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Cryptodira

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Suborder of reptiles

Cryptodira Temporal range:Late Jurassic –Present PreꞒ Ꞓ O S D C P T J K Pg N

Aldabra giant tortoise (Aldabrachelys gigantea)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Testudines
Suborder:	Cryptodira Cope, 1868^[1]
Subgroups
See text
Synonyms^[1]^[2]
CryptoderesDuméril and Bibron, 1834 CryptoderaLichtenstein, 1856 CryptodiraCope, 1868 CryptodiramorphaLee, 1995 PancryptodiraJoyce, Parham, and Gauthier, 2004

Skull of a cryptodiran turtle from the familyEmydidae

Dorsal view ofskull andcervical vertebrae of a cryptodiran turtle from the familyEmydidae. Not all cervical vertebrae are featured due to the dissection cut.

TheCryptodira (Greek:hidden neck) are asuborder ofTestudines that includes most livingtortoises andturtles. Cryptodira is commonly called the "Hidden-Neck Turtles" or the "Inside-Neck Turtles". Cryptodira differ fromPleurodira (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 speciesfreshwater turtles,snapping turtles,tortoises,softshell turtles, andsea turtles.

Neck retraction

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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 ofcervical vertebrae. In all recent turtles, the cervical column consists of nine joints and eight vertebrae.^[4] Compared to the narrow vertebrae and the closely positionedzygapophyses 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 calledginglymoidy, 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]

A comparison of cryptodiran neck retraction compared toPleurodiran neck retraction.

Cryptodiran neck retraction is also dependent on associated cervical musculature for its characteristic motions. A study that focused solely on the mechanism of neck retraction inChelodina (pleurodire) versus that ofApalone (cryptodire), found an absence of thelongissimus andiliocostalis systems and reduced epaxial musculature.^[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 developedtransverse 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.^{[citation needed]}

Systematics and evolution

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Cryptodiresevolved from pleurodires during the earlyJurassic period, originating fromSouth America andSoutheast Asia.^[6] By the end of the Jurassic, cryptodires 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 theCretaceous toEocene ofEurope,^[7] and produced a family of marine species, theBothremydidae.

The Cryptodira suborder has four livingsuperfamilies, theChelonioidea (sea turtles),Testudinoidea (tortoises and pond turtles),Kinosternoidea (Central American river turtle and mud turtles) andTrionychoidea (soft-shell turtles and relatives).Chelydridae (snapping turtles) form a sister group toKinosternoidea. The former three subfamilies (and Chelydridae) are classified in the cladeDurocryptodira, while the latter is classified in the cladeTrionychia. These two clades likely diverged in the middle of the Jurassic.^[6]^[8]

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

The alternate concept restricts the use of the term "Cryptodira" to thecrown clade (i.e. Polycryptodira). The Cryptodira as understood here are calledCryptodiramorpha in this view.A recent study placedPlesiochelyidae as anAngolachelonia and outsideTestudines, thus Cryptodira.^[9]

As per the system used here, the Cryptodira can be classified as:^[8]^[10]

†Hangaiemys
†Bashuchelyidae
Pan-Cryptodira
- Family †Macrobaenidae?Sukhanov 1964
- F amily †Xinjiangchelyidae?Nesov, 1990
- Family †Sinemydidae?Yeh, 1963
- CladePandurocryptodira
  - CladeAmerichelydia
    - CladeChelydroidea
      - FamilyChelydridae (snapping turtles)
      - SuperfamilyKinosternoidea
        FamilyDermatemydidae (river turtles)
        FamilyKinosternidae (mud turtles)
    - CladePanchelonioidea
      - Clade †Angolachelonia?
        Clade †Thalassochelydia?
        Family †Eurysternidae?
        Family †Plesiochelyidae?
        Family †Thalassemydidae?
        †Owadowia?
        Family †Sandownidae? (may also belong in Thalassochelydia)
      - Family †Protostegidae? (may also belong in Thalassochelydia)
      - Family †Toxochelyidae
      - Family †Ctenochelyidae
      - SuperfamilyChelonioidea (sea turtles)
        FamilyCheloniidae (green sea turtles and relatives)
        FamilyDermochelyidae (leatherback sea turtles)
  - CladePantestudinoidea
    - Family †Lindholmemydidae?
    - SuperfamilyTestudinoidea
      - Family †Haichemydidae
      - Family †Sinochelyidae
      - CladeEmysternia
        FamilyPlatysternidae (big-headed turtle)
        FamilyEmydidae (pond,box andwater turtles)
      - CladeTestuguria
        FamilyGeoemydidae (Asian river turtles,Asian leaf turtles,Asian box turtles androofed turtles)
        FamilyTestudinidae (tortoises)
- CladePantrionychia
  - Clade †Adocusia
    - Family †Adocidae
    - Family †Nanhsiungchelyidae
  - SuperfamilyTrionychia
    - FamilyCarettochelyidae (pignose turtles)
    - FamilyTrionychidae (softshell turtles)

Ordosemys liaoxiensis was a member ofSinemydidae.
Kinixys belliana from theTestudinidae
TheIndian flapshell turtle (Lissemys punctata) from theTrionychidae is a highly advanced eucryptodire.

Distribution

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Trionychidae (softshell turtles) are found from North America, Africa, South and East Asia to New Guinea.
Kinosternidae (mud and musk turtles) are found from eastern North America to the Amazon drainage of South America.
Dermatemydidae (Mesoamerican river turtles) are found in the Caribbean-Gulf drainage of Mesoamerica.
Emydidae (cooters, sliders, American box turtles, and Allies) are found from Europe to Ural Mountains and North America southward to Eastern Brazil.

References

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^^a ^bRhodin 2011, p. 000.171
^Rhodin 2008, p. 000.3
^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.
^^a ^bWyneken, Jeanette; Bels, V. L. (Vincent L.); Godfrey, Matthew H., eds. (2008).Biology of Turtles. Boca Raton, Florida: CRC Press.ISBN 9780849333392.OCLC 144570900.
^"Oxford Index: ginglymoidy". Archived fromthe original on 2020-01-31. Retrieved2018-05-05.
^^a ^bPereira, Anieli G.; Sterli, Juliana; Moreira, Filipe R.R.; Schrago, Carlos G. (August 2017)."Multilocus phylogeny and statistical biogeography clarify the evolutionary history of major lineages of turtles".Molecular Phylogenetics and Evolution.113:59–66.Bibcode:2017MolPE.113...59P.doi:10.1016/j.ympev.2017.05.008.hdl:11336/41137.ISSN 1055-7903.PMID 28501611.
^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.S2CID 15652309.
^^a ^bJoyce, Walter G.; Anquetin, Jérémy; Cadena, Edwin-Alberto; Claude, Julien; Danilov, Igor G.; Evers, Serjoscha W.; Ferreira, Gabriel S.; Gentry, Andrew D.; Georgalis, Georgios L.; Lyson, Tyler R.; Pérez-García, Adán (2021-02-09)."A nomenclature for fossil and living turtles using phylogenetically defined clade names".Swiss Journal of Palaeontology.140 (1): 5.Bibcode:2021SwJP..140....5J.doi:10.1186/s13358-020-00211-x.hdl:11336/155192.ISSN 1664-2384.
^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.Bibcode:2019Palgy..62...93E.doi:10.1111/pala.12384.S2CID 134736808.
^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.S2CID 85295987.

External links

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Data related toCryptodira at Wikispecies

Testudines

Kingdom:Animalia
Phylum:Chordata
Class:Reptilia
Clade:Diapsida
Order:Testudines

Genus

Chelonioidea
(Sea turtles)

Cheloniidae	†Allopleuron Caretta †Carolinochelys Chelonia †Eochelone Eretmochelys †Gigantatypus †Glarichelys †Itilochelys Lepidochelys †Mexichelys †Miocaretta Natator †Pacifichelys †Syllomus †Tasbacka
Dermochelyidae	†Arabemys †Corsochelys †Cosmochelys Dermochelys †Eosphargis †Mesodermochelys †Psephophorus
	†Euclastes †Peritresius †Procolpochelys †Protosphargis †Puppigerus

Kinosternoidea

Dermatemydidae	Dermatemys
Kinosternidae	Claudius †Hoplochelys Kinosternon Staurotypus Sternotherus

Testudinoidea

Emydidae	†Acherontemys Chrysemys Clemmys Deirochelys Emys Actinemys Emydoidea Glyptemys Graptemys Malaclemys Pseudemys Terrapene Trachemys †Wilburemys
Geoemydidae	Batagur †Banhxeochelys Cuora Cyclemys Geoclemys Geoemyda Hardella Heosemys Leucocephalon Malayemys Mauremys Melanochelys Morenia Notochelys Orlitia Pangshura Rhinoclemmys Sacalia Siebenrockiella Vijayachelys
Platysternidae	Platysternon
Testudinidae	Aldabrachelys Astrochelys Centrochelys Chelonoidis Chersina Cylindraspis †Cymatholcus †Floridemys Geochelone Gopherus †Hadrianus †Hesperotestudo Homopus Indotestudo Kinixys Malacochersus Manouria †Megalochelys †Oligopherus Psammobates Pyxis †Solitudo Stigmochelys †Stylemys Testudo

Trionychia

Carettochelyidae	†Allaeochelys †Anosteira Carettochelys
Trionychidae	Amyda Apalone †Axestemys Chitra Cyclanorbis Cycloderma †Drazinderetes Dogania †Gilmoremys †Hutchemys †Khunnuchelys Lissemys Nilssonia Palea †Palaeoamyda Pelochelys Pelodiscus Rafetus Trionyx
	†Basilochelys †Sinaspideretes

Chelydridae	Chelydra †Chelydrops †Chelydropsis †Emarginachelys †Macrocephalochelys Macrochelys †Planiplastron †Protochelydra
†Nanhsiungchelyidae	†Anomalochelys †Basilemys †Jiangxichelys
†Protostegidae	†Alienochelys †Archelon †Atlantochelys †Bouliachelys †Calcarichelys †Cratochelone †Desmatochelys †Iserosaurus †Notochelone †Ocepechelon †Pneumatoarthrus †Protostega †Rhinochelys †Santanachelys †Terlinguachelys
	†Adocus †Argillochelys †Bashuchelys †Ctenochelys †Prionochelys †Toxochelys

Pleurodira

†Araripemydidae	†Araripemys
†Bothremydidae	†Araiochelys †Arenila †Azabbaremys †Bothremys †Cearachelys †Chedighaii †Chupacabrachelys †Eotaphrosphys †Foxemys †Galianemys †Ilatardia †Inaechelys †Itapecuruemys †Jainemys †Kinkonychelys †Kurmademys †Labrostochelys †Nigeremys †Phosphatochelys †Polysternon †Puentemys †Rosasia †Rhothonemys †Sankuchemys †Taphrosphys †Ummulisani †Zolhafah
Chelidae	Acanthochelys Chelodina Chelus Elseya Elusor Emydura Hydromedusa †Lomalatachelys Mesoclemmys Myuchelys Phrynops Platemys †Prochelidella Pseudemydura Rheodytes Rhinemys †Yaminuechelys
Pelomedusidae	Pelomedusa Pelusios
Podocnemididae	†Albertwoodemys †Bauruemys †Brontochelys †Caninemys †Carbonemys †Cerrejonemys †Cordichelys Erymnochelys †Lapparentemys †Latentemys Peltocephalus Podocnemis †Stupendemys
†Sahonachelyidae	†Sahonachelys †Sokatra

	†Caribemys †Caririemys †Tacuarembemys

Phylogenetic arrangement of turtles based onTurtles of the World 2017 Update: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status. † = extinct.

See alsoList of Testudines families

Taxon identifiers
Cryptodira	Wikidata:Q499621 Wikispecies:Cryptodira BioLib:16332 CoL:87BW5 EoL:46559463 Fauna Europaea:15550 Fauna Europaea (new):a657826b-f777-4743-8b09-2a7d3deae97b iNaturalist:474372 ITIS:948943 NBN:NBNSYS0000180887 NCBI:8464 Paleobiology Database:37622 WoRMS:148741