Marine Biology

, 163:19 | Cite as

Habitat use and diving behaviour of macaroni Eudyptes chrysolophus and eastern rockhopper E. chrysocome filholi penguins during the critical pre-moult period

  • T. O. WhiteheadEmail author
  • A. Kato
  • Y. Ropert-Coudert
  • P. G. Ryan
Original paper


After the breeding season, penguins must replenish body condition and accumulate sufficient energy stores before their annual moult ashore; failure to do so may lead to starvation. Knowing where and how adult penguins find adequate resources during this energy-intensive stage is vital to understanding their susceptibility to ecosystem changes. GPS and TDR loggers were used to track movements and record diving behaviour of macaroni Eudyptes chrysolophus and eastern rockhopper E. chrysocome filholi penguins from Marion Island (46°S, 37°E) during the pre-moult foraging trip in 2012, 2013 and 2014. Both species consistently travelled in a southerly direction to forage in cooler (~3.5 °C) Antarctic Zone waters south of the Antarctic Polar Front where they associated with mesoscale eddies and sub-mesoscale filaments. Dives were predominantly to depths of 30 to 60 m, but macaroni penguins dived deeper more often. Mean trip durations of both species were similar (33 ± 6 days), but maximum foraging ranges of macaroni penguins (903 ± 165 km) were greater than eastern rockhopper penguins (696 ± 152 km). Spatial overlap of core foraging areas between species was high, but a 2- to 3-week difference in departure dates reduced potential interspecific competition at sea. Trip durations were longer in 2014 compared to 2013, when decreased productivity may have reduced prey availability, forcing penguins to remain longer at sea. Continued monitoring is vital to understand how crested penguins at Marion Island adapt to the predicted southward shift of major frontal boundaries.


Travel Speed Trip Duration Diving Behaviour Dive Depth Eddy Kinetic Energy 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



We thank Maëlle Connan, Ben Dilley, Delia Davies, Stefan Schoombie and Kim Stevens for assistance with fieldwork; Francis Crenner, Matthieu Brucker, Nicolas Chatelain for customising loggers; Pierre Pistorius for providing additional TDR loggers; and Maëlle Connan and three anonymous reviewers for valuable comments on the manuscript. The South African National Antarctic Programme provided funding and logistical support; additional funding was provided by the SA Department of Science and Technology, National Research Foundation, University of Cape Town and World Wide Fund for Nature, Australia.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

All procedures performed in this study were in accordance with the ethical standards of the Faculty of Science Animal Ethics Committee (SFAEC), University of Cape Town (2013/V5/NEW).

Supplementary material

227_2015_2794_MOESM1_ESM.pdf (2.7 mb)
Supplementary material 1 (PDF 2809 kb)


  1. Adams NJ, Brown CR (1990) Energetic of molt in penguins. In: Davis LS, Darby JT (eds) Penguin biology. Academic Press, London, pp 297–315Google Scholar
  2. Ansorge IJ, Lutjeharms JRE (2005) Direct observations of eddy turbulence at a ridge in the Southern Ocean. Geophys Res Lett 32:1–4CrossRefGoogle Scholar
  3. Ansorge IJ, Pakhomov EA, Kaehler S, Lutjeharms JRE, Durgadoo JV (2009) Physical and biological coupling in eddies in the lee of the South-West Indian Ridge. Polar Biol 33:747–759CrossRefGoogle Scholar
  4. Ansorge IJ, Jackson JM, Reid K, Durgadoo JV, Swart S, Eberenz S (2015) Evidence of a southward eddy corridor in the south-west Indian ocean. Deep Sea Res II 119:69–76CrossRefGoogle Scholar
  5. Atkinson A, Siegel V, Pakhomov EA, Rothery P, Loeb V, Ross RM, Quetin LB, Schmidt K, Fretwell P, Murphy EJ, Tarling GA, Fleming AH (2008) Oceanic circumpolar habitats of Antarctic krill. Mar Ecol Prog Ser 362:1–23CrossRefGoogle Scholar
  6. Bernard ATF, Ansorge IJ, Froneman PW, Lutjeharms JRE, Bernard KS, Swart NS (2007) Entrainment of Antarctic euphausiids across the Antarctic Polar Front by a cold eddy. Deep Sea Res I 54:1841–1851CrossRefGoogle Scholar
  7. BirdLife International (2015) IUCN Red List for birds. on 14/10/2015
  8. Bon C, Della Penna A, d’Ovidio F, Arnould J, Poupart T, Bost CA (2015) Influence of oceanographic structures on foraging strategies: macaroni penguins at Crozet Islands. Mov Ecol 3:32–42CrossRefGoogle Scholar
  9. Bost CA, Georges JY, Guinet C, Cherel Y, Pütz K, Charrassin JB, Handrich Y, Zorn T, Lage J, Le Maho Y (1997) Foraging habitat and food intake of satellite-tracked king penguins during the austral summer at Crozet Archipelago. Mar Ecol Prog Ser 150:21–33CrossRefGoogle Scholar
  10. Bost CA, Cotté C, Bailleul F, Cherel Y, Charrassin JB, Guinet C, Ainley DG, Weimerskirch H (2009) The importance of oceanographic fronts to marine birds and mammals of the Southern Oceans. J Mar Syst 78:363–376CrossRefGoogle Scholar
  11. Brown CR (1986) Feather growth, mass loss and duration of moult in macaroni and rockhopper penguins. Ostrich 57:180–184CrossRefGoogle Scholar
  12. Brown CR (1987) Traveling speed and foraging range of macaroni and rockhopper penguins at Marion Island. J Field Ornithol 58:118–125Google Scholar
  13. Brown CR (1989) Energy requirements and food consumption of Eudyptes penguins at the Prince Edward Islands. Antarct Sci 1:15–21CrossRefGoogle Scholar
  14. Calenge C (2006) The package ‘‘adehabitat’’ for the R software: a tool for the analysis of space and habitat use by animals. Ecol Model 197:516–519CrossRefGoogle Scholar
  15. Calenge C, Dray S, Royer-Carenzi M (2009) The concept of animals’ trajectories from a data analysis perspective. Ecol Inform 4:34–41CrossRefGoogle Scholar
  16. Charrassin JB, Bost CA (2001) Utilization of the oceanic habitat by king penguins over the annual cycle. Mar Ecol Prog Ser 221:285–297CrossRefGoogle Scholar
  17. Cherel Y, Charrassin JB, Challet E (1994) Energy and protein requirements for molt in the king penguin Aptenodytes patagonicus. Am J Physiol 266:R1182–R1188Google Scholar
  18. Cherel Y, Hobson KA, Guinet C, Vanpe C (2007) Stable isotopes document seasonal changes in trophic niches and winter foraging individual specialisation in diving predators from the Southern Ocean. J Anim Eco 76:826–836CrossRefGoogle Scholar
  19. Cotté C, Park YH, Guinet C, Bost CA (2007) Movements of foraging king penguins through marine mesoscale eddies. Proc R Soc Lond B 274:2385–2391CrossRefGoogle Scholar
  20. Cotté C, d’Ovidio F, Dragon AC, Guinet C, Lévy M (2015) Flexible preference of southern elephant seals for distinct mesoscale features within the Antarctic Circumpolar Current. Prog Oceanogr 131:46–58CrossRefGoogle Scholar
  21. Crawford RJM, Cooper J, Dyer BM (2003a) Population of the macaroni penguin Eudyptes chrysolophus at Marion Island, 1994/95–2002/03, with information on breeding and diet. Afr J Mar Sci 25:475–486CrossRefGoogle Scholar
  22. Crawford RJM, Cooper J, Dyer BM, Greyling MD, Klages NTW, Nel DC, Nel JL, Petersen SL, Wolfaardt AC (2003b) Decrease in numbers of the eastern rockhopper penguin Eudyptes chrysocome filholi at Marion Island, 1994/95–2002/03. Afr J Mar Sci 25:487–498CrossRefGoogle Scholar
  23. Crawford RJM, Whittington PA, Upfold L, Ryan PG, Petersen SL, Dyer BM, Cooper J (2009) Recent trends in numbers of four species of penguins at the Prince Edward Islands. Afr J Mar Sci 31:419–426CrossRefGoogle Scholar
  24. de Brooke ML (2004) The food consumption of the world’s seabirds. Proc R Soc Lond B 271:S246–S248CrossRefGoogle Scholar
  25. Development Core Team R (2015) R: a language and environment for statistical computing. R Foundation for Statistical Analysis, ViennaGoogle Scholar
  26. d’Ovidio F, López C, Hernández-García E, Fernández V (2004) Mixing structures in the Mediterranean sea from finite-size Lyapunov exponents. Geophys Res Lett 31:L17203Google Scholar
  27. Durgadoo JV, Ansorge IJ, De Cuevas BA, Lutjeharms JRE, Coward AC (2011) Decay of eddies at the South-West Indian Ridge. S Afr J Sci 107:1–10CrossRefGoogle Scholar
  28. Fieberg J, Kochanny CO (2005) Quantifying home-range overlap: the importance of the utilization distribution. J Wildl Manage 69:1346–1359CrossRefGoogle Scholar
  29. Frenger (2013) On Southern Ocean eddies and their impacts on biology and the atmosphere. Ph.D. Thesis, University in Zurich, ZurichGoogle Scholar
  30. Gille ST (2002) Warming of the Southern Ocean since the 1950s. Science 295:1275–1277CrossRefGoogle Scholar
  31. Green JA, Boyd IL, Woakes AJ, Warren NL, Butler PJ (2005) Behavioural flexibility during year-round foraging in macaroni penguins. Mar Ecol Prog Ser 296:183–196CrossRefGoogle Scholar
  32. Green JA, Boyd IL, Woakes AJ, Warren NL, Butler PJ (2009a) Evaluating the prudence of parents: daily energy expenditure throughout the annual cycle of a free-ranging bird, the macaroni penguin Eudyptes chrysolophus. J Avian Biol 40:529–538CrossRefGoogle Scholar
  33. Green JA, Wilson RP, Boyd IL, Woakes AJ, Green CJ, Butler PJ (2009b) Tracking macaroni penguins during long foraging trips using ‘behavioural geolocation’. Polar Biol 32:645–653CrossRefGoogle Scholar
  34. Horswill C (2015) The relative importance of opposing drivers in determining population change in macaroni penguins Eudyptes chrysolophus. Ph.D. thesis, University of Glasgow, GlasgowGoogle Scholar
  35. Hull CL (2000) Comparative diving behaviour and segregation of the marine habitat by breeding Royal Penguins, Eudyptes schlegeli, and eastern Rockhopper Penguins, Eudyptes chrysocome filholi, at Macquarie Island. Can J Zool 78:333–345CrossRefGoogle Scholar
  36. Hull CL, Hindell M, Le Mar K, Scofield P, Wilson J, Lea MA (2004) The breeding biology and factors affecting reproductive success in rockhopper penguins Eudyptes chrysocome at Macquarie Island. Polar Biol 27:711–720CrossRefGoogle Scholar
  37. Hulley PA (1990) Family Myctophidae. In: Gon O, Heemstra PC (eds) Fish of the Southern Ocean. J.L.B. Smith Institute of Ichthyology, Grahamstown, pp 146–178Google Scholar
  38. Hunt BPV, Pakhomov EA, McQuaid CD (2001) Short-term variation and long-term changes in the oceanographic environment and zooplankton community in the vicinity of a sub-Antarctic archipelago. Mar Biol 138:369–381CrossRefGoogle Scholar
  39. Jaeger A, Cherel Y (2011) Isotopic investigation of contemporary and historic changes in penguin trophic niches and carrying capacity of the Southern Indian Ocean. PLoS One 6:e16484CrossRefGoogle Scholar
  40. Kahru M, Mitchell BG, Gille ST, Hewes CD, Holm-Hansen O (2007) Eddies enhance biological production in the Weddell-Scotia Confluence of the Southern Ocean. Geophys Res Lett 34:L14603CrossRefGoogle Scholar
  41. Kareiva P, Odell G (1987) Swarms of predators exhibit ‘preytaxis’ if individual predators use area-restricted search. Am Nat 130:233–270CrossRefGoogle Scholar
  42. Keymer IF, Malcolm HM, Hunt A, Horsley DT (2001) Health evaluation of penguins (Sphenisciformes) following mortality in the Falklands (South Atlantic). Dis Aquat Org 45:159–169CrossRefGoogle Scholar
  43. Lapeyre G, Klein P (2006) Impact of the small-scale elongated filaments on the oceanic vertical pump. J Mar Res 64:835–851CrossRefGoogle Scholar
  44. Lavielle M (2005) Using penalized contrasts for the change-point problem. Signal Process 85:1501–1510CrossRefGoogle Scholar
  45. Lévy M, Ferrari R, Franks PJS, Martin AP, Rivière P (2012) Bringing physics to life at the submesoscale. Geophys Res Lett 39:L14602Google Scholar
  46. Lowther AD, Lydersen C, Biuw M, De Bruyn PJN, Hofmeyr GJG, Kovacs KM (2014) Post-breeding at-sea movements of three central-place foragers in relation to submesoscale fronts in the Southern Ocean around Bouvetøya. Antarct Sci 26:533–544CrossRefGoogle Scholar
  47. Ludynia K, Dehnhard N, Poisbleau M, Demongin L, Masello JF, Voigt CC, Quillfeldt P (2013) Sexual segregation in rockhop- per penguins during incubation. Anim Behav 85:255–267CrossRefGoogle Scholar
  48. MacArthur RH, Pianka ER (1966) On optimal use of a patchy environment. Am Nat 100:603–609CrossRefGoogle Scholar
  49. Massie PP, McIntyre T, Ryan PG, Bester MN, Bornemann H, Ansorge IJ (2015) The role of eddies in the diving behaviour of female southern elephant seals. Polar Biol. doi: 10.1007/s00300-015-1782-0 Google Scholar
  50. Mélice JL, Lutjeharms JRE, Rouault M, Ansorge IJ (2003) Sea-surface temperatures at the sub-Antarctic islands Marion and Gough during the past 50 years. S Afr J Sci 99:363–366Google Scholar
  51. Nel DC, Lutjeharms JRE, Pakhomov EA, Ansorge IJ, Ryan PG, Klages NTW (2001) Exploitation of mesoscale oceanographic features by grey-headed albatrosses Thalassarche chrysostoma in the southern Indian Ocean. Mar Ecol Prog Ser 217:15–26CrossRefGoogle Scholar
  52. Nordstrom CA, Battaile BC, Cotté C, Trites AW (2013) Foraging habitats of lactating northern fur seals are structured by thermocline depths and submesoscale fronts in the eastern Bering Sea. Deep Sea Res II 88:78–96CrossRefGoogle Scholar
  53. Pakhomov EA, Froneman PW (2000) Composition and spatial variability of macroplankton and micronekton within the Antarctic Polar Frontal Zone of the Indian Ocean during austral autumn 1997. Polar Biol 23:410–419CrossRefGoogle Scholar
  54. Pakhomov EA, Perisinotto R, McQuaid CD (1994) Comparative structure of the macrozooplankton/micronekton communities of the Subtropical and Antarctic Polar Fronts. Mar Ecol Prog Ser 111:155–169CrossRefGoogle Scholar
  55. Péron C, Weimerskirch H, Bost CA (2012) Projected pole- ward shift of king penguins’ (Aptenodytes patagonicus) foraging range at the Crozet Islands, southern Indian Ocean. Proc R Soc Lond B 279:2515–2523CrossRefGoogle Scholar
  56. Phillips RA, Croxall JP, Silk JRD, Briggs DR (2008) Foraging ecology of albatrosses and petrels from South Georgia: two decades of insights from tracking technologies. Aquat Conserv Mar Freshw Ecosyst 17:S6–S21CrossRefGoogle Scholar
  57. Pichegru L, Ropert-Coudert Y, Kato A, Takahashi A, Dyer BM, Ryan PG (2011) Diving patterns of female macaroni penguins breeding on Marion Island, South Africa. Polar Biol 34:945–954CrossRefGoogle Scholar
  58. Pinheiro J, Bates D, DebRoy S, Sarkar D, the R Development Core Team (2013) nlme: Linear and nonlinear mixed effects models. R package version 3.1-113. CRAN package repository. Accessed 23 Feb 2014Google Scholar
  59. Poisbleau M, Demongin L, van Noordwijk HJ, Strange IJ, Quillfeldt P (2010) Sexual dimorphism and use of morphological measurements to sex adults, immatures and chicks of rockhopper Penguins. Ardea 98:217–224CrossRefGoogle Scholar
  60. Pütz K, Raya Rey A, Huin N, Schiavini A, Pütz A, Lüthil BH (2006) Diving characteristics of southern rockhopper penguins (Eudyptes c. chrysocome) in the Southwest Atlantic. Mar Biol 149:125–137CrossRefGoogle Scholar
  61. Raya Rey A, Pütz K, Luna-Jorquera G, Lüthi B, Schiavini A (2009) Diving patterns of breeding female rockhopper penguins (Eudyptes chrysocome): Noir Island, Chile. Polar Biol 32:561–568CrossRefGoogle Scholar
  62. Ropert-Coudert Y, Kato A, Chiaradia A (2009) Impact of small-scale environmental perturbations on local marine food resources: a case study of a predator, the little penguin. Proc R Soc B 276:4105–4109CrossRefGoogle Scholar
  63. Ryan P (2009) The seabirds of the Conrad Rise, Southern Ocean. Bull Afr Bird Club 16:67–77Google Scholar
  64. Ryan PG, Bester MN (2008) Pelagic predators. In: Chown SL, Froneman PW (eds) The Prince Edward Islands—land-sea interactions in a changing ecosystem. Sun Press, Stellenbosch, pp 121–164Google Scholar
  65. Sakamoto KQ, Sato K, Ishizuka M, Watanuki Y, Takahashi A, Daunt F, Wanless S (2009) Can ethograms be automatically generated using body acceleration data from free-ranging birds? PLoS One 4:e5379CrossRefGoogle Scholar
  66. Santora JA, Sydeman WJ, Schroeder ID, Reiss CS, Wells BK, Field JC, Cossio AM, Loeb VJ (2012) Krill space: a comparative assessment of mesoscale structuring in polar and temperate marine ecosystems. ICES J Mar Sci 69:1317–1327CrossRefGoogle Scholar
  67. Sato K, Charrassin JB, Bost CA, Naito Y (2004) Why do macaroni penguins choose shallow body angles that result in longer descent and ascent durations? J Exp Biol 207:4057–4065CrossRefGoogle Scholar
  68. Schiavini A, Raya Rey A (2004) Long days, long trips: foraging ecology of female rockhopper penguin brooding chicks at Tierra del Fuego. Mar Ecol Prog Ser 275:251–262CrossRefGoogle Scholar
  69. Sokolov S, Rintoul SR (2007) On the relationship between fronts of the Antarctic Circumpolar Current and surface chlorophyll concentrations in the Southern Ocean. J Geophys Res 112:C07030CrossRefGoogle Scholar
  70. Strange IJ (1982) Breeding ecology of the Rockhopper penguin (Eudyptes crestatus) in the Falkland Islands. Gerfaut 72:137–188Google Scholar
  71. Strass VH, Naveira Garabato AC, Pollard RT, Fischer HI, Hense I, Allen JT, Read JF, Leach H, Smetacek V (2002) Mesoscale frontal dynamics: shaping the environment of primary production in the Antarctic Circumpolar Current. Deep Sea Res II 49:3735–3769CrossRefGoogle Scholar
  72. Swart S, Speich S, Ansorge IJ, Lutjeharms JRE (2010) An altimetry-based gravest empirical mode south of Africa: 1. Development and validation. J Geophys Res 115:C03002Google Scholar
  73. Thiebot JB, Cherel Y, Trathan PN, Bost CA (2012) Coexistence of oceanic predators on wintering areas explained by population-scale foraging segregation in space or time. Ecology 93:122–130CrossRefGoogle Scholar
  74. Thiebot JB, Cherel Y, Crawford RJM, Makhado AB, Trathan PN, Pinaud D, Bost CA (2013) A space oddity: geographic and specific modulation of migration in Eudyptes penguins. PLoS One 8:e71429CrossRefGoogle Scholar
  75. Thiebot JB, Cherel Y, Acqueberge M, Prudor A, Trathan PN, Bost CA (2014) Adjustment of pre-moult foraging strategies in macaroni penguins Eudyptes chrysolophus according to locality, sex and breeding status. Ibis 156:511–522CrossRefGoogle Scholar
  76. Trathan PN, Bishop C, Maclean G, Brown P, Fleming A, Collins MA (2008) Linear tracks characterise penguin foraging pathways. Mar Ecol Prog Ser 370:285–294CrossRefGoogle Scholar
  77. Tremblay Y, Cherel Y (2003) Geographic variation in the foraging behaviour, diet and chick growth of rockhopper penguins. Mar Ecol Prog Ser 251:279–297CrossRefGoogle Scholar
  78. Trivelpiece WZ, Trivelpiece SG, Volkman NJ (1987) Ecological segregation of Adelie, Gentoo and Chinstrap penguins at King George Island, Antarctica. Ecology 68:351–361CrossRefGoogle Scholar
  79. Waluda CM, Collins MA, Black AD, Staniland IJ, Trathan PN (2010) Linking predator and prey behaviour: contrasts between Antarctic fur seals and macaroni penguins at South Georgia. Mar Biol 157:99–112CrossRefGoogle Scholar
  80. Warham J (1974) The Fiordland crested penguin. Ibis 116:1–27CrossRefGoogle Scholar
  81. Warham J (1975) The crested penguins. In: Stonehouse B (ed) The biology of penguins. Macmillan, LondonGoogle Scholar
  82. Weavers BW (1992) Seasonal foraging ranges and travels at sea of little penguins Eudyptula minor, determined by radiotracking. Emu 91:302–317CrossRefGoogle Scholar
  83. Williams TD (1995) The Penguins. Oxford University Press, OxfordGoogle Scholar
  84. Williams TD, Croxall JP (1991) Annual variation in breeding biology of macaroni penguins, Eudyptes chrysolophus, at Bird Island, South Georgia. J Zool Lond 223:189–202CrossRefGoogle Scholar
  85. Wilson RP, Putz K, Peters G, Culik B, Scolaro JA, Charrassin JB, Ropert-Coudert Y (1997) Long-term attachment of transmitting and recording devices to penguins and other seabirds. Wildl Soc Bull 25:101–106Google Scholar
  86. Wood SN (2006) Generalized additive models: an introduction with R. Chapman & Hall, Boca RatonGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2016

Authors and Affiliations

  • T. O. Whitehead
    • 1
    Email author
  • A. Kato
    • 2
    • 3
  • Y. Ropert-Coudert
    • 1
    • 2
    • 3
  • P. G. Ryan
    • 1
  1. 1.Percy FitzPatrick Institute of African Ornithology, DST-NRF Centre of ExcellenceUniversity of Cape TownRondeboschSouth Africa
  2. 2.Station d’Écologie de Chizé-La Rochelle, UMR 7372, CNRSCentre d’Etudes Biologiques de ChizéVilliers-en-BoisFrance
  3. 3.UMR 7178CNRSStrasbourgFrance

Personalised recommendations