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The rise and fall of the Old World savannah fauna and the origins of the African savannah biome

An Author Correction to this article was published on 15 January 2018

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Despite much interest in the ecology and origins of the extensive grassland ecosystems of the modern world, the biogeographic relationships of savannah palaeobiomes of Africa, India and mainland Eurasia have remained unclear. Here we assemble the most recent data from the Neogene mammal fossil record in order to map the biogeographic development of Old World mammalian faunas in relation to palaeoenvironmental conditions. Using genus-level faunal similarity and mean ordinated hypsodonty in combination with palaeoclimate modelling, we show that savannah faunas developed as a spatially and temporally connected entity that we term the Old World savannah palaeobiome. The Old World savannah palaeobiome flourished under the influence of middle and late Miocene global cooling and aridification, which resulted in the spread of open habitats across vast continental areas. This extensive biome fragmented into Eurasian and African branches due to increased aridification in North Africa and Arabia during the late Miocene. Its Eurasian branches had mostly disappeared by the end of the Miocene, but the African branch survived and eventually contributed to the development of Plio–Pleistocene African savannah faunas, including their early hominins. The modern African savannah fauna is thus a continuation of the extensive Old World savannah palaeobiome.

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Fig. 1: GFRI results of all the localities from the Old World Neogene used in this study.
Fig. 2: Maps of tooth crown height for the Old World Neogene.
Fig. 3: Raup–Crick genus-level faunal similarity to the Lower Nawata.
Fig. 4: Three-dimensional spatiotemporal Raup–Crick GFRI interpolations with values above 0.7 from 12 to 1.8 Ma for the Nawatian (blue), Pikermian (red) and Baodean (yellow) faunas.
Fig. 5: Climate model simulations for precipitation over the Old World in the present day.

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  • 15 January 2018

    In the version of this Article originally published, each of the five panels in Fig. 5 incorrectly contained a black diagonal line across the plot. This has now been corrected.


  1. Watson, J. E. M. et al. Catastrophic declines in wilderness areas undermine global environment targets. Curr. Biol. 26, 1–6 (2016).

    Article  Google Scholar 

  2. Ceballos, G. & Ehrlich, P. R. Global mammal distributions, biodiversity hotsposts, and conservation. Proc. Natl Acad. Sci. USA 103, 19374–19379 (2006).

    Article  CAS  Google Scholar 

  3. Kingdon, J. Field Guide to African Mammals (Academic Press, San Diego, CA, 1997).

    Google Scholar 

  4. White, F. The Vegetation of Africa: A Descriptive Memoir to Accompany the UNESCO/AETFAT/UNSO Vegetation Map of Africa (United Nations Educational, Scientific and Cultural Organization, 1984).

  5. Sankaran, M. et al. Determinants of woody cover in African savannas. Nature 438, 846–849 (2005).

    Article  CAS  Google Scholar 

  6. Eronen, J. et al. Distribution history and climatic controls of the Late Miocene Pikermian chronofuana. Proc. Natl Acad. Sci. USA 106, 11867–11871 (2009).

    Article  CAS  Google Scholar 

  7. Solounias, N., Plavcan, J. M., Quade, J. & Witmer, L. in The Evolution of Neogene Terrestrial Ecosystems in Europe (eds Agusti, J., Rook, L. & Andrews, P.) 436–453 (Cambridge Univ. Press, Cambridge, 1999).

  8. Solounias, N., Semprebon, G. M., Mihlbachler, M. & Rivals, F. in Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology (eds Wang, X., Fortelius, M. & Flynn, L.) 676–692 (Columbia Univ. Press, New York, 2013).

  9. Zachos, J., Pagani, M., Sloan, L., Thomas, E. & Billups, K. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292, 686–693 (2001).

    Article  CAS  Google Scholar 

  10. Mirzaie Ataabadi, M., et al. in Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology (eds Wang, X., Fortelius, M. & Flynn, L.) 546–565 (Columbia Univ. Press, New York, 2013).

  11. Fortelius, M. et al. Evolution of Neogene mammals in Eurasia: environmental forcing and biotic interactions. Annu. Rev. Earth Planet. Sci. 42, 579–604 (2014).

    Article  CAS  Google Scholar 

  12. Solounias, N., Rivals, F. & Semprebon, G. Dietary interpretation and paleoecology of herbivores from Pikermi and Samos (late Miocene of Greece). Paleobiology 36, 113–136 (2010).

    Article  Google Scholar 

  13. Bernor, R. L. in New Interpretations of Ape and Human Ancestry (eds Ciochon, R. L. & Corruccini, R. S.) 21–64 (Plenum Press, New York, 1983).

  14. Janis, C. M. & Fortelius, M. On the means whereby mammals achieve increased functional durability of their dentitions, with special reference to limiting factors. Biol. Rev. 63, 197–230 (1988).

    Article  CAS  Google Scholar 

  15. Jernvall, J. & Fortelius, M. Common mammals drive the evolutionary increase of hypsodonty in the Neogene. Nature 417, 538–540 (2002).

    Article  CAS  Google Scholar 

  16. Agustí, J. in Handbook of Paleoanthropology (eds Henke, W. & Tattersal, I.) 979–1010 (Springer Books, Berlin/Heidelberg, 2015).

  17. Koufos, D. G. in Handbook of Paleoanthropology (eds Henke, W. & Tattersal, I.) 1761–1790 (Springer Books, Berlin/Heidelberg, 2015).

  18. Begun, D. R., Nargolwalla, M. C. & Kordos, L. European Miocene hominids and the origin of the African ape and human clade. Evol. Anthr. 21, 10–23 (2012).

    Article  Google Scholar 

  19. Nakaya, H. Faunal change of late Miocene Africa and Eurasia: mammalian fauna from the Namurungule Formation, Samburu Hills, Northern Kenya. Afr. Stud. Monogr. 20, 1–112 (1994).

    Google Scholar 

  20. Thomas, H. Les bovidae (Artiodactyla: Mammalia) du miocene du sous-continent indien, de la peninsule arabique et de l’afrique: biostratigraphie, biogeographie et ecologie. Palaeogeogr. Palaeoclimatol. Palaeoecol. 45, 251–299 (1984).

    Article  Google Scholar 

  21. Bernor, R. L., Rook, L. & Haile-Selassie, Y. in Ardipithecus kadabba; Late Miocene Evidence from the Middle Awash, Ethiopia (eds Haile-Selassie, Y. & WoldeGabriel, G.) 549–563 (Univ. California Press, Berkeley, CA, 2009).

  22. Katoh, S. et al. New geological and palaeontological age constraint for the gorilla–human lineage split. Nature 530, 215–218 (2016).

    Article  CAS  Google Scholar 

  23. Bibi, F. Mio–Pliocene faunal exchanges and African biogeography: the record of fossil bovids. PLoS ONE 6, e16688 (2011).

    Article  CAS  Google Scholar 

  24. Leakey, M. G. & Harris, J. M. in Lothagam: The Dawn of Humanity in Eastern Africa (eds Leakey, M. G. & Harris, J. M.) 625–655 (Columbia Univ. Press, New York, 2003).

  25. Kaya, F. Paleobiogeographic and Paleoecologic Development of the Old World Savanna Paleobiome. PhD thesis, Univ. Helsinki (2017).

  26. Boaz, N. T. A view to the south: Eo-Sahabi palaeoenvironments compared and implications for hominid origins in Neogene North Africa. Garyounis Sci. Bull. Spec. Issue 5, 291–308 (2008).

    Google Scholar 

  27. Bibi, F., Hill, A., Beech, M. & Yasin, W. in Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology (eds Wang, X., Fortelius, M. & Flynn, L.) 583–594 (Columbia Univ. Press, New York, 2013).

  28. Gilbert, C. C., Bibi, F., Hill, A. & Beech, M. J. Early guenon from the late Miocene Baynunah Formation, Abu Dhabi, with implications for cercopithecoid biogeography and evolution. Proc. Natl Acad. Sci. USA 111, 10119–10124 (2014).

    Article  CAS  Google Scholar 

  29. Schuster, M. et al. The age of the Sahara desert. Science 311, 821 (2006).

    Article  CAS  Google Scholar 

  30. Ségalen, L., Lee-Thorp, J. A. & Cerling, T. E. Timing of C4 grass expansion across sub-Saharan Africa. J. Hum. Evol. 53, 549–559 (2007).

    Article  Google Scholar 

  31. Cerling, T. E. Development of grasslands and savannas in East Africa during the Neogene. Palaeogeogr. Palaeoclimatol. Palaeoecol. 97, 241–247 (1992).

    Article  Google Scholar 

  32. Bernor, R. L. New apes fill the gap. Proc. Natl Acad. Sci. USA 104, 19661–19662 (2007).

    Article  CAS  Google Scholar 

  33. Rook, L. & Bernor, R. L. Ancestry of the African ape–human clade. Palaeontogr. Ital. 89, 30–31 (2003).

    Google Scholar 

  34. Kaya, F. Anadolu’nun Neojen Donem memeli paleobioyocografyasi ve paleoekolojisi. Kebikec 43, 157–176 (2017).

    Google Scholar 

  35. Bernor, R. L. A zoogeographic theater and a biochronologic play: the time/biofacies phenomena of Eurasian and African Miocene mammal provinces. Paléob. Cont. 14, 121–142 (1984).

    Google Scholar 

  36. Beden, M. & Brunet, M. Faunes de mammifères et paléobio-géographie des domaines indiens et péri-i ndiens au Néogène. Sci. Terre 47, 61–87 (1986).

    Google Scholar 

  37. Fortelius, M., et al. in The Evolution of Western Eurasian Neogene Mammal Faunas (eds Bernor, R. L., Fahlbusch, V. & Mittmann, H. W.) 414–448 (Columbia Univ. Press, New York, 1996).

  38. The NOW Community. New and Old Worlds Database of Fossil Mammals (NOW). Licensed under CC BY 4.0. (2017).

  39. Werdelin, L. & Sanders, W. J. Cenozoic Mammals of Africa (Univ. California Press, Berkeley, CA, 2010).

  40. Hilgen, F., Lourens, L. & Van Dam, J. in The Geologic Time Scale 2012 (eds Gradstein, F., Ogg, J., Schmitz, M. & Ogg, G.) 923–978 (Elsevier, Amsterdam, 2012).

  41. Werdelin, L. in Cenozoic Mammals of Africa (eds Werdelin, L. & Sanders, W. J.) 27–44 (Univ. California Press, Berkeley, CA, 2010).

  42. Steininger, F. F., et al. in The Evolution of Western Eurasian Neogene Mammal Faunas (eds Bernor, R. L., Fahlbusch, V. & Mittmann, H. W.) 7–46 (Columbia Univ. Press, New York, 1996).

  43. Hammer, Q., Harper, D. A. T. & Ryan, P. D. PAST: paleontological statistics software package for education and data analysis. Palaeontol. Electron. 4, 4 (2001).

    Google Scholar 

  44. Raup, D. & Crick, R. E. Measurement of faunal similarity in paleontology. J. Paleo. 53, 1213–1227 (1979).

    Google Scholar 

  45. Aranz Geo. LeapFrog Geo Software v.3.1 64 bit (Aranz Geo Ltd., 2016).

  46. Fortelius, M. et al. Fossil mammals resolve regional patterns of Eurasian climate change during 20 million years. Evol. Ecol. Res. 4, 1005–1016 (2002).

    Google Scholar 

  47. Liu, L. et al. Dental functional traits of mammals resolve productivity in terrestrial ecosystems past and present. Proc. R. Soc. B 279, 2793–2799 (2012).

    Article  Google Scholar 

  48. Micheels, A. et al. Analysis of heat transport mechanisms from a late Miocene model experiment with a fully-coupled atmosphere–ocean general circulation model. Palaeogeogr. Palaeoclimatol. Palaeoecol. 304, 337–350 (2011).

    Article  Google Scholar 

  49. Tang, H., Eronen, J. T., Micheels, A. & Ahrens, B. Strong interannual variation of the Indian summer monsoon in the late Miocene. Clim. Dyn. 41, 135–153 (2013).

    Article  CAS  Google Scholar 

  50. Kottek, M., Grieser, J., Beck, C., Rudolf, B. & Rubel, F. World map of the Koppen–Geiger climate classification updated. Meteorol. Z. 15, 259–263 (2006).

    Article  Google Scholar 

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We thank A. Karme and G. Berni for their guidance with the three-dimensional geology software and the Department of Geosciences and Geography at the University of Helsinki for providing LeapFrog Geo. We are grateful to A. H. Kaya for language improvement. J.T.E. acknowledges support from the Marie Curie Actions of the EC and Kone Foundation. M.F. acknowledges funding from the Academy of Finland and an award from the Alexander von Humboldt Foundation. The work of F.K. was supported by an Academy of Finland grant to M.F.

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Authors and Affiliations



F.K., M.F., J.T.E. and F.B. designed the research. F.K. updated the taxonomic identifications of African localities in the NOW database, performed the computational similarity and mean ordinated hypsodonty analyses and designed the figures. I.Z. performed the sensitivity tests and analysis of the computational methodology. H.T. (T.H.) performed the climate modelling. All authors participated in the interpretation of the results and wrote the paper. M.F. supervised the study. Some of the same content, including an earlier manuscript version of this paper, was included in the PhD thesis of Ferhat Kaya25. A single-authored article published by Ferhat Kaya in the Turkish social sciences journal Kebikec34 used the same data and methods to analyse the Anatolian subset of localities. The Turkish text is closely focused on Anatolia but the English abstract of that paper mentions some of the main conclusions of this paper.

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Correspondence to Ferhat Kaya.

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Supplementary Table 1

List of some selected localities with their age information and GFRI values to the Lower Nawata in Figs. 1, 3 and 4. Abbreviations; PDL stands for Present Day Location, N symbolize stands for sample size, and GFRI stands for genus level faunal resemblance indices

Supplementary Table 4

List of selected early and middle Miocene sites with age information and GFRI values used for Raup-Crick GFRI analysis in Figs. 1, 3 and 4.

Supplementary Table 5

List of selected late Miocene sites with age information and GFRI values used for Raup-Crick GFRI analysis in Figs. 1, 3 and 4.

Supplementary Table 6

List of selected Pliocene sites with age information and GFRI values used for Raup-Crick GFRI analysis in Figs. 1, 3 and 4.

Supplementary Table 7

The locality list with age information and mean hypsodonty values used for mean ordinated hypsodonty analysis in Figs. 1 and 2.

Supplementary Figure 11

Biogeographical development of the OWSP during the late Miocene. Video animation of the Raup-Crick GFRI with values above 0.7 from 12 to 1.8 Ma for the Nawatian (blue), Pikermian (red), and Baodean (yellow). The increase in the similarity among the Eurasian and African early late Miocene faunas coincides with the parallel expansion of the Nawatian, the Pikermian and the Baodean resulted in the birth of the Old World savanna paleobiome that reaches its climax during the middle late Miocene. The Pikermian decreases suddenly in western and central Eurasia, the Baodean chronofauna survives into the Pliocene and disappear, and the Nawatian eventually evolved to the East African modern savanna fauna.

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Kaya, F., Bibi, F., Žliobaitė, I. et al. The rise and fall of the Old World savannah fauna and the origins of the African savannah biome. Nat Ecol Evol 2, 241–246 (2018).

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