The Trace-Fossil Record of Major Evolutionary Events: Volume 1: Precambrian and Paleozoic PDF
Preview The Trace-Fossil Record of Major Evolutionary Events: Volume 1: Precambrian and Paleozoic
Topics in Geobiology 39 M. Gabriela Mángano Luis A. Buatois Editors The Trace-Fossil Record of Major Evolutionary Events Volume 1: Precambrian and Paleozoic Topics in Geobiology Volume 39 The Topics in Geobiology series covers the broad discipline of geobiology that is devoted to documenting life history of the Earth. A critical theme inherent in addressing this issue and one that is at the heart of the series is the interplay between the history of life and the changing environment. The series aims for high quality, scholarly volumes of original research as well as broad reviews. Geobiology remains a vibrant as well as a rapidly advancing and dynamic fi eld. Given this fi eld’s multidiscipline nature, it treats a broad spectrum of geologic, biologic, and geochemical themes all focused on documenting and understanding the fossil record and what it reveals about the evolutionary history of life. The Topics in Geobiology series was initiated to delve into how these numerous facets have infl uenced and controlled life on Earth. Recent volumes have showcased specifi c taxonomic groups, major themes in the discipline, as well as approaches to improving our understanding of how life has evolved. Taxonomic volumes focus on the biology and paleobiology of organisms – their ecology and mode of life – and, in addition, the fossil record – their phylogeny and evolutionary patterns – as well as their distribution in time and space. Theme-based volumes, such as predator-prey relationships, biomineralization, paleobiogeography, and approaches to high-resolution stratigraphy, cover specifi c topics and how important elements are manifested in a wide range of organisms and how those dynamics have changed through the evolutionary history of life. Comments or suggestions for future volumes are welcomed. Series Editors Neil H. Landman American Museum of Natural History, New York, USA Peter J. Harries Tampa, Florida, USA More information about this series at h ttp://www.springer.com/series/6623 M. Gabriela Mángano • Luis A. Buatois Editors The Trace-Fossil Record of Major Evolutionary Events Volume 1: Precambrian and Paleozoic Editors M. Gabriela Mángano Luis A. Buatois Department of Geological Sciences Department of Geological Sciences University of Saskatchewan University of Saskatchewan Saskatoon , Saskatchewan , Canada Saskatoon , Saskatchewan, Canada ISSN 0275-0120 Topics in Geobiology ISBN 978-94-017-9599-9 ISBN 978-94-017-9600-2 (eBook) DOI 10.1007/978-94-017-9600-2 Library of Congress Control Number: 2016951934 © Springer Science+Business Media Dordrecht 2016 T his work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifi cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfi lms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. T he use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specifi c statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. T he publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. Printed on acid-free paper This Springer imprint is published by Springer Nature The registered company is Springer Science+Business Media B.V. Dordrecht To Dolf Seilacher, who, as usual, thought about these issues long before us. Fore word I magine a world much like our own: an oxygen-rich atmosphere, dynamic tectonic activity, and a rich and diverse biota of plants, animals, and teeming associations of microbes. Indeed a world just like ours, with a similarly deep history of life, but with one slight difference: no body fossils. No shark teeth eroding from cliffs, no trilobites, no dinosaur bones cluttering up museums and the dreams of impression- able 5-year-olds (and Hollywood moguls). Worst of all, of course, we would be missing the remarkable schnozzle of O pabinia . Much of the rich morphological detail provided by body fossils would be irretrievably lost, but how much of the his- tory and diversity of life could we recover? Would we be able to identify the explo- sive evolutionary dynamism of the Cambrian diversifi cation? Would the paleontologists of this imaginary world be able to chart changing patterns of animal diversity and morphological disparity? Identify the invasion of land or the great mass extinctions? Chronicle advances in behavioral patterns? Would changing cli- mate patterns be evident, or the rise of great plodding vertebrates? I f we stipulate that tracks, trails, and burrows were preserved in all the detail documented in the chapters of this volume and the richly informative photographs and diagrams, then a remarkable detailed view of animal diversity, morphological disparity, and behavior emerges. For those not intimately acquainted with the exten- sive strides made over the past couple of decades of research in ichnology, this volume illustrates the range of information which can be recovered from the pri- mary record of organismal behavior to analysis of changing patterns of diversity and disparity. A s a graduate student during the early 1980s the focus of ichnology seemed to be on describing new structures and providing them (mystifyingly to those of us who did not study trace fossils) Latin binomials. Yet ichnology was already undergoing a conceptual shift in concert with changes within the broader fi eld of paleobiology. As facies concepts spread through sedimentology, the concept of ichnofacies was adopted. The increased emphasis on taphonomy, preservation, and the quality of the fossil record beginning in the 1980s found a similar expression in ichnology in the recognition that the same animal could produce very different structures depending on the environment and the nature of the sediment, and that tracks can look very vii viii Foreword different depending on what level one examines within their stratigraphy. One result of such studies has been considerable revision in the diversity of ichnotaxa. As described in Chap. 2 of this volume, many fossils that we happily accepted as bur- rows in the mid-1990s have now been recognized as components of a diverse assem- blage of latest Ediacaran tubes, and not trace fossils at all. Restudy of Ediacaran traces has drastically reduced the number of accepted ichnogenera. To a non-ichnologist, what is particularly striking about this book is how faith- fully the general outlines of evolutionary dynamics are visible with trace fossils alone. As illustrated by Chaps. 2 and 3 , the integration of trace fossils has long been standard for studies of the Ediacaran and Cambrian diversifi cation. It is not just that the base of the Cambrian is currently defi ned by the fi rst occurrence of the ichno- taxon T reptichnus pedum, rather it is hard to imagine any survey of the Ediacaran– Cambrian diversifi cation n ot including a discussion of the trace-fossil record. Data from ichnological studies has been essential to revealing patterns of morphological novelty and innovation, which is why several generations of workers, from Dolf Seilacher and Peter Crimes to Mary Droser, Soren Jensen and the editors of this volume, have played critical roles in expanding our integrated understanding of this interval. Indeed, Chap. 3 notes that in important ways the trace-fossil record may provide a more reliable picture of the diversity dynamics during the Fortunian, the fi rst stage of the Cambrian, than do other fossils. T he pattern continues with later chapters. The Ordovician biodiversifi cation event reveals breakthroughs in paleoecology including movement into infaunal habitats with increased tiering, increased bioturbation, and colonization of new environments (Chap. 4 ). Unlike the Cambrian, the Ordovician increase in the diver- sity of ichnotaxa was not accompanied by an increase in ichnodisparity. Indeed a thread throughout the chapters is the frequency of “early burst” patterns of diversi- fi cation. The exploration and exploitation of new habitats is a consistent theme throughout this volume. The invasion of land, discussed in Chap. 5 , reveals a pattern of colonization, a rapid exploration of new behaviors and architectural designs, fol- lowed by variation on the established themes. Other episodes that are addressed are the expansion of terrestrial ecosystems, the Mesozoic marine revolution, and lacus- trine revolutions. Chapter 1 4, on the Cenozoic mammalian radiation, suggests that the tracks of this episode, like the traces of the Cambrian, may provide a richer record of evolutionary change than does skeletal elements, and in addition provides unique information on locomotion, body size, and ecology. I must confess that I had never considered the nature of trace fossils associated with soils, but Chap. 1 5 illu- minates the unexpected complexity in the evolutionary exploitation of paleosols. Chapter 1 6 provides an insightful analysis of patterns of ecospace occupation through the Phanerozoic, advances in ecosystem engineering, and patterns of ichno- diversity and ichnodisparity. I was particularly struck by the proposal that a space of all possible “ichno-structures” that was explored relatively early by marine inverte- brate clades, with similar architectures subsequently discovered independently by different clades. Not surprisingly, this pattern matches studies of ecospace by Bambach and colleagues, and of morphological disparity by many writers. Foreword ix B ut of course the diversity and disparity of these structures varied with events in the history of life, as demonstrated by ichnological studies associated with the end- Permian, end-Triassic, and end-Cretaceous mass extinctions, as discussed in Chaps. 7 , 8 , and 1 2. The early Triassic aftermath of the end-Permian mass extinction reveals as complicated a pattern of ichnofaunal re-emergence as has been found looking just at body fossils. Returning to the counterfactual musings with which I began this foreword, the evidence presented in Chap. 7 makes it clear that we would be able to recover much of the complexity of the biotic recovery from the trace-fossil record alone. (I must confess to a certain pleasure in Chap. 7 as a proposal I made in 1993 which had been discarded by later workers is resurrected therein.) Sadly of course, the trace-fossil record on my mythical, alternate Earth would be no more complete than the record we possess today. Trace fossils might hint at mor- phologies not found among extant animals, but phylogenetic analysis, to take one example, would be greatly hampered by a loss of character information from the early history of many clades: the origins of turtles might forever remain a mystery. There would be many questions accessible with our extant fossil record that would be unanswerable. But of course this is just the strength of the fossil record that we possess. The trace and body fossil records each have their strength, and their weak- nesses. This volume wonderfully elucidates the power of the ichnofossil record, properly interpreted, to reveal much of the ecological and evolutionary dynamics of life on this Earth. As students read this volume (and I hope many do), the chapters here should provide a plethora of questions to pursue. While I am sure that this book will attract many students to ichnology, in some ways the most signifi cant impact of the book would be to convince paleontologists to more fully integrate ichnological data and analyses into projects on evolutionary diversifi cations and radiations, mass extinctions, and paleoecological dynamics. Douglas H. Erwin Department of Paleobiology National Museum of Natural History Washington , DC , USA Contents 1 The Conceptual and Methodological Tools of Ichnology .................... 1 Nicholas J. Minter , Luis A. Buatois , and M. Gabriela Mángano 2 Ediacaran Ecosystems and the Dawn of Animals ................................ 27 Luis A. Buatois and M. Gabriela Mángano 3 The Cambrian Explosion ....................................................................... 73 M. Gabriela Mángano and Luis A. Buatois 4 The Great Ordovician Biodiversifi cation Event ................................... 127 M. Gabriela Mángano , Luis A. Buatois , Mark Wilson , and Mary Droser 5 The Prelude to Continental Invasion .................................................... 157 Nicholas J. Minter , Luis A. Buatois , M. Gabriela Mángano , Robert B. MacNaughton , Neil S. Davies , and Martin R. Gibling 6 The Establishment of Continental Ecosystems .................................... 205 Nicholas J. Minter , Luis A. Buatois , M. Gabriela Mángano , Neil S. Davies , Martin R. Gibling , and Conrad Labandeira 7 The End-Permian Mass Extinction ....................................................... 325 Richard Hofmann Index ................................................................................................................. 351 xi Prol ogue “A beginning has been made and the discipline has been advanced to the point where some of the highlights in the history of life may be examined from a paleoecological perspective” (Valentine 1973, Evolutionary Paleoecology of the Marine Biosphere) In a remarkable book on the meaning of geologic time, Stephen Jay Gould explored the dichotomy between time’s arrow and time’s cycle (Gould 1987). According to this view, time’s arrow encompasses history as an irreversible sequence of unrepeat- able events, whereas time’s cycle refers to a nondirectional time characterized by repetition according to a recurrent pattern. As we have emphasized elsewhere (e.g., Mángano and Buatois 2012, 2015), ichnology can be viewed as an expression of the tension between these two faces of geologic time. The remarkable success of ichnology to solve problems in facies analysis, paleoenvironmental reconstruction, and sequence stratigraphy is clearly an expres- sion of the emphasis on recurrence. In fact, the very same defi nition of archetypal ichnofacies, as trace-fossil suites that record responses of benthic organisms to a given set of environmental conditions and that recur through geologic time, epito- mizes time’s cycle. In this view, regardless of the age of the unit animals should respond to ecological parameters in a similar way, refl ecting their adaptation to the environment they inhabit. This nomothetic program for ichnology, with its power as a predictive conceptual tool, is the reason that sedimentary geologists now typically include trace fossils in their toolkit to scrutinize the stratigraphic record. H owever, this success when dealing with practical issues, often of economic importance, should not prevent us from the realization that ichnology has much to contribute to our understanding of the changing ecology of the past by recovering a time’s arrow perspective. While working on a previous book (Buatois and Mángano 2011), it became increasingly obvious to us that this other face of ichnology has xiii
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