2018: Habilitation University of Tuebingen / Germany (habil.)

since 2016: Curator of the Palaeontological Collection in Tübingen / Germany 

2015: Postdoc, Museum für Naturkunde Berlin / Germany 

2013-2014: Postdoc, Paläontologisches Institut und Museum der Universität Zürich 2012-2013: Postdoc, Geowissenschaften Universität Tübingen / Germany 

2011: Postdoc, RIKEN Center for Developmental Biology Kobe/Japan 

2010-2011: Postdoc, Paläontologisches Institut und Museum der Universität Zürich 

2007-2010: PhD-student, Paläontologisches Institut und Museum der Universität Zürich / Switzerland (Dr. sc. nat.) 

2006-2007: Diploma, Institut für Spezielle Zoologie und Evolutionsbiology mit Phyletischem Museum Friedrich Schiller Universität Jena / Germany 2001-2006: Studies in Biology and Prehistory, Friedrich Schiller Universität Jena / Germany (Dipl. Biol.)

Using the ‘comprehensive organism concept’, a holistic understanding of evolutionary processes is possible. Therein, the whole life of an organism - from fertilization to dead - is understood as a process of development on which selection can act through evolution. Heterochronies - changes in the developmental timing of characters - are hypothesized to have fundamental impact on shaping an adult structure. In mutual dependence with those, spatial reorganizations take place during development, so called heterotopies. Both mechanisms, together with allometric change, enable the evolution of new morphotypes.

With the help of standardized developmental characters, I study the patterns of embryonic timing in a variety of land vertebrate species, including turtles, snakes, lizards, and diverse mammalian groups. Given the recent advances in molecular systematics, a comprehensive (falcifiable) phylogenetic framework is provided, which I use to trace evolutionary developmental changes and discuss those in the context of morphological diversification. Besides organogenesis, also patterns of skeletogenesis and some genetic aspects of development and evolution are observed.

On a histological scale, development of particular organs is studied – a classic comparative embryological approach – in order to detect homologies, to uncover patterns of recapitulation, and to trace spatial developmental changes that lead to adult morphology. For adult anatomy, similar to my embryological studies, I concentrate on the craniocervical system of amniotes and some fish. One focus lies on the morphology of the musculature and other soft tissues, their anatomical diversity, phylogenetic distribution, and functional integration. Another focus lies on the interrelationship of head and neck morphology.

My lab-members can be found here: LINK

Visiting Scientists

• 2025-26: Leonardo Kerber (Brazil)

• 2025ff: María Victoria Fernandez Blanco (Argentina)

• 2018: Xenia Schlindwein (Germany)

• 2017-18: Juliane Mittag (Germany)

• 2015-16: Oleksandr Yaryhin (Ukraine)

Postdocs

• 2025-27: Kai Ito (Japan)

• 2024-25: Franziska Wagner  (Germany)

• 2023-24: María Victoria Fernandez Blanco (Argentina)

• 2022-26: Mario Bronzati (Brazil)

• 2021-23: Omar Rafael Regalado Fernandez (Mexico)

• 2021-22: Anna Krahl (Germany)

• 2021: Andreas Matzke (Germany)

• 2018-21: Gerardo Antonio Cordero (USA, Venezuela)

• 2017-20: Oleksandr Yaryhin (Ukraine)

PhD-students

• 2024-26: Kuan-Yu Chou (Taiwan)

• 2021-26: Zitong Zhang (China)

• 2019-22: Pascal Abel (Germany)

• 2017-19: Gabriel de Souza Ferreira (Brazil)

Master's students

• 2020-21: Zitong Zhang (China)

• 2019-20: Yannick Pommery (France)

• 2016-17: Thomas Lechner (Germany)

• 2018-19: Parima Parsi-Pour (Iran)

• 2015: Alexander Janns (Germany)

• 2015: Stephan Spiekman (The Netherlands)

• 2013: Katja Polachowski (Switzerland)

Bachelor's students

• 2023-24: Tim Knauer (Germany)

• 2023: Carola Mauel (Germany)

• 2022: Christian Baun (Germany)

• 2022: Luca Leicht (Germany)

• 2020-21: Benedikt Kästle (Germany)

• 2020-21: Franziska Zepf (Germany)

• 2018-19: Xenia Schlindwein (Germany)

• 2016: Parima Parsi-Pour (Iran)

• 2016: Kim Bleher (Germany)

• 2016: Jennifer Storm (Germany

Undergratuate students (Scientific Practical)

• 2011: Alther, R. (Switzerland)

• 2011: Schwery, O. (Switzerland) 

• 2011: Fecker, D. (Switzerland)

• 2008: Bösch A. (Switzerland)

• 2008: Hüsser, C. (Switzerland)

• 2008: Fierz. J.-M. (Switzerland)

• 2008: Camenisch, D. (Switzerland)

• 2008: Signer, M. (Switzerland)

Current teaching

Bachelor

• 2016ff: Leit & Faziesfossilien (Invertebrate Fossil Taxonomy) - Lecture + practical (2 SWS) 

• 2021ff: Wirbeltierpaläontologie (Vertebrate Paleontology; part of module "Paläontologie") - Lecture (2SWS) 

Master

• 2024ff: Paleodiversity (part of module "Biodiversity") (1 SWS) 

• 2025ff: Vertebrate Evolutionary Morphology (part of Module "Anatomy of Vertebrates") - Lecture (3 SWS) 

• 2025ff: Vertebrate Taxonomy (part of module "Evolution of Organisms") - Practical/Seminar (3SWS) 

Previous teaching

• 2019-2025: “Vertebrate Evolutionary Morphology” (6 SWS) - Lecture and practical (MA)

• 2017-19: “Zoomorphology” (6 SWS) - Lecture and practical in (MA)

pre-Tübingen

• 2015-2016: Lectures and practicals on Comparative Anatomy in Humboldt Uni Berlin (BA)

• 2011: Morphology course in Kobe/Japan (MA)

• 2008-2011, 2014-15: Teaching assistant in different courses on “Evolutionary Morphology” in Zürich University (BA)

• 2004-2007: Teaching assistant Zoological ground courses (dissection & taxonomy course) for biology students in Jena University (BA)

Academic Distinctions

• 2025ff: Vice president of Gesellschaft für Naturkunde in Württemberg e.V.

• 2025ff: Board Member Stiftung Tübinger Crafoord-Vorträge

• 2024: Walter-Schall-Price; Gesellschaft für Naturkunde in Württemberg e.V.; for innovative functional morphological explanations on the origin of temporal openings in land vertebrates

• 2023: Nominations as Secretary and as Treasurer of the International Society of Vertebrate Morphology (ISVM); declined with thanks

• 2015: „Q-Team“; BmBF-funding of research-oriented teaching (Bologna.lab, Berlin); Development of jaw musculature in land vertebrates

• 2011: Bernhard Rensch Price; Gesellschaft für Biologische Systematik (GfBS); Awarded together with the Rensch-medal at the Biosystematics meeting, 26.2.2011 in Berlin, for “outstanding work in the area of Biological Systematics“. Laudation by Olivier Rieppel

• 2010: Faculty Price; Mathematisch-Naturwissenschaftlichen Fakultät der Universität Zürich, Switzerland; For exceptional scientific work during the PhD-thesis

• 2010: Writing Competition Price „Science of the future“; Internationale Bodenseehochschule

• 2009: Price for best student presentation (out of 40 applicants); 1. BioSyst EU Meeting in Leiden/The Netherlands

Honorary academic positions 

• 2017-2018: Paleontology Tübingen, voluntary curator Paläontologische Sammlung Tübingen

• 2016, 2019ff: Paleontology Tübingen, Prof. Dr. Madelaine Böhme

• 2013/11-2016/08: Naturkundemuseum Berlin/Germany, Prof. Dr. Johannes Müller, Dr. Nadia Fröbisch

• 2012ff: Spezielle Zoologie Uni Tübingen/Germany, Prof. Dr. Wolfgang Maier

• 2011/10-2011/12: Paläontologie Uni Zürich/CH, Prof. Dr. Marcelo R. Sánchez-Villagra

• 2009/12: UCL London/UK, Prof. Dr. Susan Evans, Dr. Marc Jones

• 2009/03: RIKEN Center for Developmental Biology Kobe/Japan, Prof. Dr. Shigeru Kuratani, Dr. Hiroshi Nagashima

PhD committee member

• 2025: Japser Ponstein (Netherlands)

• 2024: Senay Amalia Ozkaya de Juanas (Spain/Portugal)

• 2024: Tobias Massonne (Germany)

• 2024: Thomas Lechner (Germany)

• 2023: Pascal Abel (Germany)

• 2023: Gabriel Jorgewich-Cohan (Brazil/Switzerland)

• 2022: Nikolaos Kargopoulos (Greece/Germany)

• 2020: Juliane Mittag (né Hinz) (Germany)

• 2019: Gabriel S. Ferreira (Brasil).

Reviewer

• 2021-2023: Peer-Reviewer-Circle board member of Alexander-von-Humboldt-Gesellschaft

• 2018: Reviewer for Gesellschaft für Säugetierkunde (Fritz-Frank-Award)

• 2017: Book review for Garland Science (Taylor & Francis Group)

• 2007ff: Manuscript-reviews for different journals

Editorials: 

• 2022: Guest-editor of a special issue in Frontiers in Ecology and Evolution

• 2021-2022: Guest-editor of a virtual issue in Vertebrate Zoology

• 2019-2020: Guest-editor of a virtual issue in Vertebrate Zoology

• 2015ff: Associate Editor, journal MorphoMuseuM

• 2018ff: Associate Editor, journal Acta Zoologica

• 2016ff: Editor in chief of CHELYOPS – Berichte aus der Paläontologischen Sammlung in Tübingen

• 2014: Guest-editor of a special issue in Journal of Experimental Zoology B

• 2009ff: Editor in chief of Scidinge Hall Verlags Zürich/Tübingen

Edited books

• Vorlesung über Vergleichende Anatomie von Johannes Müller - Die Vorlesungsmitschrift von Ernst Zeller aus dem Sommersemester 1851 

• CHELYOPS - Berichte aus der Paläontologischen Sammlung in Tübingen, Band 1, 2016-2022 

• Die Taten des Lichts - Leben und Werk von Walter Werneburg [1922-1999] 

• Begleitheft zur Ausstellung "Die Taten des Lichts". 2. Juni bis 25. Juli 2022 in Tübingen 

• Vorlesung über Paleontologie von Ernst Haeckel - Die Vorlesungsmitschrift von Nikolai Nikolajewitsch Miklucho-Maclay aus dem Sommersemester 1966 

• Aus der Tiefenzeit. Die Paläontologische Sammlung der Universität Tübingen 

• Phylogenie, Funktionsmorphologie und Bionik - Schriften zum 60. Phylogenetischen Symposium in Tübingen 

• 60. Phylogenetisches Symposium. Funktionsmorphologie und Bionik. Programm und Abstracts. 23. - 25. November 2018 

• Schlüsselereignisse der organismischen Makroevolution 

• Symposium on Turtle Evolution. Program and Abstracts 

Edited Volumes

• Vergleichende Entwicklungsgeschichte – A Festschrift on the occasion of the 80th birthday of Prof. Dr. Wolfgang Maier, Tübingen 

• Tetrapod Water-Land Transition: Reconstructing Soft Tissue Anatomy and Function 

• Recent Advances in Chondrocranium Research 

• Quantifying evolutionary development: integrating metrical frameworks, gene expression, and morphology 

Books (single author)

• Als Kaiserin Jito nach Yoshino ging - Gedichte nach Kakinomoto no Hitomaro 

• Flucht nach Minorca 

• SØREN

• Minoisches Licht - und andere morphologische Reisen 

• Irings Falke. Radegundes Klage um Thüringen. Ein Trauerspiel. 

• Die Chronik der Familie Werneburg. Anno 2013. 

• Die Mexikanische Reise. Eine Morphologie der Neuen Welt. 

• Die Söhne der Sonne. 100 Gedichte nach bronzezeitlichen Felsbildern in Westschweden. 

• Urworte Panisch. Gedichte. 

Morphologie im erweiterten Sinne

• Die Bestimmung der Morphologie - Haeckel, Darwin und ein Homologie-Problem 

• Von Hardenbergs morphologischer Auftrag 

• Rekapitulation des goetheschen Geistes. Zum 100. Todestag von Ernst Haeckel am 9. August 2019 oder ‚Was wir Ernst Haeckel sonst noch verdanken‘ 

• Was den Schädel im Innersten Zusammenhält / Keeping the Skull Together 

• Versteinerte Natur. Erdgeist-Deutungen mit Walter Werneburg 

Comparative Anatomy

• Vergleichende Entwicklungsgeschichte – A Festschrift on the occasion of the 80th birthday of Prof. Dr. Wolfgang Maier, Tübingen (book)

• Einführung: Zur Methode der organismischen Evolutionsbiologie 

• Editorial: Vergleichende Entwicklungsgeschichte – A Festschrift on the occasion of the 80th birthday of Prof. Dr. Wolfgang Maier, Tübingen 

Embryology

• A Standard System to Study Vertebrate Embryos 

• Quantifying evolutionary development: integrating metrical frameworks, gene expression, and morphology (book)

• Quantifying Evolutionary Development Using Non-Model Organisms: Integrating Morphology, Metrical Frameworks, and Gene Expression (Editorial)

Bionik

• Phylogenie, Funktionsmorphologie und Bionik - Schriften zum 60. Phylogenetischen Symposium in Tübingen (book)

• Vorwort: Funktionsmorphologie und Bionik im Kontext der Phylogenetik 

• Das 60. Phylogenetische Symposium in Tübingen: Funktionsmorphologie und Bionik - Einführung, Programm und Abstracts 

• 60. Phylogenetisches Symposium. Funktionsmorphologie und Bionik. Programm und Abstracts. 23. - 25. November 2018 

• Zukunftsvisionen rund um Funktionsmorphologie und Bionik - 60. Phylogenetisches Symposium fand in Tübingen statt 

Taxonomy

• Zum Geleit (Als noch Arten-Kenner Schule machten) 

Evolutionary thinking

• Ganzheitliche Naturschau. Tübinger Paläontologie und die Evolution 

Haeckeliana

• Vorlesung über Paleontologie von Ernst Haeckel - Die Vorlesungsmitschrift von Nikolai Nikolajewitsch Miklucho-Maclay aus dem Sommersemester 1966 (book)

• Vorwort: Vorlesungen und Konzeptionen zur Paläontologie bei Ernst Haeckel 

• Discovery of rare lecture notes from 1866 provides exceptional insights to the conceptualization and visualization of paleontology by Ernst Haeckel 

• Stammbäume im ewigen Wandel - Ernst Haeckel, gepanzerte Kröten und die Transparenz der Wissenschaft 

• Flying turtles in the earliest trees of life: scientific transparency in the phylogenetic works of Ernst Haeckel 

• Echinoderm evolution: raise the historical treasure! 

• Ein Leitbild für den deutschen Staat 

• Die Bestimmung der Morphologie - Haeckel, Darwin und ein Homologie-Problem 

• Morgendämmerung der Evolutionsforschung 

• Darwin, Haeckel, and the “Mikluskan gas organ theory” 

• To the roots: back to Jena 

• Rekapitulation des goetheschen Geistes. Zum 100. Todestag von Ernst Haeckel am 9. August 2019 oder ‚Was wir Ernst Haeckel sonst noch verdanken‘. In: Palmbaum - Literarisches Journal aus Thüringen 

Wall charts / Wandtafeln

• Brüchiges Erbe - Wandtafeln der Tübinger Paläontologie wiederentdeckt 

• Palaeontologische Wandtafeln von Karl Alfred von Zittel und seinen Kollegen 

Johannes Müller

• Vorlesung über Vergleichende Anatomie von Johannes Müller - Die Vorlesungsmitschrift von Ernst Zeller aus dem Sommersemester 1851 (book)

• A life dedicated to research and ideal: Johannes Müller between empirical universality and idealistic vitalism mirrored in lecture notes from 1851 

• Ein Leben für Forschung und Ideal: Johannes Müller im Spannungsfeld zwischen empirischer Universalität und idealistischem Vitalismus [Einführung] 

Holger Preuschoft

• Nachruf auf Holger Preuschoft - 2. Januar 1932 - 8. April 2025 

• Organismen ganz neu, biomechanisch verstehen - Zum Tode von Prof. Dr. Dr. h.c. Holger Preuschoft (1932–2025) [incl. translation: "Understanding organisms in a completely new, biomechanical way – On the death of Prof. Dr. Dr. h.c. Holger Preuschoft (1932-2025)"] 

• Holger Preuschoft - Funktionelle Morphologie und universitäre Lehre 

Wolfgang Maier

• Vergleichende Entwicklungsgeschichte – A Festschrift on the occasion of the 80th birthday of Prof. Dr. Wolfgang Maier, Tübingen (book)

• Editorial: Vergleichende Entwicklungsgeschichte – A Festschrift on the occasion of the 80th birthday of Prof. Dr. Wolfgang Maier, Tübingen 

• Festschrift und Symposium für Tübinger Zoologie-Professor 

• Festsymposium zur „Vergleichenden Entwicklungsgeschichte“ - Der Tübinger Zoologie-Professor Wolfgang Maier wird geehrt 

Adolf Seilacher

• Eine Spur Glückseligkeit - Zum 100. Geburtstag des Naturforschers Dolf Seilacher (1925–2025)

• Adolf Seilacher zum 100. Geburtstag 

Frank Westphal

• Sein Herz schlug für die Geologie - und für Tübingen (Nachruf Frank Westphal) 

• Der vielleicht letzte seiner Art - Leben und Wirken von Prof. Dr. Frank Westphal (1930-2022) 

Others

• Aus dem Kaliflöz »Thüringen«: Der Geologe Otfried Konitz (1935–1977) 

• Laudation to Gabriel S. Ferreira, Bernhard Rensch-Preis 2020 

• Karl Rau (1870–1966) - Ein Forstmann und Geologe aus Tübingen und sein Belthle-Mikroskop 

• Anno 2019. Zum 50. Todestag von Friedrich Richard Freiherr von Hoyningen, genannt Huene 

• Friedrich August von Quenstedt 

General history

• Aus der Tiefenzeit. Die Paläontologische Sammlung der Universität Tübingen (book)

• Paläontologische Sammlung Tübingen 

• Ein Stück Kulturgeschichte. Zur Entwicklung der Paläontologischen Sammlung Tübingen 

• Lebendige Forschungssammlung – eine Einführung 

• Behind the Scenes at the Museum - The Palaeontological Collection, Eberhard Karls University, Tübingen, Germany 

• The Palaeontologial Collection of Tübingen 

• Denkschrift zur Bedeutung und zum Zustand der Senckenbergverwalteten Paläontologischen Sammlung an der Eberhard-Karls- Universität in Tübingen, anno 2021, nebst einem Konvoluten-Inventar ihrer Petrefakte, ihres Literaturbestandes sowie einer möglichen Zukunftsperspektive zum Erhalt und zur Erschließung derselben 

Specific topics

• Ganzheitliche Naturschau. Tübinger Paläontologie und die Evolution 

• Hidden ammonites: an insight into the Upper Cretaceous Wiedmann Collection at the Paläontologische Sammlung of Tübingen (Germany) 

• Fossile Pflanzen - Die Paläobotanische Schausammlung in Tübingen (1983 bis 2017) 

• Ein „Buch in drei Dimensionen“ - Die Lehrsammlung von Otto Schindewolf in Tübingen 

• Vorwort und Jahresberichte 2016 bis 2022 (in: CHELYOPS - Berichte aus der Paläontologischen Sammlung in Tübingen, S. I-XX) 

• CHELYOPS - Berichte aus der Paläontologischen Sammlung in Tübingen, Band 1, 2016-2022 

• Anno 2002. Die Institutsausstellung „Kunstformen der Erde“ auf Wanderschaft. In: CHELYOPS - Berichte aus der Paläontologischen Sammlung in Tübingen 1, 2021: 105-115. 

• Woher stammen die Konvolute der Paläontologischen Sammlung in Tübingen? 

• Raumzeitlich-paläontologischer Ursprung der GPIT-Konvolute. In: CHELYOPS - Berichte aus der Paläontologischen Sammlung in Tübingen 

Specific taxa / objects

• New long‑horned caddisflies in Eocene Baltic amber (Insecta, Trichoptera) 

• The morphology and evolutionary history of the temnospondyl genus Cyclotosaurus with a focus on material from Germany 

• Tomography of large fossils at NECTAR facility

• Neutron Imaging in Cultural Heritage Research at the FRM II Reactor of the Heinz Maier-Leibnitz Center  

• Neutron Tomography and X-ray Tomography as Tools for the Morphological Investigation of Non-mammalian Synapsids 

• Menschenaffe ‚UDO‘ – ein Paradigmenwandel 

• 哺乳類の顔を作ったダイナミックな進化過程 ～哺乳類の鼻は祖先の口先だった～ [The dynamic evolutionary process that created the mammalian face: The mammalian nose belonged to the mouthpart of our ancestors] 

• Mammalian face as an evolutionary novelty 

• Ausgewählte Objekte der Sammlung. Confuciusornis: Blick auf Einzelheiten 

• The Tübingen collection of ichthyosaurs from the Lower Jurassic (Lower Toarcian) Posidonienschiefer Formation of Württemberg: a historical and curatorial perspective 

• Der Tübinger Bernsteinschrank. Jüdisches Mäzenatentum im deutschen Kaiserreich 

• Objekte und Inventare. Erläuterungen 

• The Historical Archive of the Paleontological Collection Tübingen, Germany 

• Novel record of placodont remains including a Henodus cranium from the Upper Triassic Silves Group of the Algarve, southern Portugal 

• Complex dental wear analysis reveals dietary shift in Triassic placodonts (Sauropsida, Sauropterygia) 

• Dentition and feeding in Placodontia: Tooth replacement in Henodus chelyops 

• Henodus – Eine Tübinger Weltsensation 

• The Tübingen collection of ichthyosaurs from the Lower Jurassic (Lower Toarcian) Posidonienschiefer Formation of Württemberg: a historical and curatorial perspective 

• Diversity and taxonomy of the Late Triassic sauropodomorphs (Saurischia, Sauropodomorpha) stored in the Palaeontological Collection of Tübingen, Germany, historically referred to Plateosaurus 

• A new massopodan sauropodomorph from Trossingen Formation (Germany) hidden as ‘ Plateosaurus’ for 100 years in the historical Tübingen collection 

• The craniomandibular anatomy of the early archosauriform Euparkeria capensis and the dawn of the archosaur skull: Cranial anatomy of Euparkeria 

• Der Steinkern von Proganochelys quenstedtii – die erste Schildkröte der Welt 

• A surface scan of the "Tübingen Steinkern", Holotype of Proganochelys quenstedtii (Testudinata), with some historical remarks 

• An occurrence of fossil eggs from the Mesozoic of Madagascar and a detailed observation of eggshell microstructure 

GPIT-Events

• Eine Spur Glückseligkeit - Zum 100. Geburtstag des Naturforschers Dolf Seilacher (1925–2025)

• Die „Hilgendorf Lecture“, EVEREST und die Paläontologie in Tübingen 

• Mini-Symposium in the Tübingen Paleontological Collection “Research in Museums” 

GPIT-Persons

• Eine Spur Glückseligkeit - Zum 100. Geburtstag des Naturforschers Dolf Seilacher (1925–2025)

• Adolf Seilacher zum 100. Geburtstag 

• Sein Herz schlug für die Geologie - und für Tübingen (Nachruf Frank Westphal) 

• Der vielleicht letzte seiner Art - Leben und Wirken von Prof. Dr. Frank Westphal (1930-2022) Karl Rau (1870–1966) - Ein Forstmann und Geologe aus Tübingen und sein Belthle-Mikroskop 

• Anno 2019. Zum 50. Todestag von Friedrich Richard Freiherr von Hoyningen, genannt Huene 

• Friedrich August von Quenstedt 

General

• Schlüsselereignisse der organismischen Makroevolution (book)

• Konvergente Evolution herbivorer Landwirbeltiere 

Embryology

• Webpage: Domestikations-Lernspiele / Domestication education game 

• Webpage: Embryologie-Lernspiel / Embryology educational game 

Hair/scales

• O que o desenvolvimento dos pelos pode nos dizer sobre nossos antigos ancestrais 

• What fur development can tell us about our ancient ancestors 

• Hatten Dinosaurier ein Fell? / Did dinosaurs have fur coats? 

• Biomechanische Grundlagen der Schläfenevolution bei Landwirbeltieren 

• Terrestrialisation and the cranial architecture of tetrapods 

• Evolution of the temporal skull openings in land vertebrates: A hypothetical framework on the basis of biomechanics 

• The temporal region of the tetrapod skull: a textbook example on integrative morphology 

• Modeling Skull Network Integrity at the Dawn of Amniote Diversification With Considerations on Functional Morphology and Fossil Jaw Muscle Reconstructions 

• Morphology of the temporal skull region in tetrapods: research history, functional explanations, and a new comprehensive classification scheme 

• Morphofunctional Categories and Ontogenetic Origin of Temporal Skull Openings in Amniotes 

• The Tendinous Framework in the Temporal Skull Region of Turtles and Considerations About Its Morphological Implications in Amniotes: A Review

Turtle

• Neck motion in turtles and its relation to the shape of the temporal skull region 

• Temporal Bone Arrangements in Turtles: An Overview 

General

• Recent Advances in Chondrocranium Research (Book) 

• Recent Advances in Chondrocranium Research (Editorial) 

• Character definition and tempus optimum in comparative chondrocranial research 

• Chondrification and Character Identification in the Skull Exemplified for the Basicranial Anatomy of Early Squamate Embryos 

• Limb reduction in squamate reptiles correlates with the reduction of the chondrocranium: A case study on serpentiform anguids 

Mammalia

• Ontogeny and phylogeny of the mammalian chondrocranium: The cupula nasi anterior and associated structures of the anterior head region 

Tuatara

• Morphological association between muscle attachments and ossification sites in the late cartilaginous skull of tuatara embryos 

• The origin of orbitotemporal diversity in lepidosaurs: insights from tuatara chondrocranial anatomy 

Squamates

• Limb reduction in squamate reptiles correlates with the reduction of the chondrocranium: A case study on serpentiform anguids 

• Tracing the developmental origin of a lizard skull: Chondrocranial architecture, heterochrony, and variation in lacertids 

• Chondrification and Character Identification in the Skull Exemplified for the Basicranial Anatomy of Early Squamate Embryos 

Crocodiles

• Structure and evolution of the embryonic cartilaginous skull of crocodilians 

Turtles

• Variation and development of the turtle chondrocranium, with a description of the common musk turtle (Sternotherus odoratus, Kinosternidae, Cryptodira, Testudines) 

• Chondrocranial anatomy of Testudo hermanni (Testudinidae, Testudines) with a comparison to other turtles 

Birds

• Evolution and development of the bird chondrocranium 

• A Standard System to Study Vertebrate Embryos 

Mammalia

• Timing of organogenesis underscores the evolution of neonatal life histories and powered flight in bats 

• Comparative embryogenesis in ungulate domesticated species 

• Embryonic staging of bats with special reference to Vespertilio sinensis and its cochlear development 

• An irregular hourglass pattern describes the tempo of phenotypic development in placental mammal evolution 

• Ontogeny of domestic dogs and the developmental foundations of carnivoran domestication 

• Mammalian organogenesis in deep time: tools for teaching and outreach 

• Evolution of organogenesis and the origin of altriciality in mammals 

• Development and embryonic staging in non-model organisms: The case of an Afrotherian mammals 

• The early development of the echidna, Tachyglossus aculeatus (Mammalia: Monotremata), and patterns of mammalian development 

• Mammalian embryology and organogenesis 

Squamata

• Embryology and Life History of Reptilia with a Focus on the Organogenesis of Squamates 

Turtles

• Three Ways to Tackle the Turtle: Integrating Fossils, Comparative Embryology, and Microanatomy 

• Developmental differences between two marine turtle species and potential consequences for their survival at hatching 

• Zum Ursprung der Schildkröten: Ansätze der Vergleichenden Embryologie 

• Embryogenesis and Ossification of Emydura subglobosa (Testudines, Pleurodira, Chelidae) and Patterns of Turtle Development 

• Timing of organogenesis support basal position of turtles in the amniote tree of life 

Birds

• Domestication and the Comparative Embryology of Birds 

• Shifts in growth, but not differentiation, foreshadow the formation of exaggerated forms under chicken domestication 

Mammalia

• Mammalian skull heterochrony reveals modular evolution and a link between cranial development and brain size 

• Patterns in the bony skull development of marsupials: High variation in onset of ossification and conserved regions of bone contact 

• Development and embryonic staging in non-model organisms: The case of an Afrotherian mammals 

• Prenatal cranial bone development of Thomas's horseshoe bat (Rhinolophus thomasi): With special reference to petrosal morphology 

• Embryonic staging of bats with special reference to Vespertilio sinensis and its cochlear development 

Snakes + Lizards

• Embryonic development and cranial ossification of the Japanese Aodaishō, Elaphe climacophora (Serpentes: Colubridae): with special reference to the prootic bone and auditory evolution in snakes 

• Skeletal heterochrony is associated with the anatomical specializations of snakes among squamate reptiles 

• Late embryos and bony skull development in Bothropoides jararaca (Serpentes, Viperidae) 

Turtle

• Embryogenesis and Ossification of Emydura subglobosa (Testudines, Pleurodira, Chelidae) and Patterns of Turtle Development 

Mammalia

• Palaeo- & Neocauda: The ‘tail-hindlimb-complex‘ and the evolution of mammals 

• Remarks on the functional morphology of the earliest mammals 

• From sprawling to parasagittal locomotion in Therapsida: A preliminary study of historically collected museum specimens 

Turtles

• Deep time invention and hydrodynamic convergences through amniote flipper evolution 

• The hooked element in the pes of turtles (Testudines): A global approach to exploring primary and secondary homology 

• Betz's Ecomorphology of Insects 

• Ein Leitbild für den deutschen Staat 

• To the roots: back to Jena 

• ✝ Testudoolithus oosp. (Reptilia, Testudines) LAWVER, RASOAMIARAMANANA & INGMAR WERNEBURG 2015

• Electroadicella kuenowi sp. nov. (Insecta, Trichoptera) WICHARD, NEUMANN & WERNEBURG 2018

• Triaenodes simoni sp. nov. (Insecta, Trichoptera) WICHARD, NEUMANN & WERNEBURG 2018

• ✝ Tuebingosaurus gen nov. (Reptilia, Dinosauria) REGALADO FERNÁNDEZ & WERNEBURG 2022

• ✝ Tuebingosaurus maierfritzorum sp. nov.  (Reptilia, Dinosauria) REGALADO FERNÁNDEZ & WERNEBURG 2022

• ✝ Henodus sp. (Reptilia, Placodontia) RUCINSKI, CAMPOS, MATEUS, WERNEBURG 2025

• New long‑horned caddisflies in Eocene Baltic amber (Insecta, Trichoptera) 

• Betz's Ecomorphology of Insects

• Darwin, Haeckel, and the “Mikluskan gas organ theory” 

• Morphology of the jaw, suspensorial, and opercle musculature of Beloniformes and related species (Teleostei: Acanthopterygii), with a special reference to the m. adductor mandibulae complex 

• Head Morphology of the Ricefish, Oryzias latipes (Teleostei: Beloniformes) 

• Vergleichende Morphologie der Kiefermuskulatur der Beloniformes (Teleostei, Atherinomorpha) 

• Tetrapod Water-Land Transition: Reconstructing Soft Tissue Anatomy and Function 

• Contributions to the Evolution and Development of Land Vertebrates 

Fossils

• The morphology and evolutionary history of the temnospondyl genus Cyclotosaurus with a focus on material from Germany 

• The Palatal Interpterygoid Vacuities of Temnospondyls and the Implications for the Associated Eye‐ and Jaw Musculature 

• Palate anatomy and morphofunctional aspects of interpterygoid vacuities in temnospondyl cranial evolution 

Caudata

• Contributions to the functional morphology of caudate skulls: kinetic and akinetic forms 

• Tomography of large fossils at NECTAR facility

• Neutron Imaging in Cultural Heritage Research at the FRM II Reactor of the Heinz Maier-Leibnitz Center  

• Neutron Tomography and X-ray Tomography as Tools for the Morphological Investigation of Non-mammalian Synapsids 

Humans

• Menschenaffe ‚UDO‘ – ein Paradigmenwandel 

• Vom Ursprung des Menschen bis zur Neolithischen Revolution 

Comparative embryology

• 哺乳類の顔を作ったダイナミックな進化過程 ～哺乳類の鼻は祖先の口先だった～ [The dynamic evolutionary process that created the mammalian face: The mammalian nose belonged to the mouthpart of our ancestors] 

• Mammalian face as an evolutionary novelty 

• Mammalian embryology and organogenesis 

Organogenesis

• Timing of organogenesis underscores the evolution of neonatal life histories and powered flight in bats 

• Comparative embryogenesis in ungulate domesticated species 

• Embryonic staging of bats with special reference to Vespertilio sinensis and its cochlear development 

• An irregular hourglass pattern describes the tempo of phenotypic development in placental mammal evolution 

• Ontogeny of domestic dogs and the developmental foundations of carnivoran domestication 

• Mammalian organogenesis in deep time: tools for teaching and outreach 

• Evolution of organogenesis and the origin of altriciality in mammals 

• Development and embryonic staging in non-model organisms: The case of an Afrotherian mammals 

• The early development of the echidna, Tachyglossus aculeatus (Mammalia: Monotremata), and patterns of mammalian development 

Skeletogenesis

• Embryonic staging of bats with special reference to Vespertilio sinensis and its cochlear development 

• Patterns in the bony skull development of marsupials: High variation in onset of ossification and conserved regions of bone contact 

• Prenatal cranial bone development of Thomas's horseshoe bat (Rhinolophus thomasi): With special reference to petrosal morphology 

• Mammalian skull heterochrony reveals modular evolution and a link between cranial development and brain size 

• Development and embryonic staging in non-model organisms: The case of an Afrotherian mammals 

Chondrocranium

• Ontogeny and phylogeny of the mammalian chondrocranium: The cupula nasi anterior and associated structures of the anterior head region 

Locomotion

• Palaeo- & Neocauda: The ‘tail-hindlimb-complex‘ and the evolution of mammals 

• Remarks on the functional morphology of the earliest mammals 

• From sprawling to parasagittal locomotion in Therapsida: A preliminary study of historically collected museum specimens 

Captorhinus

• 3D models related to the publication: Skull sutures and cranial mechanics in the Permian reptile Captorhinus aguti and the evolution of the temporal region in early amniotes 

• Skull Sutures and Cranial Mechanics in the Permian Reptile Captorhinus aguti and the Evolution of the Temporal Region in Early Amniotes 

• Modeling Skull Network Integrity at the Dawn of Amniote Diversification With Considerations on Functional Morphology and Fossil Jaw Muscle Reconstructions 

Henodus

• Novel record of placodont remains including a Henodus cranium from the Upper Triassic Silves Group of the Algarve, southern Portugal 

• Complex dental wear analysis reveals dietary shift in Triassic placodonts (Sauropsida, Sauropterygia) 

• Dentition and feeding in Placodontia: Tooth replacement in Henodus chelyops 

• Henodus – Eine Tübinger Weltsensation 

Ichthyosauria

• The Tübingen collection of ichthyosaurs from the Lower Jurassic (Lower Toarcian) Posidonienschiefer Formation of Württemberg: a historical and curatorial perspective 

Dinosauria & allies

• Diversity and taxonomy of the Late Triassic sauropodomorphs (Saurischia, Sauropodomorpha) stored in the Palaeontological Collection of Tübingen, Germany, historically referred to Plateosaurus 

• A new massopodan sauropodomorph from Trossingen Formation (Germany) hidden as ‘ Plateosaurus’ for 100 years in the historical Tübingen collection 

• The craniomandibular anatomy of the early archosauriform Euparkeria capensis and the dawn of the archosaur skull: Cranial anatomy of Euparkeria 

• Unique skull network complexity of Tyrannosaurus rex among land vertebrates 

Fossils (general)

• Homeotic effects, somitogenesis and the evolution of vertebral numbers in recent and fossil amniotes 

Ontogeny

• Zur Evolution lebendgebärender Reptilien. Ein Weg ohne Rückkehr? 

• Dynamic evolutionary interplay between ontogenetic skull patterning and whole-head integration 

• Skeletal heterochrony is associated with the anatomical specializations of snakes among squamate reptiles 

• Embryology and Life History of Reptilia with a Focus on the Organogenesis of Squamates 

Limbs

• Deep time invention and hydrodynamic convergences through amniote flipper evolution 

• Historically transposed flipper pairs in a mounted plesiosaurian skeleton 

Axial skeleton

• Vertebral Comparative Anatomy and Morphological Differences in Anguine Lizards With a Special Reference to Pseudopus apodus 

Ear

• Trifold origin of the reptilian ear ossicle and its relation to the evolutionary modification of the temporal skull region 

• Wider das Artensterben: Ein verbessertes Gehör als Schlüsselentwicklung 

• Enhanced hearing as key innovation at a time of mass extinction 

• Deep-time origin of tympanic hearing in crown reptiles 

Ontogeny

• Embryonic development and cranial ossification of the Japanese Aodaishō, Elaphe climacophora (Serpentes: Colubridae): with special reference to the prootic bone and auditory evolution in snakes 

• Skeletal heterochrony is associated with the anatomical specializations of snakes among squamate reptiles 

• Late embryos and bony skull development in Bothropoides jararaca (Serpentes, Viperidae) 

• Morphological association between muscle attachments and ossification sites in the late cartilaginous skull of tuatara embryos 

• The origin of orbitotemporal diversity in lepidosaurs: insights from tuatara chondrocranial anatomy 

Ontogeny

• Zur Evolution lebendgebärender Reptilien. Ein Weg ohne Rückkehr? 

• Discrete embryonic character variation uncovers hidden ecological adaptations in lacertid lizards 

• Phylogenetic patterns and ontogenetic origins of limb length variation in ecologically diverse lacertine lizards 

• Bony skull development in the Argus monitor (Squamata, Varanidae, Varanus panoptes) with comments on developmental timing and adult anatomy 

Chondrocranium

• Limb reduction in squamate reptiles correlates with the reduction of the chondrocranium: A case study on serpentiform anguids 

• Tracing the developmental origin of a lizard skull: Chondrocranial architecture, heterochrony, and variation in lacertids 

• Chondrification and Character Identification in the Skull Exemplified for the Basicranial Anatomy of Early Squamate Embryos 

Chondrocranium

• Structure and evolution of the embryonic cartilaginous skull of crocodilians 

General

• Chelonia: Morphology of a stiffened body 

• Konsequenzen der Panzer-Produktion auf die Radkettenmechanik und deren Wirkung auf terminale Greifarm-Effizienz bei Schildkröten 

• Three Ways to Tackle the Turtle: Integrating Fossils, Comparative Embryology, and Microanatomy 

• Evolution and Development of Turtles: Organogenesis and Cranial Musculature 

Fossil

• Der Steinkern von Proganochelys quenstedtii – die erste Schildkröte der Welt 

• A surface scan of the "Tübingen Steinkern", Holotype of Proganochelys quenstedtii (Testudinata), with some historical remarks 

• An occurrence of fossil eggs from the Mesozoic of Madagascar and a detailed observation of eggshell microstructure 

Growth patterns

• Correlations between local geoclimatic variables and hatchling body size in the sea turtles Caretta caretta and Chelonia mydas 

Ontogeny

• Considerations on the development of the akinetic skull in pleurodire and cryptodire turtles 

• Diverging development of akinetic skulls in cryptodire and pleurodire turtles: An ontogenetic and phylogenetic study 

• Embryonic remnants of intercentra and cervical ribs in turtles 

Organogenesis

• Developmental differences between two marine turtle species and potential consequences for their survival at hatching 

• Zum Ursprung der Schildkröten: Ansätze der Vergleichenden Embryologie 

• Embryogenesis and Ossification of Emydura subglobosa (Testudines, Pleurodira, Chelidae) and Patterns of Turtle Development 

• Timing of organogenesis support basal position of turtles in the amniote tree of life 

Chondrocranium

• Variation and development of the turtle chondrocranium, with a description of the common musk turtle (Sternotherus odoratus, Kinosternidae, Cryptodira, Testudines) 

• Chondrocranial anatomy of Testudo hermanni (Testudinidae, Testudines) with a comparison to other turtles 

Skeletogenesis

• Embryogenesis and Ossification of Emydura subglobosa (Testudines, Pleurodira, Chelidae) and Patterns of Turtle Development 

Skull

• Cranial anatomy of turtles (CT-scans, PTA-stained) 

• Cranial turtle CT scans 

• Evolution of the rostral vasculature in turtles 

• Morphological Diversity of Turtle Hyoid Apparatus Is Linked to Feeding Behaviour 

• Skull osteology, neuroanatomy, and jaw‐related myology of the pig‐nosed turtle Carettochelys insculpta (Cryptodira, Trionychia) 

• Independent origin of large labyrinth size in turtles 

• Neck motion in turtles and its relation to the shape of the temporal skull region 

• Feeding behaviour in a ‘basal’ tortoise provides insights on the transitional feeding mode at the dawn of modern land turtle evolution 

• The Head and Neck Anatomy of Sea Turtles (Cryptodira: Chelonioidea) and Skull Shape in Testudines 

• Temporal Bone Arrangements in Turtles: An Overview 

Head soft tissue

• PTA-CT scans of extant turtle skulls 

• The Tendinous Framework in the Temporal Skull Region of Turtles and Considerations About Its Morphological Implications in Amniotes: A Review 

• Jaw musculature during the dawn of turtle evolution 

• The cranial musculature of turtles 

Head-neck-interface

• Feeding biomechanics suggests progressive correlation of skull architecture and neck evolution in turtles 

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

• Deep time perspective on turtle neck evolution: Chasing the Hox code by vertebral morphology 

• Modeling Neck Mobility in Fossil Turtles 

• Evolution of Neck Vertebral Shape and Neck Retraction at the Transition to Modern Turtles: an Integrated Geometric Morphometric Approach 

• Reconstructing the evolution of neck retraction in turtles 

• Biomechanical analyses suggest relation between neck-retraction and the trochlear mechanism in extant turtles 

Brains / Endocasts

• Contrasting Brains and Bones: Neuroanatomical Evolution of Turtles (Testudinata) 

• On the "cartilaginous rider" in the endocasts of turtle brain cavities (Festschrift Wolfgang Maier) 

• 3D models related to the publication: On the “cartilaginous rider” in the endocasts of turtle brain cavities 

• The endocranial anatomy of the stem turtle Naomichelys speciosa from the Early Cretaceous of North America 

• 3D model related to the publication: The endocranial anatomy of the stem turtle Naomichelys speciosa from the Early Cretaceous of North America 

• Neurovascular anatomy of the protostegid turtle Rhinochelys pulchriceps and comparisons of membranous and endosseous labyrinth shape in an extant turtle 

• Sensory Evolution and Ecology of Early Turtles Revealed by Digital Endocranial Reconstructions 

Other topics

• Deep time invention and hydrodynamic convergences through amniote flipper evolution 

• The hooked element in the pes of turtles (Testudines): A global approach to exploring primary and secondary homology 

• Symposium on Turtle Evolution. Program and Abstracts 

Embryology

• Domestication and the Comparative Embryology of Birds 

• Shifts in growth, but not differentiation, foreshadow the formation of exaggerated forms under chicken domestication 

• Evolution and development of the bird chondrocranium