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