Foraging behaviour of sympatrically breeding macaroni (Eudyptes chrysolophus) and chinstrap (Pygoscelis antarcticus) penguins at Bouvetøya, Southern Ocean

  • Audun Narvestad Norwegian Polar Institute, Fram Centre, Tromsø, Norway
  • Kit M. Kovacs Norwegian Polar Institute, Fram Centre, Tromsø, Norway
  • Christian Lydersen Norwegian Polar Institute, Fram Centre, Tromsø, Norway
  • Andrew D. Lowther Norwegian Polar Institute, Fram Centre, Tromsø, Norway
Keywords: Ecological niche, niche overlap, central place foraging, competition, stable isotope analysis, biotelemetry


Species with similar ecological requirements that overlap in range tend to segregate their niches to minimize competition for resources. However, the niche segregation possibilities for centrally foraging predators that breed on isolated Subantarctic islands may be reduced by spatial constraints and limitations in the availability of alternative prey. In this study we examined spatial and trophic aspects of the foraging niches of two sympatrically breeding penguin species, macaroni (Eudyptes chrysolophus; MAC) and chinstrap (Pygoscelis antarcticus; CHIN) penguins, at Bouvetøya over two breeding seasons. To measure at-sea movements and diving behaviour, 90 MACs and 49 CHINs were equipped with GPS loggers and dive recorders during two austral summer breeding seasons (2014/15 and 2017/18). In addition, blood samples from tracked birds were analysed for stable isotopes to obtain dietary information. CHINs displayed marked interannual variation in foraging behaviour, diving deeper, utilizing a larger foraging area and displaying enriched values of δ15N in 2014/15 compared to the 2017/18 breeding season. In contrast, MACs dove to similar depths and showed similar δ15N values, while consistently utilizing larger foraging areas compared to CHINs. We suggest that low krill abundances in the waters around Bouvetøya during the 2014/15 season resulted in CHINs shifting toward a diet that increased their niche overlap with MACs. Our findings may partly explain the decreasing number of breeding CHINs at the world’s most remote island, Bouvetøya, while also highlighting the importance of characterizing niche overlap of species using multi-season data sets.


Download data is not yet available.


Adams N.J. & Brown C.R. 1989. Dietary differentiation and trophic relationships in the sub-Antarctic penguin community at Marion Island. Marine Ecology Progress Series 57, 249–258, doi: 10.3354/meps057249.

Atkinson A., Hill S.L., Pakhomov E.A., Siegel V., Reiss C.S., Loeb V.J., Steinberg D.K., Schmidt K., Tarling G.A., Gerrish L. & Sailley S.F. 2019. Krill (Euphausia superba) distribution contracts southward during rapid regional warming. Nature Climate Change 9, 142–147, doi: 10.1038/s41558-018-0370-z.

Atkinson A., Shreeve R.S., Hirst A.G., Rothery P., Tarling G.A., Pond D.W., Korb R.E., Murphy E.J. & Watkins J.L. 2006. Natural growth rates in Antarctic krill (Euphausia superba): II. Predictive models based on food, temperature, body length, sex, and maturity stage. Limnology and Oceanography 51, 973–987, doi: 10.4319/lo.2006.51.2.0973.

Atkinson A., Siegel V., Pakhomov E. & Rothery P. 2004. Long-term decline in krill stock and increase in salps within the Southern Ocean. Nature 432, 100–103, doi: 10.1038/nature02996.

Atkinson A., Siegel V., Pakhomov A.E., Rothery P., Loeb V., Ross M.R., Quetin B.L., Schmidt K., Fretwell P., Murphy J.E., Tarling A.G. & Fleming H.A. 2008. Oceanic circumpolar habitats of Antarctic krill. Marine Ecology Progress Series 362, 1–23, doi: 10.3354/meps07498.

Barbosa A., Benzal J., de León A. & Moreno J. 2012. Population decline of chinstrap penguin (Pygoscelis antarctica) in Deception Island, South Shetlands, Antarctica. Polar Biology 35, 1453–1457, doi: 10.1007/s00300-012-1196-1.

Barlow K.E., Boyd I.L., Croxall J.P., Reid K., Staniland I.J. & Brierley A.S. 2002. Are penguins and seals in competition for Antarctic krill at South Georgia? Marine Biology 140, 205–213, doi: 10.1007/s00227-001-0691-7.

Barlow K.E. & Croxall J.P. 2002a. Seasonal and interannual variation in foraging range and habitat of macaroni penguins Eudyptes chrysolophus at South Georgia. Marine Ecology Progress Series 232, 291–304, doi: 10.3354/meps232291.

Barlow K.E. & Croxall J.P. 2002b. Provisioning behavior of macaroni penguins Eudyptes chrysolophus. Ibis 144, 248–258, doi: 10.1046/j.1474-919X.2002.00046.x.

Bearhop S., Teece M.A., Waldron S. & Furness R.W. 2000. The influence of lipid and uric acid upon δ13C and δ15N values of avian blood: implications for trophic studies. The Auk 117, 504–507, doi: 10.1093/auk/117.2.504.

BirdLife International 2019a. Species factsheet. Macaroni penguin Eudyptes chrysolophus. Accessed on the internet at on 12 April 2019.

BirdLife International 2019b. Species factsheet. Chinstrap penguin Pygoscelis antarcticus. Accessed on the internet at on 12 April 2019.

Biuw M., Lydersen C., de Bruyn N.P.J., Arriola A., Hofmeyr G.J.G., Kritzinger P. & Kovacs K.M. 2010. Long-range migration of a chinstrap penguin from Bouvetøya to Montagu Island, South Sandwich Islands. Antarctic Science 22, 157–162, doi: 10.1017/S0954102009990605.

Blanchet M.-A., Biuw M., Hofmeyr G.J.G., de Bruyn N.P.J., Lydersen C. & Kovacs K.M. 2013. At-sea behavior of three krill predators breeding at Bouvetøya—Antarctic fur seals, macaroni penguins and chinstrap penguins. Marine Ecology Progress Series 477, 285–302, doi: 10.3354/meps10110.

Calenge C. 2015. Home range estimation in R: the adehabitatHR Package. R package version 04. Statistics software accessed on the Internet at on 15 May 2019.

Cherel Y. & Hobson K.A. 2007. Geographical variation in carbon stable isotope signatures of marine predators: a tool to investigate their foraging areas in the Southern Ocean. Marine Ecology Progress Series 329, 281–287, doi: 10.3354/meps329281.

Cherel Y., Hobson K.A., Bailleul F. & Groscolas R. 2005. Nutrition, physiology, and stable isotopes: new information from fasting and molting penguins. Ecology 86, 2881–2888, doi: 10.1890/05-0562.

Clewlow H.L., Takahashi A., Watanabe S., Votier S.C., Downie R. & Ratcliffe N. 2019. Niche partitioning of sympatric penguins by leapfrog foraging appears to be resilient to climate change. Journal of Animal Ecology 88, 223–235, doi: 10.1111/1365-2656.12919.

Cristofari R., Liu X., Banodonna F., Cherel Y., Pistorius P., Le Maho Y., Raybaud V., Stenseth N.C., Le Bohec C. & Trucchi E. 2018. Climate-driven range shifts of the king penguin in a fragment ecosystem. Nature Climate Change 8, 245–251, doi: 10.1033/s41558-018-0084-2.

Croxall J.P. & Davis L.S. 1999. Penguins: paradoxes and patterns. Marine Ornithology 27, 1–12.

Croxall J.P., Davis R.W. & O’Connell M.J. 1988. Diving patterns in relation to diet of gentoo and macaroni penguins at South Georgia. The Condor 90, 157–167, doi: 10.2307/1368444.

Dann P. & Norman F.I. 2006. Population regulation in little penguins (Eudyptula minor): the role of intraspecific competition for nesting sites and food during breeding. Emu 106, 289–296, doi: 10.1071/MU06011.

Davis L.S. & Darby J.T. 1990. Penguin biology. San Diego, CA: Academic Press.

de Brooke M.L. 2004. The food consumption of the world’s seabirds. Proceedings of the Royal Society of London B 271, 246–248, doi: 10.1098/rsbl.2003.0153.

Elliot K.H., Woo K.J., Gaston A.J., Benvenuti S., Dall’Antonia L. & Davoren G.K. 2009. Central-place foraging in an Arctic seabird provides evidence for Storer-Ashmole’s halo. The Auk 126, 613–625, doi: 10.1525/auk.2009.08245.

Forcada J. & Trathan P.N. 2009. Penguin responses to climate change in the Southern Ocean. Global Change Biology 15, 1618–1630, doi: 10.1111/j.1365-2486.2009.01909.x.

Gille S.T. 2002. Warming of the Southern Ocean since the 1950s. Science 295, 1275–1277, doi: 10.1126/science.1065863.

Green J.A., Butler P.J., Woakes A.J. & Boyd I.L. 2002. Energy requirements of female macaroni penguins at South Georgia. Functional Ecology 16, 671–681, doi: 10.1046/j.1365-2435.2002.00670.x.

Haftorn S. 1986. A quantitative analysis of the behaviour of the chinstrap penguin Pygoscelis antarctica and macaroni penguin Eudyptes chrysolophus on Bouvetøya during the late incubation and early nestling periods. Polar Research 4, 33–45, doi: 10.1111/j.1751-8369.1986.tb00516.x.

Hart T., Mann R., Coulson T., Pettorelli N. & Trathan P. 2010. Behavioural switching in a central place forager: patterns of diving behaviour in the macaroni penguin (Eudyptes chrysolophus). Marine Biology 157, 1543–1553, doi: 10.1007/s00227-010-1428-2.

Hindell M.A., Robertson G.G. & Williams R. 1995. Resource partitioning in four species of sympatrically breeding penguins. In P. Dann et al. (eds.): The penguins, ecology and management. Pp. 196–215. Chipping Norton, Australia: Surrey Beatty & Sons.

Hobson K.A. & Welch H.E. 1992. Determination of trophic relationships within a High Arctic marine food web using δ15N and δ13C analysis. Marine Ecology Progress Series 84, 9–18, doi: 10.3354/meps084009.

Hofmeyr G.J.G., Krafft B.A., Kirkman S.P., Bester M.N., Lydersen C. & Kovacs K.M. 2005. Population changes of Antarctica fur seals at Nyrøysa, Bouvetøya. Polar Biology 28, 725–731, doi: 10.1007/s00300-005-0732-7.

Horswill C., Trathan P.N. & Ratcliffe N. 2017. Linking extreme interannual changes in prey availability to foraging behavior and breeding investment in a marine predator, the macaroni penguin. PLoS One 12, e0184114, doi: 10.1371/journal.pone.0184114.

Ichii T., Bengston J.L., Boveng P.L., Takao Y., Jansen J.K., Hiruki-Raring L.M., Cameron M.F., Okamura H., Hayashi T. & Naganobu M. 2007. Provisioning strategies of Antarctic fur seals and chinstrap penguins produce different responses to distribution of common prey and habitat. Marine Ecology Progress Series 344, 277–297, doi: 10.3354/mesps06873.

Inger R. & Bearhop S. 2008. Applications of stable isotope analyses to avian ecology. Ibis 150, 447–461, doi: 10.1111/j.1474-919X.2008.00839.x.

Isaksen K., Huyser O., Kirkman S., Wanless R. & Wilson W. 2000. Studies of seabirds and seals on Bouvetøya 1998/99. Norsk Polarinstitutt Internrapport 2. Tromsø: Norwegian Polar Institute.

Jackson A.L., Inger R., Parnell A.C. & Bearho S. 2011. Comparing isotopic niche widths among and within communities: SIBER—stable isotope Bayesian ellipses in R. Journal of Animal Ecology 80, 595–602, doi: 10.1111/j.1365-2656.2011.01806.x.

Jansen J.K., Russell R.W. & Meyer W.R. 2002. Seasonal shifts in the provisioning behavior of chinstrap penguins, Pygoscelis antarctica. Oecologia 131, 306–318, doi: 10.1007/s00442-002-0880-1.

Johnson D.S., London J.M., Lea M.-A. & Durban J.W. 2008. Continuous-time correlated random walk model for animal telemetry data. Ecology 89, 1208–1215, doi: 10.1890/07-1032.1.

Kirkman S.P., Wilson W., Klages N.T.W., Bester M.N. & Isaksen K. 2000. Diet and estimated food consumption of Antarctic fur seals at Bouvetøya during summer. Polar Biology 23, 745–752, doi: 10.1007/s003000000145.

Krafft A.B., Melle W., Knutsen T., Bagøien E., Broms C., Ellertsen B. & Siegel V. 2010. Distribution and demography of Antarctic krill in the Southeast Atlantic sector of the Southern Ocean during the austral summer 2008. Polar Biology 33, 957–968, doi: 10.1007/s00300-010-0774-3.

Lishman G.S. & Croxall J.P. 1983. Diving depths of the chinstrap penguin Pygoscelis antarctica. British Antarctic Survey Bulletin 61, 21–25.

Lowther A.D., Lydersen C., Biuw M., de Bruyn N.P.J., Hofmeyr G.J.G. & Kovacs K.M. 2014. Post-breeding at-sea movements of three central-place foragers in relation to submesoscale fronts in the Southern Ocean around Bouvetøya. Antarctic Science 26, 533–544, doi: 10.1017/S0954102014000170.

Luque S.P. & Fried R. 2011. Recursive filtering for zero offset correction of diving depth time series with GNU R package diveMove. PLoS One 6, e15850, doi: 10.1371/journal.pone.0015850.

Lynnes A.S., Reid K., Croxall J.P. & Trathan P.N. 2002. Conflict or co-existence? Foraging distribution and competition for prey between Adélie and chinstrap penguins. Marine Biology 141, 1165–1174, doi: 10.1007/s00227-002-0899-1.

McConnell B.J., Chambers C. & Fedak M.A. 1992. Foraging ecology of southern elephant seals in relation to the bathymetry and productivity of the Southern Ocean. Antarctic Science 4, 393–398, doi: 10.1017/S0954102092000580.

Meyer A., Polzin K.L., Sloyan B.M. & Phillips H.E. 2015. Internal waves and mixing near the Kerguelen Plateau. Journal of Physical Oceanography 46, 417–437, doi: 10.1175/JPO-D-15-0055.1.

Miller A.K., Kappes M.A., Trivelpiece S.G. & Trivelpiece W.Z. 2010. Foraging-niche separation of breeding gentoo and chinstrap penguins, South Shetland Islands, Antarctica. The Condor 112, 683–695, doi: 10.1525/cond.2010.090221.

Miller A.K. & Trivelpiece W.Z. 2008. Chinstrap penguins alter foraging and diving behavior in response to the size of their principle prey, Antarctic krill. Marine Biology 154, 201–208, doi: 10.1007/s00227-008-0909-z.

Mori Y. & Boyd I.J. 2004. Segregation of foraging between two sympatric penguin species: does rate maximisation make the difference? Marine Ecology Progress Series 275, 241–249, doi: 10.3354/meps275241.

Niemandt C., Kovacs K.M., Lydersen C., Dyer B.M., Isaksen K., Hofmeyr G.G.J., Mehlum F. & de Bruyn N.P.J. 2016. Chinstrap and macaroni penguin diet and demography at Nyrøysa, Bouvetøya. Antarctic Science 28, 91–100, doi: 10.1017/S0954102015000504.

Park Y.H., Fuda J.L., Durand I. & Naveira Garabato A.C. 2008. Internal tides and vertical mixing over the Kerguelen Plateau. Deep-Sea Research Part II 55, 582–593, doi: 10.1016/j.dsr2.2007.12.027.

Petry M.V., Valls F.C.L., Petersen E.S., Finger J.V.G. & Krüger L. 2018. Population trends of seabirds at Stinker Point, Elephant Island, Maritime Antarctica. Antarctic Science 30, 220–226, doi: 10.1017/S0954102018000135.

Polito M.J., Reiss C.S., Trivelpiece W.Z., Patterson W.P. & Emslie S.D. 2013. Stable isotopes identify an ontogenetic niche expansion in Antarctic krill (Euphausia superba) from the Southern Shetland Islands, Antarctica. Marine Biology 160, 1131–1323, doi: 10.1007/s00227-013-2182-z.

Polito M.J., Trivelpiece W.Z., Patterson W.P., Karnovsky N.J., Reiss C.S. & Emslie S.D. 2015. Contrasting specialist and generalist patterns facilitate foraging niche partitioning in sympatric populations of Pygoscelis penguins. Marine Ecology Progress Series 519, 221–237, doi: 10.3354/meps11095.

Post D.M. 2002. Using stable isotopes to estimate trophic position: models, methods and assumptions. Ecology 83, 703–718, doi: 10.1890/0012-9658(2002)083[0703:USITET]2.0.CO;2.

QGIS Development Team 2019. QGIS Geographic Information System. Open Source Geospatial Foundation Project. Statistics software accessed on the Internet at on 15 March 2019.

Ratcliffe N., Adlard S., Stowasser G. & McGill R. 2018. Dietary divergence is associated with increased intra-specific competition in a marine predator. Nature 8, article no. 6827, doi: 10.1038/s41598-018-25318-7.

Reid K. & Croxall P.J. 2001. Environmental response of upper trophic-level predators reveals a system change in an Antarctic marine ecosystem. Proceedings of Royal Society of London B 268, 377–384, doi: 10.1098/rspb.2000.1371.

Rombolá E.F., Marschoff E. & Coria N. 2010. Inter-annual variability in chinstrap penguin diet at South Shetland and South Orkneys islands. Polar Biology 33, 799–806, doi: 10.1007/s00300-009-0757-4.

Scrucca L., Fop M., Murphy T.B. & Raftery A.E. 2016. mclust 5: clustering, classification and density estimation using Gaussian finite mixture models. The R Journal 8, 289–317, doi: 10.32614/RJ-2016-021.

Strycker N., Wethington M., Borowicz A., Forrest S., Witharana C., Hart T. & Lynch H.J. 2020. A global population assessment of the chinstrap penguin (Pygoscelis antarctica). Scientific Reports 10, 19474, doi: 10.1038/s41598-020-76479-3.

Tarroux A., Lowther A.D., Lydersen C. & Kit M.K. 2016. Temporal shift in the isotopic niche of female Antarctic fur seals from Bouvetøya. Polar Research 35, 31335, doi: 10.3402/polar.v35.31335.

Thiebot J.-B., Lescroël A., Pinaud D., Trathan P.N. & Bost C.-A. 2011. Larger foraging range but similar habitat selection in non-breeding versus breeding sub-Antarctic penguins. Antarctic Science 23, 117–126, doi: 10.1017/S0954102010000957.

Thorpe S.E., Murphy E.J. & Watkins J.L. 2007. Circumpolar connections between Antarctic krill (Euphausia superba Dana) populations: investigating the roles of ocean and sea ice transport. Deep-Sea Research Part I 54, 792–810, doi: 10.1016/j.dsr.2007.01.008.

Tierney M., Southwell C., Emmerson L.M. & Hindell M.A. 2008. Evaluating and using stable-isotope analysis to infer diet composition and foraging ecology of Adélie penguins Pygoscelis adeliae. Marine Ecology Progress Series 355, 297–307, doi: 10.3354/meps07235.

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. & Wienecke B. 2015. Pollution, habitat loss, fishing, and climate change as critical threats to penguins. Conservation Biology 29, 31–41, doi: 10.1111/cobi.12349.

Trivelpiece W.Z., Hinke J.T., Miller A.K., Reiss C.S., Trivelpiece S.G. & Watters G.M. 2011. Variability in krill biomass links harvesting and climate warming to penguin population changes in Antarctica. Proceedings of the National Academy of Sciences of the United States of America 108, 7625–7628, doi: 10.1073/pnas.1016560108.

Trivelpiece W.Z., Trivelpiece S.G. & Volkman N.J. 1987. Ecological segregation of Adelie, gentoo, and chinstrap penguins at King George Island, Antarctica. Ecology 68, 351–361, doi: 10.2307/1939266.

Vaughan D.G., Marshall G.J., Connolley W.M., Parkinson C., Mulvaney R., Hodgson D.A., King J.C., Pudsey C.J. & Turner J. 2003. Recent rapid regional climate warming on the Antarctic Peninsula. Climatic Change 60, 243–274, doi: 10.1023/A:1026021217991.

Waluda C.M., Collins M.A., Black A.D., Staniland I.J. & Trathan P.N. 2010. Linking predator and prey behaviour: contrasts between Antarctic fur seals and macaroni penguins at South Georgia. Marine Biology 157, 99–112, doi: 10.1007/s00227-009-1299-6.

Whitehead O.T., Connan M., Ropert-Coudert Y. & Ryan P.T. 2017. Subtle but significant segregation in the feeding ecology of sympatric penguins during the critical pre-moult period. Marine Ecology Progress Series 565, 227–236, doi: 10.3354/meps12017.

Williams T.D., Briggs D.R., Croxall J.P., Naito Y. & Kato A. 1992. Diving pattern and performance in relation to foraging ecology in the gentoo penguin, Pygoscelis papua. Journal of Zoology 227, 211–230, doi: 10.1111/j.1469-7998.1992.tb04818.x.

Wilson R.P., Pütz K., Peters G., Culik B., Scolaro J.A., Charrassin J.-B. & Ropert-Coudert Y. 1997. Long-term attachment of transmitting and recording devices to penguins and other seabirds. Wildlife Society Bulletin 25, 101–106.

Wilson R.P. & Wilson M.P. 1989. Tape: a package attachment technique for penguins. Wildlife Society Bulletin 17, 77–79, doi: 10.2307/3801309.
How to Cite
NarvestadA., KovacsK. M., LydersenC., & LowtherA. D. (2022). Foraging behaviour of sympatrically breeding macaroni (<em>Eudyptes chrysolophus</em>) and chinstrap (<em>Pygoscelis antarcticus</em&gt;) penguins at Bouvetøya, Southern Ocean. Polar Research, 41.
Research Articles

Most read articles by the same author(s)