Turtles are reptiles of the order Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira and Cryptodira, which differ in the way the head retracts. There are 360 living and recently extinct species of turtles, including land-dwelling tortoises and freshwater terrapins. They are found on most continents, some islands and, in the case of sea turtles, much of the ocean. Like other amniotes they breathe air and do not lay eggs underwater, although many species live in or around water.
The Chelydridae is a family of turtles that has seven extinct and two extant genera. The extant genera are the snapping turtles, Chelydra and Macrochelys. Both are endemic to the Western Hemisphere. The extinct genera are Acherontemys, Chelydrops, Chelydropsis, Emarginachelys, Macrocephalochelys, Planiplastron, and Protochelydra.
Emydidae is a family of testudines (turtles) that includes close to 50 species in 10 genera. Members of this family are commonly called terrapins, pond turtles, or marsh turtles. Several species of Asian box turtles were formerly classified in the family; however, revised taxonomy has separated them to a different family (Geoemydidae). As currently defined, the Emydidae are entirely a Western Hemisphere family, with the exception of two species of pond turtle.
Chelidae is one of three living families of the turtle suborder Pleurodira, and are commonly called Austro-South American side-neck turtles. The family is distributed in Australia, New Guinea, parts of Indonesia, and throughout most of South America. It is a large family of turtles with a significant fossil history dating back to the Cretaceous. The family is entirely Gondwanan in origin, with no members found outside Gondwana, either in the present day or as a fossil.
Kinosternoidea is a superfamily of aquatic turtles, which includes two families: Dermatemydidae, and Kinosternidae.
Trionychia is a superfamily of turtles which encompasses the species that are commonly referred to as softshelled turtles as well as some others. The group contains two families, Carettochelyidae, which has only one living species, the pig-nosed turtle native to New Guinea and Northern Australia, and Trionychidae, the softshelled turtles, containing numerous species native to Asia, North America and Africa. These families likely diverged during the late Jurassic. The oldest known stem-trionychian is Sinaspideretes from the Late Jurassic of China.
The Pleurodira are one of the two living suborders of turtles, the other being the Cryptodira. The division between these two suborders represents a very deep evolutionary divide between two very different types of turtles. The physical differences between them, although anatomical and largely internal, are nonetheless significant, and the zoogeographic implications of them are substantial. The Pleurodira are known more commonly as the side-necked turtles and the name Pleurodira quite literally translates to side neck, whereas the Cryptodira are known as hidden-necked turtles. The Pleurodira turtles are currently restricted to freshwater habitats in the Southern Hemisphere, largely to Australia, South America, and Africa. Within the Pleurodira, three living families are represented: Chelidae, also known as the Austro-South American side-necked turtles, the Pelomedusidae, also known as the African mud terrapins, and the Podocnemididae, also known as the American side-neck river turtles. However, they were cosmopolitan clade during the Cretaceous and most of the Cenozoic, and even occurred in marine environments around the world.
Meiolaniidae is an extinct family of large, probably herbivorous stem-group turtles with heavily armored heads and clubbed tails known from South America and Australasia. Though once believed to be cryptodires, they are not closely related to any living species of turtle, and lie outside crown group Testudines, having diverged from them around the Middle Jurassic. They are best known from the last surviving genus, Meiolania, which lived in Australia from the Miocene until the Pleistocene, and insular species that lived on Lord Howe Island and New Caledonia during the Pleistocene and possibly the Holocene for the latter. Meiolaniids are part of the broader grouping of Meiolaniformes, which contains more primitive turtles species lacking the distinctive morphology of meiolaniids, known from the Early Cretaceous-Paleocene of South America and Australia.
Testudinoidea is a superfamily within the suborder Cryptodira of the order Testudines. It includes the pond turtles, Asian turtles, the monotypic big-headed turtle, and the tortoises.
Paracryptodira is an extinct group of reptiles in the clade Testudinata, known from the Jurassic to Paleogene of North America and Europe. Initially treated as a suborder sister to Cryptodira, they were then thought to be a very primitive lineage inside the Cryptodira according to the most common use of the latter taxon. They are now often regarded as late-diverging stem-turtles, lying outside the clade formed by Cryptodira and Pleurodira. Paracryptodires are divided into three main groups, Compsemydidae, known from the Late Jurassic to Paleocene of North America and Europe, Pleurosternidae, known from the Late Jurassic to Early Cretaceous of North America and Europe, and Baenidae, known from the Early Cretaceous to Eocene of North America. The latter two groups are more closely related to each other than to Compsemys, forming the clade Baenoidea.
Thalassemys is a genus of extinct thalassochelydian turtle from the Late Jurassic of western and central Europe. While the genus was originally named by Rütimeyer in 1859 for a large carapace and other associated fragments from the late Kimmeridgian of the Reuchenette Formation of Switzerland, although the taxon was not validly named until 1873 when Rütimeyer designated the type species T. hugii. Rütimeyer also named T. gresslyi from the Reunchenette Formation in the same paper as T. hugii, but it cannot be differentiated from the type material of T. hugii and is therefore a junior synonym. A large assemblage of shell and postcranial material from the Reunchenette was named as a species of Eurysternum, E. ignoratum, by Bräm in 1965. While originally distinguished based on the presence of fontanelles on the plastron, the feature was later identified on T. hugii and E. ignoratum was designated a junior synonym. Additional material from the Kimmeridge Clay of the United Kingdom has also been referred to T. hugii.
Kayentachelys is an extinct genus of turtle known only from the "silty facies" of the Lower Jurassic Kayenta Formation in northeastern Arizona on the lands of the Navajo Nation.
The Plesiochelyidae are an extinct family of turtles in the clade Thalassochelydia originally classified within the Cryptodira suborder, mostly belonging from the Jurassic period. An alternate study placed the clade Thalassochelydia in the Angolachelonia and outside the Testudines.
Cyrtura is a dubious genus of extinct Testudinata from the Late Jurassic of the Solnhofen Formation of Bavaria, Germany. Cyrtura was originally described as a temnospondyl amphibian by Otto Jaekel in 1904 on the basis of MNB 1890, the distal portion of a tail with 14 caudal vertebrae. Most authors overlooked the genus, although those who mentioned Cyrtura dismissed it as either undiagnostic or referable to Testudines rather than Temnospondyli. Warren and Hutchinson (1983) rejected the temnospondyl classification of the genus based on examination of a cast of the holotype, and subsequent studies showed that Cyrtura is actually a marine turtle, although the lack of diagnostic characters renders it a nomen dubium.
Americhelydia is a clade of turtles that consists of sea turtles, snapping turtles, the Central American river turtle and mud turtles, supported by several lines of molecular work. Prior to these studies some morphological and developmental work have considered sea turtles to be basal members of Cryptodira and kinosternids related to the trionychians in the clade Trionychoidea. Americhelydia and Testudinoidea, both clades within Durocryptodira, split a part during the early Cretaceous.
Perichelydia is a clade within Pantestudines known from the Middle Jurassic to Holocene. Alongside crown group Testudines, it also contains Helochelydridae, which is known from the Cretaceous of Europe and North America, Sichuanchelyidae from the Middle Jurassic to Paleocene of Asia and Europe, Meiolaniformes, which is known from the Cretaceous to Holocene of South America, Australia and Oceania, and Spoochelys, known from the Mid-Cretaceous Griman Creek Formation of Australia. Kallokibotion from the Late Cretaceous of Europe is also considered part of this group. Several other groups, including the proposed clade Angolachelonia, Paracryptodira, Macrobaenidae, Sinemydidae and Xinjiangchelyidae, which are sometimes considered members of Cryptodira, have also been found outside crown Testudines in several analyses. These groups are usually considered to be closer to the crown group than the other members of Perichelydia.
Thalassochelydia is a clade of extinct marine turtles from the Late Jurassic and earliest Cretaceous of Europe and South America. The group is defined as including Eurysternum, Plesiochelys and Thalassemys to the exclusion of Pelomedusa, Testudo and Protostega. While a clade uniting the families Eurysternidae, Plesiochelyidae and Thalassemydidae had been supported by phylogenetic evidence, a name was not given for the clade until 2017, when Jérémy Anquetin and colleagues coined Thalassochelydia.
Angolachelonia is a clade of extinct turtles from the Late Jurassic to Paleogene of Eurasia. The group is defined as all taxa derived from the ancestor of the type genus Angolachelys and Solnhofia, a definition that could potentially encompass a clade of entirely marine turtles. Angolachelonia was originally inclusive of only Solnhofia, Angolachelys and Sandownia when originally conceived by Octavio Mateus and colleagues in 2009, but later phylogenetic analyses by Serjocha Evers and Roger Benson in 2018 unites the family Sandownidae, including Angolachelys and Sandownia among other taxa, with the entirely Late Jurassic clade Thalassochelydia, where Solnhofia may be a basal member. While the placement of Solnhofia is weak and the clade that Angolachelonia represents may change with further analysis, the clade of Sandownidae and Thalassochelydia is well-supported, and does not collapse despite the uncertain evolutionary history of the group. Three alternative potential origins of Angolachelonia sensu Evers and Benson are shown below.
Platychelyidae is an extinct family of pan-pleurodiran turtles, known from the Late Jurassic and Early Cretaceous of Europe, South America, North America, and the Caribbean. It represents the oldest known clade of stem-pleurodires. All known members have been found in marine or coastal deposits. Despite this, their limb morphology suggests that they were not adapted for open marine conditions, but were likely inhabitants of shallow water environments, including brackish and saline waters, and they likely never inhabited environments more marine than lagoons. Their tolerance for saline environments likely aided their dispersal during the breakup of Pangea. Unlike modern pleurodires, which retract their necks to the sides, Platychelys retracted its neck inwards, similar to modern cryptodire turtles. Platychelys is strongly morphologically similar to mata mata and snapping turtles, suggesting that it had a similar ecology as a ram or suction feeder.
