Bothremydidae

{{Short description|Extinct family of turtles}} {{Automatic taxobox | image = Galianemys cf. emringeri.jpg | image_caption = ''Galianemys'' Skull | image2 = Taphrosphys sulcatus Carapace Fossil.jpg | image2_caption = Shell of ''Taphrosphys'' | fossil_range = {{fossilrange|Aptian|Miocene}} | taxon = Bothremydidae | authority = Baur, 1891 | subdivision_ranks = Subfamilies | subdivision = See text }}

'''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,<ref name="paiutemys" /><ref name=":0">{{Cite journal |last1=Hermanson |first1=Guilherme |last2=Benson |first2=Roger B. J. |last3=Farina |first3=Bruna M. |last4=Ferreira |first4=Gabriel S. |last5=Langer |first5=Max C. |last6=Evers |first6=Serjoscha W. |date=November 2022 |title=Cranial ecomorphology of turtles and neck retraction as a possible trigger of ecological diversification |journal=Evolution |language=en |volume=76 |issue=11 |pages=2566–2586 |doi=10.1111/evo.14629 |issn=0014-3820 |pmc=9828723 |pmid=36117268}}</ref> with some probably capable of suction feeding.<ref name=":0" /> Unlike modern pleurodires, which are exclusively freshwater, bothremydids inhabited freshwater, marine and coastal environments.<ref name="paiutemys"> {{cite journal|author1=Joyce, WG |author2=Lyson, TR |author3=Kirkland, JI |date=September 28, 2016|title=An early bothremydid (Testudines, Pleurodira) from the Late Cretaceous (Cenomanian) of Utah, North America|journal=PeerJ|volume=4 |article-number=e2502 |doi=10.7717/peerj.2502|pmc=5045886|pmid=27703852|doi-access=free }}</ref> Their marine habits allowed bothremydids to disperse across oceanic barriers into Europe and North America during the early Late Cretaceous (Cenomanian).<ref>{{Cite journal|last1=Pérez-García|first1=A.|last2=Antunes|first2=M.T.|last3=Barroso-Barcenilla|first3=F.|last4=Callapez|first4=P.M.|last5=Segura|first5=M.|last6=Soares|first6=A.F.|last7=Torices|first7=A.|date=October 2017|title=A bothremydid from the middle Cenomanian of Portugal identified as one of the oldest pleurodiran turtles in Laurasia|url=https://linkinghub.elsevier.com/retrieve/pii/S019566711730099X|journal=Cretaceous Research|language=en|volume=78|pages=61–70|doi=10.1016/j.cretres.2017.05.031|bibcode=2017CrRes..78...61P |url-access=subscription}}</ref> The youngest records of the group are indeterminate remains from Saudi Arabia and Oman, dating to the Miocene.<ref>{{Cite journal|last1=de Lapparent de Broin|first1=France|last2=Murelaga|first2=Xabier|last3=Pérez-García|first3=Adán|last4=Farrés|first4=Francesc|last5=Altimiras|first5=Jacint|date=2018-09-28|title=Supplementary information:The turtles from the upper Eocene, Osona County (Ebro Basin, Catalonia, Spain): new material and its faunistic and environmental context|url=https://fr.copernicus.org/articles/21/237/2018/|journal=Fossil Record|language=English|volume=21|issue=2|pages=237–284|doi=10.5194/fr-21-237-2018|s2cid=55783731 |issn=2193-0066|doi-access=free}}</ref>

== Taxonomy == The family is split into two subfamilies and a number of tribes.<ref>[http://pbdb.org/cgi-bin/bridge.pl?action=checkTaxonInfo&taxon_no=67318&is_real_user=0 The Paleontology Database Bothremydidae entry] accessed on 26 January 2011</ref>

'''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'' Stromer, 1934 Bahariya Formation, Egypt, Late Cretaceous (Cenomanian) * Subfamily '''Bothremydinae''' Gaffney et al., 2006 ** ''Abalakemys'' Pérez-García, 2023 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 *****''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)<ref name="2016PG">{{cite journal |author=Pérez-García, Adán |year=2016 |title=A new turtle taxon (Podocnemidoidea, Bothremydidae) reveals the oldest known dispersal event of the crown Pleurodira from Gondwana to Laurasia |journal=Journal of Systematic Palaeontology |volume=15 |issue=9 |pages=709–731 |doi=10.1080/14772019.2016.1228549 |s2cid=88840423|quote=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. }}</ref> *****''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, 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, 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, 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 (Ypresian)<ref>{{Cite journal|last=Pérez-García|first=Adán|date=2018-08-03|title=New insights on the only bothremydid turtle (Pleurodira) identified in the British record: Palemys bowerbankii new combination|url=https://palaeo-electronica.org/content/2018/2268-british-bothremydid-turtle|journal=Palaeontologia Electronica|language=English|volume=21|issue=2|pages=1–12|doi=10.26879/849|issn=1094-8074|doi-access=free}}</ref> **** ''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 ***** ''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) **** ''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:<ref name="gaffney2006"> {{cite journal |last1 = Gaffney|first1=E. S.|author-link= Eugene S. Gaffney |last2= Tong |first2= H.|last3= Meylan|first3=P. A.|title= Evolution of the side-necked turtles: The families Bothremydidae, Euraxemydidae, and Araripemydidae |journal= Bulletin of the American Museum of Natural History |volume= 300|pages= 1–700|publisher= American Museum of Natural History |location= New York |date= 2006-11-17| doi = 10.1206/0003-0090(2006)300[1:EOTSTT]2.0.CO;2|hdl=2246/5824|s2cid=85790134 |url=http://doc.rero.ch/record/15686/files/PAL_E2036.pdf |quote=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.}} </ref>

{{clade| style=font-size:85%; line-height:85% |1=Chelidae |2={{clade |1=''Pelomedusoides'' |2={{clade |1={{clade |1=''Araripemys'' |2=Pelomedusidae }} |2={{clade |label1=Euraxemydidae |1={{clade |1=''Dirqadium'' |2=''Euraxemys'' }} |label2=Podocnemidioidea |2={{clade |1=Podocnemidinura |label2=Bothremydidae |2={{clade |label1=Kurmademydini |1={{clade |1=''Kurmademys'' |2=''Sankuchemys'' }} |label2=Bothremydinae |2={{clade |label1=Cearachelyini |1={{clade |1=''Galianemys'' |2=''Cearachelys'' }} |label2=Bothremyodda |2={{clade |label1=Bothremydini |1={{clade |1={{clade |1=''Foxemys'' (=''Polysternon'') |2=''Polysternon'' }} |2={{clade |1=''Zolhafah'' |2={{clade |1=''Rosasia'' |2={{clade |1=''Araiochelys'' |2={{clade |1={{clade |1=''Chedighaii hutchisoni'' |2=''Chedighaii barberi'' }} |2={{clade |1={{clade |1=''Bothremys cooki'' |2=''Bothremys magrhebiana'' }} |2={{clade |1=''Bothremys kellyi'' |2=''Bothremys arabicus'' }} }} }} }} }} }} }} |label2=Taphrosphyini |2={{clade |1={{clade |1=''Nigeremys'' |2=''Arenila'' }} |2={{clade |1=''Azabbaremys'' |2={{clade |1=''Labrostochelys'' |2={{clade |1=''Taphrosphys'' |2={{clade |1=''Rhothonemys'' |2={{clade |1=''Phosphatochelys'' |2=''Ummulisani'' }} }} }} }} }} }} }} }} }} }} }} }} }} }}

==References== {{Reflist}}

Category:Bothremydidae Category:Extinct turtles †