EVOLUTION OF THE SIDE-NECKED TURTLES: THE FAMILIES BOTHREMYDIDAE, EURAXEMYDIDAE, AND ARARIPEMYDIDAE

Abstract

Although pleurodires have been considered significantly less diverse than their sister group, the cryptodires, current discoveries show that pleurodires had a more complex and extensive evolutionary history than had been realized. Previously unknown radiations, particularly in the near-shore marine realm, are revealed by taxa with diverse cranial morphology, indicating many different feeding and sensory strategies. The pleurodire group that is changed the most by the new discoveries is its largest group, the hyperfamily Pelomedusoides. The hyperfamily Pelomedusoides now consists of the families Pelomedusidae, Podocnemididae, Bothremydidae, Araripemydidae, and Euraxemydidae, new family. The families Bothremydidae, Araripemydidae, and Euraxemydidae, new family, are documented with descriptions of skulls, lower jaws, and shells. The relationships of the family Podocnemididae to its sister taxa Hamadachelys and Brasilemys are recognized by placing them in the epifamily Podocnemidinura. The epifamily Podocnemidinura is the sister group to the family Bothremydidae, and together they form the superfamily Podocnemidoidea.The family Araripemydidae consists of one taxon, Araripemys barretoi, from the Aptian-Albian of Brazil. Description of new cranial material suggests that it is the sister group to all other Pelomedusoides or the sister group to the Pelomedusidae, but these relationships are only weakly supported. There is strong support for a multichotomy of Araripemys, Pelomedusidae, and remaining Pelomedusoides. Araripemys is characterized by very thin triturating surfaces and by a shell that lacks mesoplastra and has the first costals reaching the shell margin.The new family Euraxemydidae consists of two new genera: Euraxemys essweini, n. gen. et sp., from the Albian Santana Formation of Brazil, and Dirqadim schaefferi, n. gen. et sp., from the Cenomanian Kem Kem beds of Morocco. Members of the Euraxemydidae are united by the unique possession of a medial process of the quadrate partially covering the prootic and narrowly contacting a ventral process of the exoccipital, in contrast to all other pleurodires, which have either complete exposure or complete covering of the prootic ventrally. Furthermore, members have a ventral process of the exoccipital that is exposed at the lateral margin of the basioccipital in an elongate foot. The Euraxemydidae is hypothesized as the sister group to the superfamily Podocnemidoidea.The family Bothremydidae and the epifamily Podocnemidinura (consisting of the family Podocnemididae, Hamadachelys, and Brasilemys) are united as the superfamily Podocnemidoidea based on the possession of a quadrate-basioccipital contact, the complete or nearly complete ventral covering of the prootic, and the extension of the pectoral scales onto the entoplastron.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. Although there is a diversity of triturating surfaces within the family, primitively bothremydids have a posteriorly wide triturating surface with a significant palatine contribution in the upper jaw.The family Bothremydidae consists of four newly recognized, monophyletic groups: the tribes Kurmademydini, Cearachelyini, Bothremydini, and Taphrosphyini. The tribe Kurmademydini consists of two taxa: Kurmademys kallamedensis, from the Maastrichtian Kallamedu Formation of India, and Sankuchemys sethnai, from the Maastrichtian Intertrappean beds of India. The tribe Kurmademydini is characterized by extensive temporal and cheek emargination, a large fossa precolumellari