International Journal of Primatology

, Volume 29, Issue 3, pp 723–741 | Cite as

A Taxonomic Reassessment of Cacajao melanocephalus Humboldt (1811), with the Description of Two New Species

  • Jean P. BoubliEmail author
  • Maria Nazareth F. da Silva
  • Manuella V. Amado
  • Tomas Hrbek
  • Francisco Boavista Pontual
  • Izeni P. Farias


The author of the last published systematic review of Cacajao recognized 2 subspecies of black-headed uakaris (black uakaris): Cacajao melanocephalus melanocephalus and C. m. ouakary. As a result of a series of black uakari surveys and collecting expeditions to several tributaries of the Rio Negro and of morphological and molecular analyses of museum specimens and specimens we collected during field expeditions, we reassess their taxonomy. We describe a newly discovered species of black uakari from the Rio Aracá, a left bank tributary of the Rio Negro, Amazonas, Brazil. We also show that ouakary is a junior synonym of melanocephalus and provide a new name and a new description for Cacajao melanocephalus melanocephalus in the Pico da Neblina region of Brazil and Venezuela. Based on genetic, morphological, and ecological evidence, we propose that there are 3 species of black uakaris. We named the Rio Aracá species Cacajao ayresi sp. nov. (Ayres uakari) in honor of the late José Márcio Ayres, a pioneer in uakari research and conservation. We named the Neblina black uakari Cacajao hosomi, after the Yanomami word for uakaris. The new taxonomic arrangement provided here implies that the conservation status of black uakaris needs to be reassessed.


Amazonia black-headed uakari Brazil Cacajao melanocephalus cytochrome b geographic distribution New World primates Pithecines Pleistocene rain forest speciation phylogeny 



We thank the Yanomami people and the ribeirinho communities of Rio Negro Basin for their help and support during our surveys. We also thank the Catholic and evangelic missionaries who provided assistance to our team in their remote outposts. We thank Drs. Bruce Patterson (FMNH), James F. Whatton (USNM), Al Gardner (USNM), and Richard Thorington (USNM) for providing photographs of black uakari specimens deposited under their care. We thank Drs. Colin Groves, Jim Patton, and 2 anonymous reviewers for their in-depth review of the manuscript. We thank Drs. José de Souza e Silva Júnior and Colin Groves for suggesting that the taxa examined here should receive species status. Pictures of uakari skins were kindly provided by Glenn Shepard. We thank the Brazilian Army, FUNAI, and IBAMA for research permits and logistical support. The Sustainable Development of the Brazilian Biodiversity Program (PROBIO/MMA/BIRD/GEF/CNPq), the Zoological Society of San Diego, and the University of Auckland funded the surveys. International Foundation for Science (IFS) grants to IPF funded in part the molecular analyses. IBAMA granted permission to conduct fieldwork and to collect tissue samples (license no. 005/2005 - CGFAU/LIC). This study is dedicated to the memory of Arnaldo Cabeludo, our friend and dedicated field assistant.


  1. Ayres, J. M. (1989). Comparative feeding ecology of the uakari and bearded saki, Cacajao and Chiropotes. Journal of Human Evolution, 18, 697–716.CrossRefGoogle Scholar
  2. Barnett, A., & Cunha, A. C. D. (1991). The golden-backed uakari or the upper Rio Negro Brazil. Oryx, 25, 80–88.Google Scholar
  3. Bonvicino, C. R., Boubli, J. P., Otazú, I. B., Almeida, F. C., Nascimento, F. F., Coura, J. R., & Seuanez, H. N. (2003). Morphologic, karyotypic and molecular evidence of a new form of Chiropotes (Primates, Pitheciinae). American Journal of Primatology, 61, 123–133.PubMedCrossRefGoogle Scholar
  4. Boubli, J. P. (1993). Southern expansion of the geographical distribution of Cacajao melanocephalus melanocephalus. International Journal of Primatology, 14, 933–937.CrossRefGoogle Scholar
  5. Boubli, J. P. (1997). A study of the black uakari, Cacajao melanocephalus, in the Pico da Neblina National Park, Brazil. Neotropical Primates, 5, 113–115.Google Scholar
  6. Boubli, J. P. (1999). The feeding ecology of the black-headed uakari Cacajao melanocephalus, in Pico da Neblina National Park, Brazil. International Journal of Primatology, 20, 719–749.CrossRefGoogle Scholar
  7. Boubli, J. P. (2002). Western Extension of the range of bearded sakis: a possible new taxon of Chiropotes sympatric with Cacajao in the Pico da Neblina National Park, Brazil. Neotropical Primates, 10(1), 1–4.Google Scholar
  8. Brown, W. M., George Jr., M., & Wilson, A. C. (1979). Rapid evolution of animal mitochondrial DNA. Proceedings of the National Academy of Sciences of the United States of America, 76, 1967–1971.PubMedCrossRefGoogle Scholar
  9. Clement, M., Posada, D., & Crandall, K. A. (2000). TCS: A computer program to estimate gene genealogies. Molecular Ecology, 9, 1657–1659.PubMedCrossRefGoogle Scholar
  10. Colinvaux, P. A., De Oliveira, P. E., Moreno, J. E., Miller, M. C., & Bush, M. B. (1996). A long pollen record from lowland Amazonia: Forest and cooling in glacial times. Science, 274, 85–88.CrossRefGoogle Scholar
  11. Groves, C. (2001). Primate taxonomy. Washington, DC: Smithsonian Institution Press.Google Scholar
  12. Haffer, J. (1997). Alternative models of vertebrate speciation in Amazonia: An overview. Biodiversity and Conservation, 6, 451–476.CrossRefGoogle Scholar
  13. Hall, T. (1999). BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.Google Scholar
  14. Handley, C. O. (1976). Mammals of the Smithsonian Venezuelan Project. Brigham Young University Science Bulletin, Biology Series, 20, 1–91.Google Scholar
  15. Hernandéz-Camacho, J., & Cooper, R. W. (1976). The nonhuman primates of Colombia. In R. W. Thorington Jr., & P. G. Heltne (Eds.) Neotropical Primates. Field Studies and Conservation (pp. 35–69). Washington, DC: National Academy of Science.Google Scholar
  16. Hershkovitz, P. (1987). Uacaris, New World monkeys of the genus Cacajao: A preliminary taxonomic review with a description of a new subspecies. American Journal of Primatology, 12, 1–53.CrossRefGoogle Scholar
  17. Hilton-Taylor, C. (compiler) (2000). 2000 IUCN Red List of Threatened Species. Switzerland and Cambridge, U.K.: IUCN, Gland.Google Scholar
  18. Ho, S. Y. W., Phillips, M. J., Cooper, A., & Drummond, A. J. (2005). Time dependency of molecular rate estimates and systematic overestimation of recent divergence times. Molecular Biology and Evolution, 22, 1561–1568.PubMedCrossRefGoogle Scholar
  19. Humboldt, A. (1811). Recueil d’observations de zoologie et d’anatomie compare, faites dans l’ocean atlantique dans l’interior du nouveau continent et dans la mer du sud pendant les années 1799, 1800, 1801, 1802 et 1803. Part 2. Humboldt, A de, Bonpland, A. (eds.) Levrault Schoell, Paris, 1811–1812.Google Scholar
  20. Humboldt, A., & Bompland, A. (1907). Personal narrative of travels to the equinoctial regions of America during the years, 1799–1804/by Alexander von Humboldt and Aimé Bonpland; translated from the French of Alexander von Humboldt and edited by Thomasina Ross. 3 vols. Bell, London.Google Scholar
  21. Irwin, D. M., Kocher, T. D., & Wilson, A. C. (1991). Evolution of the cytochrome b gene of mammals. Journal of Molecular Evolution, 32, 128–144.PubMedCrossRefGoogle Scholar
  22. Kocher, T. D., Thomas, W. K., Meyer, A., Edwards, S. V., Pääbo, S., & Villablanca, F. X. (1989). Dynamics of mtDNA evolution in animals: Amplification and sequencing with conserved primers. Proceedings of the National Academy of Sciences of the United States of America, 86, 6196–6200.PubMedCrossRefGoogle Scholar
  23. Moritz, C. (2002). Strategies to protect biological diversity and the evolutionary processes that sustain it. Systematics Biology, 51, 238–254.CrossRefGoogle Scholar
  24. Rodríguez, F. J., Oliver, J. L., Marín, A., & Medina, J. R. (1990). The general stochastic model of nucleotide substitution. Journal of Theoretical Biology, 142, 485–501.PubMedCrossRefGoogle Scholar
  25. Ronquist, F., & Huelsenbeck, J. P. (2003). MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.PubMedCrossRefGoogle Scholar
  26. Sambrook, J., Fritsch, E. F., & Maniatis, T. (1989). Molecular Cloning: A Laboratory Manual. Cold Spring Harbor, NY: Cold Spring Harbor Laboratory Press.Google Scholar
  27. Spix, J. B. (1823). Simiarum et vespertilliarum brasiliensis species novae; ou histoire naturelle des espéces nouvelles des singes et de chauve-souris observée et recueilies pendant le voyage dans l’interieur du Bresil. Monaco.Google Scholar
  28. Swofford, D. L. (2002). PAUP*. Phylogenetic Analysis Using Parsimony (* and Other Methods), Beta Version 4b10. Sunderland, MA: Sinauer Associates.Google Scholar
  29. Thompson, J. D., Higgins, D. G., & Gibson, T. J. (1996). CLUSTAL W: Improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position specific gap penalties and weight matrix choice. Nucleic Acids Research, 22, 4673–4680.CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media, LLC 2008

Authors and Affiliations

  • Jean P. Boubli
    • 1
    Email author
  • Maria Nazareth F. da Silva
    • 2
  • Manuella V. Amado
    • 3
  • Tomas Hrbek
    • 4
  • Francisco Boavista Pontual
    • 5
  • Izeni P. Farias
    • 3
  1. 1.Department of AnthropologyUniversity of AucklandAucklandNew Zealand
  2. 2.Coleção de Mamíferos, CPECInstituto Nacional de Pesquisas da AmazôniaManausBrazil
  3. 3.Laboratório de Evolução e Genética Animal, Departamento de Biologia, ICBUniversidade Federal do AmazonasManausBrazil
  4. 4.Department of BiologyUniversity of Puerto RicoSan JuanPuerto Rico
  5. 5.Department of Environmental Science, Policy and ManagementUniversity of CaliforniaBerkeleyCalifornia, USA

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