Bothremydidae is an extinct

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

of side-necked turtles (Pleurodira) known from the

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...

and

Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...

. 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,

piscivorous A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted rept ...

and possibly herbivorous grazing diets, with some probably capable of

suction feeding Aquatic feeding mechanisms face a special difficulty as compared to feeding on land, because the density of water is about the same as that of the prey, so the prey tends to be pushed away when the mouth is closed. This problem was first identifi ...

. Unlike modern pleurodires, which are exclusively freshwater, bothremydids inhabited freshwater, marine and coastal environments. 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 International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...

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

Miocene The Miocene ( ) is the first epoch (geology), 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 mea ...

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 (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...

(

Rupelian The Rupelian is, in the geologic timescale, the older of two age (geology), ages or the lower of two stage (stratigraphy), stages of the Oligocene epoch (geology), Epoch/series (stratigraphy), Series. It spans the time between . It is preceded b ...

) *'' 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), ...

(

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 ...

) *'' Apertotemporalis'' Stromer, 1934 Bahariya Formation, Egypt, Late Cretaceous (

) * Subfamily Bothremydinae Gaffney et al., 2006 ** '' Abalakemys'' Pérez-García, 2023 Farin-Doutchi Formation, Niger, Late Cretaceous (

Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...

) ** 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. Campa ...

) **'' Khargachelys'' AbdelGawad et al. 2021 Quseir Formation, Egypt, 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, Amminadav Formation, Palestine, Late Cretaceous (Cenomanian) *****''Algorachelus peregrinus'' Pérez-García 2017 Utrillas Formation, Spain, Nazaré, Portugal, 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'' Gaffney et al., 2006 Ouled Abdoun Basin, Morocco,

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

) ***** ''Bothremys cooki'' Leidy, 1865 Merchantville Formation, Navesink Formation, New Jersey, Tar Heel Formation, North Carolina, Late Cretaceous (Campanian-

) ***** ''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, Mexico, Late Cretaceous (Campanian) Kirtland Formation, 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, Brasil, Paleocene ***** '' Inaechelys pernambucensis'' Carvalho, 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 Marnes is the name or part of the name of three communes of France: * Marnes, Deux-Sèvres in the Deux-Sèvres ''département'' * Marnes-la-Coquette in the Hauts-de-Seine ''département'' * Saint-Jouin-de-Marnes in the Deux-Sèvres ''départemen ...

, 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, England,

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

(

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 Laño () 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 territo ...

, 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 Department, La Guajira and Cesar Department, Cesar. The fo ...

, 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 (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...

) **** ''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'' Batista et al., 2020 Itapecuru Formation, Brasil, Early Cretaceous (

Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...

) **** ''Itapecuruemys amazoniensis'' Batista et al., 2020 *** '' Galianemys'' Gaffney et al., 2002 Kem Kem Group, 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, 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 †