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Aulacostephanus Im Oberjura (Taxionomie, Stratigraphie, Biologie)

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Aulacostephanus Im Oberjura (Taxionomie, Stratigraphie, Biologie) SONDER-ABDRUCK aus PALAEONTOGRAPHICA BEITRÄGE ZUR NATURGESCHICHTE DER VORZEIT Band 119. Abt. A. MITTEILUNGEN AUS DEM PALÄONTOLOGISCHEN INSTITUT DER UNIVERSITÄT ZÜRICH, Nr. 17 DIE AMMONITEN-GATTUNG AULACOSTEPHANUS IM OBERJURA (TAXIONOMIE, STRATIGRAPHIE, BIOLOGIE) VON BERNHARD ZIEGLER (PALÄONTOLOGISCHES INSTITUT DER UNIVERSITÄT ZÜRICH) MIT 22 TAFELN SOWIE 85 ABBILDUNGEN UND 5 TABELLEN IM TEXT UND AUF 12 BEILAGEN STUTTGART E. SCHWEIZERBART'SCHE VERLAGSBUCHHANDLUNG (NÄGELE u. OBERMILLER) 1962 Alle Rechte, auch das der Übersetzung, vorbehalten Printed in Germany Jegliche Vervielfältigung einschließlich photomechanischer Wiedergabe nur mit ausdrücklicher Genehmigung durch den Verlag Satz und Druck: Druckerei E. Schwend KG, Schwäbisch Hall Palaeontographica Abt. A 119 Liefg. 1-4 1—172 Stuttgart, Juni 1962 DIE AMMONITEN-GATTUNG AULACOSTEPHANUS IM OBERJURA (TAXIONOMIE, STRATIGRAPHIE, BIOLOGIE) VON BERNHARD ZIEGLER (PALÄONTOLOGISCHES INSTITUT DER UNIVERSITÄT ZÜRICH) MIT 22 TAFELN SOWIE 85 ABBILDUNGEN UND 5 TABELLEN IM TEXT UND AUF 12 BEILAGEN Inhalt Seite Vorwort 2 Einleitung ... 3 Material . 3 Historisches 6 Grundbegriffe 9 Stratigraphie ... 9 Großbritannien 9 Frankreich 14 Mitteleuropa 20 Osteuropa 24 Mexico 25 Parallelisierung ... 26 Diskussion des Artbegriffes 28 Morphologie und Anatomie 32 Taxionomie 42 Aulacostephanites 44 Aul. eulepidus 44 Aul. peregrinus 48 Aulacostephanoides 50 Aul. desmonotus 50 Aul. variocostatus 52 Aul. linealis . 54 Aul. circumplicatus 57 Aul. oblongatus 60 Aul. mutabilis 62 Aul. sosvaensis 66 Aul. attenuatus 67 Aulacostephanoceras 68 Aul. pusillus . 69 Aul. mammatus 73 Aul. volongensis 74 Aul. undorae .... 75 Aul. rigidus .... 77 Aul. phorcus 78 Habilitationsschrift zur Erlangung der venia legendi der Philosophischen Fakultät II der Universität Zürich. Seite Aul. eudoxus 80 Aul. volgensis 90 Aul. kirghisensis 95 Aul. jasonoides 97 Aul. autissiodorensis 99 Aulacostephanus 104 Aul. pseudomutabilis .... 104 Aul. i/o 113 Aul. pinguis 117 Aul. fallax ... 119 Pararasenia 121 Aul. zacatecanus . 121 Aul. hibridus 122 Aul. crassicosla 124 Aul. quenstedti 125 Aul. semieudoxus 130 Aul. calvescens 132 Xenostephanus . 136 Bestimmungsschlüssol 136 Ergebnisse 138 Chronologie 138 Phylogenie 140 Zoogeographie 146 Biologie 153 Zusammenfassung 156 Literatur 158 Vorwort Die vorliegende Revision der Gattung Aulacostephanus war ursprünglich nur als Bearbeitung der im südwestdeutschen Malm vorkommenden Arten gedacht. Schon bald wurde es indessen notwendig, auch die Formen der angrenzenden Räume mit zu berücksichtigen. Studien in französischen und eng­ lischen Museen sowie eigene Geländearbeiten in Westeuropa förderten dann eine Fülle von Problemen zutage: Wo und wie hat sich die Gattung Aulacostephanus entwickelt? Ist sie überall gleichzeitig ausgestorben? Wie weit erstreckt sich das Verbreitungsgebiet der einzelnen Atilacostep/xanus-Arten? Gibt es innerhalb des Vorkommens einer Art geographische Rassen? Wenn ja, wie ist dies mit einer schwimmenden Bewegungsweise der Ammonoideen zu verein­ baren? Wie haben die Aulacostephanen gelebt? Kennt man Jugendformen? Diese und andere Fragen ließen sich indessen nur angehen, wenn es möglich war, die Gesamtheit der Gattung in ihrem ganzen Verbreitungsgebiet zu untersuchen. In dieser Hinsicht unterscheidet sich die vorliegende Arbeit von anderen Monographien, die in den vergangenen Jahren erschienen sind (z. B. R. CASEY 1957, H. HOLDER 1955, M. K. HOWARTH 1958, G. WESTERMANN 1954, B. ZIEGLER 1958 a). All diese Untersuchungen — mit Ausnahme der Arbeit R. WENGER'S (1957) — gingen von einem beschränkten Arbeitsgebiet aus und berührten das kongenerische Material und die Formen benachbarter oder ent­ legener Gebiete nur am Rande. Phylogenetische Folgerungen litten daher oft daran, daß es nicht möglich war, das Wandern der Tiere eingehend zu berücksichtigen. Nun eignet sich die Gattung Aulacostephanus aus dem unteren Kimeridgien aus mehreren Gründen gut für eine quantitative Studie, in der biologische Fragen besonders berücksichtigt werden. Ihr Ver­ breitungsgebiet ist im wesentlichen auf das außertethydische Europa begrenzt. Es ist indessen in einzelne Teilräume differenziert, die sich durch recht unterschiedliche Faunen auszeichnen. Zwar sind die Formen in weiten Bereichen ziemlich selten — gerade diese Seltenheit gestattet es jedoch, das vorhandene Material wohl nahezu vollständig zu überblicken. Allerdings wurde es hierdurch auch weitgehend un­ möglich, die Variationsbreite einzelner Arten und die Entwicklung der Formen aus ihren Vorläufern statistisch zu erfassen. Weiterhin war es außerordentlich schwierig, Material aus den entlegenen Rand­ gebieten der Verbreitung der Gattung (Ural, Kurdistan, Mexico, Grönland) zu erhalten. Die gewonnenen Ergebnisse werden somit noch in manchen Punkten modifiziert und ergänzt werden müssen. * Ohne weitreichende Hilfe von den verschiedensten Seiten hätte sich die Arbeit nicht durchführen lassen. Mein Dank gilt in erster Linie den Vorständen und Betreuern der Museen und Sammlungen, die mir ihre Bestände zur Durchsicht und zur Bearbeitung zugänglich gemacht haben oder die mir über ihr Material bereitwillig Auskunft er­ teilten. Ganz besonders danke ich all denjenigen Fachgonossen, die meine Untersuchungen durch sachdienliche Aus­ künfte, Hinweise und Ratschläge gefördert haben, allen voran Dr. W. J. ARKELI, (t), Cambridge; Dr. K. W. BARTHEL, München; Prof. Dr. J. BIEGKRT, Zürich; Dr. J. H. CM.LOMON, London; J. M. EDMUNDS, Oxford; R. ENAY, Lyon; Prof. Dr. K. H. EBBEN, Bonn; Dr. C. L. FORBES, Cambridge; Dr. E. GASOIIE, Basel; Dr. O. F. GEYER, Stuttgart; Prof. Dr. W. GHOSS, Tübingen; J. GUYADF.R, Le Havre; Prof. Dr. H. HOLDER, Tübingen; Dr. V/. G. KÜHNE, Berlin; Prof. Dr. J. ORLOV, Moskau; Dr. T. 0nviu, Stockholm; M. RIOULT, Caen; Dr. N. T. SASONOV, Moskau; Prof. Dr. O. H. SCHINDEWOI.F, Tübingen; Prof. Dr. H. SCHMIDT, Göttingen; Dr. H. SCHMIDT-KALK«. Erlangen; H. TINTANT. Dijon; Prof. Dr. R. Tit'jurv, Zürich; Dr. Cn. D. WATF.IISTON, Edinburgh; Dr. A. ZEISS, Erlangen. Den wärmsten Dank schulde ich jedoch Professor Dr. E. KUIIN-SCIINYDEII. Zürich, der mir sämtliche Einrichtungen des Paläontologischen Institutes der Universität Zürich zur Verfügung stellte und den Fortgang der Arbeit durch wert­ volle Hilfe erleichtert hat. Die Geländearbeiten wurden ermöglicht durch Zuwendungen von Seiten der Deutschen Forschungsgemeinschaft, des British Council (durch Vermittlung des Deutschen Akademischen Austauschdienstes) und des Paläontologischen Institutes der Universität Zürich. Auch diesen Institutionen gebührt mein aufrichtiger Dank. Die beigefügten Photographien verdanke ich den Herren J. AICHINGER, Zürich, und W. WETZEL, Tübingen, sowie dem British Museum (Natural History) London. Die Zeichnungen fertigte Frau S. PLETSCIIER-SCIIWIHZER an. Ihnen und allen weiteren ungenannten Helfern danke ich aufs beste. Einleitung Material Ganz besondere Sorgfalt galt der Beschaffung des Untersuchungsmaterials. Da eigene Aufsamm­ lungen im Gelände bei der Seltenheit der Formen keinen repräsentativen Querschnitt durch die Fauna erbracht hatten, war es notwendig, auch Museumsmaterial in die Untersuchungen einzubeziehen. Es war dabei mein Bestreben, die in den zugänglichen Sammlungen liegenden Stücke möglichst quantitativ zu erfassen. Das Material folgender Sammlungen lag der Arbeit zugrunde: Großbritannien: Sedgwick Museum, Cambridge Dorset Museum, Dorchester The Royal Scottish Museum, Edinburgh British Museum (Natural History), London Geological Survey and Museum, London University College, Department of Geology, London Department of Geology, University of Nottingham University Museum, Oxford Department of Geology, University of R e a d i n g Yorkshire Museum, York — 4 — Frankreich: Musee d'A uxerre Musee des Beaux Arts et d'Archeologie, Boulogne-sur-mer Laboratoire de Geologie, Universite de D i j o n Musee d'Histoire Naturelle, D i j o n Laboratoire de Geologie, Universite de G r e n o b 1 e Collection J. GUYADER, L e H a v r e Laboratoire de Geologie, Universite de Lille Laboratoire de Geologie, Universite de Lyon Musee du Chäteau, Montbeliard Ecole Nationale Superieure de Geologie Appliquee et de Prospection Miniere, Universite de Nancy Ecole Nationale Superieure des Mines, Paris Museum National d'Histoire Naturelle, Laboratoire de Geologie, Paris Museum National d'Histoire Naturelle, Laboratoire de Paleontologie, Paris Laboratoire de Geologie, Universite de Paris Laboratoire de Geologie, Universite de P o i t i e r s Museum d'Histoire Naturelle, La Rochelle Laboratoire de Geologie, Universite de Strasbourg Musee de Tonnerre Musee de T r o y e s Deutschland: Biologisches Institut der Philosophisch-Theologischen Hochschule Bamberg Kreisnaturaliensammlung Bayreuth Staatliche Geologische Kommission, Zentraler Geologischer Dienst, Berlin Staatliches Museum für Mineralogie und Geologie, Dresden Naturwissenschaftliches Institut der Philosophisch-Theologischen Hochschule Eichstätt Museum auf der Willibaldsburg, Eichstätt Geologisches Institut der Universität Erlangen Geologisch-Paläontologisches Institut der Universität Frankfurt am Main Naturmuseum „Senckenberg", Frankfurt am Main Geologisch-Paläontologisches Institut der Universität Freiburg im Breisgau Geologisches Landesamt in Baden-Württemberg, Freiburg im Breisgau Sammlung GREINER, G e i s 1 o h e (Mittelfranken) Sammlung ENGEL. Göppingen (Württemberg) Geologisch-Paläontologisches Institut der Universität Göttingen Geologisches Staatsinstitut Hamburg Geologisch-Paläontologisches Institut der Universität Kiel Sammlung R. OECHSLE,
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