The Vertebrates References

Chelonii: References

Abbreviated Dendrogram
Anapsida ? │ └─○Chelonii ├─Odontochelys └─┬─Proganochelys └─┬─Proterochersis └─┬─Pleurodira ├─Paracryptodira └─○Eucryptodira ├─Protostegidae └─┬─Angolachelonia └─┬─Xinjiangchelyidae ╘═╤═Sinemydidae ├─Trionychoidea └─┬─┬─Chelonioidea │ ├─Chelydridae │ └─Kinosternoidea └─Testudinoidea

Chelonian origins
Derived Stem Testudines


Abel, Othenio. 1919. Die Stämme der Wirbeltlere. Verinigung Wissenschaftlicher Verleger. Berlin und Leipzig.

John Acorn, 2007. Deep Alberta: Fossil Facts and Dinosaur Digs. Edmonton: University of Alberta Press

Allendorf, F. W., R. F. Leary, P. Spruell & J. K. Wenburg. 2001. The problems with hybrids: setting conservation guidelines. Trends in Ecology and Evolution 16 (11): 613-622.
Tortoise Resurrection

Anon. January 2009, St. Catherines Island Sea Turtle Conservation Program. web page

Anquetin, J. (2009) A new stem turtle from the Middle Jurassic of the Isle of Skye, Scotland, and a reassessment of basal turtle relationships. Doctoral thesis, UCL (University College London). pdf, mirror.
Chelonian phylogenetics and systematics, Baenidae, Baenodda, Chelydridae, Eileanchelys, Platysternon, Protostegidae

Jérémy Anquetin (2012). "Reassessment of the phylogenetic interrelationships of basal turtles (Testudinata)". Journal of Systematic Palaeontology 10 (1): 3–45. doi:10.1080/14772019.2011.558928.

Anquetin, J., Barrett, P. M., Jones, M. E. H., Moore-Fay, S. & 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 of London: Biological series: Vol. 276, pp. 879-886 [doi: 10.1098/rspb.2008.1429] pdf.
Chelonian phylogenetics and systematics, Eileanchelys

Barley, A.J., Spinks, P.Q., Thomson, R.C., Shaffer, H.B.., 2010, Fourteen nuclear genes provide phylogenetic resolution for difficult nodes in the turtle tree of life. Molecular Phylogenetics and Evolution 55 (2010) 1189–1194 pdf
Morphology or Molecules?

Baur, George (1890). "On the Classification of the Testudinata". The American Naturalist 24 (282): 530–536. doi:10.1086/275138. JSTOR 2450882.

Baur, G. 1891. Notes on some little known American fossil tortoises. Proc. Acad. Nat. Sci. Philadelphia, vol. 43, pp. 411-430.
Chelonian phylogenetics and systematics

Benton, M. J., Vertebrate Paleontology, 2nd ed. 2000, 3rd ed. 2004, Blackwell Science Ltd,
Australochelys, Casichelydia

Bhullar, B-A.S. and Bever, G.S. 2009. An Archosaur-Like Laterosphenoid in Early Turtles (Reptilia: Pantestudines). Breviora. vol.518:1-10. pdf
Mystery of Chelonian Origins

Gale A. Bishop. 2007. Handbook for SeaTurtle Interns especially on St. Catherines Island. part 5, Natural history of sea turtles pdf
Allopleuroninae, Dermochelyinae, Syllomus

Böhme, W. 2005. Handbuch der Reptilien und Amphibien Europas. Band 3/IIIB: Schildkröten (Testudines) II (Cheloniidae, Dermochelyidae, Fossile Schildkröten Europas) (Handbook of reptiles and amphibians of Europe, Volume 3/IIIB: Turtles (Testudines) II (Cheloniidae, Dermochelyidae, Fossil turtles of Europe)). Edited by Uwe Fritz. Illustrations: Igor G. Danilov and Jiri Moravec. Wiebeisheim: AULA-Verlag pdf (in German)

Borsuk-Bialynicka, M, & Evans, S.E. (2009). A long-necked archosauromorph from the Early Triassic of Poland. Paleontologica Polonica 65: 203–234. pdf
Mystery of Chelonian Origins

Boule, Marcellin. Priviteau, Jean. Les Fossiles. Éléments De Paléontologie. Libraries de L'académie Médecine. Paris, 1935. p. 433

Bour, R & Dubois, A. 1986, Nomenclature ordinale et familiale des tortues (Reptilia). Note complémentaire. Bulletin Mensuel de la Société Linnéenne de Lyon 55:87-90..
Classification, Meiolanoidea

Brian W. Bowen, 2003, What Is a Loggerhead Turtle? - The Genetic Perspective
Caretta, Cheloniinae

Donald B. Brinkman, 2009, A sea turtle skull (Cheloniidae: Carettini) from the Lower Miocene Nye formation of Oregon, U. S. A. Paludicola 7(2): 39-46 May 2009 pdf
Procolpochelys, Syllomus

Brochu, C.A. 2001. Progress and future directions in archosaur phylogenetics J. Paleont., 75(6), 2001, pp. 1185–1201 ArchosauriaBROCHU01.pdf">pdf
Mystery of Chelonian Origins

Edwin A. Cadena, Daniel T. Ksepka and Mark A. Norell (2013) New material of Mongolemys elegans Khosatzky and Mlynarski, 1971 (Testudines: Lindholmemydidae), from the Late Cretaceous of Mongolia with Comments on Bone Histology and Phylogeny. American Museum Novitates 3766 :1-28. 2013 doi: pdf

Carroll, R. L., 1988: Vertebrate paleontology and evolution. W. H. Freeman and company, New York,
Chelonian phylogenetics and systematics

Cherepanov GO. 1997. The origin of the bony shell of turtles as a unique evolutionary model in reptiles. Russ J Herpet. 4:155–162
Mystery of Chelonian Origins

Ckhikvadze, Viacheslav M. 1971. First find of a Tertiary turtle of the family Platysternidae. Paleontal. Zhur., 1971, no. 4, pp. 137-139.

Colbert, E.H., 1969 Evolution of the Vertebrates, John Wiley & Sons, 2nd ed.
Chelonian phylogenetics and systematics

Collins, J. 1972. The chelonian Rhinochelys Seeley from the Upper Cretaceous of England and France. Palaeontolgy 13: 355 -378

Danilov, I.G.; Averianov, A.O; & Yarkov, A.A. (2010). Itilochelys rasstrigin gen. et sp. nov., a new hard-shelled sea turtle (Cheloniidae sensu lato) from the Lower Palaeocene of Volgograd Province, Russia. Proceedings of the Zoological Institute RSA 314 (1): 24–41.

Danilov I.G., Claude J., Sukhanov V.B. A redescription of Elkemys australis (Yeh, 1974), a poorly known basal testudinoid turtle from the Paleocene of China. Proc. Zool. Inst. RAS. 2012. V. 316. ? 3. P. 223-238. pdf

Danilov, I. G. & Parham, J. F. 2006. A redescription of Plesiochelys tatsuensis from the Late Jurassic of China, with comments on the antiquity of the Extant Turtles Journal of Vertebrate Paleontology 28(2):306-318.
Morphology or Molecules?, Pantrionychia

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: 306–318
Morphology or Molecules? - abstract and text figures

deBraga, M & O Rieppel (1997) Reptile phylogeny and the interrelationships of turtles, Zool. J. Linnean Soc. 120: 281-354.
Mystery of Chelonian Origins

Dryden, L. S. 1988. Paraphyly of the Cryptodira and phylogenetic systematics of turtles. Unpublished Master's thesis, University of Kansas, Lawrence, KS.
Chelonian phylogenetics and systematics

Alain Dubois & Roger Bour, 2010 The distinction between family-series and class-series nomina in zoological nomenclature, with emphasis on the nomina created by Batsch (1788, 1789) and on the higher nomenclature of turtles, Bonn zoological Bulletin 57 (2): 149–171

Ehret, Dana Joseph 2004 "Skeletochronology as a method of aging Oligocene Gopherus laticuneus and Stylemys nebrascensis, using Gopherus polyphemus as a modern analog" Thesis, University of Florida. pdf

Fong JJ, Brown JM, Fujita MK, Boussau B (2012) A Phylogenomic Approach to Vertebrate Phylogeny Supports a Turtle-Archosaur Affinity and a Possible Paraphyletic Lissamphibia. PLoS ONE 7(11): e48990. doi:10.1371/journal.pone.0048990 online

Fossil Turtle Newsletter January, 2004. Compiled by Walter Joyce and James Parham pdf

Fujita, M.K.; Tag N. Engstrom, David E. Starkey and H. Bradley Shaffer (2004). Turtle phylogeny: insights from a novel nuclear intron. Molecular Phylogenetics and Evolution 31 (3): 1031-1040

Benjamin P. Kear, and Michael S. Y. Lee, 2006, A primitive protostegid from Australia and early sea turtle evolution Biol. Lett. 2, 116-119 doi:10.1098/rsbl.2005.0406 pdf
Bouliachelys, Toxochelyidae

Krenz, James G.; Gavin J.P. Naylor; H. Bradley Shaffer and Fredric J. Janzen (2005) Molecular phylogenetics and evolution of turtles. Molecular Phylogenetics and Evolution, Volume 37 (1):178-191.
Morphology or Molecules?

R. Kohler (1995): A new species of the fossil turtle Psephophorus (Order Testudines) from the Eocene of the South Island, New Zealand, Journal of the Royal Society of New Zealand, 25:3, 371-384 pdf

Gadagkar, S.R., Rosenberg, M.S., & Kumar, S. (2005). Inferring Species Phylogenies From Multiple Genes: Concatenated Sequence Tree Versus Consensus Gene Tree. Journal of Experimental Zoology (Mol Dev Evol) 304B:64–74 pdf
Morphology or Molecules?

Gaffney, E.S. 1972. The systematics of the North American family Baenidae (Reptilia Crytodira). Bull. Amer. Nat. Hist. 147(5):241-320.

Gaffney, E.S. (1975): A phylogeny and classification of higher categories of turtles. Bulletin of the American Museum of Natural History 155(5): 387-436. abstract, pdf
Testudinata/Testudines, Chelonian phylogenetics and systematics, Paracryptodira

Gaffney, Eugene S. (1975b). Phylogeny of the chelydrid turtles. Fieldiana, Geology (Chicago: Field Museum of Natural History) 33 (9): 166 (16). OCLC 1991971. online
Chelydrinae, Macrocephalochelys, Platysternidae, Platysternon

Gaffney, E.S. 1979. Comparative cranial morphology of Recent and fossil turtles. Bull. Amer. Mus. Natur. Hist. 164; 65-376

Gaffney, E.S (1984) Historical Analysis of Theories of Chelonian Relationship. Systematic Zoology 33. 3. 283-301.
Chelonian phylogenetics and systematics

Gaffney, E.S (1996) The postcranial morphology of Meiolania platyceps and a review of of the Meiolaniidae. Bulletin of the American Museum of Natural History ; no. 229 abstract and full pdf .
Chelydridae, Meiolania, Meiolaniidae, Ninjemys, Niolamia

Gaffney, E. S., Balouet, J. C., & Broin, F. (1984). New occurrences of extinct meiolaniid turtles in New Caledonia. American Museum novitates ; no. 2800 pdf

Gaffney and Kitching 1994, The most ancient African turtle, Nature 369: 55-58, [doi:10.1038/369055a0]

Gaffney and Kitching 1995, The Morphology and Relationships of Australochelys, an Early Jurassic Turtle from South Africa, Am. Mus. Novitates 3130: 29 pp.

Gaffney, E. S., Kool, L., Brinkman, D. B., Rich, T. H. V., & Rich, Pat Vickers. 1998. Otwayemys, a new cryptodiran turtle from the early Cretaceous of Australia. American Museum novitates ; no. 3233 abstract & pdf

Gaffney, ES & PA Meylan (1988), A phylogeny of turtles, in MJ Benton (ed.), The Phylogeny and Classification of the Tetrapods, 1: 157-219.
Pancheloniidae, Platysternon, Podocnemoidae, Testudinoidae, Trionychoidea, Xerobatinae

Gaffney E.S., Meylan, P. A., & Wyssz, A.R. 1991. A computer assisted analysis of the relationships of the higher categories of turtles. Cladistics 1991 7: 313-335 pdf
Chelonian phylogenetics and systematics, Chelydridae

Gauthier, J, AG Kluge & T Rowe (1988), The early evolution of the Amniota, in MJ Benton (ed.) The Phylogeny and Classification of the Tetrapods, Volume 1: Amphibians, Reptiles, Birds. Clarendon Press, pp. 103-155.

Gauthier, J., D. C. Cannatella, K. De Queiroz, A. G. Kluge, & T. Rowe. 1989. Tetrapod phylogeny. In B. Fernholm, K. Bremer, and H. Jornvall (eds.) The Hierarchy of Life: 337-353. New York: Elsevier Science Publishers B. V. (Biomedical Division). pdf
Chelonian phylogenetics and systematics

J. Gerlach 2001 Tortoise phylogeny and the 'Geochelone' problem. Phelsuma, Volume 9 (supplement A) pdf
Megalochelyini, Xerobatinae

Gingerich, P. D., M. Haq, I. S. Zalmout, I. H. Khan, and M. S. Malkani. 2001. Origin of whales from early artiodactyls: hands and feet of Eocene Protocetidae from Pakistan. Science, 293: 2239-2242. pdf.
Mystery of Chelonian Origins

Jean-Michel Guillon, Loreleï Guéry, Vincent Hulin, & Marc Girondot, 2012, A large phylogeny of turtles (Testudines) using molecular data, Contributions to Zoology, 81 (3) 147-158
Megalochelyini, Xerobatinae

S. H. Haughton and A. S. Brink. 1954. A bibliographical list of Reptilia from the Karroo Beds of South Africa. Palaeontologia Africana 2:1-187.
Chelonian phylogenetics and systematics

Hay, 0. P. 1905. On the group of fossil turtles known as the Amphichelydia, with remarks on the origin and relationships of the suborders, superfamilies and families of Testudines. Bull. Amer. Mus.Nat. Hist., vol. 21, art. 9, pp. 137-175.
Chelonian phylogenetics and systematics

Hay, O.P. 1908. The fossil turtles of North America. Publications of the Carnegie Institution 75:1-568.
Chelonian phylogenetics and systematics, Archelon, Platysternon

Hedges, S. Blair, and Poling, Laura L. 1999, A Molecular Phylogeny of Reptiles. Science, Vol. 283, No.5404, pp.998-1001. pdf.
Mystery of Chelonian Origins

Hill, Robert V.. 2005. Integration of Morphological Data Sets for Phylogenetic Analysis of Amniota: The Importance of Integumentary Characters and Increased Taxonomic Sampling. Systematic Biology 54(4):530-547. abstract pdf.
Mystery of Chelonian Origins

Hirayama, R. 1984. Cladistic analysis of batagurine turtles (Batagurinae: Emydidae: Testudinoides); a preliminary result. Stud. Geol. Salamanticesia. 1; 141-157

Hirayama, Ren (1994). "Phylogenetic systematics of chelonioid sea turtles". Island Arc 3 (4): 270–284. doi:10.1111/j.1440-1738.1994.tb00116.x. Retrieved 2007-09-04.
Protostegidae, Rhinochelys

Hirayama, R. 1998. "Oldest known sea turtle". Nature 392 (6677): 705–708. doi:10.1038/33669
Protostegidae, Santanachelys

Hirayama, R. 2012 Review of fossil chelonioid sea turtles (Class Reptilia: Order Testudines: Chelonioidea) from Japan, Japan Geoscience Union Meeting 2012 pdf

Ren Hirayama, Donald B. Brinkman, and Igor G. Danilov, 2000, Distribution and Biogeography of Non-marine Cretaceous Turtles, Russian Journal of Herpetology Vol. 7, No. 3, 2000, pp. 181-198, pdf,
Plesiochelyidae, Pleurosternidae

Hirayama, R., Sakurai, K., Chitoku, T., Kawakami, G., and Kito, N. 2001. Anomalochelys angulata, an unusual land turtle of family Nanhsiungchelyidae (Superfamily Trionychoidea; Order Testudines) from the Upper Cretaceous of Hokkaido, North Japan. Russian Journal of Herpetology 8: 127-138.

Honegger, R. E. 1981. List of amphibians and reptiles either known or thought to have become extinct since 1600. Biological Conservation 19: 141-158.
Tortoise Resurrection

Hooks, III, G. E., 1998: Systematic revision of the Protostegidae, with a redescription of Calcarichelys gemma Zangerl, 1953. Journal of Vertebrate Paleontology: Vol. 18, #1, pp. 85-98 abstract
Protostegidae, Protosteginae

Hugall, A. F., Foster, R. & Lee, M. S. Y. 2007 Calibration choice, rate smoothing, and the pattern of tetrapod diversification according to the long nuclear gene RAG-1. Syst. Biol. 56, 543–563.
Mystery of Chelonian Origins

Hutchison, J. H. 1992. Western North American reptile and amphibian record across the eocene/Oligocene boundary and its climatic implications. Prothero, D. R. & Berggren, W. A. (eds.), Eocene-Oligocene Climatic and Biotic Evolution 451-463.

Hutchison, J.H., 1996. Testudines. Cambridge University Press, Cambridge

Iverson J.B., Brown R.M., Akre T.M., Near T.J., Le M., Thomson R.C., Starkey D.E. 2007. In search of the tree of life of turtles.In: Shaffer H.B., FitzSimmons N.N., Georges A., Rhodin A.G.J., editors. Defining Turtle Diversity: Proceedings of a Workshop on Genetics, Ethics, and Taxonomy of Freshwater Turtles and Tortoises. Cambridge (MA). Lunenburg (MA): Chelonian Research Foundation. p. 85¨C106 (Chelonian research monographs).
Chelonian phylogenetics and systematics, Geoemydidae, Natator

Jackson, A.A.; 2008: Bedrock geology UK south. An explanation of the bedrock geology map of England and Wales - 1:625,000 edition, Keyworth, Nottingham, British Geological Survey,

Jamniczky, HA & Russell, AP (2007). Reappraisal of patterns of nonmarine cryptodiran turtle carotid circulation: evidence from osteological correlates and soft tissues. J Morphol. 2007 July; 268(7):571-87. pdf

Janke, A. D., Erpenbeck, D., Nilsson, M., and Arnason, U. 2001. The mitochondrial genomes of the iguana (Iguana iguana) and the caiman (Caiman crocodylus): implications for amniote phylogeny. Proc. R. Soc.B 268: 623-631.
Mystery of Chelonian Origins

Jehle, M. 2006, Paleocene mammals of the world - Turtles, Business as Usual

Jones MEH, Werneburg I, Curtis N, Penrose R, O'Higgins P, et al. (2012) The Head and Neck Anatomy of Sea Turtles (Cryptodira: Chelonioidea) and Skull Shape in Testudines. PLoS ONE 7(11): e47852. doi:10.1371/journal.pone.0047852 online
Crown Chelonioidea

Joyce W.G. (2000) The first complete skeleton of Solnhofia parsonsi (Cryptodira, Eurysternidae) from the Upper Jurassic of Germany and its taxonomic implications J. Paleont., 74(4), 2000, pp. 684-700

Joyce W.G. (2007) Phylogenetic relationships of Mesozoic turtles. Bull. Peabody Mus Nat Hist 48(1):3–102
Chelonian phylogenetics and systematics, Baenidae, "Baenini",, Chelydridae, Pancheloniidae, Platysternon, Protostegidae

Joyce, WG, & Gauthier, JA, (2004), Palaeoecology of Triassic stem turtles sheds new light on turtle origins, Proc. R. Soc. Lond. B (2004) vol. 271 no. 1534, 1–5, © 2003 The Royal Society DOI 10.1098/rspb.2003.2523. pdf

Joyce, W.G., Jenkins, F.A., Jr., and Rowe, T. 2006. The presence of cleithra in the basal turtle Kayentachelys aprix; pp. 93-103 in Danilov, I.G. and Parham, J.F. (eds.), Fossil Turtle Research, Volume 1. Russian Journal of Herpetology 13 (suppl.). pdf
Mystery of Chelonian Origins

Joyce, W. G., Lucas, S. G., Scheyer, T. M., Heckert, A. B., and Hunt, A. P., 2008. A thin-shelled reptile from the Late Triassic of North America and the origin of the turtle shell Proceedings of the Royal Society B Published online (FirstCite). DOI: 10.1098/rspb.2008.1196. pdf.
Chelonian phylogenetics and systematics, Chinlechelys

Joyce, W. G. and Lyson, T. R., 2010, A neglected lineage of North American turtles fills a major gap in the fossil record. Palaeontology, 53: 241–248. doi: 10.1111/j.1475-4983.2009.00929.x abstract.

Joyce, W.G., Parham, J.F., & Gauthier, J.A., (2004), Developing a protocol for the conversion of rank-based taxon names to phylogenetically defined clade names, as exemplified by turtlesJ. Paleont., 78(5), 2004, pp. 989–1013. 0022-3360/04/0078-989$03.00 pdf.
Chelonian phylogenetics and systematics, Pancheloniidae, Pancheloniodea

Walter G. Joyce, Ariel Revan, Tyler R. Lyson and Igor G. Danilov (2009). Two New Plastomenine Softshell Turtles from the Paleocene of Montana and Wyoming. Bulletin of the Peabody Museum of Natural History 50 (2): 307–325. pdf

H. F. Kaddumi (2006). "A new genus and species of gigantic marine turtles (Chelonioidea: Cheloniidae) from the Maastrichtian of the Harrana Fauna-Jordan". Vertebrate Paleontology (journal) 3 (1): 1–14.

H.-V. Karl - Zur Taxonomie der känozoischen Weichschildkröten Österreichs und Deutschlands (Trionychidae: Trionychinae).- Mitt. Geol. Und Paläont. Landesmuseum Joanneum, 56: 273- 328; Graz (1998).
Trionyx messelianus

Karl, H.-V. (2002): Übersicht über die fossilen marinen Schildkrötenfamilien Zentraleuropas (Reptilia, Testudines). Mauritiana (Altenburg), 18 (2): 171-202.

Karl, H.-V. (2005): The homology of supramarginals in turtles (Reptilia:Chelonii). Studia Geologica Salmanticensia, 41, pp. 63-75.
Mystery of Chelonian Origins

Hans-Volker Karl & Bent E. K. Lindow, 2009, First evidence of a Late Cretaceous marine turtle (Testudines: Chelonioidea) from Denmark Studia Geologica Salmanticensia, 45 (2): pp. 175-180

S. A. Karl, B. W. Bowen, and J. C. Avise,1995, Hybridization among the ancient mariners: Identification and characterization of marine turtle hybrids with molecular genetic assays. Journal of Heredity 86:262-268.pdf

Killian JK, Buckley TR, Stewart N, Munday BL, Jirtle RL. 2001. Marsupials and Eutherians reunited: genetic evidence for the Theria hypothesis of mammalian evolution. Mamm. Genome 12:513–17 pdf
Mystery of Chelonian Origins

Kordikova, E.G. Comparative morphology of the palate dentition in Proganochelys quenstedti Baur 1887 from the Upper Triassic of Germany and chelonian ancestry. - N. Jb. Geol. Paläont. Abh., 225, 2: 195-249.

Kullberg, M., Hallström, B. M., Arnason, U. and Janke, A. (2008), Phylogenetic analysis of 1.5 Mbp and platypus EST data refute the Marsupionta hypothesis and unequivocally support Monotremata as sister group to Marsupialia/Placentalia. Zoologica Scripta, 37: 115–127. doi: 10.1111/j.1463-6409.2007.00319.x
Mystery of Chelonian Origins

Kumazawa, Y. and Nishida, M. 1999. Complete Mitochondrial DNA Sequences of the Green Turtle and Blue-Tailed Mole Skink: Statistical Evidence for Archosaurian Affinity of Turtles. Mol. Biol. Evol. vol.16(6):784-792
Mystery of Chelonian Origins

Lambe, L. M. (1902). "New genera and species from the Belly River Series (Mid-Cretaceous)". Contributions to Canadian Palaeontology 3: 23–81.

Minh Le, Christopher J. Raxworthy, William P. McCord, & Lisa Mertz (2006), A molecular phylogeny of tortoises (Testudines: Testudinidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 450 (2):
Megalochelyini, Testudinidae, Testudininae, Testudinini

Lee, M.S.Y. 1993. The Origin of the Turtle Body Plan: Bridging a Famous Morphological Gap. Science. vol.261: 1716 1720.
Mystery of Chelonian Origins

Lee, MSY (1996), Correlated progression and the origin of turtles. Nature 379: 812-815.

Lee, M.S.Y. 1997. Pareiasaur Phylogeny and the Origin of Turtles. Zoo. J. Linn. Soc. vol.120:197-280
Mystery of Chelonian Origins

Lee, M. S. Y. 1997a. Reptile relationships turn turtle. Nature, 389:245–246.
Mystery of Chelonian Origins

Lee, M. S. Y. 2001. Molecules, morphology, and the monophyly of diapsid reptiles. Contrib. Zool. 70: 121–138.
Mystery of Chelonian Origins

Li, Chun; Xiao-Chun Wu, Olivier Rieppel, Li-Ting Wang & Li-Jun Zhao (2008). "An ancestral turtle from the Late Triassic of southwestern China". Nature 456: 497-501. doi:10.1038/nature07533.
Odontochelys, Chelonian phylogenetics and systematics, Mystery of Chelonian Origins

Lipka, T. R., F. Therrien, D. B. Weishampel, H. A. Jamniczky, W.G. Joyce, M. W. Colbert, AND D. B. Brinkman. 2006. A new turtle from the Arundel Clay facies (Potomac Formation, Early Cretaceous) of Maryland, U.S.A. Journal of Vertebrate Paleontology, 20:300-307.
Arundelemys, Paracryptodira

Liu, L., Pearl, D.K., 2007. Species trees from gene trees: reconstructing Bayesian posterior distributions of a species phylogeny using estimated gene tree distributions. Syst. Biol. 56 (3), 504–514.
Morphology or Molecules?

Lucas, S.G. 2001, Chinese Fossil Vertebrates, Columbia University Press, New York
Chengyuchelys, Nanhsiungchelys

Lynch, S. C., and J. F. Parham. 2003. The first report of hard-shelled sea turtles (Cheloniidae sensu lato) from the Miocene of California, including a new species (Euclastes hutchisoni) with unusually plesiomorphic characters. PaleoBios 23:21-25.

Lyson, T.R., Bever, G.S., Bhullar, B-A.S.,Joyce, G.J., Gauthier, J.A. 2010. Transitional fossils and the origin of turtles. Biol. Lett. doi:10.1098/rsbl.2010.0371 pdf
Mystery of Chelonian Origins

Lyson, T. & Gilbert, S.F. (2009). Turtles all the way down: loggerheads at the root of the chelonian tree. Evolution & Development 11:2, 133-135 (2009) DOI: 10.1111/j.1525-142X.2009.00325.x pdf
Mystery of Chelonian Origins

Tyler R. Lyson and Walter G. Joyce (2009). "A Revision of Plesiobaena (Testudines: Baenidae) and an Assessment of Baenid Ecology Across the K/T Boundary". Journal of Paleontology 83 (6): 833–853. doi:10.1666/09-035.1.
Baenidae, Plesiobaena

Tyler R. Lyson and Walter G. Joyce (2011). Cranial anatomy and phylogenetic placement of the enigmatic turtle Compsemys victa Leidy, 1856 Journal of Paleontology, 85(4), 2011, p. 789–801

Marmi, J., Vila, B., and Galobart, A. (2009) Solemys (Chelonii, Solemydidae) remains from the Maastrichtian of Pyrenees: evidence of a semi-aquatic lifestyle: Cretaceous Research, v. 30, p. 1307-1312.

Octávio Mateus; Louis Jacobs; Michael Polcyn; Anne S. Schulp; Diana Vineyard; André Buta Neto; and Miguel Telles Antunes (2009). "The oldest African eucryptodiran turtle from the Cretaceous of Angola". Acta Palaeontologica Polonica 54 (4): 581–588. doi:10.4202/app.2008.0063

Alan Riverstone McCulloch 1908 A new genus and species of turtle from north Australia. Records of the Australian Museum, vol. 7, n. 2, p. 126-128

Merck, J. W. 1997. A phylogenetic analysis of the euryapsid reptiles. Unpublished Ph.D. dissertation, University of Texas at Austin, 785 pp.
Mystery of Chelonian Origins

Meyer, A., and Zardoya, R. 2003. Recent advances in the (molecular) phylogeny of vertebrates. Ann. Rev. Ecol. Syst. 34: 311-338. pdf
Mystery of Chelonian Origins

Meylan, PA (1996), Skeletal morphology and relationships of the early Cretaceous side-necked turtle, Araripemys barretoi (Testudines: Pelomedusoides: Araripemyidae), from the Santana formation of Brazil. J. Vert. Paleontol. 16: 20-33.

Meylan, P. A. 2001. Testudines. Turtles, tortoises and terrapins. Version 31 May 2001. in The Tree of Life Web Project,
Chelonian phylogenetics and systematics

Meylan, P. A. & Gaffney, E. S. 1989 The skeletal morphology of the Cretaceous cryptodiran turtle, Adocus, and the relationships of the Trionychoidea. Am. Mus. Novit. 2941, 1-60.

Meylan, Peter A. & Ren Hirayama (2000). "The Paleontology and Phylogenetics of "Sea Turtles"". Proceedings of the 19th Annual Symposium on Sea Turtle Conservation and Biology 19: 104. NOAA Technical Memorandum NMFS-SEFSC-443.

Meylan, P. A., R. T. J. Moody, C. A. Walker, and S. D. Chapman. 2000. Sandownia harrisi, a highly derived trionychoid turtle (Testudines: Cryptodira) from the Early Cretaceous of the Isle of Wight, England. Journal of Vertebrate Paleontology 20:522–532. pdf

Meylan, P. A. & Sterrer, W. (2000). Herperotestudo (Testudines: Testudinidae) from the Pleistocene of Bermuda, with comments on the phylogenetic position of the genus. Zoological Journal of the Linnean Society 128 (1): 51–76. doi:10.1111/j.1096-3642.2000.tb00649.x pdf
Hesperotestudo, Stylemys, Xerobatinae

Milner, Andrew R. 2004. The Turtles of the Purbeck Limestone of Dorset, Southern England. Palaeontology. Vol. 47. part 6. pp. 1446-1447

Mlynarski, M. 1976: Testudines. Handbuch der Paläoherpetologie/Handbook of Paleoherpetology, Part 7, Gustav Fischer Verlag
Testudinidae, Toxochelys

Müller, J. 2003. Early loss and multiple return of the lower temporal arcade in diapsid reptiles. Naturwissenschaften 90, 473-476.
Mystery of Chelonian Origins

Nagashima, H., Sugahara, F., Takechi, M., Ericsson, R., Kawashima-Ohya, Y., Narita, Y., & Kuratani, S. (2009). Evolution of the Turtle Body Plan by the Folding and Creation of New Muscle Connections. Science, 325 (5937), 193-196 DOI: 10.1126/science.1173826

Eugenia Naro-Maciel, Minh Lea, Nancy N. FitzSimmons, George Amato, 2008, Evolutionary relationships of marine turtles: A molecular phylogeny based on nuclear and mitochondrial genes, Molecular Phylogenetics and Evolution 49 (2008) 659–662 pdf

Neaverson, E. 1955. Stratigraphical Palaeontology: a Study of Ancient Life-Provences. Second Edition. Oxford. Clarendon Press.

Elizabeth L. Nicholls, 1992, Note on the occurrence of the marine turtle Desmatochelys (Reptilia: Chelonioidea) from the Upper Cretaceous of Vancouver Island, Canadian Journal of Earth Sciences, 29(2): 377-380, 10.1139/e92-033

Nielsen, E., (l959), Eocene Turtles from Denmark Medd. fra Dansk Geol. Forening. Kobenhavn. Bd. 14 pdf

Owen, Richard. Report on British Fossil Reptilia. Report of the British Association for the Advancement of Science. P. p43- 126. 1841

Owen, R. 1853, A Monograph of the Fossil Chelonian Reptiles of the Wealden Clays and Purbeck Limestones. Palaeontographical Society, Vol. VII, 1853

Parham, J.F. 2005. A Reassessment of the referral of sea turtle skulls to the genus Osteopygis (Late Cretaceous, New Jersey, USA) Journal of Vertebrate Paleontology 25(1):71–77, March 2005 pdf

Parham, J.F. & Hutchison, J.H. 2003. A new eucryptodiran turtle from the Late Cretaceous of North America (Dinosaur Provincial Park, Alberta, Canada). J. Vertebr. Paleontol. 23 (4): 783–798. pdf

Parham, J.F., & Irmis, R.B. (2008). Caveats on the Use of Fossil Calibrations for Molecular Dating: A Comment on Near et al. Am. Nat. 2008. Vol. 171, pp. 132–136. pdf
Morphology or Molecules?

James F. Parham; Nicholas D. Pyenson (2010). "New Sea Turtle from the Miocene of Peru and the Iterative Evolution of Feeding Ecomorphologies since the Cretaceous". Journal of Paleontology 84 (2): 231–247. doi:10.1666/09-077R.1. pdf
Caretta, Euclastes, Mexichelys, Pancheloniidae, Syllomus

Paul, G.S., and Leahy, G.D. (1994). Terramegathermy in the time of the titans: Restoring the metabolics of colossal dinosaurs. Paleontol. Soc. Spec. Publ. 7, 177-198

Pérez-García, A. 2012. The European Early Cretaceous cryptodiran turtle Chitracephalus dumonii and the diversity of a poorly known lineage of turtles. Acta Palaeontologica Polonica 57 (3): 575–588.
Chelonioidea, Chitracephalus, MesodermochelysPanchelonioidea, Toxochelyidae

Pérez-García, A., Gasulla, J.M., and Ortega, F. 2013. A new species of Brodiechelys Testudines, Pan-Cryptodira) from the Early Cretaceous of Spain: Systematic and palaeobiogeographic implications. Acta Palaeontologica Polonica 5X (X): xxx-xxx.
Chengyuchelys, Xinjiangchelyidae

Adán Pérez-García & Xabier Murelaga (2012): Larachelus morla, gen. et sp. nov., a new member of the little-known European Early Cretaceous record of stem cryptodiran turtles, Journal of Vertebrate Paleontology, 32:6, 1293-1302

A. Pérez García, F Ortega & F Escaso. A small pleurosternid turtle from the Upper Jurassic of Santa Rita (Torres Vedras, Portugal): Juvenile or new form? 7th Annual EAVP Meeting 2009. 56

Pidoplichko, Ivan G. and B. I. Tarashchuk, 1960. New genus of large-headed (macrocephalous) turtle from the Pontion beds in the environs of Odessa. [In Ukranian with Russian summary.] Zbirn. Prats. Zool. Mus. Akad. URSR, no. 29, pp. 105-110.

Poulakakis, N., S. Glaberman, M. Russello, L. B. Beheregaray, C. Ciofi, J. R. Powell & A. Caccone. 2008. Historical DNA analysis reveals living descendants of an extinct species of Galápagos tortoise. Proceedings of the National Academy of Sciences of the USA 105 (40): 15464-15469.
Tortoise Resurrection

Márton Rabi and Walter G. Joyce, 2012, New Insights into the Evolution of the Carotid Circulation System in Eucryptodiran Turtles, Symposium on Turtle Evolution, University of Tübingen, Germany, 2-4 June 2012, p.36
Overview, Macrobaenidae, Sinemydidae

Márton Rabi, Walter G. Joyce, & Oliver Wings(2010) A review of the Mesozoic turtles of the Junggar Basin (Xinjiang, Northwest China) and the paleobiogeography of Jurassic to Early Cretaceous Asian testudinates, Palaeobio Palaeoenv 90:259–273 DOI 10.1007/s12549-010-0031-3
Sinemydids and Macrobaenids, Xinjiangchelyidae

Radley, Jonathan D. Distribution and Environmental Significance of Molluscs in the Late Jurassic-Early Cretaceous Purbeck Formation of Dorset, Southern England: a Review. Life and Environments in Purbeck Times. (Special Papers in Palaeontology No. 68).

Rieppel, O (1995), Studies on skeleton formation in reptiles: implications for turtle relationships. Zoology -- Analysis of Complex Systems 98: 298-308.

Rieppel, O. 2000. Turtles as diapsid reptiles. Zool. Scr. 29: 199-212. pdf
Mystery of Chelonian Origins

Rieppel O. & M. deBraga. 1996. Turtles as diapsid reptiles. Nature 384: 453-455.
Mystery of Chelonian Origins

Rieppel, O. and Reisz, R.R. 1999. The Origin and Early Evolution of Turtles. Ann. Rev. Eco. Syst. vol.30:1-22. pdf
Mystery of Chelonian Origins

Reisz, Robert R., & Jason J. Head (2008-11-27). "Palaeontology: Turtle origins out to sea". Nature 456: 450-451. doi:10.1038/nature07533. pdf
Odontochelys, Chinlechelys, Mystery of Chelonian Origins

Reisz, R.R. & Laurin, M. (1991). Owenetta and the Origin of Turtles. Nature 349(6307): 324-326.
Mystery of Chelonian Origins

Rest, J.S., Ast, J.C., Austin, C.C., Waddell, P.J., Tibbetts, E.A., Hay, J.M., Mindell, D.P. (2003). Molecular Systematics of Primary Reptilian Lineages and the Tuatara Mitochondrial Genome. Molecular Phylogenetics and Evolution vol.29 (2): 289-297. pdf.
Mystery of Chelonian Origins

Romer, A. S. 1956. The osteology of the reptiles. Chicago, Univ. of Chicago Press.
Chelonian phylogenetics and systematics

Rougier, G. W., Fuenta de la M. S., and Arcucci A. B., 1995, Late Triassic Turtles from South America: Science, v. 268, p. 855-858. abstract

Scheyer, T. M. 2007. Comparative bone histology of the turtle shell. (carapace and plastron): implications for turtle systematics, functional morphology and turtle origins. Bonn, Univ., Diss..
Mystery of Chelonian Origins

Scheyer, T M (2008). Aging the oldest turtles: the placodont affinities of Priscochelys hegnabrunnensis. Naturwissenschaften, 95(9):803-810.
Mystery of Chelonian Origins

Shaffer, H. B. 2009. Turtles (Testudines). In The Timetree of Life. S. B. Hedges and S. Kumar, Eds. (Oxford University Press, New York), pp. 398-401. pdf
Morphology or Molecules?, Trionychoidea

Shaffer, HB, P Meylan & ML McKnight (1997), Tests of turtle phylogeny: molecular, morphological, and paleontological approaches. Syst. Biol. 46: 235-268. pdf
Meiolaniidae, Trionychoidea, Testudines.

Shedlock, A.M. & Edwards, S.V. 2009 . Amniotes (Amniotes). In The Timetree of Life. S. B. Hedges and S. Kumar, Eds. (Oxford University Press, New York), pp. 375-9 .
Mystery of Chelonian Origins

Shimada, Kenshu; M.J. Everhart, G. E. Hooks III (2002). "Ichthyodectid fish and protostegid turtle bitten by the Late Cretaceous lamniform shark, Cretoxyrhina mantelli". Journal of Vertebrate Paleontology 22 (3): 106.

Shimada, Kenshu; G. E. Hooks III (January 2004). "Shark-bitten protostegid turtles from the upper Cretaceous Mooreville chalk, Alabama". Journal of Paleontology 78 (1): 205–210.

Simpson, G. G. 1938. Crossochelys, Eocene horned turtle from Patagonia. Bull. Amer. Mus. Nat. Hist., vol. 74, pp. 221-254.
Chelonian phylogenetics and systematics

Slater, BJ, Reolid, M, Schouten, R, & Benton, MJ, 2011, A new Late Jurassic turtle from Spain: phylogenetic implications, taphonomy and palaeoecology. Palaeontology, Vol. 54, Part 6, 2011, pp. 1393–1414 pdf

Spotila, James R. (2004). Sea Turtles: A Complete Guide to their Biology, Behavior, and Conservation. Baltimore, Maryland: The Johns Hopkins University Press and Oakwood Arts.

Sterli, J., 2008: A new, nearly complete stem turtle from the Jurassic of South America with implications for turtle evolution. Biology Letters: Vol. 2008, #4, pp. 286-289 [doi: 10.1098/rsbl.2008.0022] pdf.
Chelonian phylogenetics and systematics, Condorchelys

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 (3) 93-106 pdf
Chelonian phylogenetics and systematics, Morphology or Molecules?, Pleurodira, Protostegidae

Sterli, J., (2012). New Findings of Meiolaniid and Meiolaniid-like Taxa from Patagonia, Argentina, Abstracts, Symposium on Turtle Evolution, University of Tübingen, June 01-04, 2012 pdf

Juliana Sterli and Marcelo de la Fuente (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 (1): 63–78. doi:10.1111/j.1475-4983.2010.01002.x.

Sukhanov, V.B. 2003. Mesozoic turtles from Mongolia. in Benton, Michael J.; Shishkin, Mikhail A.; Unwin, David M.; and Kurochkin, Evgenii N. (eds.). The Age of Dinosaurs in Russia and Mongolia. Cambridge: Cambridge University Press.

Sukhanov, V.B., Danilov, I.G, & Syromyatnikova, E.V. 2008.The description and phylogenetic position of a new nanhsiungchelyid turtle from the Late Cretaceous of Mongolia. Acta Palaeontol. Pol. 53 (4): 601–614, pdf
, Hanbogdemys, Kharakhutulia, "Nanhsiungchelyini", Nanhsiungchelyidae, Zangerlia

Suzuki, S. & Narmandakh, P., 2004 Change of the Cretaceous Turtle Faunas in Mongolia. Hayashibara Museum of Natural Sciences Research Bulletin 2, p.7-14 (2004) pdf

Syromyatnikova, Elena, 2012, Fossil turtle families Adocidae and Nanhsiungchelyidae of Asia: morphology, taxonomy, phylogeny and extension. Phd thesis dissertation On-line (in Russian)

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, RAS, v. 313, n. 1, p. 74-94. pdf.

Thomson, Robert C. & H. Bradley Shaffer. 2010. Sparse Supermatrices for Phylogenetic Inference: Taxonomy, Alignment, Rogue Taxa, and the Phylogeny of Living Turtles. Systematic Biology. 59(1): pp. 42-58 pdf
Morphology or Molecules?, Xerobatinae

Haiyan Tong, Julien Claude, Wilailuck Naksri, Varavudh Suteethorn, Eric Buffetaut, Sasidhorn Khansubha, Kamonrak Wongko and Phisit Yuangdetkla, 2009, 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

Tong, Haiyan & Ren Hirayama (2008). A new species of Argillochelys (Testudines: Cryptodira: Cheloniidae) from the Ouled Abdoun phosphate basin, Morocco. Bulletin de la Société Géologique de France 179 (6): 623–630. doi:10.2113/gssgfbull.179.6.623.

Haiyan Tong, Hirayama R., Makhoul, E. Escuillié, F. 2006. Rhinochelys (Chelonioidea: Protostegidae) from the Late Cretaceous (Cenomanian) of Nammoura, Lebanon. Atti della società italiana di scienze naturali e storia del Museo Civico di naturale di Milano 147 (1): 113-138

Watson, D.M.S. (1914). "Eunotosaurus africanus Seeley and the ancestors of the Chelonia". Proceedings of the Zoological Society of London 11: 1011-1020.
Mystery of Chelonian Origins, Chelonian phylogenetics and systematics

Wieland, G. R. 1896. Archelon ischyros: a new gigantic cryptodire testudinate from the Fort Pierre Cretaceous of South Dakota. American Journal of Science, 4th Series 2(12):399-412, pl. v., text-figs. I, 2.

Wieland, 1909, Amer. Jour. Sci., (4), 27, p. 104.

Werneburg, Ingmar & Marcelo R Sánchez-Villagra 2009 . Timing of organogenesis support basal position of turtles in the amniote tree of life. BMC Evolutionary Biology. 9: pp. 1-9. pdf
Mystery of Chelonian Origins

Wilkinson, M., Thorley, J., and Benton, M. J. 1997. Uncertain turtle relationships. Nature 387: 466.
Mystery of Chelonian Origins

Williston, Univ. Gaol. Surv. Kansas, 11, 1897, p. 246.
Archelon, Protostega

Willmore, K.E. (2010) Development Influences Evolution, American Scientist, Vol. 98, no. 3. May-June 2010
Mystery of Chelonian Origins

Zangerl, Rainer (1953a). "The vertebrate fauna of the Selma Formation of Alabama, Part III. The turtles of the Family Protostegidae". Chicago Field Museum Mem. 3 (3). internet archive
Calcarichelys, Chelosphargis, Protostega, Rhinochelys

Zangerl, Rainer (1953b). "The vertebrate fauna of the Selma Formation of Alabama, Part IV. The turtles of the Family Toxochelyidae". Chicago Field Museum Mem. 3 (4). internet archive
Toxochelyidae, Toxochelys

Zardoya, R., and Meyer, A. 1998. Complete mitochondrial genome indicates diapsid affinities of turtles. Proc. Natl. Acad. Sci. USA 95: 14226-14231.
Mystery of Chelonian Origins

Zardoya, R., and Meyer, A. 2001. The evolutionary position of turtles revised. Naturwissenschaften 88 (2001), pp. 193-200.
Mystery of Chelonian Origins

Zelenitsky et al. , Electronic Supplementary Material, Phylogenetic affinities of Adocus pdf
Adocidae, Trionychoidea

Zhang, Zhi-Qiang (2011), Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness, Zootaxa, Volume 3148, pp. 1-237 pdf

Zhou, C-F, 2010, A second specimen of Manchurochelys manchoukuoensis Endo & Shikama, 1942 (Testudines: Eucryptodira) from the Early Cretaceous Yixian Formation of western Liaoning, China, Zootaxa 2534: 57–66. pdf

Zittel, K. A. von. Text-Book of Palaeontology Volume 2 . Translated & edited by Charles R. Eastman. Vol. II. First English edition 1902, Public Domain (copyright expired), Internet archive; Second English edition revised, with additions, by Sir Arthur Smith Woodward, 1932. London, Macmillan and Co., Ltd.; New York, Macmillan
Allopleuron, Argillochelys, Caretta, Chelonia, Chitracephalus, Corsochelys, Dermochelyinae, Dermochelys, Eochelone , Eosphargis, Euclastes, Hadrianus, Plesiochelyidae, Protosphargis, Psephophorus, Stylemys, Toxochelys

Zug, G. R. 2001 Turtles of the Lee Creek mine (Pliocene: North Carolina). Smithsonian Contributions to Paleobiology 90:203–218.

checked ATW031014, MAK100926. Page last modified MAK110922.

All material by ATW is public domain and may be freely used in any way (also any material jointly written by ATW and MAK). All material by MAK is licensed Creative Commons Attribution License Version 3.0, and may be freely used provided acknowedgement is given. All Wikipedia material is either Gnu Open Source or Creative Commons (see original Wikipedia page for details). Other graphics are copyright their respective owners