Cephalopods Present and Past: New Insights and Fresh Perspectives Cephalopods Present and Past: New Insights and Fresh Perspectives

Edited by Neil H. Landman Division of Paleontology (Invertebrates) American Museum of Natural History New York, NY, USA Richard Arnold Davis Department of Biology College of Mount St. Joseph Cincinnati, OH, USA Royal H. Mapes Department of Geological Sciences Ohio University Athens, OH, USA A C.I.P. Catalogue record for this book is available from the Library of Congress.

ISBN 978-1-4020-6461-6 ISBN 978-1-4020-6806-5 (e-book)

Published by Springer, P.O. Box 17, 3300 AA Dordrecht, The Netherlands. www.springer.com

Cover illustration: Reconstruction of the life cycle of Manticoceras, depicting the orientations of the aperture of four representative growth stages. Figure by Christian Klug, Universität Zürich.

Printed on acid-free paper

All Rights Reserved © 2007 Springer No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Preface

Cephalopods are diverse, highly developed molluscs capable of swimming and jet propulsion. These animals are an important component of present-day marine ecosys- tems throughout the world and comprise approximately 900 species. They also have an extraordinary fossil record, extending back to the Cambrian Period, with as many as 10,000 extinct species. Throughout their long history, they have experienced spec- tacular radiations and near-total extinctions. Because of their superb fossil record, they also serve as ideal index fossils to subdivide geologic time. This book touches on many of these themes, and it treats both fossil and present-day cephalopods. The chapters are outgrowths of presentations at the Sixth International Symposium “Cephalopods – Present and Past,” at the University of Arkansas in Fayetteville, September 16–19, 2004. The Symposium was organized principally by Walter L. Manger of the Department of Geology, University of Arkansas. The editors gratefully acknowledge Walter for his terrific job in putting together this symposium and for making it such an intellectual, and social, success. Other publications related to this Symposium include the abstract volume, assembled by W. L. Manger, and two field- trip guidebooks, one written by W. L. Manger, and the other by R. H. Mapes. Because this symposium was held in North America, it honored four cephalopod workers from this continent: William A. Cobban (US Geological Survey, Denver, Colorado), Brian F. Glenister (University of Iowa, Iowa City, Iowa), William M. Furnish (University of Iowa, Iowa City, Iowa), and Gerd E. G. Westermann (McMaster University, Hamilton, Ontario). These four workers are giants in their fields, and through their research on the biology, systematics, and biostratigraphy of fossil cephalopods, they have enormously expanded our understanding of these animals and the history of planet Earth. This volume is dedicated to them – in recognition of their phenomenal accomplishments. This volume contains 20 chapters covering a wide range of topics about both fossil and present-day cephalopods. We have grouped these chapters into three sections, although we recognize that many of the subjects overlap:

● Phylogeny and Systematics (Chapters 1–7) ● Morphology of Soft and Hard Tissues (Chapters 8–14) ● Biogeography, Biostratigraphy, Ecology, and Taphonomy (Chapters 15–20)

v vi Preface

Within each section, ammonoids are treated first, followed by coleoids, in order of geologic time. Every chapter was examined by at least two outside reviewers, and their suggestions and other comments, together with those of the editors, were forwarded to the authors. The reviewers made many helpful suggestions; this resulted in substantially improving the quality of the manuscripts. In addition, authors were encouraged to follow General Recommendation 10 of the International Code of Zoological Nomenclature, which suggests that the author and date of every taxon in a publication be cited at least once in that publication. The editors extend their sincere thanks to the following people who reviewed the manuscripts: Emily G. Allen (Bryn Mawr College, Bryn Mawr, Pennsylvania), Roland Anderson (Seattle Aquarium, Seattle, Washington), R. Thomas Becker (WWU, Geologisch-Paläontologisches Institut, Münster, Germany), Hugo Bucher (Paläontologisches Institut und Museum der Universität Zürich, Zürich, Switzerland), Antonio G. Checa (Universidad de Granada, Granada, Spain), William A. Cobban (US Geological Survey, Denver, Colorado), Régis Chirat (Université Claude Bernard Lyon 1, Villeurbanne Cedex, France), Larisa A. Doguzhaeva (Palaeontological Institute of the Russian Academy of Sciences, Moscow, Russia), Jean-Louis Dommergues (Centre des Sciences de la Terre, Université de Bourgogne, Dijon, France), Desmond T. Donovan (University College London, London, United Kingdom), Dirk Fuchs (Freie Universität Berlin, Berlin, Germany), Roger A. Hewitt (Leigh-on-Sea, Essex, United Kingdom), W. James Kennedy (University Museum, Oxford, United Kingdom), William T. Kirchgasser (SUNY, Potsdam, New York), Christian Klug (Paläontologisches Institut und Museum der Universität Zürich, Zürich, Switzerland), Dieter Korn (Museum für Naturkunde der Humboldt- Universität zu Berlin, Berlin, Germany), Cyprian Kulicki (Polska Akademia Nauk, Warsaw, Poland), Neal L. Larson (Black Hills Museum of Natural History, Hill City, South Dakota), George R. McGhee (Rutgers University, New Brunswick, New Jersey), Lisa K. Meeks (Exxon Mobil Development Company, Houston, Texas), Pascal Neige (Centre des Sciences de la Terre, Université de Bourgogne, Dijon, France), W. Bruce Saunders (Bryn Mawr College, Bryn Mawr, Pennsylvania), Dolf Seilacher (Yale University, New Haven, Connecticut), Kazushige Tanabe (University of Tokyo, Tokyo, Japan), Janet R. Voight (The Field Museum, Chicago, Illinois), Frank Weise (Freie Universität Berlin, Berlin, Germany), Wolfgang Weitschat (Geologische-Paläontologisches Institut und Museum der Universität Hamburg, Hamburg, Germany), Gerd E. G. Westermann (Hamilton, Ontario, Canada), and Margaret M. Yacobucci (Bowling Green State University, Bowling Green, Ohio). The editors also thank Susan M. Klofak, Kathy B. Sarg, Steve Thurston, and Stephanie Crooms (American Museum of Natural History) for help in working with the manuscripts (proofing, mailing, word processing, and scanning images), and Judith Terpos (Springer) for guidance in putting the book together.

Neil H. Landman New York, New York Richard Arnold Davis Cincinnati, Ohio Royal H. Mapes Athens, Ohio Contents

Preface ...... v

Part I • Phylogeny and Systematics

Chapter 1 • Phylogenetic Practices Among Scholars of Fossil Cephalopods, with Special Reference to Cladistics Pascal Neige, Isabelle Rouget, and Sebastien Moyne 1 Introduction ...... 3 2 Sampling Phylogenetic Practices: Review of Paleontological Literature from 1985 to 2003 ...... 4 3 Discussion...... 9 Acknowledgments ...... 12 Appendix ...... 12 References ...... 13

Chapter 2 • Patterns of Embryonic Development in Early to Middle Devonian Ammonoids Susan M. Klofak, Neil H. Landman, and Royal H. Mapes 1 Introduction ...... 15 2 Material and Methods ...... 19 3 Results ...... 20 4 Discussion...... 30 5 Conclusions ...... 35 Acknowledgments ...... 36 Appendix ...... 36 References ...... 53

vii viii Contents

Chapter 3 • Conch Form Analysis, Variability, Morphological Disparity, and Mode of Life of the Frasnian (Late Devonian) Ammonoid Manticoceras from Coumiac (Montagne Noire, France) Dieter Korn and Christian Klug 1 Introduction ...... 57 2 Material ...... 60 3 Conch Parameters ...... 61 4 Conch of Manticoceras ...... 64 5 Comparisons with Other Samples of Manticoceras ...... 69 6 PCA Analysis ...... 74 7 Orientation of the Aperture in Manticoceras ...... 77 8 Life Cycle of Manticoceras ...... 79 9 Toward a Reconstruction of the Manticoceras Animal ...... 81 10 Conclusions ...... 82 Acknowledgments ...... 82 References ...... 82

Chapter 4 • GONIAT – The Current State of the Paleontological Database System on Paleozoic Ammonoids Jürgen Kullmann 1 Introduction ...... 86 2 Scope of the Database System GONIAT ...... 87 3 Data Model ...... 88 4 Applications ...... 90 5 Problems and Limitations ...... 92 6 Future Aspects ...... 92 7 Summary ...... 95 Acknowledgments ...... 95 References ...... 95

Chapter 5 • Ornamental Polymorphism in Placenticeras kaffrarium (Ammonoidea; Upper Cretaceous of India): Evolutionary Implications Tapas K. Gangopadhyay and Subhendu Bardhan 1 Introduction ...... 97 2 Ornamental Polymorphism in Placenticeras kaffrarium ...... 99 3 Evolutionary Mechanisms of Polymorphism in Placenticeras kaffrarium ...... 107 4 Paleobiogeography and Paleoecology of Placenticeras kaffrarium ...... 107 5 Remarks ...... 112 Contents ix

Acknowledgments ...... 117 References ...... 117

Chapter 6 • A Late Carboniferous Coleoid Cephalopod from the Mazon Creek Lagerstätte (USA), with a Radula, Arm Hooks, Mantle Tissues, and Ink Larisa A. Doguzhaeva, Royal H. Mapes, and Harry Mutvei 1 Introduction ...... 121 2 Studied Material, State of Preservation, and Methods ...... 122 3 Comparative Morphology ...... 124 4 Systematic Paleontology ...... 135 5 Morphological Plasticity and Evolutionary Trends in Carboniferous Coleoids ...... 139 Acknowledgments ...... 140 References ...... 140

Chapter 7 • On the Species Status of Spirula spirula (Linné, 1758) (Cephalopoda): A New Approach Based on Divergence of Amino Acid Sequences Between the Canaries and New Caledonia Kerstin Warnke 1 Introduction ...... 144 2 Taxonomy ...... 145 3 DNA Sequence Data ...... 147 4 Material and Methods ...... 148 5 Results ...... 150 6 Discussion...... 150 Acknowledgments ...... 151 References ...... 151

PART II • Morphology of Soft and Hard Tissues

Chapter 8 • Understanding Ammonoid Sutures: New Insight into the Dynamic Evolution of Paleozoic Suture Morphology Emily G. Allen 1 Introduction ...... 159 2 Assessing Suture Morphology ...... 160 3 Material and Methods ...... 167 4 Results ...... 168 5 Discussion...... 172 6 Summary ...... 177 Acknowledgments ...... 177 References ...... 177 x Contents

Chapter 9 • Cameral Membranes in Carboniferous and Permian Goniatites: Description and Relationship to Pseudosutures Kristin Polizzotto, Neil H. Landman, and Royal H. Mapes 1 Introduction ...... 181 2 Material ...... 183 3 Methods ...... 188 4 Observations ...... 189 5 Discussion...... 195 Acknowledgments ...... 202 References ...... 202

Chapter 10 • Soft-tissue Attachment of Middle Triassic Ceratitida from Germany Christian Klug, Michael Montenari, Hartmut Schulz, and Max Urlichs 1 Introduction ...... 205 2 Methods ...... 206 3 Material ...... 207 4 Soft-tissue Attachment Structures ...... 208 5 Conclusions ...... 217 Acknowledgments ...... 218 References ...... 218

Chapter 11 • The Preservation of Body Tissues, Shell, and Mandibles in the Ceratitid Ammonoid Austrotrachyceras (Late Triassic), Austria Larisa A. Doguzhaeva, Royal H. Mapes, Herbert Summesberger, and Harry Mutvei 1 Introduction ...... 221 2 Previous Work on Soft Tissues and Hard Parts ...... 222 3 Locality and Material ...... 223 4 Purpose of this Study ...... 224 5 Ultrastructure and Preservation of the Soft Tissue, Hard Parts, and Skeleton in Austrotrachyceras ...... 224 6 Conclusions ...... 236 References ...... 237

Chapter 12 • Connecting Ring Ultrastructure in the Jurassic Ammonoid Quenstedtoceras with Discussion on Mode of Life of Ammonoids Harry Mutvei and Elena Dunca 1 Introduction ...... 239 2 Material and Methods ...... 240 3 Description ...... 240 4 Discussion...... 245 Contents xi

5 Conclusions ...... 252 Acknowledgments ...... 253 References ...... 253

Chapter 13 • Jaws and Radula of Baculites from the Upper Cretaceous (Campanian) of North America Neil H. Landman, Neal L. Larson, and William A. Cobban 1 Introduction ...... 257 2 Previous Work ...... 258 3 List of Localities ...... 259 4 Geologic Setting ...... 260 5 Conventions ...... 262 6 Description of Jaws ...... 264 7 Discussion...... 288 8 Conclusions ...... 293 Acknowledgments ...... 294 References ...... 294

Chapter 14 • Ultrastructural Analyses on the Conotheca of the Genus Belemnotheutis (Belemnitida: Coleoidea) Dirk Fuchs, Helmut Keupp, Vasilij Mitta, and Theo Engeser 1 Introduction ...... 299 2 Previous Studies ...... 300 3 Material and Methods ...... 301 4 Ultrastructural Observations on the Conotheca of Belemnotheutis ...... 305 5 Discussion...... 309 6 Conclusions ...... 310 References ...... 313

PART III • Biogeography, Biostratigraphy, Ecology, and Taphonomy

Chapter 15 • New Data on the Clymeniid Faunas of the Urals and Kazakhstan Svetlana Nikolaeva 1 Introduction ...... 317 2 Geological Setting ...... 318 3 Facies and Taphonomy ...... 319 4 Ammonoid Assemblages ...... 322 5 Changes in Diversity ...... 330 6 Distribution of Ammonoid Faunas in the Uralian Ocean ...... 331 7 Conclusions ...... 338 Acknowledgments ...... 339 References ...... 339 xii Contents

Chapter 16 • Deformities in the Late Callovian (Late Middle Jurassic) Ammonite Fauna from Saratov, Russia Neal L. Larson 1 Introduction ...... 344 2 Material ...... 346 3 Previous Reports of Epizoa on Ammonites ...... 349 4 Terminology ...... 351 5 Epizoa ...... 353 6 Deformities Caused by Epizoa ...... 355 7 Healed Shell Fractures...... 362 8 Distorted Shapes of Unknown Origin ...... 368 9 Discussion...... 369 10 Conclusions ...... 370 Acknowledgments ...... 371 References ...... 372

Chapter 17 • Biogeography of Kutch Ammonites During the Latest Jurassic (Tithonian) and a Global Paleobiogeographic Overview Subhendu Bardhan, Sabyasachi Shome, and Pinaki Roy

1 Introduction ...... 375 2 Upper Tithonian Assemblages of Different Faunal Provinces ...... 376 3 Affinity of Kutch Assemblage ...... 382 4 Migrational Routes and Paleolatitudinal Disposition of Kutch ...... 385 5 Paleobiogeography of Mass Extinction ...... 386 Acknowledgments ...... 391 References ...... 392

Chapter 18 • Ammonite Touch Marks in Upper Cretaceous (Cenomanian- Santonian) Deposits of the Western Interior Seaway Neil H. Landman and William A. Cobban

1 Introduction ...... 396 2 Localities ...... 397 3 Description of Ammonite Touch Marks ...... 402 4 Discussion ...... 406 5 Conclusions ...... 418 Acknowledgments ...... 420 References ...... 420 Contents xiii

Chapter 19 • Some Data on the Distribution and Biology of the Boreal Clubhook Squid Moroteuthis robusta (Verrill, 1876) (Onychoteuthidae, Teuthida) in the Northwest Paciﬁ c Alexei M. Orlov 1 Introduction ...... 423 2 Material and Methods ...... 424 3 Results ...... 425 4 Discussion...... 427 5 Conclusions ...... 431 References ...... 431

Chapter 20 • Habitat Ecology of Enteroctopus dofleini from Middens and Live Prey Surveys in Prince William Sound, Alaska D. Scheel, A. Lauster, and T. L. S. Vincent

1 Introduction ...... 434 2 Methods ...... 437 3 Results ...... 439 4 Discussion...... 449 Acknowledgments ...... 455 References ...... 455

Index ...... 459 List of Contributors

Emily G. Allen Fachrichtung Paläontologie Department of Geology Freie Universität Berlin Bryn Mawr College, Bryn Mawr Malteserstrasse 74-100 PA 19010, USA D-12249 Berlin, Germany E-mail: [email protected] Dirk Fuchs Subhendu Bardhan Institut für Geologische Wissenschaften Department of Geological Sciences Fachrichtung Paläontologie Jadavpur University Freie Universität Berlin Kolkata 700032, India Malteserstrasse 74-100 E-mail: [email protected] D-12249 Berlin, Germany E-mail: [email protected] William A. Cobban 70 Estes Street Lakewood Tapas K. Gangopadhyay CO 80226, USA Department of Geology Bengal Engineering and Science Larisa A. Doguzhaeva University Shibpur Paleontological Institute of the Russian Howrah 711103, India Academy of Sciences E-mail: [email protected] St. Profsoyuznaya 123 Moscow 117997, Russia Helmut Keupp E-mail: [email protected] Institut für Geologische Wissenschaften Fachrichtung Paläontologie Elena Dunca Freie Universität Berlin Department of Palaeozoology Malteserstrasse 74-100 Swedish Museum of Natural History D-12249 Berlin, Germany S-10405, Stockholm, Sweden E-mail: [email protected] Susan M. Klofak Division of Paleontology (Invertebrates) Theo Engeser American Museum of Natural History Institut für Geologische Wissenschaften 79th Street and Central Park West

xv xvi List of Contributors

New York, NY 10024 A. Lauster and Department of Biology Environmental Sciences, Alaska City College of the City University Paciﬁ c University of New York Anchorage, AK 99508, USA Convent Avenue and 138th Street New York, NY 10031, USA Royal H. Mapes E-mail: [email protected] Department of Geological Sciences 316 Clippinger Laboratories Christian Klug Ohio University, Athens, OH 45701 Paläontologisches Institut USA und Museum, Universität Zürich E-mail: [email protected] Karl Schmid-Strasse 4 CH-8006 Zürich, Switzerland Vasilij Mitta E-mail: [email protected] Russian Academy of Sciences Paleontological Institute Dieter Korn Profsojuznaja 123, Moscow 117997 Museum für Naturkunde der Russia Humboldt-Universität zu Berlin E-mail: [email protected] Invalidenstrasse 43, D-10115 Berlin Germany Michael Montenari E-mail: [email protected] Earth Sciences and Geography Keele University Jürgen Kullmann William Smith Building Institut für Geowissenschaften Newcastle (Staffordshire) Universität Tübingen, Sigwartstrasse 10 ST5 5BG United Kingdom D-72076 Tübingen, Germany E-mail: juergen.kullmann@ Sebastien Moyne uni-tuebingen.de UMR CNRS 5561 Biogéosciences 6 bd Gabriel, F-21000 Dijon, France Neil H. Landman E-mail: [email protected] Division of Paleontology (Invertebrates) American Museum of Natural History Harry Mutvei 79th Street and Central Park West Department of Palaeozoology New York, NY 10024, USA Swedish Museum of Natural History E-mail: [email protected] S-10405, Stockholm, Sweden E-mail: [email protected] Neal Larson Black Hills Institute of Geological Pascal Neige Research UMR CNRS 5561 Biogéosciences PO Box 643 6 bd Gabriel, F-21000 Dijon 117 Main Street France Hill City SD 57745 and E-mail: [email protected] Black Hills Museum of Natural History Hill City SD 57745, USA Svetlana Nikolaeva E-mail: [email protected] Paleontological Institute List of Contributors xvii

Russian Academy of Sciences Hartmut Schulz Profsoyuznaya 123, Moscow 117997 Institut für Geowissenschaften Russia Eberhard-Karls-Universität E-mail: [email protected]; Sigwartstrasse 10, D-72076 Tübingen [email protected] Germany

Alexei M. Orlov Sabyasachi Shome Russian Federal Research Institute Geological Survey of India of Fisheries and Oceanography 15A&B Kyd Street (VNIRO) Kolkata 700016, India 17, V. Krasnoselskaya Moscow 107140, Russia Herbert Summesberger E-mail: [email protected] Naturhistorisches Museum Wien A-1014, Wien, Burgring 7, Austria Kristin Polizzotto E-mail: herbert.summesberger@ Division of Paleontology (Invertebrates) nhm-wein.ac.at American Museum of Natural History 79th Street and Central Park West Max Urlichs New York, NY 10024 and Staatliches Museum für Naturkunde Department of Biological Sciences Rosenstein 1 Kingsborough Community College D-70191 Stuttgart, Germany 2001 Oriental Boulevard E-mail: urlichs.smns@naturkunde Brooklyn, NY 11235, USA museum-bw.de E-mail: kpolizzotto@ kingsborough.edu T. L. S. Vincent Environmental Sciences Isabelle Rouget Alaska Paciﬁ c University UMR CNRS 5143, Case 104 Anchorage, AK 99508 T. 46-56, 5éme E USA 4 place Jussieu, F- 75252 Paris Cedex 05, France Kerstin Warnke E-mail: [email protected] Freie Universität Berlin FR Paläontologie Pinaki Roy Malteserstrase 74-100, Haus D-12249 Department of Geological Sciences Berlin, Germany Jadavpur University E-mail: [email protected] Kolkata 700032, India

David Scheel Environmental Sciences Alaska Paciﬁ c University Anchorage, AK 99508, USA E-mail: dscheel@alaskapaciﬁ c.edu