Temporal range: Late Cretaceous to present, 84.9–0 Ma
Yellow-spotted river turtle (Podocnemis unifilis)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Testudines
Suborder: Pleurodira
Clade: Podocnemidoidae
Family: Podocnemididae
Cope, 1868[1]
Extant genera

For fossil genera see text



Podocnemididae is a family of pleurodire (side-necked) turtles, once widely distributed. Most of its 41 genera and 57 species are now extinct. Seven of its eight surviving species are native to South America: the genus Peltocephalus, with two species, only one of which is extant (P. dumerilianus, the Big-headed Amazon River turtle); and the genus Podocnemis, with six living species of South American side-necked river turtles and four extinct. There is also one genus native to Madagascar: Erymnochelys, the Madagascan big-headed turtle, whose single species E. madagascariensis.

Like other pleurodire turtles, podocs have a "side-necked" defensive posture, turning the head sideways to hide it under the shell. Another characteristic of pleurodires is that the pelvis is fused to the shell which prevents pelvic motion, making it difficult to walk on land.[2][3] Podocnemididae turtles live in aquatic environments and have shells streamlined to aid in swimming.[4]

Taxonomy and systematics

Podocnemis skulls, ventral and side view, with pterygoid and basisphenoid bones labeled "pt" and "bs" respectively.

According to Ferreira et al. (2015), the family name derives from two Greek words: "podos" (foot) and "cnemis" (leg armor worn by Roman soldiers.)[5]

To clarify some closely related names:

According to Gaffney et al. (2011), the family Podocnemididae can be diagnosed from its cranial traits including "the unique possession of a cavum pterygoidei formed by the basisphenoid, pterygoid, prootic, and quadrate [bones], underlain by the pterygoid and basisphenoid."[8]

The pocnemid family dates to the late Cretaceous; it includes 20 genera and 30 species. Only three genera (and eight species) survive.[8]

The three living genera of Podocnemididae (one of which is monotypic) are:


Fossils show that Podocnemidids were once found in Europe, Asia, North America, and Africa. Stupendemys lived around 5.5 million years ago in northern South America, and was the largest freshwater turtle with a carapace length of 2.4 metres (7.9 ft), the largest of any known turtle and is the largest pleurodire known.[10] While Peltocephalus and Erymnochelys have often been recovered as more closely related to each other than to Podocnemis in morphological analyses,[11][12][9] genetic studies have found Erymochelys to be more closely related to Podocnemis than to Peltocephalus.[13][14]


Morphology based cladogram after Ferreira et al. 2024[22]






Neochelys arenarum

Neochelys franzeni



Neochelys fajumensis





†UCMP 42008

Peltocephalus dumerilianus

Peltocephalus maturin






Bairdemys healeyorum






Bairdemys spp.


  1. ^ Cope, Edward Drinker (1868). "On the Origin of Genera". Proceedings of the Academy of Natural Sciences of Philadelphia. 20 (106): 242–300. JSTOR 4059902. Three phalanges on most digits; Zygomatic arch; no parieto-mastoid .. Podocnemididae. Temporal fossa overroofed by parietal.. Podocnemis.
  2. ^ Wise, Taylor B.; Stayton, C. Tristan (2017-03-01). "Side-necked Versus Hidden-necked: A Comparison of Shell Morphology Between Pleurodiran and Cryptodiran Turtles". Herpetologica. 73 (1): 18. doi:10.1655/HERPETOLOGICA-D-15-00038. ISSN 0018-0831. S2CID 90226667.
  3. ^ Sánchez-Villagra, Marcelo R.; Winkler, Jasmin D.; Wurst, Linda (2007-04-19). "Autopodial skeleton evolution in side-necked turtles (Pleurodira): Pleurodire autopodial evolution" (PDF). Acta Zoologica. 88 (3): 199–209. doi:10.1111/j.1463-6395.2007.00267.x. S2CID 83721738.
  4. ^ Mayerl, Christopher J.; Brainerd, Elizabeth L.; Blob, Richard W. (2016-06-23). "Pelvic girdle mobility of cryptodire and pleurodire turtles during walking and swimming". The Journal of Experimental Biology. 219 (17): 2650–2658. doi:10.1242/jeb.141622. ISSN 0022-0949. PMID 27340204.
  5. ^ a b Ferreira, Gabriel S.; Rincón, Ascanio D.; Solórzano, Andrés; Langer, Max C. (2015). "The last marine pelomedusoids (Testudines: Pleurodira): a new species of Bairdemys and the paleoecology of Stereogenyina" (PDF). PeerJ. 3: 1–29. doi:10.7717/peerj.1063. PMC 4493680. PMID 26157628. Archived from the original (PDF) on 2017-12-02. Retrieved 2019-02-20.
  6. ^ a b Gaffney, E. S.; Tong, H.; Meylan, P. A. (2006-11-17). "Evolution of the side-necked turtles: The families Bothremydidae, Euraxemydidae, and Araripemydidae" (PDF). Bulletin of the American Museum of Natural History. 300. New York: American Museum of Natural History: 1–700. doi:10.1206/0003-0090(2006)300[1:EOTSTT]2.0.CO;2. hdl:2246/5824. S2CID 85790134. The relationships of the family Podocnemididae to its sister taxa Hamadachelys and Brasilemys are recognized by placing them in the epifamily Podocnemidinura. The epifamily Podocnemidinura is the sister group to the family Bothremydidae, and together they form the superfamily Podocnemidoidea.
  7. ^ Fritz Jürgen Obst (1998). "Pelomedusinae". In H. G. Cogger; R. G. Zweifel (eds.). Encyclopedia of Reptiles and Amphibians. San Diego: Academic Press. pp. 112–113. ISBN 0-12-178560-2.
  8. ^ a b c Eugene S. Gaffney; Peter A. Meylan; Roger C. Wood; Elwyn Simons; Diogenes De Almeida Campos (2011). "Evolution of the Side-Necked Turtles: The Family Podocnemididae". Bulletin of the American Museum of Natural History. 350: 1–237. doi:10.1206/350.1. hdl:2246/6110. S2CID 83775718. The family Podocnemididae consists of 20 genera and 30 species considered here as valid and diagnosable by cranial characters. Three of these genera and eight species persist into the Recent fauna, barely reflecting the evolutionary diversity and distribution of the group. The family extends from the late Cretaceous to the Recent and occurs in North and South America, Europe, Asia, and Africa.
  9. ^ a b Ferreira, G. S.; Nasciemento, E. R.; Cadena, E. A.; Cozzuol, M. A.; Farina, B. M.; Pacheco, M. L. A. F.; Rizzutto, M. A.; Langer, M. C. (2024). "The latest freshwater giants: a new Peltocephalus (Pleurodira: Podocnemididae) turtle from the Late Pleistocene of the Brazilian Amazon". Biology Letters. 20 (3). 20240010. doi:10.1098/rsbl.2024.0010. PMC 10932709. PMID 38471564.
  10. ^ Cadena, E.-A.; Scheyer, T. M.; Carrillo-Briceño, J. D.; Sánchez, R.; Aguilera-Socorro, O. A; Vanegas, A.; Pardo, M.; Hansen, D. M.; Sánchez-Villagra, M. R. (February 2020). "The anatomy, paleobiology, and evolutionary relationships of the largest extinct side-necked turtle". Science Advances. 6 (7): eaay4593. Bibcode:2020SciA....6.4593C. doi:10.1126/sciadv.aay4593. ISSN 2375-2548. PMC 7015691. PMID 32095528.
  11. ^ de la Fuente, Marcelo S.; Maniel, Ignacio J.; Ruiz, Pablo González; Ledesma, Jonathan; Deraco, Maria V.; del Papa, Cecilia; Herrera, Claudia (2022-12-31). "A new podocnemidid (Pleurodira: Pelomedusoides) from the Eocene of north-western Argentina, with comments on its evolutionary relationships and palaeoenvironmental settings". Journal of Systematic Palaeontology. 20 (1). doi:10.1080/14772019.2022.2081939. ISSN 1477-2019.
  12. ^ Ferreira, Gabriel S.; Bronzati, Mario; Langer, Max C.; Sterli, Juliana (March 2018). "Phylogeny, biogeography and diversification patterns of side-necked turtles (Testudines: Pleurodira)". Royal Society Open Science. 5 (3): 171773. Bibcode:2018RSOS....571773F. doi:10.1098/rsos.171773. ISSN 2054-5703. PMC 5882704. PMID 29657780.
  13. ^ Vargas-Ramírez, Mario; Castaño-Mora, Olga V.; Fritz, Uwe (2008-12-20). "Molecular phylogeny and divergence times of ancient South American and Malagasy river turtles (Testudines: Pleurodira: Podocnemididae)". Organisms Diversity & Evolution. 8 (5): 388–398. doi:10.1016/j.ode.2008.10.001. ISSN 1439-6092.
  14. ^ Selvatti, Alexandre Pedro; Moreira, Filipe Romero Rebello; Carvalho, Daniel Cardoso; Prosdocimi, Francisco; Russo, Claudia Augusta de Moraes; Junqueira, Ana Carolina Martins (June 2023). "Phylogenomics reconciles molecular data with the rich fossil record on the origin of living turtles". Molecular Phylogenetics and Evolution. 183: 107773. doi:10.1016/j.ympev.2023.107773. PMID 36977459.
  15. ^ De Lapparent, France. "The oldest pre-Podocnemidid turtle (Chelonii, Pleurodira), from the Early Cretaceous, Ceara State, Brasil, and its environment". Treballs del Museu de Geologia de Barcelona. 9: 43–95. Retrieved December 4, 2021. In the Podocnemidoidea, Brasilemys n.g. represents the earliest known specimen of the Podocnemidoidae, immediately after the divergence of the Bothremydidae. It is part of the formidable radiation in the Pelomedusoides which occurs during the early Cretaceous when South America separated from Africa.
  16. ^ Hermanson, Guilherme; Iori, Fabiano V.; Evers, Serjoscha W.; Langer, Max C.; Ferreira, Gabriel S. (2020). "A small podocnemidoid (Pleurodira, Pelomedusoides) from the Late Cretaceous of Brazil, and the innervation and carotid circulation of side-necked turtles". Papers in Palaeontology. 6 (2): 329–347. Bibcode:2020PPal....6..329H. doi:10.1002/spp2.1300. ISSN 2056-2802.
  17. ^ Ferreira, Gabriel S.; Iori, Fabiano V.; Hermanson, Guilherme; Langer, Max C. (September 2018). "New turtle remains from the Late Cretaceous of Monte Alto-SP, Brazil, including cranial osteology, neuroanatomy and phylogenetic position of a new taxon". PalZ. 92 (3): 481–498. Bibcode:2018PalZ...92..481F. doi:10.1007/s12542-017-0397-x. ISSN 0031-0220. S2CID 134701879.
  18. ^ Broin, France de Lapparent de; Chirio, Laurent; Bour, Roger (October 2020). "The oldest erymnochelyine turtle skull, Ragechelus sahelica n. gen., n. sp., from the Iullemmeden basin, Upper Cretaceous of Africa, and the associated fauna in its geographical and geological context". Geodiversitas. 42 (25): 455–484. doi:10.5252/geodiversitas2020v42a25. ISSN 1280-9659. S2CID 226238494.
  19. ^ a b Pérez-García, A. (2021). "New shell information and new generic attributions for the Egyptian podocnemidid turtles "Podocnemis" fajumensis (Oligocene) and "Podocnemis" aegyptiaca (Miocene)". Fossil Record. 24 (2): 247–262. Bibcode:2021FossR..24..247P. doi:10.5194/fr-24-247-2021.
  20. ^ Pérez-García, Adán (2019-03-16). "New information and establishment of a new genus for the Egyptian Paleogene turtle 'Stereogenys' libyca (Podocnemididae, Erymnochelyinae)". Historical Biology. 31 (3): 383–392. Bibcode:2019HBio...31..383P. doi:10.1080/08912963.2017.1374383. ISSN 0891-2963. S2CID 90915135.
  21. ^ Ferreira, Gabriel S.; Bandyopadhyay, Saswati; Joyce, Walter G. (2018-11-15). "A taxonomic reassessment of Piramys auffenbergi, a neglected turtle from the late Miocene of Piram Island, Gujarat, India". PeerJ. 6: e5938. doi:10.7717/peerj.5938. ISSN 2167-8359. PMC 6240434. PMID 30479901.
  22. ^ Ferreira, G. S.; Nascimento, E. R.; Cadena, E. A.; Cozzuol, M. A.; Farina, B. M.; Pacheco, M. L. A. F.; Rizzutto, M. A.; Langer, M. C. (March 2024). "The latest freshwater giants: a new Peltocephalus (Pleurodira: Podocnemididae) turtle from the Late Pleistocene of the Brazilian Amazon". Biology Letters. 20 (3). doi:10.1098/rsbl.2024.0010. ISSN 1744-957X. PMC 10932709. PMID 38471564.