Bothremydidae is an extinct

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

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 ...

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 configu ...

. They are closely related to Podocnemididae, 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 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 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 t ...

( Rupelian) *'' Sindhochelys ragei'' Lapparent de broin et al., 2021

Khadro Formation Khadro ( ur, کھڈرو کڏڙو, Sindhi) is a town that is located in Sub-District Sinjhoro, Sanghar District of Sindh Province Sindh (; ; ur, , ; historically romanized as Sind) is one of the four provinces of Pakistan. Located in th ...

, Pakistan,

Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...

(

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 Cretac ...

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

Amminadav Formation The Amminadav 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 fo ...

, 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 Nac ...

, 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 ...

, Late Cretaceous (Cenomanian) *****''Algorachelus tibert'' (Joyce et al. 2016) Naturita Formation, 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 The Oulad Abdoun Basin (also known as the Ouled Abdoun Basin or Khouribga Basin) is a phosphate sediment ...

'' Gaffney et al., 2006 Ouled Abdoun Basin, Morocco,

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

Bothremys ''Bothremys'' is an extinct genus of bothremydid pleurodiran turtle that was discovered near Gloucester, New Jersey.Leidy, J., 1865 "Cretaceous reptiles of the United States." Smithsonian Contributions to Knowledge 192:1-135 https://doi.org ...

'' 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 (geology), 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 ...

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 Navesi ...

, New Jersey, Tar Heel Formation, 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 inte ...

) ***** ''Bothremys kellyi'' Gaffney et al., 2006 Ouled Abdoun Basin, 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, New Jersey, Tar Heel Formation, North Carolina, Late Cretaceous (Campanian) ***** ''Chedighaii hutchisoni'' Gaffney et al., 2006

Cerro del Pueblo Formation The Cerro del Pueblo Formation is a geological formation in Coahuila, Mexico whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel et al., 2004, pp.574-588 T ...

, 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, Tar Heel Formation, 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 li ...

, 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'' Tong et al., 1998 Csehbánya Formation, Hungary, Late Cretaceous (Santonian) Argiles et Grès à Reptiles Formation, Marnes d'Auzas Formation, Marnes Rouges Inferieures Formation, Lestaillats Marls Formation, France, Late Cretaceous (Campanian-Maastrichtian), Villalba de la Sierra Formation, Spain, Late Cretaceous (Campanian) ***** ''Foxemys mechinorum'' Tong et al., 1998 ****'' Palemys'' Gray, 1870 *****''Palemys bowerbankii'' (Owen, 1842)

London Clay Formation The London Clay Formation is a marine geological formation of Ypresian (early Eocene Epoch, c. 56–49 million years ago) age which crops out in the southeast of England. The London Clay is well known for its fossil content. The fossils from ...

, England,

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

(

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

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

Laño Laño ( eu, Lañu) is a hamlet and '' concejo'' (a small administrative subdivision) in Condado de Treviño within the Treviño enclave; which is administratively part of the Spanish province of Burgos, but which is completely surrounded by the ...

, Tremp Formation, Sierra Perenchiza 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'' Pérez-García et al. 2012 Vegas de Matute Formation, Villalba de la Sierra Formation, Spain, Campanian Marnes d'Auzas Formation, France, Maastrichtian *****'' Iberoccitanemys'' ''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'' 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, 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'' Gaffney et al., Santana Formation, 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 ...

) **** ''Cearachelys placidoi'' Gaffney et al., 2001 *** '' Elkanemys'' Maniel, de la Fuente and Canale, 2021

Candeleros Formation The Candeleros Formation (formerly known as the Candeleros Member of the "Río Limay Formation") is a geologic formation that crops out in the Río Negro, Neuquén, and Mendoza provinces of northern Patagonia, Argentina. It is the oldest formati ...

, Argentina, Late Cretaceous (Cenomanian) **** ''Elkanemys pritchardi'' Maniel, de la Fuente and Canale, 2021 *** '' Itapecuruemys'' 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 formation ...

, Brasil, Early Cretaceous (

Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ag ...

) **** ''Itapecuruemys amazoniensis'' Batista et al., 2020 *** '' Galianemys'' 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, India, Late Cretaceous (Maastrichtian) **** ''Jainemys pisdurensis'' Joyce & Bandyopadhyay, 2020 *** '' Kinkonychelys'' Gaffney et al., 2009 Maevarano Formation, 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