Tortoises (Chelonii, Testudinidae)

  • Terry HarrisonEmail author
Part of the Vertebrate Paleobiology and Paleoanthropology Series book series (VERT)


Two species of tortoises are known from Pliocene sites on the Eyasi Plateau, including Laetoli. The most common species is a medium-sized tortoise, Stigmochelys brachygularis, which is well represented in the Laetolil Beds (∼3.6–4.4 Ma) and Upper Ndolanya Beds (∼2.66 Ma). The giant tortoise, “Geochelone” laetoliensis, is known only from the Laetolil Beds, and is much less common than S. brachygularis. Stigmochelys brachygularis is represented by a number of relatively complete and partial shells, as well as numerous isolated and associated shell fragments, some postcranial remains and eggs. It is generally similar in size and overall morphology to the extant leopard tortoise, S. pardalis, but a number of features serve to distinguish the two species. Stigmochelys brachygularis and S. pardalis are inferred to be closely related, and are most likely sister taxa that represent time-successive species of a single lineage. The age structure of the fossil sample, in conjunction with evidence of carnivore damage on the shells, indicates that S. brachygularis was subject to relatively high levels of predation in comparison with modern-day S. pardalis. If S. brachygularis is presumed to have been ecologically similar to modern S. pardalis, it would have been capable of living in a wide range of habitats from semi-desert and savanna to open woodland. The material attributed to “Geochelonelaetoliensis is more fragmentary, and as a consequence it is not possible to determine its precise phylogenetic or taxonomic relationships. However, it likely represents a distinct genus, possibly with affinities to Astrochelys from Madagascar. A more thorough assessment of the relationships of “G.” laetoliensis with Miocene and Pliocene giant tortoises from Africa and with extant genera will have to await the recovery of more complete material from Laetoli. Giant tortoises, such as “G.” laetoliensis, became extinct on mainland Africa during the late Pliocene, possibly associated with the appearance of early Homo and stone tool using behaviors at 2.6 Ma.


Stigmochelys Geochelone Centrochelys Astrochelys Africa phylogeny paleoecology paleobiology 



The author is grateful to the Tanzania Commission for Science and Technology and the Unit of Antiquities in Dar es Salaam for permission to conduct research in Tanzania. Special thanks go to Paul Msemwa (Director) and Amandus Kweka, as well as to all of the staff at the National Museum of Tanzania in Dar es Salaam, for their support and assistance. The Government of Kenya and the National Museums of Kenya are thanked for permission to study the collections in Nairobi. I am especially grateful to Amandus Kweka for help with excavation and preparation of EP 2472/03. Thanks go to Emma Mbua, Mary Muungu, Meave Leakey (Kenya National Museum) and Oliver Hampe and Wolf-Dieter Heinrich (Humboldt-Universität Museum für Naturkunde, Berlin) for access to specimens in their care. I am grateful to F. de Lapparent de Broin, C. Crumley and P. Meylan for reviewing the manuscript and for providing many helpful comments and suggested improvements. For their advice, discussion, help and support I gratefully acknowledge the following individuals: P. Andrews, C. Jolly, T.S. Harrison, D.M.K. Kamamba, M.G. Leakey, S. Odunga, M. Pickford and D. Su. Research on the Laetoli fauna was supported by grants from the National Geographic Society, the Leakey Foundation, and NSF (Grants BCS-9903434 and BCS-0309513).


  1. Andrews, C. W. (1903). On some pleuodiran chelonians from the Eocene of the Fayum, Egypt. Annals and Magazine of Natural History Series, 7(11), 115–122.Google Scholar
  2. Andrews, C. W. (1906). A descriptive catalogue of the Teriary Vertebrata of the Fayum, Egypt. London: Trustees of the British Museum.Google Scholar
  3. Andrews, C. W. (1914). On the Lower Miocene vertebrates from British East Africa, collected by Dr. Felix Oswald. Quarterly Journal Geological Society of London, 70, 163–186.CrossRefGoogle Scholar
  4. Andrews, C. W., & Beadnell, H. J. L. (1903). A preliminary notice of a land tortoise from the Upper Eocene of the Fayum, Egypt. Cairo: Survey Department, National Printing Department.Google Scholar
  5. Archer, W. H. (1948). The mountain tortoise. Geochelone pardalis. African Wild Life, 2, 75–78.Google Scholar
  6. Arambourg, C. (1947). Contribution à l’etude géologique and paléontologique du basin du Lac Rodolphe et de la basse vallée de l’Omo. In: Mission Scientifique de l’Omo 1932–1933. 1. Géologie-Anthropologie, 3, 2ème partie, Paléontologie (pp. 231–562). Paris: Muséum national d’Histoire naturelle.Google Scholar
  7. Arambourg, C. (1979). Vertébrés Villafranchiens d’Afrique du Nord (Artiodactyles, Carnivores, Primates, Reptiles, Oisseaux). Paris: Foundation Singer-Polignac.Google Scholar
  8. Auffenberg, W. (1964). A redefinition of the fossil tortoise genus Stylemis Leidy. Journal of Paleontology, 38, 316–324.Google Scholar
  9. Auffenberg, W. (1974). Checklist of fossil land tortoises (Testudinidae). Bulletin of the Florida State Museum, 18, 121–251.Google Scholar
  10. Auffenberg, W. (1981). The fossil turtles of Olduvai Gorge, Tanzania, Africa. Copeia, 1981(3), 509–522.CrossRefGoogle Scholar
  11. Austin, J. J., Arnold, E. N., & Bour, R. (2003). Was there a second adaptive radiation of giant tortoises in the Indian Ocean? Using mitochondrial DNA to investigate speciation and biogeography of Aldabrachelys (Reptilia, Testudinidae). Molecular Ecology, 12, 1415–1424.CrossRefGoogle Scholar
  12. Bergmann, P. (2001). The natural history of the central Asian tortoise. The Newsletter of the Colorado Herpetological Society, 28(10), 1–4.Google Scholar
  13. Bour, R. (1980). Essai sur la taxonomie des Testudinidae actuels (Reptilia, Chelonii). Bulletin du Muséum National d’Histoire Naturelle, 2(4), 541–546.Google Scholar
  14. Bour, R. (1984). L’identité de Testudo gigantea Schweigger, 1812 (Reptilia, Chelonii). Bulletin du Muséum national d’histoire naturelle 4e série, 6, A(1), 159–175.Google Scholar
  15. Bour, R. (1985). Les tortues terrestres géantes des Iles de l’Océan Indien Occidental: Donnée géographiques, taxonomiques et phylogénétiques. Studia Geologica Salmanticensia. Vol. esp. 1. Studia Paleocheloniologica, 1, 17–76.Google Scholar
  16. Branch, W. (2008). Tortoises, terrapins and turtles of Africa. Cape Town: Struik.Google Scholar
  17. Broadley, D. G. (1962). Some fossil chelonian fragments from Makapansgat. Nature, 194, 791–792.CrossRefGoogle Scholar
  18. Broadley, D. G. (1997). A new species of Psammobates (Reptilia: Testudinidae) from the Early Pleistocene of South Africa. Palaeontologia Africana, 34, 111–114.Google Scholar
  19. Burleigh, R., & Arnold, E. N. (1979). Radiocarbon dating of eggshell of giant tortoise from Denis Island, Seychelles. Philosophical Transactions of the Royal Society of London, B, 286, 160–161.Google Scholar
  20. Caccone, A., Amato, G., Gratry, O. C., Behler, J., & Powell, J. R. (1999). A molecular phylogeny of four endangered Madagascar tortoises based on mtDNA sequences. Molecular Phylogenetics and Evolution, 12, 1–9.CrossRefGoogle Scholar
  21. Claude, J., & Tong, H. (2004). Early Eocene testudinoid turtles from Saint-Papoul, France, with comments on the early evolution of modern Testudinoidea. Oryctos, 5, 3–45.Google Scholar
  22. Congdon, J. D., & Gibbons, J. W. (1985). Egg components and reproductive characteristics of turtles: Relationships to body size. Herpetologia, 41, 194–205.Google Scholar
  23. Congdon, J. D., & Tinkle, D. W. (1982). Reproductive energetics of the painted turtle (Chrysemys picta). Herpetologica, 37, 228–237.Google Scholar
  24. Coulson, I. M., & Hailey, A. (2001). Low survival rate and high predation in the African hingeback tortoise Kinixys spekii. African Journal of Ecology, 39, 383–392.CrossRefGoogle Scholar
  25. Crumly, C. R. (1982). A cladistic analysis of Geochelone using cranial osteology. Journal of Herpetology, 16, 215–234.CrossRefGoogle Scholar
  26. Crumly, C.R (1984). The evolution of land tortoises (family testudinidae). Ph.D. dissertation, Rutgers University, Newark.Google Scholar
  27. de Broin, F. (1979). Chéloniens du Miocène et du Plio-Pléistocène d’Afrique orientale. Bulletin de la Societe Geologique de France, 21, 323–327.CrossRefGoogle Scholar
  28. Ditchfield, P., & Harrison, T. (2011). Sedimentology, lithostratigraphy and depositional history of the Laetoli area. In T. Harrison (Ed.), Paleontology and geology of Laetoli: Human evolution in context (Geology, geochronology, paleoecology and paleoenvironment, Vol. 1, pp. 47–76). Dordrecht: Springer.Google Scholar
  29. Fritz, U., & Bininda-Emonds, O. R. P. (2007). When genes meet nomenclature: Tortoise phylogeny and the shifting generic concepts of Testudo and Geochelone. Zoology, 110, 298–307.CrossRefGoogle Scholar
  30. Fryer, J. C. F. (1911). The structure and formation of Aldabra and neighbouring islands – with notes on their flora and fauna. Transaction of the Linnean Society of London, Zoology, 14, 397–442.CrossRefGoogle Scholar
  31. Gaffney, E. S., & Meylan, P. A. (1988). A phylogeny of turtles. In M. J. Benton (Ed.), The phylogeny aqnd classification of tetrapods. Vol. 1: Amphibians, reptiles, birds, 5. Systematics Association Special Volume N 35A (pp. 157–219). Oxford: Clarendon.Google Scholar
  32. Geraads, D. (1989). Vertébrésfossiles du Miocène supérieur de Djebel Krechem El Artsouma (Tunisie Centrale). Comparaisons biostratigraphiques. Geobios, 22, 777–801.CrossRefGoogle Scholar
  33. Gerlach, J. (2001). Tortoise phylogeny and the ‘Geochelone’ problem. Phelsuma, 9, 1–24.Google Scholar
  34. Greig, J. C., & Burdett, P. D. (1976). Patterns in the distribution of southern African terrestrial tortoises (Cryptodira: Testudinidae). Zoologica Africana, 11, 249–273.Google Scholar
  35. Hailey, A., & Loumbourdis, N. S. (1988). Egg size and shape, clutch dynamics, and reproductive effort in European tortoises. Canadian Journal of Zoology, 66, 1527–1536.CrossRefGoogle Scholar
  36. Harris, J. M., Leakey, M. G., & Cerling, T. E. (2003). Early Pliocene tetrapods from Kanapoi, Lake Turkana Basin, Kenya. In Geology and vertebrate paleontology of the Early Pliocene Site of Kanapoi, Northern Kenya. Contributions in Science 498 (pp. 39–113). Los Angeles: Natural History Museum of Los Angeles County.Google Scholar
  37. Harrison, T., & Kweka, A. (2011). Paleontological localities on the Eyasi Plateau, including Laetoli. In T. Harrison (Ed.), Paleontology and geology of Laetoli: Human evolution in context (Geology, ­geochronology, paleoecology and paleoenvironment, Vol. 1, pp. 17–45). Dordrecht: Springer.Google Scholar
  38. Highfield, A. C. (1990). Keeping and breeding tortoises in captivity. London: Serpent’s Tail.Google Scholar
  39. Hirsch, K. F. (1983). Contemporary and fossil chelonian eggshells. Copeia, 1983(2), 382–397.CrossRefGoogle Scholar
  40. Hirsch, K. F., & Lopez-Jurado, L. F. (1987). Pliocene chelonian fossil eggs from Gran Canaria, Canary Islands. Journal of Vertebrate Paleontology, 7, 96–99.CrossRefGoogle Scholar
  41. Holroyd, P. A., & Parham, J. F. (2003). The antiquity of African tortoises. Journal of Vertebrate Paleontology, 23, 688–690.CrossRefGoogle Scholar
  42. Honegger, R. (1966). Beobachtungen an der herpetofauna der Seychellen. Salamandra, 1–2, 20–36.Google Scholar
  43. Hutterer, R., García-Talavera, F., López-Martínez, N., & Michaux, J. (1997). New chelonian eggs from the Tertiary of Lanzarote and Fuerteventura, and a review of fossil tortoises of the Canary Islands (Reptilia, Testudinidae). Vieraea, 26, 139–161.Google Scholar
  44. Iverson, J. B. (1992). A revised checklist with distribution maps of the turtles of the world. Richmond: Privately Printed.Google Scholar
  45. Kabigumila, J. (2000). Growth and carapacial colour variation of the leopard tortoise, Geochelone pardalis babcocki, in northern Tanzania. African Journal of Ecology, 38, 217–223.CrossRefGoogle Scholar
  46. Kabigumila, J. (2001a). Size composition and sex ratio of the leopard tortoise (Geochelone pardalis) in northern Tanzania. African Journal of Ecology, 39, 393–395.CrossRefGoogle Scholar
  47. Kabigumila, J. (2001b). Incidence of injuries and mortality in the leopard tortoise, Geochelone pardalis, in northern Tanzania. Tanzanian Journal of Science, 27, 101–106.Google Scholar
  48. Kent, P. E. (1941). The recent history and Pleistocene deposits of the plateau north of Lake Eyasi, Tanganyika. Geological Magazine, 78, 173–184.CrossRefGoogle Scholar
  49. Kimbel, W. H., Walter, R. C., Johanson, D. C., Reed, K. E., Aronson, J. L., Assefa, Z., Marean, C. W., Eck, G. G., Bobe, R., Hovers, E., Rak, Y., Vondra, C., Yemane, T., York, D., Chen, Y., Evensen, N. M., & Smith, P. E. (1996). Late Pliocene Homo and Oldowan tools from the Hadar Formation (Kada Hadar Member), Ethiopia. Journal of Human Evolution, 31, 549–561.CrossRefGoogle Scholar
  50. Lapparent de Broin, F. de (2000). African chelonians from the Jurassic to the present: Phases of development and preliminary catalogue of the fossil record. Palaeontologia Africana, 36, 43–82.Google Scholar
  51. Lapparent de Broin, F. de (2003). Miocene chelonians from southern Namibia. In M. Pickford & B. Senut (Eds.), Geology and palaeobiology of the Central and Southern Namibia, palaeontology of the Orange River Valley (Vol. 2, pp. 67–102). Namibia: Geological Survey of Namibia, Memoir 19.Google Scholar
  52. Lapparent de Broin, F. de (2008). Miocene chelonians from south-western Namibia. Memoir of the Geological Survey of Namibia, 20, 107–145.Google Scholar
  53. Lapparent de Broin, F. de, & van Dijk, P. P. (1999). Chelonia from the Late Miocene Baynunah Formation, Emirate of Abu Dhabi, United Arab Emirates: Palaeogeographic implications. In P. J. Whybrow & A. Hill (Eds.), Fossil Vertebrates of Arabia (pp. 136–162). New Haven: Yale University Press.Google Scholar
  54. Le, M., Raxworthy, C. J., McCord, W. P., & Mertz, L. (2006). A molecular phylogeny of tortoises (Testudines: Testudinidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 40, 517–531.CrossRefGoogle Scholar
  55. Leakey, L. S. B. (1935). East African Archaeological Expedition. Fourth Season 1934–5. Eight monthly field report: May 24th–June 23rd. Unpublished report, The Natural History Museum, London.Google Scholar
  56. Leakey, L. S. B. (1965). Olduvai Gorge, 1951–1961. Vol. 1. A preliminary report on the geology and fauna. Cambridge: Cambridge University Press.CrossRefGoogle Scholar
  57. Lehmann, U. (1957). Eine jungpleistozän Wirbeltierfauna aus Östafrika (Material der Expedition Kohl-Larsen 1938 von der Mumba-Höhle). Mitteilungen aus dem Geologischen Staatsinstitut in Hamburg, 26, 100–140.Google Scholar
  58. Loehr, V. J. T., Henen, B. T., & Hofmeyr, M. D. (2004). Reproduction of the smallest tortoise, the Namaqualand speckled padloper, Homopus signatus signatus. Herpetologica, 60, 444–454.CrossRefGoogle Scholar
  59. Loveridge, A. (1935). Scientific results of an expedition to rain forest regions in eastern Africa. I. New reptiles and amphibians from East Africa. Bulletin of the Museum of Comparative Zoology, 79, 1–19.Google Scholar
  60. Loveridge, A., & Williams, E. E. (1957). Revision of the African tortoises and turtles of the suborder Cryptodira. Bulletin of the Museum of Comparative Zoology, 115, 163–557.Google Scholar
  61. Macau-Vilar, F. (1958). Contribución al estudio del mioceno canario. Boletin de la Real Sociedad Española de Historia Natural (Geologica), 56, 477–486.Google Scholar
  62. Marquez, C., Wiedenfeld, D. A., Landázuri, S., & Chávez, J. (2007). Human-caused and natural mortality of giant tortoises in the Galapagos Islands during 1995–2004. Oryx, 41, 337–342.CrossRefGoogle Scholar
  63. Meylan, P., & Auffenberg, W. (1986). New land tortoises (Testudines: Testudinidae) from the Miocene of Africa. Zoological Journal of the Linnean Society, 86, 279–307.CrossRefGoogle Scholar
  64. Meylan, P. A., & Auffenberg, W. (1987). The chelonians of the Laetolil Beds. In M. D. Leakey & J. M. Harris (Eds.), Laetoli: A Pliocene site in northern Tanzania (pp. 62–78). Oxford: Clarendon.Google Scholar
  65. Morris, L. (1994). Western hinge-back tortoise, Kinixys belliana nogueyi. Tortuga Gazette, 30(1), 1–3.Google Scholar
  66. Palkovacs, E. P., Gerlach, J., & Caccone, A. (2002). The evolutionary origin of Indian Ocean tortoises (Dipsochelys). Molecular Phylogenetics and Evolution, 24, 216–227.CrossRefGoogle Scholar
  67. Preston, F. W. (1969). Shapes of birds’ eggs: Extant North American families. The Auk, 86, 246–264.CrossRefGoogle Scholar
  68. Raynal, J. P., Texier, J. P., Geraads, D., & Sbihi-Alaoui, F.-Z. (1990). Un nouveau gisement paléontologique Plio-Pléistocene en Afrique du Nord: Alh Al Oughlam (ancienne carrière Deprez) à Casablanca (Maroc). Comptes Rendus de l’Académie des Sciences, Paris 310, Séries II (3), 315–320.Google Scholar
  69. Rodriquez Bayona, L. O., & Rylander, M. K. (1984). Notes on the biology of the tortoise Geochelone denticulata L. in Peru. Amphibia-Reptilia, 5, 323–327.Google Scholar
  70. Roger, J., Pickford, M., Thomas, H., de Lapparent de Broin, F., Tassy, P., van Neer, W., Bourdillon-de-Grissac, C., & AI-Busaldi, S. (1994). Découverte de vértebrés fossils dans le Miocene de la region du Huqf au Sultanat d’Oman. Annales de Paléontologie, 80, 253–273.Google Scholar
  71. Rothe, P., & Klemmer, K. (1991). Fossil eggs of terrestrial tortoises (Family Testudinindae) from Pliocene calcarenites of Fuerteventura (Canary Islands, Spain). Senckenbergiana Lethaea, 71, 307–317.Google Scholar
  72. Semaw, S., Rogers, M. J., Quade, J., Renne, P. R., Butler, R. F., Dominguez-Rodrigo, M., Stout, D., Hart, W. S., Pickering, T., & Simpson, S. W. (2003). 2.6-Million-year-old stone tools and associated bones from OGS-6 and OGS-7, Gona, Afar, Ethiopia. Journal of Human Evolution, 45, 169–177.CrossRefGoogle Scholar
  73. Spawls, S., Howell, K., Drewes, R., & Ashe, J. (2001). A field guide to the reptiles of East Africa. Princeton: Princeton University Press.Google Scholar
  74. Stromer, E. V. (1926). Land- und Süsswasser Bewohnender Wirbeltierereste aus dem Diamentfeldern Deutsch-Südwestafrikas. In E. Kaiser (Ed.), Die Diamentenwüste Südwestafrikas (Vol. 2, pp. 107–153). Berlin: Reimer.Google Scholar
  75. Thomas, H., Taquet, P., Ligabue, G., & De l’Agnola, C. (1978). Découverte d’un gisements de vertébrés dans the depots continentaux dans le Miocène moyen du Hasa (Arabie Saoudite). Comptes Rendus sommaires de la Société Géologique de France, 2, 69–72.Google Scholar
  76. Thomas, H., Sevket, S., Khan, M., Battail, B., & Ligabue, G. C. (1982). The Lower Miocene fauna of Al-Sarrar (Eastern Province, Saudi Arabia). Atlal. The Journal of Saudi Arabian Archaeology, 5, 109–136.Google Scholar
  77. Thomas, H., Coppens, Y., Thibault, C., & Weidmann, M. (1984). Découverte de vertébrés fossils dans le Plestocène inférieur de la République de Djibouti. Comptes Rendus de l’Académie des Sciences, Paris 299, Série, 2(1), 43–48.Google Scholar
  78. Wallis, I. R., Henen, B. T., & Nagy, K. A. (1999). Egg size and annual egg production by female desert tortoises (Gopherus agassizii): The importance of food abundance, body size, and date of egg shelling. Journal of Herpetology, 33, 394–408.CrossRefGoogle Scholar
  79. Werdelin, L., & Dehghani, R. (2011). Carnivora. In T. Harrison (Ed.), Paleontology and geology of Laetoli: Human evolution in context (Fossil hominins and the associated fauna, Vol. 2, pp. 189–232). Dordrecht: Springer.Google Scholar
  80. Wilson, V. J. (1968). The leopard tortoise, Testudo pardalis babcocki in eastern Zambia. Arnoldia, 40, 1–11.Google Scholar
  81. Wood, R. C. (1987). Fossil turtles from the Sahabi Formation. In N. T. Boaz, A. El-Arnauti, A. W. Gaziry, J. de Heinzelin, & D. D. Boaz (Eds.), Neogene paleontology and geology of sahabi (pp. 107–112). New York: Alan R. Liss.Google Scholar
  82. Wood, R. C. (2003). Fossil turtles from Lothagam. In M. G. Leakey & J. M. Harris (Eds.), Lothagam: The dawn of humanity in eastern Africa (pp. 115–134). New York: Columbia University Press.Google Scholar

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Authors and Affiliations

  1. 1.Center for the Study of Human Origins, Department of AnthropologyNew York UniversityNew YorkUSA

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