Abstract

BackgroundThe phylogenetic placement of Cretaceous marine turtles, especially Protostegidae, is still under debate among paleontologists. Whereas protostegids were traditionally thought to be situated within the clade of recent marine turtles (Chelonioidea), some recent morphological and molecular studies suggest placement along the stem of Cryptodira. The main reason why the evolution of marine turtles is still poorly understood, is in part due to a lack of insights into the cranial anatomy of protostegids. However, a general availability of high-quality fossil material, combined with modern analysis techniques, such as X-ray microtomography, provide ample opportunity to improve this situation. The scope of this study is to help resolve its phylogenetic relationships by providing a detailed description of the external and internal cranial morphology of the extinct protostegid sea turtle Desmatochelys lowii Williston, 1894.Material and MethodsThis study is based on the well-preserved holotype of Desmatochelys lowii from the Late Cretaceous (middle Cenomanian to early Turonian) Greenhorn Limestone of Jefferson County, Nebraska. The skulls of two recent marine turtles, Eretmochelys imbricata (Linnaeus, 1766) (Cheloniidae) and Dermochelys coriacea Lydekker, 1889 (Dermochelyidae), as well as the snapping turtle Chelydra serpentina (Linnaeus, 1758) (Chelydridae) provide a comparative basis. All skulls were scanned using regular or micro CT scanners and the scans were then processed with the software program Amira to create 3D isosurface models. In total, 81 bones are virtually isolated, figured, and described, including the nature of their contacts. The novel bone contact data is compiled and utilized in a preliminary phenetic study. In addition, an update phylogenetic analysis is conduced that utilizes newly obtained anatomical insights.ResultsThe detailed examination of the morphology of the herein used specimens allowed to explore some features of the skull, to refine the scoring of Desmatochelys lowii in the recent global matrix of turtles, and develop five new characters. The alleged pineal foramen in the type skull of Desmatochelys lowii is shown to be the result of damage. Instead, it appears that the pineal gland only approached the skull surface, as it is in Dermochelys coriacea. Whereas the parasphenoid in confirmed to be absent in hard-shelled sea turtles, ist possible presence in Desmatochelys lowii is unclear. The results of the phenetic study show that Desmatochelys lowii is least similar to the other examined taxa in regards to the nature of its bone contacts, and therefore suggests a placement outside Americhelydia for this protostegid sea turtle. The phylogenetic study results in a placement of Protostegidae along the stem of Chelonioidea, which is a novel position for the group.

Highlights

  • All recent marine turtles are currently accepted to form a monophyletic group, Chelonioidea Baur, 1893, that consists of two clades: the hard-shelled Cheloniidae Bonaparte, 1832 and the leathery-shelled Dermochelyidae Lydekker, 1889 (Crawford et al, 2015; Pereira et al, 2017)

  • The type skull of D. lowii from the late Cretaceous Greenhorn Limestone of Jefferson County, Nebraska, is redescribed in detail using mCT scans to provide new data that may help resolve the conundrum surrounding the phylogenetic placement of the Cretaceous marine turtle group Protostegidae and the origin of extant marine turtles

  • The detailed external and internal morphology of this specimen are compared bone by bone with the extant marine turtles D. coriacea and E. imbricata and the snapping turtle C. serpentina

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Summary

Introduction

All recent marine turtles (i.e., turtles that permanently live in fully marine environments) are currently accepted to form a monophyletic group, Chelonioidea Baur, 1893, that consists of two clades: the hard-shelled Cheloniidae Bonaparte, 1832 and the leathery-shelled Dermochelyidae Lydekker, 1889 (Crawford et al, 2015; Pereira et al, 2017). Some studies suggest that protostegids are situated within crown Chelonioidea (Gaffney & Meylan, 1988; Hirayama, 1998; Cadena & Parham, 2015), while others place them outside of crown Cryptodira (Joyce, 2007; Anquetin, 2012; Rabi et al, 2013) It has to be mentioned, that these studies base on the same dataset for sea turtles and their sampling of Protostegidae is limited to Santanachelys gaffneyi Hirayama, 1998. Material and Methods: This study is based on the well-preserved holotype of Desmatochelys lowii from the Late Cretaceous (middle Cenomanian to early Turonian) Greenhorn Limestone of Jefferson County, Nebraska. The results of the phenetic study show that Desmatochelys lowii is least similar to the other examined taxa in regards to the nature of its bone contacts, and suggests a placement outside

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