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Beta Diversity in Triassic and Modern Reef Basin Assemblages Beta-Diversität in Triassischen Und Rezenten Riffbeckenfaunen

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Beta Diversity in Triassic and Modern Reef Basin Assemblages Beta-Diversität in Triassischen Und Rezenten Riffbeckenfaunen Beta diversity in Triassic and modern reef basin assemblages Beta-Diversität in triassischen und rezenten Riffbeckenfaunen Der Naturwissenschaftlichen Fakultät der Friedrich-Alexander-Universität Erlangen-Nürnberg zur Erlangung des Doktorgrades Dr. rer. nat. vorgelegt von Vanessa Julie Roden Als Dissertation genehmigt von der Naturwissenschaftlichen Fakultät der Friedrich-Alexander-Universität Erlangen-Nürnberg Tag der mündlichen Prüfung: 20. Dezember 2019 Vorsitzender des Promotionsorgans: Prof. Dr. Georg Kreimer Gutachter: Prof. Dr. Wolfgang Kiessling PD Dr. Kenneth De Baets Summary Changes in alpha and gamma diversity throughout the Phanerozoic have long been of interest in paleontology, but beta diversity remains understudied, particularly in the post-Paleozoic. Beta diversity – the compositional variation among communities or assemblages – is a key aspect of bio- diversity and is crucial to revealing the principles of diversity assembly. Studying the biodiversity patterns of well-preserved fossil and modern assemblages can provide insight into how these patterns arise, how they change through time, and whether there are unifying principles in community composition and assembly. Taphonomic effects and sampling biases complicate the assessment of changes in biodiversity over time. Intensively sampled and well-preserved fossil assemblages are ideal to study biodiversity patterns and compare them with modern assemblages. The Middle to Late Triassic (Ladinian–Carnian) Cassian Formation, exposed in the Dolomites, Southern Alps, northern Italy, is characterized by high diversity and excellent preservation of fossils. The Cassian Formation comprises shallow and deeper water sediments deposited between carbonate platforms in a warm, tropical setting in the Western Tethys, comparable with modern tropical environments. A wide breadth of depositional environments are recorded in the Cassian Formation: a nearshore back-reef area with patch reefs and the sediments intercalated between these, the carbonate platform, and the shallow and deeper reef basin. With 1421 invertebrate species, the Cassian Formation yields the highest species richness reported from any known spatially constrained pre-Quaternary formation. This is due to (1) excellent presser- vation of fossils and the ease with which they are extracted from the poorly lithified sediments as well as (2) the high primary diversity, which is probably due to the tropical reef-associated setting, high alpha diversity, and a wide breadth of habitat types, driving beta diversity. Beta diversity is partitioned into a large species turnover and a small nestedness component. Particularly notable for a fossil assem- blage is the high proportion of molluscs, particularly gastropods. Molluscs comprise 67% of all inver- tebrate species in the Cassian Formation, and the proportion of gastropods (39%) is almost level with modern tropical settings. Studying more 'liberation lagerstätten' like the Cassian Formation can greatly contribute to our understanding of biodiversity of reef basin assemblages and other habitats, and may lead us to rethink the concept of a substantial rise of gastropods in the Cenozoic. Testing which factors drive biodiversity, specifically beta diversity, in the Cassian Formation and other comparable assemblages is the main focus of this thesis. Mean pairwise proportional dissim- ilarity is a very suitable measure to express beta diversity, as it is relatively robust to differences in sample size and grain and can be used to quantify overall beta diversity in a region or between samples as well as along gradients. Beta diversity as pairwise proportional dissimilarity is driven by dominant species. The role of rare species can be neglected in large-scale assessments of beta diversity. This finding not only makes it easier to accumulate larger amounts of data to be used in beta diversity studies but also makes incorrect taxon identification less significant. To disentangle the drivers of biodiversity patterns in the Cassian Formation and another reef- associated soft-bottom environment from the modern Bay of Safaga in the Red Sea, Egypt, beta diversity is evaluated with regard to age, water depth, and geographic distance. Results are compares with a null model to evaluate the stochasticity of community assembly. Only the ten most abundant species per sample are included to determine mean pairwise proportional dissimilarity. As in the previous studies, overall beta diversity is found to be very high in the Cassian Formation. The Bay of Safaga yields a similarly high value. The variation in community composition is found to be independent of geographic or temporal distance. Beta diversity between shallow- and deep-water communities is relatively high. Samples from deeper-water settings yield a slightly higher beta diversity than those from shallower areas. A similar pattern is seen in both assemblages. Although water depth has been a driving factor in other studies on beta diversity, it is not considered to be a major driver in the studied assemblages. Priority effects are postulated to be the main driver of beta diversity in these reef basin assemblages. In summary, results from this thesis show that dominant species drive beta diversity. Species turnover, not species loss, are the ecological cause of beta diversity in the Cassian Formation, probably resulting from the evolution of new species within this heterogeneous environment. The high diversity in the Cassian Formation results from high local diversity and large differences in community composition between assemblages driving beta diversity. Reasons for the high preserved diversity are ascribed to taphonomic conditions. Biodiversity in the Cassian Formation is similar to comparable modern environments. The high number of species and the large proportion of gastropods in the Cassian Formation are close to those found in modern soft-bottom assemblages. Values of alpha, beta and gamma diversity are similarly high. And while the heterogeneity and breadth of environments is thought to increase beta diversity, priority effects may also play an important role. Zusammenfassung Die Entwicklung von Alpha- und Gamma-Diversität während des Phanerozoikums wurde bereits in verschiedenen Studien untersucht, jedoch wurde die Beta-Diversität in paläontologischen Arbeiten meist vernachlässigt. Beta-Diversität – die Variation in der Zusammensetzung zwischen Lebensgemeinschaften oder Ansammlungen von Taxa – ist ein Schlüsselaspekt der Biodiversität und von besonderer Bedeutung, um die Mechanismen von Biodiversitätsmustern aufzudecken. Biodiversitätsmuster von gut erhaltenen fossilen sowie rezenten Lebensgemeinschaften können Rückschlüsse auf ihre Entstehung und Entwicklung über die Zeit ermöglichen, sowie einheitliche Mechanismen der Etablierung von Artengemeinschaften aufdecken. Erschwert wird die Erforschung von Biodiversität über die Zeit durch Unterschiede in der Taphonomie oder in der Probengröße und -intensität. Intensiv beprobte und gut erhaltene Fossilgemeinschaften eignen sich ideal, um Biodiversitätsmuster zu untersuchen und sie mit heutigen Lebensgemeinschaften zu vergleichen. Die Cassian-Formation (Ladinium bis Karnium, Mittel- bis Obertrias), aufgeschlossen in den Dolomiten (südliche Kalkalpen, Nord-Italien), zeichnet sich durch hohe Diversität und sehr gute Erhaltung aus. Sie umfasst Flachwassersedimente und Sedimente aus tieferen Bereichen, abgelagert zwischen Karbonatplattformen in warmen, tropischen Gewässern der westlichen Tethys, vergleichbar mit heutigen tropischen Habitaten. Die Cassian-Formation verfügt somit über eine große Bandbreite an Ablagerungsräumen: Ein küstennahes Rückriff mit Fleckriffen und dazwischen eingeschalteten Sedimenten, eine Karbonatplattform sowie ein flaches und tieferes Riffbecken. Mit 1421 Invertebraten-Arten verfügt die Cassian-Formation über das größte Artenreichtum unter allen bekannten räumlich begrenzten, präquartären Formationen. Dies liegt einerseits an der sehr guten Fossilerhaltung und der Leichtigkeit, mit der diese aus den geringlithifizierten Sedimenten erschlossen werden können, gleichzeitig aber auch an der hohen primären Diversität. Die hohe primäre Diversität resultiert vermutlich aus dem tropischen, riffnahen Habitat, der hohen Alpha-Diversität sowie der Bandbreite an Habitattypen, welche die Beta-Diversität erhöhen. Die Beta-Diversität besteht aus einer großen species turnover Komponente und einer kleinen nestedness Komponente. Besonders hervorzuheben ist der hohe Anteil an Mollusken, insbesondere Gastropoden. Mollusken umfassen 67 % aller Invertebraten-Arten in der Cassian Formation, und der Anteil an Gastropoden (39 %) ist fast so hoch wie in heutigen tropischen Habitaten. Mehr Forschung an sogenannten “Liberation Lagerstätten” wie der Cassian-Formation kann dazu beitragen, unser Wissen um Biodiversität in Riffbeckenfaunen und anderen Habitaten zu erweitern. Dies wiederum kann dazu führen, das Konzept eines erheblichen Anstiegs der Gastropoden im Känozoikum zu überdenken. Das Hauptaugenmerk dieser Arbeit liegt auf der Untersuchung der Faktoren, welche die Biodiversität, insbesondere die Beta-Diversität, in der Cassian-Formation und vergleichbaren Faunengemeinschaften bestimmen. Die mittlere pairwise proportional dissimilarity eignet sich gut als Index für die Beschreibung der Beta-Diversität. Sie ist relativ robust gegenüber Unterschieden in der Probengröße und -aufteilung und kann verwendet werden, um die gesamte Beta-Diversität einer Region oder zwischen Proben sowie
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