Evolution, Diversity, and Development of the Craniocervical System in Turtles with Special Reference to Jaw Musculature

  • Gabriel S. FerreiraEmail author
  • Ingmar WerneburgEmail author
Part of the Fascinating Life Sciences book series (FLS)


Turtles are one of the most enigmatic groups of vertebrates with their highly modified “body plan” and, as such, attracted the attention of researchers for a long time. Aside from the unusual turtle shell, the skull in this group shows great changes in comparison to that of other amniote groups. Because the skull has been considered one of most important body regions when analyzing the phylogenetic relationships of amniotes, the distinct turtle skull morphology is one of the key features in defining their position among reptiles. Here, we review the current knowledge of the turtle head, summarizing the general morphology of the skull and neck as well as the different anatomical modifications characteristic of the main lineages of extant and extinct turtles. We explore the main questions that have been raised while studying those issues, for instance, the origin and diversity of the temporal emarginations (dermal bone reductions), the different neck retraction mechanisms and their influence on the shape of the skull, and the anatomy and development of the jaw adductor musculature and its relation to some characteristic features of the turtle skull, such as akinesis and the divergent trochlear mechanisms in cryptodires and pleurodires. Based on 3D reconstructions, we propose a hypothetical model for ancestral states and gross morphology of the jaw adductor musculature in Proganochelys quenstedti (the earliest turtle with a complete shell), an important step toward the understanding of the evolution of those muscles in turtles. Finally, we suggest that more integrative approaches that consider anatomical, developmental, and paleontological data and that employ modern techniques in morphological and functional anatomic analyses (such as μCT scanning and finite element analysis) have a greater potential to answer the still numerous open questions about the evolution of the turtle head.


Testudinata Pleurodira Cryptodira Proganochelys quenstedti Diapsida Neck retraction Emargination Fenestration M. zygomaticomandibularis Trochlear mechanism 



Janine Ziermann, Raul Diaz Jr., and Rui Diogo are thanked for the invitation to write this chapter. We would like to thank Juliane Hinz (Tübingen) for help with the 3D models and Wolfgang Maier for access to the histological sections. We also thank two anonymous reviewers for their suggestions. GSF was supported by FAPESP (Fundação de Amparo à Pesquisa do Estado de São Paulo) grants 2016/03934-2 and 2014/2539-5. IW was supported by SNF advanced postdoc mobility grant P300PA_164720.


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© Springer Nature Switzerland AG 2019

Authors and Affiliations

  1. 1.Senckenberg Center for Human Evolution and PalaeoenvironmentEberhard Karls UniversitätTübingenGermany
  2. 2.Fachbereich GeowissenschaftenEberhard-Karls-UniversitätTübingenGermany
  3. 3.Laboratório de Paleontologia de Ribeirão Preto, FFCLRPUniversidade de São PauloRibeirão PretoBrazil
  4. 4.Museum für Naturkunde, Leibniz-Institut für Evolutions- und BiodiversitätsforschungHumboldt-Universität zu BerlinBerlinGermany

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