Assessments of phenotypic variations and variability as a tool for understanding evolutionary processes in echinoids Dissertation zur Erlangung des mathematisch-naturwissenschaftlichen Doktorgrades "Doctor rerum naturalium" der Georg-August-Universität Göttingen im Promotionsprogramm Geowissenschaften / Geographie der Georg-August University School of Science (GAUSS) vorgelegt von Nils Schlüter aus Berlin Göttingen 2016 Betreuungsausschuss: PD Dr. Frank Wiese, Abteilung Geobiologie, Geowissenschaftliches Zentrum der Universität Göt- tingen PD Dr. Mike Reich, SNSB - Bayerische Staatssammlung für Paläontologie und Geologie, München Mitglieder der Prüfungskommission Referent: Prof. Dr. Joachim Reitner, Abteilung Geobiologie, Geowissenschaftliches Zentrum der Universität Göttingen 1. Korreferent: PD Dr. Frank Wiese, Abteilung Geobiologie, Geowissenschaftliches Zentrum der Universität Göt- tingen 2. Korreferent: PD Dr. Mike Reich, SNSB - Bayerische Staatssammlung für Paläontologie und Geologie, München Weitere Mitglieder der Prüfungskommission: PD Dr. Gernot Arp, Abteilung Geobiologie, Geowissenschaftliches Zentrum der Universität Göttingen PD Dr. Michael Hoppert, Abteilung Allgemeine Mikrobiologie, Institut für Mikrobiologie und Genetik der Universität Göttingen Prof. Dr. Joachim Reitner, Abteilung Geobiologie, Geowissenschaftliches Zentrum der Universität Göttingen Prof. Dr. Volker Thiel, Abteilung Geobiologie, Geowissenschaftliches Zentrum der Universität Göt- tingen Tag der mündlichen Prüfung: 14.04.2016 ii Acknowledgments First of all, I would like to thank my principal supervisors Dr. Frank Wiese (Göttingen) and Dr.Mike Reich (München), as well as the board of examiners. Further, I thank several curators and colleagues who granted access to collections and loans of material, or donated important material for the studies (in alphabetical order): Angela Ehling (BGR Berlin), Tim Ewin (Natural History Museum London), Ulrich Kothoff (Universität Hamburg), Martin Krogman, Jens Lehmann (both University of Bremen), Carsten Lüter, Christian Neumann (both Museum of Natural History, Berlin), Alexander Mironov (P.P. Shirshov Institute of Oceanology, Moscow, Russia), Heinke Schultz (Hamburg, Germany) and Ekbert Seibertz (Wolfsburg), who gave very important informations regarding the whereabouts of his collected material. Also, I am thankful for technical support from Jan Evers (Freie Universität Berlin) and Dorothea Hause-Reitner (University of Göttingen), as well as Gabriele Schmidt for her support in administrative issues. Additionally, I am grateful for the help and support of several friends and colleagues: Annina Böhme, Dr. Jahn Hornung, Dr. Andreas Kroh, Manfred Kutscher, Gwen Other, Tanja Stegemann and Dr. Ben Thuy. I thank my parents and friends for their support and especially, my beloved wife Esra. I am grateful for financial support by the German Research Foundation (DFG RE 2599/6-1, DFG RE 2599/6-2 and DFG Wi 1656/8-2 within SPP 527: IODP/ODP) and by the EU (European Union) Synthesys project (DK-TAF-3208, FR-TAF-4917, GB-TAF-4914). iii Preface This doctoral thesis consists of following published manuscripts: Schlüter, N., Wiese, F. and Reich, M. (2015) Systematic assessment of the Atelostomata (Spatangoida and Holasteroida; irregular echinoids) based on spine microstructure. Zoological Journal of the Linnean Society. 175: 510–524. doi:10.1111/zoj.12291 Schlüter, N., Wiese, F., Kutscher, M. (2016) Heterochronic evolution in the Late Cretaceous echinoid Gauthieria (Echinoidea, Phymosomatidae). Cretaceous Research. 57: 294–305. doi:10.1016/j.cretres.2015.09.005 Schlüter, N. (2016) Ecophenotypic variation and developmental instability in the Late Cretaceous echinoid Micraster brevis (Irregularia; Spatangoida). PLoS ONE. 11 (2): e0148341. doi:10.1371/journal.pone.0148341 iv Contents Chapter 1. Introduction. ...................................................................................................................... 1 1.1. Evolutionary development .......................................................................................................... 2 1.2. Robustness: Canalization and developmental stability ............................................................... 4 1.3. Phenotypic plasticity ................................................................................................................... 8 1.4. The study object: the natural history of echinoids ..................................................................... 10 1.5. Methods ..................................................................................................................................... 12 1.6. Aims and scope ......................................................................................................................... 14 1.7. References ................................................................................................................................. 16 Chapter 2. Systematic assessment of the Atelostomata (Spatangoida and Holasteroida; irregular echinoids) based on spine microstructures. ................................................................................... 25 2.1.Introduction ................................................................................................................................ 25 2.2. Material ..................................................................................................................................... 28 2.2.1. Institutional abbreviations ................................................................................................... 28 2.3. General morphology of spines in atelostomate echinoids ......................................................... 29 2.4. Methods ..................................................................................................................................... 30 2.5. Systematic assessment ............................................................................................................... 31 2.6. Results ....................................................................................................................................... 33 2.7. Conclusions ............................................................................................................................... 39 2.8. Acknowledgements ................................................................................................................... 39 2.9. References ................................................................................................................................. 40 2.10. Appendix. Supplementary data ............................................................................................... 47 Chapter 3. Heterochronic evolution in the Late Cretaceous echinoid Gauthieria (Echinoidea, Phymosomatidae) ............................................................................................................................ 63 3.1. Introduction ............................................................................................................................... 63 3.2. Material and methods ................................................................................................................ 67 3.2.1. Regression analysis ............................................................................................................. 67 3.2.2. Abbreviations ...................................................................................................................... 68 3.3. Results ....................................................................................................................................... 69 3.4. Discussion ................................................................................................................................. 77 3.5. Conclusion ................................................................................................................................. 78 3.6. Systematic remarks.................................................................................................................... 78 3.7. Acknowledgements ................................................................................................................... 80 3.8. References ................................................................................................................................. 80 3.9. Appendix. Supplementary data ................................................................................................. 84 v Chapter 4. Ecophenotypic variation and developmental instability in the Late Cretaceous echinoid Micraster brevis (Irregularia; Spatangoida) ................................................................... 92 4.1. Introduction ............................................................................................................................... 92 4.2. Material and Methods ................................................................................................................ 95 4.2.1. Material ............................................................................................................................... 95 4.2.2. Institutional abbreviations ................................................................................................... 97 4.2.3. Methods – geometric morphometrically based analyses ..................................................... 97 4.2.4. Methods – variation in non-morphometric character .......................................................