Cryptodira

The Cryptodira (Greek: hidden neck) are a suborder of Testudines that includes most living tortoises and turtles. Cryptodira differ from Pleurodira (side-necked turtles) in that they lower their necks and pull the heads straight back into the shells, instead of folding their necks sideways along the body under the shells' marginals. They include among their species freshwater turtles, snapping turtles, tortoises, softshell turtles, and sea turtles.

Cryptodira Temporal range: Early Jurassic—present, 190–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Aldabra giant tortoise (Aldabrachelys gigantea)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Testudines
Suborder:	Cryptodira Cope, 1868[1]
Subgroups
See text
Synonyms[1][2]
Cryptoderes Duméril and Bibron, 1834 Cryptodera Lichtenstein, 1856 Cryptodira Cope, 1868 Cryptodiramorpha Lee, 1995 Pancryptodira Joyce, Parham, and Gauthier, 2004

Skull of a cryptodiran turtle from the family Emydidae

Dorsal view of skull and cervical vertebrae of a cryptodiran turtle from the family Emydidae. Not all cervical vertebrae are featured due to the dissection cut.

Neck retraction

The Cryptodira are characterized by retraction of the head in the vertical plane, which permits for primarily vertical movements and restricted lateral movements outside of the shell.[3] These motions are largely due to the morphology and arrangement of cervical vertebrae. Of all recent turtles, the cervical column consists of nine joints and eight vertebrae.[4] Compared to the narrow vertebrae and the closely positioned zygapophyses of the pleurodires, the cryptodires’ vertebrae take on the opposite shape. Their cervical vertebrae are more distended, and their zygapophyses (processes that interlock adjacent vertebrae) are much more widely spaced - features allowing for a condition called ginglymoidy, and ultimately, their “hidden” neck retraction. Ginglymoidy refers to the double articulation where articulation between the sixth and seventh vertebrae and the seventh and eighth vertebrae allows for bending of the neck into an S shape. Formation of this S shape occurs in one plane that enables retraction into the shell.[5]

Cryptodiran neck retraction is also dependent on associated cervical musculature for its characteristic motions. In a study that focused solely on the mechanism of neck retraction in Chelodina (Pleurodire) versus that of Apalone (Cryptodire), an absence of the longissimus system, the Iliocostalis system and minimization of the epaxial musculature was found.[4] Absence of longissimus musculature, which primarily functions in moving the neck via ipsilateral flexion and contralateral rotation, contributes to the backwards retraction of the neck into the shell. Lack of this muscular system also results in poorly developed transverse processes (the lateral processes of a vertebra), forcing them to be developed in a more cranial direction. The iliocostalis system, used for lateral flexion and extension of the vertebral column, is commonly absent in all turtles. With the presence of a shell, these muscular movements are no longer possible. Epaxial musculature that functions in alternated forms of stepping and walking is minimized in turtles, due to their restricted stride lengths and heavily weighted shells.

Systematics and evolution

Cryptodires evolved during the Jurassic period, and by the end of the Jurassic had almost completely replaced pleurodires in the lakes and rivers, while beginning to develop land-based species. Meanwhile, pleurodires became the dominant freshwater testudines in the Cretaceous to Eocene of Europe,[6] and produced a family of marine species, the Bothremydidae.

The Cryptodira suborder has three living superfamilies, the Chelonioidea (sea turtles), Testudinoidea (tortoises and pond turtles), and Trionychoidea (soft-shell turtles and relatives). The families within the Kinosternoidea are now recognized as a paraphyletic assemblage of mostly primitive Trionychoidea; they do not form a natural group.

Two circumscriptions of the Cryptodira are commonly found. One is used here; it includes a number of primitive extinct lineages known only from fossils, as well as the Eucryptodira. These are, in turn, made up from some very basal groups, and the Centrocryptodira contain the prehistoric relatives of the living cryptodires, as well as the latter, which are collectively called Polycryptodira.

The alternate concept restricts the use of the term "Cryptodira" to the crown clade (i.e. Polycryptodira). The Cryptodira as understood here are called Cryptodiramorpha in this view. A recent study placed Plesiochelyidae as an Angolachelonia and outside Testudines, thus Cryptodira.[7]

As per the system used here, the Cryptodira can be classified as:[8]

Manchurochelys liaoxiensis was a member of Sinemydidae.

Kinixys belliana from the Testudinidae

The Indian flapshell turtle (Lissemys punctata) from the Trionychidae is a highly advanced eucryptodire.

• †Hangaiemys
• †Bashuchelyidae
• Eucryptodira
  • Clade Americhelydia
    • †Emarginachelys
    • Family Chelydridae (snapping turtles)
    • Superfamily Kinosternoidea
      • Family Dermatemydidae (river turtles)
      • Family Kinosternidae (mud turtles)
    • Clade Panchelonioidea
      • Family †Sandownidae
      • Family †Xinjiangchelyidae
      • Family †Sinemydidae
      • Family †Toxochelyidae
      • †Ctenochelys
      • Clade Thalassochelydia
        • †Owadowia
        • Family †Eurysternidae
        • Family †Plesiochelyidae
        • Family †Thalassemydidae
      • Superfamily Chelonioidea (sea turtles)
        • Family †Protostegidae
        • Family Cheloniidae (green sea turtles and relatives)
        • Family Dermochelyidae (leatherback sea turtles)
  • Superfamily Testudinoidea
    • Family †Haichemydidae
    • Family †Lindholmemydidae
    • Family †Sinochelyidae
    • Family Platysternidae (big-headed turtle)
    • Family Emydidae (pond, box and water turtles)
    • Family Geoemydidae (Asian river turtles, Asian leaf turtles, Asian box turtles and roofed turtles)
    • Family Testudinidae (tortoises)
  • Clade Pantrionychia
    • Clade †Adocusia
      • Family †Adocidae
      • Family †Nanhsiungchelyidae
    • Superfamily Trionychia
      • Family Carettochelyidae (pignose turtles)
      • Family Trionychidae (softshell turtles)

References

1. Rhodin 2011, p. 000.171
2. Rhodin 2008, p. 000.3
3. Anquetin, Jérémy; Tong, Haiyan; Claude, Julien (16 February 2017). "A Jurassic stem pleurodire sheds light on the functional origin of neck retraction in turtles". Scientific Reports. 7 (1): 42376. Bibcode:2017NatSR...742376A. doi:10.1038/srep42376. PMC 5312562. PMID 28206991.
4. Biology of turtles. Wyneken, Jeanette, 1956-, Bels, V. L. (Vincent L.), Godfrey, Matthew H. Boca Raton: CRC Press. 2008. ISBN 9780849333392. OCLC 144570900.CS1 maint: others (link)
5. "ginglymoidy - oi". Cite journal requires |journal= (help)
6. Pérez-García, Adán (4 June 2016). "A new turtle confirms the presence of Bothremydidae (Pleurodira) in the Cenozoic of Europe and expands the biostratigraphic range of Foxemydina". The Science of Nature. 103 (7–8): 50. Bibcode:2016SciNa.103...50P. doi:10.1007/s00114-016-1375-y. PMID 27262289.
7. Evers, Serjoscha W.; Benson, Roger B. J.; Smith, Andrew (January 2019). "A new phylogenetic hypothesis of turtles with implications for the timing and number of evolutionary transitions to marine lifestyles in the group". Palaeontology. 62 (1): 93–134. doi:10.1111/pala.12384.
8. Anquetin, Jérémy (9 November 2011). "Reassessment of the phylogenetic interrelationships of basal turtles (Testudinata)". Journal of Systematic Palaeontology. 10 (1): 3–45. doi:10.1080/14772019.2011.558928.

Further reading

• Baur, George (June 1890). "On the Classification of the Testudinata". The American Naturalist. 24 (282): 530–536. doi:10.1086/275138. JSTOR 2450882.
• Rhodin, Anders G.J.; van Dijk, Peter Paul; Inverson, John B.; Shaffer, H. Bradley; Roger, Bour (2011-12-31). "Turtles of the world, 2011 update: Annotated checklist of taxonomy, synonymy, distribution and conservation status" (PDF). Chelonian Research Monographs. 5. Archived from the original (PDF) on 2012-01-31.
• Rhodin, Anders G.J.; van Dijk, Peter Paul; Parham, James F. (2008-12-08). "Turtles of the world, 2008 checklist" (PDF). Chelonian Research Monographs. 5. Archived from the original (PDF) on 2012-04-23. Retrieved 2012-05-22.