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Complete mitochondrial genome sequences of two extinct moas clarify ratite evolution

Nature volume 409, pages 704707 (08 February 2001) | Download Citation



The origin of the ratites, large flightless birds from the Southern Hemisphere, along with their flighted sister taxa, the South American tinamous, is central to understanding the role of plate tectonics in the distributions of modern birds and mammals. Defining the dates of ratite divergences is also critical for determining the age of modern avian orders1,2,3,4,5,6. To resolve the ratite phylogeny and provide biogeographical data to examine these issues, we have here determined the first complete mitochondrial genome sequences of any extinct taxa— two New Zealand moa genera—along with a 1,000-base-pair sequence from an extinct Madagascan elephant-bird. For comparative data, we also generated 12 kilobases of contiguous sequence from the kiwi, cassowary, emu and two tinamou genera. This large dataset allows statistically precise estimates of molecular divergence dates and these support a Late Cretaceous vicariant speciation of ratite taxa, followed by the subsequent dispersal of the kiwi to New Zealand. This first molecular view of the break-up of Gondwana provides a new temporal framework for speciation events within other Gondwanan biota and can be used to evaluate competing biogeographical hypotheses.

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  1. 1.

    Phylogeny and evolution of the ratite birds. Ibis 116, 494–521 (1974).

  2. 2.

    & Phylogeny and Classification of Birds (Yale Univ. Press, London, 1990).

  3. 3.

    et al. Independent origins of the New Zealand moas and kiwis. Proc. Natl Acad. Sci. USA 89, 8741–8744 (1992).

  4. 4.

    & Mass survival of birds across the Cretaceous–Tertiary: Molecular evidence. Science 275, 1109–1113 (1997).

  5. 5.

    The Origin and Evolution of Birds (Harvard Univ. Press, Cambridge, Massachusetts, 1997).

  6. 6.

    , & Phylogeny and biogeography of ratite birds inferred from DNA sequences of the mitochondrial ribosomal genes. Mol. Biol. Evol. 15, 370–376 (1998).

  7. 7.

    in Avian Biology Vol. VIII (eds Farner, D. S., King, J. R. & Parkes, K. C.) 79–238 (Academic, Orlando, 1985).

  8. 8.

    , & in Avian Molecular Evolution and Molecular Systematics (ed. Mindell, D.) 173–208 (Academic, New York, 1997).

  9. 9.

    Ostrich ancestors found in the Northern Hemisphere suggest new hypothesis of ratite origins. Nature 324, 563–565 (1986).

  10. 10.

    A phylogenetic analysis of postcranial skeletal characters of the ratite birds. Ann. Carnegie Mus. 57, 73–90 (1988).

  11. 11.

    in Avian Molecular Evolution and Molecular Systematics (ed. Mindell, D.) 345–373 (Academic, New York, 1997).

  12. 12.

    , , & The retrieval of ancient human DNA sequences. Am. J. Hum. Genet. 59, 368–376 (1996).

  13. 13.

    et al. Neandertal DNA sequence and the origin of modern humans. Cell 90, 19–30 (1997).

  14. 14.

    in Ancient DNA (eds Herrmann, B. & Hummel, S.) 149–165 (Springer, New York, 1993).

  15. 15.

    Hindlimb proportions and locomotion of Emuarius gidju (Patterson & Rich, 1987) (Aves: Casuariidae). Memoirs of the Queensland Museum 41, 235–240 (1997).

  16. 16.

    , , & in Geological Evolution of Antarctica (eds Thomson, M. R. A., Crame, J. A. & Thomson, J. W.) 533–539 (Cambridge Univ. Press, Cambridge, 1991).

  17. 17.

    & The New Zealand biota: Historical background and new research. Trends Ecol. Evol. 8, 429–433 (1993).

  18. 18.

    The Geological History of New Zealand and its Life (Univ. Auckland Press, Auckland, 1979).

  19. 19.

    Lands in collision. N. Z. Dept Sci. Ind. Res. Inf. Serv. 161 (1985).

  20. 20.

    in The Africa–South America Connection (eds George, W. & Lavocat, R.) 76–86 (Clarendon, Oxford, 1993).

  21. 21.

    & Using sequences of rbcL to study phylogeny and biogeography of Nothofagus species. Aust. Syst. Bot. 6, 441–447 (1993).

  22. 22.

    et al. Reinga Basin and its margins. N. Z. J. Geol. Geophys. 40, 425–451 (1997).

  23. 23.

    F. Ratite eggshells and phylogenetic questions. Bonn Zool. Beitr. 23, 3–48 (1972).

  24. 24.

    , , , & Cosmopolitanism among Gondwanan Late Cretaceous mammals. Nature 390, 504–507 (1997).

  25. 25.

    et al. Predatory dinosaur remains from Madagascar: Implications for the Cretaceous biogeography of Gondwana. Science, 280, 1048–1051 (1998).

  26. 26.

    & Ancient DNA: Do it right or not at all. Science 289, 1139 (2000).

  27. 27.

    PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods) (Sinauer, Sunderland, Massachusetts, 1999).

  28. 28.

    , & in Molecular Zoology: Strategies and Protocols (eds Ferraris, J. & Palumbi, S.) 19–45 (Wiley, New York, 1996).

  29. 29.

    & Estimating divergence dates from molecular sequences. Mol. Biol. Evol. 15, 442–448 (1998).

  30. 30.

    Evolutionary trees from DNA sequences: a maximum likelihood approach. J. Mol. Evol. 17, 368–376 (1981).

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We thank W. Boles, R. Cooper, R. Herzer, P. Houde, C. Mourer-Chauviré, D. Penny and T. Worthy for valuable comments, and M. Sorenson for allowing us access to unpublished rhea and ostrich sequences. We are grateful to T. Worthy and the staff of the Museum of New Zealand for the moa samples. Modern samples were kindly provided by A. C. Wilson (deceased), M. Potter and M. Braun, and laboratory space by R. Thomas, J. Bertranpetit and the Oxford University Museum. A.C. was supported by the NERC, the Leverhulme Fund, the New Zealand Marsden Fund and the Royal Society. C.L.F. was supported by the Comissionat per a Universitats i Recerca (Catalan Autonomous Government), and A.R. was supported by the Wellcome Trust.

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  1. *Department of Biological Anthropology and Henry Wellcome Ancient Biomolecules Centre, University of Oxford, Oxford OX1 6UE, UK

    • Alan Cooper
    • , Carles Lalueza-Fox
    • , Simon Anderson
    •  & Ryk Ward
  2. †Department of Zoology, University of Oxford, Oxford OX2 3PS, UK

    • Alan Cooper
    •  & Andrew Rambaut
  3. ‡Seccio Antropologia, Facultat de Biologia, Universitat de Barcelona, Barcelona 08028, Spain

    • Carles Lalueza-Fox
  4. §Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK

    • Jeremy Austin


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Correspondence to Alan Cooper.

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