Scientific article
| 
18 Mar 2026
Scientific article |
| 18 Mar 2026
| 18 Mar 2026
From fossils to conservation: an overview of Antarctic and sub-Antarctic penguins
Raven Quilestino-Olario
CORRESPONDING AUTHOR
Institut für Geologie und Mineralogie, Universität zu Köln, 50674 Köln, Germany
Cited articles
Ainley, D.: The Adélie penguin: bellwether of climate change, Columbia University Press, https://doi.org/10.7312/ainl12306, 2002. a, b
Ainley, D. G., Morandini, V., Barton, K., Lyver, P. O. B., Elrod, M., LaRue, M. A., and Pennycook, J.: Varying population size of the Cape Royds Adélie penguin colony, 1955–2020: a synthesis, Antarct. Sci., 36, 127–140, https://doi.org/10.1017/S0954102024000051, 2024. a
Álvarez-Varas, R., Morales-Moraga, D., González-Acuña, D., Klarian, S. A., and Vianna, J. A.: Mercury exposure in Humboldt (Spheniscus humboldti) and Chinstrap (Pygoscelis antarcticus) penguins throughout the Chilean coast and Antarctica, Arch. Environ. Con. Tox., 75, 75–86, https://doi.org/10.1007/s00244-018-0529-7, 2018. a
Amaral, A. V. R., Sykulski, A. M., Fielding, S., and Cavan, E.: Navigating challenges in spatio-temporal modelling of Antarctic krill abundance: addressing zero-inflated data and misaligned covariates, Spat. Stat., 100937, https://doi.org/10.1016/j.spasta.2025.100937, 2025. a
Barbraud, C. and Weimerskirch, H.: Emperor penguins and climate change, Nature, 411, 183–186, https://doi.org/10.1038/35075554, 2001. a, b
Barbraud, C., Delord, K., Bost, C. A., Chaigne, A., Marteau, C., and Weimerskirch, H.: Population trends of penguins in the French Southern Territories, Polar Biol., 43, 835–850, https://doi.org/10.1007/s00300-020-02691-6, 2020. a
Barreau, E., Ropert-Coudert, Y., Delord, K., Barbraud, C., and Kato-Ropert, A.: Scale matters: sea ice and breeding success of Adélie penguins, Polar Biol., 42, 1405–1410, https://doi.org/10.1007/s00300-019-02531-2, 2019. a
Barrionuevo, M., Lisovski, S., Morgenthaler, A., Millones, A., and Frere, E.: Different migration strategies under warming ocean anomalies in penguins: a study of spatial, oceanographic, and isotopic niche segregation, J. Ornithol., 1–13, https://doi.org/10.1007/s10336-025-02330-z, 2025. a, b
Bas, M., Tivoli, A. M., i Godino, I. B., Salemme, M., Santiago, F., Belardi, J. B., Borella, F., Vales, D. G., Crespo, E. A., and Cardona, L.: Changing diets over time: knock-on effects of marine megafauna overexploitation on their competitors in the southwestern Atlantic Ocean, Paleobiology, 49, 176–190, https://doi.org/10.1017/pab.2022.19, 2023. a
Belyaev, O., Román, A., Belliure, J., Navarro, G., Barbero, L., and Tovar-Sánchez, A.: Assessing topographic features and population abundance in an Antarctic penguin colony through UAV-based deep-learning models, Int. J. Appl. Earth Obs., 134, 104124, https://doi.org/10.1016/j.jag.2024.104124, 2024. a
Blair, J. J.: Tracking penguins, sensing petroleum: “Data gaps” and the politics of marine ecology in the South Atlantic, Environ. Plann. E, 5, 60–80, https://doi.org/10.1177/2514848619882938, 2022. a, b
Boersma, P. D.: Penguins as marine sentinels, Bioscience, 58, 597–607, https://doi.org/10.1641/B580707, 2008. a, b, c, d
Borowicz, A., Forrest, S., Wethington, M., Strycker, N., and Lynch, H. J.: Presence of king penguins (Aptenodytes patagonicus) on Elephant Island provides further evidence of range expansion, Polar Biol., 43, 2117–2120, https://doi.org/10.1007/s00300-020-02760-w, 2020. a
Bost, C.-A., Cotté, C., Bailleul, F., Cherel, Y., Charrassin, J.-B., Guinet, C., Ainley, D. G., and Weimerskirch, H.: Large-scale climatic anomalies affect marine predator foraging behaviour and demography, Nat. Commun., 6, 8220, https://doi.org/10.1038/ncomms9220, 2015. a
Brisson-Curadeau, É., Elliott, K., and Bost, C.-A.: Contrasting bottom-up effects of warming ocean on two king penguin populations, Glob. Change Biol., 29, 998–1008, https://doi.org/10.1111/gcb.16519, 2023. a, b, c, d
Brooks, C. M., Ainley, D. G., Abrams, P. A., Dayton, P. K., Hofman, R. J., Jacquet, J., and Siniff, D. B.: Antarctic fisheries: factor climate change into their management, Nature, 558, 177–180, https://doi.org/10.1038/d41586-018-05372-x, 2018. a
Cajiao, D., Leung, Y. F., Tejedo, P., Barbosa, A., Reck, G., and Benayas, J.: Behavioural responses of two penguin species to human presence at Barrientos Island, a popular tourist site in the Antarctic Peninsula region, Antarct. Sci., 34, 107–119, https://doi.org/10.1017/S0954102021000559, 2022. a, b
Cannone, N., Convey, P., and Malfasi, F.: Antarctic Specially Protected Areas (ASPA): a case study at Rothera Point providing tools and perspectives for the implementation of the ASPA network, Biodivers. Conserv., 27, 2641–2660, https://doi.org/10.1007/s10531-018-1559-1, 2018. a
Clapperton, C. M. and Sugden, D. E.: Holocene glacier fluctuations in South America and Antarctica, Quaternary Sci. Rev., 7, 185–198, https://doi.org/10.1016/0277-3791(88)90005-4, 1988. a, b
Cole, T. L., Zhou, C., Fang, M., Pan, H., Ksepka, D. T., Fiddaman, S. R., Emerling, C. A., Thomas, D. B., Bi, X., Fang, Q., and Ellegaard, M. R.: Genomic insights into the secondary aquatic transition of penguins, Nat. Commun., 13, 3912, https://doi.org/10.1038/s41467-022-31508-9, 2022. a
Constable, A. J., de la Mare, W. K., Agnew, D. J., Everson, I., and Miller, D.: Managing fisheries to conserve the Antarctic marine ecosystem: practical implementation of the Convention on the Conservation of Antarctic Marine Living Resources (CCAMLR), ICES J. Mar. Sci., 57, 778–791, https://doi.org/10.1006/jmsc.2000.0725, 2000. a
Cordonnery, L.: Environmental protection in Antarctica: drawing lessons from the CCAMLR model for the implementation of the Madrid Protocol, Ocean Dev. Int. Law, 29, 125–146, https://doi.org/10.1080/00908329809546120, 1998. a
Coues, E.: Material for a monograph of the Spheniscidae, P. Acad. Nat. Sci. Phila., 24, 170–212, https://www.jstor.org/stable/4624226 (last access: 16 March 2026), 1872. a
Cristofari, R., Liu, X., Bonadonna, F., Cherel, Y., Pistorius, P., Le Maho, Y., Raybaud, V., Stenseth, N. C., Le Bohec, C., and Trucchi, E.: Climate-driven range shifts of the king penguin in a fragmented ecosystem, Nat. Clim. Change, 8, 245–251, https://doi.org/10.1038/s41558-018-0084-2, 2018. a
Croxall, J. P. and Prince, P. A.: Food, feeding ecology and ecological segregation of seabirds at South Georgia, Biol. J. Linn. Soc., 14, 103–131, https://doi.org/10.1111/j.1095-8312.1980.tb00101.x, 1980. a, b, c, d
Dakwa, F. E., Ryan, P. G., Dyer, B. M., Crawford, R. J. M., Pistorius, P. A., and Makhado, A. B.: Long-term variation in the breeding diets of macaroni and eastern rockhopper penguins at Marion Island (1994–2018), Afr. J. Mar. Sci., 43, 187–199, https://doi.org/10.2989/1814232X.2021.1901248, 2021. a
De Roy, T., Jones, M., and Cornthwaite, J.: Penguins: The Ultimate Guide, Princeton University Press, Princeton, NJ, 2nd Edn., ISBN 9780691233574, https://doi.org/10.2307/j.ctv22rth6w, 2022. a, b
Ducklow, H. W., Baker, K., Martinson, D. G., Quetin, L. B., Ross, R. M., Smith, R. C., Stammerjohn, S. E., Vernet, M., and Fraser, W.: Marine pelagic ecosystems: the west Antarctic Peninsula, Philos. T. Roy. Soc. B, 362, 67–94, https://doi.org/10.1098/rstb.2006.1955, 2007. a
Dupuis, B., Chimienti, M., Angelier, F., Raclot, T., Delord, K., Barbraud, C., Ropert-Coudert, Y., and Kato, A.: Energetics link long-term environmental variations to breeding success in a wild penguin population, J. Animal Ecol., https://doi.org/10.1111/1365-2656.70219, 2026. a
Elliott, C.: Antarctica, Scott Base and its environs, New Zealand Geogr., 61, 68–76, https://doi.org/10.1111/j.1745-7939.2005.00005.x, 2005. a
Ellis, D. S., Cipro, C. V., Ogletree, C. A., Smith, K. E., and Aronson, R. B.: A 50-year retrospective of persistent organic pollutants in the fat and eggs of penguins of the Southern Ocean, Environ. Pollut., 241, 155–163, https://doi.org/10.1016/j.envpol.2018.05.003, 2018. a
Emslie, S. D.: Radiocarbon dates from abandoned penguin colonies in the Antarctic Peninsula region, Antarct. Sci., 13, 289–295, https://doi.org/10.1017/S0954102001000414, 2001. a, b, c
Emslie, S. D.: Ancient Adélie penguin colony revealed by snowmelt at Cape Irizar, Ross Sea, Antarctica, Geology, 49, 145–149, https://doi.org/10.1130/G48230.1, 2021. a, b, c
Emslie, S. D. and Woehler, E. J.: A 9000-year record of Adélie penguin occupation and diet in the Windmill Islands, East Antarctica, Antarct. Sci., 17, 57–66, https://doi.org/10.1017/S0954102005002427, 2005. a
Emslie, S. D., McKenzie, A., and Patterson, W. P.: The rise and fall of an ancient Adélie penguin supercolony at Cape Adare, Antarctica, Roy. Soc. Open Sci., 5, 172032, https://doi.org/10.1098/rsos.172032, 2018. a
Emslie, S. D., Romero, M., Juáres, M. A., and Argota, M. R.: Holocene occupation history of pygoscelid penguins at Stranger Point, King George (25 de Mayo) Island, northern Antarctic Peninsula, Holocene, 30, 190–196, https://doi.org/10.1177/0959683619875814, 2020. a
Flynn, C. M., Hart, T., Clucas, G. V., and Lynch, H. J.: Penguins in the anthropause: COVID-19 closures drive gentoo penguin movement among breeding colonies, Biol. Conserv., 286, 110318, https://doi.org/10.1016/j.biocon.2023.110318, 2023. a, b, c
Flynn, C. M., Wethington, M., Hilton, M., Clucas, G. V., Hart, T., Humphries, G. R. W., Nichol, C., and Lynch, H. J.: Revisiting “Penguins in the anthropause: Covid-19 closures drive gentoo penguin movement among breeding colonies”, Biol. Conserv., 315, 111715, https://doi.org/10.1016/j.biocon.2026.111715, 2026. a, b
Forcada, J. and Trathan, P. N.: Penguin responses to climate change in the Southern Ocean, Glob. Change Biol., 15, 1618–1630, https://doi.org/10.1111/j.1365-2486.2009.01909.x, 2009. a, b, c
Fraser, W. R. and Trivelpiece, W. Z.: Factors controlling the distribution of seabirds: winter-summer heterogeneity in the distribution of Adélie penguin populations, Foundations for ecological research west of the Antarctic Peninsula, 70, 257–272, https://doi.org/10.1029/AR070p0257, 1996. a, b, c
Freer, J. J., Warwick-Evans, V., Skaret, G., Krafft, B. A., Fielding, S., and Trathan, P. N.: A new dynamic distribution model for Antarctic krill reveals interactions with their environment, predators, and the commercial fishery in the south Scotia Sea region, Limnol. Oceanogr., 70, 833–849, https://doi.org/10.1002/lno.12809, 2025. a
Freire, R., Massaro, M., McDonald, S., Trathan, P., and Nicol, C. J.: A citizen science trial to assess perception of wild penguin welfare, Frontiers in Veterinary Science, 8, 698685, https://doi.org/10.3389/fvets.2021.698685, 2021. a
Fretwell, P. T.: A 6 year assessment of low sea-ice impacts on emperor penguins, Antarct. Sci., 36, 3–5, https://doi.org/10.1017/S0954102024000130, 2024. a, b
Fretwell, P. T. and Trathan, P. N.: Penguins from space: faecal stains reveal the location of emperor penguin colonies, Global Ecol. Biogeogr., 18, 543–552, https://doi.org/10.1111/j.1466-8238.2009.00467.x, 2009. a, b
Fretwell, P. T. and Trathan, P. N.: Discovery of new colonies by Sentinel-2 reveals good and bad news for emperor penguins, Remote Sens. Ecol. Conserv., 7, 139–153, https://doi.org/10.1002/rse2.176, 2021. a, b, c
Fretwell, P. T., LaRue, M. A., Morin, P., Kooyman, G. L., Wienecke, B., Ratcliffe, N., Fox, A. J., Fleming, A. H., Porter, C., and Trathan, P. N.: An emperor penguin population estimate: the first global, synoptic survey of a species from space, PLOS ONE, 7, e33751, https://doi.org/10.1371/journal.pone.0033751, 2012. a, b, c, d, e
Fretwell, P. T., Boutet, A., and Ratcliffe, N.: Record low 2022 Antarctic sea ice led to catastrophic breeding failure of emperor penguins, Commun. Earth Environ., 4, 273, https://doi.org/10.1038/s43247-023-00927-x, 2023. a, b, c, d
Fretwell, P. T., Bamford, C., Skachkova, A., Trathan, P. N., and Forcada, J.: Regional emperor penguin population declines exceed modelled projections, Commun. Earth Environ., 6, 436, https://doi.org/10.1038/s43247-025-02345-7, 2025. a
Gallagher, K. L., Dinniman, M. S., and Lynch, H. J.: Quantifying Antarctic krill connectivity across the West Antarctic Peninsula and its role in large-scale Pygoscelis penguin population dynamics, Sci. Rep., 13, 12072, https://doi.org/10.1038/s41598-023-39105-6, 2023. a
Gallagher, K. L., Cimino, M. A., Dinniman, M. S., and Lynch, H. J.: Quantifying potential marine debris sources and potential threats to penguins on the West Antarctic Peninsula, Environ. Pollut., 347, 123714, https://doi.org/10.1016/j.envpol.2024.123714, 2024. a
Gao, Y., Yang, L., Wang, J., Xie, Z., Wang, Y., and Sun, L.: Penguin colonization following the last glacial-interglacial transition in the Vestfold Hills, East Antarctica, Palaeogeogr. Palaeoclim., 490, 629–639, https://doi.org/10.1016/j.palaeo.2017.11.053, 2018. a
Gimeno, M., Ramírez, F., Coll, M., Chiaradia, A., and Artana, C.: Cumulative extreme events threaten penguin habitats across the Southern Hemisphere, Glob. Change Biol., 31, e70562, https://doi.org/10.1111/gcb.70562, 2025. a, b
Goldsmith, R. and Sladen, W. J. L.: Temperature regulation of some Antarctic penguins, J. Physiol., 157, 251–262, https://doi.org/10.1113/jphysiol.1961.sp006719, 1961. a
Golubev, S.: Seabirds in conditions of local chronic oil pollution, Davis Sea, Antarctica, Birds, 2, 275–284, https://doi.org/10.3390/birds2030020, 2021. a
Gray, G. R.: The Genera of Birds: Comprising Their Generic Characters, a Notice of the Habits of Each Genus, and an Extensive List of Species Referred to Their Several Genera, Longman, Brown, Green, and Longmans, London, https://doi.org/10.5962/bhl.title.60796, 1844. a
Green, K. and Johnstone, G. W.: Changes in the diet of Adélie penguins breeding in east-Antarctica, Wildlife Res., 15, 103–110, https://doi.org/10.1071/WR9880103, 1988. a
Guímaro, H. R., Thompson, D. R., Morrison, K. W., Fragão, J., Matias, R. S., and Xavier, J. C.: Evidence of eastern rockhopper penguin feeding on a key commercial arrow squid species, Polar Biol., 48, 34, https://doi.org/10.1007/s00300-025-03358-w, 2025. a
Hallet, M. and Lynch, H. J.: Update on the abundance and distribution of Macaroni Penguins (Eudyptes chrysolophus) in the Antarctic Peninsula region, Polar Biol., 47, 607–615, https://doi.org/10.1007/s00300-024-03253-w, 2024. a
Handley, J., Rouyer, M. M., Pearmain, E. J., Warwick-Evans, V., Teschke, K., Hinke, J. T., Lynch, H., Emmerson, L., Southwell, C., Griffith, G., and Cárdenas, C. A.: Marine important bird and biodiversity areas for penguins in Antarctica, targets for conservation action, Front. Mar. Sci., 7, 602972, https://doi.org/10.3389/fmars.2020.602972, 2021. a
Hemmings, A. D.: The 47th Antarctic Treaty Consultative Meeting, Milan, Italy in 2025, Polar J.l, https://doi.org/10.1080/2154896X.2025.2545729, 2025. a
Hinke, J. T., Salwicka, K., Trivelpiece, S. G., Watters, G. M., and Trivelpiece, W. Z.: Divergent responses of Pygoscelis penguins reveal a common environmental driver, Oecologia, 153, 845–855, https://doi.org/10.1007/s00442-007-0781-4, 2007. a, b, c, d
Hinke, J. T., Santos, M. M., Korczak-Abshire, M., Milinevsky, G., and Watters, G. M.: Individual variation in migratory movements of chinstrap penguins leads to widespread occupancy of ice-free winter habitats over the continental shelf and deep ocean basins of the Southern Ocean, PLOS ONE, 14, e0226207, https://doi.org/10.1371/journal.pone.0226207, 2019. a, b
Holdgate, M. W.: Biological research by the British Antarctic Survey, Polar Record, 12, 553–573, https://doi.org/10.1017/S0032247400058769, 1965. a, b
Holt, K. A. and Boersma, P. D.: Unprecedented heat mortality of Magellanic penguins, Condor, 124, duab052, https://doi.org/10.1093/ornithapp/duab052, 2022. a
Hombre, J. D., Arias, M., Carrasco, M. F., González, R. A., and Crespo, E. A.: Status and global population trend of the Magellanic penguin Spheniscus magellanicus along the Argentine coast, Sci, Rep,, 16, 3743, https://doi.org/10.1038/s41598-025-33756-3, 2026. a
Huang, T., Sun, L., Wang, Y., Liu, X., and Zhu, R.: Penguin population dynamics for the past 8500 years at Gardner Island, Vestfold Hills, Antarct, Sci,, 21, 571–578, https://doi.org/10.1017/S0954102009990332, 2009. a, b, c, d
Huddart, D. and Stott, T.: Adventure tourism in Antarctica, in: Adventure Tourism: Environmental Impacts and Management, Springer International Publishing, Cham, 11–50, https://doi.org/10.1007/978-3-030-18623-4_2, 2019. a
Hughes, K. A., Santos, M., Caccavo, J. A., Chignell, S. M., Gardiner, N. B., Gilbert, N., Howkins, A., van Vuuren, B. J., Lee, J. R., Liggett, D., Lowther, A., Lynch, H., Quesada, A., Shin, H. C., Soutullo, A., and Terauds, A.: Ant-ICON “Integrated Science to Inform Antarctic and Southern Ocean conservation”: a new SCAR Scientific Research Programme, Antarct. Sci., 34, 446–455, https://doi.org/10.1017/S0954102022000402, 2022. a
Humphries, G. R., Naveen, R., Schwaller, M., Che-Castaldo, J., McDowall, P., Schrimpf, M., and Lynch, H. J.: Mapping application for penguin populations and projected dynamics (MAPPPD): data and tools for dynamic management and decision support, Polar Record, 53, 160–166, https://doi.org/10.1017/S0032247417000055, 2017. a
Huxley, T. H.: On a fossil bird and a fossil cetacean from New Zealand, Q. J. Geol. Soc. Lond., 15, 670–677, https://doi.org/10.1144/GSL.JGS.1859.015.01-02.73, 1859. a, b
Iles, D. T., Lynch, H., Ji, R., Barbraud, C., Delord, K., and Jenouvrier, S.: Sea ice predicts long-term trends in Adélie penguin population growth, but not annual fluctuations: results from a range-wide multiscale analysis, Glob. Change Biol., 26, 3788–3798, https://doi.org/10.1111/gcb.15085, 2020. a
Jenouvrier, S., Holland, M., Iles, D., Labrousse, S., Landrum, L., Garnier, J., Caswell, H., Weimerskirch, H., LaRue, M., Ji, R., and Barbraud, C.: The Paris Agreement objectives will likely halt future declines of emperor penguins, Glob. Change Biol., 26, 1170–1184, https://doi.org/10.1111/gcb.14864, 2020. a
Jenouvrier, S., Che-Castaldo, J., Wolf, S., Holland, M., Labrousse, S., LaRue, M., Wienecke, B., Fretwell, P., Barbraud, C., Greenwald, N., and Stroeve, J.: The call of the emperor penguin: legal responses to species threatened by climate change, Glob. Change Biol., 27, 5008–5029, https://doi.org/10.1111/gcb.15806, 2021. a
Jenouvrier, S., Eparvier, A., Şen, B., Ventura, F., Che-Castaldo, C., Holland, M., Landrum, L., Krumhardt, K., Garnier, J., Delord, K., and Barbraud, C.: Living with uncertainty: using multi-model large ensembles to assess emperor penguin extinction risk for the IUCN Red List, Biol. Conserv., 305, 111037, https://doi.org/10.1016/j.biocon.2025.111037, 2025. a, b, c
Juarez Martinez, I., Kacelnik, A., Jones, F. M., Hinke, J. T., Dunn, M. J., Raya Rey, A., Lynch, H. J., Owen, K., and Hart, T.: Record phenological responses to climate change in three sympatric penguin species, J. Animal Ecol., https://doi.org/10.1111/1365-2656.70201, 2026. a
Kalvakaalva, R., Clucas, G., Herman, R. W., and Polito, M. J.: Late Holocene variation in the hard prey remains and stable isotope values of penguin and seal tissues from the Danger Islands, Antarctica, Polar Biol., 43, 1571–1582, https://doi.org/10.1007/s00300-020-02728-w, 2020. a
Kawaguchi, S., Atkinson, A., Bahlburg, D., Bernard, K. S., Cavan, E. L., Cox, M. J., Hill, S. L., Meyer, B., and Veytia, D.: Climate change impacts on Antarctic krill behaviour and population dynamics, Nat. Rev. Earth Environ., 5, 43–58, https://doi.org/10.1038/s43017-023-00504-y, 2024. a
Kim, Y., Kim, H., Jeong, M.-S., Kim, D., Kim, J., Jung, J., Seo, H.-M., Han, H.-J., Lee, W.-S., and Choi, C.-Y.: Microplastics in gastrointestinal tracts of gentoo penguin (Pygoscelis papua) chicks on King George Island, Antarctica, Sci. Rep., 13, 13016, https://doi.org/10.1038/s41598-023-39844-6, 2023. a, b
Kirkwood, R. and Robertson, G.: Seasonal change in the foraging ecology of emperor penguins on the Mawson Coast, Antarctica, Mar. Ecol. Prog. Ser., 156, 205–223, https://doi.org/10.3354/meps156205, 1997. a
Korczak-Abshire, M., Hinke, J. T., Milinevsky, G., Juáres, M. A., and Watters, G. M.: Coastal regions of the northern Antarctic Peninsula are key for gentoo populations, Biol. Lett., 17, https://doi.org/10.1098/rsbl.2020.0708, 2021. a
Kristan, A. K.: Penguins past and present: trace elements, stable isotopes, and population dynamics in Antarctic and sub-Antarctic penguins and seals, Master's thesis, Louisiana State University and Agricultural and Mechanical College, proQuest Dissertations & Theses Global (Doc ID 2714864648), https://doi.org/10.31390/gradschool_theses.5351, 2021. a
Krüger, L.: Decreasing trends of chinstrap penguin breeding colonies in a region of major and ongoing rapid environmental changes suggest population level vulnerability, Diversity, 15, 327, https://doi.org/10.3390/d15030327, 2023. a
Krüger, L., Huerta, M. F., Santa Cruz, F., and Cárdenas, C. A.: Antarctic krill fishery effects over penguin populations under adverse climate conditions: implications for the management of fishing practices, Ambio, 50, 560–571, https://doi.org/10.1007/s13280-020-01386-w, 2021. a
Ksepka, D. T. and Clarke, J. A.: The basal penguin (Aves: Sphenisciformes) Perudyptes devriesi and a phylogenetic evaluation of the penguin fossil record, B. Am. Mus. Nat. Hist., 1–77, https://doi.org/10.1206/653.1, 2010. a, b, c
Ksepka, D. T., Fordyce, R. E., Ando, T., and Jones, C. M.: Data from: New fossil penguins (Aves, Sphenisciformes) from the Oligocene of New Zealand reveal the skeletal plan of stem penguins, Dryad [data set], https://doi.org/10.5061/dryad.93j174jd, 2012. a
Labrousse, S., Fraser, A. D., Sumner, M., Tamura, T., Pinaud, D., Wienecke, B., Kirkwood, R., Ropert-Coudert, Y., Reisinger, R., Jonsen, I., and Porter-Smith, R.: Dynamic fine-scale sea icescape shapes adult emperor penguin foraging habitat in East Antarctica, Geophys. Res. Lett., 46, 11206–11218, https://doi.org/10.1029/2019GL084347, 2019. a, b
Labrousse, S., Iles, D., Viollat, L., Fretwell, P., Trathan, P. N., Zitterbart, D. P., Jenouvrier, S., and LaRue, M.: Quantifying the causes and consequences of variation in satellite-derived population indices: a case study of emperor penguins, Remote Sens. Ecol. Conserv., 8, 151–165, https://doi.org/10.1002/rse2.233, 2022. a
Labrousse, S., Nerini, D., Fraser, A. D., Salas, L., Sumner, M., Le Manach, F., Jenouvrier, S., Iles, D., and LaRue, M.: Where to live? Landfast sea ice shapes emperor penguin habitat around Antarctica, Sci. Adv., 9, eadg8340, https://doi.org/10.1126/sciadv.adg8340, 2023. a, b
LaRue, M., Iles, D., Labrousse, S., Fretwell, P., Ortega, D., Devane, E., Horstmann, I., Viollat, L., Foster-Dyer, R., Le Bohec, C., Zitterbart, D. P., Houstin, A., Richter, S., Winterl, A., Wienecke, B., Salas, L., Nixon, M., Barbraud, C., Kooyman, G., Ponganis, P., Ainley, D., Trathan, P., and Jenouvrier, S.: Advances in remote sensing of emperor penguins: first multi-year time series documenting trends in the global population, P. Roy. Soc. B, 291, 20232067, https://doi.org/10.1098/rspb.2023.2067, 2024. a
Le Guen, C., Kato, A., Raymond, B., Barbraud, C., Beaulieu, M., Bost, C.-A., Delord, K., MacIntosh, A. J. J., Meyer, X., Raclot, T., Sumner, M., Takahashi, A., Thiebot, J.-B., and Ropert-Coudert, Y.: Reproductive performance and diving behaviour share a common sea-ice concentration optimum in Adélie penguins (Pygoscelis adeliae), Glob. Change Biol., 24, 5304–5317, https://doi.org/10.1111/gcb.14377, 2018. a
Le Scornec, E., Allain, J., Bardon, G., Barracho, T., Chatelain, N., Delord, K., Filippi, D., Ribout, C., Le Bohec, C., and Barbraud, C.: Intrinsic and extrinsic drivers of juvenile survival in emperor penguins, Polar Biol., 48, 41, https://doi.org/10.1007/s00300-025-03369-7, 2025. a, b, c
Lee, W. Y., Park, S., Kim, K. W., Kim, J. H., Gal, J. K., and Chung, H.: Inter-specific and intra-specific competition of two sympatrically breeding seabirds, chinstrap and gentoo penguins, at two neighboring colonies, Animals, 11, 482, https://doi.org/10.3390/ani11020482, 2021. a
Levick, G. M.: Antarctic Penguins: A Study of Their Social Habits, William Heinemann, London, https://doi.org/10.5962/bhl.title.8385, 1914. a
Libertelli, M. M., Orgeira, J. L., and Alvarez, F.: The explorer king: southern report of the king penguin (Aptenodytes patagonicus) in the Antarctic Peninsula, Antarct. Sci., 36, 145–148, https://doi.org/10.1017/S0954102023000354, 2024. a
Liebeaux, J.: “Cats in congress or penguins in parliament”? Towards the just representation of more-than-humans in Antarctic environmental governance, Polar J., 14, 475–492, https://doi.org/10.1080/2154896X.2024.2414643, 2024. a
Lin, H., Cheng, X., Du, J., Kimball, J. S., Yan, Z., Li, T., Li, T., Li, Y., and Chen, Z.: Satellite mapping of emperor penguin habitat dispersal under climate extremes, Remote Sens. Environ., 330, 114984, https://doi.org/10.1016/j.rse.2025.114984, 2025. a, b
Lishman, G. S.: The food and feeding ecology of Adélie penguins (Pygoscelis adeliae) and Chinstrap penguins (P. antarctica) at Signy Island, South Orkney Islands, J. Zoology, 205, 245–263, https://doi.org/10.1111/j.1469-7998.1985.tb03532.x, 1985. a
Lois, N. A., Campagna, L., Balza, U., Polito, M. J., Pütz, K., Vianna, J. A., Morgenthaler, A., Frere, E., Sáenz-Samaniego, R., Raya Rey, A., and Mahler, B.: Metapopulation dynamics and foraging plasticity in a highly vagile seabird, the southern rockhopper penguin, Ecol. Evol., 10, 3346–3355, https://doi.org/10.1002/ece3.6127, 2020. a
Lowther, A. D., Trathan, P., Tarroux, A., Lydersen, C., and Kovacs, K. M.: The relationship between coastal weather and foraging behaviour of chinstrap penguins, Pygoscelis antarctica, ICES J. Mar. Sci.e, 75, 1940–1948, https://doi.org/10.1093/icesjms/fsy061, 2018. a
Lynch, H. J. and LaRue, M. A.: First global census of the Adélie penguin, Auk, 131, 457–466, https://doi.org/10.1642/AUK-14-31.1, 2014. a, b, c
Lynch, H. J., Naveen, R., Trathan, P. N., and Fagan, W. F.: Spatially integrated assessment reveals widespread changes in penguin populations on the Antarctic Peninsula, Ecology, 93, 1367–1377, https://doi.org/10.1890/11-1588.1, 2012. a, b, c, d
Marples, B. J.: Fossil penguins from the mid-Tertiary of Seymour Island, Falkland Islands Dependencies Survey, Scientific Reports No. 5, https://nora.nerc.ac.uk/id/eprint/511133/1/Fossil%20penguins%20from%20the%20mid-Tertiary%20of%20Seymour%20Island%20-%20FIDS%20Scientific%20Report%205.pdf (last access: 16 March 2026), 1953. a, b, c
Mayr, G., De Pietri, V. L., Love, L., Mannering, A. A., Bevitt, J. J., and Scofield, R. P.: First complete wing of a stem group sphenisciform from the Paleocene of New Zealand sheds light on the evolution of the penguin flipper, Diversity, 12, 46, https://doi.org/10.3390/d12020046, 2020. a
Mayr, G., De Pietri, V. L., Proffitt, J., Blokland, J. C., Clarke, J. A., Love, L., Mannering, A. A., Crouch, E. M., Reid, C., and Scofield, R. P.: Multiple exceptionally preserved fossils from the Paleocene Waipara Greensand inform the diversity of the oldest stem group Sphenisciformes and the formation of their diving adaptations, Zool. J. Linn. Soc., 204, zlaf080, https://doi.org/10.1093/zoolinnean/zlaf080, 2025. a
Meijers, A. J., Meredith, M. P., Murphy, E. J., Chambers, D. P., Belchier, M., and Young, E. F.: The role of ocean dynamics in king penguin range estimation, Nat. Clim. Change, 9, 120–121, https://doi.org/10.1038/s41558-018-0388-2, 2019. a, b
Michelot, C., Kato, A., Raclot, T., Shiomi, K., Goulet, P., Bustamante, P., and Ropert-Coudert, Y.: Sea-ice edge is more important than closer open water access for foraging Adélie penguins: evidence from two colonies, Mar. Ecol. Prog. Ser., 640, 215–230, https://doi.org/10.3354/meps13289, 2020. a, b, c
Michelson, C. I., Polito, M. J., Wunder, M. B., Emslie, S. D., McCarthy, M. D., Patterson, W. P., and McMahon, K. W.: Holocene climate change shifted Southern Ocean biogeochemical cycling and predator trophic dynamics, Limnol. Oceanogr., 68, 2642–2653, https://doi.org/10.1002/lno.12446, 2023. a
Millones, A., Morgenthaler, A., Gandini, P., and Frere, E.: Population numbers of the Magellanic penguin along its central-southern distribution in Argentina: an update after 25 years, Waterbirds, 44, 499–508, https://doi.org/10.1675/063.044.0411, 2021. a
Motas, M., Jerez-Rodríguez, S., Veiga-del Baño, J. M., Ramos, J. J., Oliva, J., Cámara, M. Á., Andreo-Martínez, P., and Corsolini, S.: Emerging pollutants in chinstrap penguins and krill from Deception Island (South Shetland Islands, Antarctica), Toxics, 13, 549, https://doi.org/10.3390/toxics13070549, 2025. a
Murphy, R. C.: A new zonal race of the gentoo penguin, Auk, 64, 454–455, 1947. a
Nievas El Makte, M. L., Polifroni, R., Sepúlveda, M. A., and Fazio, A.: Petroleum hydrocarbons in Atlantic Coastal Patagonia, in: Anthropogenic Pollution of Aquatic Ecosystems, Springer International Publishing, Cham, 325–352, https://doi.org/10.1007/978-3-030-75602-4_15, 2021. a
Noiret, A., Lewden, A., Lemonnier, C., Bocquet, C., Montblanc, M., Bertile, F., Hoareau, M., Marcon, E., Robin, J.-P., Bize, P., and Viblanc, V. A.: Mind the polar sun: solar radiations trigger frequent heat stress in breeding king penguins, despite relatively cool air temperatures, bioRxiv [preprint], https://doi.org/10.1101/2024.09.09.611977, 2024. a
Oehler, D. A., Marin, M., Kusch, A., Danielle, L., Weakley, L. A., and Fry, W. R.: Foraging ranges in southern rockhopper penguins (Eudyptes chrysocome chrysocome) on Isla Noir, Chile, International Journal of Avian & Wildlife Biology, 3, 320–325, https://doi.org/10.15406/ijawb.2018.03.00109, 2018. a
Oelke, H. and Steiniger, F.: Salmonella in Adélie penguins (Pygoscelis adeliae) and south polar skuas (Catharacta maccormicki) on Ross Island, Antarctica, Avian Dis., 568–573, https://doi.org/10.2307/1589155, 1973. a
Olivares, F., Tapia, R., Gálvez, C., Meza, F., Barriga, G. P., Borras-Chavez, R., Mena-Vasquez, J., Medina, R. A., and Neira, V.: Novel penguin avian avulaviruses 17, 18 and 19 are widely distributed in the Antarctic Peninsula, Transbound. Emerg. Dis., 66, 2227–2232, https://doi.org/10.1111/tbed.13309, 2019. a
Paris, J. R., Nitta Fernandes, F. A., Pirri, F., Greco, S., Gerdol, M., Pallavicini, A., Benoiste, M., Cornec, C., Zane, L., Haas, B., and Le Bohec, C.: Gene expression shifts in Emperor penguin adaptation to the extreme Antarctic environment, Mol. Ecol., 34, e17552, https://doi.org/10.1111/mec.17552, 2025. a
Park, J., Hyun, C. U., Kim, Y., Jeong, Y., Yoon, J. E., Guha, S., Kim, J. U., Lee, S. H., Kim, J. H., and Kim, H. C.: Spatial resolution impact assessment for long-term population monitoring of Adélie penguin guano, New Zeal. J. Geol. Geop., 69, e70027, https://doi.org/10.1002/jgo2.70027, 2026. a, b
Parkinson, C. L.: A 40-y record reveals gradual Antarctic sea ice increases followed by decreases at rates far exceeding the rates seen in the Arctic, P. Natl. Acad. Sci. USA, 116, 14414–14423, https://doi.org/10.1073/pnas.1906556116, 2019. a
Pascoe, P., Raymond, B., Carmichael, N., and McInnes, J.: The current trajectory of king penguin (Aptenodytes patagonicus) chick numbers on Macquarie Island in relation to environmental conditions, ICES J. Mar. Sci., 79, 2084–2092, https://doi.org/10.1093/icesjms/fsac139, 2022. a
Pelegrín, J. S. and Acosta Hospitaleche, C.: Evolutionary and biogeographical history of penguins (Sphenisciformes): review of the dispersal patterns and adaptations in a geologic and paleoecological context, Diversity, 14, 255, https://doi.org/10.3390/d14040255, 2022. a
Pertierra, L. R., Hughes, K. A., Vega, M. G., and Olalla-Tarraga, M. A.: High resolution spatial mapping of human footprint across Antarctica and its implications for the strategic conservation of avifauna, PLoS ONE, 12, e0168280, https://doi.org/10.1371/journal.pone.0168280, 2017. a
Pinkerton, M. H., Lyver, P. O. B., Stevens, D. W., Forman, J., Eisert, R., and Mormede, S.: Increases in Adélie penguins in the Ross Sea: Could the fishery for Antarctic toothfish be responsible?, Ecol. Model., 337, 262–271, https://doi.org/10.1016/j.ecolmodel.2016.07.007, 2016. a, b
Puasa, N. A., Zulkharnain, A., Verasoundarapandian, G., Wong, C. Y., Zahri, K. N. M., Merican, F., Shaharuddin, N. A., Gomez-Fuentes, C., and Ahmad, S. A.: Effects of diesel, heavy metals and plastics pollution on penguins in Antarctica: a review, Animals, 11, 2505, https://doi.org/10.3390/ani11092505, 2021. a
Puddicombe, R. A. and Johnstone, G. W.: The breeding season diet of Adélie penguins at the Vestfold Hills, East Antarctica, Hydrobiologia, 165, 239–253, https://doi.org/10.1007/BF00025593, 1988. a
Ratcliffe, N., Hill, S. L., Staniland, I. J., Brown, R., Adlard, S., Horswill, C., and Trathan, P. N.: Do krill fisheries compete with macaroni penguins? Spatial overlap in prey consumption and catches during winter, Divers. Distrib., 21, 1339–1348, https://doi.org/10.1111/ddi.12366, 2015. a
Ratcliffe, N., Deagle, B., Love, K., Polanowski, A., Fielding, S., Wood, A. G., Hill, S., Grant, S., Belchier, M., Fleming, A., and Hall, J.: Changes in prey fields increase the potential for spatial overlap between gentoo penguins and a krill fishery within a marine protected area, Divers. Distrib., 27, 552–563, https://doi.org/10.1111/ddi.13216, 2021. a
Raya Rey, A., Balza, U., Domato, I., and Zunino, F.: New Magellanic penguin Spheniscus magellanicus colony in a subantarctic island, Polar Biol., 45, 1553–1558, https://doi.org/10.1007/s00300-022-03093-6, 2022. a
Richter, S., Gerum, R. C., Schneider, W., Fabry, B., Le Bohec, C., and Zitterbart, D. P.: A remote-controlled observatory for behavioural and ecological research: a case study on emperor penguins, Methods Ecol. Evol., 9, 1168–1178, https://doi.org/10.1111/2041-210X.12971, 2018. a
Ritchie, P. A., Millar, C. D., Gibb, G. C., Baroni, C., and Lambert, D. M.: Ancient DNA enables timing of the Pleistocene origin and Holocene expansion of two Adélie penguin lineages in Antarctica, Mol. Biol. Evol., 21, 240–248, https://doi.org/10.1093/molbev/msh012, 2004. a, b
Roach, L. A., Dörr, J., Holmes, C. R., Massonnet, F., Blockley, E. W., Notz, D., Rackow, T., Raphael, M. N., O'Farrell, S. P., Bailey, D. A., and Bitz, C. M.: Antarctic sea ice area in CMIP6, Geophys. Res. Lett., 47, e2019GL086729, https://doi.org/10.1029/2019GL086729, 2020. a
Robin, J.-P., Bardon, G., Bertile, F., Carette, P., Cillard, A., Lenourry, L., and Le Bohec, C.: When the King penguin meets macroplastics: a first case reported in the Crozet archipelago, Southern Indian Ocean, Mar. Pollut. Bull., 209, 117093, https://doi.org/10.1016/j.marpolbul.2024.117093, 2024. a
Rogan-Finnemore, M.: Council of Managers of National Antarctic Programs (COMNAP): Annual General Meeting (AGM) XXX & Symposium, Polar J., 8, 417–419, https://doi.org/10.1080/2154896X.2018.1541835, 2018. a
Ropert-Coudert, Y., Kato, A., Shiomi, K., Barbraud, C., Angelier, F., Delord, K., Poupart, T., Koubbi, P., and Raclot, T.: Two recent massive breeding failures in an Adélie penguin colony call for the creation of a marine protected area in D'Urville Sea/Mertz, Front. Mar. Sci., 5, 264, https://doi.org/10.3389/fmars.2018.00264, 2018. a
Ross, J. C.: A Voyage of Discovery and Research in the Southern and Antarctic Regions, During the Years 1839–43, John Murray, London, https://doi.org/10.5962/bhl.title.98449, 1847. a, b
Rümmler, M.-C., Esefeld, J., Pfeifer, C., and Mustafa, O.: Effects of UAV overflight height, UAV type, and season on the behaviour of emperor penguin adults and chicks, Remote Sens. Appl., 23, 100558, https://doi.org/10.1016/j.rsase.2021.100558, 2021. a
Salmerón, N., Belle, S., Cruz, F. S., Alegria, N., Finger, J. V. G., Corá, D. H., Petry, M. V., Hernández, C., Cárdenas, C. A., and Krüger, L.: Contrasting environmental conditions precluded lower availability of Antarctic krill affecting breeding chinstrap penguins in the Antarctic Peninsula, Sci. Rep., 13, 5265, https://doi.org/10.1038/s41598-023-32352-7, 2023. a
Schmidt, A. E. and Ballard, G.: Significant chick loss after early fast ice breakup at a high-latitude emperor penguin colony, Antarct. Sci., 32, 180–185, https://doi.org/10.1017/S0954102020000048, 2020. a, b
Schmidt, A. E., Lescroël, A., Lisovski, S., Elrod, M., Jongsomjit, D., Dugger, K. M., and Ballard, G.: Sea ice concentration decline in an important Adélie penguin molt area, P. Natl. Acad. Sci. USA, 120, e2306840120, https://doi.org/10.1073/pnas.2306840120, 2023. a
Slack, K. E., Jones, C. M., Ando, T., Harrison, G. L., Fordyce, R. E., Arnason, U., and Penny, D.: Early penguin fossils, plus mitochondrial genomes, calibrate avian evolution, Mol. Biol. Evol., 23, 1144–1155, https://doi.org/10.1093/molbev/msj124, 2006. a
Smith, R. C., Baker, K. S., Fraser, W. R., Hofmann, E. E., Karl, D. M., Klinck, J. M., Quetin, L. B., Ross, R. M., and Trivelpiece, W. Z.: The Palmer LTER: a long-term ecological research program at Palmer Station, Antarctica, Oceanography, 8, 77–86, https://doi.org/10.5670/oceanog.1995.01, 1995. a
Smith, W. O., Ainley, D. G., Heywood, K. J., and Ballard, G.: New technologies aid understanding of the factors affecting Adélie penguin foraging, Oceanography, 34, 26–27, https://doi.org/10.5670/oceanog.2021.supplement.02-10, 2021. a
Stonehouse, B.: The general biology and thermal balances of penguins, Adv. Ecol. Res., 4, 131–196, https://doi.org/10.1016/S0065-2504(08)60321-9, 1967. a
Strugnell, J. M., McGregor, H. V., Wilson, N. G., Meredith, K. T., Chown, S. L., Lau, S. C., Robinson, S. A., and Saunders, K. M.: Emerging biological archives can reveal ecological and climatic change in Antarctica, Glob. Change Biol., 28, 6483–6508, https://doi.org/10.1111/gcb.16356, 2022. a
Strycker, N., Wethington, M., Borowicz, A., Forrest, S., Witharana, C., Hart, T., and Lynch, H. J.: A global population assessment of the Chinstrap penguin (Pygoscelis antarctica), Sci. Rep., 10, 19474, https://doi.org/10.1038/s41598-020-76479-3, 2020. a
Strycker, N., Borowicz, A., Wethington, M., Forrest, S., Shah, V., Liu, Y., Singh, H., and Lynch, H. J.: Fifty-year change in penguin abundance on Elephant Island, South Shetland Islands, Antarctica: results of the 2019–20 census, Polar Biol., 44, 45–56, https://doi.org/10.1007/s00300-020-02774-4, 2021. a, b, c
Sun, L., Xie, Z., and Zhao, J.: A 3,000-year record of penguin populations, Nature, 407, 858–858, https://doi.org/10.1038/35038163, 2000. a, b, c, d
Sutton, G. J., Bost, C. A., Kouzani, A. Z., Adams, S. D., Mitchell, K., and Arnould, J. P.: Fine-scale foraging effort and efficiency of Macaroni penguins is influenced by prey type, patch density and temporal dynamics, Mar. Biol., 168, 3, https://doi.org/10.1007/s00227-020-03811-w, 2021. a
Tejedo, P., Pertierra, L. R., Benayas, J., Convey, P., Justel, A., and Quesada, A.: Trampling on maritime Antarctica: can soil ecosystems be effectively protected through existing codes of conduct?, Polar Res., 31, 10888, https://doi.org/10.3402/polar.v31i0.10888, 2012. a
Thiebot, J.-B. and Bost, C.-A.: Exploitation of distant Antarctic habitats by juvenile macaroni penguins: conservation implications, Research Square [preprint], https://doi.org/10.21203/rs.3.rs-3035279/v1, 2023. a
Thiébot, J.-B. and Dreyfus, M.: Protecting marine biodiversity beyond national jurisdiction: a penguins' perspective, Mar. Policy, 131, 104640, https://doi.org/10.1016/j.marpol.2021.104640, 2021. a
Thiébot, J.-B., Ropert-Coudert, Y., Raclot, T., Poupart, T., Kato, A., and Takahashi, A.: Adélie penguins' extensive seasonal migration supports dynamic Marine Protected Area planning in Antarctica, Mar. Policy, 109, 103692, https://doi.org/10.1016/j.marpol.2019.103692, 2019. a
Tixier, P., Giménez, J., Reisinger, R. R., Méndez-Fernandez, P., Arnould, J. P. Y., Cherel, Y., and Guinet, C.: Importance of toothfish in the diet of generalist subantarctic killer whales: implications for fisheries interactions, Mar. Ecol. Prog. Ser., 613, 197–210, https://doi.org/10.3354/meps12894, 2019. a
Trathan, P. N., Green, C., Tanton, J., Peat, H., Poncet, J., and Morton, A.: Foraging dynamics of macaroni penguins Eudyptes chrysolophus at South Georgia during brood-guard, Mar. Ecol. Prog. Ser., 323, 239–251, https://doi.org/10.3354/meps323239, 2006. a, b, c, d
Trathan, P. N., Forcada, J., and Murphy, E. J.: Environmental forcing and Southern Ocean marine predator populations: effects of climate change and variability, Philos. T. Roy. Soc. B, 362, 2351–2365, https://doi.org/10.1098/rstb.2006.1953, 2007. a
Trathan, P. N., García-Borboroglu, P., Boersma, D., Bost, C.-A., Crawford, R. J. M., Crossin, G. T., Cuthbert, R. J., Dann, P., Davis, L. S., De La Puente, S., Ellenberg, U., Lynch, H. J., Mattern, T., Pütz, K., Seddon, P. J., Trivelpiece, W., and Wienecke, B.: Pollution, habitat loss, fishing, and climate change as critical threats to penguins, Conserv. Biol., 29, 31–41, https://doi.org/10.1111/cobi.12349, 2015. a, b
Trathan, P. N., Warwick-Evans, V., Hinke, J. T., Young, E. F., Murphy, E. J., Carneiro, A. P. B., Dias, M. P., Kovacs, K. M., Lowther, A. D., Godø, O. R., and Kokubun, N.: Managing fishery development in sensitive ecosystems: identifying penguin habitat use to direct management in Antarctica, Ecosphere, 9, e02392, https://doi.org/10.1002/ecs2.2392, 2018. a
Trathan, P. N., Wienecke, B., Barbraud, C., Jenouvrier, S., Kooyman, G., Le Bohec, C., Ainley, D. G., Ancel, A., Zitterbart, D. P., Chown, S. L., and LaRue, M.: The emperor penguin-Vulnerable to projected rates of warming and sea ice loss, Biol. Conserv., 241, 108216, https://doi.org/10.1016/j.biocon.2019.108216, 2020. a, b, c, d, e
Trivelpiece, W. and Volkman, N. J.: Nest-site competition between Adélie and Chinstrap penguins: an ecological interpretation, Auk, 96, 675–681, https://academic.oup.com/auk/article-abstract/96/4/675/5184822 (last access: 16 March 2026), 1979. a
Trivelpiece, W. Z., Hinke, J. T., Miller, A. K., Reiss, C. S., Trivelpiece, S. G., and Watters, G. M.: Variability in krill biomass links harvesting and climate warming to penguin population changes in Antarctica, P. Natl. Acad. Sci. USA, 108, 7625–7628, https://doi.org/10.1073/pnas.1016560108, 2011. a, b, c
Vianna, J. A., Fernandes, F. A., Frugone, M. J., Figueiró, H. V., Pertierra, L. R., Noll, D., Bi, K., Wang-Claypool, C. Y., Lowther, A., Parker, P., and Le Bohec, C.: Genome-wide analyses reveal drivers of penguin diversification, P. Natl. Acad. Sci. USA, 117, 22303–22310, https://doi.org/10.1073/pnas.2006659117, 2020. a
Volkman, N. J., Presler, P., and Trivelpiece, W.: Diets of pygoscelid penguins at King George Island, Antarctica, Condor, 82, 373–378, https://doi.org/10.2307/1367558, 1980. a
Warwick-Evans, V., Ratcliffe, N., Lowther, A. D., Manco, F., Ireland, L., Clewlow, H. L., and Trathan, P. N.: Using habitat models for chinstrap penguins Pygoscelis antarctica to advise krill fisheries management during the penguin breeding season, Divers. Distrib., 24, 1756–1771, https://doi.org/10.1111/ddi.12817, 2018. a
Watanabe, K.: The Japanese Antarctic Research Expedition in progress and its organization, Journal of Black Sea/Mediterranean Environment, 20, 78–91, 2014. a
Watters, G. M., Hinke, J. T., and Reiss, C. S.: Long-term observations from Antarctica demonstrate that mismatched scales of fisheries management and predator-prey interaction lead to erroneous conclusions about precaution, Sci. Rep., 10, 2314, https://doi.org/10.1038/s41598-020-59223-9, 2020. a
Wethington, M., Flynn, C., Borowicz, A., and Lynch, H. J.: Adélie penguins north and east of the “Adélie gap” continue to thrive in the face of dramatic declines elsewhere in the Antarctic Peninsula region, Sci. Rep., 13, 2525, https://doi.org/10.1038/s41598-023-29465-4, 2023. a, b, c
Wienecke, B., Lieser, J. L., McInnes, J. C., and Barrington, J. H.: Fast ice variability in East Antarctica: observed repercussions for emperor penguins, Endangered Species Res., 55, 1–19, https://doi.org/10.3354/esr01355, 2024. a, b
Willing, R. L.: Australian discoveries of emperor penguin rookeries in Antarctica during 1954–57, Nature, 182, 1393–1394, https://doi.org/10.1038/1821393a0, 1958. a
Wing, S. R., Wing, L. C., O'Connell-Milne, S. A., Barr, D., Stokes, D., Genovese, S., and Leichter, J. J.: Penguins and seals transport limiting nutrients between offshore pelagic and coastal regions of Antarctica under changing sea ice, Ecosystems, 24, 1203–1221, https://doi.org/10.1007/s10021-020-00578-5, 2021. a
Winterl, A., Richter, S., Houstin, A., Barracho, T., Boureau, M., Cornec, C., Couet, D., Cristofari, R., Eiselt, C., Fabry, B., Krellenstein, A., Mark, C., Mainka, A., Ménard, D., Morinay, J., Pottier, S., Schloesing, E., Le Bohec, C., and Zitterbart, D. P.: Remote sensing of emperor penguin abundance and breeding success, Nat. Commun., 15, 4419, https://doi.org/10.1038/s41467-024-48239-8, 2024. a, b
Woehler, E. J., Penney, R. L., Creet, S. M., and Burton, H. R.: Impacts of human visitors on breeding success and long-term population trends in Adélie penguins at Casey, Antarctica, Polar Biol., 14, 269–274, https://doi.org/10.1007/BF00239175, 1994. a
Wood, J. R., Zhou, C., Cole, T. L., Coleman, M., Anderson, D. P., Lyver, P. O. B., Tan, S., Xiang, X., Long, X., Luo, S., and Lou, M.: Sedimentary DNA insights into Holocene Adélie penguin (Pygoscelis adeliae) populations and ecology in the Ross Sea, Antarctica, Nat. Commun., 16, 1798, https://doi.org/10.1038/s41467-025-56925-4, 2025. a
WWF: Penguins, https://www.worldwildlife.org/species/penguin (last access: 10 February 2026), 2026. a
Xia, B.-Y., Pei, J.-L., and Li, Q.-G.: A new penguin fossil from Seymour Island and reassessment of taxonomy and diversity of Eocene Antarctic penguins, Palaeoworld, 33, 1668–1680, https://doi.org/10.1016/j.palwor.2024.04.007, 2024. a, b
Xu, Q.-B., Gao, Y.-S., Yang, L.-J., Yang, W.-Q., Chu, Z.-D., Wang, Y.-H., Sun, L.-G., and Xie, Z.-Q.: Abandonment of penguin subcolonies in the late nineteenth century on Inexpressible Island, Antarctica, J. Geophys. Res.-Biogeo., 125, e2020JG006080, https://doi.org/10.1029/2020JG006080, 2020. a
You, X.: China's Antarctic research drive offers hope for scientists, Nature, 643, 31, https://doi.org/10.1038/d41586-025-02157-x, 2025. a
Younger, J. L., Clucas, G. V., Kooyman, G., Wienecke, B., Rogers, A. D., Trathan, P. N., Hart, T., and Miller, K. J.: Too much of a good thing: sea ice extent may have forced emperor penguins into refugia during the last glacial maximum, Glob. Change Biol., 21, 2215–2226, https://doi.org/10.1111/gcb.12882, 2015. a
Youngflesh, C., Jones, F. M., Lynch, H. J., Arthur, J., Ročkaiová, Z., Torsey, H. R., and Hart, T.: Large-scale assessment of intra- and inter-annual breeding success using a remote camera network, Remote Sens. Ecol. Conserv., 7, 97–108, https://doi.org/10.1002/rse2.171, 2021. a
Zheng, Z., Nie, Y., Chen, X., Jin, J., Chen, Q., and Liu, X.: Historical population dynamics of the Adélie penguin in response to atmospheric-ocean circulation patterns at Beaufort Island, Ross Sea, Antarctica, Global Planet. Change, 216, 103892, https://doi.org/10.1016/j.gloplacha.2022.103892, 2022. a
Short summary
Penguins have survived for over 60 million years, yet many colonies are now changing rapidly. This overview links fossils, ancient colony remains, and modern surveys to place today’s trends in a long context. Some populations are expanding, but ice-dependent species are at high risk where sea ice breaks up earlier, and food shortages can worsen where fishing overlaps. These results support protecting key breeding and feeding areas, and limiting disturbance and catches in possible warming scenarios.
Penguins have survived for over 60 million years, yet many colonies are now changing rapidly....
