"Alle Schöpfung ist Werk der Natur." (Goethe)


Dr. Ingmar Werneburg

E-mail: ingmar.werneburg[at]senckenberg.de
Tel: 0049/(0)7071 2973068

Curator of the Palaeontological Collection
Senckenberg Center for Human Evolution
and Palaeoenvironment (HEP)
at Eberhard Karls Universität
Sigwartstr. 10, room: 308g
D-72076 Tübingen/Germany

Lecturer of Zoomorphology
Fachbereich Geowissenschaften
Eberhard Karls Universität
Hölderlinstraße 12, room: 308g
D-72074 Tübingen

Research: Ontogeny & phylogeny

  • 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.


Curriculum vitae



  • Since 2016. Curator of the Palaeontological Collection and Lecturer of Zoomorphology (Böhme-lab) in Tübingen / Germany
  • 2015. Swiss National Science Foundation postdoctoral researcher, host: Museum für Naturkunde Berlin / Germany (associated to the N. Fröbisch-lab)
  • 2013-2014. Postdoctoral researcher, Paläontologisches Institut und Museum der Universität Zürich / Switzerland (Sánchez-lab): "Mammalian and reptilian embryology" (50% position)
  • 2012-2013. Postdoctoral researcher, Geowissenschaften Universität Tübingen / Germany (Joyce-lab): "Turtle neck evolution"; incl. parental leave
  • 2011. Postdoctoral researcher, RIKEN, Center for Developmental Biology Kobe / Japan (Kuratani-lab): "Amniote skull development"
  • 2010-2011. Postdoctoral researcher, Paläontologisches Institut und Museum der Universität Zürich / Switzerland (Sánchez-lab): "Mammalian embryology"


Honorary Academic Positions

  • 2013-16. 'Guest scientists', Museum für Naturkunde Berlin / Germany (J. Müller-lab, N. Fröbisch-lab)
  • 2012-13. 'Visiting Scientist', Spezielle Zoologie der Universität Tübingen / Germany (W. Maier-lab): "Amniote craniogenesis"
  • 2009/Dec. 'Visiting Scientist', UCL London / UK (S. Evan's lab): "Craniocervical anatomy in marine turtles"
  • 2009/Mar. 'Visiting Scientist', RIKEN Center for Developmental Biology, Kobe / Japan (S. Kuratani-lab): "Turtle development"

Collaborators of finished projects

Attended conferences (with presentations)

  • 8th International Congress of Vertebrate Morphology (ICVM), 16.7.2007-21.7.2007 in Paris / France     
  • Paläontologisch-Herpetologische Tagung 2008 (PalHerp), 16.5.-18.5.2008 in Berlin / Germany     
  • 2. Euro Evo Devo meeting (EED), 29.7.2008-1.8.2008 in Ghent / Belgium    
  • 101. Jahresversammlung der Deutschen Zoologischen Gesellschaft (DZG), 19. - 22. September 2008 in Jena / Germany    
  • 7th Annual Meeting of the European Association of Vertebrate Palaeontologists (EAVP), 20.-26.7.2009 in Berlin / Germany    
  • 1st European BioSyst Meeting and 11th Tagung der Deutschen Gesellschaft für Biologische Systematik (GfBS), 10.-14.8.2009 in Leiden / Netherlands    
  • 69th Annual meeting of the Society of Vertebrate Paleontology (SVP) and 57th Symposium of Vertebrate Palaeontology and Comparative Anatomy (SVPCA), 23.-26.9.2009 in Bristol / United Kingdom   
  • Royal Tyrrell Museum: Gaffney Turtle Symposium, 17-18 October 2009. Drumheller, Alberta, Canada   
  • Swiss Systematics Society (SSS-Day) 6.11.2009 in Bern / Switzerland    
  • 9th International Congress of Vertebrate Morphology (ICVM), 26.7.-31.7.2010 in Punta del Este / Uruguay   
  • 80th Annual Meeting of the Palaeontological Society (Paläontologische Gesellschaft) 6.-8.10.2010 in München / Germany    
  • Biosystematics 2011. 21.2.-26.2.2011 in Berlin /Germany    
  • Symposium on Turtle Evolution, University of Tübingen, Germany, 2-5 June 2012    
  • 4th meeting of the European Society for Evolutionary Developmental Biology (EED), July 10–13, 2012 in Lisbon / Portugal    
  • 40. Treffen Arbeitskreis Wirbeltierpaläontologie in der Paläontologischen Gesellschaft, 15.-17.3.2013 in Laimering / Germany    
  • 10th International Congress of Vertebrate Morphology (ICVM), in Barcelona / Spain    
  • Studium Generale Tübingen 2013    
  • 74. Annual meeting of the Society of Vertebrate Paleontology (SVP), Berlin / Germany     
  • Meeting of Central European Turtle Evolutionary Biologists (CETEB), Zürich, December 1st - 2nd 2014    
  • 42. Arbeitskreistreffen Wirbeltierpaläontologie in Ahrensburg bei Hamburg, 13.3.2015-15.3.2015    
  • 6th International Meeting on Cranio-cervical Systems in Vertebrates. Ghent, July 7th-10th 2015    
  • 15th Meeting of the European Society for Evolutionary Biology. Laussanne, August 10th-14th 2015    
  • 18th European Congress of Herpetology, 7-12 September 2015, Wrocław, Poland    
  • 86. Annual meeting of Paläontologische Gesellschaft. Schiffweiler, September 14th-17th 2015    
  • 57. Phylogenetisches Symposium „Was bestimmt die Form? Die Morphologie in der Evolutionsbiologie“ organised by Stefan Richter and Christian Wirkner. Rostock November 20th-22nd 2015    
  • 11th International Congress of Vertebrate Morphology, Washington DC June 29th-July 3rd 2016
  • 6th EuroEvoDevo Meeting (EED), Uppsala 26th-29th July 2016
  • 44. Arbeitskreistreffen Wirbeltierpaläontologie in Münster, 24.3.-26.3.2017

Supervisions / Hosting

  • Polachowski, Katja (2010-2011). Master's thesis. Contributions to the embryology of Squamata. Skeletogenesis and morphometric analysis of the skull, external morphology, and head measurements in embryonic series of a snake (Bothrops jararaca), a monitor lizard (Varanus panoptes), and two skink species (Lerista bougainvillii & Saiphos equalis) in a phylogenetic context. Paläontologisches Institut und Museum der Universität Zürich, 133 pp.
  • Spiekman, Stephan N. F. (2015-2016). Master's thesis. Variation in the onset of ossification and conserved regions of bone contact in the bony skull development of marsupial mammals. Museum für Naturkunde Berlin / Germany, 79 pp.
  • Ferreira, Gabriel de Souza (2015-2018): PhD-thesis on turtle development (São Paulo/Tübingen)
  • Yaryhin, Oleksandr (2015-2016). Guest scientists working on lacertid craniogenesis (Kiev/Ukraine)
  • Janns, Alexander (2016). Master's thesis. New material of Mesopithecus pentelicus from Bulgaria: Description and evaluation of species concept in light of sexual dimorphism (Tübingen / Germany) (co-supervisor: Madelaine Böhme)
  • Parsipour, Parima (2016/17). Bachelor's thesis. Evolution of marine turtle embryology. Humboldt-University Berlin and University of Tübingen / Germany
  • Bleher, Kim (2016/2017). Bachelor's thesis. Craniogenesis in snakes
  •  Storm, Jennifer (2017/2018). Bachelor's Thesis. Morphogenesis of turtle skulls


[my citation index at Google Scholar]   [my publications on ResearchGate]

* = correspondence author

Peer Reviewed Original Articles

  • 1. Werneburg I*, Hertwig ST (2009). Head morphology of the ricefish, Oryzias latipes (Teleostei: Beloniformes). Journal of Morphology 270(9):1095-1106, doi: 10.1002/jmor.10746, free article - from Werneburg 2007: related animations

  • 3. Werneburg I*, Sánchez-Villagra MR (2009). Timing of organogenesis support basal position of turtles in the amniote tree of life. BMC Evolutionary Biology, 9:82, doi:10.1186/1471-2148-9-82 (highly accessed, open access: htmlpdf, additional files 123). Updated SES-database: Link. Updated SES-database.

  • 4. Werneburg I*, Hugi J, Müller J, Sánchez-Villagra MR (2009). Embryogenesis and ossification of Emydura subglobosa (Testudines, Pleurodira, Chelidae) and patterns of turtle development. Developmental Dynamics 238(11): 2770-2786, doi: 10.1002/dvdy.22104 (free articleFigure S1-18Table S1). Updated SES-database.

  • 5. Müller J*, Scheyer TM, Head JJ, Barrett PM, Werneburg I, Ericson PGP, Pol D, Sánchez-Villagra MR* (2010). Homeotic effects, somitogenesis and the evolution of vertebral numbers in recent and fossil amniotes. Proceedings of the National Academy of Science (PNAS) 107(5): 2118-2123, doi: 10.1073/pnas.0912622107 (free article + supporting information)

  • 6. Werneburg I*, Sánchez-Villagra MR (2011). The early development of the echidna, Tachyglossus aculeatus (Mammalia: Monotremata) and patterns of mammalian development. Acta Zoologica 82(1): 75-88, doi: 10.1111/j.1463-6395.2009.00447.x, (articleappendices). Updated SES-database.

  • 7. Werneburg I* (2011). The cranial musculature of turtles. Palaeontologia Electronica 14(2), 15A: 99 pages. (free htmlfree pdf)

  • 8. Werneburg I* (2012). Temporal bone arrangements in turtles: an overview. Journal of Experimental Zoology Part B: Molecular and Developmental Evolution 318: 235-249, link

  • 9. Jones MEH*, Werneburg I, Curtis N, Penrose R, O'Higgins P, Fagan MJ, Evans SE (2012). The head and neck anatomy of sea turtles (Cryptodira: Chelonioidea) and skull shape in Testudines. Plos One 7(11): e47852. doi:10.1371/journal.pone.0047852free pdf,  free htmlvideos S1+S2+S3 [add ".mpg" to file name].

  • 10. Werneburg I, Tzika AC, Hautier L, Asher RJ, Milinkovitch MC, Sánchez-Villagra MR* (2013). Development and embryonic staging in non-model organisms: the case of an afrotherian mammal. Invited paper to the special issue "Vertebrate Evolutionary Developmental Biology" in Journal of Anatomy 222:2-18, featured cover picturefree htmlfree pdf, Supplements 123. Updated SES-database.
  • 11. Polachowski KM, Werneburg I* (2013). Late embryos and bony skull development in Bothropoides jararaca (Serpentes, Viperidae). Zoology 116 (1): 36-63. doi: 10.1016/j.zool.2012.07.003. Htmlpdf, Appendix 1234567. Updated SES-database.
  • 12. Werneburg, I* (2013)The tendinous framework in the temporal skull region of turtles and considerations about its morphological implications in amniotes: a review. Invited paper to Zoological Science 31(3): 141-153. Free pdf

  • 13. Werneburg, I* (2013). Jaw musculature during the dawn of turtle evolution. Organisms, Diversity & Evolution 13: 225-254 Htmlpdf

  • 14. Joyce WG*, Werneburg I, Lyson TR (2013). The hooked element in the pes of turtles (Testudines): a global approach to exploring primary and secondary homology. Journal of Anatomy 223: 421-441, featured cover picture, htmlpdf

  • 15. Werneburg I*, Maier W, Joyce WG (2013)Embryonic remnants of intercentra and ribs in the neck of turtles. Biology Open 2(11):1103-1107Free pdf

  • 16. Koyabu DB*, Werneburg I, Morimoto N, Zollikofer CPE, Forasiepi AM, Endo H, Kimura J, Ohdachi SD, Truong SN, Sánchez-Villagra MR* (2014)Mammalian skull heterochrony reveals modular evolution and a link between cranial development and brain size. Nature Communications 5:3625, 1-9. ArticleSupplementary informationSupplementary Data 12

  • 17. Werneburg I*, Sánchez-Villagra MR (2015). Skeletal heterochrony is associated with the anatomical specializations of snakes among squamate reptiles. Evolution 69(1): 254–263, featured cover picture. Article, Supplement, Press releases: German, English

  • 18. Werneburg I*, Wilson LAB*, Parr WCH, Joyce WG (2015). Evolution of neck vertebral shape and neck retraction at the transition to modern turtles: an integrated geometric morphometric approach. Systematic Biology 64(2): 187-204, featured cover picture. Article, Supplementary.

  • 19. Werneburg I* (2015). 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. PeerJ 3:e769 http://dx.doi.org/10.7717/peerj.769, 53 pages

  • 20. Werneburg I*, Hinz JK, Gumpenberger M, Volpato V, Natchev N, Joyce WG (2015). Modeling neck mobility in fossil turtles. Journal of Experimental Zoology Part B: Molecular and Developmental Evolution 324(3): 230-243. Invited paper to the special issue on "Origin and Evolution of Turtles" edited by Jacqueline E. Moustakas-Verho and Scott Gilbert. Article, supplement.
  • 21. Werneburg I*, Polachowski KM, Hutchinson M.N. (2015). Bony skull development in the Argus monitor (Squamata, Varanidae, Varanus panoptes) with comments on developmental timing and adult anatomy. Zoology 118: 255-280, Article, Supplement 1, Supplements 2-4. Updated SES-database.

  • 22. Natchev N*, Tzankov N, Werneburg I, Heiss E (2015). Feeding behaviour in a ‘basal’ tortoise provides insights on the transitional feeding mode at the dawn of modern land turtle evolution. PeerJ 3:e1172; DOI 10.7717/peerj.1172, pp. 1-23, Article, Appendix

  • 23. Lawver DR*, Rasoamiaramanana A, Werneburg I (2015). An occurrence of fossil eggs from the Mesozoic of Madagascar and a detailed observation of eggshell microstructure. Journal of Vertebrate Paleontology 35 (5): e973030 (6 pages), article, supplement

  • 24. Werneburg I* (2015). Neck motion in turtles and its relation to the shape of the temporal skull region. Comptes Rendus Palevol 14 (6-7): 527-548, article, supplement, featured cover picture in Vol. 15 (5)

  • 25. Werneburg I*, Laurin M, Koyabu D, Sánchez-Villagra MR* (2016). Evolution of organogenesis and the origin of altriciality in mammals. Evolution & Development 18(4): 229-244, featured cover picture, article, supplement, comment in The Evolving Placenta-blog

  • 26. Natchev N*, Handschuh S, Lukanov S, Tzankov N, Naumov B, Werneburg I* (2016). Contributions to the functional morphology of caudate skulls: kinetic and akinetic forms. PeerJ 4: e2392, doi 10.7717/peerj.2392, 15 pp.


  • 28. Werneburg I*, Geiger M* (in press). Ontogeny of domestic dogs and the developmental foundations of carnivoran domestication. Journal of Mammalian Evolution, article
  • 29. Witzmann F*, Werneburg I (in press). The palatal interpterygoid vacuities of temnospondyls and the implications for the associated eye- and jaw musculature. Anatomical Record, article

  • 30. Lautenschlager S*, Witzmann F*, Werneburg I (2016). Palate anatomy and morphofunctional aspects of interpterygoid vacuities in temnospondyl cranial evolution. Naturwissenschaften 103: 76, p. 1-10, article

  • 31. Spiekman SNF*, Werneburg I* (2017). Patterns in the bony skull development of marsupials: high variation in onset of ossification and conserved regions of bone contact. Nature Scientific Reports 7: 43197, pp. 1-11, article

Peer Reviewed Original Book Chapters

  • 32. Scheyer TM*, Werneburg I, Mitgutsch C, Delfino M, Sánchez-Villagra MR (2013). Chapter 6: Three ways to tackle the turtle: integrating fossils, comparative embryology and mircoanatomy. In: Morphology and Evolution of Turtles. Vertebrate Paleobiology and Paleoanthropology Series. Editors: Gardner J, Brinkman D, Holroyd P. Springer Verlag, Dortrecht, Heidelberg, New York, London (Buy book at Amazon), pp. 63-70, paper
  • 33. Werneburg I*, Spiekman S. N. F. (in press, online). Mammalian embryology and organogenesis. From gametes to weaning. In: Zachos F., Asher R. (eds.). Mammalia. Series: The Handbook of Zoology / Handbuch der Zoologie. De Gruyter, Berlin, Boston (featured book cover images), Link; Retrieved 5 Dec. 2016, from http://www.degruyter.com/view/Zoology/bp_027590-2_7

Peer Reviewed Non-Original Publications
  • 34. Werneburg I* (2010). Zum Ursprung der Schildkröten: Ansätze der Vergleichenden Embryologie. Vierteljahrschrift der Naturforschenden Gesellschaft in Zürich 155(1/2):21-28 (free article)
  • 35. Wilson LAB*, Werneburg I (2014). Editorial: Quantifying evolutionary development: integrating morphology, metrical frameworks, and gene expression. Special Issue in Journal of Experimental Zoology Part B: Molecular and Developmental Evolution 322B: 555-557. Article [peer reviewed]
Non Peer Reviewed, Non-Original Books and Book Chapters
  • 36. Joyce WG, Corsini JA, Werneburg I, Rabi M (editors/organisators) (2012). Symposium on Turtle Evolution 2012. Program and Abstracts. TOBIAS-lib Hochschulserver der Universität Tübingenfree pdf
  • 37. Maier W*, Werneburg I* (2014, eds.). Schlüsselereignisse der organismischen Makroevolution. Scidinge Hall Verlag Zürich, 416 pp. (German; with contributions by Oliver Betz, Karl Forchhammer, Rüdiger Hampp, Wolfgang Maier, Michael W. Maisch, Irina Ruf, Ingmar Werneburg). 416pp. ISBN:978-3-905923-15-5. Order book here: Link. Press release: Link.
  • 38. Maier W*, Werneburg I* (2014). Einführung: Zur Methodik der organismischen Evolutionsbiologie. In: Maier W, Werneburg I (eds.). "Schlüsselereignisse der organismischen Makroevolution". Scidinge Hall Verlag Zürich, p. 11-17. Pdf: Link, order book here: Link.
  • 39. Werneburg I* (2014). Konvergente Evolution herbivorer Landwirbeltiere [ein Review]. In: Maier W, Werneburg I (Eds.) "Schlüsselereignisse der organismischen Makroevolution". Scidinge Hall Verlag Zürich, p. 295-331. Pdf: Link, Order book here: Link.
  • 40. Werneburg I* (2014). Vom Ursprung des Menschen bis zur Neolithischen Revolution [ein Review]. In: Maier W, Werneburg I (Eds.) "Schlüsselereignisse der organismischen Makroevolution". Scidinge Hall Verlag Zürichp. 361-411. Pdf: Link, order book here: Link. 
  • 41. Wilson LAB, Werneburg I (2014, guest editors). Quantifying evolutionary development: integrating metrical frameworks, gene expression, and morphology. Special Issue in Journal of Experimental Zoology Part B: Molecular and Developmental Evolution 322B: 555-653. (with contributions by Robert J. Asher, Cyril Charles, Stephen J. Gaunt, Norberto P. Giannini, Lionel Hautier, Daisuke B. Koyabu, Michel Laurin, Christian Mitgutsch, Lennart Ollson, Rainer R. Schoch, Karen E. Sears, Nguyen Truong Son, and Janine M. Ziermann)  Volume [peer reviewed]
  • 42. Werneburg I* (2016). Die Paläontologische Sammlung. In: Seidl E. (Hrsg.). Museen + Sammlungen der Universität Tübingen. Schriften des Museums der Universität Tübingen MUT 14: 92-97.


  • 43. Sánchez, M.R. Embryos in Deep Time: The Rock Record of Animal Development in Evolution (Indiana Press 2012). Translated to German by Ingmar Werneburg as: Sánchez-Villagra, M.R. (2012). Embryonen aus der Tiefenzeit. Was Fossilien über die Evolution von Wachstumsprozessen und die Entwicklungsgenetik erzählen. vdf Hochschulverlag AG an der ETH Zürich. 239 pp. ISBN 3728134694example chapter, buy at Amazon


  • 44. Werneburg I (2007). "Vergleichende Morphologie der Kiefermuskulatur der Beloniformes (Teleostei, Atherinomorpha)". Institut für Spezielle Zoologie und Evolutionsbiologie mit Phyletischem Museum. Friedrich-Schiller-Universität Jena. 170 pages. Diploma thesis (engl. abstract), published by Digitale Bibliothek Thüringen (incl. six 3d-animations). URL: http://www.db-thueringen.de/servlets/DocumentServlet?id=11300; full pdf (German, for the extended English summary, see in Werneburg 2009 above); 3D-head Oryzias latipes: animations, movie 1 (all elements), movie 2 (bones, ligaments, cartilages, muscles, nervous system), movie 3 (as movie 2, but without nervous system), movie 4 (bones, cartilages, ligaments), movie 5 (only muscles). ----- The thesis was later also published with extended English summary "Comparative Morphology of the Jaw Musculature of Beloniformes)" as: Werneburg I* (2009). Vergleichende Morphologie der Kiefermuskulatur der Beloniformes (Teleostei, Atherinomorpha). Verlag Dr. Müller, ISBN: 978-3-639-19302-2 . This published Diploma-thesis was the basis for the later studies of Werneburg & Hertwig (2009) and Werneburg (2015).

  • 45. Werneburg I* (2010). Evolution and Development of Turtles: Organogenesis and Cranial Musculature. Scidinge Hall Verlag, Zürich, ISBN: 978-3-905923-03-2. This published PhD-thesis contains the manuscripts of A. Scheyer et al. (2009, [2013]), B. Werneburg (2009), C. Werneburg & Sánchez-Villagra (2009), D. Werneburg et al. (2009), E. Werneburg & Sánchez-Villagra (2011), F.-G. Werneburg (2011, 2013), H. Müller et al. (2010) [only as supplement], I. Werneburg (2010), free pdf

Further publications

  • Werneburg, I. (2013a). Irings Falke. Radegundes Klage um Thüringen. Ein Trauerspiel. Scidinge Hall Verlag Zürich, 100 Seiten, ISBN 978-3-905923-12-4 

  • Werneburg, I. (2011). Die Mexikanische Reise. Eine Morphologie der Neuen Welt. Ein Reisebericht. Scidinge Hall Verlag Zürich, 225 Seiten, ISBN: 978-3-905923-07-0

  • Werneburg, I. (2010). Die Söhne der Sonne. 100 Gedichte nach Felsbildern der westschwedischen Bronzezeit, mit einem Reisebericht. Scidinge Hall Verlag Zürich, 147 Seiten, ISBN: 978-3-905923-01-8

  • Werneburg, I. (2009). Urworte Panisch. Gedichte. Scidinge Hall Verlag Zürich, 48 Seiten, ISBN: 978-3-905923-00-1