bertrand, S., Blackburn, J., Boersma, P.D., et al. 2018. Spatial scales of marine conservation management for breeding seabirds. Marine Policy 98: 37-46.

Rebstock, G.A., and Boersma, P.D.. 2018. Oceanographic conditions in wintering grounds affect arrival date and body condition in breeding female Magellanic penguins. Marine Ecology Progress Series. 601: 253-267.
Coverage: UW News

Wingfield, J.C., Hau, M., Boersma P.D., Romero, L.M, Hillgarth, N., Ramenofsky, M., Wrege, P., Scheibling, R., Kelley, J.P., Walker, B., Wikelsk,i M.. 2018. Effects of El Niño and La Niña South- ern Oscillation events on the adrenocortical responses to stress in birds of the Galápagos Is- lands. General and Comparative Endocrinology. 259: 20-33.


Boersma, P.D., Cappello, C.D., Merlen, G. 2017. First observations of post-fledging care in Galápagos penguins (Spheniscus mendiculus). The Wilson Journal of Ornithology. 129: 186 – 191.

Cerchiara, J.A., Risques, R.A., Prunkard,D.,  Smith, J.R., Kane, O.J., Boersma, P.D. 2017. Telomeres shorten and then lengthen before fledging in Magellanic penguins (Spheniscus magellanicus). Aging. 9: 487- 493.

Pikitch, E.K., Boersma, P.D. , Boyd, I.L., Conover, D.O., Cury, P., Essington, T.E., Heppell, S.S., Houde, E.D., Mangel, M., Pauly, D., Plaganyi, E., Sainsbury, K., Steneck, R.S. 2017. The strong connection between forage fish and their predators: A response to Hillborn et al. (2017). Fisheries Research. doi; 10.1016. https://10.1016/j.fishres.2017.07.022 

Rebstock, G.A., Boersma, P.D. 2017. Comparing reproductive success of a colonial seabird, Magellanic penguins, estimated by course- and fine-scale temporal sampling. Condor. 119(2): 225-238.


Koehn, L.E., Hard, J.J., Akst, E.P., Boersma P.D. 2016. Natural selection on morphology varies among years and by sex in Magellanic Penguins (Spheniscus magellanicus). Auk. 133: 783- 805.

Moore, S.E. 2016. Is it ‘boom times’ for baleen whales in the Pacific Arctic region? The Royal Society, 12(9): 20160251.


Boersma, P.D., Rebstock, G.A., García-Borboroglu, P. 2015. Marine protection is needed for Magellanic penguins in Argentina based on long-term data. Biological Conserv. 182: 197- 204.

Pozzi, L.M., García-Borboroglu P., Boersma P.D., Pascual, M.A. 2015. Population regula- tion in Magellanic penguins: what determines changes in colony size? PLoS ONE.

Rebstock, G.A., Boersma, P.D., García-Borboroglu, P. 2015. Changes in habitat use and nesting density in a declining seabird colony. Population Ecology 58: 105-119.


Boersma, P.D., and G.A. Rebstock. 2014. Climate Change Increases Reproductive Failure in Magellanic Penguins. PLoS ONE 9 (1): e85602.

Pikitch, E., S. Munch, K. Broms, M. Mangel, C. Perretti, P.D. Boersma, I. Boyd, D. Conover, P. Cury, T. Essington, T. Geers, N. Gownaris, S. Heppell, E. Houde, D. Pauly, K. Rountos, K. Sainsbury, C. Santora, and R. Steneck. 2014. Predator Responses to the Exploitation of Forage Fish: The PREP Equation. Conserv. Letters.

Pikitch, E.K., K.J. Rountos, T.E. Essington, C. Santora, D. Pauly, R. Watson, U.R. Sumaila, P.D. Boersma, I.L. Boyd, D.O. Conover, P. Cury, S.S. Heppell, E. D. Houde, M. Mangel, E. Plagányi, K. Sainsbury, R.S. Stenech, T.M. Geers, N. Gownaris, and S.B. Munch. 2014. The global contribution of forage fish to marine fisheries and ecosystems. Fish and Fisheries 15(1): 43–64.

Skewgar, E., P.D. Boersma, and A. Simeone. 2014. Winter Migration of Magellanic Penguins (Spheniscus magellanicus) along the Southeastern Pacific. Waterbirds 37(2):203-209.

Stokes, D.L., P.D. Boersma, and J. Lopez de Casenave. 2014. Conservation of Migratory Magellanic Penguins Requires Marine Zoning. Biological Conserv. 170:151-161.

Trathan, P.N., P. García-Borboroglu, P.D. Boersma, C.-A. Bost, R.J.M. Crawford, G.T. Crossin, R.J. Cuthbert, P. Dann, L.S. Davis, S. de la Puente, U. Ellenberg, H. Lynch, T. Mattern, K. Putz, P. J. Seddon, W. Trivelpiece, and B. Wienecke. 2014. Pollution, Habitat Loss, Fishing, and Climate Change as Critical Threats to Penguins. Conserv. Biology 29: 31-41.

Walker, Brian, P.D. Boersma, and J.C. Wingfield. 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. 2014. Can. J. Zool. 93:123-131.


Pozzi, L.M., P.D. Boersma, and M.A. Pascual. 2013. Beach Attendance in Magellanic penguins Spheniscus magellanicus. Ardeola 60(2):279-290.

Rebstock, G.A. and P.D. Boersma. 2013. Parental behavior controls incubation period and asynchrony of hatching in Magellanic Penguins: Reply to Demongin, Poisbleau, and Eens (2013). The Condor 115(1):5-7.


Boersma, P.D. 2012. Penguins and Petroleum: Lessons in Conservation Ecology. Frontiers in Ecology and the Environment 10:218-219.

Frias, J.E., M.N. Gil, J.L. Esteves, P. Garcia Borboroglu, O.J. Kane, J.R. Smith, and P. D. Boersma. 2012. Mercury levels in feathers of Magellanic penguins. Marine Pollution Bulletin 64(6): 1265-1269.

Kane, O.J., M.M. Uhart, V. Rago, A.J. Pereda, J.R. Smith, A. Van Buren, J.A. Clark, and P.D. Boersma. 2012. Avian pox in Magellanic penguins (Spheniscus magellanicus). Journal of Wildlife Disease 48(3): 790-794.

Knafler, G.J., J.A. Clark, P.D. Boersma, and J.L. Bouzat. 2012. MHC Diversity and Mate Choice in the Magellanic Penguin, Spheniscus magellanicus. Journal of Heredity 103(6): 759-768.

Lewison, R., D. Oro, B.J. Godley, L. Underhill, S. Bearhop, R.P. Wilson, D. Ainley, J.M. Marcos, P.D. Boersma, P.G. Borboroglu, T. Boulinier, M. Frederiksen, M. Genovart, J. González-Solís, J.A. Green, D. Grémillet, K.C. Hamer, G.M. Hilton, K.D. Hyrenbach, A. Martínez-Abraín, W.A. Montevecchi, R.A. Phillips, P.G. Ryan, P. Sagar, W.J. Sydeman, S. Wanless, Y. Watanuki, H. Weimerskirch, P. Yorio. 2012. Research priorities for seabirds: improving conservation and management in the 21st century. Endangered Species Research 17:93-121.

da Silva, R.R., J. Pereira, M. Cirano, C.A.D. Lentini, C. Tanajura., C. Lentinni, M. Cirano, P.D. Boersma, and P. Rodrigues. 2012. Occurrence of Magellanic Penguins along the Brazilian Northeast coast during 2008 austral winter. The Scientific World Journal pp. 1-10.

Wagner, E.L., E. Lee, and P. D. Boersma. 2012. Patterns of acceptance of artificial eggs and chicks by Magellanic penguins (Spheniscus magellanicus). Journal of Ornithology.


Rebstock, G.A., and P.D. Boersma. 2011. Parental behavior controls incubation period and asynchrony of hatching in Magellanic penguins. Condor 113(2):316-325.

Silva, Monica, J.P. Grandadeiro, P.D. Boersma, and I. Strange. 2011. Effects of predation risk on the nocturnal activity budgets of thin-billed prions Pachyptila belcheri on New Island, Falk- land Islands. Polar Biology, 34:421-429.

Wagner, E.L. and P.D. Boersma. 2011. Effects of fisheries on seabird community ecology. Re- views in Fisheries Science 19(3):157-167.


Boersma, P.D., and G.A Rebstock. 2010. Calculating egg volume when shape differs: when are equations appropriate? J. Field Ornithology 81(4):442-448.

Boersma, P.D., and G.A. Rebstock. 2010. Effects of double bands on Magellanic penguins. J. Field Ornithology 81:195-205.

Garcia-Bororoglu, P., P.D. Boersma, V. Ruoppolo, R. Pinho, A. Corrado, D. Conte, R. Velozo, C. Myiaji, G. Dutra, P. Maracini, C. Carvalho, V. Romos, L. Barbosa, and S. Serra. 2010. Magellanic penguin mortality in 2008 along the SW Atlantic coast. Marine Pollution Bulletin 60:1652-1657.

Kane, O.J., J.R. Smith, P.D. Boersma, N.J. Parson, V. Strauss, P. Garcia-Borboroglu, and C. Villanueva. 2010. Feather-loss disorder in African and Magellanic penguins. Colonial Waterbirds 33(3):415-421.

Rebstock, G., Aguero, M.L., Boersma, P.D., et al. 2010. Repeated observations of a Cape Gannet Morus capensis on the coast of Patagonia, Argentina. Journal of African Ornithology, 81(2): 167-169.


Boersma, P.D., and G.A. Rebstock. 2009. Flipper bands do not affect foraging-trip duration of Magellanic penguins. J. Field Ornithology 80:408-418.

Boersma, P.D., and G. A. Rebstock. 2009. Foraging distance affects reproductive success in Magellanic penguins. Marine Ecology Progress Series 375: 263-275.

Boersma, P.D., and G. A. Rebstock. 2009. Intraclutch egg-size dimorphism in Magellanic penguins (Spheniscus magellanicus): adaptation, constraints, or noise? Auk 126: 335-340.

Boersma, P.D., and G.A. Rebstock. 2009. Magellanic Penguin eggshell pores: does number matter? Ibis 151: 535-540.

Boersma, P.D., G. A. Rebstock, E. Frere, and S. E. Moore. 2009. Following the fish: penguins and productivity in the South Atlantic. Ecological Monographs 79(1): 59-76.

Bouzat, J.L., B.G. Walker, and P.D. Boersma. 2009. Regional genetic structure in the Magellanic penguin (Spheniscus magellanicus) suggests metapopulation dynamics. Auk 126: 326- 334.

Skewgar, E., A. Simeone, and P.D. Boersma. 2009. Marine reserve in Chile would benefit penguins and ecotourism. Ocean & Coastal Management 52:487-491.

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