Species Recognition in Middle Miocene Hominoids

  • Lawrence B. Martin
  • Peter Andrews
Part of the Advances in Primatology book series (AIPR)


This chapter addresses the issue of species recognition in fossil samples. The particular example that is examined is the extensive collection of dental and gnathic remains from the middle Miocene locality at Paşalar, Turkey. This site is unusual in that available sedimentological and taphonomic evidence indicate that it was accumulated extremely rapidly and from a very localized area (Al-pagut et al., 1990a; Andrews and Alpagut, 1990; Andrews and Ersoy, 1990; Bestland, 1990), so that the mammalian fauna is comparable to a museum collection of modern animals made from one locality. In fact, the fauna may be sampled from within 3–5 km of the site, which would include only a few hundred meters of altitude at most. The sediments may have been deposited in hours or days, so that the temporal range of the fauna is determined by the period of predepositional skeletal preservation. Therefore, it might be anticipated that a species at Paşalar might be less variable than modern comparative samples collected from a variety of geographical localities. The Paşalar sample thus provides an unusual opportunity to examine analytical methods that may be used to determine species numbers in such an assemblage.


Sexual Dimorphism Fossil Record Middle MIOCENE Species Recognition Incisal Edge 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Abel, O. 1902. Zwei neue Menscenaffen aus den Leithakalkbildungen des Wiener Beckens. Sitzungsberichte der Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Classe 111:1171–1207.Google Scholar
  2. Alpagut, B., Andrews, P., and Martin, L. 1990a. Miocene Paleoecology of Paçalar, Turkey, in: E. H. Lindsay, V. Fahlbusch, and P. Mein (eds.), European Neogene Mammal Chronology, pp. 443–459. NATO ASI Series A: Life Sciences Vol. 180. Plenum Press, New York.CrossRefGoogle Scholar
  3. Alpagut, B., Andrews, P., and Martin, L. 1990b. New hominoid specimens from the Middle Miocene site at Paşalar, Turkey. J . Hum. Evol 19:397–422.CrossRefGoogle Scholar
  4. Andrews, P. J. 1978. A revision of the Miocene Hominoidea of East Africa. Bull. Br Mus. (Nat. Hist) (Geology) 30(2):85–224.Google Scholar
  5. Andrews, P. 1981. Species diversity and diet in monkeys and apes during the Miocene, in: C. B. Stringer (ed.), Aspects of Human Evolution, pp. 25–61 Taylor and Francis, London.Google Scholar
  6. Andrews, P. 1990. Palaeoecology of the Miocene fauna from Paşalar, Turkey. J. Hum. Evol. 19:569–582.CrossRefGoogle Scholar
  7. Andrews, P., and Alpagut, B. 1990. Description of the fossiliferous units at Paşalar, Turkey.J. Hum Evol. 19:343–361.CrossRefGoogle Scholar
  8. Andrews, P., and Cronin, J. E. 1982. The relationships of Sivapithecus and Ramapithecus and the evolution of the orang-utan. Nature 297:541–546PubMedCrossRefGoogle Scholar
  9. Andrews, P., and Ersoy, A. 1990. Taphonomy of the Miocene bone accumulations at Paşalar, Turkey. J Hum. Evol. 19:379–396.CrossRefGoogle Scholar
  10. Andrews, P. J., and Martin, L. 1987. The phyletic position of the Ad Dabtiyah hominoid. Bull. Br. Mus. (Nat. Hist.) (Geology) 41(4):383–393.Google Scholar
  11. Andrews, P., and Martin, L. 1991. Hominoid dietary evolution. Philos. Trans R. Soc. Lond 334:199–209.CrossRefGoogle Scholar
  12. Andrews, P., and Tobien, H. 1977. New Miocene locality in Turkey with evidence on the origin of Ramapithecus and SivapithecusNature 268:699–701PubMedCrossRefGoogle Scholar
  13. Andrews, P., Lord, J. M., and Nesbit Evans, E. M. 1979. Patterns of ecological diversity in fossil and modern mammalian faunas. Biol. J. Linnean Soc. 11:177–205.CrossRefGoogle Scholar
  14. Bestland, E. A. 1990. Sedimentology and paleopedology of Miocene alluvial deposits at the Paşalar Hominoid site, Western Turkey. J. Hum. Evol. 19:363–377.CrossRefGoogle Scholar
  15. Cope, D. A. 1989. Systematic Variation in Cercopithecus Dental Samples. Unpublished Ph.D. dissertation, The University of Texas at Austin.Google Scholar
  16. Cope, D. A., and Lacy, M. G. (1992). Falsification of a single species hypothesis using the coefficient of variation: A simulation approach. Am. J. Phys. Anthropol. 89:359–378.PubMedCrossRefGoogle Scholar
  17. Flynn, L. J., Pilbeam, D., Jacobs, L. L., Barry, J. C., Behrensmeyer, A. K., and Kappelman, J. W. 1990. The Siwaliks of Pakistan: Time and faunas in a Miocene terrestrial setting.J. Geol. 98:589–604.CrossRefGoogle Scholar
  18. Garn, S. M., Lewis, A. B., Swindler, D. R., and Kerewsky, R. S. 1967. Genetic control of sexual dimorphism in tooth size. J. Dent Res. 46(5, Suppl.):963–973.PubMedCrossRefGoogle Scholar
  19. Gingench, P. D. 1974. Size variability of the teeth in living mammals and the diagnosis of closely related sympatric fossil species. J. Paleontol. 48:895–903.Google Scholar
  20. Gingerich, P. D. 1985. Species in the fossil record: concepts, trends, and transitions. Paleobiology 11:27–41.Google Scholar
  21. Gingerich, P. D. 1987. Evolution and the fossil record: patterns, rates, and processes. Can. J. Zool. 65:1053–1060.CrossRefGoogle Scholar
  22. Gingerich, P. D., and Schoeninger, M. 1979. Patterns of tooth size variability in the dentitions of primates. Am. J. Phys. Anthrop. 51:457–466.PubMedCrossRefGoogle Scholar
  23. Greenfield, L. O. 1979. On the adaptive pattern of “Ramapithecus”. Am J. Phys. Anthropol. 50:527–548.PubMedCrossRefGoogle Scholar
  24. Groves, C. P., Westwood, C., and Shea, B. T. 1992. Unfinished business: Mahalanobis and a clockwork orang. J . Hum. Evol. 22:327–340.CrossRefGoogle Scholar
  25. Hooijer, D. A. 1948. Prehistoric teeth of man and of the orang-utan from central Sumatra, with notes on the fossil orang-utan from Java and southern China. Zoologische Mededeelingen (Leiden) 29:175–301.Google Scholar
  26. Johanson, D. C. 1974 Some metric aspects of the permanent and deciduous dentition of the pygmy chimpanzee (Pan paniscus). Am J. Phys Anthropol. 41:39–48.CrossRefGoogle Scholar
  27. Kappelman, J. 1986. The Paleoecology and Chronology of the Middle Miocene Hominoids from the Chinji Formation of Pakistan. Unpublished Ph.D. dissertation, Harvard University.Google Scholar
  28. Kappelman, J., Kelley, J, Pilbeam, D., Sheikh, K. A., Ward, S., Anwar, M., Barry, J. C., Brown, B., Hake, P., Johnson, N. M., Raza, S. M., and Shah, S. M. I. 1991. The earliest occurrence of Sivapithecus from the middle Miocene Chinji Formation of Pakistan. J. Hum. Evol. 21:61–73.CrossRefGoogle Scholar
  29. Kay, R. F. 1981. The nut-crackers—A new theory of the adaptations of the Ramapithecinae. Am. J. Phys. Anthropol. 55:141–151.CrossRefGoogle Scholar
  30. Kay, R. F. 1982a. Sivapithecus simonsi, a new species of Miocene Hominoid, with comments on the phylogenetic status of the Ramapithecinae. Int J. Pnmatol. 3:113–172.Google Scholar
  31. Kay, R. K. 1982b. Sexual dimorphism in Ramapithecinae. Proc. Natl. Acad. Sci. USA 79:209–212.PubMedCentralPubMedCrossRefGoogle Scholar
  32. Kay, R. F., and Simons, E. L. 1983. A reassessment of the relationship between later Miocene and subsequent Hominoidea, in: R. L. Ciochon and R. S. Corruccini (eds.), New Interpretations of Ape and Human Ancestry, pp. 577–624. Plenum Press, New York.CrossRefGoogle Scholar
  33. Kelley, J. 1986. Species recognition and sexual dimorphism in Proconsul and Rangwapithecus. J. Hum. Evol. 15:461–495.CrossRefGoogle Scholar
  34. Kelley, J., and Etler, D. 1989. Hominoid dental variability and species number at the Late Miocene site of Lufeng, China. Am. J Pnmatol. 18:15–34.CrossRefGoogle Scholar
  35. Kelley, J., and Gingerich, P. D. 1991. The effects of time-accumulation on metric variability in fossil samples (abstract). Soc. Vertebr. Paleontol. 11 (Suppl.): 39A.Google Scholar
  36. Kelley, J., and Pilbeam, D. 1986. The Dryopithecines: Taxonomy, comparative anatomy, and phylogeny of Miocene large hominoids, in: D. R. Swindler and J. Erwin (eds.), Comparative Primate Biology, Volume 1: Systematica, Evolution, and Anatomy, pp. 361–411. Alan R. Liss, New York.Google Scholar
  37. Kelley, J., and Xu, Q. 1991. Extreme sexual dimorphism in a Miocene hominoid. Nature 352:151–153.PubMedCrossRefGoogle Scholar
  38. Marshall, J., and Sugardjito, J. 1986. Gibbon systematics, in: D. R. Swindler and J. Erwin (eds), Comparative Primate Biology, Vol. 1: Systematics, Evolution, and Anatomy, pp. 137–185. Alan R. Liss, New York.Google Scholar
  39. Martin, L. 1981. New specimens of Proconsul from Koru, Kenya. J . Hum. Evol. 10:139–150.CrossRefGoogle Scholar
  40. Martin, L. B. 1983. The Relationships of the Later Miocene Hominoidea. Unpublished Ph.D. dissertation, University College London.Google Scholar
  41. Martin, L. 1985. Significance of enamel thickness in hominoid evolution. Nature 314:260–263.PubMedCrossRefGoogle Scholar
  42. Martin, L., and Andrews, P. 1984. The phyletic position of Graecopithecus freybergi Koenigswald. Courier Forschungsinstitut Senckenberg 69:25–40.Google Scholar
  43. Oxnard, C. E. 1987. Fossils, Teeth and Sex. New Perspectives on Human Evolution. University of Washington Press, Seattle.Google Scholar
  44. Pickford, M. 1986. Sexual dimorphism in Proconsul, in: M. Pickford and B. Chiarelli (eds.), Sexual Dimorphism in Living and Fossil Primates, pp. 133–170. II Sedicesimo, Firenze.Google Scholar
  45. Pilbeam, D. R. 1969. Tertiary Pongidae of East Africa: Evolutionary relationships and taxonomy. Bull. Peabody Mus. Nat. Hist. 31:1–185.Google Scholar
  46. Plavcan, J. M. 1990. Sexual Dimorphism in the Dentition of Extant Anthropoid Primates. Ph.D. Dissertation, Duke University.Google Scholar
  47. Remane, A. 1921. Zur Beurteilung der fossilen anthropoiden. Zbl. Min. Geol. Paleontol. 11:335–339.Google Scholar
  48. Rose, K. D., and Bown, T. M. 1986. Gradual evolution and species discrimination in the fossil record. Contnb. Geol., Univ. Wyom., Special Paper 3:119–130.Google Scholar
  49. Ruff, C. B., Walker, A., and Teaford, M. F. 1989. Body mass, sexual dimorphism and femoral proportions of Proconsul from Rusinga and Mfangano Islands, Kenya. J. Hum Evol. 18:515–536.CrossRefGoogle Scholar
  50. Sickenberg, O., Becker-Platen, J. D., Benda, L., Berg, D., Engesser, B., Gaziry, W, Heissig, K., Hünermann, K. A., Sondaar, P. Y., Schmidt-Kittler, N., Staesche, K., Staesche, U., Steffens, P., and Tobien, H. 1975. Die Gliederung des höheren Jungtertiars und Altquartärs in der Turkei nach Vertebraten und ihre Bedeutung für die internationale Neogen-Stratigraphie. Geologisches Jahrbuch B15:1–167.Google Scholar
  51. Simpson, G. G. 1941. Range as a zoological character. Am. J. Sci. 239:785–804CrossRefGoogle Scholar
  52. Simpson, G. G., Roe, A., and Lewontin, R. C. 1960. Quantitative Zoology Harcourt and Brace, New York.Google Scholar
  53. Tekkaya, I. 1974. A new species of anthropoid (Primates, Mammalia) from Anatolia. Bull Min. Res Explor. Inst. Turkey (Ankara) 83:148–165.Google Scholar
  54. Wood, B. A., and Xu, Q. 1991. Variation in the Lufeng dental remains. J . Hum Evol. 20:291–311.CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media New York 1993

Authors and Affiliations

  • Lawrence B. Martin
    • 1
  • Peter Andrews
    • 2
  1. 1.Departments of Anthropology and Anatomical SciencesState University of New York at Stony BrookStony BrookUSA
  2. 2.Department of PalaeontologyThe Natural History MuseumLondonEngland

Personalised recommendations