Skip to main content

Thank you for visiting You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

The phylogenetic position of the ‘giant deer’ Megaloceros giganteus


The giant deer, or ‘Irish elk’, has featured extensively in debates on adaptation, sexual selection, and extinction. Its huge antlers—the largest of any deer species, living or extinct—formed a focus of much past work1,2,3,4. Yet the phylogenetic position of the giant deer has remained an enigma. On the basis of its flattened antlers, the species was previously regarded as closely related to the living fallow deer5,6,7. Recent morphological studies8, however, have challenged that view and placed the giant deer closer to the living red deer or wapiti. Here we present a new phylogenetic analysis encompassing morphological and DNA sequence evidence, and find that both sets of data independently support a sister-group relationship of giant and fallow deer. Our results include the successful extraction and sequencing of DNA from this extinct species, and highlight the value of a joint molecular and morphological approach.

This is a preview of subscription content

Access options

Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.

Figure 1: Phylogenetic relationships among deer species based on molecular and morphological analyses.
Figure 2: Examples of morphological characters.


  1. Gould, S. J. The origin and function of ‘bizarre’ structures: antler size and skull size in the ‘Irish Elk,’ Megaloceros giganteus . Evolution 28, 191–220 (1974)

    PubMed  Google Scholar 

  2. Moen, R. A., Pastor, J. & Cohen, Y. Antler growth and extinction of Irish Elk. Evol. Ecol. Res. 1, 235–249 (1999)

    Google Scholar 

  3. Geist, V. Deer of the World: Their Evolution, Behavior and Ecology (Swan Hill Press, Shrewsbury, 1999)

    Google Scholar 

  4. Stuart, A. J., Kosintsev, P., Higham, T. F. & Lister, A. M. Pleistocene to Holocene extinction dynamics in giant deer and woolly mammoth. Nature 431, 684–689 (2004)

    ADS  CAS  Article  PubMed  Google Scholar 

  5. Lydekker, R. The Deer of all Lands (Rowland Ward, London, 1898)

    Google Scholar 

  6. Millais, J. G. The Mammals of Great Britain and Ireland (Longman, Green & Co., London, 1906)

    Google Scholar 

  7. Reynolds, S. H. Monograph on the British Mammalia of the Pleistocene Period. The Giant Deer 1–58 (Palaeontographical Society Memoir, London, 1929)

    Google Scholar 

  8. Pfeiffer, T. Die Stellung von Dama (Cervidae, Mammalia) im System plesiometacarpaler Hirsche des Pleistozäns. Cour. Forsch.-Inst. Senckenberg 211, 1–218 (1999)

    Google Scholar 

  9. Groves, C. & Grubb, P. in Biology and Management of the Cervidae (ed. Wemmer, C. M.) 21–59 (Smithsonian Institution, Washington, D.C., 1987)

    Google Scholar 

  10. Randi, E., Mucci, N., Pierpaoli, M. & Douzery, E. New phylogenetic perspectives on the Cervidae (Artiodactyla) are provided by the mitochondrial cytochrome b gene. Proc. R. Soc. Lond. B 265, 793–801 (1998)

    CAS  Article  Google Scholar 

  11. Pitra, C., Fickel, J., Meijaard, E. & Groves, C. Evolution and phylogeny of Old World deer. Mol. Phylogenet. Evol. 33, 880–895 (2004)

    CAS  Article  PubMed  Google Scholar 

  12. Kishino, H. & Hasegawa, M. Evaluation of the maximum likelihood estimate of the evolutionary tree topologies from DNA sequence data, and the branching order in Hominoidea. J. Mol. Evol. 29, 170–179 (1989)

    ADS  CAS  Article  PubMed  Google Scholar 

  13. Guthrie, R. D. Frozen Fauna of the Mammoth Steppe (Univ. of Chicago Press, Chicago, 1990)

    Book  Google Scholar 

  14. Lister, A. M. in Biology and Management of the Cervidae (ed. Wemmer, C. M.) 81–98 (Smithsonian Institution, Washington, DC, 1987)

    Google Scholar 

  15. Azzaroli, A. & Mazza, P. On the possible origin of the Giant Deer genus Megaceroides . Rend. Fis. Acc. Lincei 3(9), 23–32 (1992)

    Article  Google Scholar 

  16. Abbazzi, L. Remarks on the validity of the generic name Praemegaceros Portis 1920, and an overview on Praemegaceros species in Italy. Rend. Fis. Acc. Lincei 15(9), 115–132 (2004)

    Article  Google Scholar 

  17. Croitor, R. & Kostopoulos, D. S. On the systematic position of the large-sized deer from Appolonia, Early Pleistocene, Greece. Palaeontol. Z. 78, 137–159 (2004)

    Article  Google Scholar 

  18. Vislobokova, I. & Hu, C. On the evolution of Megacerines. Vertebr. Palasiat. 28, 150–158 (1990)

    Google Scholar 

  19. Pfeiffer, T. The first complete skeleton of Megaloceros verticornis (Dawkins, 1868) (Cervidae, Mammalia) from Bilshausen (Lower Saxony, Germany): description and phylogenetic implications. Mitt. Mus. Naturk. Berl. Geowiss. 5, 289–308 (2002)

    Google Scholar 

  20. Made, J. v. d. Ungulates from Atapuerca TD6. J. Hum. Evol. 37, 389–413 (1999)

    Article  PubMed  Google Scholar 

  21. Soergel, W. Cervus megaceros mosbachensis n. sp. und die Stammesgeschichte der Riesenhirsche. Abh. Senck. Naturforsch. Ges. 39, 365–407 (1927)

    Google Scholar 

  22. IUCN, The IUCN Red List of Threatened Species (IUCN, Gland, 2004)

    Google Scholar 

  23. Vogler, U. New light upon the taxonomic status of European and Persian fallow deer on the basis of bone remains from Turkey. Tierarzl. Praxis 25, 100–105 (1997)

    CAS  Google Scholar 

  24. Lister, A. M. Evolutionary and ecological origins of British deer. Proc. R. Soc. Edinb. B 82, 205–229 (1984)

    Google Scholar 

  25. Lister, A. M. The morphological distinction between bones and teeth of fallow deer (Dama dama) and red deer (Cervus elaphus). Int. J. Osteoarchaeol. 6, 119–143 (1996)

    Article  Google Scholar 

  26. Swofford, D. L. PAUP* Phylogenetic Analysis using Parsimony (and Other Methods). Version 4.0b10 (Sinauer Associates, Sunderland, Massachusetts, 2002)

    Google Scholar 

  27. Barnes, I., Matheus, P., Shapiro, B., Jensen, D. & Cooper, A. Dynamics of Pleistocene population extinctions in Beringian brown bears. Science 295, 2267–2270 (2002)

    ADS  CAS  Article  PubMed  Google Scholar 

  28. MacHugh, D. E., Edwards, C. J., Bailey, J. F., Bancroft, D. R. & Bradley, D. G. The extraction and analysis of ancient DNA from bone and teeth: a survey of current methodologies. Anc. Biomol. 3, 81–102 (2000)

    CAS  Google Scholar 

  29. Yang, Z. PAML: a program package for phylogenetic analysis by maximum likelihood. Applics BioSci. 13, 555–556 (1997)

    CAS  Google Scholar 

  30. Ronquist, F. & Huelsenbeck, J. P. MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19, 1572–1574 (2003)

    CAS  Article  PubMed  Google Scholar 

Download references


We thank A. Currant, A. Friday, D. Hills, P. Jenkins, P. Kosintsev, L. Martin, N. Monaghan, T. Stuart, A. Vorobiev and E. Westwig for sampling and access to material; P. Grubb, D. MacHugh, C. O'hUigin and K. Wolfe for discussion; T. Burke and A. Cooper for laboratory facilities; P. Forey, J. Masters, A. Mitchell and M. Sánchez-Villagra for advice on cladistics; A. Murray and R. Rabinovich for technical assistance; and V. Geist and Stackpole Books for permission to reproduce the drawings in Fig. 2a. C.J.E. was supported by the Irish Research Council for Science, Engineering and Technology Basic Research Grant Scheme. I.A.vP was funded by BBSRC.

Author information

Authors and Affiliations


Corresponding author

Correspondence to I. Barnes.

Ethics declarations

Competing interests

Sequences are deposited in GenBank under accession numbers AM072730–AM072749. Reprints and permissions information is available at The authors declare no competing financial interests.

Supplementary information

Supplementary Notes

This file contains the Supplementary Methods and Discussion, Supplementary Figures 1 and 2, Supplementary Tables 3-9. This is a detailed account of methods used in both morphological and molecular analysis, with a discussion comparing results with previous studies. Tables and Figures showing sample and radiocarbon details, primer and cloning sequences, distance matrices, and GENBANK accession numbers. (DOC 252 kb)

Supplementary Table S1

Morphological character and character state descriptions; sample sizes and character scores on all taxa. (XLS 53 kb)

Supplementary Table S2

Morphological data transformed ready for analysis. (XLS 33 kb)

Rights and permissions

Reprints and Permissions

About this article

Cite this article

Lister, A., Edwards, C., Nock, D. et al. The phylogenetic position of the ‘giant deer’ Megaloceros giganteus. Nature 438, 850–853 (2005).

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI:

Further reading


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.


Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing