Ingmar Werneburg lab

Welcome & Projects

Vertebrate Evolutionary Morphology in Tübingen

We are interested in evolutionary vertebrate morphology with a focus on reptiles and mammals. We study, using traditional histological and gross anatomical approaches, the interrelationship of ontogenetic and phylogenetic patterns within a modern conceptual and analytical framework. For that, we follow a holistic organism concept dealing with extant adult, fossil, and embryonic material. Specifically, we observe the sequence of ontogenetic features, variation, and the homology of adult and developmental structures. The latter include the chondrocranium, head and neck musculature, skull anatomy, and postcranial features. Many studies deal with turtles, lizards, and mammals in general.

If you are interested to join the lab, please contact us: ingmar.wernebur[at}

Our research topics

Temporal skull region in tetrapods

Using Anatomical Network Analyses (AnNA), Finite Element Analyses (FEAs), classical morphological comparisions and phylogenetic methods, we investigate the evolution, mechanistics, and development of the temporal skull region in land vertebrates. We are interested in how and why temporal openings have evolved.

Publications in the lab: Works by Pascal Abel, Mario Bronzati, and Ingmar Werneburg

Example: Ingmar Werneburg*, Pascal Abel (2022). Modelling skull network integrity at the dawn of amniote diversification with considerations on functional morphology and fossil jaw muscle reconstructions. Frontiers in Ecology and Evolution. Sections Paleontology + Evolutionary Developmental Biology 9(799637): 1-25.

Feeding related morphology in land vertebrates

Jaw musculature, tendons, and related skeletal structures are analyzed in a comparative anatomical and a biomechanical manner.

Publications in the lab: Works by Pascal Abel, Gabriel Ferreira, Yannick Pommery, Ingmar Werneburg, Oleksandr Yaryhin, Zitong Zhang; collaborations with Nikolay Natchev

Example: Nikolay Natchev*, Nikolay Tzankov, Ingmar Werneburg, Egon Heiss (2015a). Feeding behaviour in a ‘basal’ tortoise provides insights to a transitional feeding mode at the dawn of modern land turtle evolution. PeerJ 3: e1172, p.1-23


We study the development of the vertebrate skull using traditional histological techniques as well as modern µCT and FEA technologies.

Publications in the lab: Works by Ingmar Werneburg, Oleksandr Yaryhin, Zitong Zhang, collaboration with Wolfgang Maier

Example: Ingmar Werneburg*, Wolfgang Maier (2019a). Diverging development of akinetic skulls in cryptodire and pleurodire turtles: an ontogenetic and phylogenetic study. Vertebrate Zoology 69(2): 113-143. Cover image,

Turtle neck and head

Neck mobility has great influence on the shape of the turtle skull. We proposed the hypothesis that the increased neck mobility in turtles resulted in the closure of the temporal region in the (likely diapsid) stem turtles and secondarily shaped the resulting anapsid skull of more crownward turtle taxa.

Publications in the lab: Works by Gabriel Ferreira and Ingmar Werneburg

Example: Gabriel S. Ferreira*, Stephan Lautenschlager, Serjoscha E. Evers, Cathrin Pfaff, Jürgen Kriwet, Irena Raselli, Ingmar Werneburg* (2020). Feeding biomechanics suggests progressive correlation of skull architecture and neck evolution in turtles. Scientific Reports 10: 5505, 11 pp.,

Comparative organogenesis in vertebrates

Heterochrony plays a major role in evolution. Changes in the timing of organ development (external and internal organs, incl. skeleton) are analyzed using the Continuous Analyses. A "Standard Event System to Study Vertebrate Embryos" was introduced to quantitatively study homologous structures in ontogeny (Werneburg 2009). Using this approach a number of vertebrate species are described and phylogenetically analyzed.

Publications in the lab: Works by Antonio G. Cordero, Parima Parsi-Pour, Katja Polachowski, Xenia Schlindwein, Stephan Spiekman, Ingmar Werneburg, Zitong Zhang; in collaboration with Marcelo R. Sánchez-Villagra

Example: Daniel Núñez-León, Gerardo A. Cordero, Xenia Schlindwein, Per Jensen, Esther Stoeckli, Marcelo R. Sánchez-Villagra, Ingmar Werneburg* (2021). Shifts in growth, but not differentiation, foreshadow the evolution of novel forms during chicken domestication. Proceedings of the Royal Society B: Biological Sciences 288(1953): 1-10.

Paleontology Collection in Tübingen

As a group housed in the amazing Paleontological Collection in Tübingen/Germany, we conduct research in the history.

Publications in the lab: Works by Antonio G. Cordero, Benedikt Kästle, Anna Krahl, Omar Rafael Regalado Ferndandez, Ingmar Werneburg; in collaboration with Madelaine Böhme

Example: Ernst Seidl, Edgar Bierende, Ingmar Werneburg (2021a, Eds.). Aus der Tiefenzeit - Die Paläontologische Sammlung der Eberhard Karls Universität Tübingen. Publikationen des Museums der Universität Tübingen (MUT). Tübingen, 526 pages. ISBN: 978-3-9821339-3-5

Sauropodomorph fauna in Europe

Tübingen Paleontological Collection has a valuable historical sample of early sauropodomorph material collected by Friedrich August Quenstedt and Friedrich von Huene. We study this material with modern taxonomic and phylogenetic approaches.

Publications in the lab: Work by Mario Bronzati, Benedikt Kästle, Omar Rafael Regalado Ferndandez, Henrik Stöhr, Ingmar Werneburg

Example: Omar Rafael Regalado Fernandez*, Ingmar Werneburg (2022a). A new massopodan sauropodomorph from Trossingen Formation (Germany) hidden as ‘Plateosaurus’ for 100 years in the historical Tübingen collection. Vertebrate Zoology 72: 771-822, (Festschrift Wolfgang Maier).

Locomotion in terrestrial and secondarily aquatic tetrapods

Using embryonic data as well as X-ray and Anatomical Network approaches, we study the functional anatomy, evolution, and devevelopment of various aspects of tetrapod axial and paraxial locomotion.

Publications in the lab: Work by Antonio Cordero, Anna Krahl, Omar Rafael Regalado Ferndandez, Ingmar Werneburg, Oleksandr Yaryhin

Example: Holger Preuschoft, Anna Krahl, Ingmar Werneburg (2022). From sprawling to parasagittal locomotion in Therapsida: A preliminary study of historically collected museum specimens. Vertebrate Zoology 72: 907-936.