Institut für Paläontologie
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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 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).
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.
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.
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.
Previous work on Jurassic bivalves from the Iberian Range is reviewed, whereby emphasis is placed on Callovian-Kimmeridgian species. The taxonomy, distribution pattern and ecology of the bivalve fauna occurring in Middle and Upper Jurassic rocks of the Aragonian Branch of the Iberian Range have been analysed. For this purpose 14 sections and 5 additional outcrops, selected according to the abundance of bivalves, were measured in detail and sampled. The rocks studied belong to the Chelva, Yátova, Sot de Chera and Loriguilla formations of Callovian-Kimmeridgian age. The distribution of species of bivalves is given for each section. More than 3000 specimens of bivalves representing 83 species that belong to 46 genera and subgenera of the subclasses Palaeotaxodonta, Pteriomorphia, Isofilibranchia. Palaeoheterodonta, Heterodonta and Anomaldesmata have been used for the taxonomic analysis. One species is new: Plagiostoma fuersichi from the Callovian of the Chelva Fm. The autecology (trophic group and life habit) of each bivalve has been discussed. 49 samples of four sections habe been selected for a quantitative palaeoecological analysis of the bivalve fraction of the benthic fauna. Five bivalve associations and two assemblages are recognised by a Q-mode hierarchical cluster analysis (Ward method). The main environmental factors controlling bivalve associations are thought to be substrate, water energy and distribution of organic matter. The bivalves exhibit a distinct spatial and temporal distribution pattern within the Aragonian Branch. Four of the bivalve associations occur in the Upper Oxfordian (Sot de Chera Fm) and one association in the Lower Callovian (Chelva Fm). In the Sot de Chera and Loriguilla formations, the abundance of bivalves decreases from NW to SE i.e., from relatively close to the shore line towards the distal-most part of the carbonate platform. In the Chelva Fm. bivalves are abundant in the Ariño region, interpreted as a palaeogeographic high. The distribution of bivalves might have been largely controlled by the availability of nutrients.