Biodiversity and Conservation

, Volume 28, Issue 11, pp 2861–2876 | Cite as

Global terrestrial distribution of penguins (Spheniscidae) and their conservation by protected areas

  • Rachel P. HickcoxEmail author
  • Manuel Jara
  • Laura A. K. Deacon
  • Lilly P. Harvey
  • Daniel Pincheira-DonosoEmail author
Original Paper


Establishing protected areas (PAs) ranks among the top priority actions to mitigate the global scale of modern biodiversity declines. However, the distribution of biodiversity is spatially asymmetric among regions and lineages, and the extent to which PAs offer effective protection for species and ecosystems remains uncertain. Penguins, regarded as prime bioindicator birds of the ecological health of their terrestrial and marine habitats, represent priority targets for such quantitative assessments. Of the world’s 18 penguin species, eleven are undergoing population declines, for which ten are classified as ‘Vulnerable’ or ‘Endangered’. Here, we employ a global-scale dataset to quantify the extent to which their terrestrial breeding areas are currently protected by PAs. Using quantitative methods for spatial ecology, we compare the global distribution of penguin colonies, including range and population size analyses, with the distribution of terrestrial PAs classified by the International Union for Conservation of Nature, and generate hotspot and endemism maps worldwide. Our assessment quantitatively reveals < 40% of the terrestrial range of eleven penguin species is currently protected, and that range size is the significant factor in determining PA protection. We also show that there are seven global hotspots of penguin biodiversity where four or five penguin species breed. We suggest that future penguin conservation initiatives should be implemented based on more comprehensive, quantitative assessments of the multi-dimensional interactions between areas and species to further the effectiveness of PA networks.


Biodiversity hotspots IUCN Macroecology Penguins Protected areas Species richness 



The data used are derived from public repositories. We thank UNEP World Conservation Monitoring Centre, IUCN World Commission on Protected Areas (World Database on Protected Areas), IUCN Global Species Programs and the Species Survival Commission, OBIS, GBIF, and Birdlife International for developing the source data. Thank you to Professor Philip Seddon, James Hunter, Saif Khan, and anonymous reviewers for manuscript comments and edits. M.J and L.A.K.D are fully funded by the University of Lincoln, School of Life Sciences. R.P.H is fully funded by the University of Otago Doctoral Scholarship. R.P.H, D.P.D designed the study. R.P.H performed data collection. R.P.H, M.J performed analysis and interpretation. R.P.H wrote manuscript. D.P.D. supervised project. All authors (R.P.H, M.J, L.A.K.D, L.P.H, D.P.D) discussed the results and implications and edited and commented on the manuscript at all stages.

Supplementary material

10531_2019_1801_MOESM1_ESM.docx (1.2 mb)
Supplementary material 1 (DOCX 1230 kb)


  1. Akçakaya HR, Mills G, Doncaster CP (2007) The role of metapopulations in conservation. In: Macdonald DW, Service K (eds) Key topics in conservation biology. Blackwell, Oxford, pp 64–84Google Scholar
  2. Bertzky B, Corrigan C, Kemsey J et al (2012) Protected planet report 2012: tracking progress towards global targets for protected areas. UNEP-WCMC, CambridgeGoogle Scholar
  3. Boersma PD (2008) Penguins as marine sentinels. Bioscience 58:597–607CrossRefGoogle Scholar
  4. Boersma PD, Parrish JK (1999) Limiting abuse: marine protected areas, a limited solution. Ecol Econ 31:287–304. CrossRefGoogle Scholar
  5. Boersma PD, Rebstock GA (2014) Climate change increases reproductive failure in Magellanic penguins. PLoS ONE 9:e85602CrossRefPubMedPubMedCentralGoogle Scholar
  6. Borboroglu PG, Boersma PD (2013) Penguins: natural history and conservation. University of Washington Press, Washinton, DCGoogle Scholar
  7. Borboroglu PG, Boersma PD, Reyes L, Skewgar E (2008) Petroleum, pollution, and penguins: marine conservation tools to reduce the problem. In: Hofer TN (ed) Marine pollution: new research. Nova Science Publishers Inc., New York, pp 339–356Google Scholar
  8. Breiner FT, Bergamini A (2018) Improving the estimation of area of occupancy for IUCN Red List assessments by using a circular buffer approach. Biodivers Conserv 27:2443–2448. CrossRefGoogle Scholar
  9. Brooks TM, Bakarr MI, Boucher TIM et al (2004) Coverage provided by the global protected area system: is it enough? Bioscience 54:1081–1091CrossRefGoogle Scholar
  10. Brooks TM, Mittermeier RA, da Fonseca GAB et al (2006) Global biodiversity conservation priorities. Science 80(313):58–61. CrossRefGoogle Scholar
  11. Brown JL (2014) SDMtoolbox: a python-based GIS toolkit for landscape genetic, biogeographic, and species distribution model analyses. Methods Ecol Evol 5:694–700CrossRefGoogle Scholar
  12. Ceballos G, Ehrlich PR, Barnosky AD et al (2015) Accelerated modern human-induced species losses: entering the sixth mass extinction. Sci Adv 1:e1400253–e1400253. CrossRefPubMedPubMedCentralGoogle Scholar
  13. Charlesworth B, Charlesworth D (2010) Elements of evolutionary genetics. Roberts & Co., Greenwood VillageGoogle Scholar
  14. Chevin L-M, Lande R, Mace GM (2010) Adaptation, plasticity, and extinction in a changing environment: towards a predictive theory. PLoS Biol 8:e1000357. CrossRefPubMedPubMedCentralGoogle Scholar
  15. Coetzee BWT, Convey P, Chown SL (2017) Expanding the protected area network in antarctica is urgent and readily achievable. Conserv Lett 10:670–680. CrossRefGoogle Scholar
  16. Crisp MD, Laffan S, Linder HP, Monro A (2001) Endemism in the Australian flora. J Biogeogr 28:183–198CrossRefGoogle Scholar
  17. Deguignet M, Arnell A, Juffe-Bignoli D et al (2017) Measuring the extent of overlaps in protected area designations. PLoS ONE 12:e0188681. CrossRefPubMedPubMedCentralGoogle Scholar
  18. Dirzo R, Young HS, Galetti M et al (2014) Defaunation in the Anthropocene. Science 80(345):401–406. CrossRefGoogle Scholar
  19. Dudley N (2008) Guidelines for applying protected area management categories. IUCN, GlandCrossRefGoogle Scholar
  20. Eken G, Bennun L, Brooks TM et al (2004) Key biodiversity areas as site conservation targets. Bioscience 54:1110–1118CrossRefGoogle Scholar
  21. Ellis S (1999) The penguin conservation assessment and management plan: a description of the process. Mar Ornithol 27:163–169Google Scholar
  22. ESRI (2018) ArcGIS Desktop: 10.2.2. 10.6.1Google Scholar
  23. Ferrière R, Dieckmann U, Couvet D (2004) Evolutionary conservation biology. Cambridge University Press, CambridgeCrossRefGoogle Scholar
  24. Frankham R (1996) Relationship of genetic variation to population size in wildlife. Conserv Biol 10:1500–1508. CrossRefGoogle Scholar
  25. Gandini P, Frere E, Boersma PD (2010) Status and conservation of Magellanic Penguins Spheniscus magellanicus in Patagonia, Argentina. Bird Conserv Int 6:307–316. CrossRefGoogle Scholar
  26. Gaston KJ (2003) The structure and dynamics of geographic ranges. Oxford University Press, OxfordGoogle Scholar
  27. Gaston KJ, Jackson SF, Cantú-Salazar L, Cruz-Piñón G (2008) The ecological performance of protected areas. Annu Rev Ecol Evol Syst 39:93–113. CrossRefGoogle Scholar
  28. GBIF (2018) The global biodiversity information facility backbone taxonomy.
  29. Geldmann J, Barnes M, Coad L et al (2013) Effectiveness of terrestrial protected areas in reducing habitat loss and population declines. Biol Conserv 161:230–238. CrossRefGoogle Scholar
  30. Gillingham PK, Bradbury RB, Roy DB et al (2015) The effectiveness of protected areas in the conservation of species with changing geographical ranges. Biol J Linn Soc 115(3):707–717CrossRefGoogle Scholar
  31. Hernández HM, Navarro M (2007) A new method to estimate areas of occupancy using herbarium data. Biodivers Conserv 16:2457–2470. CrossRefGoogle Scholar
  32. Höglund J (2009) Evolutionary conservation genetics. Oxford University Press, OxfordCrossRefGoogle Scholar
  33. Iojă CI, Pătroescu M, Rozylowicz L et al (2010) The efficacy of Romania’s protected areas network in conserving biodiversity. Biol Conserv 143:2468–2476. CrossRefGoogle Scholar
  34. Isik K (2011) Rare and endemic species: why are they prone to extinction. Turk J Bot 35:411–417. CrossRefGoogle Scholar
  35. IUCN (2001) IUCN red list categories and criteria, version 3.1. IUCN Species Survival Commission, Gland, Switzerland and CambridgeGoogle Scholar
  36. IUCN (2018) The IUCN red list of threatened species. Version 2018-1.
  37. IUCN, UNEP (2018) The world database on protected areas (WDPA).
  38. Laffan SW, Crisp MD (2003) Assessing endemism at multiple spatial scales, with an example from the Australian vascular flora. J Biogeogr 30:511–520. CrossRefGoogle Scholar
  39. Lennon JJ, Koleff P, Greenwood JJD, Gaston KJ (2003) Contribution of rarity and commonness to patterns of species richness. Ecol Lett 7:81–87. CrossRefGoogle Scholar
  40. Mace GM, Collar NJ, Gaston KJ et al (2008) Quantification of extinction risk: IUCN’s system for classifying threatened species. Conserv Biol 22:1424–1442. CrossRefPubMedGoogle Scholar
  41. Meiri S, Bauer AM, Allison A et al (2018) Extinct, obscure or imaginary: the lizard species with the smallest ranges. Divers Distrib 24:262–273. CrossRefGoogle Scholar
  42. Moilanen A, Wilson KA, Possingham HP (2009) Spatial conservation prioritization: quantitative methods and computational tools. Oxford University Press, OxfordGoogle Scholar
  43. Myers N, Mittermeier RA, Mittermeier CG et al (2000) Biodiversity hotspots for conservation priorities. Nature 403:853–858. CrossRefGoogle Scholar
  44. Newbold T, Hudson LN, Hill SLL et al (2015) Global effects of land use on local terrestrial biodiversity. Nature 520:45–50. CrossRefPubMedGoogle Scholar
  45. Orme CDL, Davies RG, Burgess M et al (2005) Global hotspots of species richness are not congruent with endemism or threat. Nature 436:1016–1019. CrossRefPubMedGoogle Scholar
  46. Pichegru L, Grémillet D, Crawford RJM, Ryan PG (2010) Marine no-take zone rapidly benefits endangered penguin. Biol Lett 6:498–501. CrossRefPubMedPubMedCentralGoogle Scholar
  47. Pimm SL, Jenkins CN, Abell R et al (2014) The biodiversity of species and their rates of extinction, distribution, and protection. Science. CrossRefPubMedGoogle Scholar
  48. Pressey RL, Taffs KH (2001) Scheduling conservation action in production landscapes: priority areas in western New South Wales defined by irreplaceability and vulnerability to vegetation loss. Biol Conserv 100:355–376CrossRefGoogle Scholar
  49. Pressey RL, Johnson IR, Wilson PD (1994) Shades of irreplaceability: towards a measure of the contribution of sites to a reservation goal. Biodivers Conserv 3:242–262CrossRefGoogle Scholar
  50. QGIS (2018) QGIS 3.2.1Google Scholar
  51. R Development Core Team (2019) R: a language and environment for statistical computingGoogle Scholar
  52. Rangel TF, Diniz-Filho JAF, Bini LM (2010) SAM: a comprehensive application for spatial analysis in macroecology. Ecography (Cop) 33:46–50CrossRefGoogle Scholar
  53. Reid WV (1998) Biodiversity hotspots. Trends Ecol Evol 13:275–280CrossRefPubMedGoogle Scholar
  54. Rivers MC, Bachman SP, Meagher TR et al (2010) Subpopulations, locations and fragmentation: applying IUCN red list criteria to herbarium specimen data. Biodivers Conserv 19:2071–2085. CrossRefGoogle Scholar
  55. Rodrigues ASL, Akcakaya HR, Andelman SJ et al (2004) Global gap analysis: priority regions for expanding the global protected-area network. Bioscience 54:1092–1100CrossRefGoogle Scholar
  56. Ropert-coudert Y, Chiaradia A, Ainley D et al (2019) Happy feet in a hostile world? The future of penguins depends on proactive management of current and predictable threats. Front Mar Sci 6:248CrossRefGoogle Scholar
  57. Secretariat of the Convention on Biological Diversity (2008) Protected areas in today’s world: their values and benefits for the welfare of the planet. Montreal 36:96Google Scholar
  58. Southwell C, Emmerson L, Takahashi A et al (2017) Large-scale population assessment informs conservation management for seabirds in Antarctica and the Southern Ocean: a case study of Adélie penguins. Glob Ecol Conserv 9:104–115. CrossRefGoogle Scholar
  59. Terauds A (2017) An update to the Antarctic Specially Protected Areas (ASPAs). In: Aust. Antarct. Data Cent. Accessed 11 Mar 2019
  60. Terauds A (2018) Antarctic Specially Protected Areas (Points and Polygons). In: Aust. Antarct. Data Cent. Accessed 11 Mar 2019
  61. Thiollay J-M (2002) Bird diversity and selection of protected areas in a large neotropical forest tract. Biodivers Conserv 11:1377–1395. CrossRefGoogle Scholar
  62. Thomas CD, Gillingham PK (2015) The performance of protected areas for biodiversity under climate change. Biol J Linn Soc 115:718–730CrossRefGoogle Scholar
  63. Trathan PN, Borboroglu PG, Boersma D et al (2014) Pollution, habitat loss, fishing, and climate change as critical threats to penguins. Conserv Biol. CrossRefPubMedGoogle Scholar
  64. UNEP-WCMC, IUCN, NGS (2018) Protected planet report. Gland, CambridgeGoogle Scholar
  65. United Nations (1991) Protocol on Environmental Protection to the Antarctic Treaty. In: Antarctic Treaty Consultative Meeting XVI, 7-18 Oct 1991. Bonn, GermanyGoogle Scholar
  66. Urban MC (2015) Accelerating extinction risk from climate change. Science 80(348):571–573. CrossRefGoogle Scholar
  67. Venter O, Fuller RA, Segan DB et al (2014) Targeting global protected area expansion for imperiled biodiversity. PLoS Biol 12:e1001891. CrossRefPubMedPubMedCentralGoogle Scholar

Copyright information

© Springer Nature B.V. 2019

Authors and Affiliations

  1. 1.Department of ZoologyUniversity of OtagoDunedinNew Zealand
  2. 2.Laboratory of Evolutionary Ecology of Adaptations, School of Life Sciences, Joseph Banks LaboratoriesUniversity of LincolnLincolnUK
  3. 3.Department of Population Health and PathobiologyCollege of Veterinary Medicine, North Carolina State UniversityRaleighUSA
  4. 4.School of Science and TechnologyNottingham Trent UniversityNottinghamUK

Personalised recommendations