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Topics in Geobiology 39

M. Gabriela Mángano
Luis A. Buatois Editors

The Trace-Fossil
Record of Major
Evolutionary Events
Volume 1: Precambrian and Paleozoic
This volume addresses major evolutionary
changes that took place during the
Ediacaran and the Paleozoic.
These include discussions on the nature of
Ediacaran ecosystems, as well as the
ichnologic signature of evolutionary
radiations, such as the Cambrian explosion
and the Great Ordovician biodiversification
event, the invasion of the land, and the end-
Permian mass extinction.
This volume set provides innovative reviews
of the major evolutionary events in the
history of life from an ichnologic
perspective.
Because the long temporal range of trace
fossils has been commonly emphasized,
biogenic structures have been traditionally
overlooked in macroevolution.
However, comparisons of ichnofaunas
through geologic time do reveal the
changing ecology of organism-substrate
interactions.
The use of trace fossils in evolutionary
paleoecology represents a new trend that is
opening a window for our understanding of
major evolutionary radiations and mass
extinctions.
Trace fossils provide crucial evidence for
the recognition of spatial and temporal
patterns and processes associated with
paleoecologic breakthroughs.
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 field.
Given this field’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 influenced and controlled life on Earth.
Recent volumes have showcased specific 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 specific
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 http://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

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of
the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,
broadcasting, reproduction on microfilms 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.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The 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
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or omissions that may have been made.

Printed on acid-free paper

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The registered company is Springer Science+Business Media B.V. Dordrecht
To Dolf Seilacher, who, as usual, thought
about these issues long before us.
Foreword

Imagine 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 Opabinia. 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 diversification? 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?
If 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.
As 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 field 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 diversification. It is not just that
the base of the Cambrian is currently defined by the first occurrence of the ichno-
taxon Treptichnus pedum, rather it is hard to imagine any survey of the Ediacaran–
Cambrian diversification not 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
first stage of the Cambrian, than do other fossils.
The pattern continues with later chapters. The Ordovician biodiversification
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-
fication. 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 14, 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. 15 illu-
minates the unexpected complexity in the evolutionary exploitation of paleosols.
Chapter 16 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.
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Foreword ix

But 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 12. 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 significant impact of
the book would be to convince paleontologists to more fully integrate ichnological
data and analyses into projects on evolutionary diversifications 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 Biodiversification 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
Prologue

“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 definition 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, reflecting 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.
However, 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
xiv Prologue

received far less attention. However, even a cursory review of the recent paleobio-
logic literature shows that there is an increased awareness of the importance of
bioturbation and bioerosion as driving forces in the history of life, playing major
roles as a connecting players between biotic and abiotic factors involved in complex
feedback loops that result in evolutionary change (e.g., Erwin and Tweedt 2012).
In its essence, this book reflects the growing interest within the ichnologic com-
munity in expanding the potential of ichnology to contribute to evolutionary paleo-
ecology. The authors of the different chapters are eclectic in their perspectives and
methodologies. However, there are some common themes and conceptual tools,
such as ichnodiversity and ichnodisparity, ecospace utilization, environmental
expansion, innovation associated to major radiation events, and the re-organizations
of ecological units, in particular communities and ecosystems, after mass extinc-
tions. What emerges is a wealth of information and the idea that the history of life
can be accurately reconstructed looking through the ichnologic glass.
There are two strategies to bring back the time’s arrow perspective of ichnology.
One would be to assess the trace-fossil record of the colonization of specific depo-
sitional settings, being the main focus on the reconstruction of major trends in
global ichnodiversity. In fact, this approach has a relatively long history, starting
with the colonization of the deep sea through geologic time due to pioneer papers
by Seilacher (1974, 1977) and Crimes (1974), continuing with more recent studies
on the same topic (Orr 2001; Uchman 2003, 2004), as well as on the colonization of
continental environments (Buatois and Mángano 1993; Buatois et al. 1998), estuar-
ies (Buatois et al. 2005), and tidal flats (Mángano and Buatois 2015). Another
approach would be to evaluate how the interactions between animals and substrates
were shaped by major macroevolutionary events, such as the Cambrian explosion
and the Permian mass extinction. These changes are analyzed from an ecological
perspective. For this book we have decided to follow this later approach.
The first volume opens with Chap. 1, where Minter et al. briefly revise the con-
ceptual and methodological tools of ichnology, with the aim of providing the reader
with the basic information necessary to explore the rest of the book. The approach
is eclectic with brief summaries of the basic principles and concepts in the field, as
well as revisions of the ichnofacies model and the ichnofabric approach. The chap-
ter also introduces a number of concepts and methods that are used in subsequent
chapters, including the notion of ichnodiversity and ichnodisparity as a novel
approach to unlock the potential of ichnofaunas to provide insights into mode of
life, ecospace colonization, and ecosystem engineering.
Chapter 2, by Buatois and Mángano, starts our journey through time by evaluat-
ing the highly controversial trace-fossil record of the Ediacaran. The chapter can be
understood as an attempt to revisit from an ichnologic perspective the so-called
Darwin’s dilemma on the supposed absence of evidence of life in the Precambrian
by critically searching for the roots of animal life in the Ediacaran. In addition, ich-
nologic information is used to reveal the nature of Ediacaran ecosystems and the
complexities involved in the transition to the Phanerozoic world.
In Chap. 3, Mángano and Buatois look at the other side of the great divide, by
exploring the trace-fossil record of the Cambrian explosion. The chapter empha-
Prologue xv

sizes the importance of trace-fossil data to calibrate the Cambrian diversification

event. Also, the profound changes that took place at ecosystem scale are analyzed
in detail, emphasizing the role of bioturbation as an agent of evolutionary change.
The chapter closes with an evaluation of the paleoenvironmental breadth of the
Cambrian explosion, the significance of the ichnofaunas associated with Burgess
shale-type deposits, and the dual nature of the Fortunian from an evolutionary
standpoint.
Chapter 4, by Mángano et al., examines the following evolutionary radiation, the
Great Ordovician Biodiversification Event. This chapter analyzes ichnodiversity
and ichnodisparity trajectories through the Ordovician, comparing patterns in
different depositional environments and paleocontinents. The contrasting nature of
diversification in soft-bottom and hardground communities is emphasized by show-
ing that innovation in macrobioerosion was significantly delayed when compared
with bioturbation.
Minter et al. explore the initial steps of life on land in Chap. 5 focusing on the
Ediacaran–Ordovician timespan. The chapter traces the early stages of animal
expansion from fully marine settings into marginal-marine coastal environments
and ultimately truly continental settings by the Late Ordovician. This prelude to
terrestrialization involves the gradual colonization of new environments followed
by rapid filling of ecospace, the establishment of new architectural designs, and
diversification within the framework of these new behavioral programs. Links
between these evolutionary innovations, and the incipient establishment of a land
flora and changes in fluvial styles are discussed.
In Chap. 6, Minter et al. continue with the analysis of terrestrialization by exam-
ining the trace-fossil record of marginal-marine and continental environments dur-
ing the remainder of the Paleozoic. Following the protracted prelude reviewed in the
previous chapter, the Silurian to the Permian was characterized by an explosion of
diversity and expansion into newly colonized environments, such as river channels,
overbanks, deserts, and lakes, coupled with increasing exploitation of the infaunal
ecospace. The chapter underscores how colonization of continental settings by ben-
thic organisms parallels changes in vegetation and fluvial styles.
Hofmann closes the first volume with Chap. 7, addressing the trace-fossil record
of the end-Permian mass extinction. By carefully evaluating ichnologic data, he
provides insights into the extinction event and the subsequent recovery. The impact
of burrowing organisms on geochemical conditions of the marine sediment is
emphasized, providing a critical evaluation of previous ideas regarding the role of
oxygen-depleted conditions as a cause of delayed recovery. It is argued that biotur-
bation was strongly reduced as a result of the end-Permian mass extinction, leading
to the collapse of the mixed layer. The large-scale consequences of such collapse are
evaluated in detail.
Volume 2 marks the passage to the post-Paleozoic world, opening with Chap. 8,
where Barras and Twitchett analyze the end-Triassic mass extinction. The authors
revised the ichnologic record of both terrestrial and marine environments through
the Triassic-Jurassic transition, outlining how the benthic faunas responded to the
xvi Prologue

mass extinction. Their review suggests that climatic and environmental change
were main factors controlling benthic communities on land and in the sea.
In Chap. 9, Buatois et al. take the reader to the new world arisen from the
Mesozoic Marine Revolution. The main group of bioturbators and bioeroders are
reviewed, and the trace-fossil record of Mesozoic and Cenozoic marine environ-
ments is revised to track evolutionary innovations, discussing the timing of Mesozoic
Marine Revolution. The environmental breadth of this major event is evaluated by
summarizing evolutionary innovations not only in shallow-marine environments but
also in marginal-marine and deep-marine settings. It is argued that infaunalization
predated an increase of predation pressures by approximately 50 Myr, pointing
towards a complex set of feedback mechanisms between the two.
In Chap. 10, Bernardi et al. explore the vertebrate radiation during the Mesozoic.
By revising in detail available ichnologic data, they show how the trackway record
yields insights into several issues, such as locomotor mechanics and behavior, there-
fore providing information that typically is not revealed by the body-fossil record.
In turn, biases inherent to the trace-fossil record are discussed, arguing that the
integration between the trackway and the skeletal record is essential to provide a
more holistic picture of the evolutionary changes underwent by terrestrial verte-
brates during the Mesozoic.
Chapter 11, by Buatois et al., discusses another series of evolutionary break-
throughs that took place in continental settings: those resulting from the Mesozoic
lacustrine revolution. Lakes are evaluated from an ichnologic perspective, contrast-
ing lacustrine ecosystems prior to this evolutionary event with those that arose after
the Mesozoic lacustrine revolution. Aspects discussed in this chapter include ichno-
diversity changes through time, the establishment of modern lacustrine food webs,
and the role of behavioral convergence on both sides of the salinity divide.
In Chap. 12, Labandeira et al. shift our attention to another extinction event: the
end-Cretaceous mass extinction. These authors evaluate both the record of plant–
arthropod interactions on land and that of invertebrate trace fossils in the sea, there-
fore providing a comprehensive picture of the extinction event and its aftermath. A
number of methodological, empirical, and theoretical advances resulting from the
use of ichnologic data are outlined, including application of innovative methods to
quantify ichnologic information, access to novel ecologic data, and evaluation of
catastrophic vs. gradualistic scenarios based on the analysis of bioturbated sedi-
ment, among many others.
Chapter 13, by Genise et al., represents a general departure to the overall struc-
ture of the book in that it does not address a specific evolutionary event, but instead
traces a series of dramatic changes in paleosols covering most of, if not all, the
Phanerozoic. In doing so, they provide an in-depth characterization of four revolu-
tions in paleosol ichnofaunas. In addition, this chapter provides an interesting twist
to the concept of ichnofacies by showing that their establishment in terrestrial set-
tings is fully linked to a series of evolutionary innovations by soil-burrowing
organisms.
In Chap. 14, Krapovickas and Vizcaino explore the evolution of mammals based
on their footprint record. Although this chapter summarizes the Mesozoic and
Cenozoic record of tracks attributed to mammals worldwide, emphasis is on changes
Prologue xvii

in South American mammals during the Cenozoic. The authors show that Eocene-
Oligocene trackway assemblages mostly consist of forms of uncertain affinity,
whereas later assemblages consist of trackways of both native South American and
North American mammals that arrived during the Great American Biotic
Interchange. This chapter emphasizes the strong provincialism of mammal faunas,
representing a starting point towards a more global examination of their ichnologic
record.
Chapter 15, by Lockley et al., deals with the trackway record of hominin evolu-
tion. Starting with the famous 3.6 Ma tracks from Laetoli, the reader is taken through
a journey that explores the expansion of our ancestors from Africa to Eurasia, the
New World, and Australasia. References to footprints on the Moon and tracks of
robotic vehicles on Mars add another dimension to the trip. Along the way, the
authors discuss various controversies, including the alternative interpretations of the
Laetoli trackways and the differences between early hominin footprints and those
from modern humans.
The book closes with Chap. 16, where Buatois and Mángano take the challenge of
trying to summarize possible recurrent trends revealed by the trace-fossil record that
may provide insights into the underlying dynamics of animal–substrate interactions
through geologic time. In particular, the chapter discusses organism–substrate inter-
actions during evolutionary radiations, benthic fauna response to mass extinctions,
patterns of ecospace colonization, and environmental shifts through time. In contrast
to the other chapters, which by reviewing individual evolutionary events are anchored
in an idiographic approach, this last chapter takes on a more nomothetic perspective
by trying to find recurrent patterns and processes in evolutionary paleoecology.
In the process of planning this book, it became clear that, although there is an
explicit attempt to bring a picture of the state of the art in the contributions of ich-
nology to the understanding of evolution at the macroevolutionary scale, our book
does not offer a comprehensive treatment or a closure on the topic. In fact, during
this process, we have identified numerous areas of interest where there is limited
ichnologic information to the point that it was not possible to include chapters on
these issues. For example, the end-Ordovician and Late Devonian mass extinctions
have received significantly less treatment than any of the other so-called Big Five.
Also, some of the chapters provide fresh approaches to our exploration of the colo-
nization of infaunal ecospace and the impact of ecosystem engineers by using ich-
nologic data in an innovative fashion. This may serve as inspiration for applying
new numerical and conceptual tools to a wider spectrum of paleobiologic issues.
This book is a tangible testimony that ichnology counts with a solid theoretical
framework and sufficient methodological tools to tackle evolutionary questions and
offer crucial pieces in the reconstruction of the puzzle of the history of life. Our
ambition with this book echoes the Gouldian aim, paraphrased in Chapter 10, of
sitting Ichnology at the High Table of Macroevolution and Paleobiology.
We would like to thank the reviewers of the many chapters, who did a great job
of providing valuable feedback. These are: Andrea Baucon, Zain Belaústegui,
Angela Buscalioni, Karen Chin, Matthew Clapham, Darin Croft, Phillip Currie, Bill
DiMichele, Tony Ekdale, Russell Garwood, Sören Jensen, Dirk Knaust, Conrad
Labandeira, Spencer Lucas, Ken McNamara, Christian Meyer, Nic Minter, Guy
xviii Prologue

Narbonne, Eduardo Olivero, Paul Olsen, Roy Plotnick, Gustavo Politis, Charles
Savrda, Thomas Servais, Alfred Uchman, Jean Vannier, Sally Walker, Mark Wilson,
and Anna Żylińska. Also we thank Doug Erwin for writing the Foreword.
Finally, as with almost all things ichnologic, Dolf Seilacher has been there
before. He should be recognized as the scientist who started to think along this line
of evidence, putting forward the idea that trace fossils represent the “other” fossil
record, underexplored and immensely valuable. Not necessarily an alternative
archive for the history of life, but more of an essential companion that should be
integrated to the other lines of evidence to decipher the complex evolutionary path-
ways in the history of life. Needless to say, Dolf’s influence has been huge in the
field of ichnology and the same can be said of his influence in our personal careers.
It all started long time ago when we were Geology students and he planted some
“seed ideas” during a Trace Fossil course that he taught at the Argentinean
Paleontological Association (APA). Sadly, he passed away during the completion of
this book. We would like to dedicate this book to his memory.
M. Gabriela Mángano
Luis A. Buatois

References

Buatois LA, Mángano MG (1993) Ecospace utilization, paleoenvironmental trends and the evolu-
tion of early nonmarine biotas. Geology 21: 595–598
Buatois LA, Mángano MG (2011) Ichnology: organism-substrate interactions in space and time.
Cambridge: Cambridge University Press
Buatois LA et al. (1998) The ichnologic record of the invertebrate invasion of nonmarine ecosys-
tems: evolutionary trends in ecospace utilization, environmental expansion, and behavioral
complexity. Palaios 13: 217–240
Buatois LA et al. (2005) Colonization of brackish-water systems through time: evidence from the
trace-fossil record. Palaios 20: 321–347
Erwin DH, Tweedt SM (2012) Ecological drivers of the Ediacaran-Cambrian diversification of
Metazoa. Evol Ecol 26: 417–433
Gould SJ (1987) Time’s arrows time’s cycle. Cambridge MA: Harvard University Press. pp 222
Mángano MG, Buatois LA (2012) A multifaceted approach to ichnology. Ichnos 19: 121-126
Mángano MG, Buatois LA (2015) The trace-fossil record of tidal flats through the Phanerozoic:
evolutionary innovations and faunal turnover. In: McIlroy D (ed) Ichnology: Papers from
ICHNIA III. Geol Assoc Canada Misc Pub 9: 157–177
Orr PJ (2001) Colonization of the deep-marine environment during the early Phanerozoic: the
ichnofaunal record. Geol J 36: 265−278
Seilacher A (1974) Flysch trace fossils: evolution of behavioural diversity in the deep-sea. Neues
Jahrb Geol Palaont Monat 1974: 233–245
Seilacher A (1977) Evolution of trace fossil communities. In: Hallam A (ed) Patterns of evolution
as illustrated by the fossil record. Dev Paleontol Stratigraphy, Vol. 5. Amsterdam, Elsevier
Uchman A (2003) Trends in diversity, frequency and complexity of graphoglyptid trace fossils:
evolutionary and palaeoenvironmental aspects. Palaeogeogr Palaeoclimatol Palaeoecol 192:
123–142
Uchman A (2004) Phanerozoic history of deep-sea trace fossils. In: McIlroy D (ed) The applica-
tion of ichnology to palaeoenvironmental and stratigraphic analysis. Geol Soc Spec Pub 228
Visit https://textbookfull.com
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Chapter 1
The Conceptual and Methodological Tools
of Ichnology

Nicholas J. Minter, Luis A. Buatois, and M. Gabriela Mángano

1.1 Introduction

Ichnology is the multidisciplinary science that focuses on the study of traces pro-
duced by organisms (both animals and plants) on or within a substrate, including all
issues related to bioturbation, bioerosion, and biodeposition (Pemberton et al. 1992;
Bromley 1990, 1996; Buatois and Mángano 2011). Ichnology has undergone an
explosive development since the 1960s, particularly after the publication in English
of the seminal papers by Dolf Seilacher (e.g., Seilacher 1964, 1967). At that time
somewhat off the radar of the English-speaking world were his earlier studies (e.g.,
Seilacher 1953a, b, 1955, 1958, 1963) and all the impressive corpus of work done
by the German school, including the development of the so-called “actuopalaeon-
tology” (e.g., Richter 1929; Abel 1935; Schäfer 1956, 1962, 1972), which delin-
eates a tradition to which Seilacher’s research program undoubtedly belongs
(Mángano and Buatois 2012).
Before exploring the potential of trace fossils in macroevolution it is necessary
to summarize the conceptual framework of ichnology, as well as to outline the vari-
ous approaches adopted by its practitioners. The aim of this introductory chapter is
to provide such an overview. Extensive coverage of these topics can be found in
several textbooks (e.g., Ekdale et al. 1984; Bromley 1990, 1996; Buatois and
Mángano 2011), whereas in-depth functional analysis of the most important groups

N.J. Minter (*)

School of Earth and Environmental Sciences, University of Portsmouth,
Burnaby Building, Burnaby Road, Portsmouth, Hampshire PO1 3QL, UK
e-mail: [email protected]
L.A. Buatois • M.G. Mángano
Department of Geological Sciences, University of Saskatchewan,
114 Science Place, Saskatoon, SK S7N 5E2, Canada

© Springer Science+Business Media Dordrecht 2016 1

M.G. Mángano, L.A. Buatois (eds.), The Trace-Fossil Record of Major
Evolutionary Events, Topics in Geobiology 39,
DOI 10.1007/978-94-017-9600-2_1
2 N.J. Minter et al.

of trace fossils was provided by Seilacher (2007). Because of this, our review in this
introductory chapter is succinct, summarizing information in tables, rather than in
an extensive text.

1.2 Basic Concepts

As with every science, ichnology is plagued by terminology that can be somewhat

intimidating. Some of these terms (e.g., track, trail) refer to observable entities and
are easier to grasp, whereas others deal with concepts of more abstract nature (e.g.,
ichnofacies, ichnoguild). The notion of “trace,” an individual and distinctive struc-
ture of biogenic origin showing some sort of link to the morphology of its producer
(Frey 1973), lies at the center of ichnology. The classification schemes for biogenic
structures have been reviewed elsewhere (Frey and Wheatcroft 1989; Pemberton
et al. 1992; Buatois and Mángano 2011), and there is no need for repetition here, so
only some brief definitions are provided.
The broad category of biogenic structures includes a wide variety of structures of
organic origin, some of which fall within the realm of ichnology, whereas others
(e.g., stromatolites, a typical biostratification structure) are commonly regarded as
outside the field. Of the structures included within the field of ichnology, bioturba-
tion and bioerosion structures are the most important ones. Bioturbation structures
are biogenic sedimentary structures reflecting the disruption of stratification fea-
tures or sedimentary fabrics by the activity of an organism, whereas bioerosion
structures comprise biogenic structures produced mechanically or biochemically by
an organism into rigid substrates, such as hardgrounds, clasts, bones, invertebrate
skeletons, wood, or rocks (Frey and Wheatcroft 1989). A third group of structures
embraced by ichnology are biodeposition structures, which reflect production or
concentration of sediment by the activities of an organism and include coprolites,
fecal pellets, pseudofeces, regurgitation pellets, and fecal castings (Frey and
Pemberton 1984; Frey and Wheatcroft 1989).
The vast majority of the biogenic structures discussed in this book are bioturba-
tion structures. These include tracks, trails and burrows. Tracks are imprints left by
an individual locomotion appendage (the related term “trackway” refers to a
repeated pattern of tracks). Trails are continuous impressions or grooves produced
during locomotion. Burrows are permanent to semipermanent structures excavated
within the sediment. Bioerosion structures are also addressed in several chapters.
These include macroborings (down to the millimeter scale) and microborings
(smaller than a millimeter) (Bromley 1994). Bioerosion structures encompass a
wide variety of morphologic types, including borings, embedment structures, rasps
and scrapes, surface etching scars, durophagous damage, drill holes, and bite marks
(Bromley 1992, 1994). This classification scheme is not without problems, as
reflected by the uncertain placement within this framework of other structures that
are prominent in some chapters of the book, such as biogenic structures preserved
in wood, leaves, and seeds (e.g., Labandeira 2002).
1 The Conceptual and Methodological Tools of Ichnology 3

1.3 The Peculiarities of Trace Fossils

Trace fossils differ in many ways from body fossils, regarding both their mode of
formation and their taphonomic histories (Seilacher 1964; Frey 1975; Ekdale et al.
1984; Frey and Pemberton 1985; Pemberton et al. 1990, 2001; Bromley 1990, 1996;
Buatois and Mángano 2011). As a result, ichnologists attempted to outline these
peculiarities in terms of a list of characteristics (Seilacher 1964; Frey 1975) or

Table 1.1 The ten main characteristics of trace fossils (based on Buatois and Mángano 2011)
Characteristic Comment
Trace fossils Analysis of the morphology and architecture of trace fossils provides
represent evidence evidence on the anatomy and ethology of their producers. Behavior
of behavior may range from simple (e.g., trace of a worm-like animal moving
through the substrate) to complex (e.g., the work of social insects).
Releasing the behavioral signal unlocked in a biogenic sedimentary
structure is central to any ichnologic analysis
The same organism A single organism may produce several ichnotaxa by changing its
may produce more behavior. Intergradational forms that show the transition of one
than one ichnotaxon ichnotaxon into another are referred to as “compound ichnotaxa.” In
addition, the same organism may produce different traces during its
different ontogenetic stages
The same ichnotaxon The same ichnotaxon can be produced by many different organisms, as
may be produced by a result of behavioral convergence. Establishing a one-to-one
more than one relationship between producer and biogenic sedimentary structure is
organism not possible in the vast majority of cases. In general, the simpler a trace
fossil is, the weaker the link between the biogenic sedimentary
structure and its producer. In any case, morphologic features commonly
may supply enough information on burrowing technique and anatomy,
so a link with a certain group of organisms can be established
Multiple architects A single biogenic sedimentary structure may reflect the activities of
may produce a single more than one producer. This may result from producers operating
structure more or less at the same time or in successive bioturbation events. The
first case is difficult to recognize in the fossil record and commonly
results from the establishment of symbiotic or commensalist
relationships. The second case is common in the fossil record and is
represented by abandoned biogenic sedimentary structures that are
reoccupied by a different animal. The term “composite ichnotaxa” has
been proposed to name forms that superficially look like a single
burrow system, but actually result from the interpenetration of
individual discrete ichnofossils
Producers are The body-fossil record for the most part reflects those groups that have
commonly soft- developed hard parts, whereas trace fossils commonly record the
bodied animals that activities of soft-bodied animals. This is because the trace-fossil record
are rarely preserved is biased towards the activities of infaunal organisms, and the presence
of skeletons is commonly detrimental for infaunal life. Exceptions to
this principle are represented by the preservation of the producer
directly associated with the biogenic sedimentary structure
(continued)
4 N.J. Minter et al.

Table 1.1 (continued)

Characteristic Comment
Trace fossils are Conditions favoring the preservation of trace fossils are remarkably
commonly preserved different from those of body fossils. Diagenetic processes conducive to
in rock units that are destruction of body fossils may favor trace-fossil preservation by
otherwise promoting mineral precipitation around burrow walls. Biogenic
unfossiliferous sedimentary structures commonly represent the only biotic evidence in
many rock units
The same biogenic Biogenic sedimentary structures reflecting identical burrowing
sedimentary structure technique by the producer may result in apparently disparate trace
may be differentially fossils if produced under contrasting substrate conditions. This is a
preserved in various serious complication for ichnotaxonomy, because underappreciating
substrates this issue may lead to oversplitting
Trace fossils The vast majority of ichnotaxa display long stratigraphic ranges,
commonly have long commonly spanning most if not all the Paleozoic and a small number
stratigraphic ranges originating in the Ediacaran. This fact reveals behavioral convergence
rather that a common producer through geologic time. Certain
behaviors were established relatively early in the history of metazoan
life and have remained essentially unchanged. However, there are many
departures to this principle, as it will become evident to the reader of
this book
Trace fossils In fairness, this characteristic applies more accurately to trace-fossil
commonly have associations rather than to individual ichnotaxa. In any case, it reveals
narrow that biogenic sedimentary structures are strongly controlled by
environmental environmental factors and, therefore, the large majority of them occur
ranges preferentially in certain depositional environments
Trace fossils are Trace fossils represent the in situ record of biogenic activity, and
rarely transported therefore typically they have not suffered secondary displacement.
Exceptions include borings transported together with the host medium
(e.g., bioeroded logs, shells, clasts) and burrows with strongly
reinforced walls that make them resistant to erosion and reworking

ichnologic principles (Ekdale et al. 1984; Bromley 1990, 1996), which are here
listed in Table 1.1, based on a recent synthesis by Buatois and Mángano (2011).
Preservational issues need to be assessed in any study involving trace fossils. In
particular, toponomy, which comprises the description and classification of bio-
genic structures with respect to their mode of preservation and occurrence, has
received most attention (Frey and Pemberton 1985; Rindsberg 2012). Mode of
occurrence is usually defined according to the position of the structure on or within
the stratum, or relative to the casting medium. Mechanical processes involved in the
fabrication of the structure (stratinomy) and its alteration (taphonomy) are included
within toponomy as well. A number of classification systems have been introduced
to address stratinomy. Those of Seilacher (1964) and Martinsson (1970) are the
ones that have met with most acceptance (Fig. 1.1).
1 The Conceptual and Methodological Tools of Ichnology 5

Fig 1.1 Block diagram illustrating the terms used in the stratinomic classifications of Seilacher
and Martinsson. In Seilacher’s scheme, full-relief structures are preserved within the stratum,
whereas semirelief structures are preserved at lithologic interfaces. Semirelief structures are in turn
subdivided into epirelief (preserved at the top) or hyporelief (preserved at the base) of the sand-
stone bed. The terms “concave” (negative) and “convex” (positive) are employed to provide a
picture of the trace-fossil relief. In Martinsson’s classification, epichnial preservation refers to
structures preserved at the upper surface of the casting strata, whereas hypichnial preservation
includes those at the lower surface of the casting strata. The terms grooves and ridges are used to
denote negative and positive reliefs, respectively. Endichnial preservation includes structures
preserved within the casting medium, whereas exichnial preservation comprises those preserved
outside the casting medium (after Buatois and Mángano 2011)

Because trace fossils are primarily evidence of animal behavior, evaluation of

the ethologic significance of trace fossils is central to any application of ichnology.
Therefore, the ethologic classification of trace fossils proposed originally by
Seilacher (1953a) is now considered an essential component of the conceptual
framework of ichnology. The original system consists of five categories: resting
traces (Cubichnia), locomotion traces (Repichnia), grazing traces (Pascichnia),
feeding traces (Fodinichnia), and dwelling traces (Domichnia), but many other cat-
egories have been added subsequently (Frey 1973; Ekdale et al. 1984; Ekdale 1985;
Bromley 1990; Genise and Bown 1994; de Gibert et al. 2004; Tapanila 2005; Genise
et al. 2007; Seilacher 2007; Lehane and Ekdale 2013; Vallon et al. 2015a, b). The
validity and utility of the more recent categories continue to be evaluated (Vallon
et al. 2015a), and only the more widely accepted ones are listed and illustrated in
this chapter (Table 1.2 and Fig. 1.2). Although it has been argued that many com-
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 Wherefore to stroy vs hee did all hee might.[971]

7.

His cursed sonne ensued his cruell path,

And kept my giltlesse cosin strayt in duraunce:
For whome my father hard entreated hath:
But liuing hopelesse of his line’s assuraunce,
Hee thought it best by pollitike procuraunce,
To priue[972] the king, and so restore his frend:
Which brought him selfe to an infamous end.[973]

8.

For whan king Henry,[974] of that name the fifte,

Had tane my father in his[975] conspiracy,[976]
Hee, from sir Edmund all the blame to shifte,
Was fayne to say the[977] French kinge’s ally[978]
Had hyred him this trayterous act to trye,
For which condemned shortly hee was slaine,
In helping right this was my father’s gaine.[979]

9.

Thus whan the linage of the Mortimers

Was made away by his[980] vsurping line,
Some hang’d, some slaine, some pined prisoners,
Because the crowne by right of lawe was mine,
They gan as fast against mee to repine,
In feare alwayes, least I should stur them to[981] strife,
For guylty hearts haue neuer quiet life.[982]

10.

Yet at the last, in Henrye’s dayes the sixt,

I was restored to my father’s landes,
Made duke of Yorke: where through my minde I fixt
To get the crowne and kingdome in my handes,
For ayde wherein I knit assured handes
With Neuil’s stocke, whose daughter was my make,
Who for no woe would euer mee forsake.[983]

11.

O Lord, what hap had I through mariage,

Fower goodly boyes in youth my wife shee bore
Right valiaunt[984] men and prudent for their age,
Such brethren[985] shee had and nephues [still] in store
As none had erst, nor any shall haue more:[986]
The earle of Salisbury, and his sonne of Warwicke,
Were matchlesse men[987] from Barbary to Barwicke.

12.

Through helpe of whome and fortune’s louely looke,

I vndertooke to claime my lawfull right,
And to abash such as against mee tooke,[988]
I raysed power at all poyntes prest to fight:
Of whome the chiefe, that chiefly bare mee spight,
Was Somerset the duke, whome to annoy
I alway sought, through spite, spite to destroy.[989]

13.

And maugre him, so choyse, lo, was my chaunce,

Yea though the quene, that all rulde, tooke his part,[990]
I twise bare rule[991] in Normandy and Fraunce,
And last lieuetenaunt in Ireland, where my hart
Found remedy for euery kinde of smart:
For through[992] the loue my doinges there did breede,
I had their helpe at all times in my neede.
 14.

This spitefull duke, his seely king[993] and queene,

With armed hoastes I thrise met in the fielde,
The first vnfought through treaty made betweene:
The second ioynde, wherein the king did yeelde,
The duke was slaine, the queene enforst to shyelde
Her selfe by flight: the third the queene did fight,
Where I was slaine being ouer matcht by might.[994]

15.

Before this last were other battayles three,

The first the earle of Salisbury led[995] alone,
And[996] fought on Bloreheath, and got[997] victory:
In the next was I and my kinsfolke[998] euery one:
But seeing our souldiers stale vnto our foen,
Wee warely brake our company on a night,
Dissolued our hoast, and tooke our selues to flight.[999]

16.

This boy and I, in Ireland did vs saue,

Mine eldest sonne, with Warwicke and his father,
To Calais got, whence, by the reade I gaue,
They came agayne to London, and did gather
An other hoast whereof I spake not[1000] rather:
And met our foes, slewe many a lord and knight,
And tooke the king, and draue the queene to flight.

17.

This done I came[1001] to England all in haste,

To make a[1002] claime vnto the realme and crowne:
And in the house while parliament did last,
I in the kinge’s seat boldly[1003] sate mee downe,
And claymed it, whereat[1004] the lordes did frowne,
But what for that? I did so well proceede,
That all at last confest it mine indeede.[1005]

18.

But sith the king had raigned now so long,[1006]

They would hee should continue till hee dyed,
And to the end, that than none did mee wrong,[1007]
Protector and heyre apparant they mee[1008] cryed:
But sith the queene and others this denyed,
I sped mee towardes the north, where then shee lay,
In minde by force to cause her to obay.[1009]

19.

Whereof shee warnd preparde a mighty powre,

And ere that mine were altogether ready,
Came swift to Sandale[1010] and besieged my bowre:
Where like a beast I was so rash and heady,
That out I would, there could be no remedy,
With scant fiue thousand souldiers, to assayle
Fowre times so many, encampt to most auayle.[1011]

20.

And so was slaine at first: and while my childe

Scarse twelue yeare old, sought secretly to parte,
That cruell Clifford, lord, nay lorell wylde,
While the infant[1012] wept, and prayed him rue his smart,
Knowing what hee was, with dagger cloue his heart:[1013]
This done, he came to th’camp where I lay dead,
Despoylde my corps and cut away my head.

21.
And when he had put a paper crowne thereon,
As a gawring stocke he sent it to the queene:[1014]
And shee for spite commaunded it anon
To be had to Yorke: where that it mought[1015] bee
seene,
They placed it where other traytors beene:
This mischiefe fortune did mee after death:
Such was my life, and such my losse of breath.[1016]

22.

Wherefore see, Baldwine, that thou set it forth,

To th’end the fraude of fortune may bee knowen,
That eke all princes well[1017] may way the worth
Of thinges, for which the seedes[1018] of warre be sowen:
No state so sure but soone is ouerthrowen:[1019]
No worldly good can counterpeyze the prise
Of halfe the paynes that may thereof arise.

23.

Far[1020] better it were to lose a piece of right,

Than lymmes and life in sousing[1021] for the same:
It is not force of friendship, nor of might,
But God that causeth thinges to fro or frame,[1022]
Not wit but lucke[1023] doth wield the winner’s game:
Wherefore if wee our follyes would refrayne,
Time would redresse all wronges, wee voyde of payne.

24.

Wherefore warne princes not to wade[1024] in war

For any cause, except the realme’s defence:[1025]
Theyr troublous titles are vnworthy far
The bloud, the life, the spoyle of innocence:
Of friendes, of foes,[1026] behold my foule[1027] expence,
And neuer the neare: best therefore tary time,
So right shall raigne, and quyet calme ech crime.[1028]
 [With that[1029] maister Ferrers shooke mee by the sleue, saying:
“Why how now man, do you forget your selfe? belike you minde our
matters very much.” “So I doe indeede,” sayd I,[1030] “for I dreame of
them.” And whan I had rehersed my dreame, wee had long talke
concerning the nature of dreames, which to stint, and to bring vs to
our matter agayne, thus sayd one of them: “I am glad it was your
chaunce to dreame of duke Richard, for it had bene pity to haue
ouerpassed him. And as concerning this lord Clyfford which so
cruely killed his sonne, I purpose to gieue you notes: who (as hee
well deserued) came shortly after to a sodayne death, and yet too
good for so cruell a tyrant. [For on Palmesonday next following,
being the xxix day[1031] of March, in the yeare of Christ a thousand
four hundreth threescore and one, this lord Clyfford with Henry Percy
the thirde earle of Northumberland, the earle of Westmerland, the
lord Dacres, the lord Welles, and other were slayn at Towton in
Yorkeshyre.][1032] Wherefore as you thought you saw and hearde
the headles duke speake through his necke, so now[1033] suppose
you see this lord Clyfford, all armed saue his head, with his brest
plate all gore bloud running from his throate wherein an headlesse
arrow sticketh through which wound hee sayth thus.”[1034]]
 How the Lorde Clyfford for his
straunge and abhominable cruelty
came to as straunge and sodayne a
death,[1035] Anno 1461.[1036]
1.

Open confession axeth open pennaunce,

And wisdome would a man his shame to hyde:[1037]
Yet sith forgieuenes commeth through repentaunce,
I thinke it best that men theyr crimes ascride,
For nought so secret but at length is spyed:
For couer fire, and it will neuer lynne
Till it breake forth, in like case shame and sinne.[1038]

2.

As for my selfe my faultes be out so playne,

And published [so] abroade[1039] in euery place,
That though I would I cannot hide a grayne.[1040]
All care is bootelesse in a curelesse case,
To learne by other’s griefe,[1041] some haue the grace:
And therefore, Baldwine, write my[1042] wretched fall,
The briefe whereof I briefly vtter shall.

3.

I am the same that[1043] slue duke Richard’s childe,

The louely babe that begged[1044] life with teares,
Whereby mine honour[1045] fouly I defilde:
Poore sely lambes the lion neuer teares,
The feeble mouse may ly among the beares,
But wrath of man, his rancour to requite,
Forgets all reason, ruth, and vertue[1046] quite.

4.

I meane by rancour the parentall[1047] wreke

Surnamde a vertue, as the vicious[1048] say,
But litle knowe the wicked[1049] what they speake,
In boldening vs[1050] our enmyes kin to slay:
To punish sinne is good, it is no nay:
They wreke not sinne, but merit wreke for sinne,
That wreke the father’s fault vpon[1051] his kinne.

5.

Because my father lord Iohn Clyfford, dyed,

Slayne at S. Albane’s, in his prince’s ayde,
Agaynst the duke my heart for malice fryed,
So that I could from wreck no way bee stayed,
But to auenge my father’s death, assayde
All meanes I might the duke of Yorke t’annoy,[1052]
And all his kin and friendes to kill and stroy.[1053]

6.

This made mee with my bloudy[1054] dagger wound

His guiltlesse sonne, that neuer agaynst mee storde:
His father’s body lying dead on ground
To pearce with speare, eke with my cruell sworde,
To part his necke, and with his head to bourd
Enuested with a royall paper[1055] crowne,
From place to place to beare it vp and downe.[1056]

7.
But cruelty[1057] can neuer scape the scourge
Of shame, of horror, or of sodayne death:[1058]
Repentaunce selfe, that other sinnes may pourge,
Doth fly from this, so sore the soule it slayeth:
Despayre dissolues the tyraunt’s bitter[1059] breath:
For sodayn vengeaunce sodaynly alightes
On cruell deedes,[1060] to quite theyr cruell spights.
[1061]

8.

This finde I true, for as I lay in stale,

To fight with[1062] duke Richard’s eldest son,
I was destroyde not far[1063] from Dintingdale,
For as I would my gorget haue vndon
To euent[1064] the heat that had mee nigh vndone,
An headles arrow strake mee through the throte,
Where through my soule forsooke his fylthy cote.[1065]

9.

Was this a chaunce? no sure, God’s iust awarde,

Wherein due[1066] iustice playnly doth appeare:
An headlesse arrow payde mee my rewarde:[1067]
For heading Richard lying on his[1068] bere,
And as I would his childe in no wise heare,
So sodayne death bereft my tongue the power
To aske for pardon at my dying[1069] hower.

10.

Wherefore, good Baldwine, warne the[1070] bloudy sort,

To leaue theyr wrath, theyr[1071] rigour to refrayne:
Tell cruell judges horror is the port
Through which they sayle to shame[1072] and sodayn
payne:[1073]
Hell haleth[1074] tyrauntes downe to death amayne:
Was neuer yet, nor shal be cruell deede,
Left vnrewarded with as[1075] cruell meede.
 [Whan this tragedy was ended: “O Lord,” sayd[1076] another,
“how horrible a thing is diuision in a realme, to how many mischiefes
is it the mother, what vice is not therby kindled, what vertue left
vnquenched? for what was the cause of the duke of Yorke’s death,
and of the cruelty of this Clyfford, saue[1077] the variaunce betwene
king Henry and the house of Yorke? which at length, besides millions
of the commons, brought to destruction all the[1078] nobility. For
Edward the duke’s eldest son immediately after his[1079] father was
slayn, through help of the Neuills, gaue the king a battayle, whereat,
besides this Clyfford, and xxxvi thousand other souldiers, were slain
theyr captaynes, the earles[1080] of Northumberland and
Westmerland, with the lordes Dacres and Welles: the[1081] winning
of which fielde brought Edward to the crowne, and the losse draue
king Henry and his wife into Scotland. But as few raignes begin
without bloud, so king Edward to keepe order,[1082] caused Thomas
Courtney earle of Deuonshyre, and Iohn Veer earle of Oxeforde, and
Aubrey Veer eldest sonne to the sayd earle, with diuers[1083] other
his enemies,[1084] to bee attaynted and put to death. And shortly
after he[1085] did execution vpon the duke of Somerset, and the
lordes Hungerford and Rosse, whom he toke prisoners at Exham
fielde. For thither they came with[1086] king Henry out of Scotland,
with an army of Scottes, and fought a battayle, which was lost, and
the most[1087] part of them slayn.[1088] And because these are all
noble men, I will leaue them to Baldwine’s discretion. But seyng the
earle of Worcester was the chiefe instrument whom king Edward
vsed as well in these men’s matters as in like bloudy affayres,
because he should not be forgotten, yee shall here what I haue
noted concerning his tragedy.”[1089]]
The infamovs end of the Lord Tiptoft
Earle of Worcester, for cruelly
executing his Prince’s butcherly
commaundementes, An. 1470.[1090]
1.

The glorious man is not so loth to lurke,

As the infamous glad to lye vnknowen:
Which makes mee, Baldwine, disalow thy worke,
Where prince’s faultes so openly be blowen:
I speake not this alonly for mine owne,
Which were my prince’s (if that they were any)
But for my peeres, in nombre very many.

2.

Or might report vprightly vse her tong,

It would lesse greue vs to augment the matter,
But suer I am thou shalt be forst among,
To frayne the truth the liuing for to flatter,
And otherwhiles in poyntes vnknowen to smatter:
For time neuer was, nor neuer[1091] I thinke shalbe
That truth vnshent should speake in all thinges free.

3.

This doth appeare (I dare say) by my story,

Which diuers writers diuersly declare:
But story writers ought for neyther glory,
Feare, nor fauour, truth of thinges to spare?
But still it fares as alway it did fare,
Affections, feare, or doubts that dayly brue,
Doe cause that stories neuer can be true.[1092]

4.

Unfruitfull Fabian followed the face

Of tyme and dedes, but let the causes slip:
Which Hall hath added, but with double grace,[1093]
For feare I thinke lest trouble might him trip:
“For this or that,” sayeth he, “he felt the whip:”
Thus[1094] story wryters leaue the causes out,
Or so rehearse them as they were in dout.[1095]

5.

But seing causes are the chiefest thinges

That should be noted of the story wryters,
That men may learne what endes all causes bringes,
They be vnworthy the[1096] name of chroniclers
That leaue them cleane out of theyr registers,
Or doubtfully report them: for the fruite
Of reading storyes standeth in the suite.

6.

And therefore, Baldwine, eyther speake vpright[1097]

Of our affayres, or touch them not at all:
As for my selfe I way all thinges so light,
That nought I passe how men report my fall:
The truth whereof yet playnly shew I shall,
That thou mayst write and others thereby rede,
What thinges I did whereof they should take heede.

7.

Thou hast heard of[1098] Tiptoft’s earles of Worcester,

I am that lord that liu’d in Edwarde’s days
The fourth, and was his friend and counsaylour,
And butcher to, as common rumor sayes:
But people’s voyce is neyther shame nor prayse,
For whom they would aliue deuoure to day,
To morow dead they will[1099] worship what they may.

8.

But though the people’s verdit go by chaunce,

Yet was there cause to call mee as they did:
For I, enforst by meane of gouernaunce,
Did execute what euer[1100] my king did bid,
From blame herein my selfe I cannot rid:
But fye vpon the wretched state, that must
Defame it selfe, to serue the prince’s lust.

9.

The cheifest crime wherewith men doe me charge,

Is death of th’[1101] earle of Desmund’s noble sonnes,
Of which the kinge’s charge doth me clere discharge,
By strayt commaundement and iniunctions:
Th’effect whereof so rigorously runnes,
That eyther I must procure to see them dead,
Or for contempt as a traytour loose my head.[1102]

10.

What would mine enemies[1103] doe in such a case,

Obay the king or proper death procure?
They may well say theyr fancy for a face,
But life is sweete, and loue hard to recure:
They would haue done as I did, I am sure,
For seldom will a welthy man at ease
For other’s cause his prince in ought displease.
 11.

How much lesse I, which was lieuetenaunt than

In th’[1104] Irish isle, preferred by the king:
But who for loue or dread of any man,
Consents t’[1105] accomplish any wicked thing,
Although chiefe fault thereof from other spring,
Shall not escape God’s vengeance for his deede,
Who scuseth none that dare doe ill for drede.

12.

This in my king and mee may well appeare,

Which for our faultes did not escape the scourge:
For whan wee thought our state most sure and clere,
The wynd of Warwicke blew vp such a sourge,
As from the realme and crowne the king did pourge,
And mee both from mine office,[1106] friendes, and
wife,
From good report, from honest death and life.

13.

For the earle of Warwicke, through a cancarde

grudge[1107]
Which to king Edward causelesse hee did beare,
Out of his realme by force did make him trudge,
And set king Henry agayne vpon his chaire:
And then all such as Edwarde’s louers were,
As traytours tane were greuously opprest,
But chiefly I, because I lou’d him best.

14.

And, for[1108] my goods and liuinges were not small,

The gapers for them bare[1109] the world in hand
For ten yeares space, that I was cause of all
The executions done[1110] within the land:
For this did such as did not vnderstand
Mine enmies drift, thinke all reportes were true:
And so did[1111] hate mee worse then any Jewe.

15.

For seldom shall a ruler lose his life,

Before false rumors openly be spred:
Wherby this prouerbe is as true as rife,
That ruler’s rumors hunt about a head,
Frowne fortune once, all good report is fled:
For present shew doth make the mayny blind,
And such as see dare not disclose their minde.

16.

Through this was I king Edward’s butcher named,

And bare the shame of all his cruell deedes:
I cleare me not, I worthely was blamed,
Though force was such I must obay him needes:
With hiest rulers seldome well it speedes,
For they be euer nerest to the nyp,
And fault who shall, for all feele they the whip.

17.

For whan I was by parliament attaynted,

King Edward’s euils all were counted mine:
No truth auayled, so lies were faste and painted,
Which made the people at my life, repine,
Crying, “Crucifige, kill that butcher’s lyne:”
That when I should haue gone to Blockham feast,
I could not passe, so sore they one me preast.

18.

And had not bene the officers so strong,

I thinke they would haue eaten[1112] me aliue,
Howbeit, hardly haled from the throng,
I was in the flete fast shrouded by the shriue:
Thus one daye’s life theyr malice did mee gieue,
Which whan they knew, for spite the next day after
They kept them calme, so suffered I the slaughter.

19.

Now tell mee, Baldwine, what fault dost thou finde

In mee, that iustly should such death deserue?
None sure except desire of honour blinde,
Which made mee seeke in offices to serue:
What minde so good that honours make not swerue?
So mayst thou see it only was my state
That caus’d my death, and brought mee so in hate.

20.

Warne therfore all men wisely to beware,

What offices they enterprise to beare:
The hiest alway most maligned are,
Of people’s grudge, and prince’s hate in feare.
For prince’s faults his faultors all men teare:
Which to auoyde, let none such office take,
Saue he that can for right his prince forsake.
 [This earle’s[1113] tragedy was not so soone finished, but one of
the company had prouided for another of a notable person, lord
Tiptoft’s chiefe enemy: concerning whom hee sayd: “Lord God what
trust is there in worldly chaunces? what stay in any prosperity? for
see the earle of Warwicke which caused the earle of Worcester to be
apprehended, attainted, and put to deth, triumphing with his old
imprisoned and new vnprisoned prince, king Henry, was by and by
after, and his brother with him, slayn at Barnet field by king Edward,
whom hee had before time damaged diuers wayes. As first by his
friendes at Banbury fielde, where to reuenge the death of his cosin
Henry[1114] Neuill, syr Iohn Coniers, and Iohn Clappam, his
seruauntes, slue fiue thousand Welshmen, and beheaded theyr
captaynes, the earle of Pembroke and syr Richard Harbert his
brother, after they were yeelded prisoners: of whom syr Richard
Harbert was the tallest gentilman both of his person and handes that
euer I read or heard of. At which time also, Robin of Ridsdale,[1115] a
rebell of the earle of Warwicke’s raysing, tooke the earle Riuers, king
Edwarde’s wiue’s father, and his sonne Iohn, at his manour of
Grafton, and carried them to Northampton, and there without cause
or proces beheaded them. Which spites to requite, king Edward
caused the lord Stafford of Sowthwike, one of Warwicke’s chiefe
frendes, to be taken at Brent march, and headed at Bridgewater.
This caused the earle shortly to raise his power, to encounter the
king which came against him with an army, beside Warwicke at
Wolney, where hee wan the field, tooke the king prisoner, and kept
him a while in Yorkeshire in Middleham castle: whence (as some
say) hee released him againe, but other thinke hee corrupted hys
keepers and so escaped. Then through the lords the matter was
taken vppe betweene them, and they brought to talke togeather, but
because they could not agree, the earle araised a new army,
whereof he made captayn the lord Welles’ sonne, which broile king
Edward minding to appease by pollicy, fouly distained his honour,
committing periury: for hee sent for the lord Welles and his brother
sir Thomas Dymocke, vnder safe conduite promising them vpon his
fayth to keepe them harmelesse. But after, because the lord Welles’
sonne would not dissolue his army, beheaded them both and went
with his power into Lincolnshire and there fought with sir Robert
Welles, and slewe ten thousand of his souldiers (yet ran they away
so fast, that casting of their clothes for the more speede, caused it to
bee called Lose coate fielde) and tooke sir Robert and other, and put
them to death in the same place. This misfortune forced the earle of
Warwicke to saile into Fraunce where hee was entertained of the
king a while, and at last with such poore helpe as hee procured there
of duke Rainer and other, hee came into England againe, and
encreased such a power in king Henrie’s name, that as the lord
Typtofte sayd in his tragedy, king Edward vnable to abide him, was
faine to flie ouer the washes in Lincolnshire to get a ship to saile out
of his kingdome to his brother in lawe the duke of Burgoine. So was
king Henry restored againe to his[1116] kingdome. All these despites
and troubles the earle wrought against king Edward. But Henry was
so infortunate that ere halfe a yeare was expired, king Edward came
backe againe, and enprisoned him and gaue the earle a fielde,
wherein hee slewe both him and his brother. I haue recounted thus
much before hand for the better opening of the story, which, if it
should haue beene spoken in his tragedy, would rather haue made a
volume then a pamphlete. For I entend onely to say in the tragedy,
what I haue noted in the earle of Warwicke’s person, wishing that
these other noble men, whome I haue by the way touched, should
not bee forgotten. And therefore imagine that you see this earle lying
with his brother in Paule’s church[1117] in his coate armoure, with
such a face and countinaunce as he beareth in portraiture ouer the
dore in Paule’s, at the going down to Iesus chappell from the south
end of the quier stayres, and saying as followeth.”]