Bothremydidae is an extinct

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

side-necked turtles 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 differen ...

(Pleurodira) known from the

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...

and

Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...

. They are closely related to

Podocnemididae Podocnemididae is a family of pleurodire (side-necked) turtles, once widely distributed. Most of its 20 genera and 30 species are now extinct. Seven of its eight surviving species are native to South America: the genus ''Peltocephalus'', with onl ...

, and are amongst the most widely distributed pleurodire groups, with their fossils having been found in Africa, India, the Middle East, Europe, North America and South America. Bothremydids were aquatic turtles with a high morphological diversity, indicative of generalist, molluscivorous, and

piscivorous A piscivore () is a carnivorous animal that eats primarily fish. The name ''piscivore'' is derived . Piscivore is equivalent to the Greek-derived word ichthyophage, both of which mean "fish eater". Fish were the diet of early tetrapod evoluti ...

diets. Unlike modern pleurodires, which are exclusively freshwater, bothremydids inhabited freshwater, marine and coastal settings. Their marine habits allowed bothremydids to disperse across oceanic barriers into Europe and North America during the early Late Cretaceous (

Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...

). The youngest records of the group are indeterminate remains from Saudi Arabia and Oman, dating to the

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

Taxonomy

The family is split into two subfamilies and a number of tribes. Bothremydidae *'' Pleurochayah appalachius'' Adrian et al., 2021

Woodbine Formation The Woodbine Group is a geological formation in east Texas whose strata date back to the Early to Middle Cenomanian age of the Late Cretaceous. It is the producing formation of the giant East Texas Oil Field (also known as the "Black Giant") fr ...

, Texas, Late Cretaceous (

) *''"Chrysemys" montolivensis'' Roman 1897 Montoulieu, France,

Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...

(

Rupelian The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/ Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chattian ...

) *'' Sindhochelys ragei'' Lapparent de broin et al., 2021 Khadro Formation, Pakistan,

Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...

(

Danian The Danian is the oldest age or lowest stage of the Paleocene Epoch or Series, of the Paleogene Period or System, and of the Cenozoic Era or Erathem. The beginning of the Danian (and the end of the preceding Maastrichtian) is at the Cretace ...

) * Subfamily Bothremydinae Gaffney et al., 2006 ** Tribe Bothremydini Gaffney et al., 2006 **'' Palauchelys'' López-Conde et al. 2021 Olmos Formation, Mexico, Late Cretaceous (

Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campanian s ...

) *** Subtribe Bothremydina Gaffney et al., 2006 **** '' Akoranemys'' Pérez-García, 2021. Madagascar, Late Cretaceous (Cenomanian) ****'' Algorachelus'' Pérez-García 2017 *****''Algorachelus parvus'' (Haas 1978)

Bet-Meir Formation The Bet-Meir Formation is a Mesozoic geologic formation in Israel. Pterosaur fossils have been recovered from the formation. See also * List of pterosaur-bearing stratigraphic units This is a list of stratigraphic units, where pterosaur fossi ...

Amminadav Formation The Amminadav Formation is a Mesozoic geologic Formation (geology), formation in Israel. Pterosaur fossils have been recovered from the formation. See also * List of pterosaur-bearing stratigraphic units References

Geologic formations of ...

, Palestine, Late Cretaceous (Cenomanian) *****''Algorachelus peregrinus'' Pérez-García 2017

Utrillas Formation Utrillas is a municipality located in the province of Teruel, Aragon, Spain. According to the 2004 census (INE INE, Ine or ine may refer to: Institutions * Institut für Nukleare Entsorgung, a German nuclear research center * Instituto Nacion ...

, Spain,

Nazaré, Portugal Nazaré () is a Portuguese town and municipality located in the Oeste region, in the historical province of Estremadura, and in the Leiria District. The municipality has a population of 14,889 in an area of 82.43 km2, while the town itself h ...

, Late Cretaceous (Cenomanian) *****''Algorachelus tibert'' (Joyce et al. 2016)

Naturita Formation The Naturita Formation is a classification used in western Colorado and eastern Utah for a Cretaceous Period sedimentary geologic formation. This name was "applied to the upper or carbonaceous part of Dakota Group" by R.G. Young in 1960, naming ...

, Utah, Late Cretaceous (Cenomanian) (alternatively considered the only member of the genus '' Paiutemys'') ****''

Araiochelys ''Araiochelys'' ("narrow turtle") is an extinct genus of bothremydid pleurodiran turtle that was discovered in the Ouled Abdoun Basin, Morocco.E. S. Gaffney, H. Tong, and P. A. Meylan. 2006. Evolution of the side-necked turtles: The families B ...

'' Gaffney et al., 2006

Ouled Abdoun Basin The Oulad Abdoun Basin (also known as the Ouled Abdoun Basin or Khouribga Basin) is a phosphate sedimentary basin located in Morocco, near the city of Khouribga. It is the largest in Morocco, comprising 44% of Morocco's phosphate reserves, and a ...

, Morocco,

***** ''Araiochelys hirayamai'' Gaffney et al., 2006 **** ''

Bothremys ''Bothremys'' is an extinct genus of bothremydid pleurodiran turtle that was discovered near Gloucester, New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north ...

'' Leidy, 1865 (jr synonym: ''Karkaemys'') ***** ''Bothremys arabicus'' (Zalmout et al., 2005) Wadi Umm Ghudran Formation, Jordan, Late Cretaceous (

Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. ...

) ***** ''Bothremys cooki'' Leidy, 1865

Merchantville Formation The Merchantville Formation is a geological formation in the northeastern United States whose strata date back to the Late Cretaceous, around the time of the Santonian and Campanian age. Dinosaur remains are among the fossils that have been recove ...

Navesink Formation The Navesink Formation is a 66 to 70 mya greensand glauconitic marl and sand geological formation in New Jersey. It is known for its Cretaceous period fossil shell beds and dinosaur bones. Description The Navesink Formation, named after Navesin ...

, New Jersey,

Tar Heel Formation The Tar Heel Formation is a geologic formation in North Carolina. It preserves fossils, including amber dating back to the Cretaceous period. A locality known as Phoebus Landing, has been dated to 78.5-77.1 Ma. Fish Cartilaginous fish Di ...

, North Carolina, Late Cretaceous (Campanian-

Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval ...

) ***** ''Bothremys kellyi'' Gaffney et al., 2006

, Morocco, Paleocene ***** ''Bothremys maghrebiana'' Gaffney et al., 2006 Ouled Abdoun Basin, Morocco, Paleocene **** '' Chedighaii'' Gaffney et al., 2006 ***** ''Chedighaii barberi'' (Schmidt, 1940) Brownstone Marl, Arkansas, Blufftown Formation, Georgia,

Marshalltown Formation The Marshalltown Formation is a Mesozoic geologic formation. Dinosaur remains diagnostic to the genus level are among the fossils that have been recovered from the formation.Weishampel, et al. (2004). "Dinosaur distribution." Pp. 517-607. Dinosau ...

, New Jersey,

, North Carolina, Late Cretaceous (Campanian) ***** ''Chedighaii hutchisoni'' Gaffney et al., 2006 Cerro del Pueblo Formation, Mexico, Late Cretaceous (Campanian)

Kirtland Formation The Kirtland Formation (originally the Kirtland Shale) is a sedimentary geological formation. Description The Kirtland Formation is the product of alluvial muds and overbank sand deposits from the many channels draining the coastal plain th ...

, New Mexico,

, North Carolina, Late Cretaceous (Campanian) **** '' Rosasia'' Costa, 1940 Argilas de Aveiro Formation, Portugal, Late Cretaceous (Campanian-Maastrichtian) ***** ''Rosasia soutoi'' Carrington da Costa, 1940 **** '' Inaechelys'' Carvalho, 2016

Maria Farinha Formation The Maria Farinha Formation is a geological formation of the Parnaíba Basin in Pernambuco, northeastern Brazil whose strata date back to the Danian stage of the Paleocene, or Tiupampan in the SALMA classification. The formation comprising lim ...

, Brasil, Paleocene ***** '' Inaechelys pernambucensis'' Caralho, 2016 **** '' Zolhafah'' Lapparent de Broin and Werner, 1998 Dakhla Formation, Egypt, Late Cretaceous (Maastrichtian) ***** ''Zolhafah bella'' Lapparent de Broin and Werner, 1998 *** Subtribe Foxemydina Gaffney et al., 2006 **** '' Elochelys'' Nopsca, 1931 France, Spain, Late Cretaceous (Campanian-Maastrichtian) ***** ''Elochelys perfecta'' Nopsca, 1931 **** ''

Foxemys ''Foxemys'' is an extinct genus of bothremydid turtle that was discovered at Fox Amphoux, France and also Hungary and Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , nati ...

'' Tong et al., 1998

Csehbánya Formation The Csehbánya Formation is a geological formation in the Transdanubian Mountains of Veszprém County, Hungary. The formation dates to the Late Santonian (around 85-84 million years ago) of the Late Cretaceous. It represents a floodplain environ ...

, Hungary, Late Cretaceous (Santonian) Argiles et Grès à Reptiles Formation,

Marnes d'Auzas Formation The Marnes d’Auzas Formation (Auzas marls) is a geological Formation in southwestern France (departments of Ariège and Haute-Garonne) whose strata date back to the Late Maastrichtian. It is about 100 metres thick and consists primarily of mar ...

, Marnes Rouges Inferieures Formation, Lestaillats Marls Formation, France, Late Cretaceous (Campanian-Maastrichtian),

Villalba de la Sierra Formation The Villalba de la Sierra Formation is a Campanian to Maastrichtian geologic Formation (geology), formation in Spain. Fossil dinosaur eggs have been reported from the formation, that comprises gypsum, gypsiferous, grey, argillaceous mudstones and ...

, Spain, Late Cretaceous (Campanian) ***** ''Foxemys mechinorum'' Tong et al., 1998 ****'' Palemys'' Gray, 1870 *****''Palemys bowerbankii'' (Owen, 1842) London Clay Formation, England,

Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...

(

Ypresian In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...

) **** '' Polysternon'' Portis, 1882 Argiles et Grès à Reptiles Formation, Rognacian Formation, France, Vitoria Formation, Laño,

Tremp Formation The Tremp Formation ( es, Formación de Tremp, ca, Formació de Tremp), alternatively described as Tremp Group ( es, Grupo Tremp), is a geological Formation (geology), formation in the comarca Pallars Jussà, Lleida Province, Lleida, Spain. Th ...

Sierra Perenchiza Formation The Sierra Perenchiza Formation is a late Campanian to late Maastrichtian geologic formation in Spain. Dinosaur, pterosaur, turtle, lissamphibian, frog and albanerpetontid remains are among the fossils that have been recovered from the formation, ...

, Spain, Late Cretaceous (Campanian-Maastrichtian) ***** ''Polysternon provinciale'' (Matheron, 1869) **** '' Puentemys'' Cadena et al., 2012

Cerrejón Formation The Cerrejón Formation is a geologic formation in Colombia dating back to the Middle-Late Paleocene. It is found in the El Cerrejón sub-basin of the Cesar-Ranchería Basin of La Guajira and Cesar. The formation consists of bituminous coal fie ...

, Colombia, Paleocene ***** ''Puentemys mushaisaensis'' Cadena et al., 2012 ****'' Tartaruscola'' Pérez-García, 2016 Saint-Papoul, France, Eocene (Ypresian) ****''

Iberoccitanemys ''Iberoccitanemys'' (meaning "Iberoccitanian turtle") is an extinct genus of turtle from the Bothremydidae that is known from the Marnes d'Auzas Formation in France, and also the Sobrepena Formation and Villalba de la Sierra Formation in Spain. ...

'' Pérez-García et al. 2012 Vegas de Matute Formation,

, Spain, Campanian Marnes d'Auzas Formation, France, Maastrichtian *****''

'' ''atlanticum'' (Lapparent de Broin and Murelaga, 1996 originally ''Polysternon atlanticum'') ''syn Iberoccitanemys convenarum'' (Laurent et al., 2002) *** Tribe Taphrosphyini Gaffney et al., 2006 ***** '' Ilatardia'' Pérez-García, 2019 Farin Doutchi Formation, Niger, Late Cretaceous (Maastrichtian) **** Subtribe Nigermydina Gaffney et al., 2006 ***** '' Arenila'' Lapparent de Broin and Werner, 1998 Dakhla Formation, Egypt, Late Cretaceous (Maastrichtian) ****** ''Arenila krebsi'' Lapparent de Broin and Werner, 1998 **** Subtribe Taphrosphyina Gaffney et al., 2006 ***** '' Azabbaremys'' Gaffney et al., 2001 Teberemt Formation, Mali, Paleocene ****** ''Azabbaremys moragjonesi'' Gaffney et al., 2001 ***** '' Eotaphrosphys'' Perez-Garcia, 2018 Mont-Aimé, France, Late Cretaceous (Maastrichtian) ****** ''Eotaphrosphys ambiguum'' (Gaudry, 1890) ***** '' Labrostochelys'' Gaffney et al., 2006 Ouled Abdoun Basin, Morocco, Paleocene ****** ''Labrostochelys galkini'' Gaffney et al., 2006 ***** '' Motelomama'' Perez-Garcia, 2018 Salina Group, Peru, Eocene (Ypresian) ****** ''Motelomama olssoni'' (Schmidt, 1931) ***** ''

Phosphatochelys ''Phosphatochelys'' ("phosphate turtle") is an extinct genus of bothremydid pleurodiran turtle that was discovered near Oued Zem, Morocco. E. S. Gaffney and H. Tong. 2003. Phosphatochelys, a new side-necked turtle (Pelomedusoides: Bothremydidae ...

'' Gaffney and Tong, 2003 Ouled Abdoun Basin, Morocco, Paleocene ****** ''Phosphatochelys tedfordi'' Gaffney and Tong, 2003 ***** '' Rhothonemys'' Gaffney et al., 2006 Ouled Abdoun Basin, Morocco, Paleocene ****** ''Rhothonemys brinkmani'' Gaffney et al., 2006 ***** '' Taphrosphys'' Cope, 1869 (jr synonyms: ''Amblypeza'', ''Bantuchelys'', ''Prochonias'') ****** ''Taphrosphys congolensis'' (Dollo, 1912) Landana Formation, Angola, Paleocene ****** ''Taphrosphys dares'' Hay, 1908 (''nomen dubium'') ****** ''Taphrosphys ippolitoi'' Gaffney et al., 2006 Ouled Abdoun Basin, Morocco, Paleocene ****** ''Taphrosphys sulcatus'' (Leidy, 1856)

Hornerstown Formation The Hornerstown Formation is a Paleogene or latest Mesozoic geologic Formation (geology), formation in New Jersey.Weishampel, et al. (2004). "Dinosaur distribution." Pp. 517-607. The age of these deposits have been controversial. While most fossi ...

, New Egypt Formation, New Jersey, Late Cretaceous (Maastrichtian) ***** '' Ummulisani'' Gaffney et al., 2006 Morocco, Eocene (Ypresian) ****** ''Ummulisani rutgersensis'' Gaffney et al., 2006 ***** '' Crassachelys'' Bergounioux, 1952 (''nomen dubium'') ****** ''Crassachelys neurirregularis'' Bergounioux, 1952 ***** '' Eusarkia'' Bergounioux, 1952 (''nomen dubium'') ****** ''Eusarkia rotundiformis'' Bergounioux, 1952 ***** '' Gafsachelys'' Stefano, 1903 (''nomen dubium'') ****** ''Gafsachelys phosphatica'' Stefano, 1903 ****** ''Gafsachelys moularensis'' Bergounioux, 1955 ** Tribe Cearachelyini Gaffney et al., 2006 *** ''

Cearachelys ''Cearachelys'' is an extinct genus of pleurodiran turtle which existed some 110 million years ago. The genus is monotypic, with only type species ''Cearachelys placidoi'' known. Etymology The genus was named for Ceará, the Brazilian state ...

'' Gaffney et al.,

Santana Formation The Santana Group is a geologic group, formerly included as the middle part of the Araripe Group, in the Araripe Basin of northeastern Brazil. The group comprises the Crato, Ipubi and Romualdo Formations and is dated to the Aptian to Albian sta ...

, Brazil, Early Cretaceous (

Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0.9 M ...

) **** ''Cearachelys placidoi'' Gaffney et al., 2001 *** '' Elkanemys'' Maniel, de la Fuente and Canale, 2021 Candeleros Formation, Argentina, Late Cretaceous (Cenomanian) **** ''Elkanemys pritchardi'' Maniel, de la Fuente and Canale, 2021 *** ''

Itapecuruemys ''Itapecuruemys'' ("Itapecuru turtle") is an extinct genus of bothremydid pleurodiran turtle that was discovered in the Itapecuru Formation of Brazil.Batista, Diego L.; de Sousa Carvalho, I.; de la Fuente, M.S. (2020). "A new Cretaceous (Pleur ...

'' Batista et al., 2020

Itapecuru Formation The Itapecuru Formation is a geological formation in Itapecuru Mirim, Maranhão, Brazil. It was formed during the Aptian to Albian stages of the Early Cretaceous. Dinosaur remains are among the fossils that have been recovered from the format ...

, Brasil, Early Cretaceous ( Aptian) **** ''Itapecuruemys amazoniensis'' Batista et al., 2020 *** ''

Galianemys ''Galianemys'' is an extinct genus of turtle in the family Bothremydidae, discovered in the Kem Kem Beds The Kem Kem Group (commonly known as the Kem Kem beds) is a geological group in the Kem Kem region of eastern Morocco, whose strata date ba ...

'' Gaffney et al., 2002

Kem Kem Group The Kem Kem Group (commonly known as the Kem Kem beds) is a geological group in the Kem Kem region of eastern Morocco, whose strata date back to the Cenomanian stage of the Late Cretaceous. Its strata are subdivided into two geological formations ...

, Morocco, Late Cretaceous (Cenomanian) **** ''Galianemys emringeri'' Gaffney et al., 2002 **** ''Galianemys whitei'' Gaffney et al., 2002 * Subfamily Kurmademydinae Gaffney et al., 2006 ** Tribe Kurmademydini Gaffney et al., 2006 *** '' Jainemys'' Joyce & Bandyopadhyay, 2020

Lameta Formation The Lameta Formation, also known as the Infratrappean Beds, is a sedimentary geological formation found in Madhya Pradesh, Gujarat, and Maharashtra, India, associated with the Deccan Traps. It is of Maastrichtian age (Late Cretaceous), and is nota ...

, India, Late Cretaceous (Maastrichtian) **** ''Jainemys pisdurensis'' Joyce & Bandyopadhyay, 2020 *** '' Kinkonychelys'' Gaffney et al., 2009

Maevarano Formation The Maevarano Formation is a Late Cretaceous sedimentary rock formation found in the Mahajanga Province of northwestern Madagascar. It is most likely Maastrichtian in age, and records a seasonal, semiarid environment with rivers that had greatly v ...

, Madagascar, Late Cretaceous (Maastrichtian) **** ''Kinkonychelys rogersi'' Gaffney et al., 2009 *** '' Kurmademys'' Gaffney et al., 2001 Kallamedu Formation, India, Late Cretaceous (Maastrichtian) **** ''Kurmademys kallamedensis'' Gaffney et al., 2001 *** '' Sankuchemys'' Gaffney et al., 2003

Intertrappean Beds The Intertrappean Beds are a Late Cretaceous geologic formation in India. The beds are found as interbeds between Deccan Traps layers. Indeterminate theropod and pterosaur remains have been recovered from the formation, as well as dinosaur eggs ...

, India, Late Cretaceous (Maastrichtian) **** ''Sankuchemys sethnai'' Gaffney et al., 2003

Phylogeny

Below is a cladogram by Gaffney et al. in 2006:

References

{{Testudines Extinct turtles Prehistoric reptile families