The taphonomic and paleoecologic aspects of the Upper Hauterivian to Lower Barremian Agua de la Mula Member of the Agrio Formation (Neuquén Basin, Argentina) were studied in the frame of the sequence stratigraphic paradigm. The Agua de la Mula Member, a ca. 600 m thick succession of highly cyclic marine sediments was surveyed at two localities. Detailed bed-by-bed sedimentologic, stratigraphic, ichnologic, taphonomic and paleoecologic data collection allowed a precise paleoenvironmental, stratigraphic, taphonomic and synecologic interpretation, in a controlled sequence stratigraphic framework. The main architectural stratigraphic component is the Starvation-Dilution Sequence, interpreted as a the effect of a sixth-order, Milankovitch precession-driven cycle. Dilution hemisequences are siliciclastic-dominated and show evidence of depth changes. Starvation hemisequences show a diverse variation of mixed carbonate-siliciclastic facies that is linked to sequence stratigraphy. Ammonite-based biostratigraphy was revised and new knowledge proposed. The stratigraphic framework was improved by combining biostratigraphy, sequence stratigraphy and event stratigraphy. Nine main sequences were described, linked to other stratigraphic markers and correlated with other sequence stratigraphic charts. Several orders of cyclicity were inferred. Third- and fourth-order sequences are the major sequences, not subordinated to higher hierarchies (lower order). Precession, obliquity, and short and long eccentricity cycles of the Milankovitch band are proposed. Among the different sequence stratigraphic models the transgression-regression model fits the majority of the sequences described in this work. The depositional-sequence model could be applied only to the first third-order sequence, in which the true sequence boundary is identifiable. Starvation-dilution sequences, however, are composed by to components that are not completely explained by those models. Starvation hemisequences developed in intermediate to deep settings record the transgressive phase as well as the earLy regressive one without visible stratigraphic boundaries. 112 samples with 22,572 individuals were grouped into fifteen fossil associations and one assemblage that reflect the interaction of different factors: age, position in major, medium and starvation dilution sequences and, linked to sequence stratigraphy, depth, oxygen availability, rate of terrigenous input, water agitation, and substrate conditions. Temporary possible reduction in oxygen content is inferred based on all sources of available evidence. Organic buildups are briefly described and their development interpreted in terms of the sequence stratigraphic framework. Vertical patterns of replacement of fossil associations are described and related to sequence stratigraphy. Five types of skeletal concentrations represent the diversity of coquinas decribed in this study. Type 1, 2, 4 and 5 correspond to starvation hemisequences deposited in progressively shallower settings, from basin to inner ramp. Type 3 is embedded into dilution hemisequences and inferred to be linked to shell bed type I of Kidwell (1985). Types 1 and 2 correspond to transgression, maximum flooding and early regression without distinction. Type 4A as well as Type 5 are interpreted as onlap shell beds (Kidwell 1991a) or early TST shell beds (Fürsich and Pandey 2003). Type 4B corresponds to the MFZ shell bed (Fürsich and Pandey 2003) or mid-cycle shell bed (Abbott 1997), while Type 4C to the downlap shell bed (Kidwell 1991a). Time-averaging of shell beds was assessed with precision as the time involved in the deposition of the starvation hemisequences could be inferred. All shell beds comprise within-habitat assemblages forming within a few thousand years, with little environmental condensation. The fossilization of the marine calcareous shells is modelled as a series of steps called windows: environmental, destructional, burial and diagenetic. The “diagenetic window” is the most relevant. Connected to this it is proposed that carbonate dissolution is the primary control on the development of shell beds, as has been proposed before (Fürsich 1982; Fürsich and Pandey 2003). The interpretative power resulting from combining several lines of evidence, e.g., facies analysis, sequence stratigraphy, biostratigraphy, trace fossil analysis, paleoecology and taphonomy, and unravelling their multiple relationships, are the most relevant conclusions of this study.

The invertebrate trace fossils from the Keuper (Upper Triassic) of the southern part of the Germanic Basin are revised. The Keuper sediments of the Germanic Basin are predominantly composed of rocks representing various nonmarine environments dominated by red-bed facies. The Würzburg Formation, the Stuttgart Formation, and the Hassberge Formation, all representing deposits of extended river systems, contain the richest ichnofauna. Trace fossil abundance is generally low and their occurrence is scattered. The studied material can be assigned to 28 ichnogenera, 38 ichnospecies, and 6 vernacular forms. Among the described trace fossils are one new ichnogenus and three new ichnospecies. Apart from the revision of the invertebrate trace fossils from the Keuper numerous related ichnotaxa from various localities and ages have been studied and revised. In the course of these studies several ichnotaxa are synonymised, lowered in rank, and new ichnogenera, subichnogenera, and ichnospecies are suggested. In addition, general guidelines for naming, and methodologies for studying invertebrate trace fossils are presented. The palaeoecology of three ichnocoenoses, one from the Würzburg Formation and two from the Hassberge Formation in Lower Franconia are briefly discussed. / Die Invertebratenspuren aus dem Keuper (Obere Trias) des südlichen Germanischen Beckens werden revidiert. Die Sedimente des Germanischen Keupers bestehen zum überwiegenden Teil aus verschiedenen nichtmarinen Rotsedimenten. Die Würzburg Formation, die Stuttgart Formation und the Hassberge Formation werden als Ablagerungen ausgedehnter Flusssysteme interpretiert und enthalten die reichhaltigsten Spurenfaunen. Die Häufigkeit von Spurenfossilien ist generell niedrig und ihre Verbreitung ist punktuell. Das untersuchte Material kann 28 Ichnogattungen und 38 Ichnoarten zugewiesen werden. Sechs Spurenfossiltypen konnten taxonomisch nicht zugewiesen werden. Unter den beschriebenen Formen sind eine neue Ichnogattung und drei neue Ichospezies. Neben der Revision der Keuperspuren wurden zahlreiche verwandte Spurentaxa verschiedener räumlicher und zeitlicher Herkunft detailiert revidiert. Im Zuge dieser Revision wurden einige Taxa synonymisiert oder in ihrem taxonomischen Rang erniedrigt sowie neue Ichnotaxa vorgeschlagen. Darüber hinaus werden Leitlinien zur Nomenklatur von Spurenfossilien sowie generelle Verfahrensweisen zum Studium von Invertebratenspurenfossilien vorgestellt. Die Palökolgie dreier Ichnozönosen aus Unterfranken, eine aus der Würzburg Formation und zwei aus der Hassberge Formation werden kurz diskutiert.

The quantification of disparity is an important aspect of recent macroevolutionary studies, and it is usually motivated by theoretical considerations about the pace of innovation and the filling of morphospace. In practice, varying protocols of data collection and analysis have rendered comparisons among studies difficult. The basic question remains, How sensitive is any given disparity signal to different aspects of sampling and data analysis? Here we explore this issue in the context of the radiation of the echinoid order Spatangoida during the Cretaceous. We compare patterns at the genus and species levels, with time subdivision into subepochs and into stages, and with morphological sampling based on landmarks, traditional morphometrics, and discrete characters. In terms of temporal scale, similarity of disparity pattern accrues despite a change in temporal resolution, and a general deceleration in morphological diversification is apparent. Different morphometric methods also produce similar signals. Both the landmark analysis and the discrete character analysis suggest relatively high early disparity, whereas the analysis based on traditional morphometrics records a much lower value. This difference appears to reflect primarily the measurement of different aspects of overall morphology. Disparity patterns are similar at both the genus and species levels. Moreover, inclusion or exclusion of the sister order Holasteroida and the stem group Disasteroida in the sampled morphospace did not affect proportional changes in spatangoid disparity. Similar results were found for spatangoid subclades vis-à-vis spatangoids as a whole. The relative robustness of these patterns implies that the choice of temporal scale, morphometric scheme, and taxonomic level may not affect broad trends in disparity and the representation of large-scale morphospace structure.

DIe Dissertationsschrift enthält mehrere teils in wissenschaftlichen Journals veröffentlichte Studien, die im Rahmen der kumulativen Dissertation entstanden. Im Fokus dieser Studien und der vorliegenden Dissertationsschrift steht die Untersuchung und Rekonstruktion Quartärer Umweltveränderungen in terrestrischen Ökosystemen. Dabei wurden Äolianit-paläobodensequenzen untersucht, anhand derer mithilfe von fossilen Gastropoden und deren stabilen Isotopensignalen Umweltinformationen abgeleitet werden.:TABLE OF CONTENT ACKNOWLEDGEMENT ABSTRACT KURZFASSUNG TABLE OF CONTENT 1 INTRODUCTION 1.1 RELEVANCE AND OBJECTIVE OF STUDYING FOSSIL GASTROPODS IN QUATERNARY DEPOSITS 1.2 MAJOR RESEARCH QUESTIONS AND METHODOLOGICAL APPROACH 1.3 STUDY SITES 1.4 REFERENCES 2 QUATERNARY GASTROPOD FAUNAS ON THE EASTERN CANARY ISLANDS AND INDICATIONS OF ENVIRONMENTAL CHANGES 2.1 INTRODUCTION 2.2 GEOGRAPHICAL SETTING 2.3 METHODS 2.4 RESULTS 2.5 DISCUSSION 2.6 CONCLUSIONS 2.7 ACKNOWLEDGEMENT 2.8 REFERENCES 2.9 SUPPLEMENTARY MATERIAL 3 QUATERNARY GASTROPOD FAUNAS IN SOUTHERN CAUCASIA AND INDICATIONS ON ENVIRONMENTAL CHANGES 3.1 INTRODUCTION 3.2 STUDY AREA 3.3 METHODS 3.4 RESULTS 3.5 DISCUSSION 3.6 CONCLUSION 3.7 ACKNOWLEDGEMENT REFERENCES 3.9 SUPPLEMENTARY MATERIAL 4 MALACOLOGICAL INVESTIGATIONS ON EASTERN CANARY SEDIMENT ARCHIVES –STABLE ISOTOPE INTERPRETATION IN AN OZEANIC SETTING 4.1 INTRODUCTION 4.2 GEOGRAPHICAL SETTING AND STATE OF KNOWLEDGE 4.3 METHODS 4.4 RESULTS 4.5 DISCUSSION 4.6 CONCLUSIONS 4.7 ACKNOWLEDGEMENT 4.8 REFERENCES 4.9 SUPPLEMENTARY MATERIAL 5 MALACOLOGICAL INVESTIGATIONS AT SOUTH CAUCASIAN SEDIMENT ARCHIVES –STABLE ISOTOPE INTERPRETATION IN A CONTINENTAL SETTING 5.1 INTRODUCTION 5.2 GEOGRAPHICAL SETTING AND STATE OF KNOWLEDGE 5.3 METHODS 5.3.1 Fieldwork 5.3.2 Laboratory Analyses 5.4 RESULTS 5.6 CONCLUSION 5.7 REFERENCES 5.8 SUPPLEMENTARY MATERIAL 6 SUMMARY AND SYNTHESIS 6.1 COMPARATIVE CONSIDERATION OF THE DIFFERENT SETTINGS 6.2 MAJOR CONCLUSIONS 6.3 PERSPECTIVE 6.4 REFERENCES

Four sections of the Galala and Maghra El Hadida formations on the footwalls of the slopes of the northern and southern Galala plateaus in Wadi Araba (Eastern Desert) have been measured and sampled in great detail. The Galala Formation is ranging in thickness from 55 to 95 meters. It unconformably overlies the Malha Formation which forms the base of the studied sections. The upper boundary of the Galala Formation is characterized by a major unconformity which separates it from the overlying the Maghra El Hadida Formation. The Galala Formation can be subdivided into five shallowing-upward cycles, each cycle starting with deep-lagoonal, marly-silty deposits at the base and grading into highly fossiliferous shallow-lagoonal limestones at the top. Only the basal part of the Galala Formation consists of unfossiliferous, greenish sandy siltstones intercalated with thin cross-bedded, bioturbated, fine- to medium-grained sandstones. Despite the lack of biostratigraphic markers in that lower part, its age can be assigned to the late Middle Cenomanian, since the conformably overlying strata contain the ammonite Neolobites vibrayeanus (D’ORBIGNY), the index marker of the early Upper Cenomanian which extends into the top of the formation. The measured thickness of the overlying Maghra El Hadida Formation is ranging from 59 to 118 meters. This formation starts with the Ghonima Member, introduced in this work to distinguish a brown, fine- to medium-grained calcareous sandstone unit in its lower part. The Ghonima Member is erosionally incised into the Galala Formation, explaining its strong lateral variability in thickness, ranging from 3 to 21 meters. It is mostly unfossiliferous except for irregular bioturbation in its upper part. The Ghonima Member is assigned to the middle Upper Cenomanian, based on its stratigraphic position between the lower Upper Cenomanian Neolobites vibrayeanus Zone and the overlying upper Upper Cenomanian Metoicoceras geslinianum and Vascoceras cauvini zones. This means that the lower part of the Maghra El Hadida Formation, about 20 – 30 m thick, accumulated during the latest Cenomanian and that the base of the formation does not coincide with the base of the Turonian as commonly believed. The overlying succession of the Maghra El Hadida Formation is characterized by an increase of carbonate content, represented by yellow, soft marls intercalated with fine-grained wacke- to packstones containing a highly fossiliferous ammonite assemblage of the upper Upper Cenomanian and Lower Turonian (zones of Vascoceras proprium, Choffaticeras spp., and Wrightoceras munieri). The Middle Turonian part of the Maghra El Hadida Formation consists of poorly fossiliferous, thick-bedded yellowish marls with upward-increasing silt content, showing occasional intercalations of medium- to coarse-grained sandstones with hummocky cross-stratification. The topmost part of the Maghra El Hadida Formation consists of brownish, medium-grained sandstones topped by fossiliferous marly limestones yielding the Upper Turonian zonal ammonite Coilopoceras requienianum (D’ORBIGNY). Based on sequence stratigraphic analyses, four complete 3rd order depositional sequences and the lower part of a fifth one, each bounded by major unconformities, can be recognized: depositional sequence DS WA 1 (upper Middle – lower Upper Cenomanian) includes the entire Galala Formation, while the Maghra El Hadida Formation comprises all the overlying depositional sequences: DS WA 2 (upper Upper Cenomanian – Lower Turonian) reaches from the base of the Metoicoceras geslinianum Zone to the top of Wrightoceras munieri Zone, DS WA 3 and DS WA 4 comprise the Middle Turonian, while Upper Turonian sequence DS WA 5 is not complete. The stratigraphic positions of the recognized sequence 2 boundaries SB WA 1 to SB WA 5 match well with contemporaneous sequence boundaries known from Europe and elsewhere. The stacking pattern of the basic cycles and bundles of the Galala Formation (5:1) and the Maghra El Hadida Formation (4:1) strongly suggest an orbital forcing by MILANKOVITCH periodicities. The Galala Formation is composed of five 5th-order bundles which equal to ~500 kyr, each bundle equals to ~100 kyr (short eccentricity). Every bundle has five basic (6th-order) cycles, each one representing ~20 kyr (precession). Based on this precession-short eccentricity syndrome, the accumulation rate of the Galala Formation therefore accounts for about 19 cm/kyr. The rate of sea-level fall at sequence boundary SB WA 2 (equivalent to the quasi-global mid-Late Cenomanian SB Ce V) estimated is with 35 cm/kyr which can be explained only by glacio-eustasy. The Upper Cenomanian and Lower Turonian part of the Maghra El Hadida Formation is considered to equal to ~1200 kyr, based on the existence of three 4th-order bundles with an inferred duration of ~400 kyr for each bundle (long eccentricity of the MILANKOVITCH Band). Every bundle consists of four basic cycles with a duration of ~100 kyr. This means that the upper Cenomanian part of the Maghra El Hadida Formation is equivalent to ~400 kyr, while the Lower Turonian (consisting of the two upper bundles) lasted 800 kyr. This matches well with the recently proposed 785 kyr duration of the Early Turonian (SAGEMAN et al., 2006; VOIGT et al., 2008) and contradicts the 1300 kyr according to the standard time scale of GRADSTEIN et al. (2004). According to this temporal constrains, the accumulation rate of the Maghra El Hadida Formation is about 4.25 cm/kyr. In addition, based on the cyclostratigraphic analysis, the range of the Early Turonian genus Choffaticeras (HYATT) is equivalent to ~325 kyr and morphological changes within its lineage can be quantified. The macrobenthos (bivalves, gastropods, echinoids) and cephalopods of the Galala and Maghra El Hadida formations were identified and illustrated in 24 figures. The ammonite taxonomy and palaeobiogeographic distribution is discussed in detail. Four genera and eight ammonite species are recorded from Egypt for the first time. The microfloral and -faunal assemblage identified in thin sections revealed two species of dasycladalean algae, two species of udoteacean algae, five species of benthic foraminifera, and two species of crustacean microcoprolites. The six facies types of the upper Middle – Upper Cenomanian Galala Formation document largely open-lagoonal, warm water conditions, while the depositional environment of the Upper Cenomanian – Turonian Maghra El Hadida Formation (16 facies types) is suggested to range from a deep-subtidal to intertidal.

The Middle and Upper Jurassic sedimentary successions of Alborz in northern Iran and Koppeh Dagh in northeastern Iran comprise four formations; Dalichai, Lar (Alborz) and Chaman Bid, Mozduran (Koppeh Dagh). In this thesis, the biostratigraphy, lithostratigraphy, microfacies, depositional environments and palaeobiogeography of these rocks are discussed with special emphasis on the abundant ammonite fauna. They constitute a more or less continuous sequence, being confined by two tectonic events, one at the base, in the uppermost part of the Shemshak Formation (Bajocian), the so-called Mid-Cimmerian Event, the other one at the top (early Cretaceous), the so-called Late-Cimmerian Event. The lowermost unit constitutes the uppermost member of a siliciclastic and partly continental depositional sequence known as Shemshak Formation. It contains a fairly abundant ammonite fauna ranging in age from Aalenian to early Bajocian. The following unit (Dalichai Formation) begins everywhere with a significant marine transgression of late Bajocian age. The following four sections were measured: The Dalichai section (97 m) with three members; the Golbini-Jorbat composite section (449 m) with three members of the Dalichai Formation (414 m) and two members of the Lar Formation (414 m); the Chaman Bid section (1556 m) with seven members, and the Tooy-Takhtehbashgheh composite section (567 m) with three members of the Chaman Bid Formation (567 m) and four members of the Mozduran Formation (1092 m). Altogether, 80 species of ammonites from the Dalichai and Chaman Bid formations belonging to 30 genera and 16 families are described. Among the taxa Phylloceratidae are most abundant, followed by Ataxioceratidae, Perisphinctidae, and Cardioceratidae. Pachyceratidae are the least common family. The ammonite fauna is of low diversity and is concentrated in several levels. Some of the ammonite genera and species are recorded from Iran for the first time. These include Pachyceras lalandei, Cardioceras praecordatum, Microbajocisphinctes sp., Geyssantia geyssanti, Larcheria schilli, Passendorferia sp., Sequeirosia sp., Phanerostephanus subsenex, Nothostephanus sp., Nannostephanus cf. subcomutus, Parawedekindia callomoni, Physodoceras sp., Extrenodites sp.. Biostratigraphically, thirty ammonite zones have been recognized for the Middle and Upper Jurassic successions at the four studied sections. Based on ammonites, the Dalichai Formation ranges from the Upper Bajocian to Callovian (Dalichai section) and from the Upper Bajocian to Lower Tithonian (Golbini-Jorbat section), the Chaman Bid Formation ranges from the ?Bathonian to Lower Tithonian (Chaman Bid section) and from the Upper Bajocian to Middle Kimmeridgian (Tooy-Takhtehbashgheh section), the Lar Formation ranges from the Middle to Upper Tithonian (Golbini-Jorbat section), and the Mozduran Formation from the Upper Kimmeridgian to ?Tithonian. Forty-four Microfacies types are briefly described. They were grouped into 16 facies associations, which then were interpreted in terms of their palaeoenvironments. They are part of a carbonate system consisting of a platform and adjacent slope to basin. Five major environments are represented: Tidal flat, shelf lagoon, and platform margin barrier as parts of the carbonate platform, and slope to basin representing open marine conditions. The sediments of the Dalichai and Chaman Bid formations are the slope and basinal sediments of the diachronous Lar and Mozduran formations, which formed an extensive carbonate platform in the Middle and Upper Jurassic. / In der sedimentären Abfolge des Mittel- und Oberjura im Alborz (nördlicher Iran) und im Koppeh Dagh (nordöstlicher Iran) werden vier Formationen ausgeschieden: Dalichai- und Lar-Formation im Alborz und Chaman Bid- und Mozduran-Formation im Koppeh Dagh. Die Biostratigraphie, Lithostratigraphie, Mikrofazies, Ablagerungsbedingungen und Paläobiogeographie der Abfolgen werden diskutiert, unter besonderer Berücksichtigung der reichen Ammonitenfauna. Die Abfolgen stellen eine mehr oder wenig kontinuierliche sedimentäre Sequenz dar, die an der Basis und am Top von bedeutenden überregionalen tektonischen Unconfomities begrenzt wird, verursacht durch die Mittel-Cimmerischen Bewegungen im Bajoc und die Spät-Cimmerischen Bewegungen in der frühen Kreide. Die folgenden vier Profile wurden bearbeitet: das Dalichai-Typprofil (97 m) mit drei Membern; das zusammengesetzte Golbini-Jorbat-Profil (449 m) mit drei Membern der Dalichai-Formation (414 m) und zwei Membern der Lar-Formation (414 m); das Chaman Bid-Typprofil (1556 m) mit sieben Membern, und das zusammengesetzte Tooy-Takhtehbashgheh-Profil (567 m) mit drei Membern der Chaman Bid-Formation (567 m) und vier Membern der Mozduran-Formation (1092 m). Insgesamt werden 80 Ammonitenarten aus der Dalichai- und der Chaman Bid-Formation beschrieben, die zu 30 Gattungen und 16 Familien gehören. Die Phylloceratidae zeigen die höchste Abundanz, gefolgt von den Ataxioceratidae, Perisphinctidae und Cardioceratidae. Die Pachyceratidae sind die am wenigsten häufige Familie. Die Ammonitenfauna ist gering divers und an bestimmte Horizonte gebunden. Die folgenden Ammoniten-Gattungen und -Arten werden zum ersten mal aus dem Iran beschrieben: Pachyceras lalandei, Cardioceras praecordatum, Microbajocisphinctes sp., Geyssantia geyssanti, Larcheria schilli, Passendorferia sp., Sequeirosia sp., Phanerostephanus subsenex, Nothostephanus sp., Nannostephanus cf. subcomutus, Parawedekindia callomoni, Physodoceras sp., Extrenodites sp.. 30 Ammoniten-Biozonen konnten in der Mittel- bis Oberjura-Abfolge erkannt werden. Basierend auf der Ammoniten-Biostratigraphie kann die stratigraphische Reichweite der Formationen präzisiert werden: die Dalichai-Formation reicht vom Oberbajoc bis zum Callov (Dalichai-Profil) und vom Oberbajoc bis zum Untertithon (Golbini-Jorbat-Profil); die Chaman Bid-Formation reicht vom ?Bathon bis ins Untertithonian (Chaman Bid-Profil) und vom Oberbajoc bis Mittelkimmeridge (Tooy-Takhtehbashgheh-Profil); die Lar-Formation reicht vom Mittel- bis ins Obertithon (Golbini-Jorbat-Profil); die Mozduran-Formation reicht vom Oberkimmeridge bis zum ?Tithon. 44 Mikrofaziestypen, gruppiert in 16 Fazies-Assoziationen, werden kurz beschrieben und im Hinblick auf ihre Bildungsbedingungen interpretiert. Sie dokumentieren ein Karbonatsystem bestehend aus einer Plattform, einem Hang und einem Becken. Fünf Hauptablagerungsbereiche können ausgeschieden werden: Karbonatwatten, Schelflagune und Plattformrand als Teile der ehemaligen Karbopnatplattform sowie Hang und Becken des offenmarinen Bereiches. Die Sedimente der Dalichai- und Chaman Bid-Formationen repräsentieren Hang- und Becken-Sedimente einer sich diachron in Mittel- und Oberjura bildenden Karbonatplattform, die durch die Lar- und Mozduran-Formationen dokumentiert wird.

The Upper Cretaceous Ajlun Group (Cenomanian-Turonian) of southern/south-eastern Jordan has been analysed in 15 detailed sections with thicknesses between 40 m and 200 m. Taxonomic, palaeoecological, taphonomic, and sedimentological aspects were taken into account. During the early Upper Cretaceous the study area was situated at the south-eastern margin of the Tethys Ocean, between the palaeo-shoreline in the south-east and an offshore carbonate platform in the west. Thus, the measured sections include a complete facies succession from terrestrial-dominated environments via marginal marine siliciclastics to an area of carbonate precipitation. So far, very little is known about the fauna and the depositional environment of the group, especially of the transitional marginal marine part. Also, in depth studies of the Cretaceous fauna of southern Jordan are very rare. Therefore, the benthic fauna of the area is described in an extensive taxonomic chapter. It consists of 117 taxa, 77 of which are bivalves, 22 gastropods, 9 echinoids, and 4 corals. The phyla Porifera, Bryozoa, and Brachiopoda are represented by 1 species each. Additionally, at least two species of decapod crustaceans were found. One bivalve species is new: Anthonya jordanica from Cenomanian claystones of the eastern study area. 41 quantitative samples of the benthic invertebrate fauna were grouped into nine associations and three assemblages by means of a Q-mode cluster analysis. These are described as remnants of former communities and their environments are discussed. Salinity and substrate consistency are assumed to have been the most important environmental parameters controlling the faunal distribution. The overall palaeo-environment is discussed on the basis of sedimentological and palaeoecological results. It was primarily influenced by the morphology of the sea floor, sediment supply, and salinity of the sea water. / Mit Hilfe von 15 detaillierten Profilen mit Mächtigkeiten zwischen 40 m und 200 m wird die oberkretazische Ajlun-Gruppe (Cenoman-Turon) im Süden/Südosten Jordaniens auf Fazies- und Faunenverteilung untersucht. Dabei wird besonderes Augenmerk auf Taxonomie, Paläoökologie, Taphonomie und Sedimentologie gelegt. Während der frühen Oberkreide lag das Untersuchungsgebiet am Südostrand der Tethys, im Bereich zwischen Küstenlinie und einer vorgelagerten Karbonatplattform. Die Profile liefern einen kompletten Schnitt vom terrestrischen Ablagerungsraum im Südosten über stark siliziklastisch geprägte randlich marine Verhältnisse zu vollmarinen Bereichen mit Karbonatsedimentation. Insgesamt kann gerade der klastisch dominierte Übergangsbereich als bislang nahezu unbearbeitet gelten. Auch paläontologische Untersuchungen fehlen für dieses Zeitintervall bis auf wenige Ausnahmen nahezu völlig. Daher wurde zunächst in einem taxonomischen Teil eine Bestandsaufnahme der Fauna (mit Schwerpunkt auf Benthosorganismen) gemacht. Für das Arbeitsgebiet konnten 2 Ammonoideen und 117 Benthosarten bestimmt werden, die sich aus 77 Muschel-, 22 Schnecken-, 9 Seeigel- und 4 Korallentaxa, sowie je einen Vertreter der Schwämme, Bryozoen und Brachiopoden zusammensetzen. Zusätzlich wurden mindestens zwei neue Arten decapoder Krebse gefunden. Bei den Bivalven konnte die neue Art Anthonya jordanica sp. nov. aufgestellt werden. Der allgemein mäßige Erhaltungszustand der Fauna ließ die Aufstellung weiterer neuer Arten nicht zu, obwohl unter den in offener Nomenklatur geführten Taxa mit Sicherheit einige neue Arten sind. Nach einer Fazies-Analyse der verschiedenen lithologischen Einheiten wurden mit Hilfe einer paläoökologischen Analyse neun Faunenassoziationen und drei Faunenvergesellschaftungen ausgeschieden. Diese statistischen Einheiten, überwiegend Überreste ehemaliger Lebensgemeinschaften, werden beschrieben und die sie bestimmenden Milieuparameter diskutiert. Die wichtigsten, limitierenden Faktoren, welche die Verteilung der Faunengemeinschaften entscheidend steuern, sind die Salinität und das Substrat.

The aim of this study was to assess distribution patterns of articulate brachiopods during the Mesozoic. Exploratory and confirmatory multivariate analyses in this study evaluate whether environmental preferences of brachiopods and bivalves are substantially distinct and whether structure of their communities significantly differ. Specifically, the hypothesis being tested is that differential abundances of Mesozoic brachiopods and bivalves are not related to varying substrate properties only, but also to varying food supply, turbidity and oxygen levels. This hypothesis was evaluated with quantitative data gathered in various field areas and time intervals. They include the Upper Triassic deposits of the West Carpathians and Eastern Alps, the Lower and Middle Jurassic deposits of Morocco, the Middle and Upper Jurassic deposits of the West Carpathians, the Upper Jurassic deposits of the Franconian and Swabian Alb, and the Upper Jurassic deposits of the Swiss Jura. The main conclusion is that brachiopod-dominated communities are characterized by a unique guild structure, with dominance of trophic groups with low metabolic requirements or adapted to nutrient-poor or oxygen-poor conditions. For example, brachiopod co-occured more commonly with epifaunal than with infaunal bivalves in soft-bottom environments. Abundances of brachiopods correlate mostly negatively with increasing proportions of terrigenous admixture (i.e., with increasing amount of land-derived nutrient supply and turbidity). / Das Ziel dieses Projekt war es, das Verbreitungsmuster im Erdmittelalter von artikulaten Brachiopoden zu interpretieren. Es basiert hauptsächlich auf der Charakterisierung der Struktur von Benthos-Gemeinschaften mit Brachiopoden und Bivalven. Ausgegangen wird dabei von der Hypothese, dass die Häufigkeit des Auftretens von Brachiopoden und Muscheln nicht allein mit Substratgradienten erklärt werden kann, sondern auch ein Zusammenhang mit Nährstoff-, Sauerstoff- und Turbiditätgradienten besteht. Diese Hypothese war mit Geländedaten aus spezifischen Gebieten (Obertrias der Westkarpaten und Ostalpen, Unterjura und Mitteljura in Marokko, Mitteljura und Oberjura in den Westkarpaten, Oberjura der Fränkische und Schäbische Alb in Deutschland, und Oberjura in Schweiz). Das wichtigste Ergebnis ist dass Brachiopoden-dominierte Gemeinschaften spezifische ökologische Strukturen zeigen und durch die Dominanz von Benthosgruppen charakterisiert sind, welche sind an nährstoffarme oder sauerstoffarme Bedingungen angepasst sind (zum Beispiel durch ein effizientes Filtriersystem) oder niedrige metabolische Ansprüche haben. Zum Beispiel kommen epifaunale Bivalven mit Brachiopoden wesentlich häufiger vor als infaunale Bivalven. Dies kann nicht nur mit unterschiedlichen Substratpräferenzen der infaunalen und epifaunalen Bivalven erklärt werden, weil diese Vergesellschaftungen häufig in mikritischen Wackestones und Floatstones vorkommen.Außerdem korreliert die Häufigkeit der Brachiopoden positiv mit der Abnahme des siliziklastischen Anteil (d.h., mit der Abnahme nährstoffreicher Partikel und Wassertrübe).

Auf Grundlage umfangreicher Neuaufsammlungen wird die Muschelfauna der Nayband-Formation monographisch dokumentiert, wobei die folgenden 21 Arten neu aufgestellt werden: Palaeonucula biacuta, Trigonucula goniocostata, Nuculana (Nuculana) naibandensis, Parallelodon tectum, Mysidiella imago, Gervillia (Cultriopsis) canalis, Isognomon repini, Indopecten uninodosus, Antiquilima hians, Weixiella lutensis, Gruenewaldia iranica, Gruenewaldia magna, Myophoricardium subquadratum, Praeconia matura, Coelopis (Coelopis) aurea, Opis (Trigonopis?) eumorpha, Opis (Trigonopis?) douglasi, Palaeocardita iranica, Palaeocardita stoecklini, Palaeocardita carinata und Antiquicorbula (gen. nov.) concentrica. 51 Muschelarten der Nayband-Formation werden als bereits beschriebene Arten identifiziert, 32 Taxa auf Grund unvollständiger Erhaltung in offener Nomenklatur geführt. Die höchste Diversität besitzen die Pteriomorphia mit 51 Arten (49 %), gefolgt von den Heterodonta mit 22 Arten (21,2 %), den Palaeoheterodonta und Anomalodesmata mit je 12 Arten (11,5 %) und schließlich den Palaeotaxodonta mit 7 Arten (6,7 %). In der Familie Permophoridae wird die neue Gattung Healeya, in der Familie Corbulidae die neue Gattung Antiquicorbula und in der Familie Trigoniidae die neue Untergattung Trigonia (Modestella) vorgeschlagen. Die Diagnosen der Gattungen Trigonucula, Mysidiella, Primahinnites, Indopecten, Gruenewaldia und Vietnamicardium werden revidiert. Die Gattung Primahinnites wird den Aviculopectinidae an Stelle der Prospondylidae zugeordnet, die Gattung Weixiella den Permophoridae an Stelle der Pachycardiidae. Die Familie Mysidiellidae ist u. a. auf Grund schalenmikrostruktureller Merkmale vermutlich zu den Ambonychioidea zu stellen und nicht, wie bisher meist angenommen, zu den Mytiloidea. Die von CARTER (1990) angeführten Argumente für eine Zugehörigkeit der Permophoridae zu den Modiomorphoidea werden durch das untersuchte Material bestätigt. Bei den Prospondylidae, Plicatulidae, Dimyidae und Ostreidae findet die von HAUTMANN (2001) gegebene Revision dieser Familien Anwendung, wobei die Diskussion über die taxonomische Relevanz der Schalenmikrostruktur vertieft wird. Aus funktionsmorphologischen Überlegungen, Vergleichen mit rezenten Arten und dem jeweiligen Fundzusammenhang wird die Lebensweise der einzelnen Taxa rekonstruiert. Obwohl die meisten Muschelarten der Nayband-Formation unverfestigte Substrate bewohnten, erreichte erstmals in der Erdgeschichte auch die Besiedlung von Hartsubstraten durch Muscheln eine größere Bedeutung. Die Erschließung dieses Lebensraumes gelang taxonomisch unabhängigen Gruppen mit Hilfe verschiedener Anpassungen (byssate Verankerung, Zementation, chemisches Bohren) und Besiedlungsstrategien (Begleitfauna in Korallen- und Schwammriffen, eigenständige Riffbildung, Epökie). Ein hoher Endemismusgrad insbesondere bei flachmarinen Muschelarten rechtfertigt es, die Tethys in der Obertrias als eigenes Faunenreich anzusehen. Die Stellung der Nayband-Muschelfauna innerhalb der Tethys lässt sich wegen der ungeklärten paläogeographischen Lage vieler Obertriasvorkommen nur im Rahmen einer Gesamtanalyse der Faunenbeziehungen verstehen, die ihrerseits wiederum eine unabhängige Bewertung konkurrierender plattentektonischer Modelle erlaubt. Das Verbreitungsmuster von Muschelgattungen und -arten zeigt die Existenz einer Westtethys- und einer Osttethys-Provinz auf, wobei letztere in eine Tethysnordrand- und Tethyssüdrand-Subprovinz zerfällt. Der Tethysnordrand-Subprovinz sind u. a. Iran, Yunnan, Vietnam und Burma zuzurechnen, nicht jedoch der Lhasa-Block, der enge Faunenbeziehungen zur Tethyssüdrand-Subprovinz zeigt. Die paläobiogeographische Analyse stützt damit die relativ neue These von der Existenz einer „Känotethys“ bzw. „Tethys III“, durch deren Öffnung der Lhasa-Block erst am Ende der Trias von Gondwana abgetrennt wurde. Die faunistische Eigenständigkeit des Westtethysraumes geht vermutlich auf die engräumige fazielle Gliederung dieses Gebietes zurück, durch die einerseits Faunenaustausch behindert, andererseits Speziation und damit letztlich auch die Neuentstehung höherer Taxa begünstigt wurde. Die Endemisten dieser Provinz waren vom Massenaussterben an der Trias/Jura-Grenze weit weniger betroffen als jene der Osttethys-Provinz und konnten sich mit der Unterjura-Transgression vielfach auch überregional ausbreiten. Insgesamt zeigt sich, dass für das Verbreitungsmuster obertriadischer Muscheln in der Tethys die Prozesse Artenneuentstehung, Ausbreitung und Vikarianz gleichermaßen, wenn auch mit lokal unterschiedlicher Bedeutung verantwortlich waren. / The Upper Triassic (Norian-Rhaetian) Nayband-Formation of east-central Iran contains an exceptionally well-preserved bivalve fauna with more than 100 species adapted to a large spectrum of habitats. Based on new material, the present thesis provides a monographic description and taxonomic discussion of this fauna, gives a reconstruction of its mode of life and clarifies its palaeobiogeographic relations. Out of 104 documented taxa, the following 21 species are new: Palaeonucula biacuta, Trigonucula goniocostata, Nuculana (Nuculana) naibandensis, Parallelodon tectum, Mysidiella imago, Gervillia (Cultriopsis) canalis, Isognomon repini, Indopecten uninodosus, Antiquilima hians, Weixiella lutensis, Gruenewaldia iranica, Gruenewaldia magna, Myophoricardium subquadratum, Praeconia matura, Coelopis (Coelopis) aurea, Opis (Trigonopis?) eumorpha, Opis (Trigonopis?) douglasi, Palaeocardita iranica, Palaeocardita stoecklini, Palaeocardita carinata, and Antiquicorbula (gen. nov.) concentrica. A total of 51 species are already known, and 32 incompletely preserved taxa are presented in open nomenclature. The most diverse group are the Pteriomorphia with 51 species (49 %), followed by the Heterodonta with 22 species (21,2 %), the Palaeoheterodonta and Anomalodesmata with 12 species (11,5 %) each, and finally the Palaeotaxodonta with 7 species (6,7 %). On the basis of the examined material, the diagnoses of the genera Trigonucula, Mysidiella, Primahinnites, Indopecten, Gruenewaldia and Vietnamicardium are revised, and the following new genera and subgenera are proposed: Healeya within the family Permophoridae, Antiquicorbula within the family Corbulidae and Trigonia (Modestella) within the family Trigoniidae. The genus Primahinnites is transferred from Prospondylidae to Aviculopectinidae and the genus Weixiella from Pachycardiidae to Permophoridae. On grounds of morphological and shell microstructural characters, the Mysidiellidae are classified with the Ambonychioidea rather than with the Mytiloidea. The recently proposed integration of the Permophoridae within the Modiomorphoidea is confirmed by the examined material. The revision of the families Prospondylidae, Plicatulidae, Dimyidae and Ostreidae by HAUTMANN (2001) is adopted and extended with respect to the taxonomic relevance of the shell microstructure. The mode of life of the various taxa is reconstructed on the basis of field observations, constructional morphology and comparisons with Recent species. Although most species of the Nayband-Formation occupied habitats characterized by soft substrates, colonization of hard substrates by bivalves achieved significant importance for the first time in Earth´s history. Different taxonomic groups succeeded in entering this habitat by means of various adaptations (byssal attachment, cementation, chemically boring) and settling strategies (dwelling within coral and sponge reefs, framework-constructing, epifaunal settling). A high degree of endemism, especially among shallow marine bivalves, supports categorization of the Tethys region as a separate faunal realm. Owing to the uncertain palaeogeographic position of many Upper Triassic formations, understanding the relations of the Nayband bivalves to other Tethyan bivalve faunas requires an overall analysis of the faunal connections, which, in turn, allows an independent judgement of different palaeogeographic reconstructions and plate tectonic models. The distribution pattern of bivalve genera and species suggests the existence of a western and eastern Tethys province, with the latter divided into a northern and southern subprovince. The northern subprovince comprises (among others) Iran, Yunnan, Vietnam and Burma, but not the Lhasa-block, which proved to belong to the southern subprovince. The palaeobiogeographic analysis therefore supports the hypothesis that this block was attached to Gondwana until the end of the Triassic and became separated later by opening of a "Ceno-Tethys" ocean. The faunal peculiarity of the western Tethys area is probably owing to its strongly differentiated facies pattern, which hindered faunal exchange, favoured speciation and finally led to the rise of higher taxa. Compared with the endemics of the Eastern Tethys Province, those of the Western Tethys Province were much less affected by the mass extinction event at the end of the Triassic, and many of them were able to disperse into wide areas during the ensuing Liassic transgression. The overall distribution pattern shown by the palaeobiogeographic analysis is not controlled by a single factor, but rather is the result of an interplay of speciation, dispersal and vicariance.

Synapsiden erscheinen erstmals im Fossilbericht im Oberkarbon (späten Pennsylvanium) und dominierten terrestrische Ökosysteme bis zum Ende des Paläozoikums. Diese Arbeit ist die erste detaillierte Betrachtung der frühen Evolution der Synapsiden. Modifizierte Versionen zuvor publizierter Vollständigkeitsmaße werden benutzt, um die Vollständigkeit von Pelycosaurier Fossilien einzuschätzen. Zudem wird eine Reihe unterschiedlicher Methoden genutzt, um die Übereinstimmung von Fossilbericht und Phylogenese zu messen. Die Vollständigkeitsanalyse der Pelycosaurier zeigt eine negative Korrelation zwischen Diversität und dem Maß der Merkmalsvollständigkeit, was darauf hindeutet, dass viele Spezies auf unvollständig erhaltenem Material basieren. Die fehlende Korrelation zwischen dem Maß zur Merkmalsvollständigkeit (basierend auf Abschätzung der Proportion phylogenetisch erfassbarer Merkmale) und der Diversität wird auf die Entdeckungsgeschichte der Gruppe zurück geführt: Die Mehrheit der Pelycosaurier-Arten wurden zwischen den 1930er und 1960er Jahren benannt, als taxonomische Zuordnungen häufig auf Körpergrösse, Fundort und Stratigraphie anstatt auf morphologischen Merkmalen basierten. Welche Schätzungen der Artenzahl über die Erdgeschichte beinflussen, produzieren die unterschiedlichen Methoden zur Diversitätsrekonstruktion sehr ähnliche Ergebnisse. Der initialen Diversifikation der Synapsiden im Oberkarbon und Unterperm (frühes Cisuralium) folgte ein Aussterbeereignis während des Sakmariums. Ein zweites Aussterben ereignete sich an der Grenze vom Kungurium zum Roadium. Die phylogenetisch Topologie-Analyse keine signifikanten Steigerungen der Diversitätsrate der Pelycosaurier relativ zu zeitgleich lebenden Taxa. Eine breiter angelegte Auswertung der Diversitätsentwicklung paläozoischer und triassischer Amnioten liefert ein mögliches Erklärungsmodell; Veränderungen der Diversitätsraten früher Amnioten tendieren dazu, zu Zeiten erhöhter Aussterberaten aufzutreten. / Synapsids first appear in the fossil record during the late Pennsylvanian, and dominated the terrestrial realm until the end of the Palaeozoic. This thesis provides the first detailed examination of the earliest evolution of synapsids. Modifications of previously published metrics are used to assess the completeness of their specimens, and a variety of methods are employed to measure the fit of the fossil record to the phylogeny. The analysis into the completeness of pelycosaurian-grade specimens reveals a negative correlation between diversity and the Skeletal Completeness Metric, assessing the bulk of material preserved, suggesting a tendency to name many species based on poor material. The lack of correlation between the Character Completeness Metric (assessing the proportion of phylogenetic characters that can be scored) and diversity is attributed to the history of discovery in the group: the majority of pelycosaurian-grade species were named between the 1930s and 1960s, when assignments were often based on size, location and stratigraphy rather than morphological characters. The different methods of assessing diversity provide very similar results. The initial diversification of synapsids in the Late Pennsylvanian and early Cisuralian was followed by an extinction event during the Sakmarian. A second extinction event occurred across the Kungurian/Roadian boundary. The tree topology analysis found no significant increases in diversification rate occurring in pelycosaurian-grade taxa relative to their contemporaries. A broader examination of diversification patterns in Palaeozoic and Triassic amniotes reveals a possible explanation; diversification rate shifts within early amniotes tend to occur during periods of elevated extinction.