A new fossil from the mid-Paleocene of New Zealand reveals an unexpected diversity of world’s oldest penguins


We describe leg bones of a giant penguin from the mid-Paleocene Waipara Greensand of New Zealand. The specimens were found at the type locality of Waimanu manneringi and together with this species they constitute the oldest penguin fossils known to date. Tarsometatarsus dimensions indicate a species that reached the size of Anthropornis nordenskjoeldi, one of the largest known penguin species. Stem group penguins therefore attained a giant size very early in their evolution, with this gigantism existing for more than 30 million years. The new fossils are from a species that is phylogenetically more derived than Waimanu, and the unexpected coexistence of Waimanu with more derived stem group Sphenisciformes documents a previously unknown diversity amongst the world’s oldest penguins. The characteristic tarsometatarsus shape of penguins evolved early on, and the significant morphological disparity between Waimanu and the new fossil conflicts with recent Paleocene divergence estimates for penguins, suggesting an older, Late Cretaceous, origin.

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  1. Acosta Hospitaleche C (2014) New giant penguin bones from Antarctica: systematic and paleobiological significance. Comptes Rendus Palevol 13:555–560

    Article  Google Scholar 

  2. Acosta Hospitaleche C, Olivero E (2016) Re-evaluation of the fossil penguin Palaeeudyptes gunnari from the Eocene Leticia formation, Argentina: additional material, systematics and palaeobiology. Alcheringa 40:373–382

    Article  Google Scholar 

  3. Ando T, Fordyce RE (2014) Evolutionary drivers for flightless, wing-propelled divers in the Northern and Southern Hemispheres. Palaeogeogr Palaeoclimatol Palaeoecol 400:50–61

    Article  Google Scholar 

  4. Baker AJ, Pereira SL, Haddrath OP, Edge KA (2006) Multiple gene evidence for expansion of extant penguins out of Antarctica due to global cooling. Proc Roy Soc London B: Biol Sci 273:11–17

    Article  Google Scholar 

  5. Browne GH, Lawrence MJF, Mortimer N, Clowes CD, Morgans HEG, Hollis CJ, Beu AG, Black JA, Sutherland R, Bache F (2016) Stratigraphy of Reinga and Aotea basins, NW New Zealand: constraints from dredge samples on regional correlations and reservoir character. New Zeal J Geol Geophys 59:396–415

    CAS  Article  Google Scholar 

  6. Chávez Hoffmeister M (2014) Phylogenetic characters in the humerus and tarsometatarsus of penguins. Pol Polar Res 35:469–496

    Google Scholar 

  7. Clarke JA, Ksepka DT, Stucchi M, Urbina M, Giannini N, Bertelli S, Narváez Y, Boyd CA (2007) Paleogene equatorial penguins challenge the proposed relationship between penguin biogeography, diversity, and Cenozoic climate change. Proc Natl Acad Sci U S A 104:11545–11550

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  8. Clarke JA, Ksepka DT, Salas-Gismondi R, Altamirano AJ, Shawkey MD, D’Alba L, Vinther J, DeVries TJ, Baby P (2010) Fossil evidence for evolution of the shape and color of penguin feathers. Science 330:954–957

    CAS  Article  PubMed  Google Scholar 

  9. Fordyce RE, Thomas DB (2011) Kaiika maxwelli, a new Early Eocene archaic penguin (Sphenisciformes, Aves) from Waihao Valley, South Canterbury, New Zealand. New Zeal J Geol Geop 54:43–51

    Article  Google Scholar 

  10. Goloboff PA (1993) NONA, version 2.0. Published by the author, S. M. de Tucumán, Argentina

  11. Jadwiszczak P (2001) Body size of Eocene Antarctic penguins. Pol Polar Res 22:147–158

    Google Scholar 

  12. Jadwiszczak P (2015) Another look at tarsometatarsi of early penguins. Pol Polar Res 36:343–354

    Google Scholar 

  13. Jadwiszczak P, Acosta Hospitaleche C, Reguero M (2013) Redescription of Crossvallia unienviella: the only Paleocene Antarctic penguin. Ameghiniana 50:545–553

    Article  Google Scholar 

  14. Jarvis ED, Mirarab S, Aberer AJ et al (2014) Whole-genome analyses resolve early branches in the tree of life of modern birds. Science 346:1320–1331

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  15. Kato A, Ropert-Coudert Y, Grémillet D, Cannell B (2006) Locomotion and foraging strategy in foot-propelled and wing-propelled shallow-diving seabirds. Mar Ecol Prog Ser 308:293–301

    Article  Google Scholar 

  16. Ksepka DT, Ando T (2011) Penguins past, present, and future: trends in the evolution of the Sphenisciformes. In: Dyke G, Kaiser G (eds) Living dinosaurs: the evolutionary history of modern birds. Wiley, Chichester, pp 155–186

    Google Scholar 

  17. Ksepka DT, Fordyce RE, Ando T, Jones CM (2012) New fossil penguins (Aves, Sphenisciformes) from the Oligocene of New Zealand reveal the skeletal plan of stem penguins. J Vert Paleontol 32:235–254

    Article  Google Scholar 

  18. Mayr G (2016) Avian evolution: the fossil record of birds and its paleobiological significance. Wiley-Blackwell, Chichester

    Google Scholar 

  19. Mitchell KJ, Cooper A, Phillips MJ (2015) Comment on “whole-genome analyses resolve early branches in the tree of life of modern birds”. Science 349:1460

    CAS  Article  PubMed  Google Scholar 

  20. Myrcha A, Jadwiszczak P, Tambussi CP, Noriega JI, Gaździcki A, Tatur A, del Valle RA (2002) Taxonomic revision of Eocene Antarctic penguins based on tarsometatarsal morphology. Pol Polar Res 23:5–46

    Google Scholar 

  21. Nixon KC (2002) WinClada, version 1.00.08. Ithaca

  22. Prum RO, Berv JS, Dornburg A, Field DJ, Townsend JP, Lemmon EM, Lemmon AR (2015) A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526:569–573

    CAS  Article  PubMed  Google Scholar 

  23. Simpson GG (1946) Fossil penguins. Bull Am Mus Nat Hist 87:1–99

    Google Scholar 

  24. Slack KE, Jones CM, Ando T, Harrison GL, Fordyce RE, Arnason U, Penny D (2006) Early penguin fossils, plus mitochondrial genomes, calibrate avian evolution. Mol Biol Evol 23:1144–1155

    CAS  Article  PubMed  Google Scholar 

  25. Tambussi CP, Reguero MA, Marenssi SA, Santillana SN (2005) Crossvallia unienwillia, a new Spheniscidae (Sphenisciformes, Aves) from the late Paleocene of Antarctica. Geobios 38:667–675

    Article  Google Scholar 

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We are indebted to Leigh Love, who collected the specimens and donated them to Canterbury Museum. We also thank Al Mannering for preparation work and Chris Clowes for data on the dinoflagellate fauna. Marío Chávez Hoffmeister provided a nexus file of his phylogenetic data set. Comments from Ursula Göhlich, Carolina Acosta-Hospitaleche, and four anonymous reviewers improved the text.

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Correspondence to Gerald Mayr.

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Mayr, G., De Pietri, V.L. & Paul Scofield, R. A new fossil from the mid-Paleocene of New Zealand reveals an unexpected diversity of world’s oldest penguins. Sci Nat 104, 9 (2017). https://doi.org/10.1007/s00114-017-1441-0

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  • Fossil birds
  • Sphenisciformes
  • Waipara Greensand
  • Waimanu
  • Crossvallia