Papers / Books
From the Columbia University Press

The Adelie Penguin: Bellwether of Climate Change

Dr. David G. Ainley, author

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The Adélie penguin is one of the best-studied birds in the world and is the subject of research programs from a dozen nations interested in monitoring changes in the environment and the food webs of the Southern Ocean. This species’ population has been changing dramatically over the past few decades coincident with a general warming of the maritime portion of Antarctica. When the sea-ice is seen to decline so does the population of Adélie penguins. Further south, however, the population is increasing.

This book summarizes our present ecological knowledge of this polar seabird. In so doing, David Ainley describes the ecological factors important to its life history and details the mechanisms by which it is responding to climate change. The author also chronicles the history of research on Adélie penguins, beginning with the heroic expeditions at the beginning of the twentieth century.

From Tumblehome

Waiting for Joey

Jean Pennycook, author

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Join a teacher-scientist as she observes Antarctic penguins over ten breeding seasons.

How do Adelie penguins live and adapt to the changing climatic conditions of Antarctica? The author relates 10 years of observation, tracing the story of one penguin she calls Joey. Children will relate to both the penguins and the author’s sense of adventure as she becomes an expert penguin scientist. Richly illustrated with photos taken on-site at the research station in Antarctica.


If you would like to have a copy, please contact David Ainley 

New Papers

Cusset, F., Bustamante, P., Carravieri, A. et al. Circumpolar assessment of mercury contamination: the Adélie penguin as a bioindicator of Antarctic marine ecosystems. Ecotoxicology 32, 1024–1049 (2023).

Schmidt AE, Lescroël A, Lisovski S, Elrod M, Jongsomjit D, Dugger KM, Ballard G. Sea ice concentration decline in an important Adélie penguin molt area. Proc Natl Acad Sci U S A. 2023 Nov 14;120(46):e2306840120. doi: 10.1073/pnas.2306840120. Epub 2023 Nov 6. PMID: 37931108; PMCID: PMC10656. view PDF

Jongsomjit D, Lescroël A, Schmidt AE, Lisovski S, Ainley DG, Hines E, Elrod M, Dugger KM, Ballard G. Going with the floe: Sea-ice movement affects distance and destination during Adélie penguin winter movements. Ecology. 2024 Feb;105(2):e4196. doi: 10.1002/ecy.4196. Epub 2024 Jan 7. PMID: 37885122.

Shah, K., Schmidt, A. E., Ballard, G., & Schwager, M. (2022). Large scale aerial multi-robot coverage path planning. Field Robotics, 2, 1971–1998.

Jennings, Scott, Dugger, K.M., Ballard,G.,  Ainley, DA,(2021) Effects of Diet and Provisioning Behavior on Chick Growth in Adélie Penguins (Pygoscelis adeliae), Waterbirds Vol. 44, Issue 1, (Dec 2021) , pgs 55-67  

Shick SRA, Elrod ML, Schmidt A, Kraberger S, Ainley DG, Ballard G, Varsani A. Genomes of Single-Stranded DNA Viruses in a Fecal Sample from South Polar Skua (Stercorarius maccormicki) on Ross Island, Antarctica. Microbiol Resour Announc. 2023 Jun 20;12(6):e0029923. doi: 10.1128/mra.00299-23. Epub 2023 May 18. PMID: 37199625; PMCID: PMC10281173.

Jennings, S., K. M. Dugger, G. Ballard, and D. G. Ainley (2023). Faster growth and larger size at creche onset are associated with higher offspring survival in Adelie Penguins. Ornithology 140:ukad006.

Morandini, V., Dugger, K. M., Schmidt, A. E., Varsani, A., Lescroël, A., Ballard, G., Lyver, P.O., Barton, K., & Ainley, D.G. (2024). Sex-specific recruitment rates contribute to male-biased sex ratio in Adélie penguins. Ecology and Evolution, 14, e10859.    

Lescroël A, Schmidt A, Ainley DG, Dugger KM, Elrod M, Jongsomjit D, Morandini V, Winquist S, Ballard G. High-resolution recording of foraging behaviour over multiple annual cycles shows decline in old Adélie penguins’ performance. Proc Biol Sci. 2023 Apr 12;290(1996):20222480. doi: 10.1098/rspb.2022.2480. Epub 2023 Apr 5. PMID: 37015277; PMCID: PMC10072935.

Lescroël A, Schmidt A, Elrod M, Ainley DG, Ballard G. Foraging dive frequency predicts body mass gain in the Adélie penguin. Sci Rep. 2021 Nov 24;11(1):22883. doi: 10.1038/s41598-021-02451-4. PMID: 34819596; PMCID: PMC8613197.

Natural history Patterns

Ainley, D.G. 2007. A letter from Cape Royds: of penguins and men. Pp210-226 in Antarctica: Life on the Ice (S.F. Rogers, Ed.). Traveler’s Tales Books, Sola House, Palo Alto CA.

Ainley, D.G., K.M. Dugger, M. La Mesa, G. Ballard, K.J. Barton, S. Jennings, B.J. Karl, A. Lescroël, P.O’B. Lyver, A. Schmidt, P. Wilson. 2018. Post-fledging survival of Adélie penguins at multiple colonies: chicks raised on fish do well. Marine Ecology Progress Series 601: 239-251.

Ainley, D.G., S.S. Jacobs, C.A. Ribic & I. Gaffney. 1998. Seabird distribution and oceanic features of the Amundsen and southern Bellingshausen seas.  Antarctic Science 10: 111-123 .

Ainley, D.G., N. Nur & E. J. Woehler. 1995.  Factors affecting the size and distribution of pygoscelid penguin colonies in the Antarctic.  Auk 112: 171-182 . view PDF

Ainley, D.G., C.T. Tynan & I. Stirling. 2003. Sea ice: a critical habitat for polar marine mammals and birds. Pp 240-266inSea Ice: An Introduction to its Physics, Biology, Chemistry and Geology (D.N. Thomas & G.S. Dieckmann, Eds.). Blackwell Science, London.

Ainley, D.G., E.J. Woehler & A. Lescroël. 2017. Birds and Antarctic sea ice. In: Sea Ice (3rd Edition) (D.N. Thomas, Ed.). John Wiley & Sons, Ltd., London.

Ballard, G., D.G. Ainley, C.A. Ribic & K.R. Barton. 2001. Effect of instrument attachment on foraging trip duration and nesting success of Adélie penguins. Condor 103: 481-490.

Bost, C.A., C. Cotté, F. Bailleul, Y. Cherel, J.B. Charrassin, C. Guinet, D.G. Ainley,  H. Weimerskirch. 2009. The importance of oceanographic fronts to marine birds and mammals of the southern oceans. Journal of Marine Systems 78: 363–376.

Dugger, K.M., D.G. Ainley, P. O’B. Lyver, K.M. Barton, G. Ballard. 2010. Survival differences and the effect of environmental instability on breeding dispersal in an Adélie penguin meta-population. Proc Natl Acad Sci, doi 10.1073 pnas 1000623107.

Dugger, K.M., G. Ballard, Ainley, D.G. & K.J. Barton. 2006.  Flipper-band effects on the foraging behavior and survival of Adélie Penguins. Auk 123: 858–869.

Grémillet, D., A. Lescroël, G. Ballard, K.M. Dugger, M. Massaro, E.L. Porzig, D.G. Ainley. 2018. Energetic fitness: Field metabolic rates assessed via 3D accelerometry complement conventional fitness metrics. Functional Ecology. 2018;1–11. DOI: 10.1111/1365-2435.13074.

Kappes, P.J., K.M. Dugger, A. Lescroël, D.G. Ainley, G. Ballard, K.J. Barton, P.O’B. Lyver, and P.R. Wilson. 2021. Age-related reproductive performance of the Adélie penguin, a long-lived seabird exhibiting similar outcomes regardless of individual life-history strategy. Journal of Animal Ecology, DOI: 10.1111/1365-2656.13422. 

Jennings, S., A. Varsani, K.M. Dugger, G. Ballard & D.G. Ainley. 2016. Sex-based differences in Adélie Penguin (Pygoscelis adeliae) chick growth rates and diet. PLoS ONE 11(3): e0149090. doi:10.1371/journal.pone.0149090.

LaRue, M., D. Iles, S. Labrousse, L. Salas, G. Ballard, D. Ainley, B. Saenz. 2019. A possible Adélie penguin sub-colony on fast ice by Cape Crozier, Antarctica. Antarctic Science, doi:10.1017/S095410201900018X. 

LaRue, M.A., H.J. Lynch, P.O’B Lyver, K. Barton, D.G Ainley, A. Pollard, W.R. Fraser & G. Ballard. 2014. A method for estimating population colony sizes of Adélie penguins using remote sensing imagery. Polar Biology, doi 10.1007/s00300-014-1451-8

Lescroël, A., K.M. Dugger, G. Ballard & Ainley, D.G. 2009. Effects of individual quality, reproductive success and environmental variability on survival of a long-lived seabird. Journal of Animal Ecology 78(4): 798-806.

Levinson, P., Schmidt, A., Morandini, V., Elrod, M., Jongsomjit, D., & Ballard, G. (2021). Breeding behaviour of colour-aberrant Adélie penguins (Pygoscelis adeliae) at Cape Crozier, Ross Island, Antarctica. Antarctic Science, 1-9. doi:10.1017/S0954102021000158

Lyver POB, Barron M, Barton KJ, Ainley DG, Pollard A, Gordon S, McNeill S, Ballard G & Wilson P.R. 2014. Trends in the breeding population of Adélie penguins in the Ross Sea, 1981–2012: a coincidence of climate and resource extraction effects. PLoS ONE, 9(3): e91188. doi:10.1371/journal.pone.00911

Morandini, V., Dugger, K.M., Lescroël, A. et al. Maintenance of nest quality in Adélie penguins Pygoscelis adeliae: an additional benefit to life in the center. Polar Biol 44, 1553–1562 (2021). 

Petersen, S.L., G.M. Branch, D.G. Ainley,  P. D. Boersma, J. Cooper & E.J. Woehler. 2005. Is flipper banding of penguins a problem? Marine Ornithology 33: 75–79.

Ribic, C.A., D.G. Ainley,  R.G. Ford, W.R. Fraser, C.T. Tynan & E.J. Woehler. 2011. Water masses, ocean fronts, and the structure of Antarctic seabird communities: putting the eastern Bellingshausen Sea in perspective. Deep-Sea Research II 58: 1695-1709,

Russell, D.G.D., Sladen, W.J.L. and Ainley, D.G. 2012. Dr. George Murray Levick (1876-1956): unpublished notes on the sexual habits of the Adélie penguin. Polar Record, Available on CJO doi:10.1017/S0032247412000216.

Santora, J.A., M.A. LaRue & D.G. Ainley. 2020. Geographic Structuring of Antarctic Penguin Populations.  Global Ecology and Biogeography, DOI: 10.1111/geb.13144.

Saenz, B.T., D.G. Ainley, K.L. Daly, G. Ballard, E. Conlisk, M.L. Elrod & S.L. Kim. 2020. Drivers of concentrated predation in an Antarctic marginal-ice-zone food web. Scientific Reports, 10:7282,

Shah, K., G. Ballard, A. Schmidt, M. Schwager, Multidrone aerial surveys of penguin colonies in Antarctica. Sci. Robot. 5, eabc3000 (2020)

Smeele, Z.E., D.G. Ainley, A. Varsani. 2017. Viruses associated with Antarctic wildlife: From serology based detection to identification of genomes using high throughput sequencing. Virus Research 243: 91-105.

Schmidt, A.E., Ballard, G., Lescroël, A. et al. (2021) The influence of subcolony-scale nesting habitat on the reproductive success of Adélie penguins. Sci Rep 11, 15380 (2021).     If you would like to see a short presentation on this paper, click HERE.

Schmidt, A., & Ballard, G. (2020). Significant chick loss after early fast ice breakup at a high-latitude emperor penguin colony. Antarctic Science, 32(3), 180-185. doi:10.1017/S0954102020000048 view PDF

Spear, L.B. & Ainley, D.G. 1998. Morphological differences relative to ecological segregation in petrels (Family: Procellariidae) of the Southern Ocean and tropical Pacific. Auk 115: 1017-1033.

Tynan, C.T., Ainley, D.G. & I. Stirling. 2010. Sea ice: a critical habitat for polar marine mammals and birds, pp 395-424. In D.N. Thomas & G. S. Dieckmann (ed.), Sea Ice, 2nd Edition. Wiley-Blackwell, Chichester, UK.

Varsani, A., S. Kraberger, S. Jennings, E.L. Porzig, L. Julian, M.Massaro, A. Pollard, G. Ballard & Ainley, D.G. 2014. A novel papillomavirus in Adelie penguin (Pygoscelis adeliae) faeces sampled at the Cape Crozier colony, Antarctica. Journal of General Virology 95: 1352–1365

Varsani, A., E.L. Porzig, S. Jennings, S. Kraberger, K. Farkas, L. Julian, M. Massaro, G. Ballard & Ainley, D.G. 2015. Identification of an avian polyomavirus associated with Adélie penguins (Pygoscelis adeliae). Journal of General Virology 96: 851–857.

Wilson, D.J., P. O’B. Lyver, T.C. Greene, A.L. Whitehead, K.M. Dugger, B.J. Karl, J.R. F. Barringer, R. McGarry, A. M. Pollard & Ainley, D.G. 2016. South Polar Skua breeding populations in the Ross Sea assessed from demonstrated relationship with Adélie Penguin numbers. Polar Biology DOI 10.1007/s00300-016-1980-4.

Youngflesh , C., S. Jenouvrier, Y. Li, R. Ji, D.G. Ainley, G. Ballard, C. Barbraud, K. Delord, K.M. Dugger, L.M. Emmerson, W.R. Fraser, J.T. Hinke, P. O’B. Lyver, S. Olmastroni, C.J. Southwell, S.G. Trivelpiece, W.Z. Trivelpiece, H. J. Lynch. 2017. Circumpolar analysis of the Adélie Penguin reveals the importance of environmental variability in phenological mismatch. Ecology 98: 940–951.

Foraging Ecology

Ainley, D.G. & G. Ballard. 2011. Non-consumptive factors affecting foraging patterns in Antarctic penguins: a review and synthesis. Polar Biology 35:1–13.

Ainley, D.G., G. Ballard, R.M. Jones, D. Jongsomjit, S.D. Pierce, W.O. Smith, Jr., S. Veloz. 2015. Trophic cascades in the western Ross Sea, Antarctica: revisited. Marine Ecology Progress Series, doi 10.3354/meps11394.

Ainley, D.G., G. Ballard, K.J. Barton, B.J. Karl, G.H. Rau, C.A. Ribic & P.R. Wilson. 2003. Spatial and temporal variation of diet composition and quality within a presumed metapopulation of Adélie Penguins. Condor 105: 95-106.

Ainley, D.G., G. Ballard & K.M. Dugger. 2006. Competition among penguins and cetaceans reveals trophic cascades in the Ross Sea, Antarctica. Ecology, 87:2080–2093.

Ainley, D.G., G. Ballard, S. Ackley, L.K. Blight, J.T. Eastman, S.D. Emslie, A. Lescroël, S. Olmastroni, S.E. Townsend, C.T. Tynan, P. Wilson, E. Woehler. 2007. Paradigm Lost, or, is top-down forcing no longer significant in the Antarctic Marine Ecosystem? Antarctic Science 19: 283-290.

Ainley, D.G., C.A. Ribic, G. Ballard, S. Heath, I. Gaffney, B.J. Karl, K.R. Barton, P.R. Wilson & S. Webb. 2004. Geographic structure of Adélie Penguin populations: size, overlap and use of adjacent colony-specific foraging areas. Ecological Monographs 74: 159-178.

Ainley, D.G., P.R. Wilson, K.R. Barton, G. Ballard, N. Nur & B. Karl. 1998.  Diet and foraging effort of Adélie penguins in relation to pack-ice conditions in the southern Ross Sea.  Polar Biology 20: 311-319.

Ainley, D.G., E.J Woehler, A. Lescroël. 2017. Birds and Antarctic sea ice. Pp. 570-582 in: Thomas DN (Ed.) Sea Ice, Third Edition. John Wiley & Sons, Ltd, London

Ballance, L.T., D.G. Ainley,  G. Ballard & K. Barton. 2009.  An energetic correlate between colony size and foraging effort in seabirds, an example of the Adélie penguinPygoscelis adeliae. Journal of Avian Biology 40: 279-288.

Ballard, G., K.M. Dugger, N. Nur & Ainley, D.G. 2010. Foraging strategies of Adélie penguins: adjusting body condition to cope with environmental variability. Marine Ecology Progress Series 405: 287–302.

Ballard, G., A.E. Schmidt, V. Toniolo, S. Veloz, D. Jongsomjit, K.R. Arrigo, D.G. Ainley. 2019. Fine-scale oceanographic features characterizing successful Adélie penguin foraging in the SW Ross Sea. Marine Ecology Progress Series 608: 263–277.

Ballerini, T., E.E. Hofmann, D.G. Ainley,  K. Daly, M. Marrari, C.A. Ribic, W.O. Smith Jr., & J.H. Steele. 2013. Productivity and linkages of the food web of the southern region of the western Antarctic Peninsula continental shelf. Progress in Oceanography,

Ford, R.G., D.G. Ainley,  A. Lescroël, P.O’B. Lyver, V. Toniolo & G. Ballard. 2015. Testing assumptions of central place foraging theory: a study of Adélie penguinsPygoscelis adeliaein the Ross Sea. Journal of Avian Biology, 46: 193-205, doi: 10.1111/jav.00491

Jennings, S., A. Varsani, K.M. Dugger, G. Ballard & D.G. Ainley. 2016. Sex-based differences in Adélie Penguin (Pygoscelis adeliae) chick growth rates and diet. PLoS ONE 11(3): e0149090. doi:10.1371/journal.pone.0149090

Karnovsky, N., Ainley, D.G. & P. Lee. 2007. The impact and importance of production in polynyas to top-trophic predators: three case histories. In Polynyas (W.O. Smith, Jr. & D.G. Barber, Eds.). Elsevier Publishers, London. Pp 391-410.

Lescroël, A., G. Ballard, M. Massaro, K. Dugger, S. Jennings, A. Pollard, E. Porzig, A. Schmidt, A. Varsani, D. Grémillet & D. Ainley. 2019. Evidence of age-related improvement in the foraging efficiency of Adélie penguins. Scientific Reports 9:3375, 1.

Lescroël, A., G. Ballard, V. Toniolo, K.J. Barton, P.R. Wilson, P. O’B. Lyver & D.G. Ainley. 2010. Working less to gain more: when breeding quality relates to foraging efficiency. Ecology 91: 2044–2055

Lescroël A, Lyver PO’B, Jongsomjit D, Veloz S and others (2020) Inter-individual differences in the foraging behavior of breeding Adélie penguins are driven by individual quality and sex. Mar Ecol Prog Ser 636:189-205.

Lyver, P.O’B., C.J. MacLeod, G. Ballard, B.J. Karl, J. Adams, D.G. Ainley & P.R. Wilson. 2011. Intra-seasonal variation in foraging behavior among Adélie Penguins (Pygoscelis adeliae) breeding at Cape Hallett, Ross Sea, Antarctica. Polar Biol 34:49–67.

Massaro, M., D.G. Ainley, J.A. Santora, P. Quillfeldt, A. Lescroël, A. Whitehead, A. Varsani, G. Ballard & P.O’B. Lyver. 2020. Diet segregation in Adélie penguins: some individuals attempt to overcome colony-induced and annual foraging challenges. Marine Ecology Progress Series 645: 205–218.

Sergio, F., O.J. Schmitz, C.J. Krebs, R.D. Holt, M.R. Heithaus, A.J. Wirsing, W.J. Ripple, E. Ritchie, D. Ainley, D. Oro, Y. Jhala, F. Hiraldo and E. Korpimäki. 2014. Towards a cohesive, holistic view of top predation: a definition, synthesis and perspective. Oikos doi: 10.1111/oik.01468

Whitehead, A.L., P.O’B. Lyver, G. Ballard, K. Barton, B.J. Karl, K.M. Dugger, S. Jennings, A. Lescroël, P.R. Wilson & D.G. Ainley. 2015. Factors driving Adélie penguin chick size, mass and condition at colonies of differing size in the southern Ross Sea. Marine Ecology Progress Series 523: 199–213.

Climate Change

Ainley, D.G., G. Ballard, S.D. Emslie, W.R. Fraser, P.R. Wilson & E.J. Woehler. 2003. Adélie Penguins and environmental change. Science 300: 429.

Ainley, D.G., E.D. Clarke, K. Arrigo, W.R. Fraser, A. Kato, K.J. Barton & P.R. Wilson. 2005. Decadal-scale changes in the climate and biota of the Pacific sector of the Southern Ocean, 1950s to the 1990s. Antarctic Science 17: 171-182.

Ainley, D.G., K.A. Hobson, X. Crosta, G.H. Rau, L.I. Wassenaar & P.C. Augustinus. 2006. Holocene variation in the Antarctic coastal food web: linking δD and δ13Cin snow petrel diet and marine sediments. Marine Ecology Progress Series, 306: 31-40.

Ainley, D.G., J. Russell, S. Jenouvrier, E. Woehler, P. O’b. Lyver, W.R. Fraser, G.L. Kooyman. 2010. Antarctic penguin response to habitat change as earth’s troposphere reaches 2°c above pre-industrial levels. Ecological Monographs 80: 49-66.

Ainley, D.G., P.R. Wilson & W.R. Fraser. 2001. Effects of climate change on Antarctic sea ice and penguins. Pp. 26-27inImpacts of climate change on wildlife (R.E. Green, M. Harley, M. Spalding & C. Zockler, eds.). RSPB, Cambridge, UK.

Arrigo, K.R., G.L. van Dijken, D.G. Ainley, M.A. Fahnestock & T. Markus. 2002. The impact of the B-15 iceberg on productivity and penguin breeding success in the Ross Sea, Antarctica. Geophys. Res. Letters 29(7), 10.1029/2001GLO14160.

Ballard, G., and D.G. Ainley. 2019. Rapidly diverging population trends of Adélie Penguins reveal limits to a flexible species’ adaptability to Anthropocene climate change. Pp. 91-96 in Biodiversity and Climate Change: Transforming the Biosphere (Editors, T.E. Lovejoy and L. Hannah). Yale University Press, New Haven.

Ballard G., V. Toniolo, D.G. Ainley, C.L. Parkinson, K.R. Arrigo & P.N. Trathan. 2010. Responding to climate change: Adélie penguins confront astronomical and ocean boundaries. Ecology 91: 2056-2069.

Dugger, K.M., G. Ballard, D.G. Ainley, P.O’B. Lyver & C. Schine. 2014. Adélie penguins coping with environmental change: results from a natural experiment at the edge of their breeding range. Frontiers in Ecology and Evolution 2:68, doi: 10.3389/fevo.2014.00068.

Emslie, S.D., P.A. Berkman, D.G. Ainley, L. Coats & M. Polito. 2003. Late-Holocene initiation of ice-free ecosystems in the southern Ross Sea, Antarctica. Marine Ecology Progress Series 262: 19-25.

Kim, S., B. Saenz, J. Scanniello, K. Daly, D. Ainley. 2018. Local climatology of fast ice in McMurdo Sound, Antarctica. Antarctic Science 2018: 1-18, doi:10.1017/S0954102017000578.

Kooyman, G.L., D.G. Ainley,  G. Ballard and P.J. Ponganis. 2007. Effects of giant icebergs on two emperor penguin colonies in the Ross Sea, Antarctica. Antarctic Science 19: 31-38.

LaRue, M.A., D.G. Ainley, M. Swanson, K.M. Dugger, P.O. Lyver, K. Barton & G. Ballard. 2013. Climate change winners: receding ice fields facilitate colony expansion and altered dynamics in an Adélie Penguin metapopulation. PLoS ONE 8(4): e60568. doi:10.1371/journal.pone.0060568.

Lescroël, A., G. Ballard, D. Grémillet, M. Authier & Ainley, D.G. 2014. Antarctic climate change: extreme events disrupt plastic response in Adélie penguins. PLoS ONE, 9(1): e85291. doi:10.1371/journal.pone.0085291.

Ropert-Coudert Y, Chiaradia A, Ainley D, Barbosa A, Boersma PD, Brasso R, Dewar M, Ellenberg U, García-Borboroglu P, Emmerson L, Hickcox R, Jenouvrier S, Kato A, McIntosh RR, Lewis P, Ramírez F, Ruoppolo V, Ryan PG, Seddon PJ, Sherley RB, Vanstreels RET, Waller LJ, Woehler EJ and Trathan PN (2019) Happy Feet in a Hostile World? The Future of Penguins Depends on Proactive Management of Current and Expected Threats. Front. Mar. Sci. 6:248. doi: 10.3389/fmars.2019.00248.

Shepherd, L.D., C.D. Millar, G. Ballard, D.G. Ainley,  P.R. Wilson, G.D. Haynes, C. Baroni & D.M. Lambert. 2005. Microevolution and mega-icebergs in the Antarctic. Proc. Natl Acad. Sci. 102 (46): 16717-16722.

Smith, R.C., E. Domack, S. Emslie, W.R. Fraser, D.G. Ainley,  K. Baker, J. Kennett, A. Leventer, E. Mosley-Thompson, S. Stammerjohn & M. Vernet. 1999. Marine ecosystem sensitivity to historical climate change: Antarctic Peninsula.  BioScience 49: 393-404.

Trathan, P. N., B. Wienecke, C. Barbraud, S. Jenouvrier, G. Kooyman, C. Le Bohec, D. G. Ainley, A. Ancel, D. P. Zitterbart, S. L. Chown, M. LaRuen, R. Cristofari, J. Younger, G. Clucas, C.-A. Bost, J. A. Brown, H. J. Gillett, P. T. Fretwell. 2019. The emperor penguin – Vulnerable to projected rates of warming and sea ice loss. Biological Conservation, 

Wilson, P.R., D.G. Ainley,  N. Nur, S.S. Jacobs, K.J. Barton, G. Ballard & J.C. Comiso. 2001. Adélie Penguin population change in the Pacific Sector of Antarctica: Relation to sea-ice extent and the Antarctic Circumpolar Current. Marine Ecology Progress Series 213: 301-309.

Ross Sea Ecology

Ainley, D.G. 2002. The Ross Sea, Antarctica: Where all ecosystem processes still remain for study. CCAMLR Document WG-EMM 02/60. Hobart, Australia.

Ainley, D.G. 2002. The Ross Sea, Antarctica: Where all ecosystem processes still remain for study, but maybe not for long. Marine Ornithology 31: 55-62.

Ainley, D.G. 2004. Acquiring a ‘base datum of normality’ for a marine ecosystem: the Ross Sea, Antarctica. CCAMLR document WG-EMM-04/20, Hobart.

Ainley D.G. 2007. Insights from study of the last intact neritic marine ecosystem. Trends in Ecology & Evolution 22: 444-445.

Ainley, D.G. 2010. A history of the exploitation of the Ross Sea, Antarctica. Polar Record46 (238): 233–243.

Ainley, D., V. Toniolo, G. Ballard, K. Barton, J. Eastman, B. Karl, S. Focardi, G. Kooyman, P. Lyver, S. Olmastroni, B.S. Stewart, J.W. Testa & P. Wilson. 2006. Managing Ecosystem Uncertainty: Critical Habitat and Dietary Overlap of Top-Predators in the Ross Sea. CCAMLR document EMM 06 – 07, Hobart.

Ainley, D.G., G. Ballard & J. Weller. 2010. ROSS SEA BIOREGIONALIZATION Part I: Validation of the 2007 CCAMLR Bioregionalization Workshop Results Towards Including the Ross Sea in a Representative Network of Marine Protected Areas in the Southern Ocean. CCAMLR WG-EMM-10/11, Hobart.

Ballard, G., D. Jongsomjit & D.G. Ainley. 2010. ROSS SEA BIOREGIONALIZATION, Part Ii: Patterns of Co-Occurrence of Mesopredators in an Intact Polar Ocean Ecosystem. CCAMLR WG-EMM-10/12, Hobart.

Ballard, G., Jongsomjit, D., Veloz, S.D., Ainley, D.G. 2012. Coexistence of mesopredators in an intact polar ocean ecosystem: The basis for defining a Ross Sea marine protected area. Biological Conservation 156: 72-82.

Smith, W.O. Jr., D.G. Ainley & R. Cattaneo-Vietti. 2006. Marine Ecosystems: the Ross Sea.InAntarctic Ecology: From Genes to Ecosystems, Phil. Trans. R. Soc. B (2007) 362, 95–111.

Smith, W.O. Jr., D.G. Ainley, R. Cattaneo-Vietti & E.E. Hofmann. 2012. The Ross Sea continental shelf: Regional biogeochemical cycles, trophic interactions, and potential future changes. In: Antarctic Ecosystems: An Extreme Environment in a Changing World (A.D. Rogers, N.M. Johnston, E.J. Murphy and A. Clarke, Eds.). Chapter 7, Blackwell Publishing, London, pp. 213-242.

Smith, W.O. Jr., D.G. Ainley,  K.R. Arrigo & M.S. Dinniman. 2014. The oceanography and ecology of the Ross Sea. Annual Review of Marine Science 6:469–487.

Whales (Penguin competitor)

Ainley, D.G., K.M. Dugger, V. Toniolo & I. Gaffney. 2007. Cetacean occurrence patterns in the Amundsen and southern Bellingshausen Sea sector, Southern Ocean. Marine Mammal Science 23: 287-305.

Ainley, D.G., G. Ballard, S. Olmastroni. 2009. An Apparent Decrease in the Prevalence of “Ross Sea Killer Whales” in the Southern Ross Sea. Aquatic Mammals 35(3), 335-347.

Ainley, D.G., G. Ballard, L.K. Blight, S. Ackley, S.D. Emslie, A. Lescroël, S. Olmastroni, S.E. Townsend, C.T. Tynan, P. Wilson, E. Woehler. 2010. Impacts of cetaceans on the structure of southern ocean food webs. Marine Mammal Science 26: 482-489.

Ainley, D.G., T.W. Joyce, B. Saenz, R.L. Pitman, J.W. Durban, G. Ballard, K. Daly & S. Kim. 2020. Foraging patterns of Antarctic minke whales in McMurdo Sound, Ross Sea. Antarctic Science, doi:10.1017/S0954102020000310.

Ainley, D.G., D. Jongsomjit, G. Ballard, D. Thiele, W.R. Fraser, C.T. Tynan. 2012. Modeling the relationship of Antarctic minke whales to major ocean boundaries. Polar Biology 35:281-290.

Ainley, D.G. & G. Ballard. 2012. Trophic Interactions and the decrease in Killer Whale (Orcinus orca) prevalence with reduced availability of large fish in the southern Ross Sea. Aquatic Mammals 38: 153-160.

Ainley, D.G., K. Lindke, G. Ballard, P.O’B. Lyver, S. Jennings, V. Toniolo, J. Pennycook, A. Lescroël, M. Massaro & J.A. Santora. Spatio-temporal occurrence patterns of cetaceans near Ross Island, Antarctica, 2002-2015: implications for foodweb dynamics. Polar Biology.

Ballard, G. & D.G. Ainley. 2005. Killer whale harassment of Adélie penguins at Ross Island. Antarctic Science17: 385-386

Lauriano, G., M. Vacchi, D. Ainley & G. Ballard. 2007. Observations of top predators foraging on fish in the pack ice of the southern Ross Sea. Ant. Sci. 19: 439-440.

Seals (Penguin competitor, Predator)

Ainley, D.G., G. Ballard, B.J. Karl & K.T. Dugger. 2005. Leopard seal predation rates at penguin colonies of different size. Antarctic Science17: 335-340.

Ainley, D. G., M.A. LaRue, I. Stirling, S. Stammerjohn & D.B. Siniff. 2015. An apparent population decrease, or change in distribution, of Weddell seals along the Victoria Land coast. Marine Mammal Science, DOI: 10.1111/mms.12220.

Ainley, D.G. & D.B. Siniff. 2009. The importance of Antarctic toothfish as prey of Weddell Seals in the Ross Sea: A Review. Antarctic Science 21: 317-327.

Fahsbender, E., J.M. Burns, S. Kim, S. Kraberger, G. Frankfurter, A.A. Eilers, M.R. Shero, R. Beltran, A. Kirkham, R. McCorkell, R.K. Berngartt, M.F. Male, G. Ballard, D.G. Ainley, M. Breitbart, A. Varsani. 2017. Diverse and highly recombinant anelloviruses associated with Weddell seals in Antarctica. Virus Evolution 3(1): vex017, doi: 10.1093/ve/vex017. 

Kim, S.Z., D.G. Ainley,  J. Pennycook & J.T. Eastman. 2011. Antarctic toothfish heads found along tide cracks of the McMurdo Ice Shelf. Antarctic Science 23: 469-470.

LaRue, M.A., D.G. Ainley, J. Pennycook, K. Stamatiou, L. Salas, N. Nur, S. Stammerjohn & L. Barrington. 2019. Engaging ‘the crowd’ in remote sensing to learn about habitat affinity of the Weddell seal in Antarctica. Remote Sensing in Ecology and Conservation 6 (1):70–78.

LaRue, M.A., J.J. Rotella, R.A. Garrott, D.B. Siniff,  D.G. Ainley,  G.E. Stauffer, C.C. Porter & P.J. Morin. 2011. Satellite imagery can be used to detect variation in abundance of Weddell seals (Leptonychotes weddellii) in Erebus Bay, Antarctica. Polar Biology 34:1727-1737.

LaRue, M.A., L. Salas, N. Nur, D.G. Ainley, S. Stammerjohn, L. Barrington, K. Stamatiou, J. Pennycook, M. Dozier, J. Saints, H. Nakamura. 2019. Physical and ecological factors explain the distribution of Ross Sea Weddell seals during the breeding season. Marine Ecology Progress Series, 612: 193–208.

Salas, L., N. Nur, D. Ainley, J. Burns, J. Rotella, G. Ballard. 2016. Coping with loss of large, energy-dense prey: A potential bottleneck for Weddell Seals in the Ross Sea. Ecological Applications, in press.

Siniff, D.B., R.A. Garrott, J.J. Rotella, W.R. Fraser & D.G. Ainley. 2008. Projecting the effects of environmental change on Antarctic seals. Antarctic Science 20: 425-435.

Antarctic Toothfish (penguin competitor)

Abrams, P.A., D.G. Ainley,  L.K. Blight, P.K. Dayton, J.T. Eastman, J.L. Jacquet. 2016. Necessary elements of precautionary management: implications for the Antarctic toothfish. Fish & Fisheries 17, 1152–1174, DOI: 10.1111/faf.12162.

Ainley, D.G., G. Ballard, J.T. Eastman, C.W. Evans, N. Nur, C.L. Parkinson. 2016. Changed prevalence, not absence explains toothfish status in McMurdo Sound. Antarctic Science, doi:10.1017/S0954102016000584

Ainley, D.G., E.L. Crockett, J.T. Eastman, W.R. Fraser, N. Nur, K. O’Brien, L.A. Salas & D.B. Siniff. 2017. How overfishing a large piscine mesopredator explains growth in Ross Sea penguin populations: a framework to better understand impacts of a controversial fishery. Ecological Modelling 349: 69-75.

Ainley, D.G., N. Nur, J.T. Eastman, G. Ballard, C.L. Parkinson, C.W. Evans, A.L. DeVries. 2013 Decadal trends in abundance, size and condition of Antarctic toothfish in McMurdo Sound, Antarctica, 1972-2011. Fish & Fisheries 14, 343–363 DOI: 10.1111/j.1467-2979.2012.00474.x.

Ainley, D.G., C.M. Brooks, J.T. Eastman, and M. Massaro. 2012. Unnatural selection of Antarctic Toothfish in the Ross Sea, Antarctica. F. Huettmann (ed.),Protection of the Three Poles, pp. 53-75. DOI 10.1007/978-4-431-54006-9_3, Springer, Tokyo.

Ainley, D.G., P.A. Cziko, N. Nur, J.J. Rotella, J.T. Eastman, M. LaRue, I. Stirling & P.A. Abrams. 2020. Further evidence that Antarctic toothfish are important in Weddell seal diet. Antarctic Science, doi:10.1017/S0954102020000437.

Ainley, D.G., J.T. Eastman, C.M. Brooks. 2015. Comments on ‘‘The Antarctic toothfish (Dissostichus mawsoni): biology, ecology, and life history in the Ross Sea region,’’ by S. Hanchet et al. Hydrobiologia, 2015, 761:397–414.

Ainley, D.G., D. Pauly. 2016. Antarctica. In: D. Pauly & D. Zeller (Eds.). Global Atlas of Marine Fisheries: A Critical Appraisal of Catches and Ecosystem Impacts. Island Press, Washington, DC. P. 188.

Barrera-Oro, E., E. Marschoff, D. Ainley. 2017. Changing status of three notothenioid fish at the South Shetland Islands (1983–2016) after impacts of the 1970–80s commercial fishery. Polar Biol DOI 10.1007/s00300-017-2125-0.

Brooks, C.M., D.G. Ainley. 2016. Fishing the bottom of the earth: the political challenges of ecosystem-based management. In: K. Dodds, A.D. Hemmings, P. Roberts, (Eds.) Handbook on the Politics of Antarctica.

Brooks, C.M., D.G. Ainley, P.A. Abrams, P.K. Dayton, R.J. Hofman, J. Jacquet, D.B. Siniff. 2018.  Watch over Antarctic waters — In a rapidly changing climate, fisheries in the Southern Ocean must be managed cautiously in response to data. Nature 158: 77-80.

Brooks, C.M., L.B. Crowder, L.M. Curran, R.B. Dunbar, D.G. Ainley, K.J. Dodds, K.M. Gjerde, U.R. Sumaila. 2016. The decline of science-based management in the Southern Ocean: Conservation or compromise? Science 354: 185-187.

Jacquet, J., E. Blood-Patterson, C. Brooks & D. Ainley. 2016. ‘Rational use’ in Antarctic waters. Marine Policy 63: 28-34.

Ecosystem Change

Ainley, D.G. & L.K. Blight. 2009. Ecological repercussions of historical fish extraction from the southern ocean. Fish and Fisheries 10: 13-38.Blight, L.K, and Ainley, D.G. 2008. Southern Ocean not so pristine. Science 321: 1443.

Ainley, D.G. & C.M. Brooks. 2013. Exploiting the Southern Ocean: rational use or return to tragedy of the commons, pp. 143-154.InLiggett, D. & Hemmings, A. (ed.) Exploring Antarctic Values. Antarctic Gateway Publication Series, No. 1031. Christchurch NZ.

Ainley, D.G. & D. Pauly. 2014. Fishing down the food web of the Antarctic continental shelf and slope. Polar Record 50: 92-107,doi:10.1017/S0032247412000757.

Ainley, D.G. & D. Pauly. 2016. Antarctica. In: Global Atlas of Marine Fisheries: A Critical Appraisal of Catches and Ecosystem Impacts (D. Pauly & D. Zeller, Eds.). Island Press, Washington, DC. P. 188.

Blight, L.K., D.G. Ainley,  S.F. Ackley, G. Ballard, T. Ballerini, R.L. Brownell Jr., C.-H. C. Cheng, M. Chiantore, D. Costa, M.C. Coulter, P. Dayton, A.L. Devries, R. Dunbar, S. Earle, J.T. Eastman, S. D. Emslie, C.W. Evans, R.A. Garrott, S. Kim, G. Kooyman, A. Lescroël, M. Lizotte, M. Massaro, S. Olmastroni, P.J. Ponganis, J. Russell, D.B. Siniff, W.O. Smith Jr., B.S. Stewart, I. Stirling, J. Willis, P.R Wilson & E.J. Woehler. 2010. Fishing for data in the Ross Sea. Science 330: 1316.

Brooks, C.M. & D.G. Ainley. 2016. Fishing the bottom of the earth: the political challenges of ecosystem-based management. In: Handbook on the Politics of Antarctica (K. Dodds, A.D. Hemmings, P. Roberts, Eds.), in press.

Brooks, C.M., L.B. Crowder, L.M. Curran, R.B. Dunbar, D.G. Ainley, K.J. Dodds, K.M. Gjerde & U.R. Sumaila. 2016. The decline of science-based management in the Southern Ocean: Conservation or compromise? Science 354: 185-187.

Christian, C., D. Ainley, M. Bailey, P. Dayton, J. Hocevar, M. LeVine, J. Nikoloyuk, C. Nouvian, E. Velarde, R. Werner & J. Jacquet. 2013. A review of formal objections to Marine Stewardship Council fisheries certifications. Biological Conservation 161: 10–17.

Christian, C., D. Ainley, M. Bailey, P. Dayton, J. Hocevar, M. LeVine, J. Nikoloyuk, C. Nouvian, E. Velarde, R. Werner & J. Jacquet. 2013. Not ‘‘the best environmental choice in seafood’’: A response to Gutiérrez and Agnew (2013). Biological Conservation 161:

Jacquet, J., D. Pauly, D. Ainley, S. Holt, P. Dayton & J. Jackson. 2010. Seafood stewardship in crisis. Nature 467: 28-29.

Jacquet, J., E. Blood-Patterson, C. Brooks & D. Ainley. 2016. ‘Rational use’ in Antarctic waters. Marine Policy 63: 28-34.

Marschoff, E.R., E.R. Barrera-Oro, N.S. Alescio & D.G. Ainley. 2012. Slow recovery of previously depleted demersal fish at the South Shetland Islands, 1983–2010. Fisheries Research 125/126: 206-213.

Tin, T., Z.L. Fleming, K.A. Hughes, D.G. Ainley, P. Convey, C.A. Moreno, S. Pfeiffer, J. Scott & I. Snape. 2009. Impacts of local human activities on the Antarctic environment. Antarctic Science 21, doi:10.1017/S0954102009001722

Woehler, E.J., D. Ainley & J. Jabour. 2013. Human impacts to Antarctic wildlife: predictions and speculations for 2060, pp. 27-60.InTin, T., D. Liggett, P.T. Maher, M. Lamers (eds.),Antarctic Futures, DOI: 10.1007/978-94-007-6582-5_2. Springer, Dordrecht GR.