King Penguin populations increase on South Georgia but explanations remain elusive

Original Paper

Abstract

While dramatic increases in populations of King Penguins (Aptenodytes patagonicus) have been documented throughout their range, population changes on the island of South Georgia have not been assessed. We reconstructed time series of population size for six major colonies across South Georgia using historical data stretching back to 1883 and new population estimates derived from direct on-the-ground censuses and oblique, high-resolution digital photographs. We find evidence for a significant increase in the population of King Penguins at all colonies examined over the 124 years of available survey data. We discuss our findings in the context of four established hypotheses explaining King Penguin population growth: (1) favorable changes in the pelagic food web; (2) climate forcing; (3) greater availability of breeding habitat; and (4) the cessation of harvesting. While we do find evidence that glacial retreat may have increased suitable breeding habitat at some colonies and facilitated population expansion, glacial retreat is not associated with all of South Georgia’s growing populations. Local anomalies in sea surface temperature have increased in parallel with King Penguin population growth rate, suggesting that climate forcing may contribute to colony growth, but a complete explanation for the island’s rapidly growing King Penguin population remains unclear.

Keywords

Southern Ocean High-resolution photography Population estimate Glacial retreat Climate forcing 

Notes

Acknowledgements

We gratefully acknowledge assistance from the US National Science Foundation Office of Polar Programs (Award No. NSF/OPP-1255058) and the Explorer’s Club. The authors would also like to thank S. Poncet for contributing data, and C. Black, A. Borowicz, G. Clucas, G. Humphries, M. Lynch, B. Goncalves, C. O’Leary, M. Schrimpf, and C. Youngflesh for help with data collection. Additionally, we would like to thank C. Bell, R. Biondo, L. Caliguri, K. Gregory, V. Kennelly, S. Vincent, A. West, and T. Wong for help with counting panoramas. We gratefully acknowledge the support of the One Ocean Expeditions, Cheeseman’s Ecological Safaris, and Quark Expeditions and their field staff for their support in the field.

Compliance with ethical standards

Ethical approval

All research was conducted with under the approval of the Stony Brook University Institutional Animal Care and Use Committee application 2011-1881-FAR-6.19.20-BI and South Georgia and South Sandwich Islands Permit #2015/018, and 2016/035.

Supplementary material

300_2018_2271_MOESM1_ESM.txt (1 kb)
Supplementary material 1 (TXT 1 kb)
300_2018_2271_MOESM2_ESM.csv (7 kb)
Supplementary material 2 (CSV 6 kb)

References

  1. Bannister JL (1964) Whaling stations in South Georgia. Polar Rec 12:207–209.  https://doi.org/10.1017/S0032247400054723 CrossRefGoogle Scholar
  2. Bart J, Droege S, Geissler P, Peterjohn B, Ralph CJ (2004) Density estimation in wildlife surveys. Wildl Soc Bull 32:1242–1247.  https://doi.org/10.2193/0091-7648(2004)032[1242:DEIWS]2.0.CO;2 CrossRefGoogle Scholar
  3. Bingham M (1998) The distribution, abundance and population trends of gentoo, rockhopper and king penguins in the Falkland Islands. Oryx 32:223–232.  https://doi.org/10.1046/j.1365-3008.1998.d01-39.x CrossRefGoogle Scholar
  4. BirdLife International (2017) Aptenodytes patagonicus. (amended version published in 2016) The IUCN Red List of Threatened Species 2017Google Scholar
  5. Boersma PD (2008) Penguins as marine sentinels. Bioscience 58:597–607.  https://doi.org/10.1641/B580707 CrossRefGoogle Scholar
  6. Bost C-A, Delord K, Barbraud C, Cherel Y, Pütz K, Cotté C, Péron C, Weimerskirch H (2013) King penguin (Aptenodytes patagonicus). In: Borboroglu PG, Boersma D (eds) Penguins: natural history and conservation. University of Washington Press, Seattle, pp 7–21Google Scholar
  7. Boyin H, Banzon VF, Freeman E, Lawrimore J, Liu W, Peterson TC, Smith TM, Thorne PW, Woodruff SD, Zhang H-M (2015) Extended reconstructed sea surface temperature (ERSST), Version 4. NOAA National Centers for Environmental Information.  https://doi.org/10.7289/v5kd1vvf
  8. Bried J, Jouventin P (2002) Site and mate choice in seabirds: an evolutionary approach. In: Schreiberm EA, Burger J (eds) Biology of marine birds. CRC Press, New York, pp 263–306Google Scholar
  9. British Antarctic Survey (2018) sg_landsat_backdrop. http://www.add.scar.org/geoserver/web/. Accessed 8 January 2018
  10. Budd GM (1970) Further population growth in the Heard Island King Penguins. Auk 87:366–367.  https://doi.org/10.2307/4083932 CrossRefGoogle Scholar
  11. Budd GM (2000) Changes in Heard Island glaciers, King Penguins and fur seals since 1947. In: Banks MR, Brown MJ (eds) Heard Island Papers, Pap and Proc R Soc of Tasmania, vol 133, pp 47–60Google Scholar
  12. Chamaillé-James S, Guinet C, Nicoleau F, Argentier M (2000) A method to assess population changes in King Penguins: the use of a geographical information system to estimate area-populations relationships. Polar Biol 23:545–549.  https://doi.org/10.1007/s003000000119 CrossRefGoogle Scholar
  13. Chapman EW, Hofmann EE, Patterson DL, Ribic CA, Fraser WR (2011) Marine and terrestrial factors affecting Adélie Penguin Pygocelis adeliae chick growth and recruitment off the western Antarctic Peninsula. Mar Ecol Prog Ser 436:273–289.  https://doi.org/10.3354/meps09242 CrossRefGoogle Scholar
  14. Clark G (1984) Report to the British Antarctic survey: totorore expedition to South Georgia, 13th August to 17th October 1984Google Scholar
  15. Clark G (1985) Report to the British Antarctic survey: totorore expedition to South Georgia, 11 June to 12 September 1985Google Scholar
  16. Clarke A, Croxall JP, Poncet S, Martin AR, Burton R (2012) Important bird areas: South Georgia. British Birds 105:118–144Google Scholar
  17. Clucas GV, Younger JL, Kao D, Rogers AD, Handley J, Miller GD, Jouventin P, Nolan P, Gharbi K, Miller KJ, Hart T (2016) Dispersal in the sub-Antarctic: King Penguins show remarkably little population genetic differentiation across their range. BMC Evolut Biol 16:211.  https://doi.org/10.1186/s12862-016-0784-z CrossRefGoogle Scholar
  18. Collins MA, Xavier JC, Johnston NM, North AW, Enderlein P, Tarling GA, Waluda CM, Hawker EJ, Cunningham NJ (2008) Patterns in the distribution of myctophid fish in the northern Scotia Sea ecosystem. Polar Biol 31:837–851.  https://doi.org/10.1007/s00300-008-0423-2 CrossRefGoogle Scholar
  19. Condit R, Le Boeuf B, Morris PA, Sylvan M (2007) Estimating population size in asynchronous aggregations: a Bayesian approach and test with elephant seal censuses. Mar Mammal Sci 23:834–855CrossRefGoogle Scholar
  20. Conroy JWH, White MG (1973) The breeding status of the King Penguin (Aptenodytes patagonicus). Br Antarct Surv Bull 32:31–40Google Scholar
  21. Constable AJ et al (2014) Climate change and Southern Ocean ecosystems I: how changes in physical habitats directly affect marine biota. Glob Change Biol 20:3004–3025.  https://doi.org/10.1111/gcb.12623 CrossRefGoogle Scholar
  22. Cook AJ, Poncet S, Paul A, Cooper R, Herbert DJ, Christie D (2010) Glacial retreat on South Georgia and implications for the spread of rats. Antarct Sci 22:255–263.  https://doi.org/10.1017/s0954102010000064 CrossRefGoogle Scholar
  23. Curran MAJ, van Ommen TD, Morgan VI, Phillips KL, Palmer AS (2003) Ice core evidence for Antarctic Sea ice declines since the 1950s. Science 302:1203–1206.  https://doi.org/10.1126/science.1087888 CrossRefPubMedGoogle Scholar
  24. Debenham F (1945) The voyage of Captain Bellingshausen to the Antarctic Seas, 1819–1821, vol 1–2. Hakluyt Soc, LondonGoogle Scholar
  25. Delord K, Barbraud C, Weimerskirch H (2004) Long-term trends in the population size of King Penguins at Crozet archipelago: environmental variability and density dependence? Polar Biol 27:793–800.  https://doi.org/10.1007/s00300-004-0651-z CrossRefGoogle Scholar
  26. Ellis JC (2005) Marine birds on land: a review of plant biomass, species richness, and community composition in seabird colonies. Plant Ecol 181:227–241.  https://doi.org/10.1007/s11258-005-7147-y CrossRefGoogle Scholar
  27. Farmer A, Durbain F (2006) Estimating shorebird numbers at migration stopover sites. Condor 108:792–807.  https://doi.org/10.1650/0010-5422(2006)108[792:esnams]2.0.co;2 CrossRefGoogle Scholar
  28. Frederick PC, Heath JA, Bennetts R, Hafner H (2006) Estimating nests not present at the time of breeding surveys: an important consideration in assessing nesting populations. J Field Ornithol 77:212–219.  https://doi.org/10.1111/j.1557-9263.2006.00043.x CrossRefGoogle Scholar
  29. Gales R, Pemberton D (1988) Recovery of the King Penguin, Aptenodytes-patagonicus, population on Heard Island. Wildl Res 15:579–585.  https://doi.org/10.1071/wr9880579 CrossRefGoogle Scholar
  30. Gille ST (2002) Warming of the Southern Ocean Since the1950s. Science 295:1275–1277.  https://doi.org/10.1126/science.1065863 CrossRefPubMedGoogle Scholar
  31. Gutt J, Bertler N, Bracegirdle TJ, Buschmann A, Comiso J, Hosie G, Isla E, Schloss IR, Smith CR, Tournadre J, Xavier JC (2015) The Southern Ocean ecosystem under multiple climate change stresses—an integrated circumpolar assessment. Glob Change Biol 21:1434–1453.  https://doi.org/10.1111/gcb.12794 CrossRefGoogle Scholar
  32. Headland R (1984) The Island of South Georgia. Cambridge University Press, CambridgeGoogle Scholar
  33. Hill SL, Reid K, Thorpe SE, Hinke J, Watters GM (2007) A compilation of parameters for ecosystem dynamics models of the Scotia Sea-Antarctic Peninsula region. CCAMLR Sci 14:1–25Google Scholar
  34. Jacka TH, Budd WF (1998) Detection of temperature and sea-ice extent changes in the Southern Ocean. Ann Glaciol 27:553–559.  https://doi.org/10.3189/1998aog27-1-553-559 CrossRefGoogle Scholar
  35. Jouventin P, Capdeville D, Cuenot-Chaillet F, Boiteau C (1994) Exploitation of pelagic resources by a non-flying seabird: satellite tracking of the King Penguin throughout the breeding cycle. Mar Ecol Prog Ser 106:11–19.  https://doi.org/10.3354/meps106011 CrossRefGoogle Scholar
  36. Juáres MA, Negrete J, Mennucci JA, Perchivale PJ, Santos M, Moreira E, Coria NR (2014) Further evidence of King Penguins’ breeding range extension in the South Shetland Islands? Antarct Sci 26:261–262.  https://doi.org/10.1017/s0954102013000655 CrossRefGoogle Scholar
  37. Juáres MA, Ferrer F, Coria NR, Santos MM (2017) Breeding events of King Penguin at the South Shetland Islands: has it come to stay? Polar Biol 40:457–461.  https://doi.org/10.1007/s00300-016-1947-5 CrossRefGoogle Scholar
  38. Lancraft TM, Torres JJ, Hopkins TL (1989) Micronekton and macrozooplankton in the open waters near ice edge zones (AMERIEZ 1983 and 1986). Polar Biol 9:225–233.  https://doi.org/10.1007/bf00263770 CrossRefGoogle Scholar
  39. Laws RM (1973) Population increases of fur seals at South Georgia. Polar Rec 16:856–858.  https://doi.org/10.1017/s003224740006397x CrossRefGoogle Scholar
  40. LeBohec C, Gauthier-Clerc M, Le Maho Y (2005) The adaptive significance of crèches in the King Penguin. Animal Behav 70:527–538.  https://doi.org/10.1016/j.anbehav.2004.11.012 CrossRefGoogle Scholar
  41. LeBohec C, Durant JM, Gauthier-Clerc M, Stenseth NC, Park Y-H, Pradel R, Grémillet D, Gendner J-P, Le Maho Y (2008) King penguin population threatened by Southern Ocean warming. Proc Natl Acad Sci USA 105:2493–2497.  https://doi.org/10.1073/pnas.0712031105 CrossRefGoogle Scholar
  42. Lewis Smith RI, Tallowin JRB (1979) The distribution and size of king penguin rookeries on South Georgia. Br Antarct Surv Bull 49:259–276Google Scholar
  43. Murphy RC (1915) Cruising in the South Atlantic. Brooklyn Museum Bulletin 1:83–110Google Scholar
  44. Murphy EJ, Watkins JL, Reid K, Trathan PN, Everson I, Croxall JP, Priddle J, Brandon MA, Brierley AS, Hofmann E (2007) Interannual variability of the South Georgia marine ecosystem: biological and physical sources of variation in the abundance of krill. Fish Oceanogr 7:381–390.  https://doi.org/10.1046/j.1365-2419.1998.00081.x CrossRefGoogle Scholar
  45. Olsson O, North AW (1997) Diet of the King Penguin Aptenodytes patagonicus during three summers at South Georgia. Ibis 139:504–512.  https://doi.org/10.1111/j.1474-919x.1997.tb04666.x CrossRefGoogle Scholar
  46. Pakhomov EA, Perissinotto R, McQuaid CD (1996) Prey composition and daily rations of myctophid fishes in the Southern Ocean. Mar Ecol Prog Ser 134:1–14.  https://doi.org/10.3354/meps134001 CrossRefGoogle Scholar
  47. Petry MV, Basler AB, Valls FCL, Krüger L (2013) New southerly breeding location of King Penguins (Aptenodytes patagonicus) on Elephant Island (Maritime Antarctic). Polar Biol 36:603–606.  https://doi.org/10.1007/s00300-012-1277-1 CrossRefGoogle Scholar
  48. Pollock KH, Kendall WL (1987) Visibility bias in aerial surveys: a review of estimation procedures. J Wildl Manag 51:502–510.  https://doi.org/10.2307/3801040 CrossRefGoogle Scholar
  49. Poncet S, Crosbie K (2005) A visitor’s guide to South Georgia. Princeton University Press, PrincetonGoogle Scholar
  50. Pusch C, Hulley PA, Kock KH (2004) Community structure and feeding ecology of mesopelagic fish in the slope waters of King George Island (South Shetland Islands, Antarctica). Deep-Sea Res 51:1685–1708.  https://doi.org/10.1016/j.dsr.2004.06.008 CrossRefGoogle Scholar
  51. Raclot T, Groscolas R, Cherel Y (1998) Fatty acid evidence for the importance of myctophid fishes in the diet of King Penguins, Aptenodytes patagonicus. Mar Biol 132:523–533.  https://doi.org/10.1007/s002270050418 CrossRefGoogle Scholar
  52. Ropert-Courdert Y, Kato A, Meyer X, Pellé M, McIntosh AJJ, Angelier F, Chastel O, Widmann M, Arthur B, Raymond B, Raclot T (2014) A complete breeding failure in an Adélie Penguin colony correlates with unusual and extreme environmental events. Ecography 37:1–3.  https://doi.org/10.1111/ecog.01182 CrossRefGoogle Scholar
  53. Rounsevell DE, Copson GR (1982) Growth rate and recovery of a King Penguin, Aptenodytes patagonicus, population after exploitation. Wildl Res 9:519–525.  https://doi.org/10.1071/wr9820519 CrossRefGoogle Scholar
  54. Saunders RA, Collins MA, Ward P, Stowasser G, Hill SL, Shreeve R, Tarling GA (2015) Predatory impact of the myctophid fish community on zooplankton in the Scotia Sea (Southern Ocean). Mar Ecol Prog Ser 541:45–64.  https://doi.org/10.3354/meps11527 CrossRefGoogle Scholar
  55. Sladen WJL (1964) The distribution of the Adélie and chinstrap penguins. In: Carrick R et al (eds) Biologie Antarctique. Hermann, Paris, pp 359–365Google Scholar
  56. Sokolov S, Rintoul SR (2009) Circumpolar structure and distribution of the Antarctic Circumpolar Current fronts 2: variability and relationship to sea surface height. J Geophys Res.  https://doi.org/10.1029/2008jc005248 Google Scholar
  57. Stier A, Viblanc VA, Massemin-Challet S, Handrich Y, Zahn S, Rojas ER, Saraux C, Le Vaillant M, Prud’homme O, Grosbellet E, Robin J-P, Bize P, Criscuolo F (2014) Starting with a handicap: phenotypic differences between early- and late-born King Penguin chicks and their survival correlates. Funct Ecol 28:601–611.  https://doi.org/10.1111/1365-2435.12204 CrossRefGoogle Scholar
  58. Stonehouse B (1956) The King Penguin of South Georgia. Nature 178:1424–1426.  https://doi.org/10.1038/1781424a0 CrossRefGoogle Scholar
  59. Summerhayes C (2009) The next 100 years in Antarctic climate change and the environment. Scientific Committee on Antarctic Research, Cambridge, pp 299–389Google Scholar
  60. Trathan PN, Daunt FJH, Murphy EJ (eds) (1996) South Georgia: an ecological atlas. British Antarctic Survey, CambridgeGoogle Scholar
  61. Trathan P, Forcada J, Murphy E (2007) Environmental forcing and Southern Ocean marine predator populations: effects of climate change and variability. Phil Trans Roy Soc B 362:2351–2365.  https://doi.org/10.1098/rstb.2006.1953 CrossRefGoogle Scholar
  62. Trathan PN, Bishop C, Maclean G, Brown P, Fleming A, Collins MA (2008) Linear tracks and restricted temperature ranges characterise penguin foraging pathways. Mar Ecol Prog Ser 370:285–294.  https://doi.org/10.3354/meps07638 CrossRefGoogle Scholar
  63. Trucchi E, Gratton P, Whittington JD, Cristofari R, Le Maho Y, Stenseth N, LeBohec C (2014) King penguin demography since the last glaciation inferred from genome-wide data. Proc R Soc B 281:20140528.  https://doi.org/10.1098/rspb.2014.0528 CrossRefPubMedPubMedCentralGoogle Scholar
  64. Turner J, Colwell SR, Marshall GJ, Lachlan-Cope TA, Carleton AM, Jones PD, Lagun V, Reid PA, Iagovkina S (2004) The SCAR READER project: toward a high-quality database of mean Antarctic meteorological observations. J Climate 17:2890–2898CrossRefGoogle Scholar
  65. Turner J, Barrand NE, Bracegirdle TJ, Convey P, Hodgson DA, Jarvis M, Jenkins A, Marshall G, Meredith MP, Roscoe H, Shanklin J, French J, Goosse H, Guglielmin M, Gutt J, Jacobs S, Kennicutt MC, Masson-Delmotte V, Mayewski P, Navarro F, Robinson SA, Scambos T, Sparrow M, Summerhayes C, Speer K, Klepikov A (2014) Antarctic climate change and the environment: an update. Polar Rec 50:237–259.  https://doi.org/10.1017/S0032247413000296 CrossRefGoogle Scholar
  66. van den Hoff J, McMahon CR, Field I (2009) Tipping back the balance: recolonization of the Macquarie Island isthmus by King Penguins (Aptenodytes patagonicus) following extermination for human gain. Antarctic Sci 21:237–241.  https://doi.org/10.1017/s0954102009001898 CrossRefGoogle Scholar
  67. Weimerskirch H, Stahl JC, Jouventin P (1992) The breeding biology and population dynamics of King Penguins Aptenodytes patagonica on the Crozet Islands. Ibis 134:107–117.  https://doi.org/10.1111/j.1474-919x.1992.tb08387.x CrossRefGoogle Scholar
  68. Whitehouse MJ, Meredith MP, Rothery P, Atkinson A, Ward P, Korb RE (2008) Rapid warming of the ocean around South Georgia, Southern Ocean, during the 20th century: forcings, characteristics and implications for lower trophic levels. Deep-Sea Res 55:1218–1228.  https://doi.org/10.1016/j.dsr.2008.06.002 CrossRefGoogle Scholar
  69. Williams R (1985) The potential impact of a krill fishery upon pelagic fish in the Prydz Bay area of Antarctica. Polar Biol 5:1–4.  https://doi.org/10.1007/bf00446039 CrossRefGoogle Scholar
  70. Woehler EJ, Croxall JP (1997) The status and trends of Antarctic and Sub-Antarctic seabirds. Mar Ornithol 25:43–66Google Scholar
  71. Woehler EJ, Cooper J, Croxall JP, Fraser WR, Kooyman GL, Miller GD, Nel DC, Patterson DL, Peter H-U, Ribic CA, Salwicka K, Trivelpiece WZ, Weimerskirch H (2001) A statistical assessment of the status and trends of Antarctic and Sub-Antarctic seabirds. Scientific Committee for Antarctic Research, CambridgeGoogle Scholar

Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  1. 1.Department of Ecology and EvolutionStony Brook UniversityStony BrookUSA
  2. 2.Department of ZoologyUniversity of OxfordOxfordUK

Personalised recommendations