Asian Miocene Great Apes: Searching for Orangutan Ancestors


  • Yaowalak Chaimanee Laboratory PALEVOPRIM, UMR 7262 CNRS, University of Poitiers, 6 rue Michel Brunet, 86073 Poitiers Cedex 9, FRANCE
  • Jean-Jacques Jaeger Laboratory PALEVOPRIM, UMR 7262 CNRS, University of Poitiers, 6 rue Michel Brunet, 86073 Poitiers Cedex 9, FRANCE


evolution, hominoids, Khoratpithecus, orangutans, pongids, Southeast Asia


Pongids are represented today by the genus Pongo (orangutan), with three species that live on islands of Southeast Asia. However, orangutans were much more widely distributed during the Pleistocene. The changes in their geographical distribution and taxonomy are discussed here since the first appearance of this genus in the early Pleistocene (⁓2.2 million years ago) in South China. During the Pleistocene, orangutans coexisted with Gigantopithecus in South China, sharing their food and environment. Between the middle and late Miocene (12.5–5.4 million years ago), four distinct large apes were evolving in South and Southeast Asia. Each genus has been proposed as a potential ancestor of extant orangutans. However, Lufengpithecus, from South China is excluded from the pongids and is interpreted as having Eurasian dryopithecines ancestors. Sivapithecus, from India-Pakistan, exhibits orangutan skull characters but its dentition is more primitive, and its postcranial skeleton is specialized in another locomotory repertoire. Indopithecus appears to be the closest relative of Sivapithecus and is considered a potential ancestor of Gigantopithecus. Khoratpithecus, from Thailand and Myanmar, is recognized as the closest sister taxon of extant orangutans because both genera share characters such as the symphysis structure, teeth and jaw morphology, the absence of anterior digastric muscles of the mandible, and the organization of the nasoalveolar clivus. However, the absence of Pliocene large apes in Southeast Asia prevents us from documenting the evolutionary transition between Khoratpithecus and orangutans.


Almécija, S., Hammond, A.S., Thompson, N.E., Pugh, K.D., Moyà-Solà, S. and Alba, D.M. 2021. Fossil apes and human evolution. Science, 372: eabb4363.

Alpagut, B., Andrews, P., Fortelius, M., Kappelman, J., Temizsoy, I., Celebi, H. and Lindsay, W. 1996. A new specimen of Ankarapithecus meteai from the Sinap Formation of central Anatolia. Nature, 382: 349–351.

Azzaroli, A., Boccaletti, M., Delson, E., Moratti, G. and Torre, D. 1986. Chronological and Paleogeographical background to the study of Oreopithecus bambolii. Journal of Human Evolution, 15: 533–540.

Bacon, A.-M. and Long, V.T, 2001. The first discovery of a complete skeleton of a fossil orangutan in a cave of the Hoa Binh Province, Vietnam. Journal of Human Evolution, 41: 227–241.

Badgley, C., Qi, G.Q., Chen, W. and Han, D.1988. Paleoecology of a Miocene, tropical, upland fauna: Lufeng, China. National Geographic Research, 4: 178–195.

Begun, D.R. 2015. Fossil Record of Miocene hominoids. In: Henke, W. and Tattersall, I. (Eds). Handbook of Paleoanthro-pology, Springer-Verlag, Berlin Heidelberg, 1261–1332.

Begun, D.R. and Güleç, E. 1998. Restoration of the Type and Palate of Ankarapithecus meteai: Taxonomic and Phylogenetic Implications. American Journal of Physical Anthropology, 105: 279–314.

Begun, D.R. and Kivell, T.L. 2011. Knuckle-walking in Sivapi-thecus? The combined effects of homology and homoplasy with possible implications for pongine dispersals. Journal of Human Evolution, 60: 158–170.

Begun, D.R., Nargolwalla, M.C. and Kordos, L. 2012. European Miocene Hominids and the Origin of the African Ape and Human Clade. Evolutionary Anthropology, 21: 10–23.

Bocherens, H., Schrenk, F., Chaimanee, Y., Kullmer, O., Mörike, D., Pushkina, D. and Jaeger, J.-J. 2017. Flexibility of diet and habitat in Pleistocene South Asian mammals: Implications for the fate of the giant fossil ape Gigantopithecus. Quaternary International, 434: 148–155.

Brown, R. and Ward, S. 1988. Basicranial and facial topography in Pongo and Sivapithecus. In: Schwartz, J.H. (Ed.). Orangutan Biology, Oxford University Press, London, 247–260.

Brunet, M., Guy, F., Pilbeam, D., Mackaya, H.T., Likius, A., Ahounta, D., Beauvilain, A., Blondel, C., Bocherens, H., Boisserie, J.-R., de Bonis, L., Coppens, Y., Dejax, J., Denys, C., Duringer, P., Eisenmann, V., Fanone, G., Fronty, P., Geraads, D., Lehman, T., Lihoreau, F., Louchart, A., Mahamat, A., Merceron, G., Mouchelin, G., Otero, O., Campomanes, P.P., Ponce De Leon, M., Rage, J.-C., Sapanet, M., Schuster, M., Sudre, J., Tassy, P., Valentin, X., Vignaud, P., Viriot, L., Zazzo, A. and Zollikofer, C. 2002. A new hominid from the Upper Miocene of Chad, Central Africa. Nature, 418: 145–151.

Cameron, D.W. 2001. The taxonomic status of the Siwalik Late Miocene hominid. Gigantopithecus (‘Indopithecus’). Hima-layan Geology, 22: 29–34.

Chaimanee, Y., Jolly, D., Benammi, M., Tafforeau, P., Duzer, D., Moussa, I. and Jaeger, J.-J. 2003. A middle Miocene hominoid from Thailand and orangutan origins. Nature, 422: 61–65.

Chaimanee, Y., Lazzari, V., Chaivanich, K. and Jaeger, J.-J. 2019. First maxilla of a late Miocene hominid from Thailand and the evolution of pongine derived characters. Journal of Human Evolution, 134: 102636.

Chaimanee, Y., Lazzari, V., Yamee, C., Suraprasit, K., Rugbum-rung, M., Chaivanich, K. and Jaeger, J.-J. 2022. New materials of Khoratpithecus, a late Miocene hominoid from Nakhon Ratchasima Province, Northeastern Thailand, confirm its pongine affinities. Palaeontographica, Abteilung A, 323(4–6): 147–186.

Chaimanee, Y., Suteethorn, V., Jintasakul, P., Vidthayanon, C., Marandat, B. and Jaeger, J.-J. 2004. A new orangutan relative from the Late Miocene of Thailand. Nature, 427: 439–441.

Chaimanee, Y., Yamee, C., Tian, P., Khaowiset, K., Marandat, B., Tafforeau, P., Nemoz, C. and Jaeger, J.-J. 2006. Khoratpithecus piriyai, a Late Miocene hominoid of Thailand. American Journal of Physical Anthropology, 131(3): 311–323.

Chang, L., Guo, Z., Deng, C., Wu, H., Ji, X., Zhao, Y., Zhang, C., Ge, J., Wu, B., Sun, L. and Zhu, R. 2015. Pollen evidence of the palaeoenvironments of Lufengpithecus lufengensis in the Zhaotong Basin, southeastern margin of the Tibetan Plateau. Palaeogeography Palaeoclimatology Palaeoecology, 435: 95–104.

Cheng, Y.M., Wang, Y.F. and Li, C.S. 2014. Late Miocene wood flora associated with the Yuanmou hominoid fauna from Yunnan, southwestern China and its palaeoenvironmental implication. Journal of Palaeogeography, 3: 323–330.

Ciochon, R., Long, V.T., Larick, R., Gonzalez, L., Grun, R., de Vos, J., Yonge, C., Taylor, L., Yoshida, H. and Reagan, M. 1996. Dated co-occurrence of Homo erectus and Gigantopi-thecus from Tham Khuyen Cave, Vietnam. Proceedings of the National Academy of Science USA, 93: 3016–3020.

Ciochon, R.L., Piperno, D.R. and Thompson, R.G. 1990. Opal phytoliths found on the teeth of the extinct ape Gigantopithecus blacki: Implications for paleodietary studies. Proceedings of the National Academy of Science USA, 87: 8120–8124.

Coster, P., Benammi, M., Chaimanee, Y., Yamee, C., Chavasseau, O., Emonet E.-G. and Jaeger, J.-J. 2010. A complete magnetic-polarity stratigraphy of the Miocene continental deposits of Mae Moh Basin, northern Thailand, and a reassessment of the age of hominoid-bearing localities in northern Thailand. The Geological Society of America Bulletin, 122(7–8): 1180–1191.

Dennell, R. and Roebroeks, W. 2005. An Asian perspective on early human dispersal from Africa. Nature, 438: 1099–1104.

DeSilva, J.M., Morgan, M.E., Barry, J.C. and Pilbeam, D. 2010. A hominoid distal tibia from the Miocene of Pakistan. Journal of Human Evolution, 58: 147–154.

Fleagle, J.G. 2013. Primate Adaptation and Evolution, 3rd edition. Academic Press, San Diego, CA, 423 pp.

Georgiou, L., Kivell, T.L., Pahr, D.H., Buck, L.T. and Skinner, M.M. 2019. Trabecular architecture of the great ape and human femoral head. Journal of Anatomy, 234: 679–693.

Gilbert, C.C., Pugh, K.D., and Fleagle, J.G. 2020. Dispersal of Miocene hominoids (and pliopithecoids) from Africa to Eurasia in light of changing tectonics and climate. In: Prasad, G.V. and Patnaik, R. (Eds.), Biological Consequences of Plate Tectonics. New Perspectives on Post-Gondwana Break-Up-Tribute to Ashok Sahni. Springer, Cham, pp. 393e412.

Habinger, S.G., Chavasseau, O., Jaeger, J.-J., Chaimanee, Y., Aung Naing Soe, Chit Sein, and Bocherens, H. 2022. Evolutionary ecology of Miocene hominoid primates in Southeast Asia. Scientific Reports, 12: 11841.

Hamon, N., Emonet, E.-G., Chaimanee, Y., Guy, F., Tafforeau, P. and Jaeger, J.-J. 2012a. Analysis of dental root apical morpho-logy: A new method for dietary reconstructions in primates. The Anatomical Record, 295(6): 1017–1026.

Hamon, N., Sepulchre, P., Donnadieu, Y., Henrot, A.-J., François, L., Jaeger, J.-J. and Ramstein, G. 2012b. Growth of subtropical forests in Miocene Europe: The roles of carbon dioxide and Antarctic ice volume. Geological Society of America, 40(6): 567–570.

Harrison, T., Ji, X. and Su, D. 2002. On the systematic status of the late Neogene hominoids from Yunan Province, China. Journal of Human Evolution, 43: 207–227.

Harrison, T., Jin, C., Zhang, Y.Y., Wang, Y. and Zhu, M. 2014. Fossil Pongo from the Early Pleistocene Gigantopithecus fauna of Chongzuo, Guangxi, southern China. Quaternary Interna-tional, 354: 59–67.

Harrison, T., Krigbaum, J.S. and Manser, J. 2006. Primate biogeography and ecology on the Sundra Shelf islands; A paleontological and zooarchaeological perspective. In: Fleagle, J. G. and Lehman, S. (Eds.). Primate Biogeography; Progress and Prospects, McGraw-Hill, New York, 331–372.

Harrison, T., Zhang, Y., Yang, L. and Yuan, Z. 2021. Evolutionary trend in dental size in fossil orangutans from the Pleistocene of Chongzuo, Guangxi, southern China. Journal of Human Evolution 161: 103019.

Huang, Y.-J., Ji, X.-P., Su, T., Deng, C.-L., Ferguson, D.K., Yu, T.-S., Yang, X., Sun, H. and Zhou, Z.K. 2017. Habitat, climate and potential plant food resources for the late Miocene Shuitangba hominoid in Southwest China: Insights from carpological remains. Palaeogeography, Palaeoclimatology, Palaeoecology, 470: 63–71.

Hunt, K.D. 2016. Why are there apes? Evidence for the co-evo-lution of ape and monkey ecomorphology. Journal of Anatomy, 228: 630–685.

Jaeger, J.-J., Aung Naing Soe, Chavasseau, O., Coster, P., Emonet, E.-G., Guy, F., Lebrun, R. Aye Maung, Aung Aung Khyaw, Hla Shwe, Soe Thura Tun, Kyaw Linn Oo, Rugbumrung, M., Bocherens, H., Benammi, M., Chaivanich, K., Tafforeau, P. and Chaimanee, Y. 2011. First Hominoid from the Late Miocene of the Irrawaddy Formation (Myanmar). PLoS ONE, 6(4): e17065.

Ji, X., Jablonski, N.G., Su, D. F., Deng, C., Flynn, L.J., You, Y. and Kelley, J. 2013. Juvenile hominoid cranium from the terminal Miocene of Yunnan, China. Chinese Science Bulletin 58 (31): 3771–3779.

Kappelman, J., Kelley, J., Pilbeam, D., Sheik, 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. Journal of Human Evolution, 21: 61–73.

Kappelman, J., Richmond, B.G., Seiffert, E.R., Maga, A.M. and Ryan, T.M. 2003. Hominoidea (Primates). In: Fortelius, M., Kappelman, J., Sen, S. and Bernor, L. (Eds.). Geology and Paleontology of the Miocene Sinap Formation, Turkey, Columbia University Press, New York, 90–124.

Kelley, J. 1988. A new large species of Sivapithecus from the Siwaliks of Pakistan. Journal of Human Evolution, 17: 305–324.

Kelley, J. 2005. Twenty-five years contemplating Sivapithecus taxonomy. In: Lieberman, D.E., Smith, R.J. and Kelley, J. (Eds.). Interpreting the past: essays on human, primate and mammal evolution in honor of David Pilbeam, Brill Academic Publishers, Boston, MA., 123–143.

Kelley, J. and Gao, F. 2012. Juvenile hominoid cranium from the late Miocene of southern China and hominoid diversity in Asia. Proceedings of the National Academy of Science USA, 109: 6882–6885.

Kelley, J. and Xu Q. 1991. Extreme sexual dimorphism in a Miocene hominoid. Nature, 352: 151–153.

Li, P., Zhang, C., Guo, Z., Deng, C., Ji, X., Jablonski, N.G., Wu, H. and Zhu, R. 2019. Clay mineral assemblages in the Zhaotong Basin of southwestern China: Implications for the late Miocene and Pliocene evolution of the South Asian monsoon. Palaeo-geography, Palaeoclimatology, Palaeoecology, 516: 90–100.

Li, P., Zhang, C., Kelley, J., Deng, C., Ji, X., Jablonski, N.G., Wu, H., Fu, Y., Guo, Z. and Zhu, R. 2020. Late Miocene climate cooling contributed to the disappearance of Hominoids in Yunnan Region, Southwestern China. Geophysical Research Letters, 47: e2020GL087741.

Liao, W., Harrison, T., Yao, Y., Liang, H., Tian, C., Feng, Y., Li, S., Bae, C.J. and Wang, W. 2022. Evidence for the latest fossil Pongo in southern China. Journal of Human Evolution, 170: 103233.

Liu, W., Gao, F. and Zheng, L. 2002. The diet of the Yuanmou hominoid, Yunnan Province, China: an analysis from tooth size and morphology. Anthropological Science, 110 (2): 149–163.

Louys, J., Braje, T.J., Chang, C.-H., Cosgrove, R., Fitzpatrick, S.M., Fujitah, M., Hawkins, S., Ingicco, T., Kawamura, A., MacPhee, R.D.E., McDowell, M.C.H., Meijer, J.M.P., Piper, J., Roberts, P., Simmons, A.H., van den Bergh, G., van der Geer, A., Kealy, S. and O'Connor, S. 2021. No evidence for widespread island extinctions after Pleistocene hominin arrival. Proceedings of the National Academy of Sciences USA, 118: e2023005118.

Lu, X., Ji, X., Hou, S., Wang, S., Shi, Q., Chen, S., Sun, B., Li, Y., Li, Y., Yu, T. and Li, W. 2017. Palaeoenvironment examination of the terminal Miocene hominoid locality of the Zhaotong Basin, southwestern China, based on the rhinocerotid remains. Historical Biology, 31(2): 234–242.

MacDonald, D. and Norris, S. 2001. The New Encyclopedia of Mammals. Oxford University Press, 961 pp.

MacLatchy, L.M. and Bossert, W.H. 1996. An analysis of the articular surface distribution of the femoral head and acetabulum in anthropoids, with implications for hip function in Miocene hominoids. Journal of Human Evolution, 31: 425–453.

Madar, S.I., Rose, M.D., Kelley, J., MacLatchy, L. and Pilbeam, D. 2002. New Sivapithecus postcranial specimens from the Siwaliks of Pakistan. Journal of Human Evolution, 42: 705–752.

Merceron, G., Taylor, S., Scott, R., Chaimanee, Y. and Jaeger, J.-J. 2006. Dietary characterization of the hominoid Khoratpithecus (Miocene of Thailand): evidence from dental topographic and microwear texture analyses. Naturwissenschaften, 93: 329–333.

Morley, R.J. 2018. Assembly and division of the South and South-East Asian flora in relation to tectonics and climate change. Journal of Tropical Ecology, 34: 209–234.

Nater, A., Mattle-Greminger, M.P., Nucahyo, A., Nowak, M.G., Meijaard, E. and Krutzen, M. 2017. Morphometric, Behavioral, and Genomic Evidence for a New Orangutan Species. Current Biology, 27: 3487–3498.

Nelson, S.V. 2003. The extinction of Sivapithecus: faunal and environmental changes surrounding the disappearance of a Miocene hominoid in the Siwaliks of Pakistan. Brill Academic Publishers, Boston, 106 pp.

Nelson, S.V. 2014. The Paleoecology of Early Pleistocene Giganto-pithecus blacki inferred from isotopic analyses. American Journal of Physical Anthropology, 155: 571–578.

Nengo, I., Tafforeau, P., Gilbert, C.C., Fleagle, J.G., Miller, E.R., Feibel, C., Fox, D.L., Feinberg, J., Pugh, K.D., Berruyer, C., Mana, S., Engle, Z. and Spoor, F. 2017. New infant cranium from the African Miocene sheds light on ape evolution. Nature, 548: 169e174.

Noerwidi, S., Siswanto and Widianto, H. 2016. Giant primate of Java: a new Gigantopithecus specimen from Semedo. Berkala Arkeologi, 36: 141–160.

Olejniczak, A.J., Smith, T.M., Wang, W., Potts, R., Ciochon, R., Kullmer, O., Schrenk, F. and Hublin, J.-J. 2008. Molar enamel thickness and dentine horn height in Gigantopithecus blacki. American Journal of Physical Anthropology, 135: 85–91.

Patnaik, R., Cerling, T.E., Uno, K.T. and Fleagle, J.G. 2014. Diet and habitat of Siwalik primates Indopithecus, Sivaladapis and Theropithecus. Annales Zoologici Fennici 51: 123–142.

Pilbeam, D. 1982. New hominoid skull material from the Miocene of Pakistan. Nature, 295: 232–234.

Pilbeam, D., Meyer, G.E., Badgley, C., Rose, M.D., Pickford, M.H.L., Behrensmeyer, A.K. and Shah, S.M.I. 1977. New hominoid primates from the Siwaliks of Pakistan and their bearing on hominoid evolution. Nature, 270: 689–695.

Pilbeam, D., Rose, M.D., Badgley, C. and Lipschutz, B. 1980. Miocene hominoids from Pakistan. Postilla, 181: 1–94.

Pilbeam, D., Rose, M.D., Barry, J.C. and Shah, S.M.I. 1990. New Sivapithecus humeri from Pakistan and the relationship of Sivapithecus and Pongo. Nature, 348: 237–239.

Pilgrim, G.E. 1915. New Siwaliks primates and their bearing on the question of the evolution of man and the Anthropoidea. Records Geological Survey of India, 45: 1–74.

Pillans, B., Williams, M., Cameron, D., Patnaik, R., Hogarth, J., Sahni, A., Sharma, J.C., Williams, F. and Bernor, R.L. 2005. Revised correlation of the Haritalyangar magnetostratigraphy, Indian Siwaliks: implications for the age of the Miocene hominids Indopithecus and Sivapithecus, with a note on a new hominid tooth. Journal of Human Evolution, 48: 507–515.

Plavcan, J.M. 2001. Sexual dimorphism in Primate evolution. Yearbook of Physical Anthropology, 44: 25–53.

Pugh, K.D. 2022. Phylogenetic analysis of Middle-Late Miocene apes. Journal of Human Evolution, 165: 103140.

Qi, G. 1993. The environment and ecology of the Lufeng hominoids. Journal of Human Evolution, 24: 3–11.

Qi, G., Dong, W., Zheng, L., Zhao, L., Gao, F., Yue, L. and Zhang, Y. 2006. Taxonomy, age and enviroment status of the Yuanmou hominoids. Chinese Science Bulletin, 51: 704–712.

Qiu, Z.-D. and Li., C.-K. 2003. Rodents from the Chinese Neogene: Biogeographic relationships with Europe and North America. In: Flynn, L. (Ed.), Bulletin of the American Museum of Natural History, Vertebrate Fossils and their Context: Contri-butions in Honor of Richard Tedford, New York, 586–602.

Schwartz, J.H. 1997. Lufengpithecus and hominoid phylogeny: Problems in delineating and evaluating phylogenetically relevant characters. In: Begun, D.R., Ward, C.V. and Rose, M.D. (Eds.). Advances in Primatology; Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations, Plenum Press, New York, 363–388.

Senut, B., Pickford, M., Gommery, D., Mein, P., Cheboi, K. and Coppens, Y. 2001. First hominid from the Miocene (Lukeino Formation, Kenya). Comptes Rendus de l'Académie des Sciences, Série II Sciences de la Terre et des Planètes, 323: 137–144.

Sépulchre, P., Jolly, D., Ducrocq, S., Chaimanee, Y., Jaeger, J.-J. and Raillard, A. 2010. Mid-Tertiary paleoenvironments in Thailand: pollen evidences. Climates of the Past 6(4): 461–473.

Simons, E.L. and Chopra, S.R.K.1969. Gigantopithecus (Pongidae, Hominoidea) a new species from North India. Postilla, 138: 1–18.

Steiper, M.E. and Young, N.M. 2006. Primate molecular diver-gence dates. Molecular Phylogenetics and Evolution, 41: 384–394.

Sun, X., and Wu, Y. 1980. Paleoenvironment during the time of Ramapithecus lufengensis. Vertebrata PalAsiatica, 18: 247–255.

Swindler, D.R. and Olshan, A.F. 1988. Comparative and evolu-tionary aspects of the permanent dentition. In: Schwartz, J. H. (Ed.). Orangutan Biology, Oxford University Press, 271–282.

Takai, M., Nyo, K., Kono, R.T., Htike, T., Kusuhashi, N. and Thein, Z.M.M. 2021. New hominoid mandible from the early Late Miocene Irrawaddy Formation in Tebingan area, central Myanmar. Anthropological Science, 129(1): 87–98.

Urciuoli, A. and Alba, D.M. 2023. Systematics of Miocene apes: State of the art of a neverending controversy. Journal of Human Evolution, 17: 103309.

von Koenigswald, G.H.R. 1935. Eine fossile Saugetierfauna mit Simia aus Sudchine. Proceeding of the Koninklijke Nederlandse Akademie van Wetenschappen, 38: 872–879.

Wall, C.E., Larson, S.G. and Stern, J.J.T. 1994. EMG of the digastric muscle in gibbon and orangutan: Functional consequences of the loss of the anterior digastric in orangutans. American Journal of Physical Anthropology, 94: 549–567.

Ward, S.C. and Kimbel, W.H. 1983. Subnasal alveolar morphology and the systematic position of Sivapithecus. American Journal of Physical Anthropology, 61: 157–171.

Welker, F., Ramos-Madrigal, J., Kuhlwilm, M., Liao, W., Gutenbrunner, P., de Manuel, M., Samodova, D., Mackie, M., Allentoft, M.E., Bacon, A.-M., Collins, M.J., Cox, J., Lalueza-Fox, C., Olsen, J.V., Demeter, F., Wang, W., Marques-Bonet, T. and Cappellini, E. 2019. Enamel proteome shows that Gigantopithecus was an early diverging pongine. Nature, 576: 262–265.

White, T.D., Asfaw, B., Beyene, Y., Haile-Selassie, Y., Lovejoy, C.O., Suwa, G. and WoldeGabriel, G. 2009. Ardipithecus rami-dus and the paleobiology of early hominids. Science, 326: 64: 75–86.

Woo, J. 1957. Dryopithecus teeth from Keiyuan, Yunnan Province. Vertrebrata PalAsiatica, 1: 25–32.

Woo, J.K. 1962. The mandibles and dentition of Gigantopithecus. Paleontologica Sinica, New Series D, 11: 1–94.

Wu, R. 1987. A revision of the classification of the Lufeng great apes. Acta Anthropologica Sinica, 6: 265–271.

Xu, Q., Lu, Q., Pan, Y., Zheng, X. and Zheng, L. 1978. Fossil mandible of the Lufeng Ramapithecus. Kexue Tongbao, 9: 554–556.

Xu, Q.H., and Lu, Q.W. 2008. Lufengpithecus lufengensis-an early member of Hominidae. Science Press, Beijing, China, 234 pp.

Yue, L.P. and Zhang, Y.X. 2006. Paleomagnetic dating of Lufeng-pithecus hudienensis localities (in Chinese). In: Qi G.Q. and Dong W. (Eds.). Lufengpithecus hudienensis Site, Science Press, Beijing, 245–255.

Zhang, C., Guo, Z., Deng, C., Ji, X., Wu, H., Paterson, G. A., Chang, L., Li, Q., Wu, B. and Zhu, R. 2016a. Clay mineralogy indicates a mildly warm and humid living environment for the Miocene hominoid from the Zhaotong Basin, Yunnan, China. Scientific Report, 6: 20012.

Zhang, W., Yan, M., Fang, X., Zhang, D., Zhang, T., Zan, J. and Song, C. 2019. High-resolution paleomagnetic constraint on the oldest hominoid-fossil-bearing sequence in the Xiaolongtan Basin, southeast margin of the Tibetan Plateau and its geologic implications. Global and Planetary Change, 182: 103001.

Zhang, X., Lin, Y., Jiang, C. and Xiao, L. 1987. A new species of Ramapithecus from Yuanmou, Yunnan. Journal of Yunnan University (Social Sciences), 3: 54–56.

Zhang, Y. and Harrison, T. 2017. Gigantopithecus blacki: a giant ape from the Pleistocene of Asia revisited. American Journal of Physical Anthropology, 162: 153–177.

Zhang, Y., Harrison, T. and Ji, X. 2020. Inferring the locomotor behavior of fossil hominoids from phalangeal curvature using a novel method: Lufengpithecus as a case study. Acta Anthropo-logica Sinica, 39: 532–554.

Zhang, Y., Jin, C., Kono, R.T., Harrison, T. and Wang, W. 2016b. A fourth mandible and associated dental remains of Gigantopithecus blacki from the Early Pleistocene Yanliang Cave, Fusui, Guangxi, South China. Historical Biology, 28: 95–104.




How to Cite

Chaimanee, Y. and Jaeger, J.-J. 2023. Asian Miocene Great Apes: Searching for Orangutan Ancestors. Tropical Natural History. 7 (Sep. 2023), 260–275.