Extinct family of turtles
Bothremydidae is an extinct family of side-necked turtles (Pleurodira) known from the Cretaceous and Cenozoic . 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 and possibly herbivorous grazing diets,[ 1] [ 2] suction feeding .[ 2] [ 1] Cenomanian ).[ 3] Miocene .[ 4]
Taxonomy
The family is split into two subfamilies and a number of tribes.[ 5]
Bothremydidae
Pleurochayah appalachius Adrian et al., 2021 Woodbine Formation , Texas, Late Cretaceous (Cenomanian )"Chrysemys" montolivensis Roman 1897 Montoulieu, France, Oligocene (Rupelian )Sindhochelys ragei Lapparent de broin et al., 2021 Khadro Formation , Pakistan, Paleocene (Danian )Apertotemporalis Bahariya Formation , Egypt, Late Cretaceous (Cenomanian )Subfamily Bothremydinae Gaffney et al., 2006
Abalakemys Farin-Doutchi Formation , Niger, Late Cretaceous (Maastrichtian )Tribe Bothremydini Gaffney et al., 2006
Palauchelys López-Conde et al. 2021 Olmos Formation , Mexico, Late Cretaceous (Campanian )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
Araiochelys Gaffney et al., 2006 Ouled Abdoun Basin , Morocco, Paleocene
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 )Bothremys cooki Leidy, 1865 Merchantville Formation , Navesink Formation , New Jersey, Tar Heel Formation , North Carolina, Late Cretaceous (Campanian-Maastrichtian )Bothremys kellyi Gaffney et al., 2006 Ouled Abdoun Basin , Morocco, PaleoceneBothremys maghrebiana Gaffney et al., 2006 Ouled Abdoun Basin, Morocco, PaleoceneChedighaii 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
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 Polysternon Portis, 1882 Argiles et Grès à Reptiles Formation , Rognacian Formation , France, Vitoria Formation , Laño , Tremp Formation , Sierra Perenchiza Formation , Spain, Late Cretaceous (Campanian-Maastrichtian)
Polysternon provinciale (Matheron, 1869) Puentemys Cadena et al., 2012 Cerrejón Formation , 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
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, PaleoceneTaphrosphys dares Hay, 1908 (nomen dubium )Taphrosphys ippolitoi Gaffney et al., 2006 Ouled Abdoun Basin, Morocco, PaleoceneTaphrosphys 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 )
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 )
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:[ 8]
References
^ a b
Joyce, WG; Lyson, TR; Kirkland, JI (September 28, 2016). "An early bothremydid (Testudines, Pleurodira) from the Late Cretaceous (Cenomanian) of Utah, North America" . PeerJ . 4 : e2502. doi :10.7717/peerj.2502 PMC 5045886 PMID 27703852 .
^ a b Hermanson, Guilherme; Benson, Roger B. J.; Farina, Bruna M.; Ferreira, Gabriel S.; Langer, Max C.; Evers, Serjoscha W. (November 2022). "Cranial ecomorphology of turtles and neck retraction as a possible trigger of ecological diversification" . Evolution . 76 (11): 2566– 2586. doi :10.1111/evo.14629 . ISSN 0014-3820 . PMC 9828723 PMID 36117268 . ^ Pérez-García, A.; Antunes, M.T.; Barroso-Barcenilla, F.; Callapez, P.M.; Segura, M.; Soares, A.F.; Torices, A. (October 2017). "A bothremydid from the middle Cenomanian of Portugal identified as one of the oldest pleurodiran turtles in Laurasia" . Cretaceous Research . 78 : 61– 70. Bibcode :2017CrRes..78...61P . doi :10.1016/j.cretres.2017.05.031 . ^ de Lapparent de Broin, France; Murelaga, Xabier; Pérez-García, Adán; Farrés, Francesc; Altimiras, Jacint (2018-09-28). "Supplementary information:The turtles from the upper Eocene, Osona County (Ebro Basin, Catalonia, Spain): new material and its faunistic and environmental context" . Fossil Record . 21 (2): 237– 284. doi :10.5194/fr-21-237-2018 ISSN 2193-0066 . S2CID 55783731 . ^ The Paleontology Database Bothremydidae entry accessed on 26 January 2011^ Pérez-García, Adán (2016). "A new turtle taxon (Podocnemidoidea, Bothremydidae) reveals the oldest known dispersal event of the crown Pleurodira from Gondwana to Laurasia". Journal of Systematic Palaeontology . 15 (9): 709– 731. doi :10.1080/14772019.2016.1228549 . S2CID 88840423 . Pan-Pleurodira is one of the two clades of extant turtles (i.e. Testudines). Its crown group, Pleurodira, has a Gondwanan origin being known from the Barremian. Cretaceous turtle fauna of Gondwana was composed almost exclusively of pleurodires. Extant pleurodires live in relatively warm regions, with a geographical distribution restricted to tropical regions that were part of Gondwana. ^ Pérez-García, Adán (2018-08-03). "New insights on the only bothremydid turtle (Pleurodira) identified in the British record: Palemys bowerbankii new combination" . Palaeontologia Electronica . 21 (2): 1– 12. doi :10.26879/849 ISSN 1094-8074 . ^
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 family Bothremydidae is a large and diverse group extending from the Albian to the Eocene in North and South America, Europe, Africa, and India. Its monophyly is supported by the presence of a wide exoccipital-quadrate contact, a eustachian tube separated from the incisura columellae auris usually by bone to form a bony canal for the stapes, absence of a fossa precolumellaris, a supraoccipital-quadrate contact (except in the tribe Taphrosphyini), and a posterior enlargement of the fossa orbitalis.
