Journal of Mammalian Evolution

, Volume 20, Issue 2, pp 115–127 | Cite as

A new species of Kolpochoerus (Mammalia: Suidae) from the Pliocene of Central Afar, Ethiopia: Its Taxonomy and Phylogenetic Relationships

  • Yohannes Haile-SelassieEmail author
  • Scott W. Simpson
Original Paper


Kolpochoerus (Mammalia: Suidae) is a suine genus represented by a number of species from Plio-Pleistocene sites in Africa. While the general trends in Kolpochoerus evolution are broadly known, gaps in the fossil record preclude an understanding of the details of its evolutionary tempo and mode. Here, we describe a new species, Kolpochoerus millensis, based on new fossil material from the Woranso-Mille and Gona sites in the Central Afar region of Ethiopia and dated to 3.5–3.8 million years ago (Ma). Third molars of K. millensis are metrically and morphologically intermediate between the early Pliocene K. deheinzelini and earliest late Pliocene K. afarensis. It appears that K. deheinzelini, K. millensis, and K. afarensis are temporally disjunct and phenetically distinguishable parts of a single evolving lineage. The recognition of these chronospecies provides additional evidence for anagenetic evolution. It demonstrates clearly the presence of transitional forms in the fossil record. The extensive and well-dated Kolpochoerus fossil record serves as one of the best documented examples of the occurrence of phyletic evolution. Moreover, K. millensis is one of the best biochronological markers in eastern Africa for the time between 3.5 and 3.8 Ma.


Suinae Kolpochoerus Taxonomy Phylogeny Phyletic evolution Woranso-Mille Gona Pliocene Ethiopia 



We thank the Authority for Research and Conservation of Cultural Heritage (ARCCH) of the Ministry of Culture and Tourism and the government of the Afar Regional State of Ethiopia for facilitating fieldwork permits, the Paleoanthropology Laboratory of the National Museum of Ethiopia for laboratory space and access to the original fossil specimens, and all field participants of the Woranso-Mille Project and Gona Project (S. Semaw, project leader). The National Science Foundation (Grant Nos. BCS-0234320, BCS-0542037), The Leakey Foundation, The National Geographic Society, and The Wenner-Gren Foundation financially supported the field and laboratory work. We also thank Liz Russell for photography. This manuscript benefited from reviews and constructive comments from H.B.S. Cooke, J.M. Harris, and D. Geraads.


  1. Arribas A, Garrido G (2008) Un nuevo jabalí del género Potamochoerus (Suidae, Artiodactyla, Mammalia) en el Plioceno superior terminal euroasiático (Fonelas p-1, cuenca de Guadix, Granada). In: Arribas A (ed.) Vertebrados del Plioceno superior terminal en el suroeste de Europa: Fonelas P-1 y el Proyecto Fonelas. Cuadernos del Museo Geominero, nº 10. Instituto Geológico y Minero de España, Madrid, pp 337–364Google Scholar
  2. Bishop LC (1999) Suid paleoecology and habitat preferences at African Pliocene and Pleistocene hominid localities. In: Bromage TG, Schrenk F (eds) African Biogeography: Climate Change and Human Evolution. Oxford University Press, Oxford, pp 216–225Google Scholar
  3. Bishop LC (2011) Suidae. In: Harrison T (ed) Paleontology and Geology of Laetoli: Human Evolution in Context. Volume 2: Fossil Hominins and the Associated Fauna. Vertebrate Paleobiology and Paleoanthropology, Springer, Dordrecht, pp 327–337Google Scholar
  4. Bishop LC, King T, Hill A, Wood B (2007) Paleoecology of Kolpochoerus heseloni (=K. limnetes): a multiproxy approach. Trans R Soc Afr 61:81–88CrossRefGoogle Scholar
  5. Brunet M, White TD (2001) Deux nouvelles espèces de Suini (Mammalia, Suidae) du continent Africain (Ethiopie; Tchad). C R Acad Sci Paris 332:51–57CrossRefGoogle Scholar
  6. Cooke HBS (1976) Suidae from the Pliocene-Pleistocene strata of the Rudolf basin. In: Coppens Y, Howell FC, Isaac GL, Leakey RE (eds) Earliest Man and Environments in the Lake Rudolf Basin. University of Chicago Press, Chicago, pp 177–192Google Scholar
  7. Cooke HBS (1978a) Suid evolution and correlation of African hominid localities: an alternative taxonomy. Science 201:460–463CrossRefPubMedGoogle Scholar
  8. Cooke HBS (1978b) Pliocene-Pleistocene Suidae from Hadar, Ethiopia. Kirtlandia 29:1–63Google Scholar
  9. Cooke HBS (1997) The status of the African fossil suids Kolpochoerus limnetes (Hopwood, 1926), K. phacochoeroides (Thomas, 1884) and ‘K.’ afarensis (Cooke, 1978). Geobios 30:121–126CrossRefGoogle Scholar
  10. Cooke HBS (2007) Stratigraphic variation in Suidae from the Shungura Formation and some coeval deposits. In: Bobe R, Alemseged Z, Behrensmeyer AK (eds) Hominin Environments in the East African Pliocene: An Assessment of the Faunal Evidence. Springer, New York, pp 107–127CrossRefGoogle Scholar
  11. Deino A (2011) 40Ar/39Ar dating of Laetoli, Tanzania. In: Harrison T (ed) Paleontology and Geology of Laetoli: Human Evolution in Context. Volume 1: Geology, Geochronology, Paleoecology and Paleoenvironment. Vertebrate Paleobiology and Paleoanthropology, Springer, Dordrecht, pp 77–97CrossRefGoogle Scholar
  12. Deino A, Scott G, Saylor B, Alene M, Angelini J, Haile-Selassie Y (2010) 40Ar/39Ar dating, paleomagnetism, and tephrochemistry of Pliocene strata of the hominid-bearing Woranso-Mille area, west-central Afar Rift, Ethiopia. J Hum Evol 58:111–126CrossRefPubMedGoogle Scholar
  13. Eldredge N, Cracraft J (1980) Phylogenetic Patterns and the Evolutionary Process. Method and Theory in Comparative Biology. Columbia University Press, New YorkGoogle Scholar
  14. Eldredge N, Gould SJ (1972) Punctuated equilibria: an alternative to phyletic gradualism. In: Schopf TM (ed) Models in Palaeobiology. Freeman, Cooper and Co., San Francisco, pp 82–115Google Scholar
  15. Fessaha N (1999) Systematics of Hadar (Afar, Ethiopia) Suidae. PhD thesis. Howard University, Washington D.C.Google Scholar
  16. Garrido G (2006) Paleontología sistemática de grandes mamíferos del yacimiento del Villafranquiense superior de Fonelas P-1 (Cuenca de Guadix, Granada). PhD thesis. Universidad Complutense de Madrid, Madrid, SpainGoogle Scholar
  17. Geraads D (1993) Kolpochoerus phacochoeroides (Thomas, 1884) (Suidae, Mammalia), du Pliocène supérieur de Ahl al Oughlam (Casablanca, Maroc). Geobios 26:731–743CrossRefGoogle Scholar
  18. Geraads D (2004) New skulls of Kolpochoerus phacochoeroides (Suidae: Mammalia) from the late Pliocene of Ahl al Oughlam, Morocco. Palaeontol Afr 40:69–83Google Scholar
  19. Gilbert WH (2008) Suidae. In: Gilbert WH, Asfaw B (eds) Homo erectus: Pleistocene Evidence from the Middle Awash, Ethiopia. University of California Press, Berkeley, pp 231–260Google Scholar
  20. Gingerich PD (1976) Paleontology and phylogeny: patterns of evolution of the species level in early Tertiary mammals. Am J Sci 276:1–28CrossRefGoogle Scholar
  21. Gingerich PD (1980) Evolutionary patterns in early Cenozoic mammals. Annu Rev Earth Planet Sci 8:407–424CrossRefGoogle Scholar
  22. Gould SJ (2002) The Structure of Evolutionary Theory. President and Fellows of Harvard College, Harvard University Press, HarvardGoogle Scholar
  23. Haile-Selassie Y (2009) Chapter 10: Suidae. In: Haile-Selassie Y, WoldeGabriel G (eds) Ardipithecus kadabba: Evidence from the Middle Awash, Ethiopia. University of California Press, Berkeley, pp 331–371Google Scholar
  24. Haile-Selassie Y, Asfaw B (2000) A newly discovered early Pliocene hominid bearing paleontological site in the Mulu Basin, Ethiopia. Am J Phys Anthropol 30 (Supplement):170Google Scholar
  25. Haile-Selassie Y, Deino A, Saylor B, Umer M, Latimer B (2007) Preliminary geology and paleontology of new hominid-bearing Pliocene localities in the central Afar region of Ethiopia. Anthropol Sci 115:215–222CrossRefGoogle Scholar
  26. Harris JM (1987) Fossil Suidae from Laetoli. In: Leakey MD, Harris JM (eds) Laetoli: A Pliocene Site in Northern Tanzania. Clarendon Press, Oxford, pp 349–357Google Scholar
  27. Harris JM, Cerling T (2002) Dietary adaptations of extant and Neogene African suids. J Zool London 256:45–54CrossRefGoogle Scholar
  28. Harris JM, Leakey MG, Cerling, TE (2003) Early tetrapod remains from Kanapoi, Lake Turkana Basin, Kenya. Contr Sci Los Angeles 498:39–113Google Scholar
  29. Harris JM, White TD (1979) Evolution of the Plio-Pleistocene African Suidae. Trans Am Phil Soc 69:1–128CrossRefGoogle Scholar
  30. Hendey QB, Cooke HBS (1985) Kolpochoerus paiceae (Mammalia, Suidae) from Skurwerug, near Saldanha, South Africa, and its palaeoenvironmental implications. Ann So Afr Mus 97:9–56Google Scholar
  31. Kalb JE (1993) Refined stratigraphy of the hominid-bearing Awash Group, Middle Awash Valley, Ethiopia. Newsl Stratig 29:21–62Google Scholar
  32. Kimbel WH, Lockwood CA, Ward CV, Leakey MG, Rak Y, Johanson DC (2006) Was Australopithecus anamensis ancestral to A. afarensis? A case of anagenesis in the hominin fossil record. J Hum Evol 51:134–152CrossRefPubMedGoogle Scholar
  33. Kimbel WH, Rak Y (1993) The importance of species taxa in paleoanthropology and an argument for the phylogenetic concept of the species category. In: Kimbel WH, Martin LB (eds) Species, Species Concepts, and Primate Evolution. Plenum, New York, pp 461–484Google Scholar
  34. Kullmer O, Sandrock O, Viola TB, Hujer W, Said H, Seidler H (2008) Suids, elephantoids, paleochronology, and paleoecology of the Pliocene hominid site Galili, Somali Region, Ethiopia. Palaios 23:452–464CrossRefGoogle Scholar
  35. Leakey MG, Feibel CS, McDougall I, Walker A (1995) New four-million-year-old hominid species from Kanapoi and Allia Bay, Kenya. Nature 376:565–571CrossRefPubMedGoogle Scholar
  36. Orliac MJ, Antoine P-O, Roohi G, Welcomme J-L (2010) Suoidea (Mammalia, Cetartiodactyla) from the early Oligocene of Bugti Hills, Balochistan, Pakistan. J Vertebr Paleontol 30:1300–1305CrossRefGoogle Scholar
  37. Pickford M (1990) Révision des suidés de la Formation de Beglia (Tunisie). Ann Paléontol 76:133–141Google Scholar
  38. Pickford M (1994) Fossil Suidae of the Albertine rift, Uganda–Zaire. CIFEG Occasional Publ 29:339–373Google Scholar
  39. Pickford M (2006) Synopsis of the biochronology of African Neogene and Quaternary Suiformes. Trans R Soc So Afr 61:51–62CrossRefGoogle Scholar
  40. Quade J, Levin NE, Simpson SW, Butler R, McIntosh WC, Semaw S, Kleinsasser L, Dupont-Nivet G, Renne P, Dunbar N (2008) The geology of Gona, Afar, Ethiopia. In: Quade J, Wynn J (eds) The Geology of Early Humans in the Horn of Africa. Geol Soc Am, Spec Pap 446:1–32Google Scholar
  41. Semaw S, Simpson SW, Quade J, Renne PR, Butler RF, McIntosh WC, Levin N, Dominiguez-Rodrigo M, Rogers MJ (2005) Early Pliocene hominids from Gona, Ethiopia. Nature 433:301–305CrossRefPubMedGoogle Scholar
  42. Suwa G, Nakaya H, Asfaw B, Saegusa H, Amzaye A, Kono RT, Beyene Y, Katoh S (2003) Plio-Pleistocene terrestrial mammal assemblage from Konso, southern Ethiopia. J Vertebr Paleontol 23:901–916CrossRefGoogle Scholar
  43. Urbanek C, Faupl P, Hujer W, Ntaflos T, Richter W, Weber G, Schaefer K, Viola B, Gunz P, Neubauer S, Stadlmayr A, Kullmer O, Sandrock O, Nagel D, Conroy G, Falk D, Woldearegay K, Said H, Assefa G, Seidler H (2005) Geology, paleontology and paleoanthropology of the Mount Galili Formation in the southern Afar Depression, Ethiopia - Preliminary results. Joannea Geol Paläont 6:29–43Google Scholar
  44. Van der Made J (1996) Listriodontinae (Suidae, Mammalia), their evolution, systematics and distribution in time and space. Contrib Tert Quater Geol 33:3–254Google Scholar
  45. Van Hoepen ECN, Van Hoepen HE (1932) “Vrystaate wilde varke.” Paleont. Navors Nas Mus Bloemfontein 2:39–62Google Scholar
  46. White TD (1995) African omnivores: global climatic change and Plio-Pleistocene hominids and suids. In: Vrba ES, Denton GH, Partridge TC, Burckle LH (eds) Paleoclimate and Evolution with Emphasis on Human Origins. Yale University Press, New Haven, pp 369–384Google Scholar
  47. White TD, Harris JM (1977) Suid evolution and correlation of African hominid localities. Science 198:13–21CrossRefPubMedGoogle Scholar
  48. White TD, Suwa G (2004) A new species of Notochoerus (Artiodactyla, Suidae) from the Pliocene of Ethiopia. J Vertebr Paleontol 24:474–480CrossRefGoogle Scholar
  49. White TD, WoldeGabriel G, Asfaw B, Ambrose S, Beyene Y, Bernor RL, Boisserie J-R, Currie B, Gilbert WH, Haile-Selassie Y, Hart WK, Hlusko LJ, Howell FC, Kono RT, Lehmann T, Louchart A, Lovejoy OC, Renne PR, Saegusa H, Vrba ES, Wesselman H, Suwa G (2006) Asa Issie, Aramis and the origin of Australopithecus. Nature 440:883–889CrossRefPubMedGoogle Scholar

Copyright information

© Springer Science+Business Media, LLC 2012

Authors and Affiliations

  1. 1.Department of Physical AnthropologyThe Cleveland Museum of Natural HistoryClevelandUSA
  2. 2.Departments of Anthropology, Cognitive Sciences, Anatomy, and Institute for the Science of OriginsCase Western Reserve UniversityClevelandUSA
  3. 3.Department of AnatomyCase Western Reserve University School of MedicineClevelandUSA

Personalised recommendations