Skip to main content Accessibility help

A divergence dating analysis of turtles using fossil calibrations: an example of best practices

  • Walter G. Joyce (a1) (a2), James F. Parham (a3), Tyler R. Lyson (a2) (a4) (a5), Rachel C. M. Warnock (a6) (a7) and Philip C. J. Donoghue (a7)...


Turtles have served as a model system for molecular divergence dating studies using fossil calibrations. However, because some parts of the fossil record of turtles are very well known, divergence age estimates from molecular phylogenies often do not differ greatly from those observed directly from the fossil record alone. Also, the phylogenetic position and age of turtle fossil calibrations used in previous studies have not been adequately justified. We provide the first explicitly justified minimum and soft maximum age constraints on 22 clades of turtles following best practice protocols. Using these data we undertook a Bayesian relaxed molecular clock analysis establishing a timescale for the evolution of crown Testudines that we exploit in attempting to address evolutionary questions that cannot be resolved with fossils alone. Some of these questions, such as whether the turtle crown originated in the Triassic or Jurassic, cannot be resolved by our analysis. However, our results generate novel age-of-origination estimates for clades within crown Testudines. Finally, we compare our fossil calibrations and posterior age estimates to those from other studies, revealing substantial differences in results and interpretation.



Hide All
Anderson, J. S. 2012. Fossils, molecules, divergence times, and the origin of Salamandroidea. Proceedings of the National Academy of Sciences, 109:55575558.
Andrews, C. W. 1914. On the lower Miocene vertebrates from British East Africa collected by Dr. Felix Oswald. Quarterly Journal of the Geological Society, 70:163187.
Anquetin, J. 2012. Reassessment of the phylogenetic interrelationships of basal turtles (Testudinata). Journal of Systematic Palaeontology, 10:345.
Anquetin, J., Barrett, P. M., Jones, M. E. H., Moore-Fay, S., and Evans, S. E. 2009. A new stem turtle from the Middle Jurassic of Scotland: New insights into the evolution and palaeoecology of basal turtles. Proceedings of the Royal Society B: Biological Sciences, 276:879886.
Barley, A. J., Spinks, P. Q., Thomson, R. C., and Shaffer, H. B. 2010. Fourteen nuclear genes provide phylogenetic resolution for difficult nodes in the turtle tree of life. Molecular Phylogenetics and Evolution, 55:11891194.
Belshaw, R. and Katzourakis, A. 2005. BlastAlign: A program that uses blast to align problematic nucleotide sequences. Bioinformatics, 21:122123.
Benton, M. J. and Donoghue, P. C. J. 2007. Paleontological evidence to date the Tree of Life. Molecular Biology and Evolution, 24:2653.
Benton, M. J., Donoghue, P. C. J., and Asher, R. J. 2009. Calibrating and constraining molecular clocks, p. 3586. InHedges, S. B. and Kumar, S.(eds.), The Timetree of Life. Oxford University Press, Oxford.
Baur, G. 1893. Notes on the classification of the Cryptodira. American Naturalist, 27:672675.
Bona, P. and de la Fuente, M. S. 2005. Phylogenetic and paleobiogeographic implications of Yaminuechelys maior (Staesche, 1929) new comb., a large long-necked chelid turtle from the early Paleocene of Patagonia, Argentina. Journal of Vertebrate Paleontology, 25:569582.
Bourque, J. R. 2013. Fossil Kinosternidae from the Oligocene and Miocene of Florida, U.S.A., p. 459475. InBrinkman, D. B., Holroyd, P. A., and Gardner, J. D.(eds.), Morphology and Evolution of Turtles. Springer.
Bräm, H. 1965. Die Schildkröten aus dem oberen Jura (Malm) der Gegend von Solothurn. Schweizerische Paläontologische Abhandlungen, 83:1190.
Brinkman, D. B. 2003. A review of nonmarine turtles from the Late Cretaceous of Alberta. Canadian Journal of Earth Sciences, 40:557571.
Brinkman, D. B. 2005. Turtles: Diversity, paleoecology, and distribution, p. 202220. InCurrie, P. J. and Koppelhus, E. B.(eds.), Dinosaur Provincial Park. Indiana University Press, Bloomington.
Brinkman, D., Aquillon-Martinez, M. C., de, C. A.Dávila, Leon, Jamnickzy, H., Eberth, D. A., and Colbert, M. 2009. Euclastes coahuilaensis sp. nov., a basal cheloniid turtle from the late Campanian Cerro del Pueblo Formation of Coahuila State, Mexico. PaleoBios, 28:7688.
Brinkman, D., Hart, M., Jamniczky, H., and Colbert, M. 2006. Nichollsemys baieri gen. et sp. nov, a primitive chelonioid turtle from the late Campanian of North America. Paludicola, 5:111124.
Brinkman, D. B., Li, J., and Ye, X. 2008. Order Testudines, p. 35103. InLi, J., Wu, X., and Zhang, F.(eds.), The Chinese Fossil Reptiles and Their Kin. Science Press, Beijing.
Brinkman, D. and de la Rosa, R. Rodriguez. 2006. Nonmarine turtles from the Cerro del Pueblo Formation (Campanian), Coahuila State, Mexico. New Mexico Museum of Natural History and Science Bulletin, 35:229233.
Brinkman, D. B., Stadtman, K., and Smith, D. 2000. New material of Dinochelys whitei Gaffney, 1979, from the Dry Mesa Quarry (Morrison Formation, Jurassic) of Colorado. Journal of Vertebrate Paleontology, 20:269274.
Brochu, C.A. 2004. Calibration age and quartet divergence date estimation. Evolution, 58:13751382.
Broin, F. de. 1980. Les tortues de Gadoufaoua (Aptien du Niger); Aperçu sur la paléobiogéographie des Pelomedusidae (Pleurodira). Mémoires de la Société Géologique de France, 139:3946.
Broin, F. de. 1987. Lower vertebrates from the early-middle Eocene Kuldana Formation of Kohat (Pakistan): Chelonia. Contributions from the Museum of Paleontology, University of Michigan, 27:169185.
Cadena Rueda, E. A. and Gaffney, E. S. 2005. Notoemys zapatocaensis, a new side-necked turtle (Pleurodira : Platychelyidae) from the early Cretaceous of Colombia. American Museum Novitates, 3470:119.
Cadena, E., Jaramillo, C., and Paramo, M. E. 2008. New material of Chelus colombiana (Testudines: Pleurodira) from the lower Miocene of Colombia. Journal of Vertebrate Paleontology, 28:12061212.
Cadena, E. W., Jaramillo, C. A., and Bloch, J. I. 2013. New material of the platychelyid turtle Notoemys zapatocaensis from the Early Cretaceous of Colombia; implications for understanding Pleurodira evolution, p. 105120. InBrinkman, D. B., Holroyd, P. A., and Gardner, J. D.(eds.), Morphology and Evolution of Turtles. Springer.
Carrano, M. T. and Velez-Juarbe, J. 2006. Paleoecology of the Quarry 9 vertebrate assemblage from Como Bluff, Wyoming (Morrison Formation, Late Jurassic). Palaeogeography, Palaeoclimatology, Palaeoecology, 237:147159.
Carroll, R. L. 1988. Vertebrate Paleontology and Evolution. Freeman, New York, 698 p.
Ceballos, C. P., Adams, D. C., Iverson, J. B., and Valenzuela, N. 2012. Phylogenetic patterns of sexual size dimorphism in turtles and their implications for Rensch's rule. Evolutionary Biology, online only as of March 2013.
Chiari, Y., Cahais, V., Galter, N., and Delsuc, F. 2012. Phylogenomic analyses support the position of turtles as the sister group of birds and crocodiles (Archosauria). BMC Biology 10:65.
Chkhikvadze, V. M. 1971. First find of the tertiar tortoise of the family Platysternidae. Paleontologichesky Zhurnal, 4:137139.
Chkhikvadze, V. M. 1983. Fossil Turtles of the Caucasus and Northern Regions of the Black Sea. Metsniereba, Tbilisi, 151p. (In Russian)
Chkhikvadze, V. M. 1987. Sur la classification et les caractères de certaines tortues fossiles d'Asie, rares et peu étudiées. Studia Palaeocheloniologica, 2:5586.
Chkhikvadze, V. M. 1990. Paleogene turtles of the USSR. Metsniereba, Tbilisi, 95p. (In Russian)
Clark, J. 1937. The stratigraphy and paleontology of the Chadron formation in the Big Badlands of South Dakota. Annals of the Carnegie Museum, 25:261350.
Clarke, J., Donoghue, P. C. J., and Warnock, R. C. M. 2011. Establishing a timescale for plant evolution. New Phytologist, 192:266301.
Claude, J. and Tong, H. 2012. A reinterpretation of the pre-Eocene Testudinoids, p. 14. InSymposium on Turtle Evolution 2012, Program and Abstracts. University of Tübingen, Tübingen.
Claude, J., Suteethorn, V., and Tong, H. 2007. Turtles from the late Eocene–early Oligocene of the Krabi Basin (Thailand). Bulletin de la Société Géologique de France, 178:305316.
Cope, E. D. 1871. A description of the genus Protostega, a form of extinct Testudinata. Proceedings of the American Philosophical Society, 12:422433.
Cope, E. D. 1873. [Description of Toxochelys latiremis]. Proceedings of the Academy of Natural Sciences of Philadelphia, 1873:10.
Croft, D. A. 2007. The middle Miocene (Laventan) Quebrada Honda Fauna, southern Bolivia and a description of its notoungulates. Palaeontology, 50:277303.
Danilov, I. G. 1999. A new lindholmemydid genus (Testudines: Lindholmemydidae) from the mid-Cretaceous of Uzbekistan. Russian Journal of Herpetology, 6:6371.
Danilov, I. G. and Parham, J. F. 2005. A reassessment of the referral of an isolated skull from the Late Cretaceous of Uzbekistan to the stem-testudinoid turtle genus Lindholmemys. Journal of Vertebrate Paleontology, 25:784791.
Danilov, I. G. and Parham, J. F. 2006. A redescription of ‘Plesiochelys' tatsuensis, a turtle from the Late Jurassic of China, and its bearing on the antiquity of the crown clade Cryptodira. Journal of Vertebrate Paleontology, 26:573580.
Danilov, I. G. and Parham, J. F. 2008. A reassessment of some poorly known turtles from the Middle Jurassic of China with comments on the antiquity of extant turtles. Journal of Vertebrate Paleontology, 28:306318.
Danilov, I. G. and Vitek, N. S. 2013. Cretaceous trionychids of Asia: An expanded review of their record and biogeography, p. 419438. InBrinkman, D. B., Holroyd, P. A., and Gardner, J. D.(eds.), Morphology and Evolution of Turtles. Springer.
Donoghue, P. C. J. and Benton, M. J. 2007. Rocks and clocks: Calibrating the Tree of Life using fossils and molecules. Trends in Ecology and Evolution, 22:424431.
Dornburg, A., Beaulieu, J. M., Oliver, J. N., and Near, T. J. 2011. Integrating fossil preservation biases in the selection of calibrations for molecular divergence time estimation. Systematik Biology, 60:519527.
dos Reis, M., Inoue, J., Hasegawa, M., Asher, R. J., Donoghue, P. C. J., and Yang, Z. 2012. Phylogenomic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny. Proceedings of the Royal Society B: Biological Sciences, 279:34913500.
Drake, R. E., van Couvering, J. A., Pickford, M. H., Curtis, G. H., and Harris, J. A. 1988. New chronology for the Early Miocene mammalian faunas of Kisingiri, Western Kenya. Journal of the Geological Society, 145:479491.
Drummond, A. J., Ho, S. Y. W., Phillips, M. J., and Rambaut A, A. 2006. Relaxed phylogenetics and dating with confidence. PLoS Biology, 4:e88.
Drummond, A. J. and Rambaut, A. 2007. BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology, 7:214.
Dryden, L. S. 1988. Paraphyly of the Cryptodira and phylogenetic systematics of turtles. Unpublished Master's Thesis, University of Kansas, 142p.
Eaton, J. G., Cifelli, R. L., Hutchison, J. H., Kirkland, J. I., and Parrish, J. M. 1999. Cretaceous vertebrate faunas from the Kaiparowits Plateau, south-central Utah, p. 345353. InGillette, D. D.(ed.), Vertebrate paleontology in Utah. Utah Geological Survey, Salt Lake City.
Edgar, R. C. 2004. MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32:17921797.
Engstrom, T. N., Shaffer, H. B., and McCord, W. P. 2004. Multiple data sets, high homoplasy, and the phylogeny of softshell turtles (Testudines, Trionychidae). Systematic Biology, 53:693710.
Erickson, B. R. 1973. A new chelydrid turtle Protochelydra zangerli from the late Paleocene of North Dakota. Scientific Publications of the Science Museum of Minnesota, 2:116.
Fara, E., Saraiva, A. A. F., Campos, D. D. A., Moreira, J. K. R., Siebra, D. D. C., and Kellner, A. W. A. 2005. Controlled excavations in the Romualdo Member of the Santana Formation (Early Cretaceous, Araripe Basin, northeastern Brazil): Stratigraphic, palaeo-environmental and palaeoecological implications. Palaeogeography, Palaeoclimatology, Palaeoecology, 218:145158.
Feldman, C. R. and Parham, J. F. 2002. Molecular phylogenetics of emydine turtles: Taxonomic revision and the evolution of shell kinesis. Molecular Phylogenetics and Evolution, 22:388398.
Feldmann, R. M, Schweitzer, C. E., Redman, C. M., Morris, N. J., and Ward, D. J. 2007. New Late Cretaceous lobsters from the Kyzylkum desert of Uzbekistan. Journal of Paleontology, 81:701713.
Fraas, E. 1913. Proterochersis, eine pleurodire Schildkroete aus dem Keuper. Jahreshefte der Gesellschaft für Naturkunde in Württemberg, 69:1390.
de la Fuente, M. S. and Bona, P. 2002. Una nueva especie del género Hydromedusa Wagler (Pleurodira Chelidae) del Paléogeno de Patagonia. Ameghiniana, 39:7783.
de la Fuente, M. S. and Iturralde-Vinent, M. 2001. A new pleurodiran turtle from the Jagua Formation (Oxfordian) of Western Cuba. Journal of Paleontology, 75:860869.
de la Fuente, M. S.,de Lapparent de Broin, F., and Manera de Bianco, T. 2001. The oldest and first nearly complete skeleton of a chelid, of the Hydromedusa sub-group (Chelidae, Pleurodira), from the Upper Cretaceous of Patagonia. Bulletin de la Société Géologique de France, 172:237244.
de la Fuente, M. S., Umazano, A. M., Sterli, J., and Carballido, J. L. 2011. New chelid turtles of the lower section of the Cerro Barcino formation (Aptian-Albian?), Patagonia, Argentina. Cretaceous Research, 32:527537
Fujita, M. 2003. Geological age and correlation of the vertebrate-bearing horizons in the Tetori Group. Memoir of the Fukui Prefectural Dinosaur Museum, 2:314.
Fujita, M. K., Engstrom, T. N., Starkey, D. E., and Shaffer, H. B. 2004. Turtle phylogeny: Insights from a novel nuclear intron. Molecular Phylogenetics and Evolution, 31:10311040.
Gaffney, E. S. 1972. The systematics of the North American family Baenidae (Reptilia, Cryptodira). Bulletin of the American Museum of Natural History, 147:243319.
Gaffney, E. S. 1975. A phylogeny and classification of the higher categories of turtles. Bulletin of the American Museum of Natural History, 155:389436.
Gaffney, E. S. 1977. The side-necked turtle family Chelidae: A theory of relationships using shared derived characters. American Museum Novitates, 2620:128.
Gaffney, E. S. 1979. The Jurassic turtles of North America. Bulletin of the American Museum of Natural History, 162:91135.
Gaffney, E. S. 1981. A review of the fossil turtles of Australia. American Museum Novitates, 2720:138.
Gaffney, E. S. 1996. The postcranial morphology of Meiolania platyceps and a review of the Meiolaniidae. Bulletin of the American Museum of Natural History, 229:1166.
Gaffney, E. S., de Almeida Campos, D., and Hirayama, R. 2001. Cearachelys, a new side-necked turtle (Pelomedusoides: Bothremydidae) from the Early Cretaceous of Brazil. American Museum Novitates, 3319:120.
Gaffney, E. S., Archer, M., and White, A. 1989. Chelid turtles from the Miocene freshwater limestones of Riversleigh Station, Northwestern Queensland, Australia. American Museum Novitates, 2959:110.
Gaffney, E. S. and Jenkins, F. A. Jr. 2010. The cranial morphology of Kayentachelys, an Early Jurassic cryptodire, and the early history of turtles. Acta Zoologica, 91:335368.
Gaffney, E. S., Hutchison, J. H., Jenkins, F. A. Jr., and Meeker, L. J. 1987. Modern turtle origins: The oldest known cryptodire. Science, 237:289291.
Gaffney, E. S. and Meylan, P. A. 1988. A phylogeny of turtles, p. 157219. InBenton, M. J.(ed.), The Phylogeny and Classification of the Tetrapods, Volume 1, Amphibians, Reptiles, Birds. Clarendon Press, Oxford.
Gaffney, E. S., Meylan, P. A., Wood, R. C., Simons, E., and de Almeida Campos, D. 2011. Evolution of the side-necked turtles: The family Podocnemididae. Bulletin of the American Museum of Natural History, 350:1237.
Gaffney, E. S., Rich, T. H., Vickers-Rich, P., Constantine, A., Vacca, P., and Kool, L. 2007. Chubutemys, a new eucryptodiran turtle from the Early Cretaceous of Argentina, and the relationships of the Meiolaniidae. American Museum Novitates, 3599:135.
Gaffney, E. S., Tong, H., and Meylan, P. A. 2006. Evolution of the side-necked turtles: The families Bothremydidae, Euraxemydidae, and Araripemydidae. Bulletin of the American Museum of Natural History, 300:1318.
Gardner, J. D., Russell, A. R., and Brinkman, D. B. 1995. Systematics and taxonomy of soft-shelled turtles (family Trionychidae) from the Judith River Group (mid Campanian) of North America. Canadian Journal of Earth Sciences, 32:631643.
Georges, A., Birrell, J., Saint, K. M., McCord, W., and Donnellan, S. C. 1998. A phylogeny for side-necked turtles (Chelonia: Pleurodira) based on mitochondrial and nuclear gene sequence variation. Biological Journal of the Linnean Society, 67:213246.
Gilmore, C. W. 1919. Reptilian faunas of the Torrejon, Puerco, and underlying Upper Cretaceous Formations of San Juan County, New Mexico. U.S. Geological Survey, Professional Paper 119, p. 168.
Gilmore, C. W. 1933. A new species of extinct turtle from the upper Pliocene of Idaho. Proceedings of the United States National Museum, 82:17.
Gradstein, F., Ogg, J., and Smith, A. 2004. A Geologic Time Scale. Cambridge University Press, Cambridge, 610p.
Gray, J. E. 1869. Notes on the families and genera of tortoises (Testudinata), and on the characters afforded by the study of their skulls. Proceedings of the Zoological Society of London, 1869:165223.
Gray, J. E. 1870. Supplement to the catalogue of shield reptiles in the collection of the British Museum, Part 1, Testudinata (Tortoises). Taylor and Francis, London, 120p.
Hart, W. K. and Brueseke, M. E. 1999. Analysis and dating of volcanic horizons from Hagerman Fossil Beds National Monument and a revised interpretation of eastern Glens Ferry Formation chronostratigraphy. National Park Service Report 1443-PX9608-97-003, p. 137.
Hay, O. P. 1904. On some fossil turtles belonging to the Marsh collection in Yale University Museum. American Journal of Sciences, 4:261276.
Hay, O. P. 1908. The fossil turtles of North America. Carnegie Institution of Washington Publication, 75:1568.
Head, J., Hutchison, J. H., and Evans, D. C. 2009. The phylogenetic interrelationships and antiquity of plastomenid turtles, p. 71. InGaffney Turtle Symposium, Abstract Volume. Royal Tyrrell Museum of Palaeontology, Drumheller.
Heath, T. A. 2012. A hierarchical Bayesian model for calibrating estimates of species divergence times. Systematic Biology, 61:793809
Hedges, S. B. and Kumar, S. 2004. Precision of molecular time estimates. Trends in Genetics, 20:242247.
Heled, J. and Drummond, A. J. 2012. Calibrated tree priors for relaxed phylogenetics and divergence time estimation. Systematic Biology, 61:138149.
Hervet, S. 2004. Systematique du groupe “Palaeochelys sensu lato - Mauremys” (Chelonii, Testudinoidea) du Tertiaire d'Europe occidentale: Principaux resultats. Annales de Paléontologie, 90:1378.
Hirayama, R. 1985. Cladistic analysis of batagurine turtles. Studia Palaeocheloniologica, 1:140157.
Hirayama, R. 1994. Phylogenetic systematics of chelonioid sea turtles. Island Arc, 3:270284.
Hirayama, R. 1997. Distribution and diversity of Cretaceous chelonioids, p. 225241. InCallaway, J. M. and Nicholls, E. L.(eds.), Ancient Marine Reptiles. Academic Press, London.
Hirayama, R. 1998. Oldest known sea turtle. Nature, 392:705708.
Hirayama, R. 2002. A preliminary report of the fossil turtles from the Kitadani formation (Early Cretaceous) of the Tetori Group of Katsuyama, Fukui Prefecture, Central Japan. Memoir of the Fukui Prefectural Dinosaur Museum, 1:2940.
Hirayama, R. 2007. Cranial morphology of Mesodermochelys (Chelonioidea; Testudines) from the Late Cretaceous of Japan. Journal of Vertebrate Paleontology, Supplement, 27:89A.
Hirayama, R., Brinkman, D. B., and Danilov, I. G. 2000. Distribution and biogeography of non-marine Cretaceous turtles. Russian Journal of Herpetology, 7:181198.
Hirayama, R. and Chitoku, T. 1996. Family Dermochelyidae (Superfamily Chelonioidea) from the Upper Cretaceous of North Japan. Transactions and Proceedings of the Palaeontological Society of Japan, 184:597622.
Hirayama, R. and Hikida, Y. 1998. Mesodermochelys (Testudines, Chelonioidea, Dermochelyidae) from the Late Cretaceous of Nakagawa-cho, Hokkaido, North Japan. Bulletin of Nakagawa Museum of Natural History, 1:6976.
Hirayama, R., Isaji, S., and Hibino, T. 2013. Kappachelys okurai gen. et sp. nov., a new stem soft-shelled turtle from the Early Cretaceous of Japan, p. 179185. InBrinkman, D. B., Holroyd, P. A., and Gardner, J. D.(eds.), Morphology and Evolution of Turtles. Springer.
Ho, S. Y. W. and Phillips, M. J. 2009. Accounting for calibration uncertainty in phylogenetic estimation of evolutionary divergence times. Systematic Biology, 58:367380.
Holman, J. A. 1977. Comments on turtles of the genus Chrysemys Gray. Herpetologica, 33:274276.
Holroyd, P. A. and Hutchison, J. H. 2002. Patterns of geographic variation in latest Cretaceous vertebrates: Evidence from the turtle component. Geological Society of America Special Paper 361, p. 177190.
Holroyd, P. A., Hutchison, J. H., Strait, S. G. 2001. Changes in turtle diversity and abundance through the earliest Eocene Willwood Formation: Preliminary results. University of Michigan Papers on Paleontology, 33:97108.
Holroyd, P. A. and Parham, J. F. 2003. The antiquity of African tortoises. Journal of Vertebrate Paleontology, 23:688690.
Holroyd, P. A., Parham, J. F., and Hutchison, J. H. 2005. A reappraisal of some Paleogene turtles from the southeastern United State. Journal of Vertebrate Paleontology, 25:979982.
Hugall, A. F., Foster, R., and Lee, M. S. Y. 2007. Calibration choice, rate smoothing, and the pattern of tetrapod diversification according to the long nuclear gene RAG-1. Systematic Biology, 56:543563.
Hutchison, J. H. 1991. Early Kinosterninae (Reptilia: Testudines) and their phylogenetic significance. Journal of Vertebrate Paleontology, 11:145167.
Hutchison, J. H. 1996. Testudines, p. 337353. InProthero, D. R. and Emry, R. J.(eds.), The Terrestrial Eocene-Oligocene Transition in North America. Cambridge University Press, Cambridge.
Hutchison, J. H. 1998. Turtles across the Paleocene/Eocene epoch boundary in west-central North America, p. 401408. InAubry, M.-R., Lucas, S. G., and Berggren, W. A.(eds.), Late Paleocene–Early Eocene Climatic and Biotic Events in the Marine and Terrestrial Records. Princeton University Press, Princeton.
Hutchison, J. H. 2013. New turtles from the Paleogene of North America, p. 477497. InBrinkman, D. B., Holroyd, P. A., and Gardner, J. D.(eds.), Morphology and Evolution of Turtles. Springer.
Hutchison, J. H. and Archibald, J. D. 1986. Diversity of turtles across the Cretaceous/Tertiary boundary in northeastern Montana. Palaeogeography, Palaeoclimatology, Palaeoecology, 55:122.
Hutchison, J. H. and Bramble, D. M. 1981. Homology of the plastral scales of the Kinosternidae and related turtles. Herpetologica, 37:7385.
Hutchison, J. H., Eaton, J. G., Holroyd, P. A., and Goodwin, M. B. 1998. Larger vertebrates of the Kaiparowits Formation (Campanian) in the Grand Staircase-Escalante National Monument and adjacent areas, p. 391398. InHill, L. M. and Koselak, J. J.(eds.), Learning from the Land. Grand Staircase-Escalante National Monument Science Symposium Proceedings. U.S. Department of the Interior, Bureau of Land Management, Washington D.C.
Hutchison, J. H. and Holroyd, P. A. 2003. Late Cretaceous and early Paleocene turtles of the Denver Basin, Colorado. Rocky Mountain Geology, 38:121142.
Hutchison, J. H. and Weems, R. E. 1998. Paleocene turtle remains from South Carolina. Transactions of the American Philosophical Society, 88:165195.
Hutchison, J. H., Knell, M. J., and Brinkman, D. B.In Press. Turtles from the Late Cretaceous (Campanian) Kaiparowits Formation, Utah. InTitus, A. L. and Loewen, M. A.(eds.), At the Top of the Grand Staircase, The Late Cretaceous of Southern Utah. Indiana University Press, Bloomington.
Inoue, J., Donoghue, P. C. J., and Yang, Z. 2010. The impact of the representation of fossil calibrations on Bayesian estimation of species divergence times. Systematic Biology, 59:7489.
Jackson, D. R. 1988. A re-examination of fossil turtles of the genus Trachemys (Testudines: Emydidae). Herpetologica, 44:317325.
Jaffe, A. L., Slater, G. J., and Alfaro, M. E. 2011. The evolution of island gigantism and body size variation in tortoises and turtles. Biology Letters,7:558561.
Joyce, W. G. 2000. The first complete skeleton of Solnhofia parsonsi (Cryptodira, Eurysternidae) from the Upper Jurassic of Germany and its taxonomic implications. Journal of Paleontology, 74:684700.
Joyce, W. G. 2003. A new Late Jurassic turtle specimen and the taxonomy of Palaeomedusa testa and Eurysternum wagleri. PaleoBios, 23:18.
Joyce, W. G. 2007. Phylogenetic relationships of Mesozoic turtles. Bulletin of the Peabody Museum of Natural History, 48:3102.
Joyce, W. G. and Bell, C. J. 2004. A review of the comparative morphology of extant testudinoid turtles (Reptilia: Testudines). Asiatic Herpetological Ressearch, 10:53109.
Joyce, W. G., Klein, N., and Mörs, T. 2004. Carettochelyine turtle from the Neogene of Europe. Copeia, 2004:405410.
Joyce, W. G. and Lyson, T. R. 2010 a. A neglected lineage of North American turtles fills a major gap in the fossil record. Palaeontology, 53:241248.
Joyce, W. G. and Lyson, T. R. 2010 b. Pangshura tatrotia, a new species of pond turtle (Testudinoidea) from the Pliocene Siwaliks of Pakistan. Journal of Systematic Palaeontology, 8:449458.
Joyce, W. G. and Lyson, T. R. 2011. New material of Gilmoremys lancensis nov. comb. (Testudines: Trionychidae) from the Hell Creek Formation and the diagnosis of plastomenid turtles. Journal of Paleontology, 85:442459.
Joyce, W. G., Parham, J. F., and Gauthier, J. A. 2004. Developing a protocol for the conversion of rank-based taxon names to phylogenetically defined clade names, as exemplified by turtles. Journal of Paleontology, 78:9891013.
Joyce, W. G., Revan, A., Lyson, T. R., and Danilov, I. G. 2009. Two new plastomenine softshell turtles from the Paleocene of Montana. Bulleting of the Peabody Museum of Natural History, 50:307325.
Joyce, W. G. and Sterli, J. 2012. Congruence and the phylogeny of basal turtles. Acta Zoologica, 93:149159.
Karl, H.-V. 1999. Die Zoogeographie der känozoischen Weichschildkröte Trionyx triunguis Forskal, 1775. Joanneum-Geologie und Paläontologie, 1:2760.
Kear, B. K. and Lee, M. S. Y. 2006. A primitive protostegid from Australia and early sea turtle evolution. Biology Letters, 2:116119.
Kishino, H., Thorne, J. L., and Bruno, W. J. 2001. Performance of a divergence time estimation method under a probabilistic model of rate evolution. Molecular Biology and Evolution, 18:352361.
Knauss, G., Joyce, W. G., Lyson, T. R., and Pearson, D. 2011. A new kinosternoid from the Late Cretaceous Hell Creek Formation of North Dakota and Montana and the origin of the Dermatemys mawii lineage. Paläontologische Zeitschrift, 85:125142.
Krenz, J. G., Naylor, G. J. P., Shaffer, H. B., and Janzen, F. J. 2005. Molecular phylogenetics and evolution of turtles. Molecular Phylogenetics and Evolution, 37:178191.
Kuhn, O. 1964. Fossilium Catalogus, Vol. 1, Animalia, Part 107, Testudines. Ysel Press, Gravenhage, 299p.
Lapparent de Broin, F. de. 2000a. African chelonians from the Jurassic to the present: Phases of development and preliminary catalogue of the fossil record. Palaeontologia Africana, 36:4382.
Lapparent de Broin, F. de. 2000b. The oldest pre-podocnemidid turtle (Chelonii, Pleurodira), from the early Cretaceous, Ceará state, Brasil, and its environment. Treballs del Museu Geologia de Barcelona, 9:4395.
Lapparent de Broin, F. de. 2001. The European turtle fauna from the Triassic to the Present. Dumerilia, 4:155217.
Lapparent de Broin, F. and de la Fuente, M. S. 2001. Oldest world Chelidae (Chelonii, Pleurodira), from the Cretaceous of Patagonia, Argentina. Earth and Planetary Sciences, 333:463470.
Lapparent de Broin, F., de la Fuente, M. S., and Fernandes, M. S. 2007. Notoemys laticentralis (Chelonii, Pleurodira), Late Jurassic of Argentina: New examination of the anatomical structures and comparisons. Revue de Paléobiologie, 26:99136.
Lapparent de Broin, F. and Molnar, R. 2001. Eocene chelid turtles from Redbank Plains, Southeast Queensland, Australia. Geodiversitas, 23:4179.
Lehman, T. M. and Tomlinson, S. L. 2004. Terlinguachelys fischbecki, a new genus and species of sea turtle (Chelonioidea: Protostegidae) from the Upper Cretaceous of Texas. Journal of Paleontology, 78:11631178.
Li, C., Wu, X.-C., Rieppel, O., Wang, L.-T., Zhao, L.-J. 2008. An ancestral turtle from the Late Triassic of southwestern China. Nature, 456:497501.
Linnaeus, C. 1758. Systema Naturae, Volume 1 (tenth edition). Laurentius Salvius, Holmia, 824p.
Linnaeus, C. 1766. Systema Naturae, Volume 1 (twelfth edition). Laurentius Salvius, Holmia, 532p.
Lipka, T. R., Therrien, F., Weishampel, D. B., Jamniczky, H. A., Joyce, W. G., Colbert, M. W., and Brinkman, D. B. 2006. A new turtle from the Arundel Clay facies (Potomac Formation, Early Cretaceous) of Maryland, U.S.A. Journal of Vertebrate Paleontology, 26:300307.
Loomis, F. B. 1904. Two new river reptiles from the Titanotherium beds. American Journal of Science, 4:427432.
Lourenço, J. M., Claude, J., Galtier, N., and Chiari, Y. 2012. Dating cryptodiran nodes: Origin and diversification of the turtle superfamily Testudinoidea. Molecular Phylogenetics and Evolution, 62:496507.
Lydekker, R. 1889. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History), Pt. 3, Chelonia. Longmans, London, 239p.
Lyson, T. R., Bever, G. S., Bhullar, B.-A. S., Joyce, W. G., and Gauthier, J. A. 2010. Transitional fossils and the origin of turtles. Biology Letters, 6:830833.
Madden, R. H., Guerrero, J., Kay, R. F., Flynn, J. J., Swisher, C. C. III, and Walton, A. H. 1997. The Laventan Stage and Age, p. 355381. InKay, R. F., Madden, R. H., Cifelli, R. L., and Flynn, J. J.(eds.), Vertebrate Paleontology in the Neotropics: The Miocene Fauna of La Venta, Colombia. Smithsonian Institution Press, Washington, D.C.
Maisey, J. G. 1990. Stratigraphy and depositional environment of the Crato member (Santana Formation), Lower Cretaceous of N.E. Brazil. Bulletin of the American Museum of Natural History, 195:1519.
Marshall, C. R. 2008. A simple method for bracketing absolute divergence times on molecular phylogenies using multiple fossil calibration points. American Naturalist, 171:726742.
Martill, D. M. 2007. The age of the Cretaceous Santana Formation fossil Konservat Lagerstätte of north-east Brazil: A historical review and an appraisal of the biochronostratigraphic utility of its palaeobiota. Cretaceous Research, 28:895920.
Meylan, P. A. 1987. The phylogenetic relationships of soft-shelled turtles (Family Trionychidae). Bulletin of the American Museum of Natural History, 186:1101.
Meylan, P. A. and Gaffney, E. S. 1989. The skeletal morphology of the Cretaceous cryptodiran turtle, Adocus, and the relationships of the Trionychoidea. American Museum Novitates, 2941:160.
Meylan, P. A. and Gaffney, E. S. 1992. Sinaspideretes is not the oldest trionychid turtle. Journal of Vertebrate Paleontology, 12:257259.
Meylan, P. A., Moody, T. J., Walker, C. A., and Chapman, S. D. 2000. Sandownia harrisi, a highly derived trionychoid turtle (Testudines: Cryptodira) from the Early Cretaceous of the Isle of Wight. Journal of Vertebrate Paleontology, 20:522532.
Meylan, P. A., Weig, B. S., and Wood, R. C. 1990. Fossil soft-shelled turtles (family Trionychidae) of the Lake Turkana Basin, Africa. Copeia, 1990:508528.
Mlynarski, M. 1976. Handbuch der Paläoherpetologie. Part 7, Testudines. Gustav Fischer Verlag, Stuttgart, 130p.
Müller, J. and Reisz, R. R. 2005. Four well-constrained calibration points from the vertebrate fossil record for molecular clock estimates. BioEssays, 27:10691075.
Nakajima, Y., Danilov, I. G., and Hirayama, R. 2009. Plywood-like structures in costal bones of mid- to late Cretaceous trionychids from Asia, p. 128. InGaffney Turtle Symposium, Abstract Volume. Royal Tyrrell Museum of Palaeontology, Drumheller.
Near, T. J., Meylan, P. A., and Shaffer, H. B. 2005. Assessing concordance of fossil calibration points in molecular clock studies: An example using turtles. American Naturalist, 165:137146.
Nessov, L. A. and Krasovskaya, T. B. 1984. Changes in the composition of turtles assemblages of Late Cretaceous of Middle Asia. Vestnik Leningradskogo Gosudarstvennogo Universiteta, 3:1525. (In Russian)
Nicholls, E. L. 1988. New material of Toxochelys latiremis Cope, and a revision of the genus Toxochelys (Testudines, Chelonioidea). Journal of Vertebrate Paleontology, 8:181187.
Nielsen, E. 1959. Eocene turtles from Denmark. Meddelelser fra Dansk Geologisk Forening, 14:96114.
Ogg, J. G., Ogg, G., and Gradstein, F. M. 2008. The Concise Geologic Time scale. Cambridge University Press, Cambridge, 177p.
Oliveira, G. R. de and Kellner, A. W. A. 2007. A new side-necked turtle (Pleurodira, Pelomedusoides) from the Santana Formation (Early Cretaceous), Araripe Basin, Northeastern Brazil. Zootaxa, 1425:5361.
Ostrom, J. H. 1970. Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of the Bighorn Basin Area, Wyoming and Montana. Bulleting of the Peabody Museum of Natural History, 35:1234.
Owen, R. and Bell, T. 1849. Monograph on the fossil Reptilia of the London Clay, and of the Bracklesham and other Tertiary beds. Part I. Chelonia. Palaeontographical Society Monographs, 2:176.
Owen, R. 1861. Palaeontology or a Systematic Summary of Extinct Animals and their Geological Relations, second edition. Edinburgh, 463p.
Parham, J. F., Donoghue, P. C. J., Bell, C. J., et al. (22 co-authors). 2012. Best practices for using paleontological data for molecular divergence dating analyses. Systematic Biology, 61:346359.
Parham, J. F., Feldman, C. R., and Boore, J. L. 2006. The complete mitochondrial genome of the enigmatic bigheaded turtle (Platysternon): Description of unusual genomic features and the reconciliation of phylogenetic hypotheses based on mitochondrial and nuclear DNA. BMC Evolutionary Biology 6 (11):111.
Parham, J. F. and Hutchison, J. H. 2003. A new eucryptodiran turtle from the Late Cretaceous of North America (Dinosaur Provincial Park, Alberta, Canada). Journal of Vertebrate Paleontology, 23:783798.
Parham, J. F. and Irmis, R. B. 2008. Caveats on the use of fossil calibrations for molecular dating. American Naturalist, 171:132136.
Parham, J. F. and Pyenson, N. D. 2010. New sea turtle from the Miocene of Peru and the iterative evolution of feeding ecomorphologies since the Cretaceous. Journal of Paleontology, 84:231247.
Peppe, D. J, McNulty, K. P., Cote, S. M., Harcourt-Smith, W. E. H., Dunsworth, H. M., and van Couvering, J. A. 2009. Stratigraphic interpretation of the Kulu Formation (early Miocene, Rusinga Island, Kenya) and its implication for primate evolution: Journal of Human Evolution, 56:447461.
Prothero, D. R. and Emry, R. J. 2004. The Chadronian, Orellan, and Whitneyan North American land mammal ages, p. 156168. InWoodburne, M. O.(ed.), Late Cretaceous and Cenozoic Mammals of North America. Columbia University Press, New York.
Pyron, R. A. 2010. A likelihood method for assessing molecular divergence time estimates and the placement of fossil calibrations. Systematic Biology, 59:185194.
Quintart, A. and Plisnier-Ladame, F. 1968. Eosphargis gigas (Owen, 1861). A propos d'une tortue geánte récemment exposée dans las salles de l'Institut Royal des Sciences Naturelles de Belgique. Les Naturalistes Belges, 49:406413.
Rabi, M., Joyce, W. G., and Wings, O. 2010. A review of the Mesozoic turtles of the Junggar Basin (Xinjiang, Northwest China) and the paleobiogeography of Jurassic to Early Cretaceous Asian testudinates. Palaeobiodiversity and Palaeoenvironments, 90:259273.
Rannala, B. and Yang, Z. 2007. Inferring speciation times under an episodic molecular clock. Systematic Biology, 56:453466.
Reisz, R. R. and Müller, J. 2004. The fossil record and molecular timescales: A paleontological perspective. Trends in Genetics, 20:237241.
Rhodin, A. G. J., van Dijk, P. P., and Parham, J. F. 2008. Turtles of the world: Annotated checklist of taxonomy and synonymy. Chelonian Research Monographs, 5:000.1000.38.
Riabinin, A. N. 1935. Remains of turtle from the late Cretaceous deposits of Kizylkum Desert. Trudy Paleozoologiceskogo Instituta, 4:6977. (In Russian)
Robinson, P., Gunnell, G. F., Walsh, S. L., Clyde, W. C., Storer, J. E., Stucky, R. K., Froehlich, D. J., Ferrusquia-Villaranca, I., and McKenna, M. C. 2004. Wasatchian through Duchesnean biochronology, p. 106155. InWoodburne, M. O.(ed.), Late Cretaceous and Cenozoic Mammals of North America. Columbia University Press, New York.
Romano, P. S., Oliveira, G. R., Azevedo, S. A. K., Kellner, A. W. A., and de Almeida Campos, D. 2013. New information about Pelomedusoides (Testudines: Pleurodira) from the Cretaceous of Brazil, p. 261275. InBrinkman, D. B., Holroyd, P. A., and Gardner, J. D.(eds.), Morphology and Evolution of Turtles. Springer.
Romer, A. 1956. Osteology of Reptiles. University of Chicago Press, Chicago, 772p.
Rougier, G. W., de la Fuente, M. S., and Arcucci, A. B. 1995. Late Triassic turtles from South America. Science, 268:855858.
Ruez, D. R. 2009. Framework for stratigraphic analysis of Pliocene fossiliferous deposits at Hagerman Fossil Beds National Monument, Idaho. Rocky Mountain Geology, 44:3370.
Sanderson, M. J. 1997. A nonparametric approach to estimating divergence times in the absence of rate constancy. Molecular Biology and Evolution, 14:12181231.
Scheyer, T. M. and Anquetin, J. 2008. Bone histology of the Middle Jurassic turtle shell remains from Kirtlington, Oxfordshire, England. Lethaia, 41:8596.
Scheyer, T. M., Sander, P. M., Joyce, W. G., Böhme, W., and Witzel, U. 2007. A plywood structure in the shell of fossil and living soft-shelled turtle (Trionychidae) and its evolutionary implications. Organisms Diversity and Evolution, 7:136144.
Seeley, H. G. 1892. On a new reptile from Welte Vreden (Beaufort West), Eunotosaurus africanus (Seeley). Quarterly Journal of the Geological Society, 48:583585.
Shaffer, H. B., Meylan, P., and McKnight, M. L. 1997. Tests of turtle phylogeny: Molecular, morphological, and paleontological approaches. Systematic Biology, 46:235268.
Shen, X.-X., Liang, D., Wen, J.-Z., and Zhang, P. 2011. Multiple genome alignments facilitate development of NPCL markers: A case study of tetrapod phylogeny focusing on the position of turtles. Molecular Biology and Evolution, 28:32373252.
Smith, A. B., Pisani, D., Mackenzie-Dodds, J. A., Stockley, B., Webster, B. L., and Littlewood, D. T. J. 2006. Testing the molecular clock: Molecular and paleontological estimates of divergence times in the Echinoidea (Echinodermata). Molecular Biology and Evolution, 23:18321851.
Smith, E. T. 2010. Early Cretaceous chelids from Lightning Ridge, New South Wales. Alcheringa, 34:375384.
Smith, M. E., Singer, B., Carroll, A. R. 2003. 40Ar/39Ar geochronology of the Eocene Green River Formation, Wyoming. Geological Society of America Bulletin, 115:549565.
Spinks, P. Q. and Shaffer, H. B. 2009. Conflicting mitochondrial and nuclear phylogenies for the widely disjunct Emys (Testudines: Emydidae) species complex, and what they tell us about biogeography and hybridization. Systematic Biology, 58:120.
Spinks, P. Q., Shaffer, H. B., McCord, W. P., and Iverson, J. B. 2004. Phylogenetic hypotheses for the turtle family Geoemydidae. Molecular Phylogenetics and Evolution, 32:164182.
Sterli, J. 2008. A new, nearly complete stem turtle from the Jurassic of South America with implications for turtle evolution. Biology Letters, 4:286289.
Sterli, J. 2010. Phylogenetic relationships among extinct and extant turtles: The position of Pleurodira and the effects of the fossils on rooting crown-group turtles. Contributions to Zoology, 79:93106.
Sterli, J. and de la Fuente, M. S. 2011. A new turtle from the La Colonia Formation (Campanian–Maastrichtian), Patagonia, Argentina, with remarks on the evolution of the vertebral column in turtles. Palaeontology, 54:6378.
Sterli, J., Pol, D., and Laurin, M. 2012. Incorporating phylogenetic uncertainty on phylogeny-based palaeontological dating and the timing of turtle diversification. Cladistics, 2012:114.
Sukhanov, V. B. 2000. Mesozoic turtles of middle and central Asia, p. 309367. InBenton, M. J., Shishkin, M. A., Unwin, D. M., and Kurochkin, E. N.(eds.), The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press, Cambridge.
Sukhanov, V. B. 2006. An archaic turtle, Heckerochelys romani gen. et sp. nov., from the Middle Jurassic of Moscow Region, Russia. Fossil Turtle Research, 1:112118.
Sukhanov, V. B., Danilov, I. G., and Syromyatnikova, E. V. 2008. The description and phylogenetic position of a new nanhsiungchelyid turtle from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica, 53:601614.
Swofford, D. L. 2002. PAUP∗. Phylogenetic Analysis Using Parsimony (∗and other methods). Version 4b10. Sinauer Associates, Sunderland, Massachusetts.
Syromyatnikova, E. V. and Danilov, I. G. 2009. New material and a revision of turtles of the genus Adocus (Adocidae) from the Late Cretaceous of Middle Asia and Kazakhstan. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 313:7494.
Taquet, P. 1976. Géologie et paléontologie du gisement de Gadoufoua (Aptien du Niger). Cahiers de Paléontologie. Centre National de la Recherche Scientifique (CNRS), Paris, 191p.
Thomas, H., Roger, J., Halawani, M., Memesh, A., Lebret, P., Bourdillon, C., Buffetaut, E., Cappetta, H., Cavelier, C., Dutheil, D., Tong, H., and Vaslet, D. 1999. Late Paleocene to early Eocene marine vertebrates from the uppermost Aruma Formation (northern Saudi Arabia): Implications for the K–T transition. Comptes Rendus l‘Académie des Sciences, Series IIA, 329:905919.
Thompson, J. D., Higgins, D. G., and Gibson, T. J. 1994. CLUSTAL W: Improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Ressearch, 22:46734680.
Thorne, J. L. and Kishino, H. 2005. Estimation of divergence times from molecular sequence data, p. 233256. InNielsen, R.(ed.), Statistical methods in molecular evolution. Springer-Verlag, New York.
Tong, H. and Hirayama, R. 2004. First Cretaceous dermochelyid turtle from Africa. Revue de Paléobiologie, 9:5559.
Tong, H. and Ouyang, H. 2012. A review of Sinaspideretes wimani Young and Chow, 1953, p. 44. InSymposium on Turtle Evolution 2012, Program and Abstracts. University of Tübingen, Tübingen.
Tong, H., Buffetaut, E., Thomas, H., Roger, J., Halawani, M., Memesh, A., and Lebret, P. 1999. A new dermochelyid turtle from the late Paleocene-early Eocene of Saudi Arabia. C Comptes Rendus l‘Académie des Sciences, Series IIA, 329:913919.
Tong, H., Claude, J., Suteethorn, V., Naksri, W., and Buffetaut, E. 2009 a. Turtle assemblages of the Khorat Group (Late Jurassic–Early Cretaceous) of NE Thailand and their palaeobiogeographical significance. Geological Society of London Special Publications, 315:141152.
Tong, H., Claude, J., Naksri, W., Suteethorn, V., Buffetaut, E., Khansubha, S., Wongko, K., and Yuangdetkla, P. 2009 b. Basilochelys macrobios n. gen. and n. sp., a large cryptodiran turtle from the Phu Kradung Formation (latest Jurassic–earliest Cretaceous) of the Khorat Plateau, NE Thailand. Geological Society of London Special Publications, 315:153173.
Vaillant, L. 1894. Essai sur la classification générale des chéloniens. Annales des Sciences Naturelles, 16:331345.
Valenzuela, N. and Adams, D. C. 2011. Chromosome number and sex determination coevolve in turtles. Evolution, 65:18081813.
Vargas-Ramíreza, M., Castaño-Morab, O. V., and Fritz, U. 2008. Molecular phylogeny and divergence times of ancient South American and Malagasy river turtles (Testudines: Pleurodira: Podocnemididae). Organisms Diversity and Evolution, 8:388398.
Vitek, N. S. 2011. Insights into the taxonomy and systematics of North American Eocene soft-shelled turtles from a well-preserved specimen. Bulletin of the Peabody Museum of Natural History, 52:189208.
Vitek, N. S. 2012. Giant fossil soft-shelled turtles of North America. Palaeontologia Electronica, 15 (13A):143.
Warnock, R. C. M., Yang, Z., and Donoghue, P. C. J. 2012. Exploring uncertainty in the calibration of the molecular clock. Biology Letters, 8:156159.
Weaver, W. G. Jr, and Rose, F. L. 1967. Systematics, evolution, and fossil record of the genus Chrysemys. Tulane Studies in Zoology, 14:6373.
Whetstone, K. N. 1978. A new genus of cryptodiran turtles (Testudinoidea, Chelydridae) from the Upper Cretaceous Hell Creek Formation on Montana. University of Kansas Science Bulletin, 51:539563.
Wieland, G. R. 1896. Archelon ischyros: A new gigantic cryptodire testudinate from the Fort Pierre Cretaceous of South Dakota. American Journal of Science, 2:399412.
Williams, E. E. 1950. Variation and selection in the cervical articulation of living turtles. Bulletin of the American Museum of Natural History, 94:509561.
Williams, E. 1954. A new Miocene species of Pelusios and the evolution of that genus. Breviora, 25:17.
Wing, S. L., Bown, T. M., and Obradovich, J. D. 1991. Early Eocene biotic and climatic change in interior western North America. Geology, 19:11891192.
Wong, R. A., Fong, J. J., and Papenfuss, T. J. 2010. Phylogeography of the African helmeted terrapin, Pelomedusa subrufa: Genetic structure, dispersal, and human introduction. Proceedings of the California Academy of Sciences, 61:575585.
Wood, R. C. 1973. A possible correlation between the ecology of living African pelomedusid turtles and their relative abundance in the fossil record. Copeia, 1973:627629.
Wood, R. C. 1976. Two new species of Chelus (Testudines: Pleurodira) from the late Tertiary of northern South America. Breviora, 435:126.
Wood, R. C., Johnson-Gove, J., Gaffney, E. S., and Maley, K. F. 1996. Evolution and phylogeny of leatherback turtles (Dermochelyidae), with descriptions of new fossil taxa. Chelonian Conservation and Biology, 2:266286.
Yang, Z. 2006. Computational molecular evolution. Oxford University Press, Oxford, 376p.
Yang, Z. 2007. PAML 4: Phylogenetic analysis by maximum likelihood. Molecular Biology and Evolution, 24:15861591.
Yang, Z. and Rannala, B. 2006. Bayesian estimation of species divergence times under a molecular clock using multiple fossil calibrations with soft bounds. Molecular Biology and Evolution, 23:212226.
Yeh, H.-K. 1963. Fossil turtles of China. Paleontologia Sinica, 150:1112.
Yeh, H.-K. 1965. New materials of fossil turtles of Inner Mongolia. Vertebrata PalAsiatica, 9:4770.
Yeh, H.-K. 1981. New fossil Cuora (Emydidae) from the early Pliocene of Yunnan. Vertebrata PalAsiatica, 19:239245.
Young, C. C. and Chow, M. C. 1953. New fossil reptiles from Szechuan, China. Acta Scientia Sinica, 2:216229.
Zangerl, R. 1960. The vertebrate fauna of the Selma formation of Alabama. Part V. An advanced cheloniid sea turtle. Fieldiana, Geology Memoirs, 3:279312.
Zug, G. 1969. Fossil chelonians, Chrysemys and Clemmys, from the upper Pliocene of Idaho. Great Basin Naturalist, 24:8287.


Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed