Article | Published:

New infant cranium from the African Miocene sheds light on ape evolution

Nature volume 548, pages 169174 (10 August 2017) | Download Citation


The evolutionary history of extant hominoids (humans and apes) remains poorly understood. The African fossil record during the crucial time period, the Miocene epoch, largely comprises isolated jaws and teeth, and little is known about ape cranial evolution. Here we report on the, to our knowledge, most complete fossil ape cranium yet described, recovered from the 13 million-year-old Middle Miocene site of Napudet, Kenya. The infant specimen, KNM-NP 59050, is assigned to a new species of Nyanzapithecus on the basis of its unerupted permanent teeth, visualized by synchrotron imaging. Its ear canal has a fully ossified tubular ectotympanic, a derived feature linking the species with crown catarrhines. Although it resembles some hylobatids in aspects of its morphology and dental development, it possesses no definitive hylobatid synapomorphies. The combined evidence suggests that nyanzapithecines were stem hominoids close to the origin of extant apes, and that hylobatid-like facial features evolved multiple times during catarrhine evolution.

Access optionsAccess options

Rent or Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.


  1. 1.

    Primate Adaptation and Evolution, 3rd edn (Academic, 2013)

  2. 2.

    in Handbook of Paleoanthropology (eds & ) 1261–1332 (Springer, 2015)

  3. 3.

    & Origin of the Hominidae: the record of African large hominoid evolution between 14 My and 4 My. Yearb. Phys. Anthropol. 31, 49–83 (1988)

  4. 4.

    & in The Primate Fossil Record (ed. ) 385–397 (Cambridge Univ. Press, 2002)

  5. 5.

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

  6. 6.

    et al. Cosmogenic nuclide dating of Sahelanthropus tchadensis and Australopithecus bahrelghazali: Mio–Pliocene hominids from Chad. Proc. Natl Acad. Sci. USA 105, 3226–3231 (2008)

  7. 7.

    et al. Applications of X-ray synchrotron microtomography for non-destructive 3D studies of paleontological specimens. Appl. Phys. Adv. Mater. 83, 195–202 (2006)

  8. 8.

    & A new pliopithecoid genus from the early Miocene of Uganda. J. Hum. Evol. 50, 568–586 (2006)

  9. 9.

    & The lacrimal fossa of Cercopithecoidea, with special reference to cladistic analysis of Old World monkey relationships. Folia Primatol. 60, 133–145 (1993)

  10. 10.

    Mosaic evolution in the origin of the Hominoidea. Folia Primatol. 70, 125–135 (1999)

  11. 11.

    & Facial anatomy of Victoriapithecus and its relevance to the ancestral cranial morphology of Old World monkeys and apes. Am. J. Phys. Anthropol. 92, 329–370 (1993)

  12. 12.

    . et al. Miocene small-bodied ape from Eurasia sheds light on hominoid evolution. Science 350, aab2625 (2015)

  13. 13.

    New fossil anthropoids from the middle Miocene of East Africa and their bearing on the origin of the Oreopithecidae. Am. J. Phys. Anthropol. 71, 265–284 (1986)

  14. 14.

    in The Primate Fossil Record (ed. ) 311–338 (Cambridge Univ. Press, 2002)

  15. 15.

    in Cenozoic Mammals of Africa (eds & ) 429–469 (Univ. California Press, 2010)

  16. 16.

    & Catarrhines from the Middle Miocene (ca. 14.5 Ma) of Kipsaraman, Tugen Hills, Kenya. Anthropol. Sci. 113, 189–224 (2005)

  17. 17.

    & Nondestructive imaging of hominoid dental microstructure using phase contrast X-ray synchrotron microtomography. J. Hum. Evol. 54, 272–278 (2008)

  18. 18.

    & New visions of dental tissue research: tooth development, chemistry, and structure. Evol. Anthropol. 17, 213–226 (2008)

  19. 19.

    et al. Lamellar bone is an incremental tissue reconciling enamel rhythms, body size, and organismal life history. Calcif. Tissue Int. 84, 388–404 (2009)

  20. 20.

    Significance of enamel thickness in hominoid evolution. Nature 314, 260–263 (1985)

  21. 21.

    et al. Dental evidence for ontogenetic differences between modern humans and Neanderthals. Proc. Natl Acad. Sci. USA 107, 20923–20928 (2010)

  22. 22.

    Micro-Computed Tomography of Primate Molars. PhD thesis, Stony Brook Univ. (2006)

  23. 23.

    & Life history theory and dental development in four species of catarrhine primates. J. Hum. Evol. 53, 309–320 (2007)

  24. 24.

    , , & Cerebral complexity preceded enlarged brain size and reduced olfactory bulbs in Old World monkeys. Nat. Commun. 6, 7580 (2015)

  25. 25.

    & Comparative review of the human bony labyrinth. Am. J. Phys. Anthropol. 107 (Suppl. 27), 211–251 (1998)

  26. 26.

    . et al. Evolution of locomotion in Anthropoidea: the semicircular canal evidence. Proc. R. Soc. B 279, 3467–3475 (2012)

  27. 27.

    et al. Palaeontological evidence for an Oligocene divergence between Old World monkeys and apes. Nature 497, 611–614 (2013)

  28. 28.

    & in Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations (eds , & ) 327–362 (Plenum, 1997)

  29. 29.

    , & (eds) in Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations 389–415 (Plenum, 1997)

  30. 30.

    in Hominoid Evolution and Climate Change in Europe Vol. 2 (eds , & ) 231–253 (Cambridge Univ. Press, 2001)

  31. 31.

    , & European Miocene hominids and the origin of the African ape and human clade. Evol. Anthropol. 21, 10–23 (2012)

  32. 32.

    An oreopithecid proximal humerus from the middle Miocene of Maboko Island, Kenya. Int. J. Primatol. 13, 659–677 (1992)

  33. 33.

    , , , & Dental remains of Equatorius africanus from Kipsaramon, Tugen Hills, Baringo District, Kenya. J. Hum. Evol. 42, 39–62 (2002)

  34. 34.

    , , , & Geology and geochronology of the middle Miocene Kipsaramon site complex, Muruyur Beds, Tugen Hills, Kenya. J. Hum. Evol. 42, 11–38 (2002)

  35. 35.

    , & Miocene Cercopithecoidea from the Tugen Hills, Kenya. J. Hum. Evol. 59, 465–483 (2010)

  36. 36.

    Geology of the Area South of Lodwar, Degree Sheet 18, N.E. Quarter. Report No. 87 (Geological Survey of Kenya, 1971)

  37. 37.

    et al. in Geoscience of Rift Systems - Evolution of East Africa. AAPG Studies in Geology Vol. 44 (ed. ) 19–54 (American Association of Petroleum Geologists, 1999)

  38. 38.

    , & in The Geologic Time Scale (eds , , & ) 923–978 (Elsevier, 2012)

  39. 39.

    et al. Synchronizing rock clocks of Earth history. Science 320, 500–504 (2008)

  40. 40.

    & Subcommission on geochronology: convention on the use of decay constants in geo- and cosmochronology. Earth Planet. Sci. Lett 36, 359–362 (1977)

  41. 41.

    ., , & Irradiation of Samples for 40Ar/39Ar Dating Using The Geological Survey TRIGA Reactor. US Geological Survey Professional Paper No. 1176 (US Geological Survey, 1981)

  42. 42.

    & 40Ar/39Ar age spectra of Apollo 15 impact melt rocks by laser heating and their bearing on the history of lunar basin formation. J. Geophys. Res. 98 (E7), 13085–13095 (1993)

  43. 43.

    The Comparative Morphology and Phylogeny of The Human Bony Labyrinth. PhD thesis, Utrecht Univ. (1993)

  44. 44.

    et al. The primate semicircular canal system and locomotion. Proc. Natl Acad. Sci. USA 104, 10808–10812 (2007)

  45. 45.

    & Body mass in comparative primatology. J. Hum. Evol. 32, 523–559 (1997)

  46. 46.

    A Taxonomic Review and Phylogenetic Analysis of Homo habilis. PhD thesis, Univ. Liverpool (1987)

  47. 47.

    , , , & Postcranial estimates of body weight in Proconsul, with a note on a distal tibia of P. major from Napak, Uganda. Am. J. Phys. Anthropol. 97, 391–402 (1995)

  48. 48.

    , & PAST: paleontological statistics software package for education and data analysis. Palaeontol. Electronica 4, 4 (2001)

  49. 49.

    & New partial cranium of Dryopithecus Lartet, 1863 (Hominoidea, Primates) from the Upper Miocene of Can-Llobateres, Barcelona, Spain. J. Hum. Evol. 29, 101–139 (1995)

  50. 50.

    A revised systematic scheme for the Eurasian Miocene fossil Hominidae. J. Hum. Evol. 33, 449–477 (1997)

  51. 51.

    , , & A mandible of Mabokopithecus clarki sheds new light on oreopithecid evolution. Am. J. Phys. Anthropol. 105 (Suppl. 26), 109 (1998)

  52. 52.

    (ed.) in Phylogenetic Analysis of Morphological Data. Comparative Evolutionary Series 115–145 (Smithsonian Institution Press, 2000)

  53. 53.

    , , , & Morphological systematics of the kipunji (Rungwecebus kipunji) and the ontogenetic development of phylogenetically informative characters in the Papionini. J. Hum. Evol. 60, 731–745 (2011)

  54. 54.

    et al. Endocranial volumes of primate species: scaling analyses using a comprehensive and reliable data set. J. Hum. Evol. 55, 967–978 (2008)

Download references


We thank the Government of Kenya for permission to do this research, the National Museums of Kenya, the American Museum of Natural History, Harvard Museum of Comparative Zoology, the Musée des Confluences of Lyon, and the Digital Morphology Museum (KUPRI) for access to specimens in their care, and the Turkana Basin Institute for field support. The Leakey Foundation and trustee G. Getty, the Foothill-De Anza Foundation, De Anza College Design and Manufacturing Technologies Department, the National Geographic Society, the European Synchrotron Radiation Facility (beamlines ID 19 and BM5), and the Max Planck Society funded fieldwork or laboratory studies. We are grateful to M. Kibunjia and F. Manthi for facilitating permission to transport the specimen to Grenoble for scanning, and to D. Berthet, E. Delson, M. Hill, J.-J. Hublin, D. Plotzki, and H. Temming for facilitating computed tomography scans of comparative specimens. We are grateful to R. David, M. Gilleland, L. Gonzales, T. Harrison, J. Kelley, Y. Kunimatsu, L. Martin, M. Leakey, K. O’Maley, M. Pickford, C. Rendiger, B. Sokhi, and A. Stoessel for help with this research. We thank the field crew, A. Ekes, B. Ewoi, J. Ekusi, A. Ekuon, C. Nyete, and T. Ngundo, for their tireless efforts in the field that led to the discovery of the specimen, C. Kiarie for preparing the fossil, and B. Monroy for preparing some of the graphics. We particularly thank J. Ekusi for discovering KNM-NP 59050.

Author information


  1. Department of Anthropology, De Anza College, Cupertino, California 95104, USA

    • Isaiah Nengo
  2. Turkana Basin Institute, Social and Behavioral Sciences Building N-507, Stony Brook University, Stony Brook, New York 11794-4364, USA

    • Isaiah Nengo
  3. European Synchrotron Radiation Facility, CS-40220, F-38043 Grenoble Cedex 09, France

    • Paul Tafforeau
    •  & Camille Berruyer
  4. Department of Anthropology, Hunter College of the City University of New York, 695 Park Avenue, New York, New York 10065, USA

    • Christopher C. Gilbert
  5. The Graduate Center of the City University of New York, 365 5th Avenue, New York, New York 10016, USA

    • Christopher C. Gilbert
    •  & Kelsey D. Pugh
  6. New York Consortium in Evolutionary Primatology, New York, New York, USA

    • Christopher C. Gilbert
    •  & Kelsey D. Pugh
  7. Department of Anatomical Sciences, Health Sciences Center, Stony Brook University, Stony Brook, New York 11794-8081, USA

    • John G. Fleagle
  8. Department of Anthropology, Wake Forest University, Winston Salem, North Carolina 27109-7807, USA

    • Ellen R. Miller
  9. Department of Anthropology, Rutgers University, New Brunswick, New Jersey 08901-1414, USA

    • Craig Feibel
  10. Department of Earth and Planetary Sciences, Rutgers University, Piscataway, New Jersey 08854-8066, USA

    • Craig Feibel
    •  & Zachary Engle
  11. Department of Earth Sciences, University of Minnesota, Minneapolis, Minnesota 55455, USA

    • David L. Fox
    •  & Josh Feinberg
  12. Department of Geological Sciences, Salem State University, Salem, Massachusetts 01970, USA

    • Sara Mana
  13. Department of Cell and Developmental Biology, University College London, London WC1E 6BT, UK

    • Fred Spoor
  14. Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig 04103, Germany

    • Fred Spoor


  1. Search for Isaiah Nengo in:

  2. Search for Paul Tafforeau in:

  3. Search for Christopher C. Gilbert in:

  4. Search for John G. Fleagle in:

  5. Search for Ellen R. Miller in:

  6. Search for Craig Feibel in:

  7. Search for David L. Fox in:

  8. Search for Josh Feinberg in:

  9. Search for Kelsey D. Pugh in:

  10. Search for Camille Berruyer in:

  11. Search for Sara Mana in:

  12. Search for Zachary Engle in:

  13. Search for Fred Spoor in:


I.N. and F.S. designed the study. I.N., C.F., E.R.M., and D.L.F. did field research. J.G.F., C.C.G., F.S., and K.D.P. collected and analysed the comparative data on gross dental and cranial morphology, and J.G.F. and C.C.G. prepared the diagnosis. P.T., C.B., and F.S. did the image analyses and made virtual reconstructions, P.T. and C.B. performed the dental development study, C.F., S.M., D.L.F., J.F., and Z.E. studied the geological context, and C.C.G. and K.D.P. did the phylogenetic analyses. F.S., I.N., and E.R.M. wrote the paper with contributions from C.C.G., P.T., and J.G.F.

Competing interests

The authors declare no competing financial interests.

Corresponding author

Correspondence to Isaiah Nengo.

Reviewer Information Nature thanks B. Benefit and the other anonymous reviewer(s) for their contribution to the peer review of this work.

Publisher's note: Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Extended data

Supplementary information

PDF files

  1. 1.

    Supplementary Information

    This file contains Supplementary Note 1, Supplementary Tables 1-3 and additional references.

  2. 2.

    Reporting Summary

Zip files

  1. 1.

    Supplementary Data 1

    This zipped file contains unerupted tooth crowns of KNM-NP 59050. Three-dimensional surface models of the left and right I1, M1 and M2 are provided as 3D PDFs. Surfaces rendered from segmented synchrotron microtomographic images.

Excel files

  1. 1.

    Supplementary Data 2

    Comparative cranial and dental measurements. This file contains measurements of KNM-NP 59050 and all individual juvenile and adult specimens in the comparative sample.

  2. 2.

    Supplementary Data 3

    40Ar/39Ar incremental-heating data and analytical conditions for sample 15-NPD-03.

  3. 3.

    Supplementary Data 4

    Calculation sheet for dental development of KNM-NP 59050. The calculations to determine the age at death and the general developmental timing of Nyanzapithecus alesi based on dental increments.

  4. 4.

    Supplementary Data 5

    Morphological Characters used in Phylogenetic Analyses. This spreadsheet lists the characters scored in this study along with character state definitions, character type designations, and the source of each character.

Text files

  1. 1.

    Supplementary Data 6

    The matrix used in phylogenetic analysis. This file contains the Nexus file with the character matrix used in this study, written with Mesquite version 2.75, build 566 ( ). The file can be read with any text editor.

About this article

Publication history





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.