A New Fossil-Lagerstätte from the Late Devonian of Morocco : Faunal Composition, Taphonomy and Paleoecology
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Zurich Open Repository and Archive University of Zurich Main Library Strickhofstrasse 39 CH-8057 Zurich www.zora.uzh.ch Year: 2019 A new fossil-Lagerstätte from the Late Devonian of Morocco : faunal composition, taphonomy and paleoecology Frey, Linda Posted at the Zurich Open Repository and Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-177848 Dissertation Published Version Originally published at: Frey, Linda. A new fossil-Lagerstätte from the Late Devonian of Morocco : faunal composition, taphon- omy and paleoecology. 2019, University of Zurich, Faculty of Science. A New Fossil-Lagerstätte from the Late Devonian of Morocco: Faunal Composition, Taphonomy and Palaeoecology Dissertation zur Erlangung der naturwissenschaftlichen Doktorwürde (Dr. sc. nat.) vorgelegt der Mathematisch-naturwissenschaftlichen Fakultät der Universität Zürich von Linda Frey von St. Ursen FR Promotionskommission Prof. Dr. Christian Klug (Leitung der Dissertation) Prof. Dr. Hugo Bucher Prof. Dr. Marcelo Sánchez Prof. Dr. Michael Coates Dr. Martin Rücklin Zürich, 2019 ABSTRACT ................................................................................................................................................2 INTRODUCTION......................................................................................................................................5 CHAPTER I Late Devonian and Early Carboniferous alpha diversity, ecospace occupation, vertebrate assemblages and bio-events of southeastern Morocco ..............................27 CHAPTER II Fossil-Lagerstätten and preservation of invertebrates and vertebrates from the Devonian in the eastern Anti-Atlas (Morocco) ...........................................................61 CHAPTER III Morphology, phylogenetic relationships and ecomorphology of the early elasmobranch Phoebodus ............................................................................................89 CHAPTER IV Functional morphology of mandibular arches in symmoriids as exemplifi ed by Ferromirum oukherbouchi gen. et sp. nov. (Late Devonian) ...............................135 CONCLUSIONS ....................................................................................................................................167 APPENDIX Appendix A Collaboration with other projects (abstracts) ............................................................171 Appendix B Additional article published during doctoral studies ................................................175 ACKNOWLEDGEMENTS ..................................................................................................................203 CURRICULUM VITAE ........................................................................................................................207 Abstract The Late Devonian (382. 7 to 358.9 Ma) was a period of intense environmental perturbations and fun- damental changes in marine ecosystems impacting also the marine vertebrates. Two extinction events, namely the Kellwasser Event (Frasnian-Famennian boundary) and the Hangenberg Crisis (latest Devoni- an) caused severe losses in diversity. Especially several groups of gnathostomes were strongly affected by the Hangenberg Crisis, which represents a bottleneck in the evolutionary history of jawed vertebrates. The marine ecosystem became reshaped and the previously diverse placoderms went extinct and the taxonomic diversity of acanthodians decreased, while other chondrichthyans and osteichthyans diversifi ed. From the stratigraphic interval between the Kellwasser and the Hangenberg crises, well-preserved remains of plac- oderms and isolated teeth were well documented from Morocco before. By contrast, articulated remains of chondrichthyans were discovered only recently from the Famennian of the Maïder region (Morocco) within this PhD project. Most fossil fi sh were found in a stratigraphic interval with abundant remains of thylacocephalan arthropods preserved in ferruginous nodules (early/ middle Famennian Thylacocephalan Layer). These gnathostomes include both known and new taxa of sarcopterygians (onychodontids), plac- oderms (Dunkleosteus, Titanichthys, Alienacanthus and Driscollaspis) and chondrichthyans (phoebodon- tids, symmoriids, and several new cladodont chondrichthyans). In the framework of this thesis, the remains of chondrichthyans were in the main focus because their skeletons are rarely preserved in the fossil record due to the fragility of their cartilaginous skeletons. The Famennian chondrichthyans of the Maïder region often preserve three-dimensional skulls, the visceral as well as postcranial skeleton (pectoral and pelvic girdles with articulated fi ns, dorsal fi ns with fi n spines, neural arches and sometimes the caudal fi n) and even mineralized soft-tissues (muscles, liver, digestive tract). Therefore, the discovery of such remains of Devonian chondrichthyans is of importance to examine their phylogenetic relationships, their morpholog- ical disparity and to reconstruct their ecological diversity including changes in the respective ecological roles of these vertebrate groups in the marine ecosystems. In chapter I, the palaeoenvironment of the gnathostome-bearing stratigraphic interval of the Maïder was quantitatively investigated by analysing the diversity and palaeocology of the accompanying inverte- brate faunas. For this purpose, 21 invertebrate associations from early Famennian to early Carboniferous age of the Maïder region (17 from Madene el Mrakib, four from Aguelmous) were sampled and deter- mined. Based on the composition of these associations, alpha diversity (changes in species richness at a single locality through time), ecospace occupation (three-dimensional modes of life including tiering, motility, and feeding mode) and trophic nucleus (dominant taxa of each association) were analysed. The species richness, ecospace occupation as well as the ratio between pelagic and benthic organisms were fl uctuating during the Famennian and Tournaisian. Those ecological changes correlate with numerous bio-events coinciding with fl uctuating global and/ or regional sea level and changes in oxygenation during the Devonian. Although the sea fl oor might have been a bit better oxygenated (extended ecospace includ- ing benthic modes of life) in some stratigraphic intervals, the Fammennian of the study area was mostly hypoxic to dysoxic because of the low taxonomic diversity of benthic invertebrates and the dominance of pelagic or benthic taxa tolerant to oxygen depletion. This is supported by the absence of benthic gnatho- stomes in the Maïder (including microremains). The taphonomy of the gnathostomes of the Famennian Thylacocephalan Layer and some other Devo- nian Fossillagerstätten was investigated in Chapter II. The mineral composition of the remains of gnatho- stomes, thylacocephalans and some other invertebrates from various localities were analysed by Raman spectroscopy and X-ray diffraction. To understand the genesis of the Famennian Fossillagerstätten, the mineral analyses were combined with the current knowledge of the palaeogeography and palaecology of the Maïder region. The fossils mainly contain iron oxides (oxidised pyrite) and phosphates. The Maïder was a basin isolated by land and two submarine ridges (Tafi lalt and Maïder Platform) from neighbouring basins, which limited water exchange and thus also oxygen supply during the Devonian. This palaeogeo- graphic setting in combination with the mineral composition of the fossils, the soft-tissue preservation and the low abundance of benthos (even a complete lack of benthic chondrichthyans), the Thylacocephalan Layer with exceptionally well-preserved chondrichthyans represents the fi rst Konservat-Lagerstätte (con- servation deposit) of the Devonian of North Africa. In chapter III, the morphology and palaeoecology of Phoebodus, a common and cosmopolitan chon- drichthyan from the Devonian (Givetian to Famennian), previously only known from isolated teeth, is 2 Abstract described. Phoebodus saidselachus sp. nov. of the Famennian of the Maïder region is the fi rst specimen of this genus preserving body, soft-tissue and skull remains. Its body and skull are slender and more elongate than those of other Palaeozoic chondrichthyans. Phylogenetic analyses based on a data matrix including 228 characters of 65 taxa and one outgroup taxon place Phoebodus saidselachus sp. nov. within the stem elasmobranchs. Therefore, the Devonian genus Phoebodus is the oldest stem elasmobranch with an elon- gate body shape. Other stem elasmobranchs with such body shapes are known from younger strata such as the phoebodontid Thrinacodus gracia from the Carboniferous of Bear Gulch and several taxa of xenacan- thids from the Permian. Therefore, chondrichthyans were morphologically and ecologically more diverse in the Devonian before the vertebrate bottleneck of the Hangenberg Crisis than previously thought. Chapter IV focuses on the jaw function of symmoriiform chondrichthyans. Although the fossil re- cord of Palaeozoic chondrichthyans improved in recent years, the functional morphology of anatomical elements such as the visceral skeleton are not known in detail due to the often fragmentary or deformed preservation. For example, there was a long debate about whether hyoids had a suspensory function or not (aphetohyoidean hypothesis) in symmoriids. The newly discovered chondrichthyan Ferromirum oukher- bouchi gen. et sp. nov. from the Maïder Basin has exceptionally well-preserved and undeformed mandib- ular and branchial arches.