The Trace Fossil MesozoicCenozoic PDF

Topics in Geobiology 40
M. Gabriela Mángano
Luis A. Buatois Editors
The Trace-Fossil
Record of Major
Evolutionary Events
Volume 2: Mesozoic and Cenozoic
Topics in Geobiology
Volume 40
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M. Gabriela Mángano • Luis A. Buatois
Editors
The Trace-Fossil Record of

Major Evolutionary Events
Volume 2: Mesozoic and Cenozoic
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 978-94-017-9596-8 ISBN 978-94-017-9597-5 (eBook)
DOI 10.1007/978-94-017-9597-5
Library of Congress Control Number: 2016951934
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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.
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
8 The Late Triassic Mass Extinction Event ............................................. 1

Colin Barras and Richard J. Twitchett
9 The Mesozoic Marine Revolution .......................................................... 19
Luis A. Buatois, Noelia B. Carmona, H. Allen Curran,
Renata G. Netto, M. Gabriela Mángano, and Andreas Wetzel
10 The Mesozoic Vertebrate Radiation in Terrestrial Settings ................ 135
Massimo Bernardi, Fabio Massimo Petti,
Laura Piñuela, José Carlos García-Ramos, Marco Avanzini,
and Martin G. Lockley
11 The Mesozoic Lacustrine Revolution .................................................... 179
Luis A. Buatois, Conrad C. Labandeira, M. Gabriela Mángano,
Andrew Cohen, and Sebastian Voigt
12 The End-Cretaceous Extinction and Ecosystem Change .................... 265
Conrad C. Labandeira, Francisco J. Rodríguez-Tovar,
and Alfred Uchman
13 The Phanerozoic Four Revolutions and Evolution
of Paleosol Ichnofacies ............................................................................ 301
Jorge F. Genise, Emilio Bedatou, Eduardo S. Bellosi,
Laura C. Sarzetti, M. Victoria Sánchez, and J. Marcelo Krause
14 The Cenozoic Radiation of Mammals ................................................... 371
Verónica Krapovickas and Sergio Vizcaíno
15 Major Events in Hominin Evolution ..................................................... 411
Martin Lockley, Jeff Meldrum, and Jeong Yul Kim
xi
xii Contents
16 Recurrent Patterns and Processes: The Significance

of Ichnology in Evolutionary Paleoecology........................................... 449
Luis A. Buatois and M. Gabriela Mángano
Index ................................................................................................................. 475

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 ichnol-
ogy has received far less attention. However, even a cursory review of the recent
xiii
xiv Prologue
paleobiologic literature shows that there is an increased awareness of the impor-

tance 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).
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.
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.
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 data is used to reveal the nature of Ediacaran ecosystems and the complexi-
ties 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 dif-
ferent 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
mass extinction. Their review suggests that climatic and environmental change were
main factors controlling benthic communities on land and in the sea.
xvi Prologue
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 structure
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 revolutions in paleosol
ichnofaunas. In addition, this chapter provides an interesting twist to the concept of
ichnofacies by showing that their establishment in terrestrial settings 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 in South
American mammals during the Cenozoic. The authors show that Eocene-Oligocene
trackway assemblages mostly consist of forms of uncertain affinity, whereas later
Prologue xvii
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 interactions 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 evo-
lutionary 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 Chap. 10, of sit-
ting 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
Narbonne, Eduardo Olivero, Paul Olsen, Roy Plotnick, Gustavo Politis, Charles
xviii Prologue
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
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Uchman A (2004) Phanerozoic history of deep-sea trace fossils. In: McIlroy D (ed) The application
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Chapter 8
The Late Triassic Mass Extinction Event
Colin Barras and Richard J. Twitchett
8.1 Introduction
The end of the Triassic (~200 Ma) was identified by Newell (1963) as marking one
of the five largest extinction episodes of the Phanerozoic. From the perspective of
ecologic impact on the biosphere, it ranks third (McGhee et al. 2004). In the marine
realm some 23 % of families and 50 % of genera were lost (Sepkoski 1981, 1993)
with ammonites, bivalves, radiolarians, and coral reefs suffering most (McElwain
et al. 2007; Keissling and Simpson 2011). On land, plant biodiversity declined at
genus and species levels in the latest Triassic, as shown by local palynologic studies
and analyses of macrofossils (e.g., in East Greenland; McElwain et al. 2007;
McElwain et al. 2009; Mander et al. 2010). Widespread floral change would have
affected other terrestrial groups such as vertebrates, which suffered a 45 % decline
at the family level in eastern North America during the final stages of the Triassic
(Olsen et al. 1987). Some common archosaurs disappeared at this time, although the
theropod dinosaurs appear to have increased in abundance and size across the
Triassic–Jurassic (T–J) boundary (Olsen et al. 2002).
For decades this event was relatively poorly studied, but a concerted effort in recent
years, especially following formal designation of the Global Stratotype Section and
Point (GSSP) for the base of the Jurassic (Hillebrandt et al. 2007), has led to a clearer
picture of the sequence of events and likely contributing factors. Competing scenarios
included sea level change (Hallam and Wignall 1997), a bolide impact (Olsen et al.
2002), and the dissociation of gas hydrates (Beerling and Berner 2002). However, it
C. Barras (*)
511 Taylor Lane, Chelsea, MI 48118, USA
e-mail: [email protected]
R.J. Twitchett
Department of Earth Sciences, The Natural History Museum,
London SW7 5BD, United Kingdom
© 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 40,
DOI 10.1007/978-94-017-9597-5_8
2 C. Barras and R.J. Twitchett
now appears likely that the principal kill mechanism was concomitant changes in
climate, atmosphere, and oceans associated with the formation of the Central Atlantic
Magmatic Province (CAMP), a large igneous province stretching roughly 5000 km in
a north–south direction and 2000 km in an east–west direction, and covering areas of
northwest Europe, West Africa, eastern North America, and Brazil (McHone 2000;
Fig. 8.8.1). CAMP is among the largest of all continental large igneous provinces
(Marzoli et al. 1999), and the extinctions in marine and terrestrial realms coincide with
the onset and early phases of volcanic activity in the latest Triassic (Marzoli et al. 2004;
Knight et al. 2004; Deenen et al. 2010; Mander et al. 2013).
At around the time of emplacement of the earliest CAMP lavas there is good
evidence that the level of atmospheric carbon dioxide rose rapidly. Studies of fossil
leaf stomatal density (McElwain et al. 1999; Retallack 2001; Beerling 2002;
Steinthorsdottir et al. 2011) and soil carbonates (Schaller et al. 2011) suggest a rise
in atmospheric CO2 across the extinction interval from 600–1000 ppm to 2000–
2500 ppm or from 1000 ppm to 2000–3000 ppm, depending on the respective proxy.
Disruption to the global carbon system is recorded by negative carbon isotope
Fig. 8.1 Approximate extent of the Central Atlantic Magmatic Province, after Wignall (2001a),
showing the two locations studied specifically for the effects of the end-Triassic extinction on the
trace-fossil record
8 The Late Triassic Mass Extinction Event 3
excursions from a number of localities worldwide, including among others example

in Austria (Ruhl et al. 2009), Nevada, USA (Guex et al. 2004), the Newark Basin of
eastern USA (Whiteside et al. 2010), East Greenland (McElwain et al. 1999), British
Columbia, Canada (Ward et al. 2004; Williford et al. 2006), Italy (Galli et al. 2005),
and England (Hesselbo et al. 2002; Whiteside et al. 2010).
Such a rapid increase in pCO2 would have led to global warming, with dramatic
effects on terrestrial climate zones, the global hydrologic cycle, and the circulation
and dissolved oxygen content of the world’s oceans (e.g., Kidder and Worsley
2010). Some authors have suggested that elevated CO2 would have lowered the pH
of surface waters, leading to ocean acidification that would have had detrimental
effects for calcareous organisms (Hautmann 2004; Schootbrugge et al. 2007;
Kiessling and Simpson 2011). Modern experiments suggest, however, that the
responses to lowered pH vary dramatically between different marine taxa (Findlay
et al. 2011), and there is evidence from the fossil record that calcification of some
bivalves actually increased at this time (Mander et al. 2008).
Some have questioned whether extinction near the end of the Triassic was
abrupt, or more gradual in nature (Hallam 2002; Cuny 1995; Tanner et al. 2004).
Reported faunal changes often coincide with facies changes that might give a
false impression of sudden extinction (Hallam 2002). For bivalves, at least, it
appears that changes in facies, preservation, and other rock record biases have not
significantly affected the extinction interval but may have led to the under-repre-
sentation of some groups, such as aragonitic, deeper infaunal taxa, in the immedi-
ate aftermath (Mander and Twitchett 2008). Recent assessment of the fossil-plant
record from East Greenland (McElwain et al. 2009) demonstrates that biodiver-
sity decline was relatively abrupt and began around the time of initial CAMP
volcanism. Furthermore, analysis of the palynologic records of East Greenland
and St Audrie’s Bay, UK, demonstrate that terrestrial ecosystems were affected at
the same time as those in the sea, and the extinctions occurred during the steep
rise in atmospheric CO2 (Mander et al. 2013).
8.2 Trace-Fossil Record Across the T–J Boundary
While the body-fossil record across the T–J boundary has been relatively well studied,
the trace-fossil record remains less well known. However, while trace fossils cannot
usually be tied readily to the trace maker their study can still contribute to a better
understanding of the nature of faunal turnover during extinction events such as the
Late Triassic extinction, making the few studies that have been conducted important.
In terrestrial settings, such as the Newark Basin of eastern North America, which
contains one of the best temporally constrained nonmarine vertebrate records
through the extinction interval, skeletal remains are much less common than foot-
prints making study of the footprints particularly useful for elucidating the nature of
the vertebrate faunal response (Olsen et al. 2002) (Fig. 8.8.2). In the marine realm,
trace fossils are important because most marine ecosystems are dominated by
Fig. 8.2 Terrestrial ichnotaxonomic trends through the Triassic–Jurassic interval of the Eastern
USA and shallow marine ichnotaxonomic trends of Southern England. Gray horizontal line shows
approximate location of the initial carbon isotope excursion that marks the extinction interval.
Graphs show trend in maximum length of theropod tracks in the Eastern USA, and trend in average
diameter of Diplocraterion in southern England (X’s mark the maximum diameter of
Diplocraterion). H. = Hettangian. Sin. = Sinemurian. T–J = Triassic–Jurassic boundary. After Olsen
et al. (2002) and Barras and Twitchett (2007)
nonmineralized taxa, which rarely fossilize (Allison and Briggs 1991). The traces
that they leave may be more readily preserved, which makes the marine trace-fossil
record the best, and sometimes only, window into the responses of these dominant,
soft-bodied organisms to catastrophic extinction (Twitchett and Barras 2004).
Finally, in both marine and terrestrial regimes, the trace-fossil record is spared from
such taphonomic effects as transport or reworking (Barras and Twitchett 2007),
making its study potentially less ambiguous than some body fossil studies.
8.2.1 Terrestrial Record
Terrestrial trace-fossil studies around the T–J boundary often focus on specific short
sequences within either the Triassic or Jurassic, rather than longer successions that
span the boundary and the extinction event itself. Low-diversity invertebrate trace-
fossil assemblages associated with lake margins have been reported for both the
Late Triassic (Lucas and Lerner 2006) and Early Jurassic (Lucas et al. 2006b) of the
western United States, the Triassic of Greenland (Bromley and Asgaard 1979) and
the Late Triassic of Germany (Schlirf et al. 2001).
Elsewhere in northern Europe the terrestrial record of invertebrate trace fossils is
more extensive. In southern Sweden, for example, Late Triassic and Early Jurassic
fluvio-deltaic deposits with some evidence of brackish marine influence preserve a
moderately diverse ichnofauna, including horizontal (Cochlichnus, Palaeophycus,
Planolites, Rhizocorallium, Teichichnus) and vertical (Diplocraterion,
Monocraterion, Skolithos) burrows and grazing traces, as well as arthropod track-
ways (Diplichnites), and the bivalve resting trace Lockeia (Pieńkowski 1991a,b;
Gierliński and Ahlberg 1994; Ahlberg 1994; Ahlberg and Arndorff 1994).
Further south, in Poland, exposures of Early Jurassic rocks formed in lacustrine
and fluvial environments with some subaerial exposure contain a diverse ichnofauna.
Finds here include Lockeia, arthropod resting traces (Rusophycus), arthropod track-
ways, burrows and borings (Cruziana, Diplichnites, Linckichnus, Spongeliomorpha,
Xylonichnus) equilibrichnia formed by freshwater bivalves (Calceoformites,
Scalichnus), and horizontal or vertical burrows (Cochlichnus, Helminthoidichnites,
Scoyenia, Planolites, Palaeophycus). There are also possible arthropod nest struc-
tures. In brackish marine influenced deposits, the limulid trackway Kouphichnium
also occurs (Pieńkowski 2004; Pieńkowski and Niedźwiedzki 2008).
Lacustrine assemblages associated with Triassic deltas in Argentina have been
studied for their trace fossils (Melchor 2004). Ichnofaunas typically include horizon-
tal or vertical burrows and grazing traces (e.g., Palaeophycus, Skolithos, Planolites,
Helminthoidichnites, Helminthopsis, Treptichnus) and backfilled burrows (e.g.,
Scoyenia, Taenidium) with rare arthropod walking traces, such as Diplichnites (e.g.,
Lucas et al. 2006b). However, the invertebrate trace-fossil record is too sparse to
reveal ichnotaxonomic trends, should they exist, through the extinction interval itself.
Likewise, most studies of the terrestrial vertebrate footprint fossil record often
have as their focus particular formations within the Late Triassic or Early Jurassic.
Nevertheless, there is evidence from the trace-fossil record that dinosaurs increased
in abundance, diversity, and size through the Late Triassic and into the Early
Jurassic. Hunt and Lucas (2007) summarized the record of Late Triassic tetrapod
trackways of the western United States. The Revueltian (early to mid Norian)
reveals a moderately diverse ichnofauna. Rhynchosauroides, considered by Hunt
and Lucas (2007) to represent the tracks of a sphenodontian, is found in Arizona,
New Mexico, and Utah. Apatopus, long considered the track of a phytosoaur (e.g.,
Foster et al. 2000), is found in Utah. Other ichnotaxa present in the region suggest
the presence of dinosaurs: Barrancapus, possibly representative of a prosauropod
trackway (Hunt et al. 2001), is found in New Mexico and Arizona; another
prosauropod ichnotaxon (Evazoum) is found in Utah; and Grallator, interpreted as
the tracks left by a small theropod, is also found in Utah.
The overlying Apachean (late Norian to Rhaetian) contains a diverse ichnofauna
across several states, including a more diverse dinosaurian ichnofauna (Hunt and
Lucas 2007). The possible sphenodontian track Rhynchosauroides is found in
Colorado and New Mexico; carnivorous archosaur trackways (Apatopus) are found
in New Mexico, while possibly herbivorous archosaur trackways (Brachychirotherium)
are known from Utah, Colorado, New Mexico, and Oklahoma (Lucas and Tanner
2007). Gwyneddichnium, unambiguously linked to the Late Triassic reptile
Tanytrachelos, is found in Utah and Colorado. There is also evidence of synapsid
trackways (e.g., Brasilichnium) in Colorado. The dinosaurs, represented by three
ichnotaxa in the preceding Revueltian, are more diverse and widespread in the
Apachean. The small theropod trackway Grallator is abundant in Utah, and is also
found in Wyoming, Colorado, Arizona, New Mexico, and Oklahoma. A larger thero-
pod probably left Anchisauripus tracks in Utah and New Mexico, and Eubrontes is
known from rocks that may date to the uppermost Triassic (Fig. 8.8.3e). Herbivorous
saurischian dinosaurs are represented by the prosauropod trackway Evazoum in
Utah, Colorado, and New Mexico, Eosauropus in Utah, Colorado, Arizona, and New
Mexico, and sauropodomorph trackways Pseudotetrasauropus and Tetrasauropus in
Arizona, Utah and Colorado (Lucas and Tanner 2007). Atreipus, found in Utah, may
have been produced by an ornithischian (Olsen and Baird 1986).
In the lowermost Jurassic of Utah, Arizona and Colorado, large theropod tracks
(Eubrontes) dominate, although smaller theropod tracks (Grallator) and sauropodo-
morph tracks (Otozoum; Fig. 8.8.3c) are also present. All three are also found in dino-
saur-dominated trace-fossil assemblages dating to the Sinemurian (Fig. 8.8.3d), together
with the tracks of ornithischian (Anomoepus) and prosauropod (Otozoum) dinosaurs,
synapsids (Brasilichnium) and crocodilians (Batrachopus; Lucas and Tanner 2007).
More evidence for the rise of the dinosaurs through the T–J interval comes from
Olsen et al. (2002), one of the few studies that consider the terrestrial trace-fossil
record specifically from the perspective of the Late Triassic extinction (Fig. 8.8.2).
The focus of the study is the Newark Supergroup, preserved in New York, New
Jersey, and Pennsylvania in the eastern United States. The increase in dinosaur
(ichno)taxonomic diversity that is apparent in the western United States is found
here as well; Olsen et al. (2002) also reported a concomitant increase in the size of
individual tracks, suggesting an increase in dinosaur body size in the latest Triassic.
Nondinosaurian ichnodiversity increases through the Norian and Rhaetian in the
Newark Supergroup too, with Rhynchosauroides and Gwyneddichnium common.
The probable silesaurian ichnogenus Atreipus is the most common member of the
dinosauriform clade of dinosauromorphs (as opposed to dinosaur: Irmis et al. 2007)
until the middle Rhaetian. Trackways attributed to theropods include Grallator,
which occurs throughout the Carnian, Norian, and Rhaetian, and Anchisauripus,
which first appears in the Norian.
In the Newark Basin, the Late Triassic extinction is marked by a facies change indi-
cating wetter conditions, a coal bed, an apparent iridium anomaly, and a fern spore spike
(Olsen et al. 2002). This horizon is correlated with the earliest phase of CAMP activity,
Fig. 8.3 Marine and terrestrial trace fossils of the Triassic–Jurassic interval. (a) Small
Diplocraterion burrows in the angulata Zone (Hettangian) of Pinhay Bay, Dorset, UK; (b) Large
Diplocraterion in the semicostatum Zone (Sinemurian) of Helwell Bay, Somerset, UK; (c) Track
of a prosauropod dinosaur (Otozoum) in Wingate Sandstone (?Hettangian) of Gateway, Colorado;
(d) Footprint of a theropod dinosaur (Grallator) in Kayenta Formation (Sinemurian to
Pliensbachian) of St. George, Utah; (e) Footprint of a large theropod dinosaur (Eubrontes) in upper
part of Dinosaur Canyon Member of Moenave Formation (?Late Triassic) of St. George, Utah.
Images (c), (d), and (e) reproduced with permission of Spencer G. Lucas, New Mexico Museum
of Natural History and Science, Albuquerque
recorded in Morocco, and the marine extinction event (Deenen et al. 2010; Mander et al.
2013). It marks a decline in the diversity of nondinosaurian tracks and an increase in
both diversity and size of dinosaurian tracks (Olsen et al. 2002). The nondinosaurian
ichnotaxa Gwyneddichnium, Apatopus, and Brachychirotherium parvum disappear,
whereas Rhynchosauroides and Batrachopus span the extinction, the latter left by a
crocodylomorph. Theropod dinosaurs survived the extinction, as evidenced by the
records of Grallator and Anchisauripus, with new ichnotaxa such as Eubrontes appear-
ing afterwards. Slightly higher in the section—an estimated >100 ky after the extinc-
tion—tracks belonging to prosauropods (Otozoum) and ornithischians (Anomoepus)
appear. The prosauropod tracks increase in frequency, and the ornithischian tracks
increase in size, through the remaining succession.
The Late Triassic extinction level also marks the first appearance of much
larger theropod tracks (>25 cm in length) belonging to the ichnospecies
Eubrontes giganteus, considered by some to be simply larger representatives of
Grallator (e.g., Lucas et al. 2006a), despite differences in relative elongation of
members of the two ichnogenera (e.g., Olsen 1980). Eubrontes may have been
left by a ceratosaur such as Dilophosaurus (Lucas et al. 2006a), although some
(Weems 2003) have argued the trackmaker was a Plateosaurus-like
prosauropod.
Olsen et al. (2002) suggested that the increase in size of theropod tracks may
represent either dispersal from an unknown location or indicate an evolutionary
response to reduced competitive pressure and ecologic release in the aftermath
of extinction. Lucas et al. (2006a) rejected this ecologic release hypothesis on
the basis that the ichnogenus Eubrontes has been recorded from as far back as the
Carnian. However, they failed to demonstrate that large-sized examples of the
specific ichnospecies Eubrontes giganteus are known from preextinction strata,
or that there is no size change through the extinction event, which would be a
better test of the Olsen et al. (2002) hypothesis.
The vertebrate footprint record from Europe is broadly consistent with the North
American picture. The theropod ichnotaxon Grallator has been reported from the
Rhaetian of southern Sweden (Gierliński and Ahlberg 1994). Otherwise, Late
Triassic vertebrate footprints are rare in Europe, although nondinosaurian tracks
(possibly Rhynchosauroides) are seen in a Late Triassic lagoon in southwest
England that evidently experienced occasional periods of emergence (Allington-
Jones et al. 2010).
As in North America, Grallator is also found in Lower Jurassic European
deposits—in Poland and southern Sweden—indicating that theropod dinosaurs
in the region survived the extinction interval (Ahlberg 1994; Gierliński and
Ahlberg 1994; Gierliński and Pieńkowski 1999). Indeed, the Early Jurassic
(Hettangian) of Poland preserves a diverse dinosaur ichnofauna (Gierliński and
Pieńkowski 1999; Pieńkowski 2004). Here, sauropod tracks (Parabrontopodus)
and medium to large theropod footprints (Anchisauripus and Kayentapus) are
found in rocks formed in an inland environment. A shoreline environment pre-
serves ornithischians (Anomoepus, Atreipus and Moyenisauropus) and thero-
pods (Megalosauripus, Grallator, and Anchisauripus), while Plesiornis, a
possible protoavian track, is found in rocks formed under fluvial influence
(Gierliński and Pieńkowski 1999; Pieńkowski 2004; Gierliński et al. 2010).
Early mammalian prints also occur within Poland's nonmarine Lower Jurassic
deposits (Pieńkowski 2004; Gierliński et al. 2010).
The Polish record, like that of North America, contains its first large theropod
dinosaur footprints after the Late Triassic extinction. Eubrontes footprints (>34 cm
in length) occur in the Hettangian, together with a single, unusually large (55 cm in
length) footprint that may be Megalosauripus (Gierliński et al. 2010). Large sauro-
pods are present too; some of the Parabrontopodus tracks found in the Early Jurassic
of Poland are to 42 cm in length (Gierliński et al. 2010).
8.2.2 Marine Record
8.2.2.1 Deep-sea Ichnofauna Across the T–J Boundary
Nearly 30 ichnogenera inhabited the deep-sea in the Carboniferous, before a halv-

ing of ichnotaxonomic diversity in the Permian (Uchman 2004). Numbers
remained low through the Triassic and Jurassic, rising again to Carboniferous
levels only in the Cretaceous. However, a recent analysis of Upper Triassic deep-
sea fan deposits in the Al Ayn Formation of Oman suggested ichnofaunal diversity
was abundant at this time. Wetzel et al. (2007) recorded 32 ichnogenera, making
this the most diverse deep-sea ichnofauna known until the Early Cretaceous. The
ichnofauna includes predepositional forms that are cast by turbidites (Belorhaphe,
Circulichnis, Desmograpton, Glockerichnus, Gordia, Helminthopsis, Lorenzinia,
Megagrapton, Paleodictyon, Strobiloraphe, and Treptichnus) and postdeposi-
tional forms that penetrate the turbidites (Arenicolites, Protovirgularia, Zoophycos,
Thalassinoides, and Ophiomorpha).
The deep-sea ichnofauna may have reduced in diversity in response to Permo-
Carboniferous glaciation (Uchman 2004)—deep-sea ichnotaxonomic diversity also
fell during episodes of glaciation in the Quaternary (Cronin and Raymo 1997).
Warmer deep-sea conditions in the Late Triassic may then have contributed to an
increase in ichnotaxonomic diversity. However, there is too little data as yet to
search for any effects of the end-Triassic extinction on the deep-sea ichnofauna.
8.2.2.2 Shallow-Marine Ichnofauna Across the T–J Boundary
As with the terrestrial record, a number of studies have described shallow marine
trace fossils from Upper Triassic or Lower Jurassic strata (e.g., Swift and Duffin
1999; Moghadam and Paul 2000), but few have studied ichnologic change through
the extinction event.
A Late Triassic (Rhaetian) to Early Jurassic (Sinemurian) near-shore marine suc-
cession in southern Poland contains a moderately diverse ichnofauna including
arthropod traces (Cruziana, Thalassinoides), asterozoan traces (Asteriacites), hori-
zontal burrows (Planolites, Palaeophycus), and vertically branching Chondrites
burrows (Uchman 1991).
Lower Jurassic lagoonal deposits in southern Sweden contain the bivalve traces
Lockeia and Protovirgularia (Ahlberg 1994). Elsewhere in the region, the Early
Jurassic is characterized by muds deposited in a tidally influenced environment.
Horizontal and vertical burrows (Cylindrichnus, Diplocraterion, Monocraterion,
Phycodes, Planolites, Rhizocorallium, Skolithos, Teichichnus, Thalassinoides),
bivalve traces (Lockeia), and arthropod traces (Kouphichnium, Merostomichnites)
are found here (Pieńkowski 1991a; Ahlberg 1994). In deposits dating to the
Sinemurian, the ichnofauna comprises Diplocraterion, Planolites, Rhizocorallium,
and Teichichnus (Pieńkowski 1991a). Shales deposited in relatively deep and often
dysaerobic conditions contain common Chondrites, with rare Rhizocorallium

indicating occasional oxygenation (Pieńkowski 1991a). Hettangian to Sinemurian
marine storm deposits in southern Sweden contain a burrow-dominated ichnofauna
of Teichichnus, Thalassinoides, Rhizocorallium, and rare Diplocraterion
(Pieńkowski 1991a). Storm-influenced shallow marine successions from the Early
Jurassic (Sinemurian) of Poland are rich in Diplocraterion (Pieńkowski 1991b).
The T–J shallow-marine trace-fossil records of the UK, Austria, and Nevada,
USA, have been examined by Twitchett and Barras (2004), with a more detailed
analysis of the trace-fossil record in England by Barras and Twitchett (2007;
Fig. 8.8.2). A key section is at St Audrie's Bay, Somerset, UK, which had been con-
sidered as a candidate for the Global Stratotype Section and Point for the base of the
Jurassic (e.g., Warrington et al. 1994). Although the first appearance of the ammonite
Psiloceras spelae in the Kuhjoch section of Austria was ultimately chosen as the
GSSP (Hillebrandt et al. 2007), the St Audrie’s Bay section remains critical for cor-
relating the marine and terrestrial records (Deenen et al. 2010; Mander et al. 2013).
The Late Triassic extinction event correlates with an isotopic excursion in the mid-
dle of the Lilstock Formation in St Audrie’s Bay, just prior to the boundary between
the Cotham and Langport members (Hesselbo et al. 2004; Mander et al. 2013). Trace
fossils formed prior to and possibly during the extinction event, in the Rhaetian
Westbury and Lilstock formations, have been documented by a number of researchers
(Wang 1993; Swift and Duffin 1999; Barras and Twitchett 2007; Allington-Jones et al.
2010). To the east of St Audrie's Bay, at Westbury-on-Severn, the Westbury Formation
begins with siltstones formed in relatively oxygenated shallow marine conditions.
These contain vertical burrows (Diplocraterion and Skolithos; Wang 1993). Above
these marine siltstones, conditions become more restricted and indicative of a shallow
lagoonal setting with lower oxygen. Bioturbation in discrete horizons here probably
records storm events that brought in animals, sediment, and oxygen to the restricted
environment (Wang 1993; Allington-Jones et al. 2010). Some trace fossils here
(Cruziana, Rusophycus, and Selenichnites) probably show the activity of xiphosurids
(Wang 1993). Allington-Jones et al (2010) report a diverse ichnofauna within these
storm layers, including burrows (Chondrites, Planolites, Rhizocorallium, Taenidium,
Thalassinoides), bivalve (Lockeia, Protovirgularia), and arthropod (Merostomichnites,
Monomorphichnus, Oniscoidichnus) traces, and a previously undescribed grazing
trail, Radichnus. At St Audrie’s Bay, the Rhaetian Westbury Formation is mostly thor-
oughly bioturbated, although few discrete trace fossils are preserved in the mudstone
facies (Swift and Duffin 1999). Elsewhere in the region, shallower, sandier facies
preserve a suite of trace fossils such, as Arenicolites, Diplocraterion, Kouphichnium,
Lockeia Palaeophycus, and Planolites (Swift and Duffin 1999).
Following the extinction event, the Langport Member of the Lilstock Formation
is well laminated and devoid of trace fossils at St Audrie's Bay, but elsewhere a low
diversity, small-sized assemblage may be recorded. At Long Itchington,
Warwickshire, for example, the Langport Member is moderately bioturbated with
the beds measuring 2–3 on Droser and Bottjer's (1986) ichnofabric index for verti-
cal sections (vii), while bedding planes assessed using Miller and Smail's (1997)
ichnofabric index for horizontal exposures (hii) measure 2–4 (Barras and Twitchett
2007). Rare examples of Arenicolites, Diplocraterion, Palaeophycus, and
Rhizocorallium are found, although these are small in size; Arenicolites burrow
diameter is generally 1–2 mm. Larger diameter Diplocraterion are locally abundant
at the boundary between the Langport Member and the overlying Blue Lias
Formation at Pinhay Bay in Dorset (Barras and Twitchett 2007).
The boundary between the Lilstock Formation and overlying Blue Lias Formation
represents a significant deepening of the marine environment, probably linked to
sea level rise during global warming, which needs to be borne in mind when com-
paring the ichnofaunas of the two formations. The lowermost beds of the Blue Lias
Formation are termed the “Pre-Planorbis Beds,” and were deposited during peak
global warming around the T–J boundary. They are unbioturbated (vii1) in their
lower part, but by the upper “Pre-Planorbis Beds” extensive bioturbation (vii4-5) is
recorded. Milankovitch-scale lithologic cyclicity is evident in the Blue Lias
Formation, and throughout the formation there is a small-scale alternation of lami-
nated and bioturbated beds (Moghadam and Paul 2000). From the base of the pla-
norbis Zone upwards, however, the latter are all well bioturbated (vii4-6) through to
at least the Sinemurian bucklandi Zone (Barras and Twitchett 2007).
The lowest recorded ichnotaxa in the Blue Lias Formation are Arenicolites and
Thalassinoides from the upper Pre-Planorbis Beds of Pinhay Bay. During the over-
lying Hettangian there is a stepwise increase in ichnotaxonomic diversity, which
shows a similar trend at Pinhay Bay and St Audrie's Bay (Twitchett and Barras
2004; Barras and Twitchett 2007). At other locations, the rock record may be less
complete but the trends are similar. At Long Itchington, for example, the ichnofauna
of the Rugby Limestone Member is similar to that recorded in limestones of the
same age (i.e. upper angulata and bucklandi zones) at Pinhay Bay and St Audrie's
Bay. At its most diverse, in the upper angulata Zone and above, the Blue Lias
Formation ichnofauna includes Arenicolites, Chondrites, Diplocraterion,
Palaeophycus, Planolites, Rhizocorallium, and Thalassinoides. It is only within and
above the angulata Zone that the four ichnotaxa recorded in the Triassic Lilstock
Formation co-occur again (Barras and Twitchett 2007).
When they first (re)appear in the angulata Zone, Diplocraterion burrows are signifi-
cantly (p > 0.95) smaller than are similar burrows in the Langport Member at Pinhay Bay.
Both Diplocraterion and Arenicolites significantly (p > 0.95) increase in size from the
angulata Zone through to the bucklandi Zone (Barras and Twitchett 2007; Fig. 8.8.3a, b).
As burrow size is correlated to body size (Savrda and Bottjer 1986), Barras and
Twitchett (2007) inferred that the soft-bodied trace-making infauna was small in the
“Pre-Planorbis Beds,” the planorbis and liasicus zones, and increased in size from the
later Blue Lias Formation. Given the link between small body size and low oxygen
concentration (e.g., Rhoads and Morse 1971), and evidence of anoxic and euxinic con-
ditions in the shales of the lower “Pre-Planorbis Beds” (e.g., Wignall 2001b), Barras and
Twitchett (2007) further suggested that the ichnofauna records a gradual return to oxy-
gen rich conditions following a period of anoxia in the “Pre-Planorbis Beds.”
The ichnofaunal record in Austria and Nevada is less extensive than that in England,
and has been little studied (Hallam and Wignall 2000, Twitchett and Barras 2004).
Limestones belonging to the Rhaetian-aged Kössen Formation, exposed at Gaissau and
Kendelbach Formation in Central Austria, are well bioturbated (vii3–4) and contain
Diplocraterion, Planolites, Rhizocorallium, Skolithos, and Zoophycos. These ichnotaxa
are absent from the uppermost beds of the Kössen Formation, and from the overlying
Kendelbach Formation—a roughly 10-m-thick succession of limestones thought to
encompass the Hettangian (Twitchett and Barras 2004). However, the shelly fossil
record suggests any extinction here was followed by swift recovery before the first
appearance of Psiloceras in the Kendelbach Formation (Hallam 1990).
In the New York Canyon area of the Gabbs Valley Ranges, Nevada, there is a
modestly diverse ichnofauna in the Rhaetian-aged Mount Hyatt Member of the
Gabbs Formation. Arenicolites, Planolites, Rhizocorallium, Skolithos, and
Thalassinoides are found in the limestone-dominated succession, although the rocks
are relatively poorly bioturbated (vii2; Twitchett and Barras 2004). Within the lower
reaches of the overlying Muller Canyon Member of the Gabbs Formation, Guex
et al. (2004) found evidence of a negative carbon excursion that they related to
emplacement of CAMP. Locally intense bioturbation and thoroughly mottled hori-
zons with the horizontal trace Helminthoida (now Nereites irregularis) have been
reported (Hallam and Wignall 2000), but these were not located during a later study
and in general trace fossils are rare within this member (Twitchett and Barras 2004).
The Ferguson Hill Member of the Sunrise Formation lies above and is dated as
late Hettangian to early Sinemurian (Taylor et al. 1983). Arenicolites, Planolites,
Rhizocorallium, Skolithos, and Thalassinoides all reappear in the Ferguson Hill
Member of the Sunrise Formation, and Chondrites and Diplocraterion are also
reported (Twitchett and Barras 2004). The reappearance of an abundant ichnofauna
is associated with, and possibly the result of, facies and palaeoenvironmental
changes; the Ferguson Hill Member is limestone dominated, and similar in appear-
ance to the Mount Hyatt Member.
In summary, there are clear local changes in marine ichnofaunas through the T–J
boundary interval in England, Austria, and Nevada that may relate to CAMP-induced
warming-related environmental changes. While there is no evidence for extinction of
trace makers in the Late Triassic, unlike the terrestrial record, there are clear ecologic
changes in terms of burrow size, depth, and diversity during this interval. Locally, it
appears that diverse Late Triassic ichnofaunas, representing a healthy benthic eco-
system, disappear around the extinction level and only reappear from the latest
Hettangian onwards (Twitchett and Barras 2004). In between, although there are
local differences, it appears that shallow marine ichnofaunas are typically of rela-
tively low diversity, with small-sized and shallow penetrating burrows.
8.3 Discussion
It now seems likely that the Late Triassic extinction was largely a result of global
warming and related environmental effects, associated with the widespread volcanism
that led to the formation of CAMP. Furthermore, recent correlations (e.g., Mander et al.
2013) imply that atmospheric CO2 remained high through the T–J boundary before
declining to preextinction levels sometime in the later Hettangian, following cessation
of CAMP activity. Although the data are currently sparse, and there is a clear need for
more high resolution analyses from other localities worldwide, current evidence
suggest that CAMP-related environmental changes and the associated Late Triassic
extinction event are both reflected in the T–J trace-fossil record.
On land, the onset of CAMP-related volcanism is associated with the extinction
of several groups of terrestrial vertebrate, as recorded in the disappearance of their
footprints (Olsen et al. 2002). This warming-related event led to a dramatic increase
in the dominance of dinosaur taxa and an associated increase in footprint, and there-
fore body, size during the Hettangian. While it is possible that the increase in maxi-
mum body size is simply a consequence of increased diversity and variance (Gould
1988), Olsen et al. (2002) suggested that it may be due to ecologic release following
extinction of competing taxa. It is therefore intriguing that during the Hettangian a
similar size trend is recorded in marine infaunal invertebrates, as evidenced by an
increase in burrow diameter (Twitchett and Barras 2004; Barras and Twitchett 2007).
The diameter of marine trace fossils reached a minimum during the peak warming
interval spanning the T–J boundary, probably because of the combined stresses of
elevated temperatures and an expansion in hypoxic or euxinic conditions at that time
(Kidder and Worsley 2010). Although more data are required, it is possible that size
increase in animals, on land and in the sea, is a response to the decline in atmospheric
CO2 back to preextinction levels coupled with a rise in atmospheric oxygen levels.
In the marine realm, the trace-fossil show as presently known does not show
evidence of extinction of any tracemakers, but does indicate that there were signifi-
cant ecologic changes to the marine benthic ecosystem. Apart from a reduction in
burrow diameter, a reduction in diversity and maximum burrow depth is also
recorded during peak global warming following the extinction event. Although this
is an expected consequence of the associated environmental changes, such as
reduced ocean circulation and expanding anoxia, these changes also have implica-
tions for ecosystem functioning. The depth and size of infaunal organisms affects
the amount of bioturbation and a number of key nutrient cycles (Solan et al. 2004).
The lower levels of bioturbation during peak global warming would have reduced
efficient nutrient cycling, with consequences for the entire marine ecosystem.
The T–J interval was a time of major crisis, related to emplacement of a large
igneous province and associated global warming. As a greater number of sedimen-
tary successions across the T–J boundary are studied for their trace fossils, the
impact of these major environmental changes on the marine and terrestrial realms
should become clearer, providing a more complete picture of the degree to which
Late Triassic global warming affected Earth’s biosphere.
8.4 Conclusion
The trace-fossil record across the T–J boundary appears to preserve evidence of
ecological and taxonomic change associated with the Late Triassic extinction event.
There are well-documented changes in trace-fossil size and diversity through the
interval—both in terrestrial and in shallow marine environments. Such findings are
consistent with the current understanding that the Late Triassic extinction event was
triggered by global warming and related effects of the emplacement of CAMP.
However, while there is evidence of (ichno)taxonomic extinction in the terres-

trial realm, studies to date have yet to find evidence of extinction in the marine
trace-fossil record. This might be more a reflection of a lack of study rather than a
lack of evidence, because few trace-fossil studies have been conducted specifically
from the perspective of documenting and understanding the extinction event.
Acknowledgments The constructive comments of Alfred Uchman and Paul Olsen helped to
improve an earlier draft of this text.
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Chapter 9
The Mesozoic Marine Revolution
Luis A. Buatois, Noelia B. Carmona, H. Allen Curran, Renata G. Netto,

M. Gabriela Mángano, and Andreas Wetzel
9.1 Introduction
Modern-marine ecosystems are the result of numerous evolutionary innovations

that took place during the Mesozoic, commonly referred to as the Mesozoic Marine
Revolution (MMR; Vermeij 1977, 1987). This major evolutionary episode was
responsible for the large-scale restructuring of shallow-marine benthic communi-
ties, including increases in the energy budgets of marine ecosystems (Finnegan
et al. 2011) and predation levels, the latter resulting in a number of coevolutionary
developments (Vermeij 1987). In particular, the intensification of grazing and the
diversification of durophagous predators were conducive to increases in prey sturdi-
ness and frequency of shell repair (Vermeij 1987; Kelley and Hansen 2001).
The MMR also signaled the rise to dominance of the Modern Evolutionary Fauna
(MEF). The MEF actually had its origins in the early Paleozoic, but true displacement
of Paleozoic faunas was not complete until after the end-Permian mass extinction
L.A. Buatois (*) • M.G. Mángano

Department of Geological Sciences, University of Saskatchewan,
114 Science Place, Saskatoon, SK, Canada
N.B. Carmona
CONICET-UNRN, Instituto de Investigación en Paleobiología y Geología, Universidad
Nacional de Río Negro, Isidro Lobo y Belgrano, 8332 Roca, Río Negro, Argentina
H.A. Curran
Department of Geosciences, Smith College, Northampton, MA 01063, USA
R.G. Netto
UNISINOS, PPGeo, Av. Unisinos, 950, 93022-000 São Leopoldo, RS, Brazil
A. Wetzel
Geologisch-Paläontologisches Institut, Universität Basel,
Bernoullistrasse 32, CH-4056 Basel, Switzerland

DOI 10.1007/978-94-017-9597-5_9
20 L.A. Buatois et al.
(Sepkoski and Sheehan 1983; Sepkoski and Miller 1985; see Chap. 7). This shift
from the Paleozoic Evolutionary Fauna (PEF) to the MEF involved the transition
from sedentary epifaunal suspension feeders to mobile, energetic (high-metabolism)
infaunal suspension feeders, deposit feeders, and predators (Wagner et al. 2006;
Leighton et al. 2013). These changes included the appearance of ecologic guilds not
represented within the Cambrian Evolutionary Fauna (CEF) and PEF (Thayer 1983;
Bambach 1983). The number of utilized modes of life increased to present levels by
the late Cenozoic, with all 20 Bambachian megaguilds being filled (Sheehan 2001;
Bambach et al. 2007). At this point, marine paleocommunities had a much greater
representation of infaunal organisms and a higher proportion of motile animals than
mid-Paleozoic communities (Bush et al. 2007).
Not surprisingly, the majority of studies published on the MMR have been based
on the body-fossil record. However, the trace-fossil record provides valuable addi-
tional evidence of the profound nature of this episode, particularly with respect to
the degree of infaunalization, complexity of infaunal tiering structures, and preda-
tion intensity (e.g., Bertling 1999; Harper 2003; Bromley 2004; Glaub and Vogel
2004; Wilson 2007; Knaust 2007; Carmona et al. 2008; Buatois and Mángano
2011). In this chapter, the ichnologic record of the MMR is reviewed. The chapter
is divided into three parts: first, the main groups of tracemakers involved in the rise
of the MEF are analyzed; second, the trace-fossil record of post-Paleozoic shallow-
marine environments is reviewed in order to provide insights into the timing of this
evolutionary event; and, third, the ichnologic record is discussed with respect to
how it can illuminate our knowledge of the MMR by assessing aspects, such as its
environmental breadth, evolution of behavior, evolution of the mixed layer, poten-
tial paleogeographic patterns, and secular changes in infaunalization.
9.2 The Cast of Characters
9.2.1 The Main Groups of Burrowers
The MEF is dominated by bivalves, gastropods, echinoids, crustaceans, and marine

vertebrates; other invertebrate groups, including gymnolaemate bryozoans, demo-
sponges, corals and ammonites, were also members of this fauna (Sepkoski 1981).
Of these groups, bivalves, echinoids, and crustaceans were important tracemakers
during the MMR, and identification of their trace fossil “fingerprints” is relatively
straightforward to detect in bioturbation structures, based on neoichnological obser-
vations and functional morphology analysis. Various types of worm-like organisms
should be added to this list, as many are active bioturbators, although establishing a
direct link between a given trace fossil and its producer(s) commonly is tenuous at
best. If necessary, alternative interpretations are offered. In this section, some of the
most important trace fossils produced by these groups of organisms are described
(Table 9.1). Although not all of these ichnotaxa occurred for the first time during the
MMR, most of them are particularly abundant in post-Paleozoic marine deposits.
9
Table 9.1 Common bioturbation ichnogenera in shallow-marine post-Paleozoic deposits (see main text for references). For explanation of modes of sediment
interaction, see Chap. 1
Mode of sediment
Ichnogenera Stratigraphic range Potential producer Ethology Trophic type interaction
Arachnostega Cambrian–Recent Errant polychaetes Fodinichnia Deposit-feeding Biodiffusive
Arcichnus Devonian–Recent Echinoderms (Ophiuroids) Repichnia Deposit-feeding/suspension Biodiffusive
feeding/predation
Arenicolites Cambrian–Recent Worm-like organisms Domichnia Suspension-feeding Gallery biodiffusive
(polychaetes)
Artichnus Eocene–Recent Echinoderms (Holothurians) Domichnia Deposit-feeding/suspension feeding Conveyor
Asteriacites Cambrian–Recent Echinoderms (Asterozoans) Cubichnia Deposit-feeding/suspension Biodiffusive
feeding/predation
Asterosoma Cambrian–Recent Worm-like organisms Fodinichnia Deposit-/detritus-feeding Conveyor
(probably polychaetes)
Balanoglossites Ordovician–Recent Enteropneusts or polychaetes Domichnia Suspension-feeding Gallery biodiffusive
Bichordites Eocene–Recent Echinoderms Pascichnia Deposit-feeding Conveyor
Cardioichnus Cretaceous–Recent Echinoderms Cubichnia Deposit-feeding Biodiffusive
Chondrites Ordovician–Recent Sipunculids, polychaetes Fodinichnia Detritus feeding/chemosymbionts Gallery biodiffusive
Coenobichnus Oligocene–Recent Crustaceans (hermit crabs) Repichnia Scavenging Biodiffusive
Curvolithus Cambrian–Recent Plathelminthes, nemerteans, Repichnia Predation Biodiffusive
gastropods
Cylindrichnus Cambrian–Recent Polychaetes Domichnia Deposit-/detritus-feeding Gallery biodiffusive
Diplocraterion Cambrian–Recent Polychaetes Equilibrichnia Suspension-feeding Gallery biodiffusive
Ereipichnus Cretaceous–Recent Worm-like organisms Pascichnia Deposit-feeding Gallery biodiffusive/
conveyor
Euflabella Cretaceous–Recent Worm-like organisms Fodinichnia Deposit-/detritus-feeding Conveyor
Foersterichnus Cretaceous–Recent Brachyuran crustaceans Repichnia Omnivory Biodiffusive
Gastrochaenolites Ordovician–Recent Bivalves Domichnia Suspension-feeding Gallery biodiffusive
Glyphichnus Jurassic–Recent Crustaceans Domichnia Suspension-feeding Regenerator
21
(continued)
Table 9.1 (continued)
22
Mode of sediment
Ichnogenera Stratigraphic range Potential producer Ethology Trophic type interaction
Gyrochorte Ordovician–Recent Worm-like organisms Fodinichnia Detritus-feeding Conveyor
Gyrolithes Cambrian–Recent Crustaceans Fodinichnia Deposit-feeding/gardening. Gallery biodiffusive
Haentzschelinia Jurassic–Recent Worm-like organisms Fodinichnia Detritus/deposit feeding Conveyor
Helicodromites Devonian–Recent Worm-like organisms Fodinichnia Deposit-feeding Gallery biodiffusive
Hillichnus Cretaceous–Recent Bivalves Fodinichnia Deposit-feeding/chemosymbiosis Biodiffusive/conveyor
Korymbichnus Jurassic–Recent Polychaetes Fodinichnia Deposit-feeding Biodiffusive
Lapispira Jurassic–Recent Crustaceans or worm-like Domichnia/ Deposit feeding/gardening/ Gallery biodiffusive
organisms, probably Fodinichnia omnivory
enteropneusts
Lockeia Cambrian–Recent Bivalves Cubichnia/ Deposit feeding Gallery biodiffusive
Domichnia
Lophoctenium Devonian–Recent Bivalve Fodinichnia Deposit-feeding Gallery biodiffusive
Macanopsis Jurassic–Recent Brachyuran crustaceans Domichnia Omnivory Regenerator
Macaronichnus Permian–Recent Polychaetes (opheliids) Pascichnia Deposit-feeding Conveyor
Maiakarichnus Miocene–Recent Crustaceans Calichnia Suspension-/detritus-feeding Regenerator
Nereites Cambrian–Recent Worm-like organisms, Pascichnia Deposit feeding Conveyor
probably enteropneusts
Ophioichnus Cretaceous–Recent Echinoderms (Ophiuroids) Repichnia Deposit-feeding/suspension Biodiffusive
feeding/predation
Ophiomorpha Triassic–Recent Crustaceans Domichnia Detritus-feeding Regenerator
Palaeophycus Ediacaran–Recent Glycerid and nereid Domichnia Predation/suspension-feeding Gallery biodiffusive
polychaetes
Parmaichnus Pleistocene–Recent Crustaceans Domichnia Suspension-feeding Regenerator
Patagonichnus Miocene–Recent Polychaetes (e.g., maldanids Fodinichnia Deposit-feeding/predation/ Gallery biodiffusive
and capitellids) omnivory
Pentichnus Carboniferous–Recent Echinoderms Domichnia Predation/suspension-feeding Gallery biodiffusive
L.A. Buatois et al.
9
Phoebichnus Jurassic-Pleistocene Worm-like organisms, Fodinichnia Deposit-feeding Gallery biodiffusive

crustaceans
Pholeus Triassic–Recent Crustaceans Domichnia Suspension-/detritus-feeding Regenerator
Phycosiphon Ordovician–Recent Worm-like organisms Fodinichnia Deposit-feeding Conveyor
Planolites Cambrian–Recent Worm-like organisms Fodinichnia Deposit-feeding Conveyor
(probably polychaetes)
Polykladichnus Devonian–Recent Worm-like organisms (probably Domichnia Suspension-feeding Gallery biodiffusive
polychaetes), and anemones
Protovirgularia Ordovician– Recent Bivalves Repichnia Deposit-feeding Biodiffusive
Psilonichnus Late Jurassic onward Crustaceans Domichnia Suspension-feeding Regenerator
Rhizocorallium Cambrian–Recent Worm-like organisms/ Fodinichnia Suspension-feeding/ Gallery biodiffusive
crustaceans deposit-feeding
Rosselia Cambrian–Recent Polychaetes Domichnia Deposit-feeding Gallery biodiffusive
Saronichnus Pleistocene–Recent Bivalves Fodinichnia Chemosymbiosis Gallery biodiffusive
Scalichnus Pliocene–Recent Bivalves Equilibrichnia Suspension-feeding Biodiffusive
Schaubcylindrichnus Carboniferous–Recent Polychaetes Domichnia Deposit-feeding Gallery biodiffusive
Scolicia Jurassic–Recent Echinoderms Pascichnia Deposit-feeding Conveyor
Sinusichnus Cretaceous-Pliocene Crustaceans Fodinichnia Deposit-feeding/farming Regenerator
Siphonichnus Carboniferous– Bivalves Equilibrichnia Suspension-feeding Gallery biodiffusive
Recent
Skolithos Cambrian–Recent Worm-like organisms (e.g., Domichnia Suspension-feeding Gallery biodiffusive
phoronids, polychaetes)
Solemyatuba Ordovician–Recent Bivalves Domichnia Farming Gallery biodiffusive
Spongeliomorpha Triassic–Recent Crustaceans Domichnia Suspension-feeding/farming Regenerator
Taenidium Cambrian–Recent Worm-like organisms Fodinichnia Deposit-feeding Conveyor
Teichichnus Cambrian–Recent Polychaetes, sipunculan Fodinichnia Deposit-feeding Conveyor
worms, arthropods
Thalassinoides Cambrian–Recent Crustaceans Fodinichnia Detritus-feeding Regenerator
Tubotomaculum Cretaceous–Recent Crustaceans Fodinichnia Detritus-feeding/farming Regenerator
23
Zoophycos Cambrian–Recent Sipunculids, echiuran worms Fodinichnia Detritus feeding/food cache/farming Conveyor
9.2.1.1 Crustaceans
Crustaceans of the Order Decapoda (Superclass Crustacea, Class Malacostraca) are

among the most important burrowers in the modern marine realm, constructing dis-
tinctive structures (Fig. 9.1a–g) that commonly extend to considerable depth within
the substratum and occurring in abundance in the fossil record (e.g., Fürsich 1973;
Schlirf 2000; Carmona et al. 2004). Seilacher (2007) described such branching
decapod burrows with the general name “ophiomorphids” to comprise those struc-
tures that, although heterogeneous in taxonomy, share features such as having shaft
and tunnel systems that tend to branch, forming a boxwork at depth, with enlarge-
ments at turning points and having preferential preservation due to their emplace-
ment in deeper tiers than other burrows. Such burrows occur from supratidal to
deep-marine settings. Most “thalassinidean” species (now gebiideans and axiide-
ans, following De Grave et al. 2009) dwell in shallow-marine sediments, and espe-
cially for callianassids and upogebiids, the great majority live in water depths of less
than 20 m at tropical to subtropical latitudes (Dworschak 2000, 2005; Dworschak
et al. 2012). Less abundant in the fossil record are trackways attributed to crusta-
ceans, such as Coenobichnus (Walker et al. 2003) and Foersterichnus (Pirrie et al.
2004). Other relatively common ichnotaxa, most notably Rhizocorallium, may have
been produced by crustaceans (e.g., Rodríguez-Tovar et al. 2012), although worms
cannot be disregarded in many cases (Knaust 2013). The same disagreement per-
sists regarding Phoebichnus, traditionally regarded as produced by worms (Bromley
and Asgaard 1972), but recently attributed to crustaceans (Evans and McIlroy 2016).
Integration of paleoecological data provided by trace fossils with information
inferred from the body-fossil record (e.g., origin, radiation, and extinction patterns)
is important for macroevolutionary analysis of these organisms, which have rela-
tively poor preservation potential (especially for decapods that have weakly calci-
fied exoskeletons). However, due to the presence of hard appendages, some decapods
can excavate burrows in compacted mud, producing open tunnels as permanent
domiciles. In looser sediments, some decapods also reinforce their burrow walls
with characteristic linings. There are a number of distinctive trace fossils produced
by decapods (Figs. 9.2a–k, 9.3a–f, 9.4a–f, 9.5a–e). Forms common in the ichno-
logic record are discussed below, approximately in order of most common
occurrence.
Thalassinoides consists of large burrow systems comprising horizontal and
vertical components, lacking a lining, and with characteristic “Y” or “T” branch-
ing patterns (Frey and Howard 1985; Figs. 9.2b and 9.3a–f). Such burrows usually
are interpreted as dwelling or feeding structures of selective detritus-feeding crus-
taceans (Ekdale 1992), occurring commonly in fine-grained sediments of mar-
ginal- and shallow-marine environments, and also in deeper-marine settings. In a
few instances, Thalassinoides has been reported from continental environments
(e.g., Kim et al. 2002, 2005), but in this case a completely different set of produc-
ers was involved. Thalassinoides is known since the Cambrian (Myrow 1995),
although a decapod origin is most likely only for post-Paleozoic occurrences
(Carmona et al. 2004).
Fig. 9.1 Examples of typical decapod crustaceans and the structures they produce in modern
shallow-marine environments. (a) Intertidal carbonate-sand flat completely re-engineered by cal-
lianassid burrowing, Pigeon Creek lagoon, San Salvador Island, Bahamas; (b) Callichirus major
from the clastic northern Atlantic coast of Florida, USA, shrimp is 15 cm in length; (c) Ghost crab
burrow in a Bahamian beach, opening is 4 cm in diameter; (d) The ghost crab Ocypode quadrata,
on a Bahamian beach, Lee Stocking Island, front of carapace about 5 cm in length; (e) Male and
female fiddler crabs, Uca speciosa, Pigeon Creek margin; (f) Fiddler crab burrows with radiate
pattern of feeding pellets, sandy inlet area, Cockburn Town fossil reef quarry, San Salvador, pen is
15 cm in length; (g) Subtidal callianassid mound, Pigeon Creek lagoon, about 1 m in diameter
Fig. 9.2 Schematic reconstructions of burrow architecture of decapod ichnogenera. (a) Ophiomorpha;
(b) Thalassinoides; (c) Psilonichnus; (d) Macanopsis; (e) Gyrolithes; (f) Spongeliomorpha; (g)
Pholeus; (h) Maiakarichnus; (i) Parmaichnus; (j) Sinusichnus; (k) Glyphichnus. Scale bars in (a, b,
d, f and j) are 5 cm; scale(s) in (c) is 10 cm; in (e) and (h) are 3 cm; in (g) is 2 cm; in (i) is 4 cm and
in (k) is 20 cm
Ophiomorpha refers to simple to complex, branching burrows with distinctive,

thick walls formed of agglutinated-sediment pellets, with the walls mamillated on
the exterior and smooth on the interior (Figs. 9.2a and 9.4a–f). These structures
represent the dwelling burrows of primarily selective detritus-feeding decapods,
such as modern callianassids (Dworschak 2000; Dworschak et al. 2012, and refer-
ences therein). The reinforcement of burrows in order to prevent substratum col-
lapse is an adaptation to living in energetic commonly sandy sedimentary
environments. Ophiomorpha occurs in both siliciclastic and carbonate deposits,
typically attributed to shallow-marine, high-energy environments (Frey et al.
1978; Curran 2007) and also in deep-marine settings (Uchman 2009 and references
9 The Mesozoic Marine Revolution 27
Fig. 9.3 Examples of Thalassinoides in Mesozoic–Cenozoic shallow-marine deposits. (a) Vaca

Muerta Formation, Jurassic, Neuquén Basin, Argentina; (b) Quintuco Formation, Jurassic–
Cretaceous, Neuquén Basin, Argentina; (c) Mulichinco Formation, Cretaceous, Neuquén Basin,
Argentina; (d) Río Turbio Formation, Eocene, Austral Basin, Argentina; (e) Urumaco Formation,
Miocene, Venezuela, scale is 20 cm; (f) Monte León Formation, Miocene, Austral Basin, Argentina,
scale bar is 25 cm; (g) Thalassinoides isp. with tidal fill, Río Negro Formation, Miocene-Pliocene,
Argentina
Fig. 9.4 Examples of Ophiomorpha in Mesozoic–Cenozoic shallow-marine deposits. (a)

Lajas Formation, Jurassic, Neuquén Basin, Argentina; (b) La Vela Formation, Upper Miocene-
Lower Miocene, Venezuela, scale bar is 1 cm; (c) Monte León Formation, Miocene, Argentina;
(d) Chenque Formation, Lower Miocene, San Jorge Basin, Argentina, scale bar is 4 cm; (e)
Chuí Formation, Pleistocene, Brazil, scale bar is 7 cm; (f) Upper Pleistocene, Pehuen-Co,
Argentina
Fig. 9.5 Various decapod ichnotaxa in Mesozoic–Cenozoic shallow-marine deposits. All photos
are from siliciclastic deposits with the exception of b, which is from carbonates. (a) Spongeliomorpha
isp., Middle Miocene, Vallès-Penedès Basin, Spain, scale bar is 10 cm; (b) Psilonichnus upsilon,
Holocene, San Salvador, Bahamas; (c) Gyrolithes isp., Pliocene, Araya Peninsula, Venezuela,
scale bar is 3 cm; (d) Macanopsis plataniformis, lower Cretaceous, La Tejería de Josa, Spain, scale
bar is 3 cm; (e) Sinusichnus sinuosus, Middle Miocene, Socorro Formation, Venezuela, scale bar
is 11 cm; (f) Maiakarichnus isp., Miocene-Pliocene, Río Negro Formation, Argentina
therein). Although Paleozoic examples have been recorded since the Carboniferous
(Driese and Dott 1984; Buatois et al. 2002), uncontroversial occurrences are in post-
Paleozoic strata.
Spongeliomorpha consists of unlined, branching burrow systems with distinctive
scratch marks (bioglyphs) on the walls (Figs. 9.2f and 9.5a). These burrows have been
interpreted as domiciles produced by suspension-feeding decapods (Asgaard et al.
1997) and also as structures for gardening or breeding (D’Alessandro and Bromley
1995). Spongeliomorpha occurs in shallow- and deep-marine environments (Uchman
1998; Muñiz and Mayoral 2001a). There also are some records from non-marine
deposits (Melchor et al. 2009). The presence of bioglyphs on the walls indicates that
the tracemakers used a hard appendage (e.g., a chela) to excavate these burrows and
that the sediment was firm at the time of construction. This ichnogenus is known since
the Triassic (MacNaughton and Zonneveld 2010); specimens reported from the
Permian (Carey 1979) do not display the diagnostic features (e.g., striated walls) of this
ichnogenus.
Psilonichnus comprises predominantly vertical, cylindrical, unlined, J-, Y-, or
U-shaped burrows, with passive fill, commonly interpreted as dwelling structures of
upogebiid shrimp or ocypodid crabs (see Nesbitt and Campbell 2002, 2006; and
Figs. 9.1c–d, 9.2c, and 9.5b herein). These burrows occur most typically in upper-
most foreshore and backshore environments (Frey et al. 1984; Curran and White
1991; Netto and Grangeiro 2009; Seike and Curran 2014), and also in outer-estuarine
and bay-mouth settings (Campbell and Nesbitt 2000; Nesbitt and Campbell 2002).
This ichnogenus is present from the Late Jurassic onward (Fürsich 1981).
Gyrolithes consists of vertical, coiled burrows, forming a regular spiral, unlined
to indistinctly lined, with bioglyphs and/or pellets (Figs. 9.2e and 9.5c) and, in some
Mesozoic and younger examples, connecting to Ophiomorpha or Thalassinoides
burrow forms. Gyrolithes is known from the Cambrian (e.g., Fritz 1980; Fedonkin
1981, 1983; Liñán 1984; Crimes and Anderson 1985; Hein et al. 1991; Jensen 1997;
Jensen and Grant 1998; Stanley and Feldmann 1998) to the Recent (Dworschak and
de Rodrigues 1997). However, worm producers have been indicated for the
Paleozoic occurrences, particularly for the typical Cambrian ichnospecies (see
Carmona et al. 2004; Netto et al. 2007). The robust forms of Gyrolithes from
Mesozoic and younger deposits have been interpreted as resulting from a specific
feeding strategy of decapods having a combined function of deposit feeding and
possibly gardening (Mayoral and Muñiz 1993; Netto et al. 2007). Today, incipi-
ent Gyrolithes-like burrows made by axiidean shrimps are known to occur in
modern mangrove and estuarine areas (Dworschak and de Rodrigues 1997;
Wetzel et al. 2010).
Macanopsis refers to subhorizontal to horizontal, elongated, slightly curved bur-
row chambers, unbranched, and with a subvertical shaft (Fig. 9.2g); bioglyphs may
be present. This ichnogenus has been interpreted as a dwelling structure constructed
by decapods (probably brachyurans) and is known to occur in both shallow-marine
and fluvial settings (Muñiz and Mayoral 2001b) since the Late Jurassic (Neto de
Carvalho et al. 2010).
Pholeus consists of single or complex U-shaped, lined burrows with a longitudinal
axis parallel to bedding, generally leading into an oblique shaft toward the surface
and in the opposite direction with a smaller, rising vertical to oblique shaft (modi-
fied from Knaust 2002; Figs. 9.2d and 9.5d). These structures are interpreted as
dwelling burrows of suspension- and detritus-feeding decapods (“thalassinidean”
shrimps and lobsters), with the function of the small shaft related to ventilation of
the burrow (Knaust 2002). This ichnogenus is typically associated with shallow-
marine to lagoonal (intertidal to shallow subtidal) paleoenvironments and is known
since the Middle Triassic (Knaust 2002).
Tubotomaculum consists of spindle-shaped burrows, displaying a spreite and

containing ellipsoidal pellets (García-Ramos et al. 1984, 2014). This ichnogenus is
interpreted as a feeding structure produced to store pellets that are subsequently
used as a bacteria-enriched resource during times of limited food supply (García-Ramos
et al. 2014). The presence of bifid bioglyphs on the basal and lateral surfaces of
the structure suggests production by crustaceans (García-Ramos et al. 2014).
Tubotomaculum is known from deep-marine environments, ranging in age from the
Cretaceous to Miocene (García-Ramos et al. 1984, 2014).
Sinusichnus comprises distinctive burrow systems consisting of horizontal and
regularly sinuous branches (Gibert 1996; Figs. 9.2j and 9.5e). Shafts and oblique
burrow segments are rare, short, and tend to occur on top of the branching points.
Walls are smooth and unlined, and vertical retrusive spreite are locally present.
There are several interpretations for the function of these structures (e.g., burrows
produced by selective deposit-feeding decapods; open burrows built to act as traps
for meiofauna, and/or to induce microbial growth, i.e., combined feeding strategies
(fodinichnion/agrichnion). These burrows occur in shallow-marine (mainly stressed
settings) and deep-marine environments (as doomed-pioneers), and are known from
the Late Cretaceous (Buatois et al. 2009) to early Pliocene (Belaústegui et al. 2013).
Maiakarichnus consists of subspherical chambers, with or without a clay-rich
lining, preserved in full relief, with numerous shafts radiating in an upward direc-
tion (mainly from the chamber upper hemisphere; Figs. 9.2h and 9.5f). Usually,
this form intergrades or connects to Ophiomorpha or Thalassinoides tunnels.
Maiakarichnus has been interpreted as callianassid brood structures (Curran 1976;
Verde and Martinez 2004) or fossil pantries (Nesbitt 2006). This form occurs in
marginal-marine to shallow-marine deposits and is known since the Miocene
(Verde and Martinez 2004).
Parmaichnus refers to vertical to oblique tubular burrows, composed of a
U-shaped upper part and a basal shaft (Fig. 9.2i). Distinctive swellings or turn-
around chambers are present in the upper part of the burrow and are considered a
diagnostic character for this ichnogenus. Parmaichnus is interpreted as the burrow
of suspension- and filter-feeding decapods, with the presence of turning chambers
being typical for modern upogebiid burrows (Pervesler and Uchman 2009). This
trace fossil occurs in nearshore deposits from early Pleistocene to Recent
(Pervesler and Uchman 2009).
Glyphichnus consists of arcuate, vertical, and probably U-shaped burrows
with deeply incised bioglyphs in fan-like groups, subparallel to burrow length
(Bromley and Goldring 1992; Fig. 9.2k); rarely, apertural necks have been observed
(Goldring et al. 2002). These structures are interpreted as produced by suspension-
feeding decapods. The presence of apertural necks is common in structures used for
suspension feeding and serves to inhibit intruders. This ichnogenus occurs in firm
substrates in both shallow- and deep-marine settings from the late Mesozoic to
Recent (Goldring et al. 2002; Wetzel et al. 2008).
Coenobichnus comprises asymmetrical trackways having left tracks larger than
the right tracks, both being crescent-shaped, with the interior of the crescent
pointing toward the interior of the trackway. This ichnogenus has been attributed to
land hermit crabs (Walker et al. 2003). Coenobichnus occurs in coastal dune settings,
from the Oligocene to Recent (Walker et al. 2003; Zonneveld et al. 2012).
Foersterichnus is a trackway comprising straight, or slightly curving, paired
rows of elongate to tear-shaped impressions, parallel or subparallel to its long axis.
This ichnogenus has been attributed to brachyurans and is only known from
Cretaceous shallow-marine deposits (Pirrie et al. 2004).
As stated previously, burrow systems produced by decapods are among the most
important components of post-Paleozoic shallow-marine ichnofaunas, reflecting the
dominance of the Modern Evolutionary Fauna. In a study documenting the trace-
fossil record of decapod-like gallery systems, Carmona et al. (2004) constructed a
database of all occurrences through the Phanerozoic to analyze changes in abun-
dance and ichnodiversity. During the Paleozoic, there are, in general, a low number
of “decapod-like” burrow systems, and the identity of their tracemakers is uncer-
tain, most likely reflecting behavioral convergence by groups other than decapod
crustaceans. The scenario changed significantly with increase in the number of bur-
rows attributed to decapods during the Mesozoic. This probably reflects an increas-
ing pace of decapod radiation, as it is also indicated by the Mesozoic body-fossil
record (Förster 1985; Schweitzer and Feldmann 2015; Klompmaker et al. 2015).
However, Triassic trace-fossil first occurrences are only slightly higher than
those recorded in the late Paleozoic (Carmona et al. 2004). This is also confirmed
by the body-fossil record, which indicates that decapod diversity was low in the
Triassic (Klompmaker et al. 2013; Schweitzer and Feldmann 2015). In any case,
there were changes in decapod ichnodiversity with Thalassinoides and Ophiomorpha
being the most common ichnotaxa, along with Gyrolithes (resembling modern
records) and Pholeus added to the list. Spongeliomorpha was also well established
in shallow-marine environments. This suggests the overall expansion of behavioral
modes through the Triassic.
During the Jurassic, decapod trace-fossil occurrences show a slight increase with
respect to previous periods (Carmona et al. 2004). This is clearly related to the
appearance of the body fossils of callianassids in the Late Jurassic (Glaessner 1969;
Förster 1985; Schweitzer and Feldmann 2015). Interestingly, the basic ethological
program recorded by these dwelling systems seems not to have changed signifi-
cantly since the Jurassic.
The abundance of decapod trace-fossil forms underwent a major increase toward
the end of the Mesozoic (Carmona et al. 2004). Crustacean dominance in the marine
realm during the Cretaceous is indicated by the great diversity of body fossils, as well
as by the abundance of decapod burrowing activity recorded in shallow-marine deposits.
A decapod origin is supported by the preservation of claws within some of these bur-
rows (e.g., Mángano and Buatois 1991; Swen et al. 2001) and by the presence of
diagnostic features, such as pelleted walls, bioglyphs reflecting the use of hard append-
age parts, and enlargements at burrow turnaround points. Decapod excavations are
particularly abundant in middle- and lower-shoreface clastic environments, as well as
in shallow-marine carbonate settings. The increase in the number of decapod trace-
fossil occurrences accelerated by the end-Cretaceous, including addition of the oldest
examples of Sinusichnus in Cretaceous deposits (Buatois et al. 2009).
Interestingly, Klompmaker et al. (2013) found that during the Mesozoic there
was a long-term shift in diversity of the different crustacean groups based on the
body-fossil record, with dominance of lobsters and shrimps during the Paleozoic
and Triassic, whereas during the Jurassic onward, anomurans and brachyurans
underwent rapid diversification. In particular, their study showed that the Late
Jurassic decapod radiation was associated with the expansion of reef ecosystems,
with a high percentage of generic origination of reef-dwelling brachyurans and ano-
murans (Klompmaker et al. 2013). This is in agreement with the trace-fossil record,
confirming that the Mesozoic was a highly important time in the evolution of deca-
pods (Schweitzer and Feldmann 2015).
During the Paleogene, the numbers of individual decapod trace fossils appear to
be considerably reduced compared with that found in similar environmental
settings in the Cretaceous (Carmona et al. 2004). This could be related to the end-
Cretaceous mass extinction. However, the effect of this extinction on decapods
(mostly of the Southern Hemisphere) was probably not so severe (Feldmann and
Schweitzer 2006). In the same vein, Swen et al. (2001) analyzed the demise of the
subfamily Protocallianassinae and the rise of other subfamilies (among them
Callianassinae) in nearshore deposits, which occurred below the K−Pg boundary.
They postulated that the demise of Protocallianassinae and the rise of Callianassinae
were related to the worldwide emergence of seagrasses at the end of the Cretaceous
and to the strong competition that took place globally in shallow-marine settings.
These authors also thought that seagrasses may have favored the rise of families
dominated by detritus-feeding shrimp (such as Corallianassa) over those of sus-
pension feeders (e.g., Protocallianassa). The abundance of individual decapod
trace fossils during the Eocene is apparently higher than for the other epochs of the
Paleogene. The Eocene was a time of evolutionary radiation for decapods, and this
appears to be reflected in the trace-fossil record.
Finally, during the Neogene, the abundance of decapod trace fossils underwent
another major increase, probably reaching the highest level of the entire Phanerozoic
(Carmona et al. 2004). Body-fossil data conform to the Neogene trace-fossil record,
which shows primacy of decapod burrows in shallow-marine environments, with
Maiakarichnus occurring for the first time in the Miocene. Interestingly, the sum-
mary study of Schweitzer (2001) indicated that the Miocene was a time of high
origination rates within the decapod fauna, with first appearance of the modern
genera that dominate the present seas.
Decapod burrows are widespread in modern coastal settings, being preferentially
distributed in tropical and temperate shallow-marine and marginal-marine settings.
The similarity between the modern callianassid burrows formed by Sergio mirim in
the lower foreshore of Cassino Beach (Rio Grande do Sul State, southernmost
Brazil) and Ophiomorpha nodosa specimens preserved in Pleistocene deposits of
the Chuí Formation (Gibert et al. 2006), which represent an ancient beach line on
the Rio Grande do Sul Coastal Plain, suggests that these faunas were not impacted
by Quaternary glaciation, at least in the Southern Hemisphere. The presence of
large Ophiomorpha specimens in Pleistocene shallow-marine deposits from
Pehuen-Có, Argentina, further supports this interpretation (Mouzo et al. 1989).
9.2.1.2 Mollusks
Mollusks have a long evolutionary history and include several groups, with gastro-
pods and bivalves being the most numerous, diverse, and common bioturbators
from the early Paleozoic onward. The development of key external features, such as
the shell, foot, and radula, helped mollusks acquire diverse ecological preferences.
This section focuses on trace fossils attributed to gastropods and bivalves.
Gastropods and Polyplacophorans
Locomotion and feeding structures produced by gastropods and chitons can be

observed in modern settings. Almost all gastropods and chitons use their foot during
locomotion. Typically, the sole of the foot is flat and broad, ciliated and with abun-
dant secretory glands, allowing gastropods to move over a variety of substrates
(Ruppert et al. 2004). These glands produce mucus that forms a mucus trail. Most
gastropods living on stable substrates move by pedal waves with the aid of the
mucus trail (glide crawling, Schäfer 1962, 1972; see also Buatois and Mángano
2011). Gastropods are mostly microphagous browsers that feed on microscopic
algae and scrape the substrate with their radula (a flexible longitudinal ribbon hav-
ing rows of small and transverse chitinous teeth; Ruppert et al. 2004). The com-
bined actions of the foot and radula generate distinctive patterns on the substrate,
which have also been recognized in the fossil record, although some uncertainty
persists regarding the attribution of these structures to gastropods. For example, the
Cambrian ichnogenus Climactichnites is interpreted as having been produced by an
elongate, dorso-ventrally flattened organism with a soft foot, which may have been
a primitive gastropod, although other possibilities exist (Abel 1935; Getty and
Hagadorn 2008, 2009; see Chap. 3). In modern intertidal environments, structures
slightly similar to this trace fossil can be found, although their preservation poten-
tial is very low (except in settings with microbial mats; Carmona et al. 2010).
Other trace fossils (e.g., Psammichnites) have been interpreted as locomotion
structures of gastropods, but other producers cannot be discarded (see discussions in
Mángano et al. 2002a). In any case, neither Climactichnites nor Psammichnites are
recorded in post-Paleozoic rocks, probably due to increased levels of bioturbation and
obliteration of shallow-tier traces or extinction of their producers. The much simpler
trail Archaeonassa, ranging from the Cambrian to the Recent, has been attributed to
gastropods as well (Fenton and Fenton 1937), but there is no agreement in this regard
(Buckman 1994; Yochelson and Fedonkin 1997). Therefore, the fossil record of bio-
turbation structures produced by gastropods remains uncertain.
Bivalves
Along with gastropods, bivalves constitute an important and extremely diverse mol-
luscan class, with species living in both marine and freshwater environments. Within
this group, a pronounced post-Paleozoic increase in infaunal representatives, also
exhibited by several other major groups (e.g., echinoids), is widely recognized

(Stanley 1970). The evolutionary diversification of most bivalve shells is related to
different styles of burrowing in soft sediment and of boring in hard substrates, which
provide escape from predators (Stanley 1970; Seilacher 1998). This tendency is not
only reflected by the hard part anatomy, but also by the soft parts of bivalves, which
show corresponding trends in the development of a lateral flat burrowing foot, fused
mantle edges, and pallial siphons (Stanley 1970; Seilacher 1998; Kelley and Hansen
2003; Ruppert et al. 2004 and references therein). The replacement of epifaunal and
semi-infaunal taxa by infaunal forms from the Cretaceous onward is well recog-
nized (Vermeij 1977, 1987).
In contrast to crustacean ichnotaxa, which for the most part are represented by
dwelling structures, the ichnogenera most commonly assigned to bivalve tracemak-
ers comprise various ethologies (Fig. 9.6a–i), such as locomotion (Protovirgularia,
Ptychoplasma, Oravaichnium), resting/dwelling (Lockeia), dwelling (Solemyatuba),
feeding (Lophoctenium, Saronichnus), equilibrium (Siphonichnus, Scalichnus), and
a complex combination of behaviors (Hillichnus), all in soft sediments (Figs. 9.7a–d
and 9.8a–h). The dwelling ichnogenus Gastrochaenolites may occur in both hard
and firm substrates (Fig. 9.8h) and will be addressed under bioerosion.
Protovirgularia comprises horizontal to inclined structures characterized by the
presence of V-shaped or U-shaped sediment pads transverse to a longitudinal axis
(Figs. 9.6a and 9.7a–d). This ichnogenus has been interpreted as formed by locomo-
tion of protobranch bivalves, moving through the sediment by means of a split foot
that employs a push-and-pull mechanism (Trueman 1966, 1975; Stanley 1970;
Seilacher and Seilacher 1994; Mángano et al. 1998). In addition, Seilacher and
Seilacher (1994) suggested that Protovirgularia can also be produced by scaph-
opods (another molluscan group with a cleft-foot), especially if the structures are
not related to the resting trace Lockeia (Mángano et al. 1998). This trace fossil,
recorded in marginal-, shallow-, and deep-marine environments, ranges from
Ordovician to Recent (Seilacher and Seilacher 1994; Uchman 1998).
Ptychoplasma consists of irregularly meandering, looping, winding or straight,
discontinuous or continuous, hypichnial crests having an amygdaloid, carinate, or
blocky cross-section, with poorly developed or absent chevronate structure. This
ichnogenus has been interpreted as a locomotion trace of wedge-foot bivalves
(Pieńkowski and Uchman 2009; Uchman et al. 2011). Recorded from non-marine,
marginal-, shallow-, and deep-marine environments, this ichnogenus ranges from
Ordovician to Recent (Rodríguez-Tovar et al. 2014).
Oravaichnium consists of irregularly meandering, looping or winding, continu-
ous, hypichnial crests having a box-like (non-carinate) cross-section. As with
Ptychoplasma, this ichnogenus is regarded as a locomotion trace of wedge-foot
bivalves (Uchman et al. 2011). In contrast to Ptychoplasma, however, Oravaichnium
is less understood, having been recorded only from Eocene deep-marine deposits
(Plička and Uhrová 1990; Uchman et al. 2011).
Lockeia commonly comprises oval- to almond-shaped structures, typically tapering
at one end and being more rounded on the opposite end (Mángano et al. 1998;
Figs. 9.6a and 9.8a). Lockeia is interpreted as a resting structure of bivalves, although
Fig. 9.6 Schematic reconstructions of bivalve-produced ichnogenera. (a) Reconstruction of the

locomotion activity of a cleft-foot bivalve, producing Protovirgularia and Lockeia (modified from
Ekdale and Bromley 2001a), scale bar is 5 cm; (b) Reconstruction of Ptychoplasma (redrawn from
Uchman et al. 2011), scale bar is 1 cm; (c) Interpretation of Siphonichnus (modified from
Stanistreet et al. 1980), scale bar is 5 cm; (d) Schematic representation of the pholadids during
excavation and production of Gastrochaenolites (see Carmona et al. 2007), scale bar is 2 cm; (e)
Interpretation of the complex behavior of a deposit-feeding bivalve illustrated by Hillichnus
(modified from Bromley et al. 2003), scale bar is 5 cm; (f) Reconstruction of Scalichnus (modi-
fied from Hanken et al. 2001), scale bar is 5 cm; (g) Reconstruction of Solemyatuba (based on
Seilacher 1990a), scale bar is 5 cm; (h) Reconstruction of Saronichnus (modified from Pervesler
and Zuschin 2004), scale bar is 5 cm; (i) Reconstruction of Teredolites (based on Bromley et al.
1984), scale bar is 5 cm
Fig. 9.7 Preservational variants of Protovirgularia in Mesozoic–Cenozoic shallow-marine depos-

its. (a) Forest Marble Formation, Jurassic, England, coin is 1.65 cm; (b–d) Chenque Formation,
Lower Miocene, Argentina. Coins in (b) and (c) are 2.2 cm
some specimens can represent semipermanent domiciles (Seilacher 1953; Rindsberg

1994; Mángano et al. 1998). This ichnogenus has been reported from marginal-,
shallow-, and deep-marine settings since the late Cambrian/Early Ordovician
(Mángano et al. 2002b and references therein). In contrast to most bivalve ichno-
taxa, Lockeia is also known from freshwater settings.
Solemyatuba consists of endichnial, vertically oriented, U-shaped burrows, ellip-
tical in cross-section and with a lower extension tube in one of its ichnospecies
(Seilacher 1990a; Mángano and Buatois 2003; Figs. 9.6g and 9.8d). Solemyatuba
has been interpreted as the dwelling trace of a farming bivalve (Seilacher 1990a).
This form can be found in shallow-marine deposits since the Ordovician (Seilacher
1990a; Mángano and Buatois 2003).
Lophoctenium is not generally assigned to the activity of bivalves. However,
Ekdale and Bromley (2001a) interpreted Carboniferous specimens of this trace fos-
sil as a feeding structure produced by the repetitive lateral probing of the labial
palps of a protobranch along an organic-rich sediment deposit. This structure con-
sists of coarse horizontal spreite formed by a series of closely spaced, curved ridges
(Ekdale and Bromley 2001a). In particular, the specimen studied by these authors
represents a compound trace fossil with three different ichnotaxa in combination
(Protovirgularia, Lockeia, and “Lophoctenium”; see Ekdale and Bromley 2001a).
Saronichnus consists of a system of vertical or inclined tubular to blade-shaped
probes diverging from a central area of broom- or star-like form (Pervesler and Zuschin
2004; Figs. 9.6h and 9.8f). This ichnotaxon has been found in direct connection
with Thyasira shells, indicating formation by lucinoid bivalves. It is most likely a
Fig. 9.8 Various bivalve ichnotaxa in Mesozoic–Cenozoic shallow-marine deposits. (a) Several
specimens of Lockeia siliquaria, Mulichinco Formation, Cretaceous, Argentina; (b) Scalichnus
phiale, Monte León Formation, Lower Miocene, Argentina, scale bar is 5 cm; (c) Siphonichnus
eccacensis, Chenque Formation, Lower Miocene, Argentina, coin is 1.8 cm; (d) Solemyatuba ypsi-
lon, Rhaetian Sandstone, Olgahain, Upper Triassic, southern Germany, scale bar is 1 cm; (e)
Equilibrium traces of byssate bivalves, scale bar is 4 cm; (f) Saronichnus abeli, Grund Formation,
Miocene, Austria, photograph courtesy of Peter Pervesler; (g) Hillichnus lobosensis, Carmelo
Formation, Paleocene, Point Lobos, California, United States, photograph courtesy of Nic Minter;
(h) Gastrochaenolites isp., Río Turbio Formation, Eocene, Argentina
feeding structure produced as a result of a chemosymbiotic life strategy. At present,

it has only been recorded from Pleistocene shallow-marine strata (Pervesler and
Zuschin 2004).
Siphonichnus comprises vertical structures containing a backfill of concave-
downward menisci (Stanistreet et al. 1980; Figs. 9.6c and 9.8c). The laminae form-
ing the backfill are cut through centrally by a vertical tube, filled with pale, massive
sediment. The authors of this ichnogenus postulated that siphon length of a given
bivalve tracemaker should be equal to the length of backfill laminae. Siphonichnus
occurs since the Carboniferous in marginal- to shallow-marine settings (Stanistreet
et al. 1980; Carmona et al. 2008; see also discussion in Zonneveld and Gingras 2013
and Knaust 2015).
Scalichnus includes large, vertically oriented, bottle-shaped structures, formed
during retrusive and protrusive movements of the bivalve Panopea (Hanken et al.
2001; Figs. 9.6f and 9.8b). Scalichnus is regarded as an equilibrichnial trace fossil
having a general sack-like morphology and a thick lining. This structure is common
in Cenozoic marginal- to shallow-marine settings (Hanken et al. 2001; Nara and
Kondo 2012; see also discussion in Zonneveld and Gingras 2013).
Hillichnus was defined to include morphologically complex structures inter-
preted as the work of subsurface deposit-feeding tellinacean bivalves (Bromley
et al. 2003; Figs. 9.6e and 9.8g). This structure comprises several parts or levels: a
segmented component with a basal tube within, from which lateral feather-like
structures arise, alternating on either side, and vertical sand- and mud-lined tubes
that also rise from the basal structure. According to the ichnogenus authors, the
movement of the bivalve’s inhalant siphon produced the feather-like structures on
either side of the basal tube, and the siphonal excursions to the sediment surface
produced the vertical tubes. The length of these tubes suggests that feeding activity
of these bivalves occurred well below the oxygenated sediment layer, probably
20 cm or more beneath the surface, suggesting that in addition to deposit feeding,
chemosymbiosis may have taken place. Hillichnus is known from the Cretaceous
and Paleogene, in both shallow- and deep-marine environments (Bromley et al.
2003; Pazos and Fernández 2010).
Finally, it is possible that other ichnotaxa may have been produced by bivalves,
but tracemaker origins cannot always be confirmed. An example would be
Paradictyodora, which comprises vertical spreite structures that consist of subverti-
cal folded laminae produced by the migration of a subvertical J-shaped tube (Olivero
et al. 2004). This structure has been interpreted as the feeding trace of worms or
tellinid bivalves (D’Alessandro and Fürsich 2005; Serpagli et al. 2008). This ichno-
genus occurs from Late Cretaceous to Pleistocene (Olivero et al. 2004; D’Alessandro
and Fürsich 2005).
Although bivalves were key players during the MMR, the trace-fossil record
shows that some of the most representative ichnotaxa, such as Protovirgularia and
Lockeia, have been present since the early Paleozoic, albeit becoming particularly
abundant since the late Paleozoic. However, other bivalve ichnotaxa (e.g.,
Hillichnus) seem to be directly associated with the MMR, whereas the stratigraphic
ranges of other ichnotaxa (e.g., Scalichnus, Saronichnus) are still poorly known
due to sparse occurrence. Hillichnus reflects a level of behavioral complexity

unknown in older bivalve-produced ichnotaxa (Bromley et al. 2003). Particularly,
the proposed producers of this ichnogenus, the tellinid bivalves, originated in inner
shelf environments during the Middle Triassic and then commenced significant
radiation (Jablonski and Bottjer 1990). Interestingly, it seems that the earliest tel-
linaceans were suspension feeders, whereas deposit-feeding tellinids appeared
during the Early Cretaceous (Bottjer and Jablonski 1988). Although lucinids origi-
nated in the Silurian, they experienced an evolutionary radiation near the end of the
Cretaceous, coincident with the appearance of seagrasses and mangroves (Stanley
2014). It has been hypothesized that lucinids may have exploited the dysaerobic
sediments below roots and rhizomes, acquiring sulfides for their endosymbiotic
bacteria (Stanley 2014). Saronichnus, the only ichnogenus attributed confidently to
lucinids, is only known from its type locality, so using ichnologic data to track
evolutionary trends is still not possible.
9.2.1.3 Echinoderms
The Phylum Echinodermata is one of the most important and characteristic com-
ponents of modern-marine benthic communities (Smith 1984). Echinoderms com-
prise very diverse marine groups, such as sea stars, brittle stars, sea urchins, sand
dollars, sea cucumbers, and sea lilies, with the majority being bottom dwellers
(Ruppert et al. 2004).
The Class Echinoidea consists of diverse and successful groups comprising
approximately 950 living species (Smith 1984; Ruppert et al. 2004). Echinoids
adapted to live in a variety of marine environments and present different life habits
(Smith 1984). This part of the chapter focuses mostly on irregular echinoids, whose
stratigraphic range covers the Early Jurassic to Holocene, with approximately 410
genera (Parma 2008).
Irregular echinoids are detritus- or deposit-feeding organisms that ingest material
selected by their podia. This particulate material is covered by mucus and then moved
to the mouth. Some irregular echinoid species burrow deeply into the substrate and
build a vertical channel to provide oxygenated water from the sediment surface. They
also construct sanitary channels or drains to remove fecal material mixed with sea-
water. Within irregular echinoids, the spatangoid heart urchins constitute the most
diverse group in modern oceans, being found in all latitudes and at all depths, and
contributing significantly to collective bioturbation of the surrounding sediments
(Villier and Navarro 2004). In addition, the infaunal activities of these organisms
stimulate bacterial production within the substrate via transport of organic matter to
deeper sediment layers (Osinga et al. 1997). In turn, deep-burrowing irregular
echinoids can induce a back-flow of organic matter in the form of dissolved nutrients
returned to the sea (Lohrer et al. 2004).
The abundance of spatangoids increased rapidly during the Cretaceous and has
remained stable since then, although other groups of irregular echinoids show a
concurrent decline in diversity. This has been interpreted as due to the acquisition of
Fig. 9.9 Schematic reconstructions of trace fossils produced by irregular echinoids. (a)
Reconstruction of the locomotion activity of Echinocardium cordatum producing Scolicia, scale
is 1 cm; (b) Cross-sectional view of Scolicia; (c) Cross section view of Bichordites; (d) Basal
view of Cardioichnus (based on Smith and Crimes 1983), scale is 1 cm. (a–c) modified from
Bromley (1996)
a burrowing life mode, which allowed spatangoids to access new food resources and
avoid predation (Smith 1984; Villier and Navarro 2004). Three ichnogenera
(e.g., Bichordites, Scolicia, and Cardioichnus) are attributed to the activity of irreg-
ular echinoids (Figs. 9.9a–d and 9.10a–e; see Belaústegui et al. 2015a for a review).
Scolicia comprises horizontal to inclined trace fossils characterized by the
presence of menisci (Figs. 9.9a–b and 9.10a–b). In cross-section and in hypichnial
preservation, two parallel canals, functioning as drains for used respiratory water,
are observed at the base of these structures. This ichnogenus ranges from the
Jurassic onward (Uchman 1995).
Bichordites also consists of meniscate, horizontal structures, but with only one
middle channel crossing the menisci (Figs. 9.9c and 9.10c–d). Bichordites ranges
from the Eocene onward (Bernardi et al. 2010; Demírcan and Uchman 2012;
Villegas-Martín et al. 2014).
Cardioichnus consists of heart-shaped structures having an axial depression
(Figs. 9.9d and 9.10e). It is interpreted as the resting trace of irregular echinoids
(Smith and Crimes 1983) and usually is found in association with locomotion traces.
This ichnotaxon ranges from the Late Cretaceous onward (Smith and Crimes 1983).
In modern environments, two principal groups of irregular echinoids are known
to excavate sediment: the Spatangus group, which produces meniscate structures
with two basal drain canals (modern analogues for Scolicia), and the Echinocardium
group, which produces structures with only one canal, resembling the ichnogenus
Bichordites (Bromley and Asgaard 1975; Kanazawa 1995). Although Smith and
Crimes (1983) postulated that irregular echinoids fully developed their capacity to
excavate during the Early Cretaceous, when the Spatangus group first appeared,
ichnologic evidence (i.e., the presence of Scolicia in Jurassic strata) suggests an
earlier origin.
During the Paleozoic, epifaunal echinoderm taxa (mostly suspension- and detritus-
feeders) were abundant (Dornbos 2008; Schneider 2008). Several reasons have been
Fig. 9.10 Trace fossils produced by irregular echinoids. (a) Scolicia isp., López de Bertodano
Formation, Upper Cretaceous, Antarctica; (b) Scolicia isp. with their producers, Chenque
Formation, Lower Miocene, Argentina; (c–d) Bichordites kuzunensis, Mezardere Formation, Early
Oligocene, Turkey, scale bar is 1 cm; (e) Cardioichnus isp., Upper Marine Molasse, Lower
Miocene, Switzerland; (f) Asteriacites lumbricalis, Neill Klinter Formation, Lower Jurassic,
Greenland
proposed for this abundance: high productivity of the seas during the early Paleozoic,
low abundance of deep-infaunal organisms, and the absence of efficient predators
(Sprinkle and Kier 1987; Parma 2008). This situation changed dramatically after the
Paleozoic, when echinoderms developed infaunal detritus-feeding and predation strat-
egies (Sprinkle and Kier 1987). Obviously, this change is also reflected in the trace-
fossil record. Interestingly, Smith (2005) analyzed the growth and form of Paleozoic
and post-Paleozoic echinoids and recognized that there was a distinct variation in the
nature of disparity between these two groups, with an important diversification of
shapes in those from post-Paleozoic times (see also Hopkins and Smith 2015). In
particular, he found that there was a significant evolutionary innovation for echinoids
with regard to specification of the final positions of plates during early stages of
ontogeny. This early fixation of plate position favored the appearance of other
evolutionary innovations, especially in the Atelostomata (e.g., spatangoids) and
Clypeasteroida clades (Hopkins and Smith 2015). For example, the early fixation of
plates allowed diverse regions of the test to differentiate both in structure and function.
This involved also the morphological and functional differentiation of spines, espe-
cially for the spatangoids during the Cretaceous and Cenozoic, which poses important
implications for acquisition of the ability to burrow. The early fixation of plate posi-
tion also allowed for development of bands of spines called fascioles, which serve to
create water currents and mucus streams. Fascioles are particularly important for bur-
rowing in spatangoids, and their appearance permitted this group to diversify ecologi-
cally (Smith 2005). In addition to these innovations, the presence of an intestinal
caecum enclosing sulfide-oxidizing bacteria in some spatangoid species allows
them to burrow deeply within the substrate, well below the redox boundary, to ingest
sulfide-rich and anoxic sediment (Bromley et al. 1995). Therefore, echinoids are able
to avoid predation and reduce competition both for food and space, occupying a deep
tier within the substrate (Bromley et al. 1995).
There are other trace fossils interpreted as produced by echinoderms. Among
them, the most common is the resting trace Asteriacites (Fig. 9.10f–h), interpreted
as produced by asterozoans (both asteroids and ophiuroids, but more commonly
the latter; Seilacher 1953; Mángano et al. 1999). Although it has been suggested
that Asteriacites is a nomen dubium (Schlirf 2012), the rediscovery of the type
material justifies keeping the original name (Knaust & Neumann 2016; see also
Gurav et al. 2014). Asteriacites is present in marginal- to shallow-marine settings
since the Cambrian (Mikuláš 1992a), being particularly abundant in the Triassic
and Jurassic (e.g., Dam 1990a; Wilson and Rigby 2000). There are also records
from deep-marine facies (Crimes and Crossley 1991).
Some other ichnotaxa are also attributed to asterozoans, namely, Pentichnus,
Arcichnus, and Ophioichnus. Pentichnus is a plug-shaped structure with pentameral
symmetry interpreted as a dwelling burrow of ophiuroids. This ichnotaxon has been
described from the Carboniferous (Maerz et al. 1976; Seilacher 1983, 1990b;
Rindsberg 1994; Mángano et al. 2002b). Arcichnus is interpreted as the trackway of
ophiuroids, and consists of a series of horseshoe-shaped impressions produced by
the anterior arms. This trace fossil was described from the Devonian (Sutcliffe
1997). Ophioichnus consists of imprints interpreted as produced by the arms of
ophiuroids and has been recorded in the Early Cretaceous (Bell 2004). Recently,
incipient Ophioichnus were recognized on the modern sea floor (Schatz et al. 2013).
Synonymization of Ophioichnus with Biformites has been recently suggested
(Knaust and Neumann 2016).
The ichnologic record of holothurids is restricted to the ichnogenus Artichnus,
which consists of a wide, J-shaped structure with a narrow, upward tapering shaft and
the distal end tapering to a blind termination (Zhang et al. 2008). An irregular spreite
structure surrounds the burrow lumen. This ichnogenus is known since the Eocene
(Zhang et al. 2008; Ayranci and Dashtgard 2013; Ayranci et al. 2014; Belaústegui and
Domènech 2014). A comparison with modern structures suggests that potential holo-
thurid trace fossils may occur in Carboniferous rocks (Smilek and Hembree 2012). In
fact, burrows attributed to holothurians have been recorded in the early Permian (Netto
1994; 2000). Further work is needed to assess the relationship between Artichnus and
the previously defined ichnogenus Naviculichnium (Książkiewicz 1977) in order to
evaluate more completely the record of trace fossils generated by holothurians.
In sum, the trace-fossil record matches closely body-fossil information, as indi-
cated by the stratigraphic ranges of ichnotaxa attributed to irregular echinoids, such as
Scolicia, Bichordites, and Cardioichnus, which are typical structures of the Modern
Evolutionary Fauna. In contrast, structures commonly attributed to ophiuroids have a
much longer stratigraphic range (Asteriacites, Arcichnus) or are still poorly known
(Ophioichnus). Further occurrences of structures produced by holothurids in the fossil
record are needed in order to establish potential trends.
9.2.1.4 Worms
Many ichnogenera have been attributed to the activity of “worms,” without clear
distinction regarding which group of worms might have been responsible. In fact,
worm-like animals constitute a particularly important group of bioturbators in
marine environments, including principally vermiform nemerteans, polychaetes,
and hemichordates, all groups that are diverse and abundant. According to Seilacher
(2007), when worms are mentioned, the term serves to designate “all long, soft and
usually cylindrical animals belonging to many different phyla,” which have evolved
a peristaltic mode of locomotion for life in soft substrates, but other locomotion
strategies may be employed, such as undulation and bolting (Schäfer 1972; Buatois
and Mángano 2011). The objective of this section is to discuss the most common
trace fossils attributed to worm-like organisms, while relating them to the evolution-
ary history of some of their likely producers wherever possible.
Polychaetes are annelids having a pair of parapodia and associated chaetae in
each body segment. Most polychaetes (e.g., terebellids, maldanids, glycerids, oph-
elids) are burrowers, although some bore into shells and rocks, secrete tubes, move
over surfaces, and even swim (Ruppert et al. 2004). Nutrition habits of polychaetes
are closely related to their mode of life; deposit feeders are mostly burrowers and
sedentary worms living in burrows/tubes; suspension feeders are sedentary organ-
isms living in tubes in the sediment or are attached to organic or inorganic hard
substrates; and scavengers, carnivores, and herbivores are typically errant worms
(although some live in tubes or are active burrowers) (Ruppert et al. 2004). Groups
with ichnologic importance include members of the clades Scolecida and Palpata
(Gingras et al. 2008). Scolecids are deposit-feeding burrowers, similar to the pre-
sumed ancestor of annelids, without head appendages, and palpates have head
appendages and more diversified life habits (Ruppert et al. 2004). In particular,
ophelids (scolecids) include burrowers with an eversible bulbous pharynx with which
they ingest sediment in high-energy areas such as the foreshore, producing
Macaronichnus-like structures (Clifton and Thompson 1978; Ruppert et al. 2004).
Nemerteans include several species of long, slender worms resembling annelids,
although lacking their segmentation. They have a characteristic proboscis apparatus
with which they capture prey and burrow (Ruppert et al. 2004). Species that burrow
(e.g., Carinoma) commonly have a muscular body wall that they use to produce
peristaltic movement (Ruppert et al. 2004).
Echiurans (or spoon worms) comprise coelomate marine organisms that lack
segmentation. Echiurans have two main body regions: an anterior cylindrical part,
called the prostomium or proboscis, and a trunk, which commonly has three-layered
musculature used to produce peristaltic movements to ventilate their burrows. Most
echiurans are deposit-feeders, although some are suspension-feeders. Many species
live in U-shaped burrows built in sand and mud in shallow-marine settings, although
some occupy deep-marine sediments (Ruppert et al. 2004).
Sipunculids are a group of worms sharing some characters with echiurans
(e.g., both lack segmentation). These organisms are suspension- or deposit-feeders,
collecting their food with ciliated tentacles present in the oral disc. Sipunculids are
bottom-dwelling organisms, with most living in shallow-marine settings, where
they burrow into the sediment or bore into coralline rocks or wood (Ruppert et al.
2004). Although the fossil record of this group is sparse, Huang et al. (2004)
reported three species from the early Cambrian.
Priapulids are benthic marine worms characterized by an extendable, eversible
proboscis (i.e., introvert). These organisms excavate using a push-and-pull mecha-
nism, with the trunk acting as a penetration anchor and the introvert as a terminal
anchor. They live in sandy and muddy sediments in shallow- and deep-marine set-
tings (Ruppert et al. 2004). Based on their fossil record, priapulids were important
components of Cambrian marine communities (Huang et al. 2007), and it has been
convincingly argued that priapulids may have been the producers of Treptichnus
pedum (Vannier et al. 2010).
Enteropneusts are benthic worms that have a body divided into three regions:
proboscis (protosome), collar (mesosome), and trunk (metasome), and thus are tri-
coelomate animals. Generally, they use their anterior region to burrow by retrograde
peristaltic contractions, with most being deposit-feeders (Ruppert et al. 2004). They
mostly live in shallow-marine environments, and their fossil record extends back to
the Cambrian (Caron et al. 2013).
Trace fossils attributed to worms are diverse and comprise structures that are mor-
phologically simple, such as vertical or U-shaped burrows, to those that are more com-
plex, displaying sophisticated spreiten (Fig. 9.11a-i). In this section, a brief description
of most of the ichnogenera attributed to “worms” is alphabetically presented.
Ancorichnus comprises a cylindrical meniscate burrow fill surrounded by a
structureless tubular mantle (Heinberg 1974; Dam 1990a). This ichnogenus has
been interpreted as a locomotion trace (repichnion) of a soft-bodied organism
(Heinberg 1974), most likely a worm. Dam (1990a) noted transitions with
Jamesonichnites, suggesting that Ancorichnus represents the axial burrow from
which Jamesonichnites probes were made. Ancorichnus occurs in shallow-marine
environments and ranges from the Jurassic to the Cretaceous (Heinberg 1974); older
and younger records require critical re-evaluation (e.g., Singh et al. 2010).
Arachnostega consists of irregular, elongate, and net-like burrows in sediment
fills of shells and visible on the surfaces of internal molds. This ichnotaxon is attrib-
uted to errant polychaetes, but crustaceans cannot be ruled out as producers. Bertling
(1992) considered the diameter and course of the burrows to reflect an opportunistic
polychaete larval infestation in recently sediment-filled bivalve borings.

Arachnostega is known since the Cambrian (Rodrigues et al. 2005), although its
presence is more common from the Jurassic onward in shallow-marine settings.
Arenicolites is a vertical U-tube without spreite. It is interpreted as a dwelling
structure. Although it has been attributed to worm-like organisms such as poly-
chaetes (Goldring 1962; Fürsich 1974); crustaceans (Goldring 1962) and holothuri-
ans (Seilacher 1990a; Bromley 1996) have been suggested as well. Arenicolites is
present in a broad variety of shallow- to deep-marine and even continental environ-
ments, being most common in nearshore high-energy settings since the Cambrian
(Mángano and Buatois 2014).
Asterosoma consists of elongate, bulbous segments having a terminal or eccen-
trical, laminated fill; segments are oriented (sub)horizontally and are arranged
around a central vertical, somewhat twisted axis (Schlirf 2000; Fig. 9.11a). This
ichnotaxon is a feeding trace produced by deposit or detritus feeders. Its intergrada-
tion with Rosselia suggests terebellid polychaetes as possible producers, but some
authors (e.g., Neto de Carvalho and Rodrigues 2007) have proposed crustaceans as
well. Asterosoma occurs most commonly in shallow-marine settings since the
Cambrian (Mángano and Buatois 2014).
Balanoglossites consists of connected, U-shaped burrows that occur in shallow-
marine environments, especially intertidal settings and carbonate ramps (Knaust 1998
and references therein; Fig. 9.11b). This trace fossil is interpreted as the domicile of
enteropneusts or polychaetes (Mägdefrau 1932; Kaźmierczak and Pszczōłkowski
1969). Although this ichnogenus occurs since the Ordovician (Knaust and Dronov
2013), it mostly has been recorded since the Triassic (Knaust 2004, 2010; Carmona
et al. 2008; Desai and Saklani 2012; Knaust and Costamagna 2012).
Bornichnus comprises clusters of small, lined, contorted, branched tubes
(Bromley and Uchman 2003). These authors interpreted Bornichnus as an open bur-
row probably produced by farming organisms and noted that similar systems are
produced by capitellid polychaetes in modern deposits. Bornichnus has only been
reported from its type locality in the Lower to Middle Jurassic of Denmark (Bromley
and Uchman 2003).
Chondrites comprise a tree-like system of tunnels that branch downward, with
the angle of branching commonly less than 45° (Fig. 9.11c). The color of the sedi-
ment fills is invariably different from the host-rock color. This ichnotaxon has been
interpreted as the burrow of infaunal detritus feeders or chemosymbiotic organisms
(Fu 1991). Although worms, such as sipunculids and polychaetes, produce similar
structures today, bivalves cannot be completely ruled out (Kotake 1991 and refer-
ences therein). This ichnogenus is known since the Ordovician, and it is a facies-
crossing form, from offshore to abyssal environments (Mángano et al. 2002b and
references therein).
Cylindrichnus consists of broad U- or bow-shaped, concentrically laminated bur-
rows (Fig. 9.11d). According to Belaústegui and de Gibert (2013), the morphologic
characteristics of this ichnogenus display important similarities with burrows of
some terebellid polychaetes. The ichnogenus Cylindrichnus occurs since the
Cambrian (Mángano and Buatois 2014), and Belaústegui and de Gibert (2013)
reported that Cylindrichnus concentricus ichnofabrics occur commonly in Mesozoic

and Cenozoic offshore to shoreface settings.
Curvolithus comprises ribbon-like, flattened, endostratal trace fossils
(Fig. 9.11e). Three rounded lobes are present on the upper surface (Buatois et al. 1998).
Fig. 9.11 Ichnogenera produced by worms. (a) Asterosoma, scale bar is 5 cm; (b) Balanoglossites,
scale bar is 5 cm; (c) Chondrites, scale bar is 1 cm; (d) Cylindrichnus (modified from Belaústegui
and de Gibert 2009), scale bar is 1 cm; (e) Curvolithus, scale bar is 1 cm; (f) Gyrochorte, scale bar
is 1 cm; (g) Helicodromites, scale bar is 10 cm; (h) Nereites (modified from Seilacher 2007), scale
bar is 1 cm; (i) Phycosiphon (modified from Bromley 1996), scale bar is 0.2 cm; (j) Rosselia (modi-
fied from Nara 1995), scale bar is 8 cm; (k) Schaubcylindrichnus, scale bar is 1 cm; (l) Teichichnus
(transverse vertical section, modified from Nara 1995), scale bar is 1 cm
This ichnogenus is interpreted as the locomotion trace of infaunal predators, such as

flatworms or nemerteans, but gastropods cannot be completely ruled out as producers
(Buatois et al. 1998, see also Knaust 2010). It is known since the Cambrian
(Mángano and Buatois 2014).
Diplocraterion is a U-shaped, vertical burrow with protrusive (common) or retru-
sive (seldom) spreite. Openings to the seafloor are commonly funnel-shaped.
Diplocraterion is interpreted as a dwelling structure and an equilibrium structure
(Goldring 1962; Cornish 1986; Ekdale and Lewis 1991; Bromley 1996). Polychaetes
have been suggested as tracemakers (Arkell 1939), although crustaceans have been
indicated as well (Bromley 1996). Diplocraterion occurs in a wide variety of shal-
low- to deep-marine and even continental environments. However, it is more common
in nearshore high-energy settings and is known since the Cambrian (Mángano and
Buatois 2014).
Euflabella is a morphologically complex ichnogenus that comprises burrows
with single or multiple spreite bodies arranged in linear or radial patterns (Olivero
and López-Cabrera 2013). This trace fossil is interpreted as most likely produced by
an elongated worm-like organism and has been recognized only in Upper Cretaceous
and Paleogene shallow-marine settings (Olivero and López-Cabrera 2013).
Gyrochorte consists of curved to meandering, vertically penetrating burrows,
with a bilobate epichnial ridge and an underlying hypichnial groove, both with
transverse striae (de Gibert and Ekdale 2002; Fig. 9.11f). This ichnogenus is inter-
preted as produced by detritus-feeding worms, most likely annelids (de Gibert and
Benner 2002). It is known since the Early Ordovician, although most recorded
occurrences are from the Jurassic (de Gibert and Benner 2002 and references
therein).
Haentzschelinia is a vertical, radial spreite trace, having a central shaft (Fürsich
and Bromley 1985). This structure is interpreted as produced by worm-like organ-
isms with a proboscis used for reworking the sediment from a central shaft (Fürsich
and Bromley 1985; de Gibert et al. 1995; Uchman and Pervesler 2007). Such struc-
tures have been commonly referred to as Dactyloidites, but this name should be
reserved for more simple radial forms particularly common in early Paleozoic depos-
its (see Belaústegui et al. 2015b). Although it has been mentioned from the Triassic
(Beatty et al. 2008), the oldest illustrated occurrences have been documented in the
Jurassic (Agirrezabala and de Gibert 2004).
Helicodromites comprises horizontal spiral burrows, with pale fill contrasting
with the host rock (Fig. 9.11g). These trace fossils are interpreted as feeding
burrows constructed by vermiform organisms (probably capitellid polychaetes or
enteropneusts; Poschmann 2015). Helicodromites is common in both deep- and
shallow-marine deposits, especially in low-energy settings (Baldwin and McCave
1999). It occurs from the Devonian onward (Baldwin and McCave 1999;
Poschmann 2015).
Jamesonichnites consists of cylindrical tunnels with meniscate infill and well-
defined surrounding mantle (Dam 1990a). According to this author, Jamesonichnites
is transitional with Ancorichnus and it represents a feeding trace (fodinichnion) of a
soft-bodied organism. However, its possible occurrence in transition with Hillichnus
(Pazos and Fernández 2010) would be consistent with a bivalve producer. It is present
in shallow-marine settings, ranging from the Jurassic (Dam 1990a; Desai and
Saklani 2014) perhaps to the Cretaceous (Pazos and Fernández 2010).
Korymbichnus is similar to Arachnostega, although it is distinguished by filiform,
dichotomous branches that fan out distally (Damborenea and Manceñido 1996).
This structure is interpreted as produced by small deposit-feeding polychaetes that
fed at the interface between a shell and its internal mold. Korymbichnus is known
only from within mollusks molds from the Middle Jurassic of the Neuquén Basin,
Argentina (Damborenea and Manceñido 1996).
Lapispira consists of double helicoidal, concentrically arranged burrows with
coiling axes perpendicular to the bedding plane (Lanés et al. 2007; Pagani et al.
2012). This ichnogenus is interpreted as a feeding burrow, but its producer is uncer-
tain, with worms, such as polychaetes and enteropneusts, but also thalassinidean
crustaceans, as the most likely candidates (Lanés et al. 2007). It is only known from
Early Jurassic shallow-marine settings. Another possible recording of Lapispira has
been documented from the Miocene. It was attributed to crustaceans and most likely
represents a new ichnotaxon (de Gibert et al. 2012).
Macaronichnus is a mostly horizontal to sub-horizontal trace fossil with a fill char-
acteristically lighter than the host rock. It tends to occur in high densities. Modern
polychaetes (opheliids) produce structures similar to this ichnogenus by ingesting
sand to consume bacteria and organic material attached to the grains and excreting the
clean sand that fills the core of the burrow (Pemberton et al. 2001). This form is com-
monly present in upper-shoreface and foreshore deposits (Pemberton et al. 2001;
Seike 2009) and occurs since the Permian (Quiroz et al. 2010).
Nereites is a predominantly horizontal, winding to meandering trace fossil with
a central tunnel filled with relatively dark sediment and lateral lobes of reworked,
paler sediment (Fig. 9.11h). Interpreted as the structure of vermiform deposit-
feeders, probably enteropneusts (Uchman 1995 and references therein), Nereites is
commonly reported in both shallow- and deep-marine environments since the
Cambrian (Mángano and Buatois 2014).
Palaeophycus is a straight to slightly curved, inclined to horizontal burrow, with
smooth, ornamented, or annulated walls and with infill identical to the host rock.
This ichnogenus has been interpreted as the dwelling burrow of a predaceous or
suspension-feeding organism. Glycerid and nereid polychaetes form similar struc-
tures in modern environments (Pemberton and Frey 1982), but Palaeophycus is a
very simple structure that in fact can be produced by many different groups of phy-
logenetically unrelated organisms. It is a eurybathic trace fossil recorded from
shallow-marine, deep-marine, brackish, and continental paleoenvironments since
the Ediacaran (Mángano et al. 2002b and references therein).
Patagonichnus comprises a morphologically variable and complex burrow system
consisting of vertically and horizontally branched, lined tubes, with cylindrically
laminated structures surrounding the main tubes (Olivero and López-Cabrera 2005).
These authors suggested that Patagonichnus was most likely produced by poly-
chaetes, such as maldanids and capitellids. This ichnogenus is known since the
Miocene (Olivero and López-Cabrera 2005).
Phoebichnus comprises a central shaft consisting of stacked disc-shaped layers
from which several radial burrows emerged; these radial burrows display a lamination
inclined toward the shaft and an annulated mantle with annuli concave toward the shaft
(Bromley and Asgaard 1972; Dam 1990a; Evans and McIlroy 2016). Phoebichnus is a
feeding trace (fodinichnion) of a worm-like organism (Bromley and Asgaard 1972).
However, a crustacean producer has been suggested recently (Evans and McIlroy
2016). This ichnogenus is particularly common in shallow-marine settings, ranging
from the Jurassic to the Pleistocene (Bromley and Asgaard 1972; Kotake 2003).
Phycosiphon is a spreite trace fossil formed by recurving U-lobes in bedding
planes and having a dark core and a pale halo. This ichnogenus has been interpreted
as a feeding structure of vermiform organisms, probably polychaetes (Wetzel and
Bromley 1994), and it occurs in shallow- and deep-marine settings since the
Ordovician (Mángano et al. 2002b and references therein).
Planolites is an unlined, straight to slightly curved, smooth trace fossil, with striae
or annulations, being predominantly horizontal and with an infill contrasting in texture
with the host rock. It is interpreted as a feeding trace of deposit-feeder organisms, prob-
ably polychaetes (Pemberton and Frey 1982). Documented as a eurybathic trace fossil
recorded in shallow-marine, deep-marine, brackish-water, and continental paleoenvi-
ronments, Planolites is known since the Cambrian (Mángano and Buatois 2014).
Polykladichnus comprises vertical to steeply oblique tubes, lined or unlined, with
single or multiple U or Y shapes. The tubes are usually connected to the surface, and
slight enlargements can be seen at junctions. Probable producers for marine
Polykladichnus are polychaetes (Schlirf and Uchman 2005) and nemerteans (Knaust
2010), although cerianthid anemones (Schlirf and Uchman 2005) have been suggested
as well. This ichnogenus is known since the Devonian (Schlirf and Uchman 2005).
Rosselia is a vertical to inclined funnel-shaped burrow with a central tube filled
with sandy sediment and surrounded by concentric muddy laminae (Fig. 9.11j).
Specimens commonly exhibit erosional truncations. Rosselia is interpreted as the
dwelling structure of deposit-feeders, such as terebellid or trichobranchid poly-
chaetes (Nara 1995). This ichnotaxon is common in shallow-marine environments,
with normal-marine or brackish-water conditions, although it has also been reported
from deep-marine environments in a few cases (Mángano et al. 2002b and refer-
ences therein). Rosselia is known since the Cambrian (Jensen 1997).
Schaubcylindrichnus consists of oblique to horizontal bundles of congruent,
lined, circular, and pale-colored tubes (Fig. 9.11k). Some specimens show cross-
cutting relationships between successive tubes. This ichnogenus is usually inter-
preted as a dwelling structure produced by tubicolous deposit-feeders, similar to
modern maldanid polychaetes (Pemberton et al. 2001). Löwemark and Hong (2006)
considered the Schaubcylindrichnus tracemaker to be a suspension-feeding organ-
ism with the ability to construct closely spaced tubes sequentially within the sedi-
ment. This ichnogenus occurs in shoreface and upper-offshore sediments deposited
under normal salinity conditions (Pemberton et al. 2001). Although known from the
Carboniferous, it is much more common in post-Paleozoic rocks (Buckman 1997;
Löwemark and Hong 2006; Löwemark and Nara 2013).
Skolithos is a simple, vertical burrow that can be lined or unlined. In marine
environments, it is interpreted as the domicile of vermiform organisms (e.g., phoro-
nids and/or polychaetes; Schlirf and Uchman 2005). Skolithos occurs across a broad
range of shallow- to deep-marine and even continental environments, being most

common in nearshore high-energy settings since the Cambrian (Howard and Frey
1975; Mángano and Buatois 2014).
Taenidium is a cylindrical and unlined, sinuous trace fossil, with fill that consists
of meniscate segments alternately composed of fine- and coarse-grained sediment.
This ichnogenus is interpreted as a grazing or feeding structure produced by vermi-
form, deposit-feeding invertebrates that fed while moving through the sediment
(Keighley and Pickerill 1994). It occurs since the Cambrian in various marine envi-
ronments (e.g., marginal-marine, shallow- and deep-marine) and also in continental
settings (Buatois et al. 2007; Netto 2007).
Teichichnus is a horizontal to slightly inclined, unlined, simple structure, with
retrusive spreite (Fig. 9.11l). It is interpreted as the feeding structure of deposit-
feeding organisms, with each spreite formed during sediment mining, or an equilib-
rium structure with the spreite being in response to sea floor aggradation (Seilacher
1955; Corner and Fjalstad 1993). Probable tracemakers are polychaetes and sipun-
culan worms, but some arthropods may produce similar structures (Mángano et al.
2002b and references therein). This is a eurybathic form, being common in mar-
ginal-, shallow-, and deep-marine environments since the Cambrian (Mángano and
Buatois 2014).
Zoophycos is a complex trace fossil with a wide range of morphological vari-
ability (Olivero and Gaillard 2007; Fig. 9.11i). This ichnogenus ranges from simple
and isolated lobes with spreite to commonly large and very complex structures with
spiral morphology (Bromley and Hanken 2003). It is known since the Cambrian
(Olivero 2003). Various ethologic models have been invoked to explain the origin of
this structure, including strip mining, detritus feeding, refuse-dump, food cache,
and gardening (Bromley 1991; Löwemark 2015). Morphologic analysis suggests it
is the work of sipunculid or echiuran worms (Wetzel and Werner 1981; Kotake
1992; Olivero and Gaillard 2007). Olivero and Gaillard (2007) noted that the oldest
Zoophycos are usually unlobed, in contrast to more recent forms that show well
developed lobes (see also Seilacher 1986; Bottjer et al. 1988) (see Sect. 9.6).
Considering the evolutionary history of the above-mentioned groups of worms,
most of the major clades appeared during the early Paleozoic, although their body-
fossil record is patchy due to their low preservation potential. For example, unques-
tionable polychaetes had certainly appeared by the middle Cambrian (Glasby et al.
2000). Fauchald (1984) suggested that by the end of the Paleozoic, most of the
polychaete orders were already present, with groups such as opheliids (Scolecida)
recorded since the Devonian (Thompson 1979). Interestingly, some groups of poly-
chaetes became extinct by the end of the Paleozoic, and many groups recognized
today appeared during the Triassic (Glasby et al. 2000). In this regard, Vermeij
(1987) observed that lugworms (Arenicolidae), which intensely bioturbate sedi-
ments by excavating very deep burrows in modern settings, are known from the
Triassic onward. Similarly, Ippolitov (2010) studied the diversification patterns of
serpulids (calcareous tube-dwelling polychaetes) during the Mesozoic, recording an
important increase in the total biodiversity of this group during the Middle to Late
Jurassic. Knaust (2010) documented exceptionally preserved soft-bodied organisms
together with their trace fossils in a Middle Triassic mud flat. He recorded structures
produced by flatworms, nemerteans, nematodes, annelids, arthropods, and mollusks,
and concluded that these different taxa were already established in the early
Mesozoic (Knaust 2010).
Although the vast majority of the ichnotaxa attributed to worm-like organisms is
known since the early Paleozoic (e.g., Planolites, Palaeophycus, Cylindrichnus),
other more complex forms seem to have resulted from behavioral innovations asso-
ciated with the MMR (e.g., Bornichnus, Euflabella, Haentzschelinia, Lapispira,
Patagonichnus, Phoebichnus). Also, some ichnotaxa, albeit known from the
Paleozoic, are much more common in post-Paleozoic rocks (e.g., Macaronichnus,
Schaubcylindrichnus). In addition, some ichnogenera are represented by a few ich-
nospecies in the Paleozoic and then experienced marked increase in morphologic
variability, resulting in diversifications at ichnospecific rank. In addition, post-
Paleozoic ichnospecies tend to display deeper-tier positions and are morphologi-
cally more complex than Paleozoic ones. Good examples include Asterosoma and
Teichichnus. The ichnospecies A. radiciformis, A. ludwigae, and A. coxii are known
since the Devonian-Carboniferous, but the Jurassic shows the addition of A. surlyki,
A. striata, and an unnamed ichnospecies referred to by Seilacher (2007) as
Tatzelwurm (for discussion on these forms see Chamberlain 1971; Schlirf 2000;
Bromley and Uchman 2003, and Seilacher 2007). The ichnospecies Teichichnus
rectus, T. nodosus, T. flexuosus, and (probably) T. zigzag originated in the
Paleozoic, whereas T. sigmoidalis, T. spiralis, and T. patens have their earliest rep-
resentatives in the Mesozoic (for further discussion of these ichnospecies, see
Schlirf 2000 and Seilacher 2007).
9.2.2 The Main Groups of Bioeroders
Bioerosion was central to the MMR (Vermeij 1977; Bertling 1999; Perry and
Bertling 2000; Taylor and Wilson 2003; Bromley 2004; Wilson 2007). Of particular
significance is the role of sponges, gastropods, bivalves, chitons, echinoids, and
worms as bioeroders (Wilson 2007). Bryozoans and crustaceans were also impor-
tant bioerosion players in the MMR. Brachiopods, which typically produce etching
structures, were already established bioeroders in the Paleozoic and, therefore, are
not treated here. Algae, fungi, and foraminiferans, and cyanobacteria significantly
contribute to microbioerosion (Glaub and Vogel 2004; Bromley 2004; Taylor et al.
2015), but the majority of their ichnogenera have a Paleozoic origin (Glaub and
Vogel 2004; Bromley 2004). Vertebrates, such as fish and sharks, also were impor-
tant bioeroders during the MMR, as recorded by bite marks. However, in most
cases, these traces have not received ichnotaxonomic treatment. As with burrows,
the degree of precision in attribution of individual ichnotaxa to specific bioeroders
is variable, with endolithic algae, cyanobacteria, and bryozoans typically leaving
distinctive fingerprints and worm-generated structures having a higher level of
uncertainty (Bromley 2004). In this section, the most important bioerosion structures
produced by these groups of organisms are described (Table 9.2).
Table 9.2 Common bioerosion ichnogenera in shallow-marine post-Paleozoic deposits (see main
text for references)
Potential
Ichnogenera Stratigraphic range producers Ethology Trophic type
Belichnus Oligocene–Recent Stomatopods Praedichnia Predation
Caulostrepsis Devonian–Recent Spionid Domichnia Suspension
(polydorid) feeding/predation
polychaetes
Cavernula Ordovician–Recent Algae Domichnia Photoautotrophy
Centrichnus Cretaceous–Recent Anomiid bivalves, Fixichnia Suspension
balanid cirripeds feeding
Circolites Jurassic–Recent Regular echinoids Domichnia Grazing
Entobia Devonian–Recent Sponges (mostly Domichnia Suspension
clionaids) feeding
Finichnus Cretaceous–Recent Cheilostome Fixichnia Suspension
bryozoans feeding
Gastrochaenolites Ordovician–Recent Bivalves Domichnia Suspension
feeding
Gnathichnus Triassic–Recent Regular echinoids Pascichnia Grazing
Helicotaphrichnus Eocene–Recent Spionid Domichnia Commensalism
polychaetes
Meandropolydora Jurassic–Cretaceous Spionid Domichnia Suspension
polychaetes feeding
Oichnus Ediacaran–Recent Gastropods (also Praedichnia Predation
octopods)
Palaeosabella Ordovician–Recent Sipunculids and Domichnia Commensalism
polychaetes
Pennatichnus Miocene–Recent Ctenostome Domichnia Suspension
bryozoans feeding
Petroxestes Ordovician-Miocene Mytilid bivalves Domichnia Suspension
feeding
Radulichnus Jurassic–Recent Gastropods and Pascichnia Herbivory
polyplacophorans
Renichnus Cretaceous–Recent Vermetid Fixichnia Suspension
gastropods feeding
Rhopalia Carboniferous– Algae Domichnia Photoautotrophy
Recent
Rogerella Devonian–Recent Acrothoracican Domichnia Suspension
cirripeds feeding
Ropalonaria Ordovician–Recent Ctenostome Domichnia Suspension
bryozoans feeding
Saccomorpha Carboniferous– Fungi or algae Domichnia
Recent
Spirichnus Jurassic Worms Domichnia
Stellichnus Pliocene Ctenostome Domichnia Suspension
bryozoans feeding
Talpina Devonian–Recent Phoronid Domichnia
pseudocolonies
Teredolites Jurassic–Recent Wood-boring Fodinichnia Herbivory (wood
bivalves consumption)
Trypanites Cambrian–Recent Polychaetes and Domichnia Suspension
sipunculids feeding/predation
9.2.2.1 Sponges
Sponge borings typically consist of anastomosing channel networks generally form-

ing rounded chambers and commonly displaying diagnostic bioglyphs on the walls
(Bromley 1992, 2004). The most common boring produced by sponges is Entobia,
although other ichnotaxa (e.g., Clionolithes, Clionoides, Cicatricula) have been
attributed to the activity of sponges, albeit with variable degrees of certainty (Taylor
and Wilson 2003; Bromley 2004; Wilson 2007).
Entobia consists of multi-apertured and multi-chambered borings (Fig. 9.12b, d).
It is mostly produced by clionaid sponges, although other groups of modern endo-
lithic sponges, such as those in the Family Adociidae, produce similar structures
(Bromley 2004). Traditionally, Entobia was thought to range from the Jurassic to
Recent (Taylor and Wilson 2003; Bromley 2004), occurring in both shallow- and
deep-marine settings, although displaying higher ichnospecies richness in the for-
mer (Bromley and D’Alessandro 1990). However, subsequent taxonomic revisions
by Tapanila (2006) regarded the Devonian boring Topsentopsis as a junior synonym
Fig. 9.12 Bioerosion structures associated with the MMR. (a) Cliff conglomerate clast containing
deep Gastrochaenolites isp., Middle Miocene, Skotniki Quarry, Holy Cross Mountains, Poland;
(b) Entobia cracoviensis, a large chambered sponge boring formed in a Late Cretaceous abrasion
platform cut into Upper Jurassic limestone. Upper Cretaceous, Bonarka Quarry, Cracow, Poland;
(c) Teredolites clavatus, Horseshoe Canyon Formation, Upper Cretaceous, near Drumheller,
Canada, scale bar is 2 cm; (d) Gastrochaenolites isp. and Entobia isp. preserved as casts on a shell,
Rio Negro Formation, Miocene-Pliocene, Argentina
of Entobia. Therefore, the ichnospecies Entobia devonica is an early representative

of this ichnogenus, which reached a peak in ichnospecific diversity as a result of
the MMR.
9.2.2.2 Mollusks
Mollusks are well suited for the exploitation of hard substrates, producing both
dwelling and grazing structures. The focus here is on bioerosion structures attributed
to gastropods and bivalves, although chitons are also known to produce grazing
raspings and can collectively be a major source of bioerosion (Bromley et al. 1990;
Radley 2010).
Gastropods
Gastropods typically produced predatory borings, such as Oichnus (Wisshak et al.

2015) and radulation marks, primarily assigned to Radulichnus (Taylor and Wilson
2003; Bromley 2004; Wilson 2007). In addition, they may produce structures that
are intermediate between borings and surface scars, representing fixation/anchoring
traces or Fixichnia (de Gibert et al. 2004), as illustrated by the ichnogenus Renichnus.
Gastropods also produce a wide variety of etched and rasped pits and scars on
shells, but such marks have not yet received ichnotaxonomic treatment (Bromley
2004; Walker 2007). An extensive analysis of predatory gastropods and their associ-
ated structures, most of them still unnamed, is provided by Walker (2007).
Oichnus comprises circular or subcircular borings generated during predation by
gastropods (but also octopods; Bromley 1981, 1993; Nielsen and Nielsen 2001;
Taylor and Wilson 2003; Wilson 2007; Wisshak et al. 2015). This ichnotaxon is
known from the Ediacaran to Recent (Hua et al. 2003; Taylor and Wilson 2003),
occurring in both shallow- and deep-marine settings (Bromley 2005).
Radulichnus consists of a series of parallel sets of straight to curving scrape
marks produced by the radula of gastropods and chitons (Taylor and Wilson 2003).
This ichnogenus is known in shallow-marine settings since the Jurassic (Voigt 1977;
Kase et al. 1998).
Renichnus consists of etchings comprising a series of kidney-shaped depressions
produced by vermetid gastropods spiraling at an angle to the substrate surface
(Mayoral 1987a). It is known in shallow-marine settings from the Pliocene to the
Recent (Mayoral 1987a).
Bivalves
As well as being major bioturbators during the MMR, bivalves were also important
bioeroders. They produced dwelling structures not only in hardgrounds but also in
woodgrounds, as recorded by the ichnogenera Gastrochaenolites and Teredolites,
respectively.
Gastrochaenolites comprises clavate or drop-like trace fossils that are circular to

heart-shaped in cross-section, with a narrowed, neck-like upper part in most ichno-
species (Kelly and Bromley 1984; Figs. 9.6d, 9.12a, d). It is interpreted as a dwell-
ing structure produced by suspension-feeding bivalves that commonly bore into
rockgrounds, hardgrounds, and corals of shallow-marine environments (especially
intertidal to shallow subtidal environments), although they also can occur in firm
substrates (Kelly and Bromley 1984; Carmona et al. 2006, 2008). This ichnogenus
has been recorded in deposits ranging from the Ordovician to Recent (Pemberton
and Frey 1985; Ekdale and Bromley 2001b), but the earliest example confidently
attributed to bivalves is from the late Carboniferous (Wilson and Palmer 1998;
Taylor and Wilson 2003).
Teredolites consists of straight to clavate-shaped, gregarious, closely spaced bor-
ings in wood (Kelly and Bromley 1984; Figs. 9.6i and 9.12c). It is a dwelling struc-
ture produced by wood-boring bivalves (Kelly 1988). This ichnogenus ranges from
the Jurassic to Recent (Villegas-Martín et al. 2012) and has been described from
marginal-marine coal seams and woodgrounds, as well as from transported logs in
shallow- and deep-marine environments (Bromley et al. 1984; Buatois and Mángano
1992; Savrda and King 1993).
In addition, the ichnogenus Petroxestes, a boring with elongate outline and
rounded base, known from the Ordovician (Wilson and Palmer 1988) and the
Silurian (Tapanila and Cooper 2002), has been attributed to mytilids. This ichnoge-
nus represents an early example of bivalve bioerosion, significantly predating the
MMR and having been recorded also in the Cretaceous (Jagt et al. 2009) and
Miocene (Pickerill et al. 2001). The ichnospecies Centrichnus eccentricus, an
Upper Cretaceous to Recent ichnotaxon, consists of etching scars produced by
anomiid bivalves (Bromley 1999).
9.2.2.3 Echinoids
Echinoids also produce bioerosion structures, known from Mesozoic and Cenozoic
shells and hardgrounds. Most of these structures consist of a pattern of five radiating
grooves that are interpreted as tooth scratches produced by regular echinoids
(Bromley 1975).
Gnathichnus is a penta-radiate scrape mark produced by regular echinoids
(Bromley 1975, 2004; Taylor and Wilson 2003; Wilson 2007). Gnathichnus likely
reflects development of the jaw apparatus of regular echinoids (Bromley 1975). It is
known from the Triassic to Recent (Bromley 1975; Michalík 1977), occurring in
both shallow- and deep-marine settings (Bromley 2005).
Circolites comprises hemispherical pits that have been attributed to regular echi-
noids (Mikuláš 1992b; Bromley 2004). This ichnogenus ranges from the Jurassic to
Recent, occurring in shallow-marine settings (de Gibert et al. 1998a; Bromley 2004).
Ericichnus consists of series of deep and sinuous grooves and has been attributed
to regular echinoids (Santos et al. 2015). It is known from a rocky shoreline of the
Miocene (Santos et al. 2015).
9.2.2.4 Worms
Several bioerosion structures have been assigned to the activity of “worms,” especially
borings attributed to polychaetes and sipunculids (Tapanila and Hutchings 2012).
They include a wide variety of morphologies from simple borings to branching and
spiraled structures. Some of these structures are known since the Paleozoic (e.g.,
Caulostrepsis, Palaeosabella, Trypanites), whereas others appear to be a product of
the MMR (e.g., Maeandropolydora) and others appear even later (e.g.,
Helicotaphrichnus). A number of ichnotaxa (e.g., Lapispecus, Ramosulcichnus) may
be associated with the MMR, but they are only known from their type localities,
precluding further inferences; these structures are not discussed here. In addition, the
status of some worm ichnotaxa is a matter of debate (Taylor and Wilson 2003;
Bromley 2004; Wilson 2007). Finally, some of the ichnotaxa (e.g., Caulosptrepsis,
Talpina) that have their first occurrence in the Paleozoic but only achieve common
occurrence from the Mesozoic onward (Bromley 2004; Wilson 2007).
Trypanites consists of unbranched, cylindrical borings (Mägdefrau 1932). Examples
from the Ordovician contain scolecodonts within, suggesting that the producers were
polychaetes (Kobluk and Nemcsok 1982). Also, short and fat Trypanites are known to
be made today by sipunculids (Bromley 1978). Trypanites occurs since the Cambrian,
typically in shallow-marine environments (Taylor and Wilson 2003).
Maeandropolydora consists of sinuous to contorted galleries, having two or
more apertures (Voigt 1965; Bromley and D’Alessandro 1983; Taylor and Wilson
2003) and was probably produced by spionid polychaetes (Santos et al. 2003a). It is
known since the Cretaceous (Taylor and Wilson 2003), occurring in both shallow- and
deep-marine settings (Bromley 2005).
Caulostrepsis consists of a vertical gallery slightly bent to a U-shaped tube,
constituting an overall pouch- or ear-shaped structure, which can be even more
complex by the development of lobes (Bromley and D’Alessandro 1983). It has
been attributed to the activity of spionid (polydorid) polychaetes and is known since
the Devonian (Taylor and Wilson 2003), occurring in both shallow- and deep-marine
settings (Bromley 2005).
Palaeosabella comprises long, tubular borings that expand distally (Bromley
2004). This structure is attributed to sipunculids, which make similar bioerosion
structures today (Rice 1969). Palaeosabella has been regarded as a junior synonym
of Clionoides (Furlong and McRoberts 2014). However, Palaeosabella is mostly
an unbranched boring, whereas Clionoides is branched, forming complex three-
dimensional networks (Wilson 2007). This ichnotaxon is known since the
Ordovician (Bromley 2004), occurring in both shallow- and deep-marine settings
(Bromley 2005).
Talpina consists of curved and branching tunnels produced by phoronid pseu-
docolonies (Voigt 1972, 1978). It ranges from the Devonian to Recent (Rodrigues
and Gutschick 1970), occurring in both shallow- and deep-marine settings
(Bromley 2005).
Helicotaphrichnus is a helicoidal boring made in the columella of gastropod

shells occupied by hermit crabs (Kern et al. 1974). This ichnogenus is attributed to
spionid worms living in a commensal relationship with hermit crabs (Walker 1992).
It occurs in shallow-marine settings and ranges from the Eocene to the Recent
(Walker 1992).
Spirichnus is a cylindrical spiral boring that branches at regular intervals (Fürsich
et al. 1994). Although attributed to worms (Bromley 2004), no further details regard-
ing its affinities are known. It has only been recorded in Jurassic shallow-marine
settings (Fürsich et al. 1994; Bertling and Insalaco 1998).
Cunctichnus is a cylindrical boring with tapering side branches that occurs on
shells (Fürsich et al. 1994; Donovan et al. 2015). It has been attributed to sipuncu-
lids (Fürsich et al. 1994). Cunctichnus is present in shallow-marine settings and
ranges from the Jurassic to the Cretaceous (Donovan et al. 2015). However, doubts
persist regarding its taxonomic validity, and some authors have argued in favor
of synonymization with Vermiforichnus, which is known since the Devonian
(Taylor and Wilson 2003).
9.2.2.5 Bryozoans
Bryozoans are important bioeroders, although taxonomic issues (e.g., the fact that
some bryozoan biotaxa are erected based on their borings) complicate further evalu-
ation of their significance as players during the MMR (Bromley 2004; Wilson
2007). Finichnus (a senior synonym of Leptichnus; Taylor et al. 2013), arguably the
most widespread bryozoan ichnotaxon linked to the MMR, comprises surficial etch-
ings produced by cheilostome bryozoans (Taylor et al. 1999; Rosso 2008). It ranges
from the Cretaceous to Recent. Stellichnus and Pennatichnus have been attributed to
ctenostome bryozoans, but these forms have only been reported from the Pliocene
(Mayoral 1987b, 1988; Mayoral and Reguant 1995). Ropalonaria, ramifying tun-
nels that may have been produced by ctenostome bryozoans, is known since the
Ordovician, although some of its ichnospecies may have originated during the late
Mesozoic (Taylor and Wilson 2003; Bromley 2004).
9.2.2.6 Crustaceans
Crustaceans, major bioturbators of the MMR, are also important bioeroders.

Bioerosion structures are produced by acrothoracican and ascothoracican cirripeds,
brachyuran crabs, stomatopods, and thalassinideans (Seilacher 1969; Radwański
1977; Walker 1992; Taylor and Wilson 2003; Bromley 2004). Verrucid cirripeds
produce etching scars (Bromley and Martinell 1991). These are represented by the
ichnospecies Centrichnus concentricus, which is known from the Miocene to Recent
(Radwański 1977; Bromley and Martinell 1991). Rogerella, a pouch-shaped boring
produced by acrothoracican barnacles, is known from the Devonian, but is definitely
more abundant since the Mesozoic (Taylor and Wilson 2003; Bromley 2004).
Ascothoracican borings are known since the Cretaceous, but have not received
proper ichnotaxonomic treatment (Bromley 2004). Shell chipping and peeling are
commonly performed by decapod crustaceans, particularly brachyuran crabs
(Alexander and Dietl 2003). Although chipping and peeling have been known to
occur prior to the MMR, these activities are particularly common since the late
Mesozoic and are an innovation of the MMR (Huntley and Kowalewski 2007). Some
of the resulting structures have been recently included in the new ichnogenus
Caedichnus (Stafford et al. 2015). Brachyuran crabs experienced a rapid diversifica-
tion during the Late Jurassic (Klompmaker et al. 2013). Branching borings in corals
produced by thalassinideans are common, but have not received ichnotaxonomic
treatment yet (Bromley 2004). Belichnus, representing predatory holes produced by
blows struck by stomatopods, is known since the Oligocene (David 1997). However,
the finding that similar structures may be produced by sea gulls provides a caution-
ary note (Cadée and de Wolf 2013)
9.2.2.7 Algae
Endolithic, photoautotrophic organisms, such as algae, are responsible for a wide

variety of microbioerosion ichnotaxa, but most of them are known since the
Paleozoic (Bromley 2004; Glaub and Vogel 2004; Wisshak 2012). One exception is
Eurygonum, interpreted as produced by chlorophytes, which is known since the
Triassic (Wisshak et al 2005).
9.2.2.8 Fungi
A few microbioerosion ichnotaxa have been attributed to fungi. In particular, the

ichnogenera Saccomorpha and Orthogonum seem to be relatively abundant
(Schmidt 1992; Wisshak 2012). Although the possibility of an algal producer can-
not be disregarded for Saccomorpha (Glaub and Vogel 2004), recent detailed analy-
sis reinforces the idea of a fungal origin (Golubic et al. 2014). In any case, the
earliest representatives of these ichnogenera are Ordovician, whereas some ichno-
species (e.g., S. sphaerula) seem to have originated in the Cenozoic (Wisshak 2012).
9.2.2.9 Foraminifera
Foraminifera are known to produce borings of various morphologies, such as

horseshoe-shapes or rosettes (Bromley 2004; Bromley et al. 2007). Rosetted and
fan-shaped microbioerosion structures have been included in the Ichnofamily
Dendrinidae by Bromley et al. (2007). Of these, the ichnogenus Semidendrina has
been attributed to foraminifera and ranges in age from the Carboniferous to the
Recent (Bromley et al. 2007). However, this ichnogenus is most likely a junior syn-
onym of Nododendrina (M. Wisshak, written communication 2015). At present,
any connection between bioeroding foraminiferans and the MMR is unclear.
9.3 Ichnofaunal Composition, Infaunalization, and Tiering

Structure: Patterns Developed During the Mesozoic
Marine Revolution
The MMR strongly shaped the nature of animal–substrate interactions in the post-
Paleozoic world. Because trace fossils provide direct evidence of colonization of
infaunal ecospace, they represent useful tools to evaluate the degree of infaunaliza-
tion and complexity of tiering structure manifested during the MMR. In this regard,
comparing tiering structure and ichnoguilds through the Mesozoic and Cenozoic
may help to answer the question of whether or not the exploitation of deep infaunal
ecospace that characterizes the MMR was an abrupt or a protracted event (Thayer
1983; Bambach 1983). Within the bioturbated zone of soft sediments, two different
intervals are distinguished. The mixed layer occurs just beneath the sediment sur-
face and has a homogeneous appearance, whereas distinct burrows (trace fossils)
are formed within the transitional layer that lies below (Berger et al. 1979). Benthic
faunas from marginal-marine, brackish-water environments are affected by a num-
ber of stress factors (e.g., dilution of normal-marine salinities) and, therefore, their
ichnologic record may reflect departures from that of the archetypal, fully marine
ichnofaunas (Pemberton and Wightman 1992; MacEachern and Pemberton 1994;
MacEachern et al. 2007; Buatois and Mángano 2011) and will be addressed elsewhere
(see Sect. 9.5). In this section, the focus is on innovations involving trace-making
organisms and trace fossils that took place in open, shallow- and normal-marine
environments.
The timing of the MMR is a controversial issue. Central to this evolutionary
breakthrough is the replacement of brachiopod-dominated faunas to those domi-
nated by mollusks, which involved a change from primarily sedentary, epifaunal
suspension feeding to mobile suspension feeding and deposit feeding, with increased
predation (Wagner et al. 2006; Leighton et al. 2013). Certainly, this faunal turnover
started by the late Paleozoic when many groups of bivalves and gastropods, which
were previously dominant in nearshore settings, moved seaward to become part of
mixed brachiopod-mollusk communities in offshore environments (Sepkoski and
Miller 1985; Jablonski and Bottjer 1990; Olszewski and Patzkowsky 2001).
However, other major players in the MMR, such as decapod crustaceans and irregu-
lar echinoids, became dominant or appeared for the first time during the Jurassic.
From the perspective of bioerosion structures, an increase in diversity for macrobor-
ings occurred by the Devonian (Wilson 2007). Some bioerosion ichnotaxa that
appeared for the first time by the middle Paleozoic, such as Rogerella, Caulostrepsis,
and Talpina, became dominant later in the Phanerozoic (Bromley 2004; Wilson
2007). Signor and Brett (1984) noted that a rapid radiation of durophagous predators
also took place during the Devonian, and they considered this diversification event
as a precursor of the MMR. However, the middle Paleozoic diversification of bio-
eroders seems to be a distinct episode, whose link with the subsequent MMR is
tenuous at best.
9.3.1 The Triassic: Setting the Stage for the Mesozoic Marine
Revolution
The Triassic seems to represent a pivotal point in the ecology of shallow-marine

environments, being bracketed by two major mass extinctions. Unsurprisingly,
ichnologic information from the Triassic is biased toward the aftermath of the end-
Permian mass extinction and subsequent marine fauna recovery (see Chap. 7). To a
lesser extent, other ichnologic studies have focused on the end-Triassic mass extinc-
tion (see Chap. 8). Ichnofaunas from the lowermost Triassic, reflecting benthic con-
ditions during the immediate post-extinction aftermath, are typically monospecific
and consist of trace-fossil specimens that are smaller than normal and attributed to
opportunistic tracemaker organisms. During the rest of the Early Triassic, there is
evidence of recovery, as suggested by an increase in ichnodiversity, burrow size, and
depth of bioturbation (Twitchett and Barras 2004; see Chap. 7).
From an ichnologic perspective, some of the most thoroughly studied Middle to
Upper Triassic units are the Muschelkalk and Keuper of Europe (e.g., Pérez-López
1997; Knaust 1998, 2002, 2007, 2008; Knaust and Costamagna 2012; Chrząstek
2013). These studies offer a glimpse of animal–substrate interactions between the
two mass extinction events and, therefore, need to be treated in detail. In particular,
a comprehensive study by Knaust (2007) provided detailed documentation of the
ichnology of Muschelkalk carbonates in Germany. Forty ichnospecies (and 24 ich-
nogenera) were identified and described, pointing to a high-diversity community.
This is particularly remarkable given the restricted environmental conditions that
controlled deposition of the Muschelkalk carbonates. Both bioturbation and bioero-
sion structures are present. The former are dominated by trace fossils produced by
decapod crustaceans (e.g., Pholeus, Rhizocorallium, Thalassinoides), worm-like
organisms (possibly Balanoglossites, Arachnostega, and Zoophycos), actinarians
(Bergaueria), and bivalves (Lockeia, Protovirgularia), among other groups. The
presence of trace fossils interpreted as being produced by meiobenthos also was
indicated (Cochlichnus, Helminthopsis, Helminthoidichnites). Among the bioero-
sion structures (Caulostrepsis, Maeandropolydora, Palaeosabella, Trypanites),
polychaetes or sipunculans are the most likely tracemakers, although bivalve borings
(Gastrochaenolites) also have been recognized.
Tiering structure remained quite simple, but it is unclear if this was due to evolu-
tionary or environmental constraints. A remarkably similar ichnofauna to that of the
Muschelkalk in Germany has been recently documented for coeval strata in Poland,
which is thought to have been deposited in a shallow, storm-affected carbonate ramp

setting that is located within the same basin, but closer to the gateway to the Tethys
Ocean (Chrząstek 2013). The Polish ichnofauna consists of Archaeonassa,
Balanoglossites, Gastrochaenolites, Lockeia, Palaeophycus, Planolites,
Protovirgularia, Rhizocorallium, Skolithos, Thalassinoides, and Trypanites (Chrząstek
2013). A similar ichnofauna to those of Germany and Poland, albeit with lower diver-
sities, also has been recorded from Middle to Upper Triassic carbonates of Sardinia
(Knaust and Costamagna 2012).
With respect to Triassic shallow-marine clastics, a detailed study on the ichnology
of a transgressive–regressive deltaic succession of the Bravaisberget Formation in
Svalbard (Mørk and Bromley 2008) provided valuable information to compare with
the emerging picture based on carbonate ichnology. The tiering structure consists
essentially of three levels: a shallow tier dominated by Taenidium and Rhizocorallium,
a middle tier consisting of Thalassinoides, and a deeper tier represented by
Chondrites, which may occur as reburrowing of Thalassinoides fills; Polykladichnus
seems to have been associated with a late colonization event. Overall ichnodiversity
levels are moderate, and the tiering structure is quite simple. As in the case of the
Muschelkalk, discriminating between environmental constraints (e.g., salinity dilu-
tion, oxygen fluctuations) and evolutionary factors is not straightforward. The fact
that the tiers are interpreted as representing a succession of community replacements
caused by an unstable environmental regime, rather than a single stable community
(Mørk and Bromley 2008), supports the former possibility.
Despite some caveats, the emerging picture for the end-Triassic mass extinction
is also one of decreasing ichnodiversity, degree of bioturbation, and burrow size
(see Chap. 8). For example, the lower “Pre-Planorbis Beds” (uppermost Triassic) of
the Blue Lias Formation record low ichnotaxonomic diversity, low bioturbation
intensity, small burrow diameters, and an absence of deep-tier bioturbation, reflect-
ing the impact of the end-Triassic mass extinction (Barras and Twitchett 2007).
With regard to bioerosion, the bioeroders typical of modern reefs, such as endo-
lithic bivalves or clionaid sponges, were uncommon to absent prior to the Late
Triassic (Bertling 1999). By the Late Triassic, boring bivalves exhibited increased
size and abundance, accompanied by an increased importance of sponges (Perry
and Bertling 2000; Tapanila and Hutchings 2012). Although the Triassic witnessed
the appearance of some ichnotaxa (e.g., Gnathichnus) that were destined to become
dominant later, these ichnotaxa were relatively uncommon prior to the onset of the
MMR (Taylor and Wilson 2003; Bromley 2004).
Regarding microborings, although earlier studies suggested an early Mesozoic
radiation (Glaub and Vogel 2004), the subsequent finding of a superbly preserved
Carboniferous ichnocoenosis indicates that Carboniferous microendolith communi-
ties were “modern” in aspect (Wisshak et al. 2006). In fact, Mesozoic ichnodiversity
levels are similar to those of the Carboniferous (Wisshak et al. 2006).
Many Paleozoic predators became extinct by the end of the Permian, with few
groups surviving; however, some predators rebounded rapidly after the end-
Permian mass extinction (Walker and Brett 2002). According to these authors, dur-
ing the Middle Triassic, a number of new predator guilds appeared; these include
lobsters having appendages indicative of durophagy. However, an increase in
diversity of predator decapods took place later in the Late Cretaceous (Schweitzer
and Feldmann 2010). Also, some durophagous marine reptiles, boney fish, and
sharks may have been active predators since the Triassic (Cuny and Benton 1999;
Neenan et al. 2013; Brachaniec et al. 2015). In contrast, the sparse record of shell
boreholes may indicate that drilling predation was limited during the Triassic
(Kowalewski et al. 1998; Walker and Brett 2002). Also, the overall frequency of
shell repair is quite low during the Triassic (Vermeij et al. 1981; Walker and Brett
2002; Alexander and Dietl 2003).
Overall, sparse ichnologic information from the Triassic precludes generaliza-
tions. However, data from the Muschelkalk seem to suggest that by the Middle
Triassic, full recovery from the end-Permian mass extinction had taken place in
equatorial carbonate settings, particularly with respect to diversity levels (Knaust
2007), although refugia may have been present (Wetzel et al. 2007). Nonetheless,
Triassic ichnofaunas show limited infaunalization and relatively simple tiering
structure, particularly in clastic environments (e.g., Mørk and Bromley 2008).
Accordingly, the evidence from bioturbation and macrobioerosion structures
reviewed here seem to suggest that the Triassic is better regarded as a prelude to the
MMR, rather than part of this evolutionary breakthrough. However, ichnologic
information indicates that a few of the key players of this major evolutionary event,
particularly decapod crustaceans, were already dominant in carbonate settings by
the Middle to Late Triassic.
9.3.2 The Jurassic: Onset of the Mesozoic Marine Revolution
Recovery from the end-Triassic mass extinction is characterized by a gradual

increase in burrow size and the reappearance of deep-tier trace fossils, suggesting
the return to “normal” environmental conditions by the end of the Hettangian (see
Chap. 8; Barras and Twitchett 2007). In fact, an 8 % increase in global ichnodiver-
sity has been recorded in marine environments for the Early Jurassic (see Chap. 16).
The Lower to Middle Jurassic Plover Formation of the northwest Australia shelf is
a critical stratigraphic unit for assessing the degree of infaunalization and tiering
complexity at the onset of the MMR because it contains intensely bioturbated
marine deposits (Buatois et al. 2013; Burns et al. 2013). In particular, shelf, off-
shore, and shoreface deposits of Sinemurian to Aalenian age reveal quite complex
tiering patterns along the depositional profile (Fig. 9.13). For example, upper-
offshore mudstones of this unit are typically intensely bioturbated (BI 5–6), with
some intervals locally showing moderate levels of reworking (BI 3–4). The upper-
offshore ichnofauna consists of very shallow-tier Planolites montanus, shallow-
tier Asterosoma isp., mid-tier Thalassinoides isp., Palaeophycus tubularis, P.
heberti, Cylindrichnus concentricus, Rosselia socialis, Schaubcylindrichnus
upper shoreface middle shoreface lower shoreface upper offshore lower offshore
Pl Pl As
Pl Pl Pl
As
Tr T
Ta Ta Ta Ta
As Ta
Cy
Pa
Ro Pa
Tr Ro Sch
Pa As Pa
Ta Ta Ta
Ro Cy
Tr
T Z
Si Ph Ph
Op Si Ch
Di Di Ch Ch
Sk Ph Ph
Op Ch Ch
upper middle lower upper lower
Backshore Foreshore Shoreface Offshore Shelf
fair-weather wavebase
storm-wavebase
Fig. 9.13 Tiering structure along the depositional profile of a Lower to Middle Jurassic wave-
dominated shallow-marine platform, Plover Formation of the northwest Australia shelf
coronus, Teichichnus rectus, and Siphonichnus eccacensis, and deep-tier Chondrites

isp. and Phycosiphon incertum (Fig. 9.14a–f). From an ichnoguild perspective,
seven are recognized: the Planolites ichnoguild (vagile, deposit-feeding, very shal-
low tier), the Asterosoma ichnoguild (semi-vagile, deposit-feeding, shallow tier),
the Cylindrichnus-Rosselia ichnoguild (semi-vagile, deposit- to detritus-feeding,
mid-tier), the Palaeophycus-Schaubcylindrichnus ichnoguild (vagile, suspension-
to deposit-feeding, mid-tier), the Teichichnus ichnoguild (vagile, deposit-feeding,
mid-tier), the Siphonichnus ichnoguild (permanent, suspension-feeding, mid-tier),
and the Phycosiphon-Chondrites ichnoguild (vagile, deposit-feeding to chemosym-
biont, deep-tier).
The Sinemurian-Toarcian Neill Klinter Formation of Greenland is another
important unit to evaluate the onset of the MMR. Although no tiering analysis is
available from this unit, ichnologic studies by Dam (1990a, b) showed relatively
high-diversity assemblages in upper-offshore deposits, including dwelling
(Arenicolites isp., Diplocraterion parallelum, Ophiomorpha nodosa, Palaeophycus
isp., Thalassinoides isp.), locomotion (Cruziana isp., Curvolithus simplex,
Gyrochorte comosa), and feeding (Planolites beverleyensis, Rhizocorallium irregu-
lare, Taenidium serpentinum) structures. Similar ichnofaunas are known in shallow-
marine storm-affected deposits of the Lower to Middle Jurassic Bardas Blancas
Formation of Argentina (Bressan and Palma 2009).
Middle Jurassic shallow-marine ichnofaunas are particularly abundant in India,
where a number of studies have been made (e.g., Chiplonkar and Badve 1970;
Chiplonkar and Ghare 1975; Badve and Ghare 1978; Badve 1987; Fürsich 1998;
Sudan et al. 2000; Patel et al. 2008, 2014; Desai et al. 2008; Srivastava et al. 2010;
Fig. 9.14 Ichnofaunas from upper-offshore deposits of the Lower to Middle Jurassic Plover
Formation of the northwest Australia shelf. (a) Shallow-tier Asterosoma isp. (As), mid-tier
Thalassinoides isp. (Th), and deep-tier Chondrites isp. (Ch) and Phycosiphon incertum (Ph); (b)
very shallow-tier Planolites montanus (Pl), mid-tier Palaeophycus heberti (Pah), and deep-tier
Chondrites isp. (Ch) and Phycosiphon incertum (Ph); (c) mid-tier Palaeophycus heberti (Pah) and
Cylindrichnus concentricus (Cy), and deep-tier Chondrites isp. (Ch) and Phycosiphon incertum
(Ph); (d) mid-tier Thalassinoides isp. (Th) reworked by deep-tier Phycosiphon incertum (Ph); (e)
Shallow-tier Asterosoma isp. (As), mid-tier Palaeophycus heberti (Pah) and Teichichnus rectus,
and deep-tier Chondrites isp.; (f) mid-tier Siphonichnus eccacensis (Si), and deep-tier Chondrites
isp. (Ch) and Phycosiphon incertum (Ph). All scale bars are 1 cm
Joseph et al 2012; Gurav et al. 2014). With the exception of Desai et al. (2008),
these studies have focused on the broader characteristics of these ichnofaunas,
rather than tiering analysis. In any case, the available ichnologic information
indicates high diversity levels in fully marine environments, suggesting a continua-
tion of the trends displayed in the Early Jurassic. High-diversity levels also have
been noted for the Middle Jurassic Vardekløft Formation of Greenland (Heinberg
and Birkelund 1984; Fig. 9.15a–d). Relatively diverse ichnofaunas are also known
from the deltaic deposits of the Middle Jurassic Lajas Formation of Argentina
(McIlroy 2007 and Fig. 9.16a–d herein). In particular, offshore deposits were
intensely bioturbated, but lack of lithologic contrast prevented recognition of dis-
crete trace fossils (McIlroy 2007). These deposits pass upward into delta-front
deposits containing various ichnotaxa. The ichnogenus Haentzschelinia (Fig. 9.16c)
is present in some of these deposits together with crustacean burrows (Ophiomorpha,
Thalassinoides), among other forms, such as Asteriacites (Fig. 9.16a), Phycosiphon
(Fig. 9.16b), and Rhizocorallium (Fig. 9.16d) (McIlroy 2007; Canale et al. 2015).
Although the unit has been interpreted as having been deposited in a tide-dominated
delta (McIlroy 2007), recent observations indicate a greater fluvial dominance than
previously recognized (Canale et al. 2015).
Another Middle Jurassic unit interpreted as having formed in a tide-dominated
delta is the Ile Formation of offshore Norway (McIlroy 2004). Moderate ichnodiver-
Fig. 9.15 Characteristic trace fossils from Lower to Middle Jurassic shallow-marine deposits of
Greenland. (a) Haentzschelinia ottoi in bedding-plane view, unknown unit; (b) Haentzschelinia
ottoi in cross-section view, unknown unit; (c) Gyrochorte comosa and Lockeia amygdaloides,
Middle Jurassic, Vardekløft Formation, Jameson Land; (d) Phoebichnus trochoides, Lower
Jurassic, Neill Klinter Formation, Jameson Land. All scale bars are 1 cm
Fig. 9.16 Characteristic trace fossils from the Middle Jurassic Lajas Formation of Argentina. (a)
Asteriacites lumbricalis in cross section view, scale bar is 1 cm; (b) Phycosiphon incertum in cross-
section view, scale bar is 1 cm; (c) Haentzschelinia ottoi in bedding-plane view; (d) Rhizocorallium
isp. in bedding-plane view
sity was recorded in the associated offshore facies, where Phycosiphon, Phoebichnus,
Schaubcylindrichnus, Palaeophycus, and Thalassinoides are present. Gyrochorte,
otherwise a common ichnogenus in the Jurassic, is only abundant in the mouth-bar
facies, whereas archetypal crustacean burrows, such as Thalassinoides and
Ophiomorpha, occur in a wider spectrum of subenvironments (McIlroy 2004).
Available information indicates that by the Early to Middle Jurassic, shallow-
marine settings were host to diverse infaunal communities displaying complex tier-
ing patterns. Tiering and ichnoguild analysis reveals colonization of shallow-, mid-,
and deep-tiers by detritus, deposit and suspension feeders, as well as predators and
chemosymbionts. However, ichnodiversity per tier may be regarded as moderate.
Intensity of bioturbation commonly was conducive to total reworking of fine-grained
sediments, indicating the presence of efficient bioturbators.
The trends established during the Early to Middle Jurassic continued into the
Late Jurassic. Burrowing innovations are reflected by the appearance of crab bur-
rows, as recorded by the earliest occurrences of Psilonichnus (Fürsich 1981) and
Macanopsis (Neto de Carvalho et al. 2010) in coastal deposits of Portugal. From an
ichnofacies perspective, this represents the appearance of the Psilonichnus
Ichnofacies, which characterizes transitional settings, typically backshore areas,
washover fans, coastal dunes, and supratidal flats (Frey and Pemberton 1987; Curran
and White 1991; Buatois and Mángano 2011).
Upper Jurassic fully marine ichnofaunas have been documented in detail from
the North Sea (Fig. 9.17a–g). Typical elements of these Upper Jurassic deposits
are Ophiomorpha, Skolithos, Diplocraterion (Fig. 9.17g), Cylindrichnus,
Schaubcylindrichnus, Siphonichnus, Palaeophycus, Planolites, Asterosoma
(Fig. 9.17a–b), Phoebichnus (Fig. 9.17c–e), Teichichnus (Fig. 9.17f), Phycosiphon,
and Chondrites (Martin and Pollard 1996; Goldring et al. 1991, 2005; Baniak et al.
2014). In particular, the Fulmar Formation of England contains intensely biotur-
bated ichnofabrics that have been the focus of a number of studies (Taylor and
Gawthorpe 1993; Martin and Pollard 1996; Goldring et al. 1991, 2005). Pervasively
bioturbated Upper Jurassic shallow-marine deposits also have been recently
described from the Ula Formation of Norway (Baniak et al. 2014). In particular, the
offshore deposits of the Fulmar and Ula formations are remarkably similar to those
described from Lower Jurassic counterparts. For example, the ichnotaxonomic
composition of the upper-offshore deposits of the Ula Formation is almost identical
to that of the upper-offshore deposits of the Lower to Middle Jurassic Plover
Formation previously described. Although no information on tiering structure has
been provided for the Ula Formation, the available illustrations seem to indicate a
four-tier structure resembling that of the Plover Formation. A similar picture
emerges from deposits in the Fulmar Formation (e.g., Martin and Pollard 1996).
Ichnologic comparison of similar offshore deposits through the Jurassic indicates
that the main evolutionary innovations in these fully marine siliciclastic settings
took place in the Early Jurassic and persisted for the remainder of the period and, as
will be described, continued into the Cretaceous.
Jurassic nearshore sands were host to mid- to deep-tier infaunal communities
(e.g., Heinberg and Birkelund 1984; Dam 1990b; Buatois et al. 2013; Burns et al.
2013; Baniak et al. 2014). Crustacean burrows, such as Ophiomorpha, become
dominant particularly in middle- to upper-shoreface settings. In fact, Ophiomorpha
replaced Skolithos as the dominant component of the Skolithos ichnofacies in post-
Paleozoic strata (Droser and Bottjer 1993). The Sinemurian-Toarcian Neill Klinter
Formation of East Greenland contains extensive Ophiomorpha systems in strongly
storm-affected lower-shoreface deposits (Dam 1990b). This occurrence reveals that
an Ophiomorpha ichnoguild was already present in high-energy nearshore settings
by the Early Jurassic, revealing the establishment of a deep-tier malacostracan
guild. In addition to Ophiomorpha, relatively high-energy nearshore deposits are
characterized by deep-tier Diplocraterion habichi, which may reach up to 1 m
below the sediment-water interface. Because these structures can occur with high
densities, Diplocraterion habichi ichnofabrics may be regarded as a post-Paleozoic
equivalent of Skolithos pipe rock. Under slightly lower-energy conditions, most
typically in the lower shoreface, other ichnotaxa, such as Rosselia, Teichichnus,
Asterosoma, and Rhizocorallium, may become common as well (Ager and Walace
1970; Schlirf 2000, 2003).
Also, by the Jurassic, oxygen-depleted ichnofaunas of modern aspect appear to
have become well established (Savrda and Bottjer 1989; Wignall 1991; Etter 1995;
Fig. 9.17 Characteristic trace fossils from Lower to Middle Jurassic shallow-marine deposits of
the Yorkshire coast, northeast England. (a) Asterosoma isp. in bedding-plane view, Middle
Jurassic, Hundale Sandstone Member, Scarborough Formation, Hundale Point; (b) Asterosoma
isp. in cross-section view, Middle Jurassic, Hundale Sandstone Member, Scarborough Formation,
Hundale Point; (c) Phoebichnus trochoides, Middle Jurassic, Helwath Beck Member, Scarborough
Formation, Cloughton Bay; (d) Phoebichnus trochoides, Lower Jurassic, Blea Wyke Sandstone,
Blea Wyke Point; (e) Close-up of one of the radial branches shown in (d) to illustrate complex
backfilling; (f) Teichichnus rectus, Middle Jurassic, Gristhorpe Member, Scarborough Formation,
Yon’s Nab; (g) Diplocraterion parallelum, Lower Jurassic, Staithes Sandstone Formation, Staithes
Harbour. Scale bars are 1 cm, lens cap diameter is 5.5 cm, and coin is 1.8 cm
Monaco 1994, 1995; Komatsu et al. 2001; Olóriz and Rodríguez-Tovar 2002;
Uchman et al. 2003; Leonowicz 2012; Šimo and Tomašových 2013). Study of these
ichnofaunas has been favored by past occurrences of various anoxic events, which
have been particularly widespread in the European portion of the Tethys. Essentially,
all the basic aspects of the model of oxygen-related ichnocoenoses (ORI) put for-
ward by Savrda and Bottjer (1986, 1987, 1991) in their seminal papers were in
place by the Jurassic. These include a decrease in ichnodiversity, burrow diameter,
and burrowing depth in concert with decreasing oxygen content. According to this
model, structures emplaced in deep tiers in oxygenated sediments tend to move
upward as a response to the upward migration of the redox discontinuity under
dysoxic conditions. Chondrites generally has been regarded as the trace-fossil
form most tolerant to lowered oxygen content of bottom waters, showing dimin-
ished size and shallow-tier positions under extreme oxygen depletion (Bromley
and Ekdale 1984a).
Shallow-marine carbonate ichnofabrics tend to be dominated by crustacean bur-
rows, such as Thalassinoides (Fig. 9.18a–b) and Ophiomorpha (Monaco and
Garassino 2001; Monaco and Giannetti 2002; Giannetti and Monaco 2004; Goldring
et al. 2005; Neto de Carvalho et al. 2010). The latter commonly form nodular lime-
stones, a fabric in which crustacean burrows serve as nuclei for CaCO3 precipitation
(Fürsich 1972; Fig. 9.18a). Other ichnogenera common in Jurassic shallow-marine
carbonates include Rhizocorallium (Fig. 9.18c–d), Diplocraterion (Fig. 9.18e),
Phymatoderma (Fig. 9.18f), Chondrites (Fig. 9.18g), and Cylindrichnus (Fürsich
1974, 1975; Monaco et al. 1996; Monaco 2002; Goldring et al. 2005; García-Ramos
et al. 2011; Rodríguez-Tovar et al. 2012). As in the case of clastic settings, trace-
fossil assemblages in shallow-marine carbonates tend to reflect an energy gradient
along the depositional profile, with dwelling structures commonly, but not exclu-
sively, produced by suspension feeders (e.g., Diplocraterion, Arenicolites,
Ophiomorpha) occupying high-energy proximal areas, monospecific suites of
Teichichnus in protected lagoonal deposits, and a wider variety of ethologies and
trophic types (e.g., Rhizocorallium, Chondrites, Planolites, Thalassinoides,
Cylindrichnus) present in distal, low-energy shelf deposits, as documented in Upper
Jurassic carbonates of England and France (Fürsich 1974, 1975).
Similar studies in the Upper Jurassic of Saudi Arabia also show comparable
proximal-distal trends in an unrestricted carbonate shelf, displaying high ichnodi-
versities in distalmost mudstone facies (Goldring et al. 2005). These authors noted
the dominance of deep-tier Cylindrichnus typically cross-cutting shallow- to mid-tier
ichnotaxa, such as Planolites, Rhizocorallium, and Chondrites. In fact, Goldring
et al. (2005) noted a close correspondence between ichnofaunas from siliciclastics
of the Upper Jurassic Fulmar Formation of the North Sea and carbonates from the
roughly coeval Jubaila Formation of Saudi Arabia. The tiering structure of infaunal
communities in storm-affected carbonate platforms has been documented in the
Jurassic of Italy by Monaco et al. (1996) and Monaco (2002), who identified four
tiers: a shallow-tier Thalassinoides, a shallow-tier Planolites-Palaeophycus, a
Fig. 9.18 Characteristic trace fossils from Jurassic carbonates. (a) Thalassinoides nodular lime-
stone, Upper Jurassic, Coralline Oolite Formation, Carr Naze, England, lens cap diameter is
5.5 cm; (b) Extensive networks of Thalassinoides suevicus, Upper Jurassic, Coralline Oolite
Formation, Carr Naze; (c) Rhizocorallium jenense jenense, Lower Jurassic, Buerres Member,
Rodiles Formation, Punta Rodiles, Asturias, northern Spain, scale bar is 1 cm; (d) Rhizocorallium
jenense spinosus, Lower Jurassic, Buerres Member, Rodiles Formation, Punta Rodiles, Asturias,
northern Spain, scale bar is 1 cm; (e) Diplocraterion parallelum in bedding-plane view, Lower
Jurassic, Buerres Member, Rodiles Formation, Punta Rodiles, Asturias, northern Spain; (f)
Phymatoderma isp., Lower Jurassic, Rodiles Formation, Vega Beach, Asturias, northern Spain,
scale bar is 1 cm; (g) Chondrites isp., Lower Jurassic, Rodiles Formation, Vega Beach, Asturias,
northern Spain, scale bar is 1 cm
Fig. 9.19 Rhizocorallium isp. from a Jurassic carbonate firmground, Upper Jurassic, Rodiles
Formation, El Puntal Cliffs, San Martín del Mar, Villaviciosa, Asturias, northern Spain. (a) General
view of the surface with several superbly preserved specimens of Rhizocorallium isp., lens cap
diameter is 5.5 cm; (b) Close-up of a specimen of Rhizocorallium isp. showing well-preserved
bioglyphs, scale bar is 1 cm; (c) Close-up of specimens of Rhizocorallium isp. with sharp burrow
boundaries
mid-tier Helminthopsis, and a deep-tier Chondrites. The ichnogenus Rhizocorallium

is quite common in carbonate firmgrounds as illustrated by spectacular examples
in the Jurassic of Asturias (García-Ramos et al. 2011; Rodríguez-Tovar et al. 2012;
Fig. 9.19a–c).
In addition to infaunalization in soft substrates, ichnologic evidence suggests
that the character and levels of bioerosion in hard substrates also showed significant
changes by the Jurassic (Fürsich et al. 1994; Kowalewski et al. 1998; de Gibert et al.
1998a; Bertling 1999; Bromley 2004; Tapanila and Hutchings 2012). In particular,
Fig. 9.20 Gastrochaenolites isp. and Trypanites isp. demarcating the carbonate hardground at the
contact between the Middle Jurassic Upper Inferior Oolite and the underlying Carboniferous lime-
stone, England. Scale bar is 1 cm
de Gibert et al. (1998a)) noted the sharp contrast between Paleozoic and post-
Paleozoic (mostly from the Jurassic onward) bioerosion ichnofaunas, with the
former being dominated by worm borings and the latter by bivalve and sponge
borings. Middle Jurassic patch reefs from Israel have been extensively bioeroded by
bivalves, worms, barnacles, and phoronids (Wilson et al. 2010). The documented
ichnogenera include Clionolithes (produced by sponges or algae), Gastrochaenolites
(bivalves; Fig. 9.20), Oichnus (likely a soft-bodied sclerobiont), Rogerella (acrotho-
racican barnacle), Talpina (phoronids), and Trypanites (worms; Fig. 9.20).
Fürsich et al. (1994) documented a high diversity of macroborings in an Upper
Jurassic reef, noting that this is the oldest example where borers destroyed such a
large part of the reef, removing up to 40 % of reef volume. Bertling (1999) indicated
that the appearance of a modern macrobioeroding reef fauna took place by the Late
Jurassic. However, a subsequent study by Perry and Bertling (2000) downplayed the
importance of the Late Jurassic radiation of macroborers in favor of a more protracted
process throughout the Jurassic. According to these authors, cirripeds, bivalves, and
worms dominated in the Early Jurassic, with clionaid sponges becoming main players
by the Middle Jurassic. The appearance of Radulichnus in the Jurassic is particu-
larly important because gastropods and chitons may have contributed significantly
to grazing disturbance in shallow-marine settings, leading to the post-Triassic
decline of articulate brachiopods and their retreat to deep-water and/or cryptic refugia
(Radley 2010).
A Jurassic diversification of macroborers was certainly not restricted to reefs, but
can be extended to hard substrates in other settings, such as mollusk shells and
limestone rockgrounds, which supported a bioeroding fauna of modern aspect,
including polychaetes, sponges, and bivalves (Taylor and Wilson 2003; Goldring
et al. 2005). Despite these changes, Tapanila (2008) noted that, with the exception
of echinoids, no new classes of organisms adopted an endolithic strategy during the
MMR. From an ichnofacies perspective, the Jurassic marks the appearance of the
Gnathichnus ichnofacies (de Gibert et al. 2007).
Woodgrounds were colonized for the first time by the Early Jurassic, as indi-
cated by the first appearance of the ichnogenus Teredolites, which is produced by
pholadacean bivalves (Villegas-Martín et al. 2012). This occurrence predates the
first recorded appearance of pholadacean body fossils, documented from the

Middle Jurassic (Haga and Kase 2011; Villegas-Martín et al. 2012). Innovations in
microbioerosion were not apparent by the Jurassic (Glaub and Vogel 2004; Wisshak
et al. 2006).
Increase in predation pressure is one of the most important features of the
MMR. Although increased predation is reflected in the body-fossil record, the ich-
nologic record of predation provides key information for understanding the role of
predators during this evolutionary breakthrough (Huntley and Kowalewski 2007).
In particular, the roles of drilling predation, as reflected by bioerosion structures,
typically including drill holes of the ichnogenus Oichnus (Müller 1977) in shells
(e.g., Kowalewski et al. 1998, 1999; Harper et al. 1998, 1999; Walker and Brett
2002; Harper 2003) and of durophagous predation, as evidenced by shell-crushing
structures (e.g., Harper 2003; Oji et al. 2003; Zatoń and Salamon 2008), have
received considerable attention. In addition, fish bite marks in echinoid spines have
been recently documented (Wilson et al. 2014). Evaluation of the role of drilling in
the Jurassic has been controversial, essentially due to a patchy fossil record
(Kowalewski et al. 1998, 1999; Harper et al. 1998, 1999). Although the 120 Myr
gap in the presence of drill hole structures, ranging from the end of the Late Triassic
to the beginning of the late Early Cretaceous, is starting to be filled with a few
reports (see discussions in Kowalewski et al. 1998, 1999; Harper et al. 1998, 1999),
available evidence seems to suggest that drilling was not a major selection pressure
during the Jurassic (Harper 2003; Huntley and Kowalewski 2007).
In contrast, a number of animals capable of durophagous predation, including
malacostracan crustaceans with crushing chelae and several vertebrates (e.g., marine
crocodilians, ichthyosaurs, and plesiosaurs), appeared for the first time by the
Jurassic (Walker and Brett 2002). The extent of durophagous predation by the
Jurassic is still uncertain, but the presence of abundant shells displaying crushing
structures most likely produced by predatory fish, probably selachians or teleosteans,
in the Middle Jurassic of Poland points toward intense predation pressure, at least
locally (Zatoń and Salamon 2008). Interestingly, grazing by gastropods, chitons,
and regular echinoids may have been responsible for widespread dislodgement,
removal, and consumption of sessile and cemented epifauna from hardgrounds.
According to this view, grazing bioerosion was an important factor of escalation
during the Jurassic (Radley 2010). This is consistent with the ideas of Vermeij
(2008), who noted that the Early Jurassic was characterized by a remarkable increase
in escalation-related innovations, with rapid diversification of clades displaying
adaptative responses to predators.
Ichnologic data suggest that the rise to dominance of the Modern Evolutionary
Fauna is clearly evidenced since the beginning of the Jurassic. Burrow systems
attributed on actualistic grounds to malacostracan crustaceans are pervasive in
shallow-marine carbonates and siliciclastics, including Ophiomorpha,
Thalassinoides, and Spongeliomorpha (e.g., Dam 1990a, b; Monaco and Garassino
2001; Monaco and Giannetti 2002; Giannetti and Monaco 2004; Neto de Carvalho
et al. 2010). Rhizocorallium is another ichnotaxon produced by crustaceans, at least
in some cases, which has been extremely common since the Jurassic (see review in
Knaust 2013). Although present since the Cambrian, post-Paleozoic forms of

Rhizocorallium record more sophisticated feeding strategies, including the presence
of spine-shaped protuberances that may have served as caches or for farming
(Rodríguez-Tovar et al. 2012; Fig. 9.18d).
The occurrence of Scolicia in shallow-marine deposits (e.g., Dam 1990a, b),
albeit rare in comparison with younger deposits, reveals the impact of irregular
echinoids. Bivalve trace fossils, such as Protovirgularia, although known since the
early Paleozoic, are quite abundant since the Jurassic (e.g., Hallam 1970; Leonowicz
2008). Trace fossils attributed to worm-like organisms or unknown animals reflect
the appearance of new behavioral programs. This is clearly evidenced by the
ichnogenera Haentzschelinia (referred to as Dactyloidites in some studies, but see
Belaústegui et al. 2015b) and Phoebichnus (Bromley and Asgaard 1972; Heinberg
and Birkelund 1984; McIlroy 2004, 2007; Srivastava et al. 2010; Patel et al. 2014;
Canale et al. 2015; Evans and McIlroy 2016). Other worm structures, such as
Gyrochorte, although present in Paleozoic rocks, became particularly abundant in
the Jurassic (e.g., Hallam 1970; Heinberg 1973; Heinberg and Birkelund 1984;
Powell 1992; Sudan et al. 2000; de Gibert and Benner 2002; Leonowicz 2008;
Bressan and Palma 2009; Patel et al. 2014). The ichnogenus Curvolithus, probably
produced by flat worms, also seems to be particularly abundant in Jurassic shallow-
marine deposits (Häntzschel and Reineck 1968; Heinberg 1973; Heinberg and
Birkelund 1984; Buatois et al. 1998). Both Gyrochorte and Curvolithus may be
regarded as part of a relatively widespread ichnoguild in the Jurassic, comprising
vagile, shallow-tier detritus-feeder and predation structures. The spiral burrow
Lapispira, a structure that may be attributed to either worms or crustaceans, is at
present only known from Lower Jurassic shallow-marine deposits (Lanés et al.
2007; Pagani et al. 2012, but see de Gibert et al. 2012 for a possible Miocene
occurrence).
Overall, Jurassic ichnofaunas reveal a compositional turnover with respect to
older, particularly Paleozoic, ichnofaunas and give strong evidence for the rise to
dominance of the Modern Evolutionary Fauna. Comparison of shallow-marine ich-
nofaunas through the Triassic-Jurassic suggests that this faunal turnover took place
essentially by the Early Jurassic as indicated by bioturbation structures, with evi-
dence of a more protracted process for macrobioeroders.
9.3.3 The Cretaceous: Continuation of Previous Trends
Cretaceous siliciclastic shallow-marine ichnofaunas are extremely well known and

have been analyzed in detail in several studies. One area that has been explored in
great detail is the Western Interior Seaway of North America (Howard and Frey
1984; Vossler and Pemberton 1988, 1989; Frey and Howard 1982, 1985, 1990; Frey
1990; MacEachern and Pemberton 1992; Pemberton et al. 1992; Saunders et al.
1994; Pemberton and MacEachern 1997; MacEachern et al. 1999). Although these
studies emphasized ichnofacies applications in facies analysis and sequence
stratigraphy, the data presented can be easily explored from an evolutionary per-
spective. As a whole, information from the Cretaceous of the Western Interior
reveals the establishment of diverse climax communities in fully marine offshore
settings. Ichnofaunas from these settings include a wide variety of ethologic types
and feeding strategies, represented by a large number of ichnogenera, such as
Thalassinoides (Fig. 9.21f), Diplocraterion (Fig. 9.21c), Cylindrichnus,
Protovirgularia (Fig. 9.21l), Schaubcylindrichnus (Fig. 9.21g), Gyrochorte
(Fig. 9.21h), Curvolithus (Fig. 9.21i), Halopoa (Fig. 9.21k), Palaeophycus,
Planolites, Teichichnus, Taenidium, Zoophycos, Helminthopsis, Phycosiphon,
Nereites (referred to as Cosmorhaphe in some of the original studies), Phoebichnus,
Scolicia, Bichordites (Fig. 9.21j), Chondrites, Rhizocorallium, Bergaueria,
Asterosoma, and Rosselia (Fig. 9.21d). Overall, no major departures are apparent
from the trends established in the Jurassic for these settings.
Relatively high-energy nearshore deposits from Cretaceous sections elsewhere
display ichnofaunas that are essentially identical to those of the Jurassic (e.g.,
Curran 1985; Martino and Curran 1990; Pollard et al. 1993; Nielsen et al. 1996;
Weissbrod and Barthel 1998). Cretaceous nearshore sandstones are dominated by
mid- to deep-tier dwelling burrows, typically of crustacean (e.g., Ophiomorpha;
Fig. 9.21a) and worm (e.g., Skolithos) origin. Crustaceans encompass a wide variety
of feeding strategies. Among other producers, although suspension-feeding strate-
gies are dominant, other trophic types include detritus feeding (e.g., Cylindrichnus,
Rosselia) and passive predation (Conichnus, Bergaueria, Fig. 9.21e). The ichnoge-
nus Macaronichnus became particularly common in high-energy nearshore sands of
Cretaceous deposits (see review by Quiroz et al. 2010; Fig. 9.21a).
From the perspective of infaunalization, Upper Cretaceous lower-offshore
deposits of the Magallanes Formation of Patagonia, Argentina provide a glimpse
into the complexity of tiering structures by the end of the Mesozoic. These deposits
have been totally obliterated by bioturbation, mostly due to the activity of a deposit-
feeding infauna (Buatois et al. 2011; Fig. 9.22a–d). The shallowest tier consists of
an indistinct mottling (Planolites) representing background bioturbation in the
water-saturated, soupy substrate of the mixed zone. Below are shallow-tier
Thalassinoides, Phycosiphon, and Zoophycos. The mid-tier is represented by
Asterosoma, Planolites, and “Terebellina” (= Palaeophycus heberti), whereas the
deep tier is occupied by Chondrites, Teichichnus, and a second type of Thalassinoides.
The well-defined morphology and sharp burrow margins of Teichichnus, in addition
to the absence of other trace fossils cross-cutting this ichnotaxon, indicate emplace-
ment deep in compacted sediment. Six ichnoguilds are recognized: the Planolites
ichnoguild (vagile, deposit-feeding, very shallow tier), the Thalassinoides ichnoguild
I (semi-vagile and vagile, shallow-tier deposit-feeder structures), the Phycosiphon-
Zoophycos ichnoguild (vagile, deposit-feeding to chemosymbiont, shallow-tier), the
Asterosoma-Planolites-“Terebellina” ichnoguild (semi-vagile, deposit-feeding, mid-
tier), the Teichichnus-Chondrites ichnoguild (vagile, deposit-feeding to chemosym-
biont, deep-tier), and the Thalassinoides ichnoguild II (semi-vagile and vagile,
deep-tier deposit-feeder structures).
Fig. 9.21 Characteristic trace fossils from the Cretaceous Western Interior Seaway of North
America. (a) Macaronichnus segregatis, Upper Cretaceous, Horseshoe Canyon Formation, near
Drumheller, Alberta, Canada; (b) Ophiomorpha irregulaire, Upper Cretaceous, Horseshoe Canyon
Formation, near Drumheller, Alberta, Canada; (c) Diplocraterion parallelum, Lower Cretaceous,
Dakota Group, Alameda Avenue, Denver, Colorado, United States; (d) Rosselia socialis, Upper
Cretaceous, Horseshoe Canyon Formation, near Drumheller, Alberta, Canada; (e) Bergaueria isp.,
Upper Cretaceous, Wapiabi Formation, Mount Yamnuska Quarry, Alberta, Canada; (f)
Thalassinoides suevicus, Upper Cretaceous, Ferron Sandstone, Ivie Creek area, Utah, United
States; (g) Schaubcylindrichnus coronus, Upper Cretaceous, Panther Tongue Member, Star Point
Formation, Kenilworth Wash, Book Cliffs, Utah, United States; (h) Gyrochorte comosa, Upper
Cretaceous, Kenilworth Member, Blackhawk Formation, near the southern entrance to Tusher
Canyon, Book Cliffs, Utah, United States; (i) Curvolithus simplex, Upper Cretaceous, Kenilworth
Member, Blackhawk Formation, near the southern entrance to Tusher Canyon, Book Cliffs, Utah,
United States; (j) Bichordites isp., Upper Cretaceous, Panther Tongue Member, Star Point Formation,
Kenilworth Wash, Book Cliffs, Utah, United States; (k) Halopoa isp., Upper Cretaceous, Ram
Member, Cardium Formation, Seebe Dam, Alberta, Canada; (l) Protovirgularia dichotoma, Upper
Cretaceous, Kenilworth Member, Blackhawk Formation, Hutch Mesa area, Book Cliffs, Utah, United
States. Scale bars in (g, h, i, j, and l) are 1 cm, whereas those in (a, b, c, d, e, f), and (k) are 3 cm
Fig. 9.22 Intensely bioturbated Upper Cretaceous lower-offshore deposits of the Magallanes
Formation, Austral Basin, Patagonia, Argentina. (a) Shallow-tier Phycosiphon (Ph) and Zoophycos
(Zo), mid-tier Asterosoma (As), and deep tier Chondrites (Ch) and Thalassinoides (Th); (b)
Shallow-tier Phycosiphon (Ph), mid-tier “Terebellina” (T), and deep tier Chondrites (Ch) and
Teichichnus (Te); (c) Shallow-tier Phycosiphon (Ph), mid-tier Asterosoma (As) and “Terebellina”
(T), and deep tier Chondrites (Ch) and Teichichnus (Te); (d) Shallow-tier Phycosiphon (Ph) and
mid-tier Asterosoma (As). Scale bars are 1 cm
Fig. 9.23 A hardground surface demarcating the Intra-Valanginian Discontinuity (Lower

Cretaceous) from the Neuquén Basin of Argentina. (a) Close-up showing Thalassinoides isp. (Th)
penetrating well below the surface and Gastrochaenolites isp. (Ga) right below the discontinuity
surface; (b) Close-up showing Gastrochaenolites turbinatus (Gt) and Gastrochaenolites? isp. (G).
Note abundant intraclasts mantling the surface and unidentified borings. Scale bars are 1 cm
Information on Cretaceous oxygen-depleted ichnofaunas comes from the analysis

of the Oceanic Anoxic Event 2 (OAE-2) during the Cenomanian–Turonian transi-
tion in Europe (Uchman et al. 2008; Rodríguez-Tovar et al. 2009a, b; Monaco et al.
2012; Uchman et al. 2013). These trace-fossil assemblages are virtually identical to
those of the Jurassic. In fact, ichnofaunas of oxygen-depleted environments from
the Cretaceous onward seem overall to be quite conservative, and no major changes
have been recorded subsequent to the Jurassic.
As in the case of the Jurassic, Cretaceous shallow-marine carbonates tend to be
dominated by crustacean burrows, particularly in high-energy nearshore grain-
stones (e.g., Spalletti et al. 2001a, b), but also in slightly distal and lower-energy
packstones and wakestones (e.g., Mángano and Buatois 1994) and omission sur-
faces (e.g., Mángano and Buatois 1991; Schwarz and Buatois 2012; Fig. 9.23a–b).
Other less common elements, most likely produced by worms, including
Schaubcylindrichnus and Phycodes, tend to be present in more distal settings
(e.g., Mángano and Buatois 1994).
It has been noted that in modern environments, a rapid response of the deep-marine
benthic community is triggered by seasonal productivity, resulting in a pattern referred
to as the “benthic–pelagic coupling” (Graf 1989). Calcareous nannofossils, diatoms,
and dinoflagellates became the largest components of the phytoplankton during the
Cretaceous (Katz et al. 2007). As a result, primary production increased dramatically
(Kotake 2014). There is significant ichnologic evidence supporting the establishment

of a modern-style benthic–pelagic coupling during the Cretaceous (Kotake 2014;
Izumi 2015). These include the permanent occupation of the same burrow by the
Zoophycos producer as a response to high benthic-food conditions (Kotake 2014) and
the synchronization of deposit-feeding strategies with coccolithophore bloom by the
Phymatoderma animal (Izumi 2015). Therefore, ichnologic information underscores
the fact that significant changes took place in marine ecosystems by the late Mesozoic.
Another dataset comes from chalk that has been extensively studied from an
ichnologic perspective in Europe (e.g., Bromley 1967; Kennedy 1967, 1970, 1975;
Fürsich et al. 1981; Bromley and Ekdale 1984b, 1986; Ekdale and Bromley 1984,
1991) and the United States (Frey 1970, 1972; Frey and Bromley 1985; Savrda
2014). During the Cretaceous, thick chalk deposits resulted from the accumulation
of coccoliths on the seafloor, derived post-mortem from the flourishing populations
of coccolithophores that dwelled in the marine photic zone above (Stanley et al.
2005). Such deposits provide an ideal medium for ichnofabric analysis due to their
complete bioturbation as a result of very slow rates of sedimentation and full-marine
conditions (Ekdale and Bromley 1991; Savrda 2012). Coccolithophorids first
appeared in the Late Triassic and diversified rapidly during the Early Jurassic (Bown
2005), but it is not until the Cretaceous that this group became a major contributor
to ocean sedimentation (Hay 2004). The average size of individual coccoliths
increased with time, from Early Jurassic to Middle Cretaceous, and this size history
parallels the diversity history of Mesozoic coccolithophorids (Aubry et al. 2005).
In particular, shelf chalk beds, which formed between water depths of 50 and
300 m (Scholle et al. 1983), provide a window to unique infaunal tiered communities
in shallow-water settings (Figs. 9.24a–d and 9.25a–e). Crustacean burrow systems,
such as those of Thalassinoides, are dominant in these deposits, with Ophiomorpha
and Gyrolithes being less common (Ekdale and Bromley 1984). Feeding structures,
such as Chondrites, Phycosiphon, Zoophycos, Planolites, and Teichichnus, are com-
mon as well (Ekdale and Bromley 1984). Concretionary flint in chalk typically rep-
resents silicified burrow systems, such as Thalassinoides, commonly allowing for
three-dimensional preservation (Fig. 9.26a–b); Bromley and Ekdale 1984b).
Another common structure in European chalks is the large burrow structure
Bathichnus paramoudrae, which is typically encircled by ring-like paramoudra flint
(Bromley and Ekdale 1984b).
From an ichnofabric perspective, chalk hosts extremely complex, tiered ichno-
communities as demonstrated by the detailed analysis of pelagic deposits from
Denmark (Ekdale and Bromley 1991). As noted by these authors, degree of biotur-
bation in these deposits is so high that virtually every grain of sediment was pro-
cessed by several animals, resulting in composite ichnofabrics recording the activity
of multi-tiered infaunal communities. The shallowest tier corresponds to indistinct
burrow mottling onto which as many as four ichnoguilds are typically overprinted.
A complete overprinting sequence would include the Planolites ichnoguild (vagile,
shallow-tier deposit-feeder structures), the Thalassinoides ichnoguild (semi-vagile
and vagile, mid-tier deposit-feeder structures), the Taenidium-Phycosiphon ich-
noguild (vagile, middle-to-deep-tier deposit-feeder structures), and the Zoophycos-
Fig. 9.24 Chalk trace fossils, Upper Cretaceous, West Melbury Chalk Formation, Beachy Head,
southeast England. (a) General view of intensely bioturbated deposits overprinted by mid-tier
Thalassinoides paradoxicus; (b) Close-up of Thalassinoides paradoxicus; (c) Close-up of
Thalassinoides paradoxicus cross-cut by deep-tier Chondrites isp.; (d) General view of intensely
bioturbated deposits overprinted by mid-tier Thalassinoides paradoxicus. Scale bars are 1 cm
Chondrites ichnoguild (non-vagile, deep-tier deposit-feeder or chemosymbiont

structures). A similar tiering pattern has been recorded in chalk from Alabama
(Frey and Bromley 1985). The tiering structure of the Alabama deposits consists of
very shallow-tier, indistinct burrow mottling and three ichnoguilds, the
Thalassinoides ichnoguild (semi-vagile and vagile, mid-tier deposit-feeder struc-
tures), the Zoophycos-Taenidium ichnoguild (vagile, middle-to-deep-tier deposit-
feeder or farming structures), and the Chondrites ichnoguild (non-vagile, deep-tier
deposit-feeder or chemosymbiont structures).
Common macroborers in the Cretaceous include acrothoracican barnacles,
bivalves, clionaid sponges, polychaetes, and bryozoans (Taylor and Wilson 2003).
In the specific case of macroborers in reef environments, information is patchy for
the Early Cretaceous, but available data suggest a dominance of bivalves and worms
over cirripedians (Perry and Bertling 2000; Tapanila and Hutchings 2012). Bivalve
borings (Gastrochaenolites) were also common in Early Cretaceous hardgrounds
(Schwarz and Buatois 2012; Fig. 9.23a–b). During the Late Cretaceous, borer
sponges may have played a major role, particularly in rudist reefs, and to some
extent also in coral reefs, together with worms and bivalves (Perry and Bertling
2000; Tapanila and Hutchings 2012). Rocky shorelines also were affected by sig-
nificant bioerosion, with hardgrounds and carbonate rockgrounds displaying similar
borer communities (Taylor and Wilson 2003). A wide diversity of bioerosion structures
Fig. 9.25 Ichnofabrics and trace fossils in Eocene to Maestrichtian chalk cored at DSDP Site 605 (for
details about the lithology see van Hinte et al. 1987; for details about the ichnology see Wetzel 1987).
(a) Ichnofabric characterized by the vertical or steeply oriented spreite of Zoophycos isp. (Z) [note
deviating spreite (ds) and even upward shifting ones (us)], Thalassinoides isp. (Th) some no longer
identifiable burrows have been reworked by producers of Chondrites isp. (C); background sediment
has been homogenized (core 19, section 4, 20–29 cm); (b) Complex ichnofabric composed of
Planolites isp. (P), Teichichnus isp. (T), Thalassinoides isp. (Th), and Zoophycos isp. (Z); some bur-
rows have been reworked by producers of Chondrites isp. (C) (core 21, section 1, 75–81 cm); (c)
Complex ichnofabric composed of Zoophycos isp. (Z), Teichichnus isp. (T), and Planolites isp. (P) that
both have been reworked by producers of Chondrites isp. (C), and Thalassinoides isp. (Th) (core 17,
section 1, 21–29 cm); (d) Ichnofabric with dominant Chondrites isp. (core 41, section 6, 15–19 cm);
(e) Spirophyton-like burrow (S) and some Chondrites isp. tubes (C) (core 50, section 3, 64–71 cm)
Fig. 9.26 Concretionary flint in chalk representing silicified Thalassinoides isp. (a, b) Upper
Cretaceous specimens, Beachy Head, southeast England. All scale bars are 1 cm
was described from a Lower Cretaceous karst surface in the Czech Republic
(Mikuláš 1992b). The assemblage included borings by sponges (various ichnospecies
of Entobia), worms (Trypanites), acrothoracican barnacles (Rogerella), bivalves
(Gastrochaenolites), and echinoids (Circolites). Non-carbonate rocky shorelines
were characteristically of low diversity, as indicated by a monospecific suite of
bivalve borings (Gastrochaenolites) penetrating slates (Buatois and Encinas 2011).
Bioerosion in woodgrounds became much more common in the Cretaceous, as shown
by widespread occurrences of the ichnogenus Teredolites (Taylor and Wilson 2003;
Villegas-Martín et al. 2012). From a microbioerosion perspective, the Cretaceous
seems to show only a very slight increase in the number of ichnospecies recorded
(Wisshak et al. 2006).
Predation structures, drilling and, to a lesser extent, durophagous predation may
have increased in intensity during the Cretaceous (Walker and Brett 2002; Harper
2003, 2006; Huntley and Kowalewski 2007). There is overwhelming paleontologic
and ichnologic evidence that drilling predation underwent major evolutionary inno-
vation in the Cretaceous (Kelley and Hansen 2003; Huntley and Kowalewski 2007).
The earliest undisputed body-fossil record of muricid and naticid gastropods is from
the Late Cretaceous (Merle and Pacaud 2002), although drill holes identical to those
produced by these gastropods occur in the Early Cretaceous (Taylor et al. 1983) and
even the Triassic (Fürsich and Jablonski 1984). The appearance of these gastro-
pods undoubtedly increased predation pressures (Harper 2003, 2006). Octopod
cephalopods may have been important predators, but no undisputed ichnologic evi-
dence of Cretaceous drilling predation by these organisms has been documented
(Walker and Brett 2002). With respect to durophagous predation, the appearance
by the Cretaceous of many crustacean decapod families that are known to be active
predators (e.g., Palinuridae) represented a major breakthrough (Harper 2003). In
addition, many vertebrate predators were very active during the Cretaceous, with
crocodilians, ichthyosaurs, and plesiosaurs being common in the Early Cretaceous
and pliosaurids and mosasaurs as well during the Late Cretaceous (Walker and Brett
2002). Oji et al. (2003) noted that, although a number of durophagous decapod
crustaceans (as well as teleost fish) were present by the Late Cretaceous, the scarcity
of crushed shells suggests that durophagous predators did not play a major role in
Cretaceous ecosystems. However, their study is of very local scale, based on a

number of units in Japan, which may explain the discrepancies with broader compi-
lations, such as those by Vermeij et al. (1981) that noted a global peak in gastropod
shell repair frequency in the Cretaceous.
Regarding burrowers, crustaceans, bivalves, and echinoderms were among the
most important, signaling consolidation of the MMR in shallow-marine environ-
ments. Worm-like organisms also were undoubtedly key bioturbators in these settings.
Opheliid polychaetes became common in high-energy foreshore to upper-shoreface
sands, as evidenced by the abundance of Macaronichnus, particularly in mid- to high-
latitude settings (Quiroz et al. 2010). Also, in high-latitude settings, complex spreiten
burrows, namely, Paradictyodora and Euflabella, occurred for the first time (Olivero
et al. 2004; Olivero and López-Cabrera 2013; Olivero and López-Cabrera 2014).
To summarize, from the perspective of animal–substrate interactions in shallow-
marine siliciclastic settings, the Cretaceous represents a continuation of the trends
established in the Jurassic. However, by the end of the Cretaceous, a modern-style
benthic–pelagic coupling pattern was established and an important increase in
global ichnodiversity (35 %) took place in marine environments (see Chap. 16).
In carbonate settings, crustacean burrows are among the dominant forms. In the
specific case of chalk, the explosion in coccolithophores provided a new substrate
that allowed for the establishment of extremely complex tiering structures, unknown
in older deposits. Diversification of microborers was limited, but macrobioerosion
exhibited the increased activity of borers typical of modern communities (e.g.,
sponges, bivalves, gastropods).
9.3.4 The Paleogene: Rise of the Modern Evolutionary Fauna

After End-Cretaceous Mass Extinction
The end-Cretaceous mass extinction (see Chap. 12) differentially impacted the
various tracemakers involved in the MMR. Unfortunately, the present scarcity of
papers documenting Paleogene shallow-marine ichnofaunas inhibits detailed
recognition and reconstruction of patterns and trends.
The few available trace-fossil studies of shallow-marine clastic settings seem to
display similarities with late Mesozoic ichnofaunas, although the sparse record pre-
vents real evaluation of tiering structure. Crustacean burrows, such as Thalassinoides,
continued to be elite trace fossils (sensu Bromley 1990) in the Paleogene, having
been recorded in shallow-marine deposits as early as the early Paleocene (Danian;
Rodríguez and Panza 2003). Overall, typical Mesozoic elements, such as
Thalassinoides, Ophiomorpha, Psilonichnus, and Rhizocorallium, are dominant in
shallow-marine areas during the entirety of the Paleogene (Belt et al. 1997; 2005;
Myint 2001; Shelley and Lawton 2005; Uchman and Gaździcki 2006; Singh et al.
2008; Villegas-Martín et al. 2014). Other typical Mesozoic burrowers, such as irregular
echinoderms and prosobranch bivalves, also are represented by their trace fossils
in the Paleogene (Uchman and Gaździcki 2006; Demírcan and Uchman 2012;
Fig. 9.27 Characteristic trace fossils from the Middle to Upper Eocene Leticia Formation,
Patagonia, Argentina (a) Schaubcylindrichnus isp., cross-section view, coin is 1.8 cm; (b)
Gyrochorte isp., bedding-plane view; (c) Diplocraterion isp., bedding-plane view; (d) Scolicia
isp., bedding-plane view
Villegas-Martín et al. 2014). One of the most detailed analysis of Paleogene

ichnofaunas is that of López-Cabrera et al. (2008), who documented a high-diver-
sity ichnofauna from the upper Eocene of Tierra del Fuego, Argentina, comprising
the ichnogenera Diplocraterion, Schaubcylindrichnus, Palaeophycus, Planolites,
Rosselia, Asterosoma, Rhizocorallium, Taenidium, Teichichnus, Tasselia,
Curvolithus, Gyrochorte, Patagonichnus, and Paradictyodora (Fig. 9.27a–d). This
study showed that, at least by the middle Eocene, ichnodiversity levels were similar
to those attained in the late Mesozoic. Of particular importance is the appearance of
the ichnogenus Patagonichnus, a complex trace fossil attributed to gregarious poly-
chaetes (see also Olivero and López-Cabrera 2005).
As in the case of Jurassic-Cretaceous examples, Paleogene nearshore sands are
typically dominated by Ophiomorpha (Deville 1996), as well as by other vertical
burrows, such as Skolithos, Diplocraterion, and Arenicolites (Zhou 1997; Shelley
and Lawton 2005). Quiroz et al. (2010) noted that Macaronichnus, an ichnogenus
common in nearshore sands of high- to mid-latitudes, is absent in Paleocene to
lower Eocene deposits, a pattern that may reflect overall climatic conditions.
Evaluation of the macrobioerosion record is biased as a result of uneven strati-
graphic distribution of host substrates, but reports include borings in reefs, rocky
shorelines, clasts, and shells. In general, Perry and Bertling (2000) did not note any
major change in style of macrobioerosion across the Cretaceous–Paleogene boundary.

At present there are few examples of Paleocene and Eocene reef bioerosion, but the
sea-level highstand and global oligotrophic conditions during the Oligocene con-
tributed to extensive reef development (Perry and Bertling 2000). Bivalves were
apparently the most important macrobioeroders in reefs during the Paleocene, with
a subordinate presence of sponges, worms, and bryozoans (Babić and Zupanič
1981; Bernecker and Weidlich 1996; Schuster 1996). Sparse information from the
Eocene also underscores the important role of bioeroding bivalves in reefs (Gaemers
1978). Advanced teleost fish, most significantly parrotfish, were important contrib-
utors to macrobioerosion in tropical settings since the Eocene because they have a
sophisticated jaw mechanism for herbivory (Berg 1940; Steneck 1983). Perry and
Bertling (2000) noted that the interpretation of Oligocene patterns is not straightfor-
ward, with European reefs having a meager macrobioerosion record, whereas those
from the Caribbean display a high ichnodiversity.
In particular, patch and barrier reefs from Puerto Rico contain a wide variety of
ichnotaxa, including Trypanites and Uniglobites, as well as several ichnospecies of
Entobia and Gastrochaenolites (Edinger and Risk 1994). Overall, sponges domi-
nated over bivalves and worms in these reefs. In addition to reefs, rocky shorelines
were subject to significant bioerosion by clionaid sponges, bivalves, polychaete
worms, and barnacles (Roniewicz 1970; Hanna and Al-Radwany 1993). Composite
ichnofabrics typically developed in association with omission surfaces, including
both burrows in softgrounds and firmgrounds and borings in hardgrounds, leading
to complex patterns of cross-cutting relationships (Lewis and Ekdale 1992).
Bioerosion by bivalves and polychaetes has been recorded in clasts as well (Babić
and Zupanič 2000). Furthermore, bioerosion in shells was also common during the
Cretaceous–Paleogene, particularly in the Eocene-Oligocene (Taylor and Wilson
2003). Examples include borings by acrothoracican barnacles (Abletz 1993), pho-
ronids (Abletz 1994), and lithophagid bivalves (Krumm and Jones 1993; Krumm
1999). Nummulite tests breached by Oichnus have been documented from the
Eocene (Sengupta and Nielsen 2009). In addition, nummulitids were used as mate-
rials to reinforce burrow walls, as illustrated by the ichnogenus Nummipera (Hölder
1989; Jach et al. 2011; our Fig. 9.28a–b). During the Oligocene, whale skeletons for
Fig. 9.28 The ichnogenus Nummipera, characterized by the presence of nummulite shells rein-
forcing burrow walls. Middle Eocene, Soğucak Limestone Formation, Uğurlu Beach, Gökçeada
Island, western Turkey. (a) General view; (b) Close-up. Scale bars are 1 cm
the first time show evidence of bioerosion, most likely produced by the osteophagous
worm Osedax (Boessenecker and Fordyce 2014).
Although there was a slight increase in the diversity of microborings in the
Paleogene, a faunal turnover is apparent (Glaub and Vogel 2004). Change in the
dominant macrobioeroding ichnotaxa took place by this time, with the appearance
of new ichnospecies whose oldest record is Paleogene (Glaub and Vogel 2004;
Wisshak 2012).
The great end-Cretaceous mass extinction differentially impacted some groups
of predators. Many marine reptiles and the ammonites and belemnoids became
extinct, but the majority of invertebrate and fish predatory groups survived (Walker
and Brett 2002). Several predatory invertebrates (e.g., neogastropods, decapods)
and vertebrates (e.g., teleost fish, neoselachian sharks, sea birds, marine mammals)
diversified during the Paleogene. Although precise documentation of ichnologic
trends through time is difficult to evaluate, there is general agreement that drilling
predation increased during the Paleogene (Kelley and Hansen 2003; Huntley and
Kowalewski 2007). Vermeij (1987) indicated that an episode of drilling escalation
occurred between the Cretaceous and the Eocene, but the details of timing are
uncertain due to uneven stratigraphic distribution of data, including a lack of infor-
mation from the Paleocene.
A similar Cretaceous–Paleogene phase has been identified (Kowalewski et al.
1998). A decline of drilling frequencies across the Cretaceous–Paleogene boundary
followed by a subsequent increase may have resulted from the end-Cretaceous mass
extinction and subsequent recovery (Kelley and Hansen 1996, 2003). The record of
octopod shell drilling is essentially restricted to the Cenozoic (Walker and Brett
2002). An Eocene radiation of crushing predator crustaceans, including the
Portunidae, Cancridae, Calappiidea, Grapsidae, Partheopidae, and Majodae, may
have contributed to an increase in predation pressures (Walker and Brett 2002;
Harper 2003), although claw-like appendages, such as those in these groups, may
have evolved first as defensive traits (Vermeij 1982). Stomatopod crustaceans, com-
prising both gonodactyloids and squilloids, became important predators in the
Cenozoic, although only the former are durophagous predators, commonly preying
on mollusks (Walker and Brett 2002). The earliest representatives of gonodactyloid
stomatopods in the fossil record are Miocene (Hof and Briggs 1997; Hof 1998).
However, the ballistic ichnogenus Belichnus, which consists of small puncture
marks on mollusk shells (Geary et al. 1991; Pether 1995), has been recognized since
the Oligocene (David 1997). Vertebrate predators also contributed to durophagy,
including sea turtles as well as a number of marine mammals, the latter since the
Eocene (Walker and Brett 2002). Interpretation of shell repair frequency is not
straightforward, but Allmon et al. (1990) noted a drop in repair frequency for tur-
ritellid shells during the Paleocene and a rebound in the Eocene.
In short, Paleogene ichnofaunas from shallow-marine clastic settings are similar
to those of the late Mesozoic. Macrobioerosion styles seem to have persisted across
the Cretaceous–Paleogene boundary as well, although an increased role of sponge
and fish bioerosion is noted later in the Paleogene. In addition, predation pressures,
as reflected by both drilling and durophagy, likely increased during this interval
(Kowalewski et al. 1998; Kelley and Hansen 2003; Huntley and Kowalewski 2007).
9.3.5 The Neogene: Establishment of Modern Marine

Ecosystems
The Neogene records the establishment of modern-marine ecosystems. In comparison

with the Paleogene, the ichnology of Neogene shallow-marine clastic deposits has
been documented through a larger number of studies (e.g., Curran and Frey,
1977; Mayoral 1986; Ting et al. 1991; Martini et al 1995; Uchman and Krenmayr
1995, 2004; Pickerill et al. 1996, 1998; Hong 1997; de Gibert et al. 1998b, de Gibert
et al. 2013; de Gibert and Martinell 1998; Mayoral et al. 1998, 2013; Muñiz and
Mayoral 2001a; Buatois et al. 2003; Malpas et al. 2005; Campbell et al. 2006;
Cantalamessa et al. 2007; Carmona et al. 2008, 2012; Pervesler et al. 2011; Lokho
and Singh 2013).
Compositionally, Neogene shallow-marine ichnofaunas in clastic settings tend to
display similar elements to their Paleogene counterparts. Crustacean burrows, such
as Thalassinoides, Ophiomorpha, Spongeliomorpha, Psilonichnus, Macanopsis,
Gyrolithes, and Rhizocorallium, continued to be common elements (e.g., Mayoral
1986; Ting et al. 1991; Martini et al. 1995; Uchman and Krenmayr 1995, 2004;
Hong 1997; de Gibert et al. 1998b, 2013; Mayoral et al. 1998; Muñiz and Mayoral
2001a, b; Buatois et al. 2003; Carmona and Buatois 2003; Cantalamessa et al. 2007;
Carmona et al. 2004, 2008, 2012; Malpas et al. 2005; Campbell et al. 2006; Pervesler
et al. 2011; de Gibert et al. 2013). Both locomotion and resting trace fossils gener-
ated by irregular echinoids are common (e.g., Plaziat and Mahmoudi 1988; Colella
and D’Alessandro 1988; Uchman and Krenmayr 1995, 2004; Mayoral and Muñiz
2001; Buatois et al. 2003; Carmona et al. 2008, 2012).
Bivalves are also key burrowers in Neogene shallow-marine sediments, as
revealed by the abundance of the ichnogenera Lockeia, Protovirgularia, and
Siphonichnus (e.g., Buatois et al. 2003; de Gibert and Domènech 2008; Carmona
et al. 2008, 2010, 2012). Also added to this list should be two bivalve ichnogenera
that seem to have occurred for the first time by the Neogene: the equilibrium struc-
ture Scalichnus (Hanken et al 2001; Carmona et al. 2008) and the feeding trace
Saronichnus (Pervesler and Zuschin 2004). Equilibrium structures of terebellid
polychaetes, included in the ichnogenus Rosselia, are also common, sometimes dis-
playing a response to sedimentation pulses by formation of dense assemblages
(Nara 1995, 1997, 2002; Campbell et al. 2006; Frieling 2007). The highly complex
burrow system Patagonichnus, produced by gregarious polychaetes, displays
sophisticated feeding patterns expressed by its various ichnospecies (Olivero and
López-Cabrera 2005).
Neogene nearshore sands display similar ichnofaunas to those of other Early
Jurassic–Paleogene deposits. Typical ichnogenera are Ophiomorpha, Conichnus,
Bergaueria, Skolithos, Arenicolites, and Diplocraterion (e.g., Curran and Frey 1977;
Mayoral 1986; Ting et al. 1991; Martini et al. 1995; Uchman and Krenmayr 1995;
2004; Hong 1997; de Gibert et al. 1998b; Mayoral et al. 1998; Muñiz and Mayoral
2001a,b; Buatois et al. 2003; Carmona and Buatois 2003; Cantalamessa et al. 2007;
Carmona et al. 2004, 2008, 2012; Malpas et al. 2005; Pervesler et al. 2011).
Macaronichnus became common again, with records from a number of high-
energy settings (e.g., Uchman and Krenmayr 2004; Carmona et al. 2008; Quiroz
et al. 2010).
Despite the overall similarities with Paleogene ichnofaunas, Neogene ichnofau-
nas display an increase in ichnodiversity (both alpha and global sensu Buatois and
Mángano 2013; see Chap. 16) and a more complex pattern of infaunal tiering.
Detailed analysis in lower Miocene lower-shoreface to offshore-transition deposits
of Patagonia, Argentina (Fig. 9.29a–f) revealed the presence of six ichnoguilds and
nine tiers (Buatois et al. 2003; Carmona et al. 2012). This extremely complex infau-
nal tiering structure includes a mottled texture (vagile, deposit feeders, very
shallow-tier), the Thalassinoides-Asterosoma-Rosselia ichnoguild (semi-vagile,
Fig. 9.29 Characteristic trace fossils from Lower Miocene shallow-marine deposits of Patagonia,
Argentina. (a) Scolicia isp. and its producer, bedding-plane view, Chenque Formation, coin is
2.5 cm; (b) Ophiomorpha isp., bedding-plane view, Chenque Formation, lens cap diameter is
5.5 cm; (c) Helicodromites isp., bedding-plane view, Chenque Formation; (d) Scolicia isp., cross-
section view, note the two drain channels at the base of the specimens (white arrows); (e)
Ophiomorpha nodosa, cross-section view, Monte León Formation; (f) Asterosoma isp., cross-
section view
deposit feeders, shallow-tier), the Schaubcylindrichnus-Palaeophycus ichnoguild

(vagile, suspension and deposit feeders, middle-tier, the Scolicia-Phycosiphon-
Helicodromites-Teichichnus-Taenidium ichnoguild (vagile, deposit feeders,
middle-tier), the Thalassinoides ichnoguild (stationary, deposit feeders, deep-tier),
and the Chondrites ichnoguild (stationary, deposit feeders or chemosymbionts, very
deep-tier). This tiering structure reflects higher partitioning of the infaunal habitat
and an increase in degree of complexity than that found in older ichnofaunas from
siliciclastic settings. This is revealed not only by the number of ichnoguilds present,
but particularly by the increased diversity within individual ichnoguilds.
Regarding carbonate settings, an extensive review of the character and impor-
tance of ichnology to the study of shallow-marine carbonates was made by Knaust
et al. (2012). In a comprehensive study of the Bateig Limestone, Miocene of
southeastern Spain, de Gibert and Goldring (2007) identified several ichnofabrics
in a pelagic limestone, typically displaying intense bioturbation. The dominant
ichnogenera are Ophiomorpha, Bichordites, and Palaeophycus, with subordinate
presence of Planolites, Skolithos, and Teichichnus. Invariably, deeply emplaced
biogenic structures cross-cut indistinct mottling. Crustacean burrows commonly
form quite complex compound structures (de Gibert et al. 2012). More recently,
Belaústegui and de Gibert (2013) noted the abundance of Cylindrichnus, which
contributes to intensely bioturbated carbonate fabrics, and, in this case, represents
an elite trace fossil (see also Goldring et al. 2002).
Detailed information on carbonate ichnofaunas also has been derived from
Pleistocene–Holocene Bahamian-type carbonates (Curran 2007, and earlier papers
cited therein). The ichnologic study of grainstones on San Salvador and other
islands of the Bahamas Archipelago and the Miami Limestone of south Florida
defined five ichnocoenoses within the Skolithos and Psilonichnus ichnofacies
(Fig. 9.30a–g). Ophiomorpha is the dominant trace fossil in shallow subtidal grain-
stones whereas Psilonichnus upsilon, representing the burrows of the ghost crab,
Ocypode quadrata, is common in beach backshore beds. Carbonate dune deposits
(eolianites) also can exhibit a distinctive ichnocoenosis dominated by sometimes
complex insect burrows (Curran and White 2001). Similar subtidal ichnofaunas
have been recorded in the Pleistocene grainstones of the Cayman Islands (Pemberton
and Jones 1988; Jones and Pemberton 1989; Metz 2011).
As is the case with bioturbation structures, macrobioerosion during the Neogene
is essentially indistinguishable from that of the modern oceans. Modern reef bio-
erosion is dominated by grazing and, although the timing of this evolutionary
innovation is not totally clear, the onset of widespread grazing may have occurred
by the end of the Oligocene (Perry and Bertling 2000). Extensive information on
Fig. 9.30 (continued) shallow-subtidal grainstone, Upper Pleistocene, Cockburn Town Member,
Grotto Beach Formation, Clifton Pier, New Providence, pen is 15 cm in length; (f) Specimen of a
large, well-lithified Ophiomorpha nodosa shaft-terminus structure, Upper Pleistocene, Harry Cay,
Little Exuma; (g) Psilonichnus upsilon in beach-backshore grainstone, Holocene, Hanna Bay
Member, Rice Bay Formation, Moriah Harbour Cay, Little Exuma
Fig. 9.30 Characteristic trace fossils from Pleistocene–Holocene Bahamian-type carbonates.

(a) Closely-packed Ophiomorpha nodosa in shallow-subtidal, coral-rich grainstone, Upper
Pleistocene, Cockburn Town Member, Grotto Beach Formation, Cockburn Town Fossil Coral
Reef, San Salvador; (b) Ophiomorpha nodosa tunnels and shafts, bedding-plane view, same age
and location as (a), pen is 15 cm in length: (c) Shallow-subtidal grainstone with abundant
Ophiomorpha nodosa (Op), Planolites isp. (Pl), and Skolithos linearis (Sk), Upper Pleistocene,
Cockburn Town Member, Grotto Beach Formation, Harry Cay, Little Exuma, scale bar is 2.5 cm;
(d) Well-lithified Planolites, same age and location as (c); coin is 1.8 cm; (e) Conichnus conicus,
a structure commonly attributed to the resting and upward-movement activity of sea anemones, in
macrobioerosion comes from the study of rocky shorelines, which are abundantly
preserved in Miocene-Pliocene deposits. Clionaid sponges, echinoids, bivalves,
gastropods, polychaete worms, and barnacles are the dominant members of the bio-
erosion community in hardgrounds and carbonate rocky shorelines, as documented
in a large number of studies (e.g., Radwański 1970, 1977; Bromley and Asgaard
1993a, b; Mikulás and Pek 1995; Martinell and Domènech 1995; D’Alessandro and
Massari 1997; de Gibert and Martinell 1998; Domènech et al. 2001; Bromley et al.
2009; Caracuel et al. 2011; Demírcan 2012; Pineda-Salgado et al. 2015). Typical
ichnogenera include Entobia, Gastrochaenolites, Maeandropolydora, Centrichnus,
Caulostrepsis, Trypanites, Gnathichnus, Conchotrema, Oichnus, Phrixichnus,
Radulichnus, Renichnus, and Rogerella.
Shells and clasts are also host to a wide variety of bioerosion structures (e.g.,
Martinell and Domènech 1995; Kim and Heo 1997; de Gibert and Martinell 1998;
Doyle et al. 1998; Pickerill and Donovan 1998; Farinati and Zavala 2002; Santos
et al. 2003a, b, 2005, 2011). Detailed studies by Bromley and Asgaard (1993a) on
a coastal karst surface on the island of Rhodes in Greece demonstrated a complex
tiering structure with multiple ichnoguilds, namely the Oichnus ichnoguild (vagile,
predator, very shallow-tier), the Radulichnus-Gnathichnus ichnoguild (vagile,
algal grazer, surficial), the Centrichnus-Renichnus ichnoguild (permanent, preda-
tor, very shallow-tier), the Entobia ichnoguild (permanent, suspension-feeding,
mid- to deep-tier), the Trypanites-Caulostrepsis-Maeandropolydora ichnoguild
(vagile, predator, deposit feeder, and suspension feeder, deep-tier), and the
Gastrochaenolites-Phrixichnus ichnoguild (permanent, suspension-feeding, deep-
tier). This level of complexity in tiering structure remains undocumented for older
macrobioeroding communities, although it is unclear if this is real or due to a paucity
of observations.
Whale falls provided a suitable substrate for bioerosion during the Neogene
(Muñiz et al. 2010; Belaústegui et al. 2012; Higgs et al. 2012). Bioerosion structures
ascribed to the annelid Osedax (Trypanites) and bivalves (Gastrochaenolites) have
been recorded in whale skeletons (Muñiz et al. 2010; Belaústegui et al. 2012; Higgs
et al. 2012). From a microbioerosion standpoint, no major evolutionary innovations
seem to have arisen during the Neogene (Glaub and Vogel 2004). In fact, the more
recent compilation by Wisshak et al. (2006) indicated a decrease in ichnodiversity
at ichnospecies level.
Walker and Brett (2002) indicated a new phase of predator intensification during
the Neogene. Buccinid gastropods also diversified by the Neogene, further increas-
ing frequency and intensity of durophagy (Walker and Brett 2002). Sea otters, which
appeared by the Miocene, may have contributed to shell damage as well (Walker
and Brett 2002). Overall, both drilling predation on gastropods and bivalves and
shell repair frequencies seem to have increased during the Neogene, although not all
estimates in this regard are consistent (Allmon et al. 1990; Kelley and Hansen 1993;
Alexander and Dietl 2003).
To summarize, from an ichnologic perspective, the information available suggests
that the shallow-marine Neogene is indistinguishable from the modern regarding
both bioturbation and bioerosion structures. Shallow-marine ichnofaunas display a
continued ichnodiversity increase, as well as more complex tiering structures, albeit

without any significant change in overall taxonomic composition. A further increase in
predation pressure also is evident (Huntley and Kowalewski 2007), most likely as a
result of activity of predators such as stomatopod crustaceans and marine mammals.
9.4 The Paleoenvironmental Breadth of the Mesozoic

Marine Revolution
Although the MMR essentially took place in shallow-water, fully marine settings
(but see Walker and Voight 1994), its expression in other ecosystems, such as the
deep sea and marginal-marine, brackish-water settings, also can be evaluated by the
analysis of the trace-fossil record. Colonization of the deep sea has been analyzed
in detail by Uchman (2003, 2004). He noted that the end-Permian mass extinction
did not appear to have played a major role in decreasing trace-fossil diversity in
the deep sea. His studies indicated that ichnodiversity in deep-marine environments
displayed nonlinear changes, with peaks in the Late Jurassic–Early Cretaceous and
Eocene, reaching a maximum in the latter and decreasing afterward. Irregular echi-
noids and decapod crustaceans, key players of the MMR in shallow seas at least
since the Early Jurassic, apparently arrived to the deep sea by the Late Jurassic, as
indicated by the presence of Ophiomorpha (Fig. 9.31a–d) and Scolicia (Fig. 9.31f–g)
in turbidite deposits (Tchoumatchenco and Uchman 2001). The ichnogenus
Thalassinoides is also relatively common in deep-marine deposits (Fig. 9.31e). The
pellet-filled burrow Tubotomaculum seems to be restricted to deep-marine settings,
which is consistent with the cache strategy used by its crustacean tracemaker
(García-Ramos et al. 2014). Holothurids, as represented by the ichnogenus Artichnus
(and its potential senior synonym Naviculichnium; Fig. 9.31h), can be added to this
list. These bioturbators may have intensely ploughed the sea floor, increasing irriga-
tion of oxygenated waters, and thereby deepening the redox discontinuity surface,
which in turn allowed for colonization of deep tiers (Uchman 2004). Scolicia and
Ophiomorpha show an increase in frequency of occurrence from the Late Cretaceous
to the Eocene (Uchman 2004). Ichnologic information seems to suggest that the
effects of the MMR were slightly delayed in the deep sea. Body-fossil information
from deep-sea deposits is patchy, but Walker and Voight (1994) showed the exis-
tence of complex ecological interactions in Cenozoic deep-water gastropods,
including high rates of shell repair in large specimens, pointing toward intense pre-
dation pressures in the deep sea.
The temporal changes in brackish-water ichnofaunas have been addressed by
Buatois et al. (2005), who noted that, as in the case of the deep sea, the end-Permian
mass extinction apparently did not play a major role. These authors suggested that
the colonization of marginal-marine habitats did not occur at a constant rate, but
rather in a number of discrete phases. Two phases relevant to understanding the
expression of the MMR in brackish-water ecosystems were identified: Jurassic–
Paleogene and Neogene–Recent. The Jurassic–Paleogene phase was marked by an
Fig. 9.31 Ichnologic representatives of the MMR in deep-marine environments. (a) Ophiomorpha
annulata, Upper Eocene-Lower Oligocene, Ceylan Formation, Gelibolu Peninsula, western
Turkey; (b) Close-up of Ophiomorpha annulata showing the diagnostic pelletoidal walls, Upper
Eocene-Lower Oligocene, Ceylan Formation, Gelibolu Peninsula, western Turkey; (c) General
view of surface with Ophiomorpha rudis, Middle to Upper Eocene, Cerro Colorado Formation,
Cerro Colorado, Tierra del Fuego, Argentina; (d) Close-up of Ophiomorpha rudis, Middle to
Upper Eocene, Cerro Colorado Formation, Cerro Colorado, Tierra del Fuego, Argentina; (e)
Thalassinoides suevicus, Lower Cretaceous, Whisky Bay Formation, James Ross Island,
Antarctica; (f) Scolicia strozzi at base of a thick-bedded sandstone turbidite, Eocene, Piwniczna
Sandstone Member, Magura Formation, Tylmanowa-Baszta, Polish Carpathians; (g) Scolicia isp.
at base of a thin-bedded sandstone turbidite, Upper Cretaceous–Paleocene, Ropianka Formation,
Słopnice, Polish Carpathians; (h) High density of Artichnus pholeoides, a possible junior synonym
of Naviculichnium marginatum, a burrow produced by holothurians, Eocene, Magura Formation,
Tymbark-Miasto, Polish Carpathians. Lens cap diameter is 5.5 cm and scale bars are 1 cm
increase in ichnodiversity and degree of bioturbation in estuarine facies. In addition,

colonization was not restricted to softgrounds and firmgrounds, but took place in
hardground and xylic substrates as well. As in the case of fully marine ichnofaunas,
the Neogene–Recent phase recorded the rise of the modern brackish-water benthos,
accompanied by a further increase in ichnodiversity and intensity of bioturbation.
Furthermore, cemented surfaces, shells, and clasts began to be colonized by sponges,
polychaetes, gastropods, and bivalves, reflecting radiation of several groups of bor-
ers into brackish water. Most of the main players of the MMR in fully marine set-
tings, such as decapod crustaceans, bivalves, and worms, were also dominant in
brackish-water settings (Fig. 9.32a–f). Irregular echinoids, because of their lack of
tolerance to dilution of normal marine salinity, were the exception, and continued
for the most part to be restricted to fully marine settings.
Fig. 9.32 Ichnologic representatives of the MMR in marginal-marine environments. (a) Nereites
isp., bedding plane view; (b) Asterosoma isp., bedding plane view; (c) Protovirgularia isp.,
bedding-plane view; (a–c) prodelta deposits, Monte León Formation, Lower Miocene, Argentina;
(d) Rosselia socialis and Macaronichnus isp. in cross-section view; (e) Macaronichnus, bedding-
plane view; (f) Nereites isp., bedding-plane view; (d–f) delta-front deposits, Chenque Formation,
Lower Miocene, Argentina
9.5 Behavioral Evolution and Environmental Shifts
In addition to the arrival to the deep sea of crustaceans and echinoids capable of
creating extensive burrow systems during the MMR, other ichnotaxa experienced
environmental shifts coupled with behavioral changes. This is particularly well
illustrated by the trace fossil Zoophycos, which records a change in morphology
and in penetration depth and inferred behavior of the producer during the MMR
(e.g., Seilacher 1986; Chamberlain 2000; Kotake 2014; our Fig. 9.33a–e). The mor-
phology of Paleozoic and early Mesozoic Zoophycos is relatively simple, consisting
of irregularly lobate, “rooster tail”-like spreiten (Fig. 9.34a) with the tendency to
turn into a spiral (Seilacher 1986; our Fig. 9.34b–c). In addition, forms were
described with lobate spreiten deviating at several levels from a central shaft (e.g.,
Knaust 2004; Chamberlain 2000; Fig. 9.34d). Normally these Zoophycos were
emplaced in a shallow-tier position, and the producers followed a sediment-feeding
behavior (e.g., Knaust 2004). The host sediment is commonly muddy sand to sandy
mud that accumulated in shallow-marine waters from above wave base to outer
shelf settings (e.g., Bottjer et al. 1988). Some specimens, however, have been
reported from deeper-water deposits, but these occurrences are rare (e.g., Wetzel
et al. 2007). Morphologically similar Zoophycos specimens occur in fairly high
abundance until the Middle Jurassic (Seilacher 1986).
However, since the Late Triassic, Zoophycos producers tended to move to deeper-
water settings, and Zoophycos became sparse in deposits that accumulated above
storm-wave base. Late Jurassic shelf settings immediately below storm-wave base
were mostly abandoned by Zoophycos producers, and since the Oligocene outer-
shelf settings also were almost no longer utilized by these tracemakers (Bottjer et al.
1988), with just a few exceptions (e.g., Carmona et al. 2008, 2012). Instead, the
Zoophycos producers migrated to deeper-water environments (e.g., Seilacher 1986),
and today they inhabit continental slope and abyssal plain sediments virtually exclu-
sively (e.g., Wetzel 1991).
This shift in the habitat was accompanied by an evolution of behavior recorded
by the formation of multi-lobate and coiled, increasingly complex spreite of
Zoophycos (Seilacher 1986; Chamberlain 2000; Kotake 2014; our Fig. 9.34e–f),
along with an increase in size (area occupied by an individual specimen) and pene-
tration depth. The latter is documented by cross-cutting relationships with other
burrows. Cretaceous Zoophycos was commonly cut by Chondrites (e.g., Frey and
Bromley 1985), and Chondrites and Zoophycos may occupy the same tier in Upper
Cretaceous chalks (Bromley and Ekdale 1984b), whereas today Zoophycos repre-
sents the deepest tier and cross-cuts all other burrows including Chondrites (e.g.,
Wetzel and Werner 1981). Furthermore, a change in nutritional behavior can be
inferred, with a strip-mining, sediment-feeding strategy suggested for the Triassic
(Knaust 2004) and even Middle Jurassic (Olivero and Gaillard 1996). For modern
Zoophycos, a temporary-cache producer behavior can be deduced from material
within the spreite that is significantly younger than the host sediment (e.g.,
Löwemark and Grootes 2004). Some Zoophycos producers may have employed a
cache strategy as early as the Cretaceous, as some spreite exhibit obvious color
young
d e
b c
old
a
Fig. 9.33 Evolution of Zoophycos (based on Seilacher 1977 and Chamberlain 2000). (a) Basic,
protrusive “rooster tail”-shaped spreite, which occurred in the Palaeozoic and is still produced
today; (b) Continuous coiled spreite form, which occurred for the first time in the late Paleozoic to
early Mesozoic; (c) Discontinuously formed spreite consisting of several “rooster-tail”-like spreite
one attached to the side/tip of another, a morphology recorded for the first time in the late Paleozoic
to early Mesozoic; (d) Multiply coiled, helicoidal spreite typical of the Mesozoic onward; (e)
Helicoidal spreite system consisting of laterally assembled “rooster tail”-like spreite, also typical
of the Mesozoic onward (redrawn from Seilacher 2007)
changes between lamellae of the spreite and with the host sediment as well. This can
hardly be explained solely by in situ sediment sorting (for instance, see Fig. 3F in
Locklair and Savrda 1998). However, this observation has not yet been investigated
in detail. Furthermore, with time, it appears that the Zoophycos producers increas-
ingly preferred muddy substrates.
Fig. 9.34 Evolution of Zoophycos. (a) Simple, “rooster-tail”-like Zoophycos from Lower
Carboniferous limestone deposited near storm wave-base, near Ogmore-by-Sea, Great Britain (for
further details see Wu 1982); (b) Simply coiled Zoophycos from Bajocian siliciclastic fine-grained
deposits near Hauenstein (northern Switzerland); (c) Excavated coiled Zoophycos specimen from
Upper Cretaceous pelagic limestone (“Scaglia”) north of Sciacca (Sicily, Italy); (d) Lobate
Zoophycos from Upper Cretaceous Flysch showing major and minor lamellae (near Gropello,
northern Italy); (e) Zoophycos from the Paleogene Flysch in the Apennine near Florence, on dis-
play in the Institut und Museum für Geologie und Paläontologie, University of Tübingen in 1995.
The specimen was discovered by A. Seilacher, prepared by H. Luginsland and photographed by
W. Wetzel (all Tübingen at that time); courtesy of A. Seilacher (for details see Seilacher 2007,
plate 38 h); (f) Zoophycos with major and minor lamellae from Seravallian-Tortonian pelagic
Marne ad Orbulina near Monte Aquilone (Latium, Italy); courtesy of P. Belotti (Roma, Italy),
original figure (fig. 8) and further details in Bellotti and Valeri (1978)
The stratigraphic distribution of the U-shaped ichnogenus Rhizocorallium also

seems to show some connection with the MMR. Knaust (2013) reviewed the taxon-
omy of Rhizocorallium, concluding that only two ichnospecies are valid, R. commune
and R. jenense, although ichnosubspecies and varieties are recognized. Rhizocorallium
commune is known since the Cambrian and has not displayed significant morpho-
logic change, although size has shown some variation through the Phanerozoic,
essentially an increase through the Paleozoic, with a decrease after the end-Permian
mass extinction, followed by a time of new size increase (Knaust 2013). This ichno-
species has been attributed to worms (Knaust 2013). On the contrary, Rhizocorallium
jenense originated in the Triassic, reflecting an adaptation to firm substrates that was
unknown prior to the MMR, and having been produced most likely by crustaceans.
Also, as is the case for Zoophycos, Rhizocorallium seems to show environmental
shifts through time. In particular, Rhizocorallium commune is common in shallow-
marine environments during the Paleozoic and Mesozoic, becoming relatively abundant
in the deep sea during the Cenozoic (Knaust 2013).
9.6 Evolution of the Mixed Layer
The evolution of the uppermost interval of the seafloor, the mixed layer, cannot be
described step-by-step over long time spans. This layer is homogeneous in appear-
ance and lacks other diagnostic characteristics (Fig. 9.35a–d). Thus, its crucial
parameter is only its thickness, the so-called mixed layer depth (e.g., Boudreau
1998). However, the mixed layer depth cannot be measured in continuously accu-
mulating deposits, because deeper penetrating inhabitants of the transitional layer
normally overprint the homogeneous ichnofabric (Wetzel 1981; Werner and Wetzel
1982). Therefore, the preservation potential of the mixed layer is low. Only in the
case of frozen tiering resulting from abrupt de-oxygenation or non-erosive deposi-
tion of event beds will the mixed layer be preserved in the rock record (e.g., Savrda
and Bottjer 1986; Savrda and Ozalas 1992). Evidence of the mixed layer can be
detected, albeit indirectly, by the presence of burrow mottlings that are cross-cut by
discrete trace fossils emplaced in the transition layer.
Today the mixed layer of the seafloor is normally up to 10 cm thick as defined by
excess concentration of radiotracers (e.g., Boudreau 1998). In the rock record, how-
ever, the mixed layer thickness can be determined less exactly because its boundary
to the transitional layer is gradational (Fig. 9.35b). The homogeneous appearance of
the mixed layer is due to the production of biodeformational structures by meiofauna
and crypto-bioturbators, shallowly burrowing macrofauna, or even bulldozing mega-
fauna. These organisms utilize the organic matter close to or on the sediment surface
and process the deposits in a way such that no discrete traces are left. Furthermore, a
homogeneous appearance is favored by the very soft to soupy sediment consistency.
Mixed layer depth also depends on environmental conditions, in particular (1)
penetration depth of burrows; and (2) burrow diameter, with both decreasing with
Fig. 9.35 Mixed layer traces. (a) Mixed layer (h) in modern muddy sediment; upper boundary of
the transitional layer is marked by the occurrence of lined tubes (black arrows) or fairly sharp
bounded sand-filled Planolites-like burrows (white arrows). Core Sonne 220 34–1 (19° 14.06′
N/108° 9.23′ E), South China Sea, 50 m water depth; for details see Wiesner et al. 2012); (b)
Frozen-tiering structure underneath a low- to non-erosive turbidite having hydraulically light
foraminifera shells at the base. The transition between mixed layer and transitional layer is gra-
dational and occupies about ¼ of the maximum thickness of the transitional layer (~4 cm). Upper
Cretaceous flysch from the area of Gropello, northern Italy; (c) Scolicia isp. (Sc) producers bur-
rowing into the mixed layer from below and extinguishing it to a high degree; lower boundary of
a recently deposited event layer is marked by an arrow. Core Sonne 132 35–1 (13° 37.12′ N/119°
58.43′ E), South China Sea, 3202 m water depth; for details see Wiesner et al. 1998), 0–13 cm
core depth; (d) Basal part of a Thalassinoides filled with foraminifera shells (arrow) that have
been collected by the burrow producer from the seafloor, probably as additional food source. In
this way benthic food content of the mixed layer was lowered by a transitional-layer inhabitant.
Core 12345–5 (15° 28.8′ N/17° 21.6 W), off NW Africa, 966 m water depth; for details see
Wetzel 1981), 107–120 cm core depth
decreasing oxygenation and benthic food content (e.g., Wetzel and Uchman 2012).
In turn, the burial of organic matter is affected by the sedimentation rate (e.g., Müller
and Suess 1979).
Over geologic time, mixed-layer producers became capable of burrowing to
increasingly greater depths. A behavioral evolution is also evident for burrows pro-
duced by inhabitants of the transitional layer. Besides a trend to explore increas-
ingly deeper intervals within the seafloor, the burrow producers expanded their
behavioral programs to supplement limited food resources. With respect to the
mixed layer, three strategies of transitional-layer inhabitants are of importance, (1)
temporary feeding directly on the sediment surface; (2) burrowing upward into the
mixed layer from below; and (3) taking benthic food from the sediment surface to
store in a cache. Again, with time, the number and abundance of ichnogenera exhib-
iting such behavioral programs increased. For example, the producers of some
Nereites or Scolicia feed temporarily on or close to the sediment surface, respec-
tively (Fig. 9.35c; Wetzel 2008), and the producers of Zoophycos and Thalassinoides
can construct and utilize caches (Fig. 9.35d; e.g., Dworschak 1987; Löwemark and
Werner 2001).
Mixed layer thickness is affected by two long-term strategies of burrowing
organisms. The inhabitants of the mixed layer tend to expand their habitat vertically,
whereas deep-burrowing organisms living in the transitional layer utilize food
resources from the mixed layer. The mixed-layer inhabitants, however, often
respond more rapidly to environmental changes. Therefore, thickness of the mixed
layer, as evidenced by its homogeneous appearance, may fluctuate even during the
course of a year, depending on oxygenation and benthic food supply and overprint-
ing (competition) by transitional-layer inhabitants. Consequently, for an analysis of
the development of the mixed layer, it is necessary to compare mixed layers that
formed within similar environmental settings over geologic time. Given the low fos-
silization potential of the mixed layer, the data available at present are too sparse for
drawing detailed conclusions. In any case, available ichnologic information sug-
gests that the origin of the modern mixed layer can be traced back at least to the
Mesozoic.
9.7 Role of Paleogeography and Climate
There is growing evidence that trace fossils may reveal paleogeographic patterns
(e.g., Jensen et al. 2013), and that infaunalization trajectories may have been dispa-
rate in the different oceans surrounding various paleocontinents (e.g., Mángano and
Buatois 2011) during the Cambrian explosion (see Chap. 3) and the Great Ordovician
Biodiversification Event (see Chap. 4). Here we will explore potential similar ich-
nologic trends in connection with the MMR.
Understanding the role of paleogeography and climate on the MMR and its
potential ichnologic expression is still in its infancy. However, information from
modern environments provides an actualistic base that may help to detect potential
patterns in the post-Paleozoic world. Thayer (1983) indicated that deposit-feeding

taxa in warm-water settings tend to disturb sediment at a greater rate than is the case
in cold-water regimes. Cadée (2001) noted an increase in degree of bioturbation and
diversity from high-latitude Arctic coasts to low-latitude warm coasts. In addition,
he pointed to latitudinal faunal changes, with a diverse callianassid and crab fauna
in low latitudes, which is absent or has low diversity in Arctic regions. In the same
vein, irregular echinoids tend to display a preference for low- and mid-latitudes
(Goldring et al. 2004, 2007). Goldring and coworkers expanded these ideas, by inte-
grating observations from modern environments with information from the
Mesozoic and Cenozoic trace-fossil record. According to their model, the distribu-
tion of the benthos and associated biogenic structures can be summarized for the
three major climatic zones: (1) tropics and subtropics with Ophiomorpha and echi-
noid trace fossils; (2) temperate zone with echinoid trace fossils and Thalassinoides;
and (3) high latitude zone with only mollusk and worm trace fossils. Gingras et al.
(2006) extended the dominance of mollusk and worm burrows to the temperate
zone. In addition, Quiroz et al. (2010) summarized available information on the
polychaete trace fossil Macaronichnus, demonstrating its preference for high to mid
latitudes (see also Gingras et al. 2006).
Interestingly, Quiroz et al. (2010) noted the general absence of Macaronichnus
in Paleocene to lower Eocene deposits, and interpreted this pattern as a result of
overall high temperatures and expansion of subtropical belts during these times
(Zachos et al. 2001; Hollis et al. 2009). Quiroz et al. (2010) also noted that the few
middle to upper Eocene occurrences are from high latitudes (e.g., Olivero et al.
2008; Pearson et al. 2013), most likely reflecting the transition to overall colder
climates by these times.
These trends are, however, not without exceptions. Both Ophiomorpha and
Scolicia are abundant in Miocene mid-latitude shallow-marine deposits of Patagonia
under temperate to cold climates, representing a departure to the proposed pattern
(Buatois et al. 2003; Carmona et al. 2008). Also, Macaronichnus has been docu-
mented in the Neogene low-latitude nearshore deposits of Venezuela, representing
an apparent anomaly to the overall pattern (Quiroz et al. 2010). However, as noted
by these authors, this occurrence most likely reflects coastal upwelling of nutrient-
rich cold waters, which was extensive in the Caribbean prior to the final closure of
the Panama Isthmus in the Pliocene (O’Dea et al. 2007). Despite potential short-
comings, it seems that the present geographic distribution of infaunal organisms can
be traced back to the onset of the MMR.
Ongoing research by Olivero and López-Cabrera (2014) indicates that complex
spreite burrows, such as Paradictyodora and Euflabella, appear to be restricted to
southern high paleolatitudes during the Late Cretaceous. Strong seasonal variations
in primary production and cooling may have promoted the establishment of special-
ized strategies and behaviors. These authors also noted that this pattern in trace-
fossil distribution parallels the increased provincialism in bivalves and ammonites
during the Late Cretaceous in southern Gondwana, reflecting concurrent paleogeo-
graphical restrictions in the benthos and nekton.
Predation pressures in modern oceans also display a geographic component
(Vermeij 1978, 1987). He noted that predation is apparently more intense in the
Western Pacific and Indian Oceans than in the Atlantic Ocean, with the Eastern
Pacific occupying an intermediate position. From an ichnologic standpoint, this is
revealed by contrasting frequencies of shell repair in neritid and terebrid gastropods
in the different oceans. However, tracking these trends back in time to assess a
paleogeographic component of escalation is still premature. Also, information from
modern environments indicates that predation levels are higher in the tropics
(Freestone et al. 2011).
9.8 Discussion: Secular Changes in Bioturbation

and Ichnofaunas and the Mesozoic Marine Revolution
The pattern in trace-fossil distribution during the Mesozoic and Cenozoic as previ-
ously outlined provides a basis to discuss further how the MMR is expressed in the
ichnologic record. First, evaluation of the trace-fossil record may help to detect the
timing of this major evolutionary event. In this regard, the Triassic can be viewed as
setting the stage for the revolution, albeit with burrows by decapod crustaceans
already becoming dominant in carbonate settings (e.g., Knaust 2007). Interestingly,
the MMR appears to be at full speed by the Early Jurassic. This is evidenced not
only by the increased role of decapod crustaceans, but also by other key players of
the Modern Evolutionary Fauna, such as irregular echinoids and bivalves. Infaunal
tiering structure also displays a remarkable increase in complexity at this time
(Fig. 9.36). The dominant tracemakers remained essentially the same during the rest
of the Mesozoic and through the Cenozoic. Complexity of infaunal tiering struc-
tures also did not exhibit any further increase until the Neogene (Fig. 9.36). If this
chronology is correct, then infaunalization resulting from the MMR can be viewed
as a relatively rapid process followed by a long period of stasis. This pattern, essen-
tially based on the analysis of the ichnologic record, is partially consistent with that
derived from the analysis of the body-fossil record. In a series of classic studies
(Ausich and Bottjer 1982, 1985; Bottjer and Ausich 1986), the tiering history of
both infaunal and epifaunal communities of suspension feeders was assessed. These
studies demonstrated that epifaunal suspension feeders were strongly affected by
the end-Permian mass extinction, showing a relatively rapid recovery in the Triassic,
reaching a plateau of maximum diversity during the Jurassic, a subsequent decrease
through the Cretaceous, and then reaching another plateau in the Cenozoic. Infaunal
suspension feeders displayed a remarkable increase in penetration depth by the late
Paleozoic, being apparently unaffected by the end-Permian mass extinction and
reaching a plateau all through the Mesozoic and Cenozoic. Ichnogeneric compila-
tions show that the MMR was associated with an increase of global ichnodiversity
of 35 % in marine environments (see Chap. 16).
Secondly, comparing the timing of infaunalization with respect to the timing of
increase in predation pressures may yield some insights into the complexities of
escalation. Although there are still doubts regarding secular changes in predation
pressures, drilling apparently was not significant during the Jurassic (Harper 2003),
Bravaisberget Formation, MiddleTriassic, Plover Formation, Lower Jurassic,
shallow marine Ichnoguilds upper offshore Ichnoguilds
Taenidium Planolites
Pl
T T
Asterosoma
As As
Rh Rhizocorallium
Cylindrichnus-
Rh Cy Rosselia
Sch Ro
Schaubcylindrichnus-
Palaeophycus
Pa
Thalassinoides Teichichnus
Th
T
Siphonichnus
Si Si
Th
Ch Phycosiphon-
Chondrites Ph Ch
Chondrites
Ch Ch
Magallanes Formation, Upper Dania Quarry, Upper Cretaceous, Chalk

Cretaceous, lower offshore Ichnoguilds Ichnoguilds
Planolites Planolites-
Ta large Taenidium
Z Thalassinoides I Pl
Phycosiphon- Thalassinoides
Th
Ph Zoophycos Th Th
Ph
Ta Dark Planolites-
As Pl Pl Taenidium-
Planolites-
As Phycosiphon
Asterosoma-
Sch Schaubcylindrichnus
Ph
Z
Teichichnus- Zoophycos -
T Chondrites Chondrites
Ch Ch
Ch
Th Thalassinoides II
Capdevila Formation, Lower Eocene, Chenque Formation, Lower Miocene,

lower-shoreface to offshore-transition Ichnoguilds Lower-shoreface to offshore-transition Ichnoguilds
Mottled texture
Pa bm
Shallow Thalassinoides-
Palaeophycus
Shallow Thalassinoides-
As Asterosoma-Rosselia
Ro
Scolicia
Schaubcylindrichnus-
S Sch Palaeophycus
Asterosoma Pa
As Ta
Ph
Sc Scolicia-Phycosiphon-
Zoophycos-Chondrites- Helicodromites-
Ph Phycosiphon He
Z Teichichnus-Taenidium
As
T
Deep Thalassinoides
Ch Planolites-Helminthopsis Ch Chondrites
Pl
Pl
Fig. 9.36 Representative tiering structures and ichnoguilds from selected case studies illustrating
infaunal colonization during and in the aftermath of the MMR. Note that by the Early Jurassic, com-
plex tiering patterns have been established. However, an increase in complexity, particularly reflected
by the increased number of ichnotaxa per ichnoguild, is apparent by the Miocene. Reconstructions
based on Mørk and Bromley (2008); Burns et al. (2013); Buatois et al. (2013); Buatois et al. (2011);
Ekdale and Bromley (1991); Villegas-Martín et al. (2014) and Carmona et al. (2012)
although durophagous predation may have displayed an increase in importance,

albeit locally, by the Middle Jurassic (Zatoń and Salamon 2008). In any case, by the
Cretaceous, predation pressures experienced a significant increase (Vermeij et al.
1981; Kelley and Hansen 2003). Therefore, ichnologic evidence suggests that infau-
nalization predates an overall increase of predation pressures by approximately 50
Myr. The infaunal environment is typically regarded as a haven from predation
(Vermeij 1987) and, therefore, infaunalization could be perceived as an adaptive
response to increased predation (Bottjer and Ausich 1986). However, the chronol-
ogy presented above does not support this cause-effect scenario; rather a more com-
plex set of feedback mechanisms between infaunalization and predation is thought
to be the case. A similar conclusion was reached by McRoberts (2001) based on the
analysis of Triassic bivalves. Interestingly, infaunalization occurring before an
increase in predation has been noted during the Cambrian explosion as well
(MacNaughton and Narbonne 1999). In the same vein, complex geobiologic feedbacks
involving bioturbation have been invoked recently for the Cambrian diversification
event (Mángano and Buatois 2014).
Thirdly, evaluation of the Mesozoic–Cenozoic ichnologic record provides impe-
tus to revisit the bulldozing hypothesis of Thayer (1979, 1983), who proposed that
biological disturbance increased through the Phanerozoic. Sediment disturbance
tends to be greater when large, mobile bioturbators occur in dense populations
(Thayer 1983). In particular, sediment bulldozers rank as the most efficient biotur-
bators, with their ability to displace sediment, manipulate sediment in burrowing
and crawling, and manipulate sediment externally in feeding. Based on the integra-
tion of observations from modern environments with those from the fossil record,
he noted that post-Paleozoic bioturbators rework sediment faster, tend to burrow
deeper, and can generate shorter sediment-turnover times than Paleozoic taxa. The
present review of the post-Paleozoic ichnologic record supports this view. In fact, it can
be argued that a dramatic increase in sediment bulldozing is one of the diagnostic
features of the MMR. Most, if not all, of the bioturbators ranked by Thayer (1983)
as most efficient can be regarded as main players in the MMR. Examples include
decapod crustaceans [e.g., Callichirus major (formerly Callianassa major) and
many other callianassids, and the many species of Uca], clypeasteroid echinoderms
(e.g., Dendraster excentricus), irregular echinoids (e.g., Echinocardium cordatum,
Meoma ventricosa), tellinacean bivalves (e.g., Macoma balthica), protobranch
bivalves (e.g., Portlandia arctica), and arenicolid polychaetes (e.g., Abarenicola
pacifica). In addition to this list of invertebrates, it should be noted that marine
mammals are important sediment bulldozers throughout Cenozoic (Thayer 1983).
The turnover in ichnofaunal composition that took place in the early Mesozoic
fully supports an impressive increase in sediment disturbance in connection with
the MMR.
Fourth, it has long been argued that the fossil record indicates a pattern of onshore
origination of evolutionary innovations and subsequent expansion of new forms to
deeper water (Jablonski et al. 1983; Sepkoski and Miller 1985; Jablonski and Bottjer
1990; Sepkoski 1991; Jablonski 2005). Onshore-offshore trends are certainly
displayed by individual ichnotaxa, such as Zoophycos (Bottjer et al. 1988) and
Rhizocorallium commune (Knaust 2013). In the case of the innovations associated
with the MMR, the pattern of trace-fossil distribution supports a delayed arrival to
deep-marine settings. Whereas the changes of this significant evolutionary event
were in place in shallow-water settings by the Early Jurassic, they may only have
reached the deep sea by the Late Jurassic (Tchoumatchenco and Uchman 2001).
Intense bioturbation in deep-marine sediments has been recorded at least since the
Cretaceous (Ekdale 1977). In any case, biogenic mixing is even today much slower
in the deep sea than in shelf areas (Thayer 1983).
9.9 Conclusions
The ichnologic record of the MMR provides information on various aspects of this
major evolutionary event, such as the types of burrowers involved, the timing of this
breakthrough, the complexity of infaunal tiering structures, the paleoenvironmental
breadth of the event, and the interplay between predation and infaunalization,
among other points. Crustaceans, bivalves, echinoids, and worms were arguably the
most important burrowers. The MMR appears to be at full speed by the Early
Jurassic, as indicated not only by the overall dominance of these key burrower
groups, but also by the remarkable increase in complexity of tiering structures in
shallow-marine deposits. The complexity of infaunal tiering structures did not expe-
rience further increase until the Neogene, suggesting that the MMR was a relatively
rapid event followed by a long period of stasis. The MMR took place in shallow-
water settings, both fully marine and brackish, by the Early Jurassic, but this evolu-
tionary event may only have reached the deep sea by the Late Jurassic. Infaunalization
predated an increase of predation pressures by approximately 50 Myr, suggesting a
complex set of feedback mechanisms between predation and infaunalization rather
than a cause-effect between the two. Ichnologic information suggests that post-
Paleozoic bioturbators reworked sediment faster, tended to burrow deeper, and
had shorter turnover times than Paleozoic taxa. Our review strongly supports the
bulldozing hypothesis, indicating a dramatic rise in biogenic sediment distur-
bance during the MMR.
Acknowledgments We dedicate this chapter to our late colleague and friend Jordi de Gibert, who
contributed significantly to our understanding of the evolutionary aspects of ichnology, particu-
larly in connection with the origin of modern ecosystems. Many colleagues shared ichnologic
examples from throughout the geologic column and all over the world that helped us greatly in
developing our ideas on the trace-fossil record of the MMR and in illustrating representative ich-
nofaunas. These are Richard Bromley, Fiona Burns, Huriye Demírcan, Tony Ekdale, Jose Carlos
García-Ramos, Murray Gingras, Stephen Hubbard, María Isabel López-Cabrera, James
MacEachern, Francisco Medina, Eduardo Olivero, George Pemberton, Laura Piñuela, John
Pollard, Juan Ponce, Francisco Rodríguez-Tovar, Tom Saunders, Ernesto Schwarz, Andy Taylor,
and Alfred Uchman. Mark Wilson and Sally Walker greatly improved our manuscript with their
reviews. Nic Minter gave us useful feedback on the modes of sediment interaction used in
Table 9.1. The photographs used in Fig. 9.23 were provided by Ernesto Schwarz. Peter Pervesler
and Nic Minter provided photographs of Saronichnus and Hillichnus, respectively. Juan Ponce
helped us with preparation of the figures. Financial support for this study was provided by Natural
Sciences and Engineering Research Council (NSERC) Discovery Grants 311727–08 awarded to
Mángano. Netto is supported by the research grants 305208/2010-1 and 311473/2013-0 awarded
by The Brazilian Scientific and Technological Developing Council (CNPq). Carmona’s research is
funded by the Argentinean Research Council (PICT 2011–1373 and PICTO 2010–0199).
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Chapter 10
The Mesozoic Vertebrate Radiation
in Terrestrial Settings
Massimo Bernardi, Fabio Massimo Petti, Laura Piñuela, José Carlos García-
Ramos, Marco Avanzini, and Martin G. Lockley
10.1 Introduction
Given their worldwide abundance, it is not surprising that the study of Mesozoic
vertebrate tracks is the most ancient branch of vertebrate ichnology. Dinosaur tracks
were first observed and figured by Native Americans by, if not before, the seven-
teenth century (Lockley et al. 2006a), and the first document dates back to the earli-
est nineteenth century when Pliny Moody reported the discovery of large tracks in
the eastern U.S. With Edward Hitchcock (who would eventually coin the world “ich-
nology”), tracks began to be approached scientifically (e.g., Hitchcock 1858) and
subsequent papers by Wolfgang Soergel (Soergel 1925), Othenio Abel (Abel 1935)
M. Bernardi (*)
Department of Geology and Palaeontology, MUSE - Museo delle Scienze,
Corso del Lavoro e della Scienza 3, 38122 Trento, Italy
School of Earth Sciences, University of Bristol, Queen’s Road, Bristol BS8 1RJ, UK
F.M. Petti
PaleoFactory, Dipartimento di Scienze della Terra, Sapienza Università di Roma,
Piazzale Aldo Moro, 5, 00185 Rome, Italy
L. Piñuela • J.C. García-Ramos
Museo del Jurásico de Asturias [MUJA], Colunga, Asturias, España
M. Avanzini
M.G. Lockley
Dinosaur Tracks Research Group, University of Colorado Denver,
PO Box 173364, Denver, CO 80217-3364, USA

DOI 10.1007/978-94-017-9597-5_10
136 M. Bernardi et al.
and Roland Bird (Bird 1939, 1944) brought consciousness and the first scientific
methods into the discipline. Haubold (1971a, b) published two seminal papers which
constituted the base of the first reliable track-trackmakers associations.
In the late twentieth century, the discovery of numerous and vast dinosaur tracksites
contributed to the so-called “renaissance” in vertebrate ichnology (Lockley and Gillette
1987) and brought the first solid attempts to standardize study procedures (Leonardi
1987). With this new impetus the first ideas were put forward for using track data for
palebiologic (mainly behavioral and physiologic) studies (Lockley and Gillette 1987).
This continued in the last two decades with the publication of several seminal volumes,
especially on dinosaur tracks (Leonardi 1987, 1994; Gillette and Lockley 1989; Thulborn
1990; Lockley 1991; Lockley and Hunt 1995a; Lockley and Meyer 2000). In order to
fully contribute to the paleobiology of dinosaurs, however, the long-standing debate of
the relation between tracks and trackmakers needed to be addressed. No paleobiologic
inference can in fact be drawn except by interpreting tracks as the result of the activity of
a trackmaker, identified as belonging to a particular taxon. It is commonly assumed that
tracks can rarely be accurately associated with their producers at the species or genus
level (Farlow and Pianka 2000; Carrano and Wilson 2001). During its two centuries of
history, however, vertebrate ichnology built a broad consensus on the assignment of some
footprint morphogroups to broad taxonomic categories, based on clear analysis of the
morphologies of digits, hands, and feet, and comparisons of tracks with hand and foot
skeletons (e.g., Haubold 1971a, b; Demathieu and Demathieu 2004; Padian et al. 2010;
Heckert et al. 2010) or, rarely, on the direct association of track and fossil material (e.g.,
Voigt et al. 2012). Most authors agree that a trackmaker attribution at a high taxonomic
level is often possible (e.g., Baird 1980; Sarjeant 1990; Olsen et al. 2002; Carrano and
Wilson 2001; Thulborn 2006), and in fact morphofamilies or ichnofamilies have been
referred to in the literature since the early twentieth century (Lull 1904). In the 1980s,
Demathieu and colleagues began to elaborate on the possibility of implementing tracks
in phylogenetic studies (Demathieu 1981; Demathieu and Haubold 1978). Demathieu
also pioneered the use of vertebrate tracks in many other areas of paleobiologic studies
(Demathieu 1986); since then, a synapomorphy-based approach has been employed in a
number of studies (Olsen 1995; Olsen and Baird 1986; Olsen et al. 1998; Wilson and
Carrano 1999; Haubold and Klein 2000, 2002; Padian 2003; Wilson 2005; Wright 2005;
Brusatte et al. 2010b; Kubo and Kubo 2012; Bernardi et al. 2015). This perspective has
considerably constrained potential trackmakers finally allowing full hypothesis testing
and a better integration of vertebrate ichnology in paleobiologic analysis.
10.2 Trends and Events During the Mesozoic
10.2.1 Tetrapods Across the PT Boundary
A track is the result of the activity of a producer, or trackmaker, namely a track is

the imprint of an autopodium (a limb) registered during locomotion. The vertebrate
track record can be thought as an archive of the evolution of foot morphology and
10 The Mesozoic Vertebrate Radiation in Terrestrial Settings 137
posture of extinct and extant vertebrates. Here we must also remember that tracks
also record many details of soft tissue morphology, not revealed by body fossils,
which in many cases may lack foot skeletons, even when other key diagnostic ele-
ments are present. When consecutive steps are impressed as an animal moves, in
fact, a trackway is formed and its parameters (i.e., width, distance between single
tracks, angle between tracks, distance of the tracks from midline, etc.) can be used
to accurately define the walking posture of its producer. Assessing posture from a
skeleton involves observing limb articular surfaces and manipulating (physically or
virtually) bones at the hip, knee, and ankle in the hindlimb, and shoulder, elbow, and
wrist in the forelimb. This procedure is time-consuming and necessitates good pres-
ervation of the joints, which is not often the case. On the contrary, basic trackway
parameters, such as those listed here, can be easily measured in all kinds of track-
ways and are less dependent on substrate consistency and taphonomic deformation
with respect to single-track descriptive measurements (i.e., interdigital angles).
Furthermore, the track record is often much more abundant than the skeletal one,
thus giving opportunities to statistically analyze the data.
Limb postures of tetrapods vary from sprawling to erect. An erect limb can be
found within birds and cursorial mammals, where the limbs are held under the body
and move in a parasagittal plane. On the other hand, a sprawling limb posture is seen
in limbed squamates and all limbed amphibians, where the proximal limb segments
are highly abducted and swing largely in a horizontal plane (Blob 2000). Posture can
change during locomotion, a classic example being crocodiles which show a sprawl-
ing posture at low speeds and a semi-erect one when running (Zug 1974; Carpenter
2009). An erect limb posture is energetically advantageous over a sprawling limb
posture, especially for large animals. Stresses over the knee and elbow joints are
reduced and respiration is facilitated when the limbs are held vertical (Carrier 1987).
From skeletal studies, it is known that Late Permian tetrapods such as temnospon-
dyl amphibians, pelycosaurs, and pareiasaurs were largely sprawlers (Carroll 1988;
Benton 2005). Late Triassic faunas were instead dominated by erect forms such as
dinosaurs and crurotarsan archosaurs (Nesbitt 2011). The postural transition from
sprawling to erect between those times can be described by looking at skeletal remains,
but these are scanty in the Early and Middle Triassic. Vertebrate ichnology has allowed
a better understanding of this event, especially regarding the exact pattern, timing, and
possible correlation with other events by studying fossil trackways.
A key study in this respect was developed by Kubo and Benton (2009). The
authors used a classic, simple measure to distinguish sprawlers from erect walkers:
the ratio of stride length to trackway width, indicated by the pace angulation
(Peabody 1959). This is defined as the angle formed by three continuous front or
hind footprints (Peabody 1959) (Fig. 10.1). Pace angulation is dependent on body
size and speed, but limb posture is the key factor in determining pace angulation. As
exemplified by neoichnologic studies, values lower than 107°, in fact, indicate the
producer performed a sprawled locomotion while values greater than 138° indicate
erect locomotion (Kubo and Benton 2009).
By analyzing a large dataset of Late Paleozoic to Early Mesozoic tetrapod track-
ways, the authors found a clear distinction between Permian and Triassic samples
Fig. 10.1 Pace angulation, the angle formed by three consecutive footprints, can be used as a
proxy for trackmaker’s posture. Using the measuring method shown in (a), Kubo and Benton
(2009) discovered that the shift from a sprowler posture (b) to a more erect one (c) occurred across
the Permian-Triassic boundary, and may thus be related with the profound impact of the devastat-
ing mass extinction event. Modified from Kubo and Benton (2009)
when median values rise from 87.5° (Late Permian) to 139.5° (Early Triassic),
whereas within those epochs no significative change in the distributions of pace angu-
lation could be found. All tetrapods, whether synapsids or archosaurs, show the same
postural shift across the Permo-Triassic boundary and once the posture had changed,
there was no significant change in mean pace angulation through the Triassic. This
shift was mostly caused by the increase in the number of chirotheriid trackways from
the Early Triassic. The ichnofamily Chirotheriidae (Abel 1935) is a form family com-
prising the ichnogenera Chirotherium Kaup, 1835; Brachychirotherium Beurlen,
1950; Isochirotherium Haubold, 1971b; Synaptichnium Nopcsa, 1923;
Parasynaptichnium Mietto, 1987, and Protochirotherium Fichter and Kunz, 2004.
These have been consistently attributed to archosaurs, usually to crurotarsans (pseu-
dosuchians) and basal archosauriforms (Krebs 1965; Haubold 1967, 1971a, 1971b,
1984, 1986, 1999; 2006; Lockley and Meyer 2000; Demathieu and Demathieu 2004;
Gand et al. 2010; Bowden et al. 2010; Heckert et al. 2010; Lucas and Heckert 2011;
Desojo et al. 2013; Bernardi et al. 2015) and show manus (fore foot) and pes (hind
foot) imprints with a compact anterior digit group I–IV and a postero-laterally posi-
tioned, strongly reduced digit V, which are apomorphies of Archosauromorpha
(Nesbitt 2011). Thus it is clear that archosaurs (and to a lesser extent, therapsids) were
responsible for the high pace angulation of Triassic trackways. Footprints made by
amphibians, procolophonids, and small basal diapsids indicate retained sprawling
limb posture (Haubold 1971a, b). The abundance of trackways from the Permian up
to the Triassic provides resolution that the rare skeletal fossil tetrapods of this age do
not. In addition to providing a description of the pattern of this event, dating indicates
that the shift was accomplished already 6 myr after the profound Permo-Triassic mass
extinction; this is considerably earlier than the 15–20 myr suggested by the study of
locomotory evolution based on skeletal fossils (Charig 1972; Kemp 1982; Bonaparte
1984; Parrish 1987; Kubo and Benton 2009). The ichnologic record also supports a
late Permian-Early Triassic radiation of archosauriforms not well documented by
skeletal material. By analizing several chirotheriid footprints from the upper Permian
of the Southern Alps (Italy), Bernardi et al. (2015) showed that archosauriforms had
already undergone substantial taxonomic diversification by the late Permian, widen-
ing the geographical distribution of this clade before and after the P-T boundary
(Bernardi et al. 2015). Furthermore, analysis of body size, as deduced from track
length, allowed formulating a new hypothesis on the response of archosauriforms to
the end-Permian events, proposing that their body size did not change significantly
from the late Permian to the Early Triassic (Bernardi et al. 2015).
10.2.2 The Origin of Dinosauromorphs
In vertebrate ichnology synapomorphy-based studies are still rare. However, it now

appears possible to integrate this discipline into paleobiologic studies if rigorous
descriptions are coupled by thoughtful character-based evaluation of potential
trackmakers. When synapomorphic characters are found, potential trackmakers can
be constrained considerably, allowing full hypothesis testing. More simply, each
find that reveals new morphotypes of trackways or skeletal remains of limbs/feet
showing apomorphic characters, helps constrain possible track-trackmaker correla-
tions more accurately. Using this approach, tracks and trackways become “biologi-
cally informative” and can thus be of interest for paleobiologic analysis. The
synapomorphy-based technique focuses on the identification of osteologic-derived
character states in the tracks that result from the impression of corresponding (syn-
apomorphic) characters in the autopodia of the trackmaker. These characters allow
the recognition of a particular clade independent from the defined body-fossil dis-
tributions (see Olsen et al. 1998; Carrano and Wilson 2001; Wilson 2005). This
approach has been applied only recently (e.g., Olsen 1995; Olsen and Baird 1986;
Olsen et al. 1998; Wilson and Carrano 1999; Carrano and Wilson 2001; Haubold
and Klein 2000, 2002; Padian 2003; Wilson 2005; Wright 2005; Brusatte et al.
2010b; Kubo and Kubo 2013; Bernardi et al. 2015).
The most recent advances in the understanding of the origin of dinosaur lineages
are a good example of the potential for integrating ichnologic studies with cladistic
approaches. The resulting contribution is that, conceived in this way, vertebrate ich-
nology can give to a deeper understanding of patterns and processes in the evolution
of life. Dinosauromorpha is a clade including all taxa closer to dinosaurs than to the
other main archosaur clades (i.e., pterosaurs and crocodylomorphs).
Dinosauromorphs originated, and radiated in the Triassic, and several taxa are now
known that vary in size, feeding, and locomotory features, and that were spread over
most of western Pangea (Langer et al. 2013). However, both the timing and pattern
of the early radiation of this lineage are poorly understood (see Marsicano et al.
2015 for a recent update). The oldest known body fossils of dinosauromorphs are
silesaurids, which are Anisian in age (Nesbitt et al. 2010; Peecook et al. 2011;
Barrett et al. 2015). Silesaurids occupy a derived position within Dinosauromorpha
(Ezcurra 2006; Nesbitt et al. 2010; Nesbitt 2011), and therefore imply the presence
of other dinosauromorph groups such as lagerpetids, and non-silesaurid dinosauri-
formes before that time. No such Early Triassic groups are documented in the body
fossil record. However, the Early Triassic archosaur track record is rich (see Klein
and Haubold 2007 and references therein), raising the intriguing possibility that
dinosauromorph tracks can be recognized among the diverse ichnoassociations.
Rotodactylus, a common ichnogenus in the latest Olenekian–Ladinian worldwide
(Peabody 1948; Gand and Demathieu 2005; Gand et al. 2005; Klein and Haubold
2007), has been long attributed to basal dinosauromorphs (Haubold 1999; Haubold
and Klein 2002); however, a strict synapomorphy-based assessment was lacking.
Recently, Brusatte et al. (2010a) presented a formal analysis corroborating this
identification. Rotodactylus, in fact, possesses unequivocal features of dinosauro-
morphs, such as reduction of digits I and V and digitigrade posture. Moreover
Brusatte et al. (2010a, see also; Niedźwiedzki et al. 2013), described a new ichno-
genus named Prorotodactylus, discovered in the Early Triassic of Koszary and
Stryczowice, Poland. Prorotodactylus can be assigned to Dinosauromorpha thanks
to the presence of unequivocal synapomorphies such as the bunched metatarsus,
reduction of digits I and V, the posterior deflection of digit V, and the manifest digi-
tigrady. Brusatte et al. (2010a) suggested Lagerpeton as possible trackmaker, and
although this is (Niedźwiedzki et al. 2013) or might be (Langer et al. 2013) the case,
an attribution of Prorotodactylus to dinosauromorphs appears quite solid
(Niedźwiedzki et al. 2013; but see Klein and Niedźwiedzki 2012). The nearly com-
plete absence of autopodia in the skeletal record (with the notable exception of
Lagerpeton, Sereno and Arcucci 1994) prevents an evaluation of digit ratios across
other dinosauromorph taxa (see Langer et al. 2013 for a discussion) and enhance the
importance of the above cited ichnotaxa in exploring the early history of this group.
The synapomorphy-based identifications of Rotodactylus and Prorotodactylus shed
new light on the debate over the posture of the earliest dinosauromorphs; both show
that these early forms were much more digitigrade than any other closely related
archosaurs and that they walked on all fours (Fig. 10.2b). The ichnogenus
Sphingopus, which is found in numerous Olenekian to Middle Triassic sites all over
the world (Klein and Haubold 2007), shares the apomorphic characters described in
Prorotodactylus; however, the lack of associated manus imprints and high pace
angulation indicates that the trackmaker was walking bipedally. Furthermore,
Sphingopus is much larger than the other dinosauromorph tracks. The tendency
toward a full tridactyl morphology, with digit III being the longest, suggest that
Sphingopus isp. was produced by a dinosauromorph more derived than the Proroto
dactylus/Rotodactylus trackmaker, and therefore suggests that bipedality was
acquired and the trackmaker was closer to the origin of dinosaurs (Fig. 10.2c).
Fig. 10.2 Footprints attributed to dinosauromorphs and early dinosaurs can be integrated with
skeletal data to build a general timeline of early dinosaur history. Footprints, as those described by
Brusatte et al. (2010b), document the earliest phase of this radiation and predate timing of emer-
gence of the various groups (a). (b) Shows a likely reconstruction of an early dinosauromoph as
reconstructed on the base of Rotodactylus trackways. (a) Modified from Brusatte et al. (2010b), (b)
modified from Haubold (1999) and Niedźwiedzki et al. (2013)
The stratigraphically successive records of Prorotodactylus and Rotodactylus docu-

mented by Brusatte et al. (2010a) show that changes in foot anatomy were occurring in
dinosauromorphs during the Olenekian; the appearance of Sphingopus documents the
evolutionary shift from quadrupedality to bipedality, and from small to large size, in the
dinosaur lineage. Tracks, identified using synapomorphies, can thus provide new data
on the timing and pattern of character acquisition during early dinosauromorph radia-
tion (Fig. 10.2a). Such data are currently not visible in the body fossil record.
Furthermore, being older than indicated by the skeletal record, tracks might indicate
that the appearance of the dinosauromorph lineage can be linked with the Permian—
Triassic mass extinction event (Brusatte et al. 2010a; Niedźwiedzki et al. 2013).
10.2.3 The Origin and Early History of Dinosaurs
The vertebrate track record can be used to make inferences about the spatial and
temporal distribution of tetrapod taxa and can give reliable clues, although at a
coarse taxonomic level, into tetrapod diversity. If parsimoniously interpreted, it can
complement the skeletal record offering a “total evidence” approach to the study of
evolutionary patterns and timing of events.
The study of early Mesozoic vertebrate terrestrial ecosystems is a lively field in
both vertebrate paleontology and paleoichnology, mostly powered by the debates on
dinosaur origins and their early history. The appearance of dinosaurs during the
Triassic marked the onset of a faunal diversification that has dominated terrestrial
ecosystems for almost 230 Myr (Brusatte et al. 2008a). As discussed, the possible
contribution of track data to this and other taxon-based debates, depends on reliable
identification of dinosaur trackmakers.
Triassic tridactyl prints are traditionally assigned to Dinosauria (see Lucas et al.
2006 and reference therein). Supposed dinosaur tracks are recognized on the base of
few synapomorphies: (1) the prevalence of the digit II–IV group, (2) mesaxonic pat-
tern of foot structure, (3) digit I reduced and shifted backwards (and thus often not
preserved in tracks), and (4) tendency towards digitigrady (see also Thulborn 1990;
Haubold 1999; Brusatte et al. 2010b). However, this morphotype cannot be unam-
biguously assigned to Dinosauria using apomorphy-based identifications; a function-
ally tridactyl pes is found also in some non-dinosaurian dinosauriforms (Gauthier
1986; Brinkman and Sues 1987; King and Benton 1996; Novas 1992; Carrano and
Wilson 2001; Fraser et al. 2002; Dzik 2003; Marsicano et al. 2007; Nesbitt et al.
2007; Brusatte et al. 2010b). Furthermore, no synapomorphies of three-toed foot-
prints can discriminate among theropods, basal saurischian, and basal ornithischian
groups (Olsen et al. 1998). “Theropod-like” tracks can, however, be identified on the
basis of the following characters: (1) asymmetry of the track with angle between
digit III and II lower than III-IV, (2) digit III longer than IV > II, (3) sharp claw traces
on all digits, and (4) tip of digit II turned inwards (Bernardi et al. 2013). Lockley
(2001, 2007a, 2009) has also argued that narrowness and strong mesaxony distin-
guishes theropod tracks from those of tridactyl ornithischians. Nevertheless, no clear
synapomorphy can be described in tridactyl tracks to clearly distinguish between

dinosaurian- and non-dinosaurian-dinosauriforms. All Triassic functionally tridactyl
tracks with digitigrade pes produced by bipedal parasagittal posture should be loosely
attributed to an archosaur on the dinosaur line. Some derived characters, such as
those listed by Bernardi et al. (2013), can be used in support of an attribution to
Dinosauria, but a “gray zone” will always exist.
Debates on dinosaur origin thus focus around three main topics: (1) timing of
appearance, (2) pattern and pace of early diversification (body size and biogeogra-
phy), and (3) dynamics of replacement of crurotarsan archosaurs by dinosaurs. If the
first relies on new discoveries of older specimens and reliable dating of the dinosaur-
bearing formations, the second refers to dinosaurs having experienced a rapid diversi-
fication or having a deeper origin (“slow fuse”) not documented (with the possible
notable exception of Nesbitt et al. 2013) by current knowledge of the fossil record.
The last needs data from mixed assemblages to test for habitat preferences and niche
competition, and the in situ nature of footprints can be of great value in this respect.
10.2.3.1 Timing of Appearance
The exact timing of dinosaur origins remains unclear (Benton 1983; Brusatte et al.
2008a). The oldest record of dinosaurs based on skeletal remains is from the Carnian
of Argentina (Benton 1983; Sereno and Arcucci 1994; Sereno 1999; Irmis et al.
2007; Brusatte et al. 2010b; Langer et al. 2010; Ezcurra 2010; Martinez et al. 2011),
although Nesbitt et al. (2013) reported a possible earlier dinosaur from the Anisian of
Tanzania. By the Carnian-Norian transition, all three major dinosaur lineages
(Ornithischia, Sauropodomorpha, Theropoda) are documented (Brusatte et al. 2010b;
Irmis et al. 2007; Langer et al. 2010). This forces researchers to focus mostly on the
still poorly known Middle Triassic record. Furthermore, ghost lineages inferred from
phylogenetic analyses predict that Dinosauria and its immediate stem lineage should
be as old the early Middle Triassic (Sereno and Arcucci 1994; Nesbitt et al. 2010).
Early Triassic tracks previously attributed to dinosaurs have been reconsidered as
partially or badly preserved “chirotheroids” (King and Benton 1996; see also Cavin
et al. 2012). Although track-bearing formations cannot be always confidently dated,
functionally tridactyl footprints are documented in the Middle Triassic of France
(Demathieu 1989; Lockley and Meyer 2000), Germany (Haubold and Klein 2000,
2002), Italy (Avanzini 2002), and Argentina (Marsicano et al. 2007 and references
therein), and have been generally attributed to dinosauromorphs. Dinosaur tracks
have been reported from the Middle Triassic Los Rastros Formation (Argentina) by
Marsicano et al. (2004, 2007; see also Marsicano and Barredo 2004; Melchor and
Valais 2006) but have been considered ambiguously identified by others (Langer
et al. 2010; Irmis 2011). Therefore the ichnologic record, although failing to defini-
tively prove a Middle Triassic origin for dinosaurs, clearly indicates the presence of
a diverse fauna of dinosauromorphs with functionally tridactyl digitigrade pedes, and
possibly true dinosaurs during the Middle Triassic (Fig. 10.3), a condition that the
body-fossil record is only recently beginning to document (Nesbitt et al. 2013).
Fig. 10.3 A calibrated phylogeny of early dinosaurs and sister taxa based on both skeletal and
ichnological data. The possible presence of dinosaurs in the Middle Triassic as testified by contro-
versial footprints have recently received support from the description of Nyasasaurus which is
considered either within, or sister-taxon of, Dinosauria. Modified from Marsicano et al. (2007) and
Nesbitt et al. (2013)
10.2.3.2 Early Diversification
Although the first unequivocal dinosaur body fossils are known from the Carnian
(Brusatte et al. 2010b; Langer et al. 2010; Nesbitt et al. 2013), Brusatte et al. (2008a,
2010b) demonstrated that dinosaurs increased their diversification and their
morphologic disparity only from the Norian; dinosaurs experienced a burst of
diversification only after the Triassic–Jurassic transition.
At the end of the Triassic the dinosaurs experienced an increase in maximum

body size, later expressed as an increase in average body size (Sookias et al. 2012;
Turner and Nesbitt 2013). Late Triassic tridactyl footprints are well known from
USA, Europe, South America, Australia, and Africa (e.g., Tucker and Burchette
1977; Biron and Dutuit 1981; Mietto 1988; Olsen and Baird 1986; Thulborn 2000;
Gatesy 2001; Knoll 2004; Milàn and Gierliński 2004; Weishampel et al. 2004;
Lockley and Eisenberg 2006; Silva et al. 2007; Meyer et al. 2013). The Triassic
record of large tridactyl tracks (e.g., Thulborn 2003; Bernardi et al. 2013; Meyer
et al. 2013) has invalidated the hypothesis of Olsen et al. (2002) of a dramatic
increase in size at the Triassic-Jurassic boundary. However, the increasing abun-
dance toward the end of the Triassic of medium- and large-sized theropod tracks
(Eubrontes and Grallator above all) marked a crucial phase in dinosaur history,
when the main radiation or diversification of theropods and the evolution of large
species got underway during the Late Triassic (Lockley and Hunt 1995a).
Furthermore, if the attribution of the largest Late Triassic tracks is correct, the esti-
mated body length, calculated according to Thulborn’s (1990) equations
(h = 3.06 × FL1.14 and L = 4 h), would be more than 5 m. Their estimated body mass
value, following Weems (2006), would have been about 200 kg (Bernardi et al.
2013). This size is not known from dinosaur skeletal remains of Carnian age (Benton
2006; Brusatte et al. 2010b) and is approximated only by the Norian theropod dino-
saur Liliensternus (Cuny and Galton 1993; Lucas et al. 2006).
Tracks are also a valuable tool that gives us insight into the geographic distribution
of the first dinosaurs. The earliest skeletal remains of dinosaurs are from southern
Pangaea and the oldest possible dinosaur tracks are from Argentina. The earliest
dinosaur from northern Pangaea has recently been dated as “middle” Norian of west-
ern North America (Nesbitt et al. 2009). This has led to the view that the major dino-
saurian clades (Ornithischia, Sauropodomorpha, Theropoda) may have originated in
southern Pangaea (Brusatte et al. 2010b; Langer et al. 2010). The paleogeographic
importance of this distribution, however, is compromised by the absence of well-
preserved skeletal remains of Carnian age from northern localities (northern Africa,
Europe, North America) (Langer 2005; Nesbitt et al. 2007; Martinez et al. 2011).
Tracks, however, indicate that dinosaurs were already present by the Late Carnian in
northern Pangea (Bernardi et al. 2013, Fig. 10.4). With the exclusion of the possible
earliest dinosaur Nyasasaurus (Nesbitt et al. 2013), this would imply a nearly contem-
poraneous appearance of dinosaurs on all of Pangea (Bernardi et al. 2013).
10.2.3.3 Dynamics of Replacement
In the past, the Late Triassic was described as the period when the faunas dominated
by crurotarsan archosaurs were competitively replaced by those dominated by dino-
saurs (Bakker 1972; Charig 1984). A growing consensus now suggests that dino-
saurs were initially rare components of Triassic terrestrial ecosystems and that their
ascent was gradual (Brusatte et al. 2008b). It was only after the Triassic–Jurassic
transition that dinosaurs became dominant in terrestrial vertebrate ecosystems
Fig. 10.4 The ichnological record provides valuable data on both temporal and spatial distribution
of basal dinosaurs. Tracks, for example, indicate the presence of archosaur faunas, which include
dinosaurs, during the early Late Triassic both in Laurasia and Gondwana. This pattern is not (yet)
documented by body-fossil data. Modified from Brusatte et al. (2010a, 2011)
worldwide (Benton 1983; Sereno 1999 see also Brusatte et al. 2010b). The body
fossil record has shown that in the Late Triassic, during the first 25–30 million years
of dinosaur history, crurotarsans had double the variety of body plans, lifestyles,
and diets in comparison with contemporary dinosaurs (Brusatte et al. 2008a, 2010b,
2011; see also, Benton 1983; Sereno 1999). In addition to crurotarsans, Triassic
dinosaurs lived together with dicynodont synapsids (Dzik et al. 2008) and coexisted
with dinosauromorphs for at least 20 million years (Irmis et al. 2007). The scarcity
of Early to Middle Triassic terrestrial body fossils, however, leaves open questions
on taxonomic abundance, composition, diversity, and regional specificity of such
dinosaur communities (Irmis et al. 2007; Dzik et al. 2008; Nesbitt et al. 2009;
Ezcurra 2010; Irmis 2011). The track record can be integrated into this picture.
As a general depiction, while the upper Triassic ichnologic record is character-
ized by dinosaur, dinosauromorph, crurotarsan archosaurs, and nonmammalian the-
rapsid tracks (Marsicano and Barredo 2004; Bernardi et al. 2013; Lockley and
Meyer 2000; Klein and Haubold 2007; Lucas and Tanner 2007), the Lower Jurassic
record is dominated by saurischian dinosaurs (Lockley and Hunt 1995a; Lockley
and Meyer 2000; Olsen et al. 2002; Avanzini et al. 2006; Avanzini and Petti 2008).
Well-diversified tetrapod ichnoassemblages are known from the Middle to Late
Triassic of both southern and northern Pangaea (see Bernardi et al. 2013). Olsen and
Huber (1998) reported the co-occurence of Apatopus, cf. Brachychirotherium, and
small bipedal and tridactyl forms, possibly produced by “dinosaurian” trackmakers

in the Carnian of North Carolina, while Cameron and Wood (2003, 2004) described
a rich association composed of Atreipus, Brachychirotherium, Grallator, and
Rhynchosauroides from the Carnian of Canada. Similar assemblages are ubiquitous
in the western USA (Lockley and Hunt 1995a). Marsicano and Barredo (2004)
reported the presence of crurotarsal archosaurs, dinosaurs (sauropodomorphs and
theropods) and nonmammalian therapsids (small cynodonts and large dicynodonts).
Nicosia and Loi (2003) described a rich Carnian ichnoassociation from Italy with
crurotarsan and dinosaurian tracks. D’Orazi Porchetti et al. (2008) and Bernardi
et al. (2013) document the co-occurrence of large- and middle-sized dinosaurs,
dinosauriforms, and crurotarsans in the late Carnian (Tuvalian) of the southern
Alps. Therefore, the record of ichnofaunal assemblages composed of large and
small dinosaurs, dinosauriforms and non-dinosaurian archosaurs and their co-
occurrence with dinosauriforms and crurotarsal archosaurs provides support to the
non-abrupt replacement of the more archaic faunas by dinosaur-dominated ones.
Furthermore, the co-occurrence of dinosaurs, dinosauriforms, and crurotarsal archo-
saurs in the Middle to Late Triassic (Ladinian to Late Carnian) documents a pro-
longed coexistence of different groups of archosaurs, even in the same environment.
This provides support to the view of crurotarsan-dominated faunas being neither
rapidly outcompeted nor quickly opportunistically replaced by dinosaurs, but rather
substituted by a more gradual process of ecologic replacement (Irmis et al. 2007;
Nesbitt et al. 2010; Brusatte et al. 2010b; Martinez et al. 2011).
10.2.4 A Look into Sauropod Paleobiology
One key feature of tracks that neatly distinguishes them from body fossils is that they
cannot be transported post-mortem, in this case, “post-emplacement”. Tracks thus
provide a direct indication of environmental and/or habitat preferences of the pro-
ducer (Thulborn 1982; Lockley 1991; Wilson and Carrano 1999; Carrano and Wilson
2001). To better understand habits, locomotory attitudes or geographic distribution
of particular groups, it is therefore useful to integrate body fossil findings with track
data. A good example in this respect is provided by the sauropod dinosaur record.
Numerous studies of locomotory dynamics in this group attribute different foot-
print morphotypes to distinct trackmakers. Many authors (Farlow et al. 1989; Farlow
1992; Lockley et al. 1995a; Moratalla et al. 1995; Wilson and Carrano 1999;
Lockley et al. 2002, 2005; Day et al. 2002a; Henderson 2002) have observed that
sauropod trackways could be essentially subdivided into two broad categories: “nar-
row gauge” and “wide gauge” (Fig. 10.5). Sauropod trackways can also show (or
lack) claw and phalangeal impressions in the manus (Farlow et al. 1989; Lockley
et al. 1995a; Wilson and Carrano 1999; Day et al. 2004). “Wide-gauge” trackways
are those in which manus and pes prints are well separated from the midline, and
which frequently lack pollex claw impressions (e.g., Brontopodus Farlow et al.
1989); “Narrow gauge” trackways have manus and pes prints that approach or inter-
sect the trackway midline, and have pollex claw impressions (e.g., Parabrontopodus
Fig. 10.5 Many authors (see text for discussion) have observed that sauropod trackways could be
essentially subdivided into two broad categories: “narrow gauge” (a) and “wide gauge” (b). These
reflect the postural difference between non-titanosaur (c, here represented by Camarasaurs—pel-
vic girdles and hindlimbs in anterior view) and titanosaur sauropods (d, here Opisthocoelicaudia).
This inference allowed full integration of ichnogical and skeletal data, which contributed to a bet-
ter understanding of sauropod paleobiology. Modified from Lockley et al. (1995b), and Wilson and
Carrano (1999)
Lockley et al. 1995a). Lockley (1999, 2001, 2007a) also argued that there is a posi-
tive correlation between narrow gauge and small manus forms, and between
wide-gauge and large manus forms. Furthermore, large and small manus forms have
been independently identified among the various chirotherian ichnotaxa (Lockley
2007a). Such differences in turn speak to changes in the position of the center of
gravity in various archosaur clades.
Ichnologic investigations suggest that the transition from narrow- to wide-gauge
occurred during the Kimmeridgian-Tithonian (Wilson and Carrano 1999). “Wide-
gauge” trackways show a trend towards an increase of manus print dimensions
through time. This feature and the age of “wide-gauge” trackways closely match the
evolutionary trend seen in the sauropod body fossil record. Skeletal remains record a
gradual shift of the center of gravity toward the front, with the consequent growth of
the forelimbs in response to the greater weight they had to support. This evolutionary
change coincides with the appearance of Titanosauriformes Salgado, Coria and
Calvo 1997 (including Brachiosauridae Riggs 1904 and Titanosauria Bonaparte and
Coria 1993) (Wilson 1995; Upchurch et al. 2004), animals that would have produced
tracks with larger manus impressions. The lack of digital impressions in the manus
ichnites of the “wide-gauge” trackways could also be referred to the appearance of
Titanosauriformes; indeed both Brachiosauridae and Titanosauria are characterized
by the progressive reduction and loss of phalanges and claws in the manus (Wilson
and Sereno 1998; Upchurch et al. 2004; Apesteguía 2005). In particular, Titanosauria
probably walked only on the distal parts of their metacarpal bones (Salgado et al.
1997; Wilson and Sereno 1998). It is thus difficult to envisage digital or claw marks
produced by individuals belonging to Titanosauriformes. Wilson and Carrano (1999)
argued that hindlimb modifications, such as outwardly angled femora, offset knee
condyles, and a more eccentric femoral midshaft cross-section, could be responsible

for wide-gauge tracks. All these features are synapomorphies of titanosaurs, and thus
support the hypothesis that they were the producers of wide-gauge trackways.
Temporal and geographic distribution of titanosaurs is coherent with this hypothesis;
“wide-gauge” trackways predominate during the Cretaceous and are found world-
wide. After Wilson and Carrano’s (1999) proposals, other studies have cast some
doubt on this correlation. D’Emic (2012) suggested that some Late Jurassic non-
titanosaurs might have femoral cross-sections similar to those of titanosaurs.
Henderson (2006) proposed that all large sauropods (over 12.6 tonnes) would have
been constrained to adopt a wide-gauge stance in order to maintain stability during
locomotion; trackway gauge could also be influenced by the degree of lateral motion
of the trackmaker (Carpenter 2009). Recent discovery of sauropod trackways from
Switzerland, Morocco (both of Jurassic age), and Spain (Cretaceous age) show that
single sauropod trackmakers were able to adopt both narrow and wide gauge as testi-
fied by the occurrence of differently spaced tracks along single trackways (Marty
et al. 2010; Castanera et al. 2012a, b). Furthermore, wide-gauge trackways could
have appeared more than once in sauropod evolution. Wide-gauge trackways are not
exclusive to titanosauriform sauropods and caution must be applied ascribing a track-
way to an ichnotaxon on the basis of this character (Castanera et al. 2012a, b).
Sauropod tracks have provided important clues for the understanding of the
environmental preferences of this clade. Based on the analysis of a dataset of
sauropod trackways, Lockley et al. (1995b) found a good correlation between
sauropod tracksites and coastal environments, and noted that they are primarily
associated with tropical and subtropical paleolatitudes (mean average Northern
Hemisphere = 25°), and semi-arid or seasonal climatic carbonate depositional
environments (lacustrine or marine carbonate platform). Butler and Barrett
(2008) partially confirmed these results: the percentage of Cretaceous sauropod
tracks in coastal environments is remarkable (32 %) if compared with the con-
temporaneous sauropod body fossil record (7 %). Nevertheless, statistic analysis
of the ichnologic data does not indicate that sauropod trackways are overrepre-
sented if compared to those of other herbivorous dinosaurs. In contrast, the body
fossil record supports previous hypotheses of a preference for inland settings
(e.g., Lehman 1987; Hunt et al. 1995a). However, it should be noted that these
results might be hindered by preservational biases that are not yet fully under-
stood (Mannion and Upchurch 2010). Using a new large data set of sauropod
skeletal remains, Mannion and Upchurch (2010) found a significant positive
association between non-titanosaurs and coastal environments, and between
titanosaurs and inland environments. Their analysis of track records also high-
lighted a positive correlation between narrow-gauge trackways and coastal envi-
ronments, and wide-gauge trackways and inland environments; this reinforced
the hypothesis of Wilson and Carrano (1999) that wide-gauge trackways were
produced by titanosaurs. The hypothesis of Mannion and Upchurch (2010) has
been partially put into question by isolated findings of wide-gauge trackways
preserved in coastal environments (e.g., Castanera et al. 2011).
10.2.5 Clues About Theropod Paleobiology
Most famously among track features, trackways allow estimation of speed

(Alexander 1976, 1989; Thulborn 1981, 1990). In fact, debates about speeds attained
by dinosaurs led to a revival of tetrapod ichnology (Alexander 1976; Lockley 1998).
Early estimates of rapid progression of large dinosaurs were proposed in support of
the idea that dinosaurs were highly active and possibly warm-blooded (Russell and
Belland 1976; Czerkas and Olson 1987); however, later studies based on larger
samples and updated formulae concluded that most medium- to large-sized thero-
pods walked fairly slowly (Farlow 1981; Thulborn 1990). Estimates of the speeds of
theropod dinosaurs are based on the formula of Alexander (as corrected by Thulborn
1990); speed (v, velocity) is calculated on the basis of stride length (SL) where
v ≈ 0.25 g0.5 SL1.67 h−1.17, where hip height (h) of a trackmaker can be estimated from
footprint length (FL): h ≈ 8.60 FL0.85 (Thulborn 1984; Thulborn and Wade 1984).
This formula was successively slightly modified by Weems (2006) to estimate basal
theropod dinosaur speeds (v = 0.16 SL2 h−1.5 g0.5). The same formula can be used for
any bipedal dinosaurs (see Currie 1983, 1995), while no similar, reliable formulae
are available for quadrupedal dinosaur taxa. Using musculoskeletal modeling and
Computer Aided Engineering, Mallison (2011) questioned the validity of the
Alexander formula to assess speed in dinosaurs without knowing the frequency of
the stride. He suggested that speeds calculated from tracks should be better consid-
ered as the slowest speeds at which the animals moved.
Tracks and trackways have also been long used to establish dynamics of foot
movement (Avanzini 1998; Gatesy et al. 1999; Gatesy 2001, 2003; Manning 2004;
Milàn et al. 2004, 2006; Avanzini et al. 2012; Ellis and Gatesy 2013), locomotion
(Alexander 1976, 1985, 1989; Thulborn 1981, 1990) and contributed in the debate
about theropod posture (Lockley 1998). The high pace angulation exhibited by the-
ropod trackways, for example, shows that theropods placed their hind feet close to,
or on, the midline (Alexander 1976, 1985). This optimizes the efficiency by reduc-
ing the energy lost through lateral displacement of the center of gravity (Thulborn
1981, 1990), and is characteristic of a highly specialized locomotory control system.
Lockley (2007a) regarded such narrow trackway configurations as a manifestation
of an intrinsic morphodynamic organization in theropods. Furthermore, saurischi-
ans in general had narrow bodies, narrow feet, and other laterally compressed skel-
etal elements, although this contrasts with the wide-gauge trackway configurations
and wide bodies and feet seen in derived saurischians such as the titanosaurs.
Furthermore, Day et al. (2002b) showed that large theropods were able to change
their speed instantaneously, highlighting their high maneuverability and balance
control.
Theropod trackways do not typically include hand imprints, which is again an
expression of a posterior center of gravity (Lockley 2007a). Only when the trunk was
lowered toward a substrate, as in a crouched posture, could the hands potentially pro-
duce impressions. Crouching has been demonstrated by a few extraordinary discover-
ies of impressions of autopodia associated with metatarsals and pelvic girdle imprints.
Milner et al. (2009) reported unambiguous theropod manus impressions associated
with a crouching trace that indicate the avian orientation of the manus (with a medially
facing palm) evolved early within the history of Theropoda (see Gierliński et al. 2009
and references therein for further examples). The crouching trace was registered when
the animal rested on the substrate in a posture similar to that of modern birds; the traces
include well-defined impressions made by both hands, the ischial callosity, pedes, and
the tail. This trace constitutes evidence that an Early Jurassic theropod expressed two
bird-like features: anatomic restriction to a palms-medial manual posture, and sym-
metric leg positions while resting (Fig. 10.6). Inferences that theropods displayed
behavior analogous to modern birds are intriguing but often too speculative. Recently
discovered Cretaceous theropod tracks in Colorado provided extensive physical evi-
dence of substrate scraping behavior by large theropods; they were considered as com-
pelling evidence of “display arenas” or leks, and consistent with “nest scrape display”
behavior among many extant ground-nesting birds (Lockley et al. 2016).
Another iconic example of the impact that ichnology has had on postural biome-
chanic reconstructions is the case of the orientations of dinosaur trunks and tails.
Analysis of trackways have clearly shown that most dinosaurs walked with their
tails carried well off the ground, so tail-drag traces are rare (Irby and Albright 2012;
García-Ramos et al. 2004, 2006; Platt and Hasiotis 2008; Kim and Lockley 2013).
The presence of tail traces, therefore, implies either a pathologic condition of the
Fig. 10.6 Footprints provide clues to the individual behavior of trackmakers. These crouching
traces described by Milner et al. (2009), for example, demonstrate that avian orientation of the
manus, with medially facing palms, evolved by the Early Jurassic within Theropoda. Such conclu-
sion would have never been reached on the base of skeletal record alone. Modified from Milner
et al. (2009)
trackmaker or an atypical mode of locomotion because dinosaurs used their tails for
balance on land (Galton 1970; Farlow et al. 2000; Fujita et al. 2012).
Tracks can also be used to assess the effects of traumas and illness in dinosaur
bones and therefore the evolution of phatologic behavior. Tracks and trackways
indicative of pathologies (e.g., fractures in pedal phalanxes) are relatively frequent,
especially in middle to large-sized theropods (Tanke and Rothschild 1997, 2002;
Rothschild et al. 2001; Rothschild and Tanke 2005). Many of the trackways described
refer to large carnivorous dinosaurs that show missing or curled digits (Lockley
1991; Tanke and Rothschild 1997). Footprint and trackway evidence of limping the-
ropods (Ishigaki 1986; Dantas et al. 1994; Lockley et al. 1994b) suggests injury or
arthritis. Foot pathologies in large bipedal dinosaurs however have also been used to
infer more active life-styles or natural fragility of the narrow, protruding digits (Laws
1997; Tanke and Rothschild 1997). For other good examples of pathologic tracks,
see Abel (1935), Thulborn (1990) and Avanzini et al. (2005).
Tracks have also helped unveil another aspect of dinosaur behavior: their ability
to swim. No skeletal remains can help addressing this question, and tracks attribut-
able to swimming theropods have been reported, among others, by Coombs (1980),
Milner et al. (2006), Ezquerra et al. (2007) and Xing et al. (2013a) (similar reports
are available also for other dinosaur taxa; e.g., Currie 1983; Fujita et al. 2012).
Theropods could float, as demonstrated by clawing traces left in the sediment as
they swam (Ezquerra et al. 2007); the trackway shows that this theropod used a
pelvic paddle motion, i.e. they were powered by alternating movements of the two
hind limbs, like in living aquatic birds, and swam with amplified asymmetric walk-
ing movements to maintain its direction while swimming perpendicular to a water
current (Ezquerra et al. 2007). The track record, therefore, has shown that dinosaurs
(mostly theropods) colonized areas that included aquatic environments and were
therefore probably able to swim.
10.2.6 Evidence for Gregariousness
Gregariousness is a form of social interaction between individuals that probably

evolved to increase the fitness of single individuals within a group (Alexander
1974). Gregariousness allows greater defense of individuals from predators,
increases effective access to food, efficiency of breeding, effective migration, and
enhances the probability of safely raising the young (Currie and Eberth 2010). In
order to maintain group cohesion, individuals within a group must synchronize their
behavior in terms of time spent resting, foraging, and moving (Conradt 1998).
Tracksites, together with nesting sites, provide some of the most reliable proof
about the gregariousness in extinct animals, in particular among dinosaurs (Ostrom
1972; Currie 1983; Lockley et al. 1986; Farlow et al. 1989; Horner and Gorman 1990;
Carpenter 1999). Fossil trackways made by multiple individual animals can provide
important qualitative and quantitative information about their social behavior.
Evidence for gregariousness in dinosaurs is difficult to gain from the body fossil
record, and only bone bed assemblages may reflect such group behavior. Bone beds
are concentration of bones in a localized area or stratigraphically limited sedimentary
unit, in which the bones belong to more than one individual (see Rogers et al. 2007
for more details). Significant work on bone beds assemblages has provided compel-
ling evidence on the gregarious habits of some ornithischian dinosaurs and the coex-
istence of multispecific individuals in the same habitat (Rogers 1990; Varricchio and
Horner 1993; Mathews et al. 2009; Zhao et al. 2007).
In the study of dinosaur tracks, gregariousness may be deduced by the recognition
of some peculiar features, primarily by the occurrence of multiple parallel trackways,
exhibiting a preferring directionality (Ostrom 1972; Currie 1983; Lockley 1991).
However, there is no arbitrary minimum number of adjacent parallel trackways for
indicating gregarious behavior (Lockley and Matsukawa 1999). The presence of
many parallel trackways does not necessarily constitute proof of a herding behavior,
and data should be critically examined: trackways could have been made by animals
that have walked alone at different times, and selected the same direction for the
presence of a physical constraint, such as narrow river valley, shoreline, or the banks
of a river (e.g., Ostrom 1972). Depths of the tracks in each trackway must be taken
into consideration; similar-sized dinosaurs, walking contemporaneously on a sub-
strate characterized by similar physical properties, probably produced parallel track-
ways with comparable depths. By contrast, parallel trackways with different track
depths suggest that the animals walked on that substrate at different times, perhaps
months, days, or even years later one from the other. A useful parameter to recognize
gregariousness from track data is the intertrackway spacing (i.e., an index of regular-
ity of spacing between parallel trackways; Lockley 1989; Lockley et al. 1998). In
recent years, several trackways characterized by regular intertrackway spacing have
been reported (Lockley et al. 1986, 2006b; Farlow et al. 1989; Lim et al. 1989; Li
et al. 2008), suggesting animals that walked shoulder to shoulder, and thereby sub-
stantiating the hypothesis of their social behavior. Parallel trackways curving or
swerving in unison could also provide evidence of herd habits among the trackmak-
ers (Lockley and Hunt 1995a, b; Lockley and Matsukawa 1999).
Such ichnologic evidence for gregarious behavior in dinosaurs has been
reported from Jurassic and Cretaceous sites in North America (Currie 1983,
1995; Lockley et al. 1983, 1986, 1992a, 1998; Farlow 1987; Farlow et al. 1989;
Pittman and Gillette 1989; Hawthorne 1990; Carpenter 1992; Barnes and Lockley
1995; Lockley and Hunt 1995a, b; Cotton et al. 1998; Kvale et al. 2001;
Schumacher 2003; Myers and Fiorillo 2009), South America (Leonardi 1989;
Lockley et al. 2002; Fig. 10.7), East Asia (Lim et al. 1989; Matsukawa et al.
1997, 2001, 2005; Lockley et al. 2002, 2006b, 2012a, b; Hwang et al. 2004; Li
et al. 2008; Xing et al. 2013a), Europe (Kaever and Lapparent 1974; Delair 1981;
Agirrezabala et al. 1985; Viera and Torres 1992; Lockley et al. 1995c; Moratalla
et al. 1997; Day et al. 2002a, 2004; Piñuela et al. 2002; Pérez-Lorente 2003;
García-Ramos et al. 2004; Clark et al. 2005; Vila et al. 2008; Castanera et al.
2011, 2013, 2014; García-Ortiz and Pérez-Lorente 2014), Africa (Lingham-
Soliar et al. 2003) and Australia (Thulborn and Wade 1979, 1984).
Fig. 10.7 Footprints

provide clues on the social
behavior of dinosaurs.
Here spectacular parallel
trackways at the Humaca
dinosaur tracksite
(Chuquisaca, Bolivia)
interpreted as evidence of
gregarious behavior by
Lockley et al. (2002).
Direction of progression is
towards the right of
picture. From Lockley
et al. (2002)
Evidence suggests that large herbivorous dinosaurs, especially sauropods and

ornithopods often traveled in herds, and that both groups acquired gregarious ten-
dencies early in their histories (see Lockley and Matsukawa 1999 and Myers and
Fiorillo 2009 and references therein). Sauropod trackways indicate groups of 40 or
more individuals (Day et al. 2002a, 2004) and show either mixed-age or are age-
segregated ichnologic assemblages (Lockley et al. 1986, 1995c, 2002; Castanera
et al. 2011). The co-occurrence of adult and juvenile trackways may suggest pro-
tracted parental care far beyond the hatchling stage of development; groups of
small-sized trackways may be interpreted as herds of juveniles or (as in the case of
archipelago paleoenvironments) as the presence of dwarfed populations (Benton
et al. 2010). Few tracksites exhibit parallel trackways of different species or groups
(e.g., Day et al. 2002a, 2004).
Much evidence of possible social behavior in other groups of herbivorous dino-
saurs, such as ankylosaurs (McCrea and Currie 1998; McCrea 2000) and ceratop-
sians (Lockley and Hunt 1995b) has now also accumulated. Evidence of
gregariousness in carnivorous dinosaur trackways, particularly non-avian thero-
pods, is less common than in other groups of dinosaurs, and the interpretations of
such evidence remain more controversial. Some researchers support this hypothesis
(Lockley and Matsukawa 1999; Carvalho and Pedrão 2000; Smith et al. 2002;
Lingham-Soliar et al. 2003; Clark et al. 2005; Barco et al. 2006; Li et al. 2008;
Currie and Eberth 2010; McCrea et al. 2014) while others prefer to consider track-
way data as coincidental instances of normally solitary individuals converging on a

common point (e.g., food source, Roach and Brinkman 2007).
10.2.7 On the Tracks of Birds
Birds are among the most diversified and thoroughly analyzed vertebrate groups, but
both their origin and evolution remain controversial. The skeletal record of birds is
strongly biased by the nature of bones that, in this group, are mostly small and deli-
cate so that limbs are commonly lost in taphonomic decay processes (Davis and
Briggs 1998; Brand et al. 2003). This makes the bird fossil record highly incomplete,
and only a multidisciplinary approach, including paleornithology, paleoichnology,
and molecular studies, can help in the understanding avian origins and radiation.
The skeletal record indicates an initial avian radiation around the Jurassic-
Cretaceous boundary (about 145 million years ago) and an early Paleogene diversi-
fication for modern birds (Neornithes) (e.g., Clarke et al. 2005; Dyke and Kaiser
2011; Brocklehurst et al. 2012; Ksepka and Boyd 2012). Molecular studies and
recent combined molecular and morphologic analyses, instead, indicate that modern
birds began radiating in the Early Cretaceous (Cooper and Penny 1997; Haddrath
and Baker 2012; Jetz et al. 2012; Lee et al. 2014).
Tracks similar to those of modern birds (i.e., shorebirds, small ducks, small herons,
roadrunners) are only 15-20 million younger than the oldest avian skeletal remains,
which is in agreement with molecular data (Brown et al. 2008). In contrast, the oldest
body-fossil records of anseriforms and possibly charadriiforms date to the latest
Cretaceous (ca. 70 million years ago), and are even younger for ciconiiforms and cucu-
liforms (Lockley and Harris 2010; Fig. 10.8). This could be explained through the fol-
lowing two hypotheses: (1) pre-latest Cretaceous occurrence of neornithians, implying
that the body fossil record is strongly biased toward the preservation of non-neornithian
birds, or (2) neornithians converged in foot morphology with their non-neornithian,
Cretaceous ancestors. The latter appears more parsimonious suggesting that convergent
evolutionary programs, in similar niches and at different periods, produced not only
similar pedal morphologies but also similar size (Lockley and Harris 2010).
Feeding behavior has been also inferred from different fossil bird tracks by com-
paring them with traces characteristic of modern birds. For example several
Ignotornis trackways exhibit reduced pace length, interpreted as a type of “foot
stirring” behavior characteristic of modern herons “stirring” up the substrate for
feeding (Lockley et al. 2008, 2009). The most noticeable example of feeding behav-
ior was reported from the Cretaceous of Korea (Lockley and Harris 2010; Lockley
et al. 2012a; Kim et al. 2012). Traces consist of sets of fine, zigzag, arcuate to semi-
circular grooves that “sweep” back and forth across the trackway, indistinguishable
from those of modern spoonbills (Swennen and Yu 2005). But spoonbills are not
recorded in the Cretaceous fossil record, and no spoonbill morphology is known in
any Cretaceous bird. The ichnologic record thus adds new data to the Cretaceous
avian diversity.
Fig. 10.8 Paleornithology and avian paleoichnology provide complementary data on the evolu-
tion of birds. This cladogram superimoposed on geologic time scale shows both taxon ranges
based on skeletal data and associated footprint occurrences. See text for discussion. Modified from
Lockley and Harris (2010)
The primary Mesozoic bird tracksites were recently reviewed by Lockley and Harris
(2010) but there are new findings reported every year (Falk et al. 2010, 2014; Falk 2011;
Fiorillo et al. 2011; Xing et al. 2011, 2013b; Contessi and Fanti 2012; Huh et al. 2012;
Kim et al. 2012; Lockley et al. 2012c, d; Azuma et al. 2013; Martin et al. 2014). Most
fossil bird tracks exhibit sizes and morphologies similar to those of extant shorebirds
and water-birds (Greben and Lockley 1992), and are generally associated with lake
basin, fluvial floodplain, coastal plain, and marine shoreline deposits (Lockley and
Harris 2010). This bias is strictly related to the physical properties of these environments
that apparently facilitate the registration and preservation of tracks, and the creation of
what has been called the shorebird ichnofacies (Lockley et al. 1994a; Lockley 2007b).
Tracks of birds with feet adapted for perching, such as songbirds or raptors, have much
less chance to be produced and preserved; the same happens with birds that live in envi-
ronments where the burial and preservation of tracks is less frequent or improbable
(mountain, forest, open marine, etc.). Even though there are many avian skeletal and
track fossils, the two records do not match each other and few footprints were probably
produced by the feet represented by the skeletal remains (Lockley and Harris 2010).
This mismatch and all the previously mentioned data strongly indicate that morpholo-
gies and behaviors of many modern birds actually evolved convergently with many of
their extinct, Mesozoic relatives. A striking example is provided by the discovery of an
Early Cretaceous (Aptian-Albian 120–110 Ma) trackway: Shandongornipes muxiai (Li
et al. 2005). This discovery shows the evolution of a terrestrial, zygodactylous bird that
is unknown in the body fossil record; it probably occupied a roadrunner-like niche indi-
cating evolutionary convergence of an early Cretaceous bird with the locomotory habits
and possible ecology of modern birds (see Lockley et al. 2007).
The majority of the fossil bird tracksites discovered to date are in East Asia,
primarily in China and South Korea (Azuma et al. 2002, 2013; Li et al. 2002; Kim
et al. 2006, 2012; Lockley et al. 2006b, 2007, 2009, 2012c, d; Xing et al. 2011,
2013b; Contessi and Fanti 2012; Huh et al. 2012; He et al. 2013; Falk et al. 2014).
The high frequency could be related to the abundance of lake deposits in these
regions, but the contemporaneous Chinese and North Korean skeletal records are
also very rich in birds. The complementary evidence thus indicates that the distribu-
tion pattern may reflect a true center of avian radiation (Lockley and Harris 2010).
10.2.8 Pterosaur Locomotion
Pterosaurs are the earliest known vertebrates to have evolved powered flight. These
winged reptiles first appeared in the Late Triassic and went extinct at the end of the
Cretaceous period (Buffetaut and Mazin 2003). Since their first discovery, there has
been considerable debate over their terrestrial locomotor capabilities, in particular
whether they walked bipedally (Cuvier 1809) or quadrupedally with their hindlimbs
splayed (Soemmerring 1812, 1817). Some eminent paleontologists considered that
pterosaurs were unable to walk (Abel 1925) or were bad walkers with their hindlimbs
unsuitable for a quadrupedal gait (Wellnhofer 1978). The hypothesis that pterosaurs
were digitigrade bipeds with an erect stance was supported and argued with phyloge-
netic and osteologic analysis by Padian (1983, 1984, 1985, 1987, 1988, 1991), Padian
and Rayner (1993), Paul (1987) and Bennett (1990). In contrast, several researchers,
on the basis of osteologic arguments, claim that pterosaurs were quadrupedal planti-
grades, with semi-erect or possibly even sprawling stances (Pennycuick 1986; Unwin
1987, 1989; Wellnhofer and Vahldiek 1986; Wellnhofer 1988, 1991a, b).
As was the case for other extinct track-making animals, ichnology provided
unequivocal evidence in resolving this long-standing dispute. Already in 1957 Stokes
described a short trackway composed of tridactyl manus and tetradactyl pes prints
(Stokes 1957). He instituted the new ichnotaxon Pteraichnus saltwashensis that he

identified as produced by a quadrupedal pterodactyloid pterosaur. This attribution
was endorsed by subsequent findings (Stokes 1978; Wellnhofer 1978; Stokes and
Madsen 1979), but was strongly opposed by Padian and Olsen (1984) who proposed
they were made by a crocodilian trackmaker. Pteraichnus was no longer considered
a pterosaur ichnotaxon (Unwin 1986, 1989; Prince and Lockley 1989; Lockley 1991;
Wellnhofer 1991a; Bennett 1992). New discoveries provided new information
(Logue 1994; Hunt et al. 1995b; Lockley and Hunt 1995a) and Mazin et al. (1995)
and Lockley et al. (1995d) independently demonstrated that Pteraichnus could be
attributed to pterosaurs (Fig. 10.9). To date, this interpretation is accepted by most
pterosaur researchers (Lockley et al. 1996, 1997; Bennett 1997; Mazin et al. 1997,
2001; Unwin 1997a, b, 2006; Wright et al. 1997; Kubo 2008; Witton 2013). Ichnology
therefore provided definitive evidence that pterosaurs walked on all fours.
These studies also demonstrated that at low-velocity walking, the body was held
upright, with the forelimbs held more vertically and the mani brought close to the
axis of the trackway, so they could not be starched much foreword. Conversely, at
Fig. 10.9 Fossil trackways

attributed to pterosaurs
provided exceptional
insight into the functional
morphology and behavior
of these flying reptiles.
Furthermore, they solved
the long-standing debate
over pterosaur terrestrial
locomotion. The tracks
called Pteraichnus show
that pterosaurs employed a
plantigrade, quadrupedal
stance and gait with the
hind limb partially—to
fully—abducted. From
Unwin (1996)
higher velocities, the body was held less erect (possibly subhorizontally) allowing
the long forelimbs to reach a more distant anterior point (Mazin et al. 2001).
Trackways were also useful to infer the landing behavior of pterosaurs, which like
modern birds used their wings to stall before landing (Mazin et al. 2009).
Pterosaur tracks are abundant in the Upper Jurassic rocks of Arizona, Colorado,
Oklahoma, Utah and Wyoming (Stokes 1957; Lockley et al. 1995d, 1996; Lockley and
Wright 2003; Lockley et al. 2008), France (Mazin et al. 1997, 2003, 2009; Billon-
Bruyat and Mazin 2003), Poland (Pienkowski and Niedzwiedzki 2005), and Spain
(García-Ramos et al. 2000, 2002, 2006; Meijide Calvo and Fuentes Vidarte 2001;
Fuentes Vidarte et al. 2004a, b; Sánchez-Hernández et al. 2009). Tracks of Cretaceous
age are much larger than those of the Late Jurassic, which agrees with body size trends
of skeletal remains (Buffetaut and Mazin 2003). Large body size in this group can have
evolved as early as Late Jurassic as testified by abundant footprints from Spain (García-
Ramos et al. 2000, 2002, 2006) and few skeletal remains from Switzerland (Meyer and
Hunt 1999). Pterosaur tracks have been reported from the Lower Cretaceous of China
(Li et al. 2002; Zhang et al. 2006; Xing et al. 2013b, c), England (Delair 1963, re-
interpreted by Wright et al. 1997), Germany (Hornung and Reich 2013), Japan (Lee
et al. 2010), South Korea (Kim et al. 2006; Lee et al. 2008), Spain (Pascual Arribas and
Sanz Pérez 2000; Meijide Calvo 2001; Fuentes Vidarte 2001; Hernández Medrano
et al. 2006; Moratalla and Hernán 2009; Sánchez-Hernández et al. 2009), and USA
(Maryland; Lockley and Rainforth 2002). In the Upper Cretaceous, they are known
from Argentina (Calvo and Lockley 2001), Mexico (Rodríguez de la Rosa 2003),
South Korea (Lockley et al. 1997; Hwang et al. 2002) and USA (Colorado, Lockley
and Rainforth 2002; Utah, Lockley et al. 1995d). Upper Cretaceous Korean tracks,
known as Haenamichnus, are the largest pterodactyloid tracks (up to 33 cm) and were
made by a trackmaker not yet known from the body fossil record.
Pterosaur tracks have been also used as an indicator of body mass. Kubo
(2011) tried to correlate the foot area with weight. He used seven ichnospecies of
pterosaur tracks and estimated body weights of pterosaurs as ranging from 110 to
145 kg. This result indicates that large pterosaurs were ten times heavier than the
heaviest modern birds.
The pterosaur track record provides evidence about the preferred environ-
ments and behaviors of these flying reptiles. Most Upper Jurassic pterosaur
tracks are found in marginal marine, carbonate, and clastic-evaporitic lagoonal
deposits whereas Cretaceous tracksites are associated with fluvial and freshwa-
ter deposits (Lockley and Rainforth 2002). The Late Jurassic record is consis-
tent with the hypothesis, inferred also through the body fossil record, that
pterosaurs inhabited marine shoreline habitats. This view is further corrobo-
rated by the discoveries of “swimming” and “feeding traces” (García-Ramos
et al. 2000, 2002, 2006; Lockley and Wright 2003; Witton 2013; Lockley et al.
2014). They consist of scrape marks that are interpreted as traces produced by
paddling limbs, a random distribution of tracks without evidence of trackways,

and small circular paired depressions left in the substrate by beak prods of
pterosaurs in search of food. This evidence reveals that pterosaurs could prob-
ably swim, or at least float on the surface of the water, like modern seabirds
(Witton 2013; Hone and Henderson 2014; Li et al. 2015).
10.3 Conclusions
By applying rigorous analytic methods, which allow full hypothesis testing, verte-
brate ichnology can provide reliable information about paleobiology (documenting
evolutionary patterns, tempo of evolution, locomotion, posture, size, speed, social
and individual behavior), paleobiodiversity (indicating the presence of taxa undocu-
mented by the skeletal record), paleoecology (giving hints on environmental/habitat
preferences and faunal composition) and paleobiogeography (providing presence/
absence data on geographic distribution) of trackmakers, vertebrate ichnology is
finally sitting at the “high table” of paleobiology.
In this review of Mesozoic reptile tracks some milestones of paleoichnologic
research have been evaluated with the aim of highlighting the potential of these
studies in contributing to the knowledge of evolutionary history and coevolution on
the changing Earth. We have shown that vertebrate track record can be used to study
the evolution of posture. Studies of the Permian to Triassic tetrapod-track record
documents the appearance of upright stance in reptiles and shows a previously hid-
den correlation between this event and the Permo-Triassic mass extinction. Using
synapomorphic characters in tracks has been essential in developing the present
understanding of the origin of the dinosauromorph clade. Tracks show that the most
basal dinosauromorphs were quadrupedal and that bipedalism was acquired phylo-
genetically closer to the origin of dinosaurs. An earlier origin of dinosauromorphs
might imply a possible link with the Permo-Triassic mass extinction. The ichno-
logic record can be used to make hypotheses about the spatial and temporal distribu-
tion of basal dinosaurs. Tracks indicate the presence of a diverse and widespread
fauna of dinosaurs during the early Late Triassic both in Laurasia and Gondwana, a
distribution not yet documented by the body-fossil record. Track records provide
evidence that habitats were shared by dinosaurian and non-dinosaurian archosaurs
in the Late Triassic, and support the hypothesis that crurotarsan-dominated faunas
were replaced by dinosaurs through a gradual process of ecologic replacement.
Combined analyses of body and track fossils provide insights in the understanding
of both environmental preferences and evolution of locomotory mechanisms in sau-
ropod dinosaurs. Ichnologic data reveals the inland preference of wide-gauge titano-
saurs highlighting the importance of considering track data in ecologic analysis.
Trackways are used to estimate the speeds of trackmakers and provide clues to
individual behaviors of theropod dinosaurs. Empiric formulae derived from extant
animals constitute the basis of these interpretations, and show that theropods pri-
marily used walking gaits, although some were runners, and there are a few records
of individuals using trotting gaits. Crouching traces demonstrate that the avian ori-
entation of the manus, with a medially facing palm, had evolved by the Early
Jurassic within Theropoda. Footprints provide clues on social behavior in dino-
saurs. Many sites have shown that herbivorous dinosaurs were at times gregarious.
Carnivorous non-avian theropods were sometimes gregarious although interpreta-
tions are more controversial. A strong link between paleornithology and avian
paleoichnology illuminates aspects of avian evolution, particularly with regard to
paleoenvironmental preferences. Ichnology is particularly relevant when studying
shorebirds evolution, given the completeness of the track record for this group.
Footprints are crucial to understand pterosaur terrestrial locomotion and stance, to
estimate body mass, and to learn about their physiology and behavior (swimming,
feeding, landing).
Acknowledgments Our sincere thanks to the editors M.G. Mángano and L.A. Buatois for the
invitation to contribute to this volume and for the enthusiastic support during the realization of the
chapter. We thank J.D. Harris for providing bibliographic references. P. Currie and C. Meyer are
thanked for their constructive reviews. This study was supported by MUSE, Museo delle Scienze,
Trento. M.B., F.M.P. and M.A. were supported also by the Promotion of Educational Policies,
University and Research Department of the Autonomous Province of Bolzano-South Tyrol as part
of the project “The Permian–Triassic ecological crisis in the Dolomites: extinction and recovery
dynamics in Terrestrial Ecosystems” and partly as part of the project “The end-Permian mass
extinction in the Southern and Eastern Alps: extinction rates vs taphonomic biases in different
depositional environments” financed by the Euregio Science Fund (call 2014, IPN16) of the
Europaregion Euregio. L.P and J.C.G.-R. were supported by Project CGL2012-33281 of the
Spanish Ministry of Economy and Competitiveness.
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Chapter 11
The Mesozoic Lacustrine Revolution
Luis A. Buatois, Conrad C. Labandeira, M. Gabriela Mángano,

Andrew Cohen, and Sebastian Voigt
11.1 Introduction
The Mesozoic lacustrine revolution (MLR) represents a major evolutionary event

in the continental realm (Cohen 2003). The decline in taxic diversity at the family
level in lacustrine environments that took place during the late Paleozoic–Middle
Triassic was reversed later in the Mesozoic, with diversification seeming to have
continued to the Quaternary, although potential biases may have affected this
trend (Cohen 2003). Major novelties and innovations that took place in lacustrine

C.C. Labandeira
Department of Paleobiology, National Museum of Natural History; Smithsonian Institution
Washington, DC 200013-7012, USA
Department of Entomology and BEES Program, University of Maryland,
Collage Park, MD 20742, USA
College of Life Sciences, Capital Normal University, Beijing 100048, China
Department of Geology, Rhodes University, Grahamstown, 6140 South Africa
College Park, MD 21740, USA
A. Cohen
Department of Geosciences, The University of Arizona, 1040 E. 4th Street,
Tucson, AZ 85721, USA
S. Voigt
Urweltmuseum GEOSKOP, Burg Lichtenberg (Pfalz), Burgstraße 19,
66871 Thallichtenberg, Germany

DOI 10.1007/978-94-017-9597-5_11
settings during the mid-Mesozoic include the diversification and increase in mor-
phological disparity of aquatic macrophytes, aquatic insects, and teleosts. By the
mid Cretaceous (125–90 Ma), the basic ecologic structure of the MLR was well
established, defined by an ecologically significant increase of herbivory within
ecosystems which previously had been overwhelmingly composed of detritivores
and predators. This trophic readjustment was minimally altered by the end-Creta-
ceous global crisis and the subsequent ecologic rearrangement of the Paleogene
world (Dunne et al. 2014).
It is a basic premise of this chapter that a review of this formative interval from
a comparison of the body- and trace-fossil records will illuminate our understanding
of the MLR. Although direct links between individual ichnotaxa and producers
commonly are not possible, a comparison of the trace-fossil and body-fossil records
through time is valuable in detecting the timing of large-scale ecologic changes in
lacustrine ecosystems. Consequently, in this chapter we review the trace- and body-
fossil record of lake and related continental aquatic deposits to establish and assess
the importance of the MLR. As part of our examination, we discuss how ichnologic
evidence may provide insights into major evolutionary innovations within lacus-
trine ecosystems. To do so, we provide an extensive summary of the available ich-
nologic information for Mesozoic–Cenozoic lacustrine successions. In addition, a
review of the paleoecologic structure of Eurasian paleolakes in particular provide
important evidence for understanding the impact this event had on trophic relation-
ships throughout the water column (Zherikhin et al. 1999; Sinitshenkova 2002) as
well as the lacustrine benthos and associated sediments during this formative inter-
val (Buatois et al. 1998a; Mángano and Buatois 2007; Voigt and Hoppe 2010).
Although this chapter is focused on the evolutionary novelties and innovations that
took place during the Mesozoic, in order to place these changes within a broader
context, we briefly outline the basic features of lacustrine ecosystems prior to the
MLR (see Chap. 6 for a detailed discussion on the ichnology of Silurian-Permian
lakes). A more complete context for understanding the MLR involves a search for
the roots of Mesozoic MLR novelties and innovations in older deposits to establish
relevant ichnological, biotic, and physical contrasts between the late Paleozoic and
the Mesozoic worlds. Similarly, we provide a discussion of the post-Mesozoic
record in order to evaluate the aftermath of the MLR.
11.2 Methods and Terminology
For this study, all ichnotaxonomic determinations have been checked and adjusted
based on a reevaluation of the available literature. Because the literature essentially
provides two contrasting trace fossil-assemblages, a distinction is made between
lake-margin settings and fully subaqueous lacustrine conditions. Lake-margin set-
tings are characterized by periodic fluctuations of the water table, leading to highly
variable conditions in substrate consistency. Typically, a characteristic zonation from
softgrounds near the edge of the water body to drier substrates outward from the lake
develops (Scott et al. 2012a). Lake-margin ichnofaunas commonly display activity of
11 The Mesozoic Lacustrine Revolution 181
a combined terrestrial to shallow subaqueous biota. The degree of consolidation of

the substrate plays a major role for preservation of biogenic structures in lake-margin
deposits (Buatois and Mángano 2004, 2009). Ichnofaunas from lake-margin settings
typically comprise the Scoyenia Ichnofacies (Buatois and Mángano 1995, 2004,
2009) and less commonly the Camborygma Ichnofacies (see Chaps. 1 and 13).
Fully lacustrine settings (i.e. the permanent subaqueous zone), particularly in hydro-
logically open systems, may host a relatively diverse benthic fauna. In this zone a where
a lake experiences a high degree of environmental stability and appropriate ecologic
conditions, particularly high levels of oxygenation, low energy, constant food supply, the
presence of freshwater, and cohesive substrates (Buatois and Mángano 1995, 2004,
2009; Miller and White 2007; Scott et al. 2012a). Oxygenation is a first-order limiting
factor, because in lakes with year-round thermal stratification the hypolimnion becomes
anoxic or dysoxic and bioturbation is precluded in the lake bottom below the mixolim-
nion. Low-energy turbidity and underflow currents may provide oxygen and food to
lake bottoms. Colonization by macrobenthos and preservation of their trace fossils are
both unfavorable in soupy substrates and, accordingly, some degree of substrate cohe-
sion is required to allow the passage of discrete biogenic structures through the fossiliza-
tion barrier. Ichnofaunas from fully lacustrine settings typically form the Mermia
Ichnofacies (Buatois and Mángano 1995, 2004, 2009; see Chap. 1).
At the scale of the basin, we commonly refer to the scheme developed by Bohacs
et al. (2000), who recognized three different types of lake basins, namely overfilled,
balanced-fill, and underfilled. Overfilled-lake basins occur if the rate of sediment/
water input exceeds the rate of formation of potential accommodation. These sys-
tems are typically hydrologically open, and form deposits of fluvio-lacustrine, silici-
clastic sediments that display parasequences resulting from shoreline progradation
and delta-channel avulsion. Balanced-fill lake basins are formed when rates of sedi-
ment/water supply are in balance with the formation of potential accommodation.
These lakes tend to shift periodically from hydrologically open to closed and vice
versa, producing both carbonate and siliciclastic deposits which display parase-
quences that record both progradational parasequences and aggradation of chemical
sediments during periods of desiccation. Underfilled-lake basins are characterized by
rates of accommodation formation that exceed the rate of supply of sediment/water.
These are typically hydrologically closed lakes, dominated by evaporite deposits
which display parasequences that record vertical aggradation. This scheme has been
successfully used to frame both ichnologic (Buatois and Mángano, 2004, 2007,
2009) and paleobiologic (Gierlowski-Kordesch and Park, 2004) information.
In order to frame the major ecosystem changes discussed in this chapter, we have
used the distinction between novelties and innovations outlined by Erwin and
Krakauer (2004) and Erwin (2012). According to these authors, invention is the
creation of something new, whereas innovation is a successful development of an
invention. Evolutionary novelties only result in innovations when they drive eco-
logic transformation (Erwin 2012).
For better clarity in specifying the immatures of non-holometabolous versus
holometabolous insects, we employ the standard entomological terminology used in
North America (China et al. 1958). For those insects possessing egg to naiad/nymph
to adult development, the term “naiad” is used for an aquatic immature instar that
lacks holometabolous development (Snodgrass 1954). Immatures consisting of

naiads occur in the Ephemeroptera (mayflies) and Odonata (dragonflies and damsel-
flies), the extinct archaeorthopteroid Chresmodida, Plecoptera (stoneflies), and
some groups of Heteroptera (true bugs) within the Hemiptera. A terrestrial imma-
ture of these non-holometabolous insects is termed a “nymph” (Snodgrass 1954;
Davies 1958), and nymphs commonly occur within the same major groups as spe-
cies with naiad immatures. Examples of taxa with nymphs include the terrestrial
Orthoptera (grasshoppers, crickets), Blattodea (cockroaches), Isoptera (termites),
and Mantodea (mantids). We restrict the term “larva” solely to insects with holome-
tabolous development, consisting of egg to larva to pupa to adult development
(Snodgrass 1954). Holometabolous taxa that have aquatic larvae include certain
groups within the Coleoptera (beetles); the Megaloptera (alderflies and dobson-
flies); a few groups of Neuroptera such as spongillaflies; certain, especially nemato-
cerous, Diptera (true flies); and the Trichoptera (caddisflies). Restriction of the
term, larva, to holometabolous insects offers more precision than application of the
term broadly to multiple developmental modes that would encompass developmen-
tal stages such as aquatic naiads, terrestrial nymphs, and probably forms that lack
developmental change. The distinction used herein generally has been accepted by
a broad spectrum of evolutionary developmental biologists (Gilbert 2014).
The body-fossil record of ancient lacustrine basins is strongly biased as a result
of the distribution of fossil Lagerstätten. Fortunately, these spectacular deposits are
quite common in lacustrine successions (see Table 14.1 in Cohen 2003). However,
integrating body-fossil evidence from fossil Lagerstätten with ichnologic informa-
tion frequently is problematic because many of the former records accumulated as
event beds under anoxic conditions, preventing bioturbation, and therefore having a
sparse trace-fossil record. Notwithstanding this limitation, there are a number of
Mesozoic examples where exceptional body-fossil accumulations have been recov-
ered in association with trace fossils, most notably the Madygen (Voigt and Hoppe
2010), El Montsec (de Gibert et al. 2000) and Las Hoyas (Buatois et al. 2000a; de
Gibert et al. 2016) biotas.
11.3 Lacustrine Ecosystems in Deep Time
Exploration of lacustrine ecosystems in deep time is a challenging enterprise

because lakes are geologically ephemeral (Cohen 2003). However, the long-term
evolution of organisms in freshwater settings provides continuity, allowing recon-
struction, albeit tentative, of secular changes in lacustrine biotas (Anderson and
Dean 1988; Buatois et al. 1998a; Labandeira 1999; Park and Gierlowski-Kordesch
2007). Because of the patchiness and lack of connection between individual lakes in
space and time, as opposed to the more continuous record of oceans, the fossil
record of lacustrine organisms is sporadic (Cohen 2003).
As with other depositional settings, lakes are affected by a number of tapho-
nomic megabiases (Behrensmeyer and Kidwell 1985; Behrensmeyer et al. 2000).
Taphonomic megabiases are defined as “largescale patterns in the quality of the

fossil record that affect paleobiologic analysis at provincial to global levels and at
timescales usually exceeding ten million years” (Behrensmeyer et al. 2000). In par-
ticular, the fossil record of lacustrine basins is strongly affected by both intrinsic
changes and extrinsic abiotic and biotic changes. The former results from the evolu-
tion of new body plans and the impact of behavior on fossilization potential
(Behrensmeyer et al. 2000). As in the archetypal case of marine environments,
development of mineralized hard parts is one of the intrinsic factors in lake settings,
as is the establishment of fossorial behavior by vertebrates. The increased depth and
extent of bioturbation through time exhibited by the lacustrine infauna is certainly
one of the most important, extrinsic biotic factors contributing to taphonomic mega-
biases in these settings (see Sect. 11.8.2). Finally, tectonic and climatic controls on
the establishment and evolution of lacustrine basins are prime examples of extrinsic,
abiotic factors in the generation of taphonomic megabiases. These interpretive limi-
tations notwithstanding, our understanding of this record are informed by several
key observations.
(1) Lacustrine habitats have experienced a history of regular and repeated
formation, isolation, and destruction. Because of the combined effects of climate
variability and change as well as sediment infill, most individual lakes only persist
over timescales of ~102–104 yrs, with only a small percentage of lakes, generally
those originating from tectonic, volcanic, and meteoritic impact processes, persist-
ing over longer intervals of time (Cohen 2003; Cohen et al., 2015). Most of the
processes that create lakes, such as fluvial diversion from damming, glacial block-
age of drainage systems and delta-plain subsidence, are also predilect for the even-
tual destruction of these same habitats. The creation and destruction of lakes often
occur at quasi-predictable tempos scaled by the lake formation process, including
determining potential sediment accommodation space, coupled with the typical
range of sediment accumulation rates associated with lacustrine systems.
(2) The need for dispersal of organism populations among lakes is a primary
driver of both evolutionary innovations and constraints on the lacustrine biota.
The excellent dispersal adaptations of organisms are a hallmark characteristic of
most lacustrine systems. Organisms have evolved adaptations for dispersal as well
as for the constraints emplaced on them by the “typical” lacustrine environment
mentioned above. These features include adaptations for flight, such as active flight
by aquatic insects, and passive carriage by birds, fish, and wind (Boag 1985; Bilton
et al. 2001; Green and Figuerola 2005; Van Bocxlaer et al. 2011); active parasitism
on animal dispersal vectors (Graf and Cummings 2006); and desiccation resistance
(Watanabe et al. 2002; Watanabe 2006). Conversely, the very adaptations for disper-
sal between ephemeral waterbodies also promote gene exchange between popula-
tions and reduce the likelihood of population isolation and speciation (Bilton et al.
2001). These adaptations inherently constrain and scale the rates of lacustrine evo-
lution to the stability and persistence of their waterbodies, with the faunas of shorter-
lived lakes (better migrants or desiccation survivors) associated with slower
evolutionary rates and overall lower diversity. Morphological novelties arising from
new functional and behavioral modifications take advantage of changing lacustrine
food resources, of which herbivory on aquatic plants or tiered detritivory exploiting

deeper substrates are prime examples. These novelties become innovations when in
the case of dispersal, the spread of disseminules is severely impeded by a periodic
scarcity of quasipersistent lakes, and the energetic requirements of adaptations for
exploiting lacustrine resources run up against the competing need to migrate to new
habitats.
(3) Lacustrine evolution in long-lived lakes is highly iterative and lacustrine
habitats can act as refugia for evolutionary novelties and innovations.
Comparative studies of the living and fossil faunas of long-lived Neogene African
Great Lakes and early Mesozoic Newark Supergroup paleolakes show that specia-
tion and diversification trends often are highly iterative. The iteration of homolo-
gous morphological novelties in lacustrine clades evolve repeatedly in space and
time whenever lakes persist sufficiently long for the results of diversification to be
expressed (McCune 1996; Salzburger et al. 2014). Particular body plans and evolu-
tionary novelties have been observed resulting from the convergence of lineages
within isolated lakes. One such convergence is the iconic example of cichlid fish
pharyngeal jaw mechanics and overall body structure converging among species
populations in Lakes Tanganyika and Malawi (Kocher et al. 1993). Perhaps even
more remarkable, trophic interactions have also been observed to converge among
long-lived lakes, albeit occasionally with slightly different players. One example
involves patterns of escalatory predator–prey coevolution (West and Cohen 1996;
Van Damme and Pickford 2003). Occasionally, the resulting species of these radia-
tions are themselves later dispersed into other lake systems, providing the begin-
nings for more widespread spatial diversification of the original clade (Van Damme
and Pickford 1999; Anderson et al. 2010).
(4) The tempo of evolutionary innovation related to trophic interactions in
lakes is probably modulated by the history of geochronologically long-lived
lakes. If long-lived lakes can serve as incubators for diversification and evolution of
key innovations and interactions in aquatic organisms, then it is likely that during
times of such lake abundance, the most favorable opportunity exists for large scale
trophic “revolutions” within lacustrine biotas. The mid-Mesozoic breakup of
Pangaea probably represented the peak period of long-lived lake formation. These
lakes were coupled with an ever increasing provinciality as they were progressively
rafted on different continental land masses around the planet (Cohen 2003).
11.4 The Lacustrine World Prior to the Mesozoic Revolution
In this section we provide a brief review of the basic features of lacustrine commu-
nities that immediately preceded those of the Mesozoic world. Evidence of
Proterozoic lacustrine biotas is scarce, to say the least, essentially being restricted to
stromatolites most likely formed by cyanobacteria (e.g. Hoffmann et al. 1980) and
leiosphaeridian acritarchs that may have been adapted to freshwater, although this
is a contentious issue (Martín-Closas 2003). There is a paucity of evidence for
body-fossils in Cambrian through mid-Silurian continental aquatic habitats

(Labandeira 2005a). Tasmanitids (green algae) were abundant during the early
Paleozoic, but most likely inhabited brackish rather than fresh water (Martín-Closas
2003). The Cambrian-Ordovician evidence for the incipient colonization of land
originates from trace fossils of arthropods that were able to foray into intertidal
areas, coastal dunes and ponds (see Chap. 5), and stromatolites that were similar in
structure to, but commonly more diminutive than those of the Precambrian
(Ponomarenko 2009). In addition, the occurrence of a diverse cryptospore assem-
blage in Lower to Middle Ordovician deposits represents the earliest evidence of
land plants, suggesting an early origin of embryophytes (Rubinstein et al. 2010).
Chlorophytes were represented mainly by the Chlorococcales, which became domi-
nant in lacustrine plankton by the Ordovician (Martín-Closas 2003), and the closely
related Charophytes that were present in freshwater bodies at by the Late Silurian,
if not earlier (Feist et al. 2005). The presence of liverwort and embryophyte-grade
spores toward the end of this interval, and continuing into the Early Devonian, sig-
nals the establishment of an incipient subaerial plant cover, probably adjacent to
bodies of brackish and freshwater inland and along coastlines (Strother 2000).
These developments, in conjunction with the presence of lacustrine (and fluvial)
sediments, particularly during the Late Ordovician and Silurian, indicate the emer-
gence of a depauperate lacustrine biota consisting of transient and permanent micro-
organisms, plants, fungi, and invertebrate animals. The fossil history of lacustrine
zooplankton is notably poor. However, molecular clocks may help to provide some
constraints. For example, cladoceran crustaceans are thought to have originated by
the Devonian (Sacherová and Hebert 2003). The Silurian-Devonian lacustrine
trace-fossil record is restricted to lake-margin settings, and is dominated by arthro-
pod trackways (Buatois and Mángano 1993a; Buatois et al. 1998a; see Chap. 6).
The restriction of biogenic structures to lake margins and the apparent absence of
fully lacustrine ichnofaunas are consistent with extremely inefficient nutrient deliv-
ery in the absence of an extensive upland plant cover (Cohen 2003). These lakes
may have been ultraoligotrophic and limited in phosphorous (Cohen 2003), pre-
cluding the establishment of a lacustrine benthos.
With few exceptions (Grenier 1974; Clarkson et al. 1993; Rolfe et al. 1993;
Jeram and Selden 1993; Shear 1993), the Lower Carboniferous (Mississippian)
lacks a significant record of well-documented lake deposits and associated biotas.
During the Late Carboniferous (Pennsylvanian), two major types of ecosystems
appeared that were displaced by differing environmental conditions. The Wet Biome
emerged during the earlier Pennsylvanian and included a wide variety of wetlands,
notably coal-swamp communities that consisted of water-tolerant lycopods, sphe-
nopsids, medullosan seedferns, and arborescent marattialean ferns, occupying much
of humid equatorial Euramerica (Gastaldo et al. 1996). By contrast, evidence for a
Dry Biome appears toward the latter half of the Pennsylvanian, and included com-
munities dominated by a desiccation-tolerant biota, such as cordaites, noeggeran-
thialeans, conifers, gigantopterids, peltasperms, and probably cycads that occupied
more continental, extra-tropical environments (Opluštil et al. 2013). Regarding zoo-
plankton, copepods were already present by the Mississippian in glacial lakes of
Gondwana (Selden et al 2010). During the Permian, the tropical to paratropical Wet
Biome diminished significantly, at least in North America and Europe, although it
survived largely intact in China (D’Rozario et al. 2011). By contrast, the Dry Biome,
which first appeared during the latter half of the Pennsylvanian (Opluštil et al.
2013), temporally overlapped with the Wet Biome, and became prominent in more
inland, drier basins of the Permian as it assumed a distinctive biotal character in
many habitats. Aquatic herbivory was still absent, in contrast to a pulse of signifi-
cant herbivory that had evolved on land by the Late Pennsylvanian (Labandeira
2006). In addition to ichnofaunas in lake-margin deposits, Mississippian and
Pennsylvanian trace fossils also are present in fully subaqueous lacustrine deposits,
indicating a significant environmental expansion of the benthic fauna (Buatois and
Mángano 1993a; Buatois et al. 1998a) (see Chap. 6). These lacustrine deposits were
colonized by a moderately diverse, mobile, detritus-feeding epifauna. This expan-
sion was probably linked to the rapid diversification, and increase in abundance of
land plants, attributable to vegetational changes that introduced abundant organic
detritus into previously nutrient-poor, lacustrine habitats (Maples and Archer 1989).
Also, ichnologic information suggests a major diversification event during the
Mississippian and Pennsylvanian, which parallels the diversification of freshwater
organisms, such as arthropods, annelids, fish, and mollusks (Maples and Archer
1989; Buatois et al. 1998a). The ichnologic evidence of the Dry Biome during the
subsequent Permian consists of a high abundance and moderate diversity of arthro-
pod trackways emplaced in subaerially exposed sediment of playa-lake systems
(see Chap. 6). By the Permian, the presence of meniscate trace fossils (e.g. Scoyenia)
records the establishment of a mobile, shallow- to mid-tier, infauna in firm, desic-
cated substrates. However, these structures still were relatively rare and for the most
part have been recorded in overbank environments rather than in lake-margin set-
tings that are overwhelmingly dominated by arthropod and tetrapod trackways.
11.5 The Mesozoic Lacustrine Revolution
The Mesozoic trace-fossil record of lacustrine successions is uneven in time and

space. The Lower Triassic ichnologic record of lake-margin environments is
remarkably poor, and we are not aware of well-documented, fully lacustrine, Early
Triassic ichnofaunas. In contrast, Middle to Late Triassic lacustrine ichnofaunas
have been documented in great detail. The Jurassic ichnologic record is uneven,
with a number of detailed studies documenting both Early and Late Jurassic ichno-
faunas, but less information is available for the Middle Jurassic. The Early
Cretaceous record is relatively good, but the record of Late Cretaceous fully lacus-
trine ichnofaunas is particularly poor.
Unsurprisingly, body fossils of the MLR are unevenly distributed as well.
Triassic paleontologic information is essentially based on Eurasian localities and, to
a lesser extent, eastern North America and South Africa. The Jurassic body-fossil
record, particularly for aquatic insects, is still significantly centered in Eurasia, but
includes a few localities in western North America as well. In contrast, the Early
Cretaceous tends to show a more widespread distribution of lacustrine fossiliferous
localities, but still is heavily represented by Eurasian lake deposits, most likely
resulting from extensional tectonics during the breakup of Gondwana. However,
information from the Late Cretaceous is patchy.
11.5.1 Early Triassic Denouement
After the ecologic crisis at the Permian–Triassic boundary, the short-lived, five
million-year-long Early Triassic represents a period that ranges from biotal stasis to
subtle increases in the diversity of aquatic and terrestrial communities (Looy et al.
1999). The ichnology of Lower Triassic lake-margin deposits has been documented
in a few places, namely Germany (Knaust and Hauschke 2004, 2005) and the west-
ern United States (Lovelace and Lovelace 2012). A moderate diversity of inverte-
brate trace fossils has been recorded in marginal facies of an underfilled playa-lake
system in Germany (Knaust and Hauschke 2005). The playa-lake ichnofauna con-
tains arthropod trackways and trails (Diplichnites, Stiallia, Diplopodichnus), arthro-
pod bilobate structures (Cruziana, Rusophycus), vertical burrows (Skolithos),
bivalve burrows (Lockeia), branching burrow systems (Phycodes), and the horizon-
tal J-shaped ichnogenus Fuersichnus. Pseudofossils, such as Aristophycus, are pres-
ent in the same deposits (Knaust and Hauschke 2004). Body fossils have been
recovered from these deposits, including conchostracans, notostracans, and
xiphosurids, as well as indeterminate fish remains (Knaust and Hauschke 2005).
The arthropod trackway Diplichnites and the bivalve dwelling/resting structure
Lockeia also occur in Lower Triassic shallow-lacustrine deposits of the western
United States (Lovelace and Lovelace 2012).
In contrast to younger lake-margin ichnofaunas, meniscate, backfilled trace fos-
sils are absent in these Lower Triassic examples, and arthropod trackways seem to
be relatively common. In this respect, Lower Triassic lake-margin ichnofaunas
apparently are more similar to those from the Permian (see Chap. 6) than to the
archetypal Mesozoic suites, revealing the persistence of Paleozoic types of animal–
substrate interactions. Also, as is commonly the case for Paleozoic lacustrine ichno-
faunas, shallow tiers were dominant and penetrative structures, such as those
responsible for ichnofabric formation, were absent (Knaust and Hauschke 2005).
The Early Triassic continental body-fossil record is sparse as well (Shcherbakov
2008a). Freshwater insects originate mostly from Eurasian localities and generally
consist of mayfly wings (Sinitshenkova 2013), archaeorthopteran and related
orthopteroid taxa (Shcherbakov 2008b), cockroaches (Żyla et al. 2013), a few basal
hemipteran lineages (Shcherbakov 2008a), and isolated elytra of schizophorid and
permosynid beetles and related lineages (Ponomarenko 2004, 2008). Many of these
fossil taxa preferentially are represented by immatures, especially naiads, that when
compared to conspecific adults, occur at frequencies greater than is typical of the
Permian (Shcherbakov 2008b). As with Early Triassic plants (Looy et al. 2001),
contemporaneous insect faunas are characterized by four major changes when com-
pared to their Late Permian precursors (Shcherbakov 2008b). These shifts are: (1)
the evolution of new, Early Triassic dominant groups at the expense of earlier Late
Permian dominants; (2) movement into the high-latitude Triassic by groups that
formerly occurred at Permian low latitudes; (3) preferential survival of small-sized,
generalist feeding taxa; and (4) emergence of lineages that were aquatic or other-
wise tied to water bodies such as lakes. Although these changes promoted diversity,
it was not until the Anisian stage of the early Middle Triassic, 5–11 m.yr. after the
end-Permian crisis, when insect taxa, including aquatic forms, and their interactions
with plants approached the diversity that had existed during the Late Permian (Gall
1996; Labandeira 2005b).
Another important component of the Triassic benthos was charophytes, which
are particularly well known in lacustrine carbonates and marls (Martín-Closas
2003). Isoetes-related, nonflowering vascular plants were present in the littoral zone
(Cohen 2003; Moisan et al., 2012b). As in the Paleozoic, lacustrine phytoplankton
was still dominated by Chlorococcales, but an increase in diversity is apparent
(Brenner and Foster 1994; Martín-Closas 2003).
11.5.2 Rediversification During the Middle to Late Triassic
Ichnofaunas have been documented in detail in Middle to Upper Triassic lake-

margin deposits (including those of playa lakes) from the eastern (Metz 1995, 1996,
2000; Szajna and Hartline 2003) and western (Gillette et al. 2003; Lucas et al. 2010)
United States, Argentina (Melchor et al. 2003, 2006; Melchor 2004; Genise et al.
2009; Marsicano et al. 2010), Greenland (Bromley and Asgaard 1979), England
(Porter and Gallois 2008), Germany (Schlirf et al. 2001), Morocco (Hminna et al.
2015), and China (Shi et al. 2007; Li et al. 2014). These lake-margin deposits com-
monly contain a relatively wide variety of vertebrate and invertebrate trace fossils.
Vertebrate ichnofaunas in Middle to Upper Triassic lake-margin deposits typi-
cally are dominated by tetrapod trackways (e.g. Brachychirotherium, Brasilichnium,
Dicynodontipus, Characichnos, Evazoum, Grallator, Gwyneddichnium,
Rhynchosauroides, Tetrasauropus) and, more rarely, lungfish burrows
(Redondarefugium) (Melchor et al. 2006; Lucas et al. 2010). Typical invertebrate
trace fossils in lake-margin deposits are shallow-tier, ornamented, meniscate,
unbranched (Scoyenia), and branched (Spongeliomorpha) structures, as well as
meniscate structures lacking bioglyphs (Taenidium), arthropod bilobate structures
(Cruziana, Rusophycus), and simple horizontal (Palaeophycus, Planolites) and verti-
cal (Skolithos) burrows (Bromley and Asgaard 1979; Metz 1995, 1996; Schlirf et al.
2001; Gillette et al. 2003; Melchor et al. 2003, 2006; Melchor 2004; Shi et al. 2007;
Porter and Gallois 2008; Lucas et al. 2010; Li et al. 2014; Hminna et al. 2015). In
some cases, great densities of the bivalve burrow Lockeia (Lucas et al. 2010) and the
ichnogenus Fuersichnus (Bromley and Asgaard 1979) have been documented, typi-
cally forming monospecific trace-fossil suites. Arthropod trackways, although
extremely common in Paleozoic lake-margin settings (Buatois and Mángano 1993a;

Buatois et al. 1998a; see Chap. 6), and are quite rare in Middle to Upper Triassic
deposits that were formed in similar environments, typically represented as isolated
occurrences (Melchor 2004; Melchor et al. 2006). Horizontal, nonspecialized graz-
ing trails, such as Helminthopsis and Cochlichnus, may be present (Melchor 2004;
Shi et al. 2007; Li et al. 2014; Hminna et al. 2015), but they are never dominant and
tend to reflect emplacement in substrates that did not experience periodic desicca-
tion. Three-dimensional branching burrow systems (Treptichnus) may be present,
but are uncommon (Porter and Gallois 2008). These structures were typically
emplaced close to the sediment surface, the only exception being those produced by
crayfish, referred to the ichnogenus Camborygma, which display variations in archi-
tecture as a response to the depth of the water table. Complex architectures with
many branches and chambers were constructed by primary burrowers in areas of
high water table, whereas deep and simple burrows are dominant in areas of low and/
or highly fluctuating water table (Hobbs 1981; Hasiotis and Mitchell 1993).
Middle to Late Triassic, fully lacustrine ichnofaunas are known from the eastern
United States (Metz 1995, 1996, 2000), Argentina (Melchor et al. 2003, Melchor
2004), Kyrgyzstan (Voigt and Hoppe 2010), England (Porter and Gallois 2008), and
Greenland (Bromley and Asgaard 1979). Ichnofaunas in low-energy deposits are
essentially represented by invertebrate trace fossils, with a dominance of very
shallow-tier, simple, horizontal trails of deposit and detritus feeders (e.g.
Helminthopsis, Helminthoidichnites, Cochlichnus, Mermia) and of shallow-tier,
three-dimensional, branching burrow systems (Treptichnus) (e.g. Metz 1995, 1996,
2000; Melchor et al. 2003; Melchor 2004; Porter and Gallois 2008). Vertebrates are
represented by the fish trail Undichna (Melchor 2004).
One of the first lacustrine ichnofaunas studied in detail is that of the Fleming
Fjord Formation of Greenland (Bromley and Asgaard 1979) (Fig. 11.1a–f).
Integration of ichnologic and sedimentologic information indicates that three main
trace-fossil assemblages are present in these ephemeral lacustrine deposits (Bromley
and Asgaard 1979, 1991; Dam and Stemmerik 1994; Bromley 1996). Skolithos,
Arenicolites, and Polykladichnus occur in tempestites, whereas Fuersichnus
(Fig. 11.1a–b) and Lockeia (Fig. 11.1c) are present in fair-weather deposits.
Scoyenia (Fig. 11.1d–f) and Skolithos (Fig. 11.1d–f) occur in marginal-lacustrine
deposits.
Upper Triassic ichnofaunas preserved in deposits of the Newark Supergroup,
which record sedimentation in lakes within an extensive rift system in eastern North
America, formed during the initial breakup of Pangaea (Olsen 1989; Schlische 2003).
These ichnofauna have been analyzed in detail (e.g. Olsen and Flynn 1989; Metz
1995, 1996, 2000) (Fig. 11.2a–g). One of these units, the Lockatong Formation,
records sedimentation in a balanced-fill lake characterized by recurrent base-level
fluctuations. As a result, successions show well-defined, vertically stacked, trans-
gressive–regressive cycles (Olsen 1980). The invertebrate ichnofauna consists of a
combination of feeding (Planolites montanus, Scoyenia gracilis, Treptichnus pol-
lardi), dwelling (Spongeliomorpha milfordensis), grazing (Cochlichnus anguineus),
resting (Lockeia siliquaria), and locomotion (an undetermined arthropod trackway)
Fig. 11.1 Characteristic trace fossils from Upper Triassic ephemeral lacustrine deposits of the
Flemming Fjord Formation of Greenland. (a) General view of a surface displaying a high density
of Fuersichnus communis; (b) Close-up of Fuersichnus communis showing its characteristic
banana shape; (c) The almond-shape trace fossil Lockeia amygdaloides. The associated horizontal
burrows may be referred to the ichnogenus Ptychoplasma; (d) General view of a surface showing
several specimens of the meniscate ichnotaxon Scoyenia gracilis. Note the presence of circular
cross-section of vertical Skolithos isp.; (e) Scoyenia gracilis with less developed meniscate infill
and bedding-plane expression of Skolithos isp.; (f) Close-up of Scoyenia gracilis and Skolithos isp.
All scale bars are 1 cm long
Fig. 11.2 Characteristic trace fossils from Upper Triassic lake-margin deposits of the Newark
Supergroup of eastern United States. (a) General view of a surface with Scoyenia gracilis, Passaic
Formation; (b) Close-up showing bioglyphs in Scoyenia gracilis, Passaic Formation; (c) Close-up
showing meniscate infill in Scoyenia gracilis, Lockatong Formation; (d) General view of a surface
with high density of Lockeia amygdaloides, Passaic Formation; (e) Treptichnus bifurcus, Passaic
Formation; (f) Cochlichnus anguineus, Passaic Formation; (g) Helminthoidichnites tenuis, Passaic
Formation. All scale bars are 1 cm long. Photographs courtesy of Robert Metz
structures (Metz 1995). The reptile trackway Gwyneddichnium is present as well

(Olsen and Flynn 1989). Trace fossils are almost invariably restricted to lake-margin
deposits. Whereas some of these structures most likely were emplaced in wet sub-
strates (e.g. Treptichnus pollardi, Lockeia siliquaria), other ichnotaxa, such as
Scoyenia gracilis and Spongeliomorpha milfordensis, support their formation in firm
substrates resulting from desiccation (Metz 1995). Under extremely dry conditions,
all other ichnotaxa disappear, and only Spongeliomorpha and Scoyenia are present.
Collectively, the Lockatong ichnofauna represents the Scoyenia Ichnofacies (Metz
1995). A slightly more complicated picture is revealed by the Passaic Formation.
Overall, this unit displays similar ichnologic characteristics to the Lockatong
Formation, namely the widespread presence of the Scoyenia Ichnofacies in lake-
margin deposits, and the presence of Spongeliomorpha and Scoyenia as the only
ichnotaxa present in sediments deposited under extremely arid conditions (Metz
1996). However, some elements of the Mermia Ichnofacies are present in this unit as
well, illustrating transitions from subaqueous portions during the initial phase of lake
regression (Metz 1996).
Another extensively studied continental unit, containing well-exposed lacustrine

intervals and represented by abundant trace fossils, is the Middle to Upper Triassic
Agua de la Peña Group of the Ischigualasto–Villa Unión Basin of western Argentina
(Melchor et al. 2003; Melchor 2001, 2004, 2007). As in the case of the Newark
Supergroup, this rift basin developed during the breakup of Pangea (Uliana and
Biddle 1988; Milana and Alcober 1994). In particular, the most abundant and
diverse ichnofaunas occur in the Los Rastros Formation, which represents sedimen-
tation in a shallow, overfilled lake characterized by successive prograding delta
deposits (Melchor 2007). Deposits are typically arranged in coarsening-upward
parasequences encompassing prodelta, delta-front, and delta-plain facies. The high-
est ichnodiversity occurs in the distal delta-front deposits, which are dominated by
grazing trails (Helminthoidichnites tenuis, Gordia marina, Archaeonassa fossulata,
Cochlichnus anguineus) and fish trails (Undichna britannica, U. bina, U. cf. inso-
lentia). Also present are feeding (Treptichnus pollardi), dwelling (Palaeophycus
tubularis), locomotion (Bifurculapes isp., Cruziana problematica, Diplopodichnus
biformis, Didymaulichnus lyelli, Diplichnites isp., Protichnites isp.) and resting
(Rusophycus stromnessi, Avolatichnium isp.) structures (Melchor 2001). Middle
delta-front deposits display a similar ichnologic composition to distal delta-front
deposits, being dominated by grazing trails (Helminthoidichnites tenuis,
Helminthopsis abeli, Gordia indianaensis, Archaeonassa fossulata, Cochlichnus
anguineus), with fish trails (Undichna britannica) and dwelling structures
(Palaeophycus tubularis) also present (Melchor et al. 2003). Upper delta-front to
lower delta-plain deposits reflect a decrease in ichnodiversity that parallels a shal-
lowing of the lake, with only Palaeophycus tubularis, Skolithos isp., and Cochlichnus
anguineus recorded. Upper delta-plain deposits contain dwelling trace fossils, some
of which contain striations (e.g. Palaeophycus striatus) and vertebrate trackways
(Rhynchosauroides isp.). Collectively, the Los Rastros ichnofauna reflects the verti-
cal transition from the Mermia to the Scoyenia Ichnofacies as a result of shallowing
caused by deltaic progradation.
The Middle to Upper Triassic Madygen Formation of Kyrgyzstan contains abun-
dant trace fossils formed in an overfilled lake (Voigt and Hoppe 2010; Voigt et al.
2016). Lake-margin deposits are characterized by pervasive root trace fossils (Voigt
et al. 2016) (Fig. 11.3a). The most abundant trace fossils in this unit occur in perma-
nent subaqueous deposits, and consist of horizontal networks of multiple-branched
burrows (Voigt and Hoppe 2010) (Fig. 11.3b–e). These structures have been com-
pared with Thalassinoides by Voigt and Hoppe (2010), but were placed in the ich-
nogenus Virgaichnus by Knaust (2010). Further work is required to unravel the
ichnotaxonomic affinity of these burrows. Regardless of these ichnotaxonomic
complexities, these trace fossils represent feeding structures of worm-like deposit
feeders. These burrows occur at their highest densities in deposits formed around the
sublittoral–profundal boundary, probably coincident with the paleo-thermocline and
chemocline (Voigt et al. 2016). It has been speculated that the producers of these bur-
rows may have been able to tolerate dysoxic conditions, allowing them to colonize
deeper parts of the lake which may have acted as a refugium from predators (Voigt
et al. 2016). Other structures documented in these deposits are the grazing trail
Helminthoidichnites tenuis (Fig. 11.3f), unidentified sand-filled radiating burrow

systems, and ribbon-like burrows with transverse segmentation, which have been
interpreted as branchiopod locomotion traces (Voigt and Hoppe 2010; Voigt et al.
2016).
Fig. 11.3 Characteristic trace fossils from Middle to Upper Triassic lacustrine deposits of the
Madygen Formation of Kyrgyzstan. (a) Root trace fossils in lake-margin deposits; (b) General
view of a sandstone surface with horizontal networks of multiple-branched burrows in permanent
subaqueous lacustrine deposits; (c) and (d) Close-up of networks showing branching patterns; (e)
Preservational variation of horizontal networks as cleavage relief in mudstone; (f)
Helminthoidichnites tenuis in lacustrine deltaic deposits. All scale bars are 1 cm long, with the
exception of c and d, which are 0.5 cm long
Globally, Middle to Late Triassic ichnofaunas from lake-margin deposits are of

modern aspect and record the widespread establishment of the Scoyenia Ichnofacies.
In addition, the presence of trace-fossil suites dominated by the crayfish burrow
Camborygma records the appearance of the homonymous ichnofacies (see Chap. 13).
Middle to Late Triassic, fully lacustrine ichnofaunas resemble trace-fossil assem-
blages described from similar, late Paleozoic settings (Buatois and Mángano 1993a;
Buatois et al. 1998a; see Chap. 6). However, more penetrative trace fossils occur, as
revealed by the presence of networks of irregularly branched burrows in the
Madygen deposits (Voigt and Hoppe 2010). In addition, higher-energy sandy, storm,
and mouth-bar deposits tend to contain relatively deep vertical burrows, such as
Skolithos, Arenicolites, and Polykladichnus (Bromley and Asgaard 1979; Mángano
et al. 1994; Bromley 1996) (Fig. 11.4), representing freshwater equivalents of the
Skolithos Ichnofacies (Buatois and Mángano 2004, 2009).
In short, ichnologic data seem to support Sinitshenkova’s (2002) view that the
Triassic is a time of significant evolutionary innovation in lacustrine communities.
This is particularly evident in lake-margin settings which exhibit trace-fossil assem-
blages that would dominate these environments for the rest of the Phanerozoic. The
picture in fully lacustrine settings is slightly different because central lake deposits
display some ichnofaunas reminiscent of the late Paleozoic, with the addition of
more penetrative burrows typical of the rest of the Mesozoic and Cenozoic.
During the late Middle to Late Triassic there was a major, qualitative increase
in the abundance of lacustrine insect taxa. Fossil assemblages of this age contain a
prevalence of immature over adult aquatic insects, such that in many deposits
immature stages (naiads and larvae) outnumber adult specimens (Sinitshenkova
2002)—a situation that reverses that of the Permian. For this, and other reasons,
Sinitshenkova (2002) mentions that “…the Triassic [is] a starting point of a new,
Mesozoic evolutionary stage of lacustrine biocoenoses.” This novelty may be
Fig. 11.4 Arenicolites isp.

in deltaic mouth bar
deposits of the Upper
Triassic Tanzhuang
Formation of central
China. Scale bar is 1 cm
long
attributable to greater stability of the hydrological and water-chemistry conditions

in the physical environment of Eurasian, Late Triassic lakes (Kalugina 1980).
Also, part of the diversification event may be attributed to increased provinciality
from the breakup of Pangaea or to sizable increases in number and persistence of
long-lived lakes associated with the breakup (Cohen 2003).
Late Triassic aquatic taxa included new lineages representing a diversity of func-
tional feeding groups. For crustaceans, the major lineages were detritivorous ostra-
cods, conchostracans, notostracans and the bizarre central Asian lineage,
Kazakharthra. Dasyleptid bristletails were holdovers from the Permian, and per-
sisted until the Late Triassic along bodies of water. Mayflies diversified, with ben-
thic naiads assuming nektonic, epifaunal and infaunal filter-feeding strategies; some
benthic forms constructed U-shaped burrows, representing potential producers of
the biogenic structures typically found in lacustrine deposits of this age (e.g.
Bromley and Asgaard 1979; Mángano et al. 1994) (Fig. 11.4). Odonatans are repre-
sented by fewer fossil occurrences of naiads than adults, and all of the mostly ben-
thic naiads possessed a prominent, raptorial labial mask. The Plecoptera were
represented by obligately aquatic, benthic naiads, as were their Permian counter-
parts, and included the extinct benthic–lentic taxa of the Euxenoperlidae,
Mesoleuctra and Siberioperla (Sinitshenkova 2002). Grylloblattids (rock crawlers),
a surviving lineage from the Permian, may have inhabited lakeshores or even
shallow-aquatic habitats.
Most of the trophically dominant predators were streamlined, agile nectic heter-
opteran bugs and adephagan beetles. A few lineages of dominantly predatory,
aquatic heteropteran bugs appeared during or just before the Late Triassic, consist-
ing of the dominant naucoroid lineages of the extinct Triassocoridae, and the extant
Notonectidae (backswimmers) and Belostomatidae (giant water bugs) (Popov
1980). Aquatic adephagan lineages, such as the extinct Schizophoridae,
Ademosynidae, Colymbothetidae, and the extant Dytiscidae (predaceous diving
beetles) had similar dietary habits. The extant algivorous Haliplidae (crawling water
beetles) and its extinct Triassic relative, the Triaplidae, occurred on bottom sub-
strates and in aquatic plant entanglements (Fraser et al. 1996). Both adults and lar-
vae of the nectic Coptoclavidae were major predators, and had agile, active life
habits based on legs equipped for rapid movement and raptorial mouthparts, likely
feeding on small vertebrates and large insects. Megalopteran larvae are very similar
to present-day alderflies (Marchal-Papier 1998). A few taxa of Mecoptera (scorpi-
onflies) have been encountered, such as liassophilids that resemble modern aquatic
Nannochoristidae. The earliest, definitive, aquatic Trichoptera (caddisflies) are from
the Late Triassic, and include adults of the basal lineages Prorhyacophilidae and
Necrotaulidae, but also immatures that likely were predatory (Sinitshenkova 2002).
These lacustrine biotas still lacked a significant herbivore component, which
emerged later in the Mesozoic. Supporting this observation is the absence of sub-
merged or emergent aquatic macrophytes, which did not occur in significant abun-
dance until the Early Cretaceous. There is limited evidence, however, for a flora
fringing lakes and pools during the earlier Mesozoic, as indicated by the presence of
aquatic quillworts such as Isoetes, and Azolla-related ferns (Retallack 1997; Moisan
et al. 2012a, 2012b; Sun et al. 2014). In any case, the contribution was overwhelm-
ingly dead plant matter for detritivores rather than live plant tissues. For herbivores
as in the Early Triassic, charophytes continued to be the dominant macrophytes and
Chlorococcales the dominant phytoplankton (Martín-Closas 2003).
Six major community types have been recognized in Triassic Eurasian freshwa-
ter deposits (Sinitshenkova 2002). These community types are characterized by dis-
tinctive groupings of major taxa that occur in particular types of lakes and habitats.
Although Sinitshenkova’s (2002) general trophic analysis of lake ecosystems dem-
onstrated significant partitioning of available lotic and lentic resources, evidently
the herbivore guild was largely absent.
The most diverse Triassic insect assemblage is in an upper-Middle to lower-
Upper Triassic lacustrine deposit near the village of Madygen, in Kyrgyzstan, cen-
tral Asia, whose ichnofauna has been summarized above. In particular, one locality,
Dzhailoucho, and nearby outcrops have provided some of the best insights into later
Triassic aquatic and terrestrial life, including a diverse assemblage of 20 orders and
ca. 106 family-ranked lineages of insects, such as the earliest definitive occurrences
of Diptera (true flies) and Hymenoptera (sawflies, wasps, ants, and bees)
(Shcherbakov 2008c). The Madygen Biota consists of typical, early Mesozoic plant
groups, including cryptogams, sphenopsids, lycopsids, filicalean ferns, broadleaved
conifers, diverse peltasperms, corystosperms, a variety of ginkgophytes and rare
bennettitaleans (Dobruskina 1995). Aquatic invertebrates include several microcon-
chids, bivalves, gastropods, bryozoans, and a spectrum of crustaceans consisting of
phyllopods, ostracods, conchostracans, kazakharthrans, and malacostracan deca-
pods (Voigt et al. 2006). The vertebrate fauna is rather diverse, and is comprised of
a broad variety of fishes, such as lungfish, coelacanths, hybodontid, and xenacanthid
sharks, and ray-finned fish, including palaeoniscids, evenkiids, perleidids, and a
saurichthyid (Sytchevskaya 1999; Kogan et al. 2009; Fischer et al. 2011). The fauna
also includes six genera of lake-margin tetrapods, notably an early urodelan, basal
reptiliomorph, primitive cynodont, and three small diaspids including tree-climbing,
gliding, and insectivorous reptiles (Voigt et al. 2006; Schoch et al. 2010; Alifanov
and Kurochkin 2011). The lacustrine aquatic food web consisted of a variety of
lakeside plants, numerous insects and other aquatic invertebrates, fish and occa-
sional tetrapods (Shcherbakov 2008c). The aquatic portion of the biota included
eight orders of insects and ca. 25 families of primarily aquatic or amphibiotic
insects, of which a fifth of the lineages are extant. The Ephemeroptera included only
the single, extant family, Siphlonuridae (small minnow mayflies); by contrast, ten
families of Odonata were present, none of which are extant. Extinct Miomoptera
constituted one family, and Plecoptera were represented by two extinct families.
The Hemiptera included one family, the extant Ochteridae (velvet shore bugs). As
for holometabolous insects, aquatic Coleoptera were represented by five, early to
mid-Mesozoic, structurally streamlined families, but only one lineage, the
Hydrophilidae (water scavenger beetles) are extant, likely appearing as large, black,
and shining species. Similarly, the Trichoptera was represented by the basal and
extant Philopotamidae (fingernet caddisflies). Likewise, the Diptera consisted of
two new taxa, notably the diverse, extant Limoniidae (short-palped crane flies).
Neoichnologic data suggest that dipterans, in particular larval Limoniidae, pro-

duce zigzag burrows that in the fossil record may be referred to the ichnogenus
Treptichnus (Muñiz-Guinea et al. 2014). This ichnogenus is quite common in
Middle to Upper Triassic lacustrine deposits (Fig. 11.2e), which is consistent with
paleoentomologic information on the earliest definitive occurrences of Diptera.
However, Treptichnus is known in late Paleozoic lacustrine to fluvio-estuarine
deposits as well (Buatois and Mángano 1993b, c; Buatois et al. 1998b). Interestingly,
a molecular analysis placed the origin of crown group Diptera in the middle Permian
(Bertone et al. 2008).
11.5.3 Continued Jurassic Diversification and Increased

Infaunalization
Lower Jurassic, lake-margin ichnofaunas have been documented in both eastern

(Gierlowski-Kordesch 1991; Metz 1992; Collette et al. 2011) and western (Lucas
et al. 2006; Tanner and Lucas 2008) United States, whereas Upper Jurassic equiva-
lents are widespread only in the latter (Hasiotis 2004; Foster and Lockley 2006;
Hunt and Lucas 2006). Among invertebrate trace fossils, shallow-tier, meniscate,
backfilled structures with striated walls (Scoyenia) or without bioglyphs (Taenidium)
are typical, as are striated, branched burrows (Spongeliomorpha) (Gierlowski-
Kordesch 1991; Hasiotis 2004). Simple horizontal (Planolites, Palaeophycus) and
vertical (Skolithos) structures, together with crayfish burrows (Camborygma) and
banana-shaped structures (Fuersichnus), also are common (Gierlowski-Kordesch
1991; Hasiotis 2004; Tanner and Lucas 2008). Grazing trails (Helminthoidichnites)
and arthropod trackways (Diplichnites, Kouphichnium) are only abundant locally
(Lucas et al. 2006). Borings in stromatolites have been mentioned, but it is uncertain
if these were formed in lacustrine or marine coastal settings (Hasiotis 2004).
Although the earliest body-fossil record of dinosaurs is Late Triassic, dinosaur
tracks are known at least since the Middle Triassic (Marsicano et al. 2007; see Chap.
10). However, it is by the Early Jurassic that dinosaur tracks (e.g. Grallator,
Eubrontes) become the dominant vertebrate-generated structures in lake-margin
deposits (Hunt and Lucas 2006). Dinosaur tracks also occur in Lower Jurassic
deposits where the first megatracksites are recorded (Hamblin et al. 2006). These
megatracksites include trampled surfaces, although these seem to occur in wet inter-
dunes, rather than in lake margins (Seiler and Chan 2008). However, by the Late
Jurassic spectacular megatracksites are known from lake-margin deposits of the
Morrison Formation (Lockley et al. 1986; Jennings and Hasiotis 2006), among
other sites. The Morrison tracksites are associated with intense bioturbation and the
development of trampled surfaces (Lockley et al. 1986; Jennings and Hasiotis
2006). These Jurassic trampled surfaces are significant because, as demonstrated by
studies in Quaternary and modern lake-margin deposits (e.g. Laporte and
Behrensmeyer 1980; Ashley and Liutkus 2002), activities by large vertebrates play
a major role in disturbing primary sedimentary fabric.
Fully lacustrine ichnofaunas have been documented in the Lower Jurassic of the
eastern United States (Metz 1992) and China (Wu 1985; Buatois et al. 1995, 1996;
Hu et al. 1998; Uchman et al. 2011), the Middle Jurassic of England (Whyte et al.
2007), and the Upper Jurassic of the western United States (Hasiotis 2004). These
ichnofaunas show a combination of shallow-tier, simple trails and burrows
(Helminthopsis, Helminthoidichnites, Palaeophycus, Diplopodichnus, Paracan-
thorhaphe) and deeper-tier, branching burrow systems (Vagorichnus) (Metz 1992;
Buatois et al. 1996; Uchman et al. 2011). Vertical U-shaped burrows (Arenicolites)
(Hasiotis 2004) and bivalve burrows (Lockeia) (e.g. Whyte et al. 2007) are present
as well. As in the case of earlier assemblages, the fish trail Undichna is the typical
vertebrate ichnotaxa in this setting (Whyte et al. 2007).
The Lower Jurassic Anyao Formation of central China is arguably the most stud-
ied deep-lacustrine succession of this age, providing valuable insight into the colo-
nization of a fully lacustrine setting (Wu 1985; Buatois et al. 1995, 1996; Hu et al.
1998; Uchman et al. 2011). This formation represents deposition in a deep over-
filled lake system developed in a pull-apart basin (Buatois et al. 2000b). The Anyao
trace fossils are mostly present in thin-bedded turbidites which were formed in
lobe-fringe areas of lacustrine turbidite systems. This ichnofauna is relatively
diverse, encompassing both pre- (open burrows casted by the incoming turbidite
sand) and post- (burrows penetrating from a colonization surface at the top of the
turbidite sand) turbidite suites (Buatois et al. 1996) (Fig. 11.5a–i). The pre-event
suite consists of Helminthopsis abeli (Fig. 11.5a), Helminthoidichnites tenuis
(Fig. 11.5b), discrete specimens of Tuberculichnus vagans (Fig. 11.5c–d),
Monomorphichnus lineatus, Paracanthorhaphe togwunia (Fig. 11.5e–f), and thin
irregular trails. The post-event suite consists of Vagorichnus anyao (Fig. 11.5g–h),
Cochlichnus anguineus (Fig. 11.5i) and Helminthopsis hieroglyphica. Specimens of
Gordia marina and Tuberculichnus vagans intergrading with V. anyao also are part
of this suite. The ichnofauna is dominated by feeding and grazing trace fossils pro-
duced by deposit feeders. From an evolutionary perspective, the Anyao ichnofauna
is significant because it is dominated by infaunal burrows that reflect the activity of
invertebrates that were able to penetrate into discrete sandy layers for subsequently
expanding along sand–mud interfaces, a behavior unknown in older deposits in
similar environmental settings. The producer of these burrows is uncertain, although
it has been noted that in modern environments oligochaetes construct structures that
radiate from a surface tube and branch horizontally within the sediment (Chamberlain
1975; Buatois et al. 1995). Amphipods and isopods may potentially construct simi-
lar structures, but the paucity of modern analogues precludes further evaluation
(Buatois et al. 1995). In addition, this behavior shows the appearance of more orga-
nized branching patterns, such as Paracanthorhaphe togwunia, that reveal an
increase in complexity with respect to Paleozoic lacustrine ichnofaunas.
Overall, the taxonomic composition of invertebrate ichnofaunas in lake-margin
Jurassic deposits is quite similar to that of the Middle to Upper Triassic. No major
evolutionary novelties or innovations are apparent in these settings from the perspec-
tive of invertebrate ichnnology. However, the appearance of megatracksites and
trampled surfaces represents a major change with respect to older deposits, implying
substantial sediment reworking and modification by dinosaurs. This was probably

the first time in the history of the biosphere that lake-margin landscapes were signifi-
cantly transformed at a large scale by biogenic activity.
In addition, trace-fossil information from fully lacustrine environments pro-
vides a different picture to that of older deposits. Ichnodiversity levels are similar
Fig. 11.5 Characteristic trace fossils preserved at the base of Lower Jurassic lacustrine turbidites
of the Anyao Formation of central China. (a) Helminthopsis abeli; (b) Helminthoidichnites tenuis;
(c) General view of Tuberculichnus vagans displaying a meandering pattern; (d) Close-up of
Tuberculichnus vagans; (e) and (f) Paracanthorhaphe togwunia; (g) Vagorichnus anyao cross-
cutting and inorganic sole mark; (h) Close-up of Vagorichnus anyao; (i) Cochlichnus anguineus.
All scale bars are 1 cm long
to those of the Middle to Late Triassic, and even to those of the late Paleozoic, but
the principal difference is the degree of infaunalization, as revealed by the wide-
spread occupation of mid tiers in deep lacustrine sediments.
During the Early Jurassic there was further taxonomic, morphologic, and eco-
logic diversification of continental aquatic biotas. Much of the Jurassic zooplankton
was represented by diverse cladoceran crustaceans, indicated by fossil occurrences
(Zherikhin and Kalugina 1985), supported by molecular-clock data from phyloge-
netic analyses (Colbourne and Hebert 1996). One of the more distinctive differences
from Triassic aquatic insect assemblages was the greater presence of immature
stages (naiads, larvae) in Jurassic deposits, a consequence of a different preserva-
tional mode. The opposite situation of adult stages predominating over immature
stages was much less common, and may have been attributable to poor preservation
in habitats frequented by adults or the presence of adults in distant, more upland
environments that have low preservation potential. Another distinction is the
absence of virtually all Permian insect faunal elements and the first occurrences of
many modern family-level lineages (Sinitshenkova 2002).
The dominant insect groups of Jurassic lotic and lentic ecosystems represent a
spectrum of lineages that would occur in the same habitats today, although not nec-
essarily with the same ecologic tolerances or proportional abundances as their
present-day descendants. Ephemeropterans and odonatans were significantly more
speciose than their Triassic equivalent taxa. Ephemeropterans experienced a major
diversification, with the naiads of many groups, such as the Siphlonuridae and
extinct Mesonetidae and Epeorominidae having laterally positioned abdominal gills
and reaching considerably larger sizes than their Triassic confamilial antecedents.
Currently, ephemeropterans occur in high numerical abundance at local scales, are
present at significant diversities, and apparently were the major collector and filter-
feeding functional feeding groups (Merritt and Cummins 1984) in many Jurassic
localities. The Jurassic mayfly fauna is found in almost every aquatic environment,
in streams ranging from cascade-like rhythral settings, to calm water sites on flood-
plains such as oxbow lakes and overbank ponds, to lowland playas, and highland
graben-formed lakes. Like ephemeropterans, odonatans were diverse, best illus-
trated by several, large, dragonfly genera which co-occur in the Solnhofen Formation
of southern Germany, including a relict lineage of late Paleozoic meganeurid forms.
Jurassic odonatan naiads typically resemble modern damselfly naiads with a narrow
body and three, flap-like, heavily sclerotized, and terminal abdominal structures, the
cercus and paracerci. The formidable labial-masks of naiads indicate predation on
other larger aquatic insects and small vertebrates, such as fish and tadpoles. In some
lacustrine environments odonatan naiads likely were top predators in the absence of
fish (Sinitshenkova 2002).
Plecopteran nymphs occur commonly in Lower Jurassic deposits and are repre-
sented by three life-habit groups that are different from their modern representatives
taxa. The Mesoleuctridae possessed unusually long and slender legs, suggesting an
unusual detritivorous feeding habit. By contrast, the Platyperlidae bore an opposite,
flat habitus, with exceptionally squat, wide legs indicating a benthic insectivorous
diet. The Siberioperlidae, judging from robust mouthparts unusual for the Plecoptera,
were consumers of fibrous or otherwise indurated plant tissues (Sinitshenkova 2002).

An unrelated group is the extinct Chresmodidae of the Archaeorthoptera. The family
Chresmodidae includes large, water-strider-like insects presumably occurring on
water surfaces. Chresmodids have had a checkered history of being taxonomically
poorly resolved and having equally contentious dietary preferences. Some speci-
mens display mouthparts designed for active predation but possess external, saw-
tooth ovipositors that functioned for insertion of eggs into substrates such as aquatic
plant stems (Delclòs et al. 2008). Based on ovipositional lesions on preangiosper-
mous Mesozoic plants associated with lake deposits, the most likely hosts for ovi-
positing chresmodids were lycopsids (Moisan et al. 2012b) and horsetails (Kräusel
1958), or less likely a seed plant such as a bennettitalean (Pott et al. 2008).
Jurassic lotic and lentic biotal assemblages commonly are dominated by aquatic
heteropteran bugs or adephagan beetles, or both. Heteropteran aquatic bugs included
the dominant nepomorph taxa of Corixidae (water boatman) and Nepidae (water
scorpions), as well as giant water bugs, backswimmers, and the Mesoveliidae
(mesoveliid water striders), distant relatives of modern Gerridae (water striders) that
likely skimmed the water surface of lakes and streams. These taxa were predaceous,
except for mostly algivorous corixids and shurabellids (Popov 1971). Based on
mouthpart structure that employed a triturating device (Cobben 1978), these microv-
orous corixid and shurabellid microvores filter fed on unicellular or very small mul-
ticellular algae. Some heteropterans inhabited hypersaline playas and coastal
brackish lagoons or estuaries (Santiago-Blay et al. 2001). Like heteropteran bugs,
aquatic adephagan beetles were ecologically diverse and mostly predaceous,
although some were scavengers and microvorous consumers of algae. There is no
indication of macrovorous herbivorous forms.
During the Early Jurassic the dominant beetle clade is the Schizophoridae, but its
diversity and abundance decreases towards the latest Jurassic (Ponomarenko 1995).
During this time extinct aquatic lineages include large, predaceous Coptoclavidae
with active, nektonic larvae, the Liadytidae, possessing benthic larvae, and the
Parahygrobiidae, of unknown feeding habits. The extant, large Gyrinidae (whirligig
beetles) appear in significant numbers in Jurassic lentic deposits, consisting of pre-
daceous larvae and detritivorous adults. The Parahygrobiidae are known only from
larvae of mid-Mesozoic lake deposits and are presumed predaceous. In contrast to
the commonness of beetle taxa, alderfly and dobsonfly larvae of the Megaloptera are
rare and confined to lotic habitats.
Other holometabolan taxa are the larvae of scorpionflies, caddisflies, and true flies
which typically were rare at most sites. The extinct Liassophilidae belongs to a scor-
pionfly lineage that survived into the Early Jurassic, and whose modern close relatives
are the primitive, aquatic Nannochoristidae (Novokshonov 1997). Caddisflies are rep-
resented primarily by the Necrotaulidae and the larger complex of families constitut-
ing the Phryganeina. The Phryganeina appeared during the Late Jurassic (Sukatcheva
1991) and their presence is indicated by some of the earliest caddisfly cases in the
fossil record. Caddisflies were mostly microvores, and likely were algivorous, detri-
tivorous, or omnivorous, commonly occurring in eutrophic environments (Zherikhin
and Kalugina 1985). Unlike other abundant holometabolan groups, true flies represent
a major radiation throughout the Jurassic, particularly involving several nematocerous

lineages, principally the Limoniidae, Chironomidae (midges), Chaoboridae (phantom
midges), Psychodidae (moth flies), Simuliidae (black flies), Dixidae (nonbiting
midges), and the extinct Eoptychopteridae, a clade related to modern phantom crane
flies (Kalugina 1980). These seven dipteran lineages included disparate mouthpart
modifications for passive and active filter feeding, net sieving, collecting, and mandi-
ble-assisted chewing, indicating that they were dominantly microvores. Microvory in
nematocerous Diptera is typically achieved by the pumping of water currents into a
specialized mouthpart filter or sieve for capture of small particulate matter such as
diatoms, protists, ostracods, and a variety of other microorganisms.
The Jurassic also witnessed the appearance of major lineages of teleost fish,
frogs, crocodilians, and aquatic birds (Cohen 2003). These lineages represent evo-
lutionary novelties that may have been conducive to key innovations, mostly
increased predation in freshwater settings, which in turn may have resulted in the
disappearance of large-bodied branchiopods and other slow moving crustaceans
from many lacustrine settings (Webb 1979; Cohen 2003). Since the Jurassic, these
taxa seem to have been displaced to more marginal habitats, such as saline lakes and
vernal pools, characterized by low diversity of predators. Also, the coincidence of
increased predation and infaunalization of the lacustrine benthos may be significant.
As indicated by Vermeij (1987), the infaunal environment typically is regarded as a
haven from predation and, therefore, infaunalization may be regarded as an adap-
tive response to increased predation.
Relatively little is known about aquatic vegetation during the Jurassic. The domi-
nance of charophytes is apparent, with porocharaceans and nitellaceans occurring in
Early to Middle Jurassic Chinese basins (Wang et al. 1976), probably reflecting
accommodation to ephemeral lakes (Martín-Closas and Serra-Kiel 1991). By the
Late Jurassic, two charophyte families underwent diversification, the Characeae and
the Clavatoraceae, which were associated with oligotrophic and alkaline lakes
(Martín-Closas and Serra-Kiel 1991). This may have resulted in the displacement of
porocharaceans to higher latitudes and brackish-water settings (Martín-Closas
2003). Nonflowering vascular plants were represented by relatives of Azolla ferns
along littoral zones (Cohen 2003). An incipient colonization of ponds by ferns also
is apparent by the Late Jurassic (Martín-Closas 2003), particularly the
Hymenophyllaceae (filmy ferns), that preferred hydric habitats (Hennequin et al.,
2008). Lacustrine phytoplankton retained its ancestral condition, with Chlorococcales
remaining dominant (Martín-Closas 2003).
Beginning toward the end of the Early Jurassic, and culminating during the
Middle to Late Jurassic, a distinctive, fluvio-lacustrine ecosystem is recognized in
Eurasia. This ecosystem represents a continuation of earlier, similar, Late Triassic
ecosystems, but is characterized by a suite of physical, chemical, and biological
features that were distinct from other contemporaneous and later ecosystems
(Table 11.1). The overwhelmingly Jurassic biotas that occupy this ecosystem are the
“Assemblage B” of Sinitshenkova and Zherikhin (1996), characterized by a
detritivore-based food web, in which primary production consisted of coarse and
fine detritus and dead benthic algae. These sources of food were channeled to filter-
feeding and shredding invertebrates, particularly insects, but also crustaceans, mollusks,
and bryozoans (Table 11.1 and Fig. 11.6). Although this biota persisted into the
Early Cretaceous, the geochronologic acme of this community type was during the
Middle to Late Jurassic.
A prime example of an Assemblage B community is the Mesoleuctra–Mesoneta
assemblage at Ust’-Balei, in Transbaikalian Russia, representing a lacustrine deposit
from uppermost Lower Jurassic strata (Sinitshenkova and Zherikhin 1996). The
Mesoleuctra–Mesoneta assemblage was more productive, through the fixation of
organic carbon by chemotrophic and autotrophic organisms, than those from coex-
isting Assemblage A lakes. However, the Mesoleuctra–Mesoneta assemblage was
considerably less productive than Assemblage C lakes of the Early Cretaceous
(Table 11.1 and Fig. 11.6). This assemblage type may have occurred in several
Gondwanan sites, although documentation is poor. The dominant biotal features of
the Mesoleuctra–Mesoneta assemblage at Ust’-Balei was an epibenthic fauna that
consisted dominantly of mayfly and stonefly taxa, the absence of chaoborid dipteran
larvae, and the general presence of some algae, bivalves, and fish. Deposits contain-
ing an Assemblage B biota are associated with lignitic and coal-bearing strata,
paleobotanical indicators suggesting oxygen depletion (Samylina 1988), and a set-
ting in which microbial activity was strongly suppressed. At Ust’-Balei, the
Mesoleuctra–Mesoneta assemblage inhabited an aquatic environment similar to a
modern, disaerobic peat lake wherein bacterial metabolic activity is limited by
water acidity (Kuznetsov 1970).
Jurassic lacustrine deposits are present in North America as well (see review of
ichnologic literature above), but unlike those in Eurasia, they have been less studied
in a paleoentomological context, typically lacking body fossils, notably insects.
Deposits such as the deep, graben-fill deposits of the Late Triassic (Theismeyer
1939) and the more laterally persistent, epicontinental strata of the Late Cretaceous–
Early Paleogene temporally bracket the Middle Jurassic (Callovian) Sundance and
Late Jurassic (Tithonian) Morrison Formations. The Morrison Formation evidently
included a series or large, relatively shallow alkaline lakes, most of which were
intermittent and lacked significant freshwater input due to water limitation within
an arid, subtropical belt (Turner and Fishman 1991; Demko and Parrish 1998;
Engelmann et al. 2004). Although much is known of the vertebrates of landscapes
surrounding Morrison lakes, little is understood of their aquatic invertebrate faunas.
Lacustrine insects are unknown, and the only significantly studied invertebrates are
lacustrine ostracods (Schudack 1998) and bivalves occurring in associated fluvial,
floodplain and pond environments (Good 2004).
The Sundance Formation perhaps provides the sole exception of a Jurassic,
North American lacustrine biota on par with numerous Eurasian examples of the
same period (Fig. 11.7). The Sundance Formation occurs along the central-northern
Wyoming and central-southern Montana border, and provides paleoentomologi-
cally the best documented Jurassic lacustrine ecosystem in North America (Santiago-
Blay et al. 2001; Grimaldi and Engel 2005). In particular, the thinly laminated,
interbedded paper shales of the Hulett Sandstone Member provide considerable
evidence for a modestly diverse lacustrine fauna. The deposit was formed from
Table 11.1 Biological features of Mesozoic lakes before and after the Mesozoic Lacustrine Revolution (MLR) based on aquatic insect data. Modified slightly
from Sinitshenkova and Zherikhin (1996), with brown indicating more detritivore-based and green more herbivore-based lacustrine assemblages
204
Pre Mesozoic lacus trine revolution lake structure Post Mesozoic lacustrine revolution lake structure
a
Assemblage Type A B C D E
Age: Upper Lower J—Lower K Upper Lower J— Lower Ka Lower K Lower K
Lower Ka
1. Algal production Low Low High ?Moderate High
2. Macrophyte production Very low Moderate Low Low Low
3. Allochthonous detritus Low High Low ?Low ?Variable
4. Turnover rate Low High High High High
5. Microbial activity Low Low, suppres-sed by Moderate, high in Unknown ?High
antibio-tic leaf litter deep-water zone
6. Dominating trophic Detritivorous Detritivorous Herbivorous Herbivorous Herbivorous
chain
7. Trophic web complexity Low Low High Low Moderate
8. Dominating strategy K K ?r ?r r
9. Shreddersb Common Common Uncommon ?Uncommon ?Uncommon
10. Scrapersb Common Common Uncommon Uncommon Uncommon
11. Grazersb Rare Rare Common Common Common
12. Filtratorsb Very rare Rare Uncommon Common Common
13. Planktonivoresb Moderately common Rare Very abundant Very abundant Very abundant
14. Predatorsb Common Common Common Rare Common
15. Modern analogues Cold, clear-water None None Unknown None
oligo-trophic lakes
16. Ecosystem type Montane; oligotrophic to Hypotrophic (non-acid Pseudoligo-trophic ?Mesoligo-trophic ?Pseudoligo-trophic.
ultraoligotrophic dystrophic) (productive
nonsaprobic)
The vertical line separating Assemblages B and C indicates the timing of the MMLR transition
a
The five assemblages (A–E) are further detailed in Sinitshenkova and Zherikhin (1996)
b
Macroinvertebrate functional feeding groups (after Merritt and Cummins 1984)
L.A. Buatois et al.
Fig. 11.6 Trophic structure reconstruction for primary producers and arthropods of a Middle
Jurassic hypotrophic lake (Zherikhin and Kalugina 1985). This lake represents the Mesoleuctra–
Mesoneta biotal assemblage typifying Lake Ust’Balei, in the Irkutsk Region of northern Siberia,
Russia. This trophic reconstruction corresponds to the Type B lake biotal assemblage of
Sinitshenkova and Zherikhin (1996). See Assemblage B in Table 11.1 for additional physical, chem-
ical and biological details. Redrawn, with modification, from Sinitshenkova and Zherikhin (1996)
freshwater to brackish water input and includes locally a back-barrier shoal and
tidal inlet consistent with a more estuarine environment during certain time inter-
vals (Uhlir et al. 2006; Stone and Vondra 2013). Vertebrate footprints and bone
material have been found in other strata of the Sundance Formation. The biota of the
Hulett Sandstone Member consists of pollen, land-plant fragments, aquatic insects,
and a leptolepid fish, presumably of continental provenance (Fig. 11.7a–p). The
insects are modestly diverse, consisting minimally of about 15 species, and include
a variety of typically freshwater nepomorph heteropterans, especially notonectids,
corixids, belostomatids, and possibly the extinct Enicocoridae. Adephagan beetles
include dytiscids and possible parahygrobiids, and elytra assigned to Holcoptera
are suggestive of a polyphagan lineage (Santiago-Blay et al. 2001; Grimaldi and
Engel 2005). Other faunal constituents are very rare caddisfly cases. Whereas much
of these strata, particularly those containing the insects, indicate lacustrine deposi-
tion, other strata indicate a more marine influence, suggesting harsh, occasionally
hypersaline conditions and an uninhabitable benthic environment. Palynomorph
and mesofossil land-plant material reveal a xerophytic coastal vegetation of cheiro-
lepidiaceous and araucariaceous conifers and possible gnetaleans indicated by
Eucommiidites pollen. A similar, approximately coeval lacustrine deposit, the
Todilto Formation, occurs in central New Mexico, is less diverse but has a similar
depositional environment with periodic influxes of fresh and brackish water alter-
nating with marine incursions from the east, and has a similar lacustrine biota
(Bradbury and Kirkland 1966; Anderson and Lucas 1996; Ulmer-Scholle 2005).
11.5.4 Persistence of Previous Trends During the Early

Cretaceous
Early Cretaceous lake-margin ichnofaunas have been documented in England

(Goldring et al. 2005), Spain (Moratalla et al. 1995; Moratalla and Hernán 2009),
Mongolia (Johnson and Graham 2004) and Korea (Kim and Paik 1997; Kim et al.
2002, 2005, 2012a,b,c; Paik et al. 2012; Lockley et al. 2012). Dinosaur (e.g.
Ornithopodichnus, Dromaeosauripus, Caririchnium, Minisauripus), bird (e.g.
Koranornis, Jindongornipes, Uhangrichnus, Ignotomis), crocodile, turtle
(Emydiphus), and pterosaur (e.g. Pteraichnus, Haenamichnus) tracks are extremely
common, in many cases forming megatracksites (Moratalla et al. 1995; Meyer et al.
2001; Moratalla and Hernán 2009; Lockley et al. 2012; Kim et al. 2012a,b,c).
As is the case for their earlier Mesozoic counterparts, Early Cretaceous lake-
margin invertebrate ichnofaunas are dominated by shallow-tier striated or nonstriated
meniscate, backfilled structures (Scoyenia, Taenidium), and simple horizontal
(Planolites, Palaeophycus) and vertical simple or U-shaped (Skolithos,
Diplocraterion) features (Zhang 1987; Kim and Paik 1997; Kim et al. 2002, 2005;
Johnson and Graham 2004; Goldring et al. 2005). Arthropod trackways (Diplichnites),
simple grazing trails (Helminthopsis, Cochlichnus), and branching burrows
(Thalassinoides) occur locally (Kim et al. 2002, 2005; Goldring et al. 2005).
Fig. 11.7 The late Middle Jurassic (Callovian, 165 Ma) Sundance Biota from the lacustrine
Sundance Formation, central-northernmost Wyoming and adjacent central-southernmost Montana,
United States (Santiago-Blay et al. 2001). (a) Leptocerid fish (USNM-597438, Specimen 605); (b)
Leptocerid fish (USNM-597439, Specimen 1281); (c) Hemiptera: ?Belostomatidae (USNM-597440,
Specimen 554–39); (d) Hemiptera: Belostomatidae (USNM-597441, Specimen 775); (e) Hemiptera:
Corixidae (USNM-597442, Specimen 1377A); (f) Hemiptera: Corixidae (USNM-597443, Specimen
1801); (g) Hemiptera: ?Notonectidae (USNM-597444, Specimen 2688B); (h) Hemiptera:
Enicocoridae (USNM-597445, Specimen 2194); (i) Coleoptera: undetermined family (USNM-
597446, Specimen 24); (j) Coleoptera: undetermined family (USNM-597447, Specimen 2443); (k)
Coleoptera: (USNM-597448, Specimen 2501); (l) (USNM-597449, Specimen 1936B); (m) (USNM-
597450, Specimen 2398B); (n) (USNM-597451, Specimen 2260); (o) (USNM-597452, Specimen
2906); (p) Coleoptera: (USNM-597453, Specimen 2149). Scale bars: solid, 10 mm; striped, 1 mm
Early Cretaceous examples of fully lacustrine ichnofaunas have been recorded in

Spain (de Gibert et al. 1999, 2000, 2016; Buatois et al. 2000a) and Brazil (Buatois
and Mángano 1998). Information is sparse, and examples are known from quite dif-
ferent lacustrine settings, further complicating the establishment of general pat-
terns. The Spanish paleolake deposits are carbonates, and their ichnofaunas are
dominated by very shallow-tier trails and burrows (Gordia, Cochlichnus,
Steinsfjordichnus, Cruziana, Helminthoidichnites, Palaeophycus, Treptichnus,
Planolites), with a very minor contribution of arthropod trackways (Hamipes) (de
Gibert et al. 2000, 2016; Buatois et al. 2000a) (Fig. 11.8a–d). The vertebrate com-
ponent of the subaqueous biota is represented by the fish trail Undichna (de Gibert
Fig. 11.8 Characteristic trace fossils from Lower Cretaceous fully subaqueous lacustrine carbonate
deposits of Las Hoyas, Spain. (a) Cruziana isp.; (b) Helminthoidichnites tenuis; (c) Palaeophycus
tubularis; (d) Surface containing high density of poorly developed Treptichnus pollardi; (e)
Undichna unisulca. All scale bars are 1 cm wide, with the exception of e, which is 2 cm long
Fig. 11.9 Ichnofabrics from the Lower Cretaceous lacustrine shoreface deposits of the Coqueiro
Seco Formation of northeast Brazil. (a) General view of cross-bedded pebbly very coarse-grained
sandstone (foreground) interbedded with intensely bioturbated fine-grained sandstone (back-
ground); (b) Close-up of intensely bioturbated sandstone displaying mottled texture. Lens cap
diameter is 5.5 cm long
et al. 1999) (Fig. 11.8a–e). In contrast, the Brazilian ichnofauna is derived from
intensely bioturbated, lacustrine, clastic shoreface deposits, and displays mottling
attributable to Planolites (Buatois and Mángano 1998) (Fig. 11.9a–b).
Lower Cretaceous shallow-lake deposits present within the nonmarine Sindong
Group of South Korea are useful to illustrate the major characteristics of lacustrine
ichnofaunas from this age (e.g. Kim and Paik 1997; Kim et al. 2002, 2005; Paik
et al. 2012). In particular, the Jinju Formation has been the object of several ichno-
logic studies (e.g. Kim et al. 2002, 2005, 2012c). This unit was deposited in a shal-
low lake periodically affected by immersion and desiccation. The ichnofauna
contains a variety of feeding (Beaconites antarcticus, B. coronus, Planolites annu-
laris, P. beverleyensis, Taenidium barretti), locomotion (Octopodichnus cf. didacty-
lus, Diplichnites ispp., unassigned sauropod trackways), dwelling (Palaeophycus
sulcatus, P. tubularis, Skolithos magnus), and grazing (Cochlichnus anguineus,
Helminthopsis hieroglyphica) traces (e.g. Kim et al. 2005). The vast majority of
these ichnotaxa are common in the Scoyenia Ichnofacies, which is consistent with
the lake-margin interpretation for these deposits. The sharply defined margins of the
trackways suggest firm substrates. However, Kim et al. (2005) noted that some of
the grazing trails may have been emplaced in softgrounds, implying colonization
under subaqueous conditions and transition to the Mermia Ichnofacies. Potential
producers include several groups of dipterans as well as nematodes and spiders.
Overall, the Early Cretaceous, South Korean lake ichnofaunas show a similar trace-
fossil distribution to those ichnofaunas recorded in shallow-lacustrine deposits of
the Newark Supergroup. In both cases, colonization by benthic organisms reflects
changes in the degree of consolidation of the substrate as a result of progressive
subaerial exposure.
Early Cretaceous lotic and especially lentic insect taxa are derived from the same
taxonomic families as those of the Late Jurassic (Martínez-Delclòs et al. 1995;
Delclòs et al. 2008). Representatives of this faunal assemblage are a lentic, ephem-
eropteran fauna that consists of siphlonurids, hexagenitids and mesonetids.
Hexagenitid naiads, as well as ephemeroid naiads, are very common in some Early
Cretaceous lentic assemblages (Sinitshenkova 2002), including taxa that produced
the bioerosion ichnogenus Asthenopodichnium subaqueously in woods (Thenius
1989). Most of the Early Cretaceous mayfly genera did not survive into the Late
Cretaceous (Sinitshenkova 2002), and only one lineage, the Australiephemeridae,
became extinct at the end-Cretaceous (K-Pg) boundary.
The Early Cretaceous aquatic odonatan fauna is quite different from that of the
Jurassic. The diversification of modern anisopteran (dragonfly) and zygopteran (dam-
selfly) lineages appear for the first time. Curiously, the Pseudomyrmeleontidae, inter-
preted by some as related to the Paleozoic Meganeuroidea, were still present as a relict
group. Unlike ephemeropteran assemblages, odonatan assemblages appear to have
most of their evolutionary development in lotic habitats. The dominant Early Cretaceous
odonatan lineages are the Isophlebioidea, Heterophlebioidea, Calopterygoidea, and
other anisopteran clades. Among these lineages are a few specialized types, such as
naiads of the Eurasian Hemeroscopidae and Sonidae (Pritykina 1986), which include a
morphotype with long legs possessing paddling locomotion and a nectic, predatory
existence. By contrast, other taxa, such as the Nothomacromyiidae lacked swimming
structures but retained actively cursorial, long legs, indicating predatory pursuit habits.
Another ecomorph is represented by the short-legged, burrowing Gomphidae (Bechley
1998). The labial-mask mouthpart structure has been examined for a select few species
of these lineages, and no conclusions could be made regarding prey-specific relation-
ships (Sinitshenkova 2002).
Plecopteran family-level diversity increased during the Early Cretaceous.
Jurassic plecopteran clades persisted into the Early Cretaceous and were supple-
mented by the earliest occurrences of several, modern, family-level lineages. All of
the modern, plecopteran functional feeding groups were present during the Early
Cretaceous, such as collectors and shredders engaged in detritivory, algal herbivory
and insect predation. In addition, chresmodids are still present in some lacustrine
settings, but probably became competitively excluded by emerging, Early Cretaceous
gerrid lineages.
True bugs of the Hemiptera are ubiquitous and diverse throughout the Early
Cretaceous, and with the exception of the Shurabellidae, are continuations of
Jurassic lineages (Sinitshenkova 2002). By contrast, the Gerridae and Mesoveloidea
(water treaders) initially appear during the Early Cretaceous, ecologically supple-
menting and replacing an earlier convergent ecotype, the orthopteroid Chresmodidae,
which had affinities to orthopteroid insects (Martínez-Delclòs et al. 1995). This
replacement included larger-sized, surface-skimming gerrids during the Late
Cretaceous that approached in size many of the medium-sized Jurassic chresmo-
dids, the latter of which had legspans of up to 16 cm (Labandeira pers. observ.).
Corixids, notonectids, mesotrephids and notably large, predaceous belostomatids
continued into the Early Cretaceous (Delclòs et al. 2008). Although there are some
evolutionary novelties between Jurassic and Early Cretaceous aquatic bug faunas, it
appears that all of the Jurassic ecologic feeding types were present during the Early
Cretaceous and encompassed surface water and neuston-zone feeders on dead or
dying arthropods, nectic algivores, nectic predators and epibenthic predators.
There are minimal taxonomic and ecologic differences between Late Jurassic
and Early Cretaceous beetle faunas. Archostematan schizophorids became very
rare; the largely aquatic Adephaga retained a similar spectrum of major taxa as
before, although the proportions of major lineages change. The major shifts are that
the Parahygrobiidae become more abundant; the Coptoclavidae become dominated
by a newly emerging, advanced clade, the Coptoclavinae; the Gyrinidae increase in
diversity; the Liadytidae become rarer; the Dytiscidae remain rare; and the
Haliplidae have their earliest occurrence (Ponomarenko 1969). The dominantly ter-
restrial Polyphaga are represented by the same families in the Early Cretaceous as
during the Jurassic, including the Scirtiidae (marsh beetles), but especially the detri-
tivorous Hydrophilidae, represented by new taxa such as Cretotaenia.
Other holometabolous insect clades with aquatic stages include the Megaloptera,
Neuroptera (lacewings, antlions, and related forms) and Mecoptera. These clades
express a similar pattern of mostly lotic-based diversity, as do their descendants in
modern ecosystems. The predaceous larvae of megalopterans, such as the Corydalidae
(dobsonflies), neuropterans such as the Osmyloidea (net-winged lacewings and rela-
tives) and the detritivorous mecopteran larvae of the Nannochoristidae occur in Lower
Cretaceous deposits of Eurasia and Australia (Zherikhin 1978; Jell and Duncan 1986).
These lineages currently reside in the same habitats, although they have been overshad-
owed by more derived Cenozoic lineages that entered the same aquatic niches.
During the Early Cretaceous, caddisflies experienced major evolutionary novel-
ties, as new family-level lineages emerged, other groups assumed dominance, and a
profusion of larval case-making activity ensued. Evidence from adult caddisflies
indicate that the earliest Mesozoic caddisflies, necrotaulids, became rare; others,
such as the extinct phryganeoid families Dysoneuridae, Vitimotaulidae, and
Baissoferidae were common, as were the extant Phyganeoidea, the Calamoceratidae
(flat-case caddisflies), Lepidostomatidae (lepidostomatid casemaker caddisflies),
Plectrotarsidae (plectrotarsid caddisflies), and Helicopsychidae (snail-case caddis-
flies) (Sinitshenkova 2002). Of these, the Vitimotaulidae were the most numerically
abundant, or at least were common across most Eurasian localities. There was
significant differentiation of the Vitimotaulidae at the generic level across Eurasia,
and the dominance of particular local genera occurred throughout the region.
An important event in lotic and especially lentic environments was the global
diversification of caddis larval-case morphotypes (Sukatcheva 1982). This expansion
of domicile morphotypes that used a variety of mineral, plant, and animal resources
evidently commenced during the earliest Cretaceous in northern Asia, and penecon-
temporaneously in Europe, South America and Australia (Sinitshenkova 1999).
Circumstantial evidence indicates that the Vitimotaulidae were the dominant fabrica-
tors of these cases. In localities where adult body fossils and larval cases are both
abundant and diverse, it appears that each case morphotype likely is associated with
one adult species. Apparently, each case morphotype was made by an adult species
that was restricted to a particular lotic or lentic habitat, partly reflected by use of
building materials that were available for case construction. Cases were variously fab-
ricated from conchostracan shells, Karkenia ginkgoalean seeds, conifer needles, plant
twigs, quartz grains, mica flakes, and other available materials (Sukatcheva 1982).
The dominant lacustrine lineages of the Diptera during the Early Cretaceous were
the same as for the Late Jurassic. Planktivorous chaoborids were more abundant but
less speciose during the Early Cretaceous when compared to the Late Jurassic,
whereas chironomids had an opposite trend. Both groups are represented primarily
by adult and pupal fossils; larvae are relatively rare and poorly preserved. This pat-
tern of differential preservation of developmental stages also exists for limoniids,
eoptychopterids and other nematocerous fly lineages. Not only compression deposits
capture this pattern, but this pattern also is found in amber occurrences, such as
Lebanese Amber. In Lebanese Amber the most common families are, in decreasing
rank: Chironomidae, Ceratopogonidae, Psychodidae, and Eoptychopteridae
(Kalugina 1980). However, in Paleolake Baissa deposits, chaoborids were very abun-
dant and likely were a major food resource for consumers that sustained much of the
upper tier of the food-web. One interesting development in Early Cretaceous lakes
was the emergence of brachyceran fly larvae, including the Stratiomyiidae (soldier
flies), Empididae (dance flies), other asilomorphs, and possibly the Sciomyzidae
(snail-eating flies) (Whalley and Jarzembowski 1985).
Arguably, one of the major innovations by the Early Cretaceous is the appear-
ance of aquatic angiosperms which, together with ferns (Schneider et al. 2004),
started to replace charophytes as the dominant macrovegetational elements in lacus-
trine ecosystems (Martín-Closas 2003; Friis et al. 2003, 2010; Gandolfo et al. 2004;
Coiffard et al. 2007). The presence of freshwater species representing among the
first angiosperms documented in the fossil record is hard to interpret, because it may
reflect an initial evolution and diversification in aquatic habitats or a taphonomic
bias (Martín-Closas 2003). Regardless of these alternatives, the appearance of
angiosperms was an evolutionary breakthrough that may have had a considerable
impact on aquatic life in ponds and lakes (Ponomarenko 1998; Friis et al. 2010).
Specifically, aquatic angiosperms (Sun et al. 2002; Friis et al. 2003; Dilcher et al.
2007) would have been instrumental in providing substrates for epiphytic organ-
isms, increasing the release of phosphorous by decaying macrophytes, promoting
light attenuation, steepening of the vertical temperature gradient, retarding of water
flow and enhancing fine-grained sediment deposition, among other processes
(Carpenter and Lodge 1986; Granéli and Solander 1988; Ponomarenko 1998; Cohen
2003). Paralleling this pattern, aquatic ferns experienced further diversification
(Schneider et al. 2004), but lycophytes remained abundant (Martín-Closas 2003).
With respect to phytoplankton, although Chlorococcales were still abundant, the
appearance of freshwater dinoflagellates is a major evolutionary innovation which
signaled the demise of green algal dominance in freshwater phytoplankton (Martín-
Closas 2003; Leliaert et al. 2011).
As with the Triassic and Jurassic, Cretaceous lotic and lentic insect faunas have
been divided into distinctive lacustrine assemblages, each characterized by specific
dominant and ecologically important insect species occurring in a particular environ-
mental setting (Zherikhin 1978; Sinitshenkova and Zherikhin 1996; Sinitshenkova
1999; Sinitshenkova 2002). For the Cretaceous, 18 such assemblages have been
described (Sinitshenkova 2002), one of which is Assemblage 5A, or the Ephemeropsis
melanurus–Hemeroscopus baissicus Assemblage, whose environmental conditions
and trophic structure have been examined in detail (Table 11.1 and Fig. 11.10). This
Fig. 11.10 Trophic structure reconstruction of an Early Cretaceous pseudoligotrophic lake

(Zherikhin et al. 1999). This lake represents the Ephemeropsis melanurus–Hemeroscopus bassicus
biotal assemblage typifying Paleolake Baissa in southern Siberia, Russia. This trophic reconstruc-
tion corresponds to the Type C lake biotal assemblage of Sinitshenkova and Zherikhin (1996). See
Assemblage C in Table 11.1 for additional physical, chemical and biological details. Redrawn,
with modification, from Sinitshenkova and Zherikhin (1996)
assemblage, from Paleolake Baissa, occurs in the Zaza Formation, Buryat Republic,
in Transbaikalian Russia, and was deposited in a deep, intermontane, meromictic
lake. Paleolake Baissa periodically dried out in shallower regions, indicated by mass
mortality occurring as dense accumulations of the same instar of large coptoclavid
larvae that became locally confined to a few bedding surfaces. Accumulations of
chaoborid larvae are present at more widespread event horizons and at small uncon-
formities or possibly diastems separating adjacent beds (Zherikhin et al. 1999).
The Ephemeropsis melanurus–Hemeroscopus bassicus Assemblage is more tax-
onomically and ecologically diverse than earlier Jurassic Eurasian paleolakes. This
assemblage includes tetrapods, birds, and osteoglossomorph fish such as Lycoptera,
and a rare Stychopterus sturgeon; phyllopod, conchostracan and ostracod crusta-
ceans, gastropods, bivalves and freshwater bryozoans (Vinogradov 1996). Most of
the estimated 1000 species of insects in the fauna are terrestrial and allochthonous
in origin (Zherikhin et al. 1999). The autochthonous lacustrine component of the
biota consists of ca. 80–120 species of lake residents, and was dominated by the
aquatic immatures of odonatans, heteropterans, coleopterans, dipterans and numer-
ous caddisflies (as cases). Insect fossils typically are shed exuviae rather than whole,
dead individuals. The surrounding vegetation consisted of Pseudolarix forest, a
Podozamites–Czekanowskia shrubland, and groundcover of horsetails, ferns, and
very rare angiosperms of uncertain affiliation (Zherikhin et al. 1999). Although this
assemblage represents one of the earliest occurrences of angiosperms in an Eurasian
lake deposit, the effect of angiosperms were insignificant, and it was toward the end
of the Early Cretaceous, during the Aptian and Albian Stages, during which the role
of angiosperms were manifest in terrestrial ecosystems (Labandeira, 2014;
Labandeira et al. 2016) and likely in freshwater ecosystems as well.
In most Mesozoic deposits intraspecific links between growth stages, such as
naiads and adults in nonholometabolous insects and larvae and adults in holometabolous
insects are not possible (Zherikhin and Sinitshenkova 2002). However, in the
Ephemeropsis melanurus–Hemeroscopus bassicus Assemblage, some larval and
adult taxa are associated based on a variety of evidence. Strong associations include
the dragonfly adult of the hemeroscopid Hemeroscopus baissicus and a very common
species of an aeschnidiid naiad, and similarly the hydrophilid beetle adult
Hydrophilopsia baissensis with the hydrophilid larva Cretotaenia pallipes. Adults and
larvae of the dobsonfly Cretochaulus lacustris and the adephagan beetle Coptoclava
longipoda are also found in this assemblage, although the latter may be a complex of
several species (Zherikhin et al. 1999). It appears that most of the caddisfly cases were
constructed by various species of Vitimotaulidae. However, many of these fossils are
immature individuals of lentic taxa whose adults are associated with nearby, lotic,
especially rhythral, habitats that were allochthonously transported into Paleolake
Baissa. The transported lotic taxa are represented by rare odonatans, plecopterans,
simuliid dipterans, and nannochoristid mecopterans. These connections between
immatures and adults in different aquatic habitats within lakes and rivers indicate a
greater autecologic partitioning within species by life stage, as well as among species.
This twofold, intraspecific and interspecific subdivision of resources can be inter-
preted as a maturation of terrestrial aquatic habitats that became more widespread
during the Early Cretaceous (Zherikhin et al. 1999).
11.5.5 The Late Cretaceous Biotal Replacement
Ichnologic information from Late Cretaceous paleolakes is remarkably scarce, pre-

cluding a detailed discussion. Late Cretaceous examples of lake-margin ichnofaunas
are known from the southern United States (Montgomery and Barnes 2012), western
Canada (Rylaarsdam et al. 2006), Botswana (Du Plessis and Le Roux 1995), Bolivia
(Meyer et al. 2001), China (Zhang 1987), and Korea (Paik et al. 2012; Lockley et al.
2012; Minter et al. 2012). However, most of these studies do not address the ichno-
faunas in a comprehensive fashion. As for fully lacustrine deposits, we are aware of
only one study documenting Late Cretaceous ichnofaunas in this setting, namely
perennial lake deposits in Botswana (Du Plessis and Le Roux 1995).
The crayfish burrow Camborygma is present in lake-margin deposits
(Montgomery and Barnes 2012), indicating the persistence of this deep tier through-
out the Mesozoic. Vertical burrows (e.g. Skolithos) and branching structures (e.g.
Thalassinoides) occur as well (Zhang 1987; Du Plessis and Le Roux 1995).
Arthropod trackways (Lithographus) have been recorded locally (Minter et al.
2012). Dinosaur and bird tracks preserved along lacustrine paleoshorelines also are
known (Meyer et al. 2001; Rylaarsdam et al. 2006; Paik et al. 2012; Lockley et al.
2012). In particular, intense bioturbation by dinosaurs continued to have an impact
on the sedimentary fabric of lake-margin deposits throughout the Late Cretaceous
(Rylaarsdam et al. 2006).
A complex mosaic of lacustrine deposits is preserved in Upper Cretaceous gra-
bens, whose infill is recorded in the Kalahari Group of Botswana (Du Plessis and Le
Roux 1995). The succession includes deposits of ephemeral and perennial alkaline
lakes. In both systems, an irregular network of tunnels, showing Y-shaped bifurca-
tions, attributed to Thalassinoides, has been recorded (Du Plessis and Le Roux
1995). In the ephemeral lake deposits, these structures are present in sand and mud
flats flanking the lake system. In the perennial lake deposits, Thalassinoides occurs
in fully lacustrine deposits in great abundances, commonly forming intensely biotur-
bated fabrics, which most likely record less saline conditions in relatively deeper
water environments (Du Plessis and Le Roux 1995). In these deposits, Thalassinoides
commonly occurs in association with vertical shafts that have lateral tunnels, which
have been compared with the poorly known vertical burrow Lennea (Du Plessis and
Le Roux 1995), but this attribution needs further confirmation. In short, the sparse
information available suggests continuation in lake-margin environments of the
same trends evidenced earlier in this setting. As well, the presence of intensely bio-
turbated, fully lacustrine deposits resembles similar deposits of the Lower Cretaceous.
Much less is known about Late Cretaceous lacustrine biotas when compared to those
of the Early Cretaceous. Most likely this is a reflection of limited outcrop that resulted
from the end of the broad extensional-tectonic regime that dominated the mid-Mesozoic
continental breakup, which promoted the formation of widespread lacustrine rifts
(Cohen 2003). The fossil record of aquatic insects documents mid-Cretaceous extinc-
tion of some lineages that were thriving during the Early Cretaceous but are not present
during the Late Cretaceous. This event included the extinction of about 20 family- and
superfamily-level lineages (Zherikhin 1978; Sinitshenkova and Zherikhin 1996). Of

ephemeropterans, none became extinct; for odonatans, the latest meganeuroid lineage,
the Protomyrmeleontidae, became extinct, as did the Isophlebioidea and
Heterophlebioidea. For plecopterans, the family Oecanthoperlidae is extinguished, as
was the Chresmodida, a major Jurassic clade of aquatic Archaeorthoptera. With the
exception of the Shurabellidae, apparently no lineage of hemipterans became extinct, as
early originating lineages such as corixids, notonectids, belostomatids, and mesoveliids
survive to the present day. Coleopterans experienced several major extinctions, such as
the Ademosynidae, Parahygrobiidae, Schizophoridae, Coptoclavidae, and Liadytidae.
No major extinctions are known for the major lineages of the relatively undiverse neu-
ropteroid groups Megaloptera, Raphidioptera, or Neuroptera, or for the Mecoptera, of
which the extant Nannochoristidae is an aquatic representative. For the Trichoptera,
major extinctions of family-level taxa were the Dysoneuridae, Necrotaulidae,
Baissoferidae, and Vitimotaulidae, the latter of which produced a high diversity of cad-
disfly case morphologies. The Diptera remained unaffected by the mid-Cretaceous
extinction events; evidently the nematocerous Eoptychopteridae and possibly the
Hennigmatidae are the only family-level lineages that became extinct (Zherikhin 1978;
Sinitshenkova and Zherikhin 1996).
Perhaps more important are the originations of new aquatic insect lineages during
the mid Cretaceous, almost all of which survive to the present and essentially form
the modern lentic and lotic aquatic insect fauna. Approximately 19 family- and
superfamily-level lineages have their first fossil occurrences during the mid
Cretaceous, consisting of approximately the same number of lineages that go extinct.
For the Ephemeroptera, only the earliest Heptagenoidea is known to occur close to
the Early–Late Cretaceous boundary. In odonatans, modern family-level lineages of
the Zygoptera (damselflies) may have originated during this interval, but geochrono-
logical timing is poorly constrained. No known major lineages of plecopterans origi-
nated during the mid Cretaceous. As for hemipterans, the diverse clade Gerromorpha
had its earliest major bout of diversification during the mid Cretaceous, including
some of the first occurrences of sublineages within the Gerridae, Hydrometroidea
(water measurers), probably the Hebroidea (velvet waterbugs and relatives), and the
extinct Mesotrephidae. Of coleopterans, only the Haliplidae and Scirtiidae have their
earliest occurrences during the mid Cretaceous. Five lineages of the Trichoptera have
their first occurrences at this time, including the Hydrobiosidae (pincer-clawed cad-
disflies), Sericostomatidae (bushedtailed caddisflies), Odontoceridae (mortarjoint
casemakers), Calamoceratidae (comblipped casemakers), and Leptoceridae (long-
horned caddisflies). The origin of these lineages curiously is associated with a sig-
nificant decline in case morphotype diversity, as early Cretaceous cases range from
10–15 morphotypes in a typical assemblage, to ca. 5 per assemblage in the Late
Cretaceous (Sukatcheva 1991). This decline may be attributable to the disappearance
of the diverse Vitimotaulidae which became extinct at this time. Several nematocer-
ous lineages of dipterans with aquatic larvae diversified during the mid Cretaceous,
particularly the Ptychopteridae (phantom crane flies), Culicidae (mosquitoes),
Chaoboridae (phantom midges), Corethrellidae (frog-biting midges), and
Ceratopogonidae (biting midges). Interestingly, while the number of known aquatic
insect extinctions and originations for the mid Cretaceous appear to balance out with
ca. 20 lineages in each category, the totals of insect herbivore turnover is much
greater for the mid-Cretaceous terrestrial realm (Labandeira 2014).
The biologies of these aquatic insect lineages indicate that, whereas the aquatic
immatures (naiads, larvae) were well integrated into aquatic ecosystems, the adults
became more removed from aquatic habitats than in previous freshwater biotas. For
example, whereas the naiads of damselflies were major aquatic predators, their adults
preyed on aerially winged insects. The adults of aquatic heteropterans became preda-
tors on other aquatic insects (Anderson 1998). Almost all dipteran lineages were com-
prised of larvae that were aquatic filter-feeders whereas their adults—at least the
females—were obligate blood feeders on vertebrates (Labandeira 2002). As the
aquatic larvae of caddisflies increasingly became herbivorous, their terrestrial adults
targeted angiosperm nectar or similar nutritive sources (Porsch 1958), or lacked feed-
ing capabilities altogether. The disjunction of habitats between immatures (naiads,
larvae) and adults increased during the Late Cretaceous, exhibiting an even more pro-
found separation between the life habits of immatures and adults than ever before.
The Late Cretaceous witnessed the continuation of the radiation of aquatic
angiosperms and ferns (Martín-Closas 2003; Gandolfo et al. 2004; Schneider et al.
2004), the former showing increased interaction with modern insect families. The
shift from gymnosperm-dominated floras to angiosperm-dominated floras was
accompanied by a major turnover in their terrestrial insect associates (Labandeira
2014; Labandeira et al. 2016). This transition occurred during the angiosperm radia-
tion, and included episodes of extinction and origination that paralleled a similar
pattern characterizing the MLR, although it appears that there is a time shift between
the two major events. By contrast, no such pattern is seen in freshwater algal groups.
Charophyte assemblages became dominated by the Characeae (Martín-Closas
2003). During the Late Cretaceous, lacustrine phytoplankton displays a similar
composition to that of the Early Cretaceous, as indicated by the presence of
Chlorococcales and freshwater dinoflagellates (Martín-Closas 2003). However, evi-
dence of diatoms is known from the Late Cretaceous, suggesting early diversifica-
tion, albeit playing a limited role as lacustrine plankton (Chacón-Bacca et al. 2002).
Chrysophytes also are documented from the Cretaceous (Cornell 1979; Adam and
Mahood 1981), but as with diatoms, they radiated later in the Cenozoic when they
became an important phytoplankton component (Leliaert et al. 2011). In any case,
the major changes in the plankton composition involved the diversification of fresh-
water diatoms and chrysophytes.
11.6 The Aftermath of the Revolution
An examination of the trace- and body-fossil records, as well as the ecological

structure of Cenozoic lakes, provides a picture of the aftermath of the MLR,
thereby revealing whether additional evolutionary novelties or innovations took
place. Interestingly, the biological response to the MLR is not straightforward.
A review of the Cenozoic lacustrine trace-fossil record shows that Cenozoic lake-margin
invertebrate ichnofaunas are similar in composition to those of the Late Cretaceous
(Table 11.2). Apparently there was minimal effect of the Cretaceous–Paleogene
(K-Pg) events on the MLR (see Chap. 12). This is in sharp contrast, for example,
with the ongoing rapid diversification of insect nesting structures in paleosols that
took place during the Cenozoic (see Chap. 13).
Little is known of complete inventories of bulk aquatic insect faunas from amber
and compression–impression deposits during the Late Cretaceous and Paleocene,
unlike the significantly better documented terrestrial fossil insect record (Zherikhin
and Sinitshenkova 2002; Labandeira 2005b). Perhaps more importantly, paleoeco-
logical data indicate that the modern ecologic structure from one, well-documented
lake was well established by the early–middle Eocene boundary interval at Paleolake
Messel (Fig. 11.12), and resembled in all measured food-web indices that of a vari-
ety of modern lake ecosystems (Dunne et al. 2014). This study proposes that earlier,
well preserved lake deposits that postdate the MLR but predate Messel (and strad-
dling the K-Pg boundary) also be examined to determine how far back in geologic
time modern lacustrine food-web structure extends (Dunne et al. 2014). Until food-
web studies that use similar analytical techniques and trophic measurement indices
as those used for Messel are applied to well-preserved lake deposits before and after
the MLR, analytical detection of the trophic shift left by the MLR may be difficult.
Nevertheless, studies of ichnofaunas and insect biotas, particularly those from the
Paleogene may provide robust results in lieu of detailed food-web analyses of lake
deposits using modern ecological techniques. One particular approach that may be
useful is to attempt an associational approach in the freshwater, especially lacus-
trine realm that has been done for the terrestrial realm (Labandeira et al. 2002; Wilf
et al. 2006). In lieu of more complete knowledge of aquatic, arthropod body-fossil
faunas during this time interval, analyses of interactions may provide an another
approach for detecting the trophic shift toward herbivore-dominated aquatic com-
munities following the MLR.
Ichnofaunas from lake-margin deposits are known from the Paleogene of the west-
ern United States (Melchior and Erickson 1979; Lamond and Tapanila 2003; Bohacs
et al. 2007; Scott and Smith 2015), Spain (Rossi 1992; de Gibert and Sáez 2009),
China (Hsiao et al. 2010), Antarctica (Yang and Shen 1999; Perea et al. 2001), the
Neogene of the western United States (Toots 1975; Smith et al. 1982; Squires and
Advocate 1984; Scrivner and Bottjer 1986; Lucas et al. 2002), Spain (Rodriguez-
Aranda and Calvo 1998; Uchman and Álvaro 2000; Ortí et al. 2003) and Kenya
(Laporte and Behrensmeyer 1980; Cohen 1982; Ekdale et al. 1989; Cohen et al. 1991,
1993; Lamond and Tapanila 2003). Shallow-tier, horizontally striated or non-orna-
mented, meniscate, backfilled structures (Scoyenia, Taenidium, Beaconites), simple
horizontal structures (Planolites, Palaeophycus), vertical burrows (Arenicolites,
Polykladichnus), and striated or non-ornamented branching burrows (Spongeliomorpha,
Labyrintichnus) are the dominant elements (Toots 1975; Smith et al. 1982; Squires
and Advocate 1984; Rodriguez-Aranda and Calvo 1998; Uchman and Álvaro 2000;
Ortí et al. 2003; Bohacs et al. 2007; de Gibert and Sáez 2009; Hsiao et al. 2010). The
crayfish burrow Camborygma has been locally recorded (Hsiao et al. 2010). Vertical
burrows with a terminal chamber, ascribed to the ichnotaxon Liticuniculatus erectus,
Table 11.2 Summary of ichnologic, paleontologic, and paleoecologic information on the Mesozoic Lacustrine Revolution (MLR)
Lake- Permanently
Evolutionary Aquatic insect body fossils margin trace subaqueous
phases (representative families)a Other benthos Aquatic plants Nekton/plankton fossils trace fossils Ecosystem characteristics
Pre-Mesozoic Dasyleptidae, Protereismatidae, Chelicerates, Charophytes, Chlorophytes Arthropod Superficial Colonization of lake-
Backdrop to Misthodotidae, conchostracans, water-tolerant (mostly and tetrapod grazing trails margin settings by the
Diversification Syntonopteridae, Jarmilidae, leperditocopid lycopods, Chlorococcales), trackways and shallow- Silurian-Devonian and of
Oboriphlebiidae, Eraspteriidae, crustaceans, sphenopsids, agnathan, dominant tier feeding permanently submerged
Meganeuridae, shrimp-like medullosan seedferns acanthodian, trace fossils lake bottoms by the
Protomyrmeleontidae, crustaceans, and arborescent placoderm, dominant, fish Carboniferous.
Permagrionidae, Kennedyidae, oligochaetes, marattialean ferns in chondrichthyan, trails present Limited to no colonization
Perlopseidae, bivalves, gastropods, wetlands; cordaites, and osteichthyan of the infaunal ecospace.
Tschekardiperliidae, amphibians, noeggeranthialeans, fish Poorly developed food
Palaeonemouriidae reptiles conifers, webs driven by detritivores
gigantopterids, and and top predators as
peltasperms in dry near-exclusive consumers;
settings herbivory was absent
Early Triassic Protomyrmeleontidae, Decline in family- Charophytes and Chlorophytes Arthropod No data Persistence of Paleozoic
Denouement Batkeniidae, level biodiversity. Isoetes-related (mostly trackways styles of animal–substrate
Naucoridae, Conchostracans, nonflowering Chlorococcales), dominant interactions in lake-margin
Belostomatidae, shrimp-like vascular plants osteichthyan fish settings. Poorly developed
Permosynidae, crustaceans, food webs driven by
Schizophoridae, bivalves, gastropods, detritivores and top predators
Liassophilidae, oligochaetes, as near-exclusive consumers;
Tipulidae amphibians, reptiles herbivory was absent
(continued)
Lake- Permanently
Middle-Late Triassolestidae, Biodiversity Charophytes and Chlorophytes Meniscate Superficial Increase in depth and extent
Triassic Saxonagrionidae, recovery. Isoetes- and (mostly trace fossils grazing trails and of bioturbation in
Rediversification Triadophlebiidae, Conchostracans, Azolla-related Chlorococcales), dominant, shallow-tier lake-margin sediments
Siphloneuridae, Euxenoperlidae, shrimp-like nonflowering osteichthyes fish crayfish feeding trace Persistence of Paleozoic
Ochteridae, Triassocoridae, crustaceans, vascular plants burrows fossils dominant, styles of animal-substrate
Notonectidae, ostracods, common, fish trails present, interactions in permanent
Belostomatidae, kazacharthrans, reptile mid-tier feeding subaqueous lake settings, but
Schizophoridae, Ademosynidae, phyllopods, bivalves, trackways burrows locally incipient colonization of the
Colymbothetidae, gastropods, and lungfish present.Vertical infaunal ecospace locally.
Hydrophilidae, Dytiscidae, oligochaetes, burrows dwelling burrows More complex food webs,
Triaplidae, Philopotamidae, bryozoans, present in higher-energy albeit still driven by
Prorhyacophilidae, amphibians, reptiles settings detritivores and top predators
Nannochoristidae with herbivory absent
Jurassic Tarsophlebiidae, Siphlonuridae, Conchostracans, Charophytes Chlorophytes Meniscate Shallow-tier Persistence of Middle to Late
diversification and Mesonetidae, Epeorominidae, shrimp-like dominant (radiation (mostly trace fossils feeding and Triassic styles of animal-
infaunalization Mesoleuctridae, crustaceans, of Characeae and the Chlorococcales), dominant, grazing trace substrate interactions in
Siberioperlidae, ostracods, Clavatoraceae), osteichthyes fish dinoaur fossils and lake-margin settings, but
Chresmodidae, phyllopods, bivalves, Azolla-related trackways deeper-tier with the addition of intense
Belostomatidae, gastropods, nonflowering present feeding burrows bioturbation by dinosaurs.
Gyrinidae, Mesoveliidae, oligochaetes, vascular plants, (including dominant, fish Widespread occupation of
Shurabellidae, bryozoans, ferns, trampled trails present, mid tiers in deep lacustrine
Coptoclavidae, Haliplidae, amphibians, reptiles porocharaceans and surfaces) mid-tier feeding deposits, but limited
Liadytidae, Parahygrobiidae, nitellaceans locally burrows locally sediment reworking.Increase
Necrotaulidae, present. Vertical in predation pressures.
Limoniidae, dwelling Initial transformation from
Chironomidae burrows in detritivore-driven to
Eoptychopteridae, higher-energy herbivore-driven trophic
Chaoboriodae settings networks of the mesolimnion
and epilimnion
Early Cretaceous Petaluridae, Conchostracans, Aquatic angiosperms Chlorophytes Meniscate Shallow-tier Establishment of the
Persistence Libellulidae, shrimp-like and ferns common, (mostly trace fossils feeding and lacustrine mixed layer.
Hemeroscopidae, crustaceans, charophytes present Chlorococcales), dominant, grazing trace Transformation from
Hexagenitidae, ostracods, dominant, crayfish fossils, mid-tier detritivore-driven to
Chresmodidae, phyllopods, bivalves, dinoflgellates, burrows pervasive herbivore-driven trophic
Clypostemmatidae, gastropods, osteichthyan fish common, burrow networks of the
Mesotrephidae, oligochaetes, dinosaur, mottlings, fish mesolimnion and
Gerridae, Mesoveliidae, bryozoans, pterosaur, trails present epilimnion
Hydrophilidae, Dytiscidae, amphibians, reptiles bird, reptile
Coptoclavidae Scirtiidae, and turtle
Corydalidae trackways
Vitimotaulidae, Baissoferidae present
Calamoceratidae
Late Cretaceous Gomphidae, Conchostracans, Aquatic angiosperms Chlorophytes Crayfish Deeper-tier Persistence of previous
biotal replacement Petaluridae, shrimp-like and ferns dominant, (mostly burrows, feeding trends in styles of
Aeshnidae, crustaceans, charophytes present. Chlorococcales), and burrows animal–substrate
Libellulidae ostracods, and dinoflgellates dinosaur present interactions
Coenagrionidae phyllopods, bivalves, dominant, and bird
Gerridae, gastropods, osteichthyan fish trackways
Hebridae, oligochaetes, present
Hydrometridae, bryozoans,
Mesotrephidae, amphibians, reptiles
Hydrobioscidae,
Sericostomatidae,
Leptoceridae
Culicidae,
Ptychopteridae,
Chaoboridae
Ceratopogonidae
(continued)
Lake- Permanently
Cenozoic Sieblosiidae, Conchostracans, Aquatic angiosperms Chlorophytes Meniscate Shallow-tier Persistence of previous
Aftermath to MLR Zacallatidae, shrimp-like and ferns dominant. (mostly trace fossils feeding and trends in styles of
Dysagrionidae, crustaceans, Chlorococcales), dominant, grazing trace animal–substrate
Baetidae, ostracods, and dinoflgellates crayfish fossils and interactions.
Hydrophilidae, phyllopods, bivalves, dominant, burrows deeper-tier Strong impact of diatoms
Dytiscidae, gastropods, freshwater common, feeding starting in early Neogene
Hydropsychidae, oligochaetes, diatoms and insect nests burrows on the biogeochemical
Leptoceridae, bryozoans, chrysophytes, and borings dominant, fish cycling of silica, promotion
Limnephilidae, amphibians, reptiles osteichthyan fish locally, bird trails and nests of more complex trophic
Hydroptilidae, and mammals and present, webs and probable overall
Phryganeidae, mammal mid-tier increase of lacustrine
Culicidae trackways feeding productivity
Tipulidae, present burrows locally
Cylindrotomidae present.
Chironomidae Shallow-tier
trails in glacial
settings.
Vertical
dwelling
burrows in
higher-energy
settings
a
This column provides the more important aquatic forms or inhabitants of wet habitats adjacent to lakes for at least one stage in their life cycle (naiad, larva, adult)
were compared with similar structures produced by decapods (Melchior and Erickson
1979). This is a poorly known ichnotaxon that may elicit comparisons with
Camborygma. Simple grazing trails (Archaeonassa, Cochlichnus) are very rare (Smith
et al. 1982; de Gibert and Sáez 2009).
In some cases lake-margin deposits are overprinted by a suite of insect nests
(Celliforma, Roselichnus; Uchman and Álvaro 2000), which are particularly com-
mon in Cenozoic terrestrial settings (see Chap. 13). Spectacular examples of sur-
faces and beds containing several overprinted suites from fully aquatic to transitional
and fully terrestrial facies are known from a number of Quaternary localities in
Kenya (Scott et al. 2009; Owen et al. 2009). Borings (Trypanites, Sertaterebrites)
and embedment cavities have been documented in lacustrine stromatolites (Ekdale
et al. 1989; Lamond and Tapanila 2003; but see Corsetti and Grotzinger 2005 for an
alternative interpretation). Evidence of insects capable of burrowing in highly
stressed, hypersaline environments of underfilled lakes are observed for the first
time in some of the Neogene basins of Spain (e.g. Rodriguez-Aranda and Calvo
1998; Uchman and Álvaro 2000; Ortí et al. 2003).
Vertebrate trace fossils in Cenozoic lake-margin deposits include bird (e.g.
Gruipeda) and mammal tracks (Scrivner and Bottjer 1986; Yang and Shen 1999;
Perea et al. 2001; Lucas et al. 2002; de Gibert and Sáez 2009). A wealth of informa-
tion has been produced on how vertebrates modify lake-margin surfaces. Large
mammals (e.g. hippos) are known to produce trampled surfaces, actively modifying
lake-margin landscapes (Laporte and Behrensmeyer 1980; Cohen et al. 1991, 1993;
Ashley and Liutkus 2002; Deocampo 2002; Scott et al 2007, 2008) (Fig. 11.11a–c).
Additionally, flamingos produce nest mounds and alter lake-margin deposits by
trampling and churning wet clayey sediments (Scott et al. 2009, 2012b). These
authors documented that these nests may be so compacted that they contribute to the
stability of deltaic distributary channels in lake margins, leading to channelization
of flow by water diversion around the mounds. Indeed, the impact of flamingos on
lake-margin sediments results in the development of a distinctive mounded topog-
raphy that qualifies as an ichnolandscape (sensu Buatois and Mángano 2011a),
which can be preserved in the fossil record (Fig. 11.12a–b).
Ichnofaunas from fully lacustrine deposits have been recorded nearly worldwide in
Cenozoic deposits. They have been documented from the Paleogene of the western
United States (Moussa 1968, 1970; Melchior and Erickson 1979; Loewen and de
Gibert 1999; Bohacs et al. 2007; Martin et al. 2010; Scott and Smith 2015), Antarctica
(Yang and Shen 1999; Perea et al. 2001), the Neogene of the eastern (O’Brien and
Pietraskek-Mattner 1998; Benner et al. 2009; Knecht et al. 2009) and western (Smith
et al. 1982) United States, eastern Canada (Gibbard and Dreimanis 1978), Hungary
(Babinszski et al. 2003; Magyar et al. 2006; Cziczer et al. 2009), Slovakia (Starek et al.
2010; Hyžný et al. 2015), England (Gibbard and Stuart 1974), Lithuania (Uchman
et al. 2008, 2009), Finland (Gibbard 1977), Sweden (Uchman and Kumpulainen 2011),
Germany (Walter 1985; Walter and Suhr 1998), Greece (Owen et al. 2011), Turkey
(Price and McCann 1990; Uchman et al. 2007), China (Yang 1996), Indonesia
(Whateley and Jordan 1989), Thailand (Gibling et al. 1985; Flint et al. 1989), Japan
(Allison et al. 2008), Kenya (Feibel 1987) and New Zealand (Lindqvist 1994).
Fig. 11.11 Fossilized trampled surface consisting of a high-density of hippopotamus tracks,

Pleistocene, Ilosowuani horst near Logumukum, Lake Bogoria Basin, Kenya. (a) General view of
the trampled surface; (b) Close-up of one of the tracks; (c) Cross-section view showing intense
disturbance of the primary sedimentary fabric
Ichnofaunas from fully lacustrine, Cenozoic deposits are known from both nongla-
cial and glacial settings. Nonglacial lake deposits that accumulated under low energy
conditions tend to be dominated by a combination of very shallow-tier grazing trails
(Cochlichnus, Helminthopsis), shallow-tier mottlings attributed to Planolites, and
deeper-tier branching burrows (Vagorichnus) (Moussa 1968, 1970; Smith et al. 1982;
Gibling et al. 1985; Feibel 1987; Flint et al. 1989; Whateley and Jordan 1989; Yang and
Shen 1999; Perea et al. 2001; Bohacs et al. 2007; Uchman et al. 2007; Owen et al.
2011). Oligichnos limnos, an ichnotaxon erected by Melchior and Erickson (1979) and
suggested to have been made by oligochaetes, needs re-evaluation and may become a
junior synonym of Helminthopsis. Also, the presence of the bivalve burrow Lockeia
has been mentioned (Feibel 1987). Interbedded sandstone tempestites and dolomites
contain deeper-tier vertical burrows (Skolithos, Arenicolites Polykladichnus) (Price and
McCann 1990; Magyar et al. 2006). Vertical burrows similar to Trichichnus have been
Fig. 11.12 Fossilized flamingo nests, Pleistocene, High Magadi Beds, Lake Magadi, Kenya. (a)
General view of the nest-bearing surface showing a spectacular biogenic topography (i.e. ichno-
landscape); (b) Close-up of nests. Scale bar is 10 cm long
mentioned in lacustrine shales (Lindqvist 1994). Burrow systems from long-lived

brackish- to freshwater fossil Paleolake Pannon have been attributed a new ichnotaxon,
Egbellichnus jordidegiberti (Hyžný et al. 2015). These burrows have been interpreted
to have been produced by ghost shrimps, indicating that these marine organisms were
able to survive for a long time after the closure of the seaway connecting this water
body with the central Paratethys sea (Hyžný et al. 2015). Vertebrates are represented by
the fish trail Undichna (Loewen and de Gibert 1999; Martin et al. 2010) and the fish
nest Piscichnus (Feibel 1987).
Glacial lakes are dominated by very shallow-tier grazing trails (Cochlichnus,
Gordia, Helminthoidichnites) and arthropod trackways (Glaciichnium,
Warvichnium, Dendroichnites, Lusatichnium), with the branching burrow
Treptichnus and the conchostracan resting trail Surculichnus locally present
(Gibbard and Stuart 1974; Gibbard 1977; Gibbard and Dreimanis 1978; Walter and
Suhr 1998; O’Brien and Pietraskek-Mattner 1998; Uchman et al. 2009; Benner
et al. 2009; Knecht et al. 2009; Uchman and Kumpulainen 2011). Vertebrates are
represented by the fish ichnotaxa Undichna, Broomichnium and Piscichnus
(Benner et al. 2009; Uchman and Kumpulainen 2011). The ichnofaunas of
Pleistocene glacial lakes are somewhat unusual in that they closely resemble
those that are typical of the Paleozoic, displaying a dominance of very shallow-
tier ichnotaxa and an underutilized infaunal ecospace. The anachronistic nature of
Pleistocene glacial lacustrine ichnofaunas may represent another example of the
so-called déjà vu effect (Buatois and Mángano 2011b). The dominance of very
shallow-tier structures in Pleistocene glacial deposits may reflect very limited
burial of organic matter in highly oligotrophic lakes, restricting the habitable zone
to a few millimeters below the sediment–water interface.
It should be noted, however, that suppressed bioturbation is not restricted to
these types of settings, but also occur in highly productive lakes having oxygen-
ated bottom waters, as evidenced in Lake Turkana (Cohen 1984). In this lake, pri-
mary productivity is extremely high toward the lake shore, but phytoplankton
populations are markedly less productive offshore. As a result, food content is
quite low in offshore lake bottoms and the invertebrate fauna essentially consists
of epibenthic detritivores which do not disturb the primary sedimentary fabric. In
addition, the lack of sediment mixing results in a redox discontinuity surface very
close to the sediment–water interface, further restricting infaunalization (Cohen
1984). This pattern may be regarded as reminiscent of Paleozoic settings display-
ing limited colonization of infaunal ecospace.
To summarize, a review of the Cenozoic lacustrine trace-fossil record fails to show
the appearance of any major evolutionary novelty or innovation from an ichnologic
perspective. The spectacular trampled surfaces produced by large mammals that char-
acterize Neogene and modern lake margins were preceded by equally extended and
bioturbated surfaces produced by dinosaurs in the Mesozoic. Cenozoic ichnofaunas
suggest a continuation of trends that started during the MLR. However, these exten-
sive ichnologic studies need to be linked with examination of body-fossil assemblages
from lacustrine and other aquatic environments, particularly during the Paleogene, to
more comprehensively document the effects of the MLR. It is notable that, as dis-
cussed below, aquatic organisms and ecosystems were better buffered from extinction
and ecological reorganization than terrestrial ecosystems during and after the K-Pg
crisis (also see Chap. 12). This buffering has multiple sources, including (1), behav-
ioral modifications of lacustrine organisms, including infaunalization, encystment,
and planktotrophic larvae that would predispose organisms toward avoiding major
environmental vicissitude (Chap. 12); (2), more generalized and less specialized food
webs occurring in lakes than for coexisting, surrounding terrestrial communities, at
least known for the middle Eocene (Dunne et al. 2014); and (3), the temporally
ephemeral nature of lake communities that require considerable dispersal mecha-
nisms for continuity of lake biotas (Cohen 2003), Additionally, the responses to the
K-Pg event were variable by the taxonomic group involved, by their occupied habitat,
by where they occurred in regional landscapes and in their worldwide distribution,
and were variably dampened by the effects of time.
A major effect of the end-Cretaceous global crisis (see Chap. 12) was not only
extensive taxonomic extinction and subsequent origination for many non-arthropodan
terrestrial lineages, but equally important, trophic pruning of ecosystem structure.
On land, the consequences of this event meant the demise of numerous lineages,
particularly vertebrates (Archibald and Bryant 1990) and plants (Nichols and Johnson
2008), but this extinction only minimally affected arthropod groups, when evaluated
as body-fossil taxa at the family level (Labandeira 2005b). However, when plant–
insect interactions are examined at the K-Pg boundary interval, the consequences
were important, and hit especially specialized insects such as gallers and leaf miners,
while leaving taxa with generalized feeding habits relatively intact (Labandeira et al.
2002; Wilf et al. 2006). In aquatic ecosystems, there was less of an effect (but see
Bailey et al. 2005 for possible taphonomic biases). Aquatic insects were minimally
affected, at least at the family level (Labandeira 2005b), although there may have
been significant changes in food-web structure that persisted well into the Paleogene
(Dunne et al. 2014). One important pattern documented for aquatic invertebrates was
the differential effects that the K-Pg event had on organisms that inhabited the ben-
thos of water bodies (Twitchett 2006). Detritivorous organisms in or on the sediment
were considerably more buffered both from extinction and destruction of their food
resources than those trophic networks subsisting on food resources derived from
photosynthetic organisms such as algae, diatoms, or aquatic vascular plants (Sheehan
and Hansen 1986). This immunity was enhanced for those aquatic invertebrates,
such as many detritivorous arthropodan groups, that had planktic immature stages
and thus were more able to withstand extinction. For aquatic, especially lentic and
lotic–potamic habitats, there was a reduced effect from the end-Cretaceous crisis,
particularly for local trophic networks whose dietary resources were not dependent
on the herbivory of photosynthetic organisms.
The history of Paleogene insects in aquatic ecosystems is poorly known outside
of Central Europe and North America. Most aquatic insect orders have been mini-
mally documented, and many assemblages are woefully deficient in species-level
descriptions and revisions. The number of autochthonous lentic taxa is poorly
known. Better documented are exceptionally well preserved compression deposits
that include the Green River, Messel, Kishenehn, and Menat paleolakes (Nel and
Roy 1996; Smith ME et al. 2008b; Greenwalt and Labandeira 2013; Dunne et al.
2014). These well-documented deposits represent only a broad outline of Paleogene
aquatic insect history, even though their temporal proximity to the modern fauna
should allow easier identification and ecological interpretation of the insect taxa
present (Zherikhin and Sinitshenkova 2002). These and almost all Paleogene aquatic
taxa are assigned to extant families. The only significant exceptions to this general-
ization lie within the Odonata, in which the families Sieblosiidae (which disap-
peared during the Pliocene) and Zacallatidae (Paleogene only) and the subfamilies
Dysagrioninae (Paleogene only) and Eodichrominae (Paleogene only) are absent
from the modern fauna (Zherikhin and Sinitshenkova 2002). The overwhelming
majority of modern lotic and lentic aquatic families extend at least to the Paleogene,
and occur in compression and amber deposits. There are many cases of extant aquatic
genera extending to the middle Eocene, and apparently living species present in
sediments as old as the Eocene (Askevold 1990; Murray 1976; Kluge 1986).
The naiads of ephemeropterans are uncommon in Paleogene compression depo-

sits. Some amber deposits, such as Baltic Amber have provided the overwhelming
majority of known mayfly adults. A common, modern epibenthic form, the Baetidae,
is virtually absent from Paleogene aquatic faunas. Paleogene odonatan assemblages
exhibit minimal similarity with those of today when compared to other aquatic
insect lineages (Zherikhin and Sinitshenkova 2002). The relative rarity of Lestidae
(spreadwing damselflies), Coenagronidae (narrow-winged damselflies), Corduliidae
(emerald dragonflies) and Libellulidae (darner dragonflies) are in stark contrast to
the abundance of these lineages in extant aquatic faunas. Naiads are rare in Paleogene
aquatic deposits, whereas they are much more common in Neogene faunas, a pat-
tern mirrored by ephemeropterans and attributable to styles of preservation in lacus-
trine deposits rather than any intrinsic biologic cause.
Paleogene Plecoptera are represented principally by adults in resins and are
referable to extant genera. These fossil Plecoptera probably were confined to lotic
habitats, consistent with the rarity of their naiad stages in lentic deposits. Apparently,
stoneflies do not exhibit any shift in taxonomic proportions between the Paleogene
and Neogene (Zherikhin and Sinitshenkova 2002), and display a dominance similar
to corixids and notonectids. Aquatic bugs (Hemiptera), unlike paleopterous taxa,
are common in Paleogene deposits, where they are often numerically dominant.
Similarly, aquatic beetles are diverse in the Paleogene, but are marked by the
absence of the algivorous Haliplidae and the overwhelmingly presence of the
Hydrophilidae and Dytiscidae, particularly during the Oligocene. Aquatic megalop-
terans and neuropterans are represented only by extinct genera, and aquatic mecop-
terans have not been documented.
In contrast to several modern aquatic groups that were uncommon during the
Paleogene, the Trichoptera were quite common in local assemblages. All Paleogene
taxa have been assigned to modern lineages, many which consist of extant genera
(Sukatcheva 1982; Wichard and Weitschat 1996). However, there is a significant
difference in family-level dominance between the Paleogene and Neogene assem-
blages, and in compression versus resin deposits, with the Oligocene being a transi-
tional interval. During the Paleogene, the currently dominant families, the
Hydropsychidae (net-spinning caddisflies), Leptoceridae, and Limnephilidae
(northern caddisflies) are either rare or absent. Similarly, the diversity of caddisfly
cases are rare when compared to the Cretaceous or present-day levels (Zherikhin
and Sinitshenkova 2002). Some of these taxa may have colonized floating algal
mats (Ponomarenko 1996) or dwelled amid the floating leaves, as appears to be the
case for fossils from earliest middle Eocene Paleolake Messel (Lutz 1991).
The Paleogene aquatic dipteran fauna is particularly depauperate in culicids,
chironomine chironomids (nonbiting midges), and higher brachyceran taxa.
However, the abundance of the enigmatic botfly-like larvae of Lithohypoderma
continues to elude taxonomic placement and ecologic understanding (Stokes 1978;
Sinitshenkova 2002). Culicids (mosquitoes) and brachyceran taxa became more
abundant during the Oligocene. Toward the Oligocene–Miocene boundary, the
aquatic fauna attained an even more modern cast than in the previously described
Paleogene assemblages.
Paleogene aquatic angiosperms continued to be dominant in ponds and lakes,

forming diverse assemblages and displaying increased provincialism (Martín-
Closas 2003). Aquatic ferns also exhibited further diversification (Schneider et al.
2004), whereas taxonomic turnover within the Characeae took place during the
Paleogene (Martín-Closas 2003). Further extinctions and replacements occurred
within the charophytes later in the Cenozoic. Aquatic bryophytes became common
in shallow lakes and swamps (Mai 1995). The most significant changes, however,
are those experienced by lacustrine phytoplankton, essentially by expansion of
freshwater diatoms and chrysophytes during the Eocene (Martín-Closas 2003;
Wolfe and Edlund 2005; Sims et al. 2006; Leliaert et al. 2011). During the Neogene,
diatoms clearly dominated over the Chlorococcales (Martín-Closas 2003; Sims
et al. 2006). The overall ecologic impact of the colonization of freshwater bodies by
diatoms cannot be overemphasized. In particular, diatoms typically exert a signifi-
cant effect on the biogeochemical cycling of silica, promote a diversion of other
nutrients into other trophic pathways and increase overall lacustrine productivity
(Cohen 2003).
One of the most highly investigated Cenozoic lacustrine deposits is the series of
four major paleolake basins that form the Green River Formation in the Western
Interior of the United States. The Green River Formation is dated from ca. 54.0–
43.5 Ma (Smith ME et al. 2008b), and consists of a mixture of open lacustrine,
lake-margin lacustrine, shallow playa, carbonate mudflat, fluvial, deltaic and allu-
vial deposits (Ferber and Wells 1995; Chetel and Carroll 2010; Aswasereelert et al.
2013). The Green River Formation is assigned to Cenozoic Lake Assemblage Type
8 (Zherikhin and Sinitshenkova 2002), dominated by brachyceran fly larvae with
few, if any, other aquatic insects (Zherikhin and Sinitshenkova 2002). Green River
fossils overwhelmingly preserve land-based organisms, especially plants, insects,
and vertebrates, to the near exclusion of autochthonous aquatic organisms (Wilson
1978), which likely was taphonomically modulated by depositional biases favoring
terrestrial taxa (Smith, 2000, 2008). The insect fauna is very diverse, consisting of
about 26 of the ca. 35 modern recognized orders for the Piceance Creek Basin of the
Green River Formation (Pribyl et al. 1996). The apparently under-represented
aquatic insect fauna is autochthonous, but only consists of several major lineages of
mostly nematocerous Diptera, principally tipulids, cylindrotomids, culicids, chi-
ronomids, and the occasional aquatic beetle. In spite of the extensive geographic
coverage and numerous fossiliferous strata, there is only minor representation of
lotic and lentic fossils from the Green River Formation. Nevertheless, three families
of dragonflies are represented (Wilson 1978; Petrulevičius et al. 2007), two families
of Hemiptera, two families of beetles, four families of nematocerous Diptera and
the enigmatic Lithohypoderma “botfly” (Pribyl et al. 1996; Sinitshenkova 2002).
The Trichoptera includes the Hydropsychidae, Hydroptilidae (micro caddisflies),
Limnephilidae, and Sericostomatidae. The particular reason for such an under-
representation of the lentic and lotic fauna, especially when adjacent terrestrial
insects are exceedingly abundant by comparison, is likely attributable to the role of
size in the formation of live versus death assemblages of insects. In the Green River
biota, death assemblages that become incorporated in the fossil record are smaller
sized and more robust than live equivalents (Smith 2000). Although the role of size
and robustness applies to beetles (Smith 2000), it is unclear if such a relationship

also applies to softer-bodied, more gracile insects that are often disarticulated prior
to sediment burial.
The taxonomic composition and ecologic structure of Neogene paleolakes is very
similar to that of modern lakes (Zherikhin and Sinitshenkova 2002). Many modern
aquatic species occur throughout the Neogene. A few extinct insect clades are known,
such as the subfamily Electrobatinae of the Gerridae and the coral bug species
Halovelia electrodominica, both from early Miocene Dominican Amber; the coeval
dytiscid genus Palaeogyrinus from Germany; several extinct aquatic taxa of the
Megaloptera and Neuroptera, including the species Sialis strausi from the Pliocene of
Germany; and several new caddisfly case morphotypes that are difficult to attribute to
potential modern or extinct genera (Illies 1967; Galewski and Glazek 1997; Sukatcheva
1982; Anderson 1998). Throughout the Miocene, these occurrences do not represent
any significant change in aquatic insect community characterization, a conclusion
buttressed by the similarity of overall lake ecologic structure between the middle
Eocene and the present (Dunne et al. 2014), indicated in Fig. 11.13.
There is evidence for an increase in lentic habitats and their biotas at the beginning
of the Neogene. This extension is indicated by greater submergent and emergent angio-
sperm vegetation that increased habitat structural complexity, resulting in an expansion
of herbivory, greater constancy of O2 levels and nutrient cycle stability (Zherikhin
Fig. 11.13 Visualization of the middle Eocene (48 Ma) food web of Paleolake Messel, in central-
west Germany, representing a post MLR lake. Spheres designate trophic taxa and lines represent
feeding links. A trophic taxon is a species or group of species (in the case of microorganisms) that
have links to organisms that consume it and to other organisms that it consumes. Plants have links
only to their consumers. Looped links refer to cannibalism. The vertical axis corresponds to short-
weighted trophic level, with autotrophic taxa and detritus at the bottom. The colors of nodes indi-
cate the taxonomic assignment of species: green = plants, algae and diatoms; blue = bacteria, fungi
and detritus; yellow = invertebrates; orange = vertebrates. This image was produced with Network
3D software (Williams 2010), and reprinted with permission from the Royal Society
1978; Kalugina 1980). Nevertheless, special sedimentary environments occupying

spatially small areas could have hosted aquatic communities of distinctive taxonomic
composition and ecologic structure. Examples of these biotic microcosms include
small bodies of water associated with sinter structures, ponds in asphaltum depres-
sions, karst sinkholes, impact craters, and pits created by subsurface diatreme explo-
sions. Such deposits are typically spatiotemporally ephemeral and constitute a very
minor part of the lacustrine record that is represented preferentially toward the recent.
11.7 Discussion
There are four issues about the MLR that necessitate further discussion. The first
issue is whether the MLR is indirectly or directly connected to other obvious, major
biotic events during the mid Mesozoic, such as the Mesozoic marine, parasitoid,
angiosperm, and Cretaceous terrestrial revolutions. The second issue is to what
extent did the MLR involve not only a change in the diversity and abundances of
participating fossil groups, but also their effect on the sedimentological record
through the increased variety of ichnotaxa and bioturbation intensities recorded
from well sampled sections. A third issue is how modern lacustrine trophic webs
were established. These trophic webs are recorded in a well-examined Paleogene
lake deposit, but also should be present in earlier, mid-Mesozoic food webs. Finally,
what is the role of behavioral convergence in shaping the aquatic marine and conti-
nental trace fossil records, and to what extent should ichnotaxonomic nomenclature
be applied across the marine/continental divide to very similar trace fossils that
involve convergent morphologies (and presumably behaviors).
11.7.1 Possible Connections to Other Mesozoic, Biotal

Revolutions
Other than the MLR, four other, major biotic revolutions occurred during the mid to
late Mesozoic: the Mesozoic marine revolution in the marine realm and the parasit-
oid, angiosperm and Cretaceous terrestrial revolutions in the continental realm.
These mid-Mesozoic, global ecologic shifts may have had an effect on the MLR
that may be relevant to the development of lacustrine food webs throughout the later
Mesozoic. The possible connections among these five prolonged events may shed
light on the shift of detritivore- to herbivore dominated ecosystems in lacustrine and
other freshwater habitats during the mid Mesozoic.
One of the aspects involved in the Mesozoic marine revolution (see Chap. 9) was the
notable increase in mechanical sturdiness and increased robustness of other structural
features that promoted the resistance of shells, principally gastropods, to crushing by
predators (Vermeij 1977; Stanley 1977). Major structural transformations favored dura-
ble shell architecture of molluscan prey, and involved new modes by mostly arthropods
and teleost predators of pursuing, capturing, and overcoming the mechanical defenses
of their prey. Another aspect of the Mesozoic marine revolution was intensification of
marine herbivory, such as grazing on algae at relatively shallow and subsurface depths.
Added to the expansion of durophagy and grazing pressure was increased predation in
benthic communities that led to infaunalization of many former epibenthic groups, as
well as a significant reduction of the sessile life habit (Stanley 2008). The major conse-
quence of these innovations has been the relegation of their certain life forms to extinc-
tion or to refugial habitats (Oji 1996), and the occupation of their adaptive zones by
competitively superior, new, predatory groups (Vermeij 1977; Stanley 1977). This
worldwide marine reorganization has been linked to continental breakup, widening of
latitudinal belts, and the emergence of angiosperms on land (Vermeij 1977).
The parasitoid revolution is based on an observation that the first appearances of
several major clades of parasitoid insect clades occur primarily during the mid
Jurassic (Labandeira 2002). Parasitoids are a distinctive type of carnivore in which
a small, invading organism attaches to or lives on or in an animal and slowly feeds
on internal tissues and organs of their relatively long-lived but still alive host. The
much smaller parasitoid feeds on host such that the most vital organs are consumed
immediately before the host dies, followed by its emergence from the host as an
adult parasitoid (Vinson and Barbosa 1987). Parasites, by contrast, such as mosqui-
toes do not kill the host, but use host tissues such as blood to feed on. For insects,
the actively feeding parasitoid stage is always a holometabolous larva, overwhelm-
ingly a parasitoid wasp, and less frequently a brachyceran fly. Parasitoids may feed
on their hosts internally (endoparasitoids) or externally (ectoparasitoids) and attack
arthropod hosts, such as myriapods, arachnids, especially insects, and some verte-
brates (Godfray 1994). The earliest appearance of the parasitoid guild was the mid
Early Jurassic, during which several clades of parasitoid wasps appear. This was
followed and supplemented by more modest brachyceran dipteran diversification
and the massive radiation of several parasitoid superfamilies during the Late
Jurassic, and continued with the addition of occasional neuropteran, beetle, and
lepidopteran parasitoid lineages into the Paleogene (Labandeira 2002).
A major consequence of the parasitoid revolution was the increased complexity
of food webs by insertion of a significant more efficient mode of carnivory. This
greater efficiency resulted from the targeting of particular species of prey by a host-
specific parasitoid using specific chemical, behavioral, visual, and other cues ema-
nating from prey. With the proliferation of the host-specialist, parasitoid feeding
guild that originated during the mid Mesozoic, carnivory became considerably
more targeted than was the case for food webs earlier in time based solely on diffuse
generalist predation patterns (Labandeira 2002). This ecological shift resulted in
consumer-driven, top–down regulation of herbivores, rather than bottom–up,
resource-driven regulation, based on modern food-web studies (Dunne et al. 2002).
In modern ecosystems, parasitoid loads on insect herbivores are elevated (Memmott
and Godfray 1993), and are much more efficient at assimilating prey biomass into
consumer biomass than either predators such as dragonflies or parasites such as
fleas or lice (Eggleton and Belshaw 1992). In lacustrine systems, unionacean and
mytilacean bivalves are dispersed in their larval stages as ectoparasites on fish;
some copepod and isopod crustaceans are also parasitic on fish, as well as certain
flatworms and trematodes (Cohen 2003). One of the major effects of the parasitoid
revolution on lake biotas was to further separate immature aquatic stages of insects
from their adult terrestrial counterparts. Additional work needs to be done to prop-
erly address the importance of parasitoids in lacustrine food webs.
The angiosperm revolution consists of the early, major diversification interval of
flowering plants, ranging during the mid Cretaceous from 125 to 90 million years
ago (Crane et al 1995). The angiosperm revolution had a major effect on terrestrial
ecosystem structure (Wing and Boucher 1998). The emergence of numerous clades
of angiosperms over a ca. 35 million-year interval resulted in a major replacement
of fern- and gymnosperm dominated floras by angiosperms in most habitats glob-
ally (Crane 1987), including aquatic settings (Wing and Boucher 1998).
The Cretaceous terrestrial revolution is directly connected to and may be a con-
sequence of the angiosperm revolution because it encompasses not only the evolu-
tionary radiation of land-dwelling organisms, but involves the replacement of ferns
and gymnosperms by angiosperms (Lloyd et al. 2008). Therefore, the comments
outlined for the angiosperm revolution are prefatory to the Cretaceous terrestrial
revolution. Together with the rapid expansion of flowering plants, the Cretaceous
terrestrial revolution includes the diversification of herbivorous and social insects
(see Chap. 13), squamates, birds, and mammals (Lloyd et al. 2008). However, these
authors (Lloyd et al. 2008) questioned the notion that dinosaur evolution was driven
directly by angiosperm diversification. From an ichnologic standpoint, the appear-
ance of trampled surfaces is a direct consequence of the expansion of dinosaur
faunas that significantly altered lake-margin landscapes.
Both the Mesozoic marine revolution and the parasitoid revolution provided a greater
efficiency in the conversion of prey biomass into consumer biomass. The angiosperm and
Cretaceous terrestrial revolutions expanded the amount of photosynthetic biomass cre-
ated by a new group of primary producers that had a major global impact. The MLR was
characterized by increased infaunalization and a major trophic shift from detritivore-
based to herbivore-based foodwebs in lacustrine and other terrestrial aquatic ecosystems.
However, the detritivore to herbivore shift occurred much earlier in terrestrial ecosystems
than in aquatic ones (Vermeij and Lindberg 2000; Miller and Labandeira 2002) – a shift
that was already noticeable during the Permian (Labandeira 2006). This is not surprising,
as in the continental realm, aquatic ecosystems always have been lagging behind terres-
trial ecosystems in other features such as species diversity, trophic relationships, and
food-web development (Vermeij and Lindberg 2000; Sinitshenkova 2002). Nevertheless,
it is unclear whether there are satisfactory explanations invoking cause-and-effect links
among any of these four major revolutions to lacustrine ecosystems.
11.7.2 Secular Changes in Global Ichnodiversity

and Bioturbation
A systematic review of the ichnologic record indicates that the MLR is expressed
not only by body-fossil data. Ichnologic evidence demonstrates that colonization of
freshwater habitats has been a protracted process (Buatois and Mángano 1993a;
Buatois et al. 1998a), with the MLR representing a pivotal point (Cohen 2003)
(Table 11.2). Freshwater colonization may have been delayed due to the need to
develop complex osmoregulatory systems and innovative styles of reproduction and
dispersal (Miller and Labandeira 2002).
Unlocking the evolutionary significance of the lacustrine trace fossil record
requires integration of several conceptual and methodological tools. Patterns of
change in global ichnodiversity may be misleading, and should not be analyzed in
isolation (Buatois and Mángano 2013; see Chap. 16). Mesozoic global invertebrate
ichnodiversity in lacustrine environments does not show any significant increase
with respect to late Paleozoic levels (Buatois and Mángano 1993a; Buatois et al.
1998a), in contrast to what may have been expected by diversity curves based on
body fossils (Labandeira 2005b). Indeed, whereas global ichnodiversity levels for
fully lacustrine environments have remained more or less constant since the
Carboniferous, global ichnodiversity of lake-margin settings is actually higher in the
late Paleozoic than in the Mesozoic. However, it often has been suggested that the
Permian ichnodiversity peak reflects at least in part splitting tendencies in arthropod
trackway taxonomy (see discussion in Buatois et al. 1998a and Chap. 6). Also, the
fact that diversity trajectories for body and trace fossils are markedly dissimilar is not
unexpected. The three groups that diversified the most – aquatic insects, aquatic
macrophytes, and teleosts – are unevenly represented in the ichnologic record.
Aquatic insects are thought to have been widespread tracemakers; some fish (e.g.
cichlids) may contribute locally to sediment disturbance (Feibel 1987; Martin et al.
2010; Abbate et al. 2012); and aquatic macrophytes are represented by root struc-
tures but technically do not contribute to ichnodiversity. (No formal ichnotaxonomic
names are given to root trace fossils.) In addition, it long has been recognized that
equating ichnodiversity with biological diversity is fundamentally misleading, and
one-to-one correspondences between ichnotaxa and biotaxa is simply not possible
(Bromley 1996). This is particularly true for very simple structures (Buatois and
Mángano 2011a), which tend to be dominant in lacustrine settings.
However, an evaluation of changes of ichnodiversity that parallel changes in inten-
sity and depth of bioturbation may be illuminating. In both lake-margin and fully
lacustrine deposits, an increase in extent and depth of bioturbation long has been
recognized (Miller 1984; Buatois and Mángano 1993a; Buatois et al. 1996, 1998a;
Miller and Labandeira 2002; Miller et al. 2002) (Fig. 11.14). In lake-margin settings,
trace fossils of the Scoyenia ichnoguild became more abundant, leading to increased
sediment mixing since the Middle Triassic (Buatois et al. 1998a). In addition, a sta-
tionary deep crayfish infauna , referred to as the Camborygma ichnoguild, was estab-
lished by the Triassic, further contributing to destruction of primary sedimentary
fabric. This is in sharp contrast with the situation of Paleozoic lake-margin deposits,
which overwhelmingly are dominated by very shallow-tier trace fossils, mostly
arthropod trackways (Buatois and Mángano 1993a; Buatois et al. 1998a; see Chap. 6).
Arthropod trackways, although locally present in Mesozoic lake-margin deposits, are
never dominant components, and they essentially disappeared from the Cenozoic
lacustrine trace-fossil record with the exception of glacial lakes, where they tend to
occur in distal facies rather than in lake-margin deposits (Uchman et al. 2009).
Seilacher (2008) underscored the importance of bioglue for trackway preservation,
Fig. 11.14 Secular changes in bioturbation in lake-margin and fully subaqueous lacustrine
deposits. Note progressive colonization of the infaunal ecospace through time
suggesting that the scarcity of trackways in post-Paleozoic lake-margin deposits may

have resulted from the appearance of a bioturbating meiofauna, therefore precluding
the formation of coherent biofilms. In any case, it is clear that there is a negative
correlation between diversity of shallow-tier trace fossils and intensity and depth of
bioturbation. This trend also has been noted for tidal flats, which display high diver-
sity during the late Paleozoic due to a richness of shallow-tier structures and decreased
ichnodiversity during the Mesozoic–Cenozoic, with ichnofaunas characterized by
deeper-tier structures (Mángano et al. 2002; Mángano and Buatois 2015). This
decrease in tidal-flat global ichnodiversity is arguably a taphonomic product resulting
from increased colonization of infaunal ecospace (Mángano et al. 2002; Buatois and
Mángano 2011a; Mángano and Buatois 2015). A similar argument can be posited to
explain the post-Paleozoic decrease in global ichnodiversity in lake-margin deposits.
In fully lacustrine settings, a similar increase in intensity and depth of bioturba-

tion is apparent throughout the Mesozoic, although changes seem to have been
slightly more gradual, lagging behind those in lake-margin environments. During
the Middle to Late Triassic, assemblages dominated by very shallow-tier grazing
trails, not unlike those of the late Paleozoic, were still common (Metz 1995, 1996,
2000; Melchor et al. 2003; Melchor 2004). However, the simultaneous appearance
of penetrative trace fossils is evidenced by systems of irregularly branched burrows,
most likely produced by oligochaetes or insect larvae (Voigt and Hoppe 2010). The
high density of these burrows contributed locally to disruption of the primary fabric,
generating patches of intense bioturbation in lacustrine mudstones. Branching bur-
rows occur in Lower Jurassic turbidites as well, reflecting the establishment of the
Vagorichnus ichnoguild, which represents the activity of a mobile, mid-tier, deposit-
feeding infauna (Buatois et al. 1995, 1996). These mid-tier trace fossils persisted in
Cenozoic lacustrine deposits (Uchman et al. 2007). Interestingly, although Lower
Jurassic turbidites contain many ichnotaxa (e.g. Cochlichnus, Helminthopsis) com-
mon in older deposits, these are more robust and produced infaunally, reflecting
penetration into the sediment and unlike those of the late Paleozoic. In the example
of the Vagorichnus-bearing deposits, the increase in maximum bioturbation depth is
not associated with an increase in bioturbation intensity. The decoupling of biotur-
bation depth and intensity of bioturbation results from the observation that these
burrow systems were emplaced at lithologic interfaces without causing major dis-
turbance of the primary sedimentary fabric (Buatois et al. 1995, 1996).
This pattern contrasts with Cretaceous ichnofaunas, which include pervasive
mottlings reflecting establishment of a shallow-tier deposit-feeding infauna, referred
to as the Planolites ichnoguild. The high density of these structures caused major
disruption of lacustrine sedimentary fabrics (Buatois and Mángano 1998; Buatois
et al. 1998a). These observations may suggest that the establishment of the mixed
layer in these settings is a result of the MLR. Although further work needs to be
done to detect the exact timing of formation of the mixed layer, sparse data suggest
that this sediment zone was already incipiently developed in lake bottoms by the
Middle to Late Triassic and well established by the beginning of the Cretaceous. As
is the case of lake-margin deposits, the appearance of these active bioturbators was
detrimental for the preservation of very shallow-tier structures. In any case, biogenic
homogenization of the lacustrine bottom sediments is not a universal phenomenon,
as indicated by the dominance of very shallow-tier trace fossils, in pristinely pre-
served, sedimentary fabrics of other Lower Cretaceous lacustrine deposits (de
Gibert et al. 2000, 2016; Buatois et al. 2000a), as well as in modern glaciolacustrine
varves (Gibbard and Stuart 1974; Uchman et al. 2009). Studies in other modern
lakes, such as the Great Lakes in North America, indicate up to 10 cm of deep
reworking by insects, oligochaetes, bivalves, and amphipods (McCall and Tevesz
1982; Miller and Labandeira 2002), suggesting a well-established mixed layer.
The pattern of increased infaunalization during the MLR is empirically well sup-
ported, but the underlying causes remain more speculative. As a first approach, the
increase in sediment penetration and disruption during the MLR is connected with the
expansion and diversification of certain groups of benthic aquatic organisms, most

likely dipterans (such as chironomids), which together with oligochaetes, are among
the key bioturbators in modern lacustrine settings (McCall and Tevesz 1982; Duck and
McManus 1984; Wootton 1988; Evenhuis 1994; Buatois et al. 1998a). At a deeper
explanatory level, one may invoke protection from environmental disturbance, escape
from predators and increased rates of buried organics as potential causes of infaunal-
ization. It long has been known that burrowing is a key strategy intended to minimize
environmental stress, such as salinity variations and sediment disturbance by currents
or waves (see Buatois and Mángano 2011a for discussion). However, environmental
disturbance may only work as an explanation for infaunalization at a local scale. In a
situation such as the MLR, infaunalization occurred globally and no overarching dis-
turbance factor can be associated with this macroevolutionary trait, therefore preclud-
ing protection from environmental disturbance as a likely causal factor of infaunalization.
In contrast, the link between increased predation pressure and infaunalization seems to
be a more robust explanation because the former is considered a driving force in mac-
roevolution, with the infaunal ecospace serving as refugium for predation. As discussed
above, the coincidence between increased predation and infaunalization during the
MLR is consistent with a causal link. The possibility that infaunalization can be linked
to exploitation of increasing amounts of organic matter buried within the sediment can-
not be disregarded. Higher quantities of food supply may have resulted from increased
eutrophication in lacustrine systems combined with increased delivery of terrestrially
derived and aquatic organic matter resulting from macrophyte diversification. Indeed,
nutrient availability may be regarded as an overarching factor controlling lacustrine
ecospace utilization in deep time (Cohen 2003). The ultraoligotrophic conditions pre-
dominant during the early Paleozoic were most likely the key limiting factor prevent-
ing colonization of lacustrine bottoms, a situation that started to change during the late
Paleozoic, albeit with colonization limited to a narrow inhabitable zone close to the
sediment–water interface (Buatois and Mángano 1993a; Buatois et al. 1998a). Finally,
it may be argued that invoking complex geobiologic feedbacks between bioturbation
and a wide range of abiotic to biotic factors (e.g. Mángano and Buatois 2014) probably
is more realistic than strict causal linkage. In the case of the MLR, the timing of events
suggests that the interplay of increased predation pressures and food availability may
have been the casual drivers forces in lacustrine infaunalization.
To summarize, whereas the Paleozoic is characterized by an increase in global
ichnodiversity as a result of the progressive colonization of continental environ-
ments (Buatois and Mángano 1993a; Buatois et al. 1998a; see Chap. 6), the ichno-
logic expression of the MLR is one of increased colonization of infaunal ecospace
(Table 11.2). This is reflected by an increase in both degree and depth of bioturba-
tion, although these increases occurred first in lake margins and subsequently in
fully lacustrine settings. In addition, the increased intensity of bioturbation lagged
behind greater penetration in burrowing depth in both settings. For further evalua-
tion of the evolutionary significance of the MLR, we turn now our attention to
trophic webs.
11.7.3 The Establishment of Modern Lacustrine Trophic Webs
Biotic interactions are of fundamental importance to understand ecosystem struc-

ture and function in lakes. In particular, deciphering food webs, the complex net-
works among predators, herbivores, autotrophs, and detritus/deposit feeders is a
central issue in reconstructing the biotic dynamics of lakes. Experimental studies
show that lacustrine food webs reflect an interplay of both bottom–up and top–down
selective pressures. Bottom–up processes involve food/nutrient resource availability
and competition for those resources whereas top–down processes involve trophic
cascades in which predation and selective consumption are the main regulators of
community structure (Carpenter and Kitchell 1993). Various groups of benthic
invertebrates, including crustaceans, mollusks, annelids, and larval insects play
important roles in mediating energy flow, nutrient cycling, and the ingestion of
organic and inorganic detritus and fecal production (reflected to some extent by
bioturbation) that is a central component of lacustrine food webs (Charbonneau and
Hare 1998; Covich et al. 1999; Voigt and Hoppe 2010). However, from an energetic
point of view, the importance of planktonic and nektonic components of the lacus-
trine food web is probably much greater (Schweitzer et al. 2007).
Accordingly, exploring the timing of the establishment of the modern lacustrine
food web and its potential connection with the MLR may yield insights into the evo-
lutionary history of lake ecosystems. There is little known about Early Triassic lakes
or their fossils. Late Middle–early Late Triassic Paleolake Madygen, discussed ear-
lier (Voigt et al. 2006), included organisms representing at least five trophic levels
(Fig. 11.15). Phytoplankton (of which there is no fossil evidence as yet) and macro-
phytes (e.g., Ricciopsis, Neocalamites, and some lycopsids) were presumably the
major primary producers. An important external source of food must have been dead
Fig 11.15 Ecosystem of the Madygen Formation of Kyrgyzstan. Trophic levels and their respec-
tive constituents as proposed for the Triassic Madygen lake environment. Note the absence of
macro herbivores After Voigt et al. (2006, 2016)
organic matter (e.g., plants, insects, tetrapods) transported from the land into lakes.
Zooplankton (of which there is no fossil evidence as yet), microconchids, gastro-
pods, bivalves, conchostracans, ostracods, kazakharthrans, certain insects (schizo-
phorid beetles), and wormlike aquatic invertebrates such as oligochaetes or particular
insect larvae, based on indirect evidence from trace fossils, are interpreted as primary
consumers. These organisms in turn served as food for a variety of fishes including
actinopterygians (Palaeoniscidae, “Perleididae”), dipnoans (Asiatoceratodus), and
durophagous sharks (Lonchidion). Four large carnivorous fishes – Saurichthys,
Oshia, a currently unknown xenacanthid (suggested by Fayolia-type egg capsules),
and coelacanths – can be considered tertiary consumers. The semi-aquatic reptilio-
morph Madygenerpton may have been the apex predator of Paleolake Madygen. This
system shows clear evolutionary innovations when compared to Paleozoic lakes, as
it represents among the earliest known record of a well-developed, deep lacustrine
infauna (Voigt and Hoppe 2010), and documents a quantitatively important pattern of
macrophytic colonization of the shoreline (Moisan et al. 2012a).
Although energy-flow diagrams have not been constructed for a Late Triassic
lake, coarse-grained trophic webs have been estimated for the biota of a hypotrophic
(low primary production, high O2) Middle Jurassic lake (Fig. 11.6; Sinitshenkova
and Zherikhin 1996), and for a pseudoligotrophic (high O2 but with an active herbi-
vore guild) Early Cretaceous lake (Fig. 11.10; Zherikhin et al. 1999). These data
indicate that there was a major shift in lacustrine productivity and biotal complexity
later in the Mesozoic (Table 11.1). However, it appears that these changes likely
began during the Late Triassic. The trophic structure of the Jurassic Mesoleuctra–
Mesoneta Assemblage (Sinitshenkova and Zherikhin 1996) occurred throughout the
warm temperate region of Siberia and other Eurasian and perhaps Gondwanan
Jurassic localities (Sinitshenkova 2002). In the lowlands at temperate latitudes, par-
ticularly in Eurasia, these shallow, hypotrophic (type B) lakes of the Jurassic con-
sisted of depauperate lentic and lotic faunas that allowed high O2 levels to accumulate,
and with a near exclusive trophic emphasis on detritivory (Fig. 11.6). The high abun-
dance of O2 throughout these hypotrophic lakes, and particularly the benthos, is
attributable to the absence of respiring organisms to sufficiently take up the O2 that
was being produced by autotrophic microorganisms (Sinitshenkova and Zherikhin
1996). In addition, Sinitshenkova and Zherikhin (1996) state that these lakes were
littered with incoming ginkgoalean and czekanoskialean plant detritus that exerted a
negative control on productivity of the lake by having an antimicrobial effect on
decomposers, judging by the negative effect that modern Ginkgo biloba litter has on
modern lake productivity (Samylina 1988). Consequently, productivity in these lakes
had two, related features: there was poor development of a detritivore base, in part
likely hindered by the dominant terrestrial vegetation of the time that favored build-
ups of O2. In addition, there was absence of large, especially vascular, hydric plants
that would allow herbivores to expand herbivory, as seen in later Cretaceous lakes
(Fig. 11.10). The subsequent changeover is notable, particularly establishment of a
guild of grazing herbivorous organisms, present in pseudoligotrophic Paleolake
Baissa, a type C lake but also with high O2 levels, is notable. This shift during the mid
Early Cretaceous from a detritivore-based to herbivore-driven food web, occurred in
the switch from Type A and B to Type C to E lakes, documented in Eurasia, and is
probably the single most important event associated with the MLR. The ecologic
structure of Early Cretaceous Paleolake Baissa differs significantly from that of its
predecessor lacustrine biotas during the Jurassic (Table 11.1).
The basic trophic structure of Cretaceous Paleolake Baissa was a highly produc-
tive, pseudo-oligotrophic lake in which there were relatively low levels of dissolved
nutrients supporting an abundant and diverse standing crop of green plants, espe-
cially algae. Paleolake Baissa apparently is a lake type without a clear modern ana-
log. The algae was limited by high consumption levels, but promoted a complex,
herbivore-based food web within an ecosystem of de-emphasized detritivore food
chains (Sinitshenkova 2002). The elevated O2 levels were sufficiently depressed to
allow a diverse, detritivorous insect fauna (Zherikhin et al. 1999) at greater depths
in the water column, the hypolimnion. At intermediate depths, in the mesolimnion,
but below the surface-water layer of the epilimnion, there was an herbivore com-
munity of grazing and algivorous gastropods and insects, including case-bearing
caddisflies, which were supported by abundant and diverse benthic, planktonic, and
floating algae (Sinitshenkova 2002). Paleolake Baissa was an early lake ecosystem
where aquatic invertebrate herbivory played a significant trophic role. Other coeval
deposits of similar origin are the lacustrine beds of the Yixian Formation in China
(Barrett 2000; Pan et al. 2011) and the Las Hoyas wetland deposits of Spain
(Buscalioni et al. 2016 and references therein).
In particular, the Las Hoyas fossil site, which is now interpreted as a freshwater
carbonatic, lentic wetland, has been analyzed recently from a trophic-web approach
(Buscalioni et al. 2016). According to this study, hydrophytic vegetation is domi-
nated by charophytes and aquatic angiosperms. The large mass of hydrophytic
plants allowed the presence of abundant grazers, such as ostracods, gastropods, spe-
laeogryphaceans, and aquatic insects, pointing to the importance of herbivory in this
trophic web, as is the case of Paleolake Baissa. Various worm-like organisms have
been regarded as feeding on phytoplankton and zooplankton, whereas unionid
bivalves were suspension feeders and crayfish are considered omnivorous scaven-
gers. Some large aquatic insects may have preyed on fish larvae and medium-sized
fish consumed various aquatic insects. Buscalioni et al. (2016) also emphasized the
importance of insects and fish in lake productivity, which is consistent with data
from modern wetlands.
During the Cretaceous, physical and chemical lake conditions exhibit a major
shift favoring development of certain lake types based on a variety of physiochemi-
cal conditions. These features were increased aridity, greater topography, more
unstable and variable lake levels, enhanced variability in annual temperature, greater
water-column stratification, and higher water turbidity. Chemical indicators show
elevated nutrient levels, a tendency toward alkaline over acidic water pH’s, and
more variable and lower dissolved oxygen levels. These physiochemical shifts pro-
vided an opportune environment for biotal changes, such as less allochthonous plant
detritus as input, much greater turnover rates, higher microbial activity, and consid-
erably higher algal production, but anomalously, lower macrophyte production. For
food-web development, there was a shift from detritivore to herbivore processing of
primary productivity (via more intensive levels of herbivory), greater complexity of

food webs, and an emphasis from K-selected to r-selected evolutionary strategies.
Among benthic insects there was a trophic shift from shredders and scrapers to
filter-feeding and live-plant ingesters as the dominant feeding types (Sinitshenkova
and Zherikhin 1996).
Finally, a recent examination of the 48 million-year-old deposit of Paleolake
Messel, near Darmstadt, in central Germany, produced an exhaustive food-web
analysis that was made for 94, well-documented organisms, including amphibious
taxa, that constituted the lake portion (Fig. 11.13) of the total food web (Dunne et al.
2014). This study provides valuable information on food webs in the aftermath of
the MLR. The Messel lacustrine web was constructed by using highly-resolved,
well-documented data of feeding relationships among all taxa. The lacustrine food-
web data from Messel indicate an ecologic structure very similar to modern lake
webs, and notably, a stability of trophic relationships that were likely in place
shortly after the K-Pg extinction (Dunne et al. 2014). However, the fact that
Chlorococcales dominated over diatoms in Paleolake Messel represents a departure
from the situation in modern lakes.
11.7.4 Behavioral Convergence Between Marine

and Continental Benthic Fauna
A comparison between marine and lacustrine ichnofaunas suggests that use of

freshwater infaunal ecospace may have been less complete than in marine environ-
ments (Miller and Labandeira 2002). In addition, levels of ichnodiversity and com-
plexity of biogenic structures are significantly lower in lakes than in marine settings
(Buatois and Mángano 1998). Evaluation of the extent and limitations of behavioral
convergence on both sides of the salinity barrier may help to understand evolution-
ary constraints on the lake colonization process.
Ichnotaxonomic problems undoubtedly prevented an adequate recognition of the
similarities and differences between the marine and continental realms. There are two
sides to this problem: uncritical use of marine ichnotaxa in continental settings and
unsupported erection of new ichnotaxa apparently exclusive to continental settings.
Whereas the former has contributed to the overemphasis of behavioral convergence,
the latter promoted its lack of appreciation. This is essentially an issue with freshwa-
ter, rather than terrestrial trace fossils. The vast majority of the latter (e.g.
Coprinisphaera, Termitichnus, Vondrichnus, Celliforma, Eatonichnus, Castrichnus,
Quirogaichnus) are exclusively found in paleosols, and their ichnotaxonomic intrica-
cies have been clarified by extremely detailed and solid work (e.g. Genise 2000, 2004;
Laza 2006).
Freshwater trace fossils tend to be characterized by relatively simple morpholo-
gies, typically including facies-crossing ichnotaxa that occur in marine environ-
ments (e.g. Gordia, Helminthoidichnites, Cochlichnus). In contrast, there are many
ichnotaxa that are restricted to marine environments, including the typical elements
of the Nereites and Zoophycos Ichnofacies and a considerable number of those in
the Cruziana Ichnofacies (Buatois and Mángano 2007). Arguably, the best example
of this confusion is Scolicia. This ichnogenus consists of bilobate or trilobate hori-
zontal structures displaying a complex meniscate backfill and two parallel strings,
representing the feeding and locomotion activities of irregular echinoids (Smith and
Crimes 1983; Uchman 1995; Bromley et al. 1997), which are restricted to the
marine realm. Although Scolicia has been the subject of a number of taxonomic
revisions and is well understood among ichnologists, surprisingly the name contin-
ues to be applied for continental, simple epirelief furrows that lack the complex
morphology of this ichnogenus (Turner 1978; Hasiotis 2002, 2004; Lovelace and
Lovelace 2012). Less commonly, other typical marine ichnotaxa, such as
Paleodictyon, Nereites, and Chondrites, are used for much simpler freshwater
trace fossils. Structures included in Paleodictyon from freshwater settings (e.g.
Archer and Maples 1984; Wu 1985; Pickerill 1990) are quite simple, and do not
display the regular pattern that characterizes this ichnogenus in marine turbidites.
A feeding trace referred to as Nereites in lacustrine turbidites (Hu et al. 1998)
lacks the internal, complex backfill structure of this ichnogenus, displaying only
superficial similarities with Nereites. Feeding traces doubtfully assigned to
Chondrites in lacustrine deposits (Smith et al. 1982; Kim et al. 2005) may super-
ficially resemble this ichnogenus, although the dichotomous, primary successive
branching that is diagnostic of Chondrites has never been documented in conti-
nental settings.
By contrast, some names that have been introduced for freshwater trace fossils
fail to pass ichnotaxonomic validation. The classic example is Isopodichnus, a com-
bination of short, bilobate resting traces and more continuous bilobate trails.
Although Isopodichnus was frequently used in the past for continental bilobate
trace fossils, recently its use essentially has been abandoned, following convincing
demonstration that it is a junior synonym of Rusophycus and Cruziana (Bromley
1996). More recently, however, meniscate trace fossils present in continental depos-
its, previously referred informally as “adhesive meniscate burrows” (Hasiotis 2004),
were subsequently included in a new ichnogenus, Naktodemasis (Smith JJ et al. 2008).
However, Naktodemasis clearly falls within the diagnosis of Taenidium (Krapovickas
et al. 2009; Díez-Canseco et al. 2016), an ichnogenus known from marine environ-
ments as well.
Behavioral convergence may also be evaluated by examining categories of
ichnodisparity (architectural designs) rather than ichnotaxa. Of the 58 architectural
design categories defined for invertebrate bioturbation structures (see Chap. 16),
none are exclusive to freshwater settings, six are only present in terrestrial settings
(vertical to oblique simple ornamented burrows; isolated, clustered, or intercon-
nected cells; chambers with discrete thick linings; excavated chambers with thin
linings undetachable from rock matrix; interconnected chambers and boxworks;
and Holes, pits and galleries in walls and fillings), and one is shared by terrestrial
and marginal-ichnofaunas (Simple to complex burrows with terminal chambers).
Indeed, the only ichnogenus in the latter present in both marine and continental
environments is Macanopsis, which actually occurs in backshore coastal areas,

rather than in fully marine settings.
Freshwater ichnofaunas are represented by twenty architectural design catego-
ries, all of which also occur in marine environments: (1) simple horizontal trails
(Archaeonassa, Circulichnis, Cochlichnus, Gordia, Helminthoidichnites,
Helminthopsis, Herpystezoum, Mermia); (2) trails with undulating transverse bars
and furrows (Steinsfjordichnus); (3) bilobate trails and paired grooves (Cruziana,
Didymaulichnus, Diplopodichnus); (4) trackways and scratch marks (e.g.
Diplichnites, Hamipes, Keircalia, Lithographus, Siskemia, Stiallia, Stiaria,
Tasmanadia, Umfolozia); (5) bilaterally symmetrical short, scratched impressions
(e.g. Avolatichnium, Rotterodichnium, Tonganoxichnus); (6) bilaterally symmetri-
cal short, scratched burrows (e.g. Rusophycus); (7) passively filled horizontal bur-
rows (Palaeophycus); (8) simple actively filled (massive) horizontal burrows (e.g.
Planolites); (9) simple actively filled (meniscate) horizontal burrows (e.g. Scoyenia,
Taenidium, Beaconites); (10) simple actively filled (pelletoidal) horizontal burrows
(e.g. Edaphichnium, Sphaerapus); (11) complex actively filled (meniscate) horizon-
tal burrows (Scolecocoprus); (12) horizontal branching burrow systems
(Labyrintichnus, Paracanthorhaphe, Shanwangichnus, Vagorichnus); (13) horizon-
tal burrows with horizontal to vertical branches (Ctenopholeus, Treptichnus); (14)
burrows with horizontal spreiten (Fuersichnus, Rhizocorallium); (15) isolated and
serial almond-shaped burrows (Calceoformites, Lockeia, Ptychoplasma); (16) verti-
cal simple burrows (e.g. Skolithos); (17) vertical U- and Y-shaped burrows (e.g.
Arenicolites, Diplocraterion); (18) vertical multiple U- and Y-shaped burrows
(Polykladichnus); (19) Simple to complex burrows with terminal chambers
(Camborygma, Castrichnus, Katbergia, Macanopsis, Platicytes); and (20) mazes
and boxworks (Thalassinoides, Spongeliomorpha, Virgaichnus)
The common feature of this list is that these freshwater biogenic structures col-
lectively represent relatively simple behaviors. Miller and Vokes (1998) categorized
trace fossils as incidental and deliberate. Incidental trace fossils are those that record
a single or dominant behavioral activity, and typically are structurally simple.
Deliberate trace fossils are those that represent restructuring of habitats, modulation
of disturbances, and control of food resources, and are typically structurally com-
plex. Freshwater trace fossils tend to fall within the first category. In some cases, the
same groups of producers were involved (Lockeia produced by both marine and
freshwater bivalves). In other cases, true behavioral convergence can be invoked.
The U-shaped burrow Arenicolites in marine environments is produced by a wide
variety of organisms, such as polychaetes, echiuran worms, crustaceans, holothuri-
ans, and enteropneusts (e.g. Bromley 1996; Mángano et al. 2002), whereas in fresh-
water settings insects and oligochaetes are involved (e.g. McCall and Tevesz 1982;
Scott et al. 2012a). Amphipods produce U-shaped burrows in both freshwater and
marine settings. Behavioral convergence on both sides of the salinity barrier seems
to have occurred only with the simplest ethologic types. The most complex archi-
tectural categories, such as burrows with helicoidal spreiten, dichotomous branch-
ing burrows, and those included within graphoglyptids, do not have a freshwater
counterpart. This is clearly illustrated by the contrasting trace-fossils suites present

in lacustrine and marine turbidites (Buatois and Mángano 1998).
Although the term “salinity barrier” has been commonly used to contrast the
nature of freshwater and marine ichnofaunas, the term is somewhat misleading.
Differences between ichnodiversity levels and the degree of morphologic complex-
ity of trace fossils most likely are explained by the stability-time hypothesis devel-
oped by Sanders (1968), rather than salinity per se. According to this hypothesis,
species diversity – and parallel to this, the degree of complexity of biogenic struc-
tures – is a function of environmental stability or predictability of the environment.
Because lakes are considerably shorter-lived than oceanic basins, they tend to dis-
play lower taxonomic diversity (and ichnodiversity) levels and more simple struc-
tures than marine settings (Buatois and Mángano 1998). In addition, this line of
reasoning can be applied to understand contrasting ichnodiversity levels in different
lakes. For example, lakes from recently glaciated regions exhibit lower diversity
levels than those from the long-lived, large, and deep lakes such as Lakes Baikal
and Tanganyika (Saunders 1968).
Finally, regardless of the specifics of the MLR and lacustrine ecosystems in
general, there are at least two common themes between the macroevolutionary
aspects of marine and lacustrine settings. First, the overall trend in increased infau-
nalization discussed above also took place in marine basins, albeit with very differ-
ent timing, because infaunalization in lacustrine basins lagged behind the same
process in marine settings (Buatois and Mángano 1993a; Buatois et al. 1998a).
Second, an analogue of the onshore – offshore pattern recognized in marine com-
munities seems to be apparent in freshwater settings as well. An onshore origina-
tion of novelties and subsequent migration or expansion into deeper water has been
proposed in marine settings based on body fossils (e.g. Jablonski et al. 1983;
Sepkoski and Miller 1985; Jablonski 2005; Sepkoski and Sheehan 1983) and trace
fossils (Crimes and Anderson 1985; Bottjer et al. 1988; Jensen and Mens 1999). In
freshwater settings, an analogue of the onshore-offshore pattern is indicated by
increases in the depth and extent of bioturbation that took place progressively
through time, expanding from fluvial and lake-margin settings to permanent sub-
aqueous lacustrine environments (Buatois et al. 1998a). These commonalities
between the continental and marine trace-fossil records suggest the existence of
recurrent macroevolutionary patterns of animal–substrate interactions through
time (see Chap. 16).
11.8 Conclusions
Our review of the trace-fossil and body-fossil histories of the continental aquatic
record suggests that the Mesozoic Lacustrine Revolution (MLR) represents a signifi-
cant evolutionary event for lacustrine ecosystems that took place in a protracted fash-
ion in time and space. Although ichnologic data demonstrate that both lake-margin
and fully lacustrine deposits were colonized prior to the MLR, benthic activity
essentially was restricted to a very narrow zone at the sediment–water interface, leav-
ing the overwhelming portion of infaunal ecospace empty or underutilized. This situ-
ation commenced first in lake-margin environments during the Middle to Late
Triassic, as shown by widespread presence of mid-tier meniscate trace fossils and
deep-tier crayfish burrows. Colonization of the infaunal ecospace in these deposits
resulted in more intense sediment reworking, typically precluding preservation of
superficial trace fossils such as arthropod trackways. Although incipient penetration
of the substrate in fully lacustrine settings has been recorded locally during the
Middle to Late Triassic, ichnofaunas from these settings are typically reminiscent of
those from the Paleozoic, suggesting that colonization of lacustrine bottoms was
delayed in comparison with lake margins. Deeper penetration into the substrate
became more common during the Early Jurassic, but the intensity of bioturbation
remained low. The lacustrine mixed layer seems to have become well-established by
the Early Cretaceous, as indicated by intense bioturbation mottlings.
The MLR also is recorded by examination of the record of body-fossils, particularly
arthropods and angiosperms, and by trophic interactions at both the interorganismic
and entire-lake foodweb levels. From an ecosystem perspective, prior to the MLR,
lacustrine primary production by microorganisms was low and the depressed level of
invertebrate consumption was overwhelmingly detritivorous. After the MLR, lakes are
characterized by the appearance of macrophytes and a significantly more robust herbi-
vore guild of microorganisms and increasingly larger arthropods and vertebrates, espe-
cially grazers on plants other than microscopic and small algae. This transition occurred
during elevated lake oxygen levels, attributable to the insufficiency of detritivorous
microorganisms before the MLR and detritivorous plus dominant herbivorous organ-
isms after the MLR to use all available O2 for respiration. Given this context, the MLR
actually represents the incremental trophic shift from a detritivore-based to an herbi-
vore-based lacustrine biota and concomitant food-web adjustments. But this change
also represents a shift toward infaunalization that results from predatory escalation,
also seen in other parts of the lake environment postdating the MLR. Perhaps related to
predatory escalation is the distinct evolution of insect species in which their immature
aquatic stages become increasingly more environmentally decoupled from their con-
specific, exclusively terrestrially occurring adult stages. Based on the arthropod body-
fossil record, this switch took place during the Late Jurassic to Early Cretaceous, but
may have had a variable spatiotemporal occurrence in other mid-Mesozoic continents.
Information from both trace fossils and body fossils suggests that this shift to more
mixed trophic strategy consisting of detritivores, herbivores, predators, and other mac-
roguilds that form animal communities has persisted to the present since initiation of
the MLR, as demonstrated by the Paleolake Messel food web.
Acknowledgements Tony Ekdale, Leif Tapanila, and Mark Wilson provided useful feedback on
the Kenyan stromatolites. Richard Bromley and Ulla Asgaard showed to some of us (LAB and
MGM) the wonderful Flemming Fjord trace-fossil collection. Nilo Azambuja Filho and Adali
Spadini showed one of us (LAB) the lacustrine outcrops in the Sergipe-Alagoas Basin of Brazil.
Robin Renaut and Bernie Owen guided LAB through the impressive lakes of the Kenya Rift
Valley, resulting in a greater appreciation of the complexities of these systems. Robert Metz made
available trace-fossil photos from the Newark Supergroup. Ángela Buscalioni and Francisco
Poyato-Ariza provided information on the ecology of Las Hoyas. Ángela Buscalioni and Nic
Minter reviewed the chapter, offering valuable suggestions to improve it. Thanks go to Jorge
Santiago-Blay for photography of specimens in Fig. 11.7 and assistance with identifications.
Jennifer Dunne and Richard Williams rendered the food web in Fig. 11.13, and Finnegan Marsh
produced Figs. 11.6, 11.7, and 11.10. This is contribution 276 of the Evolution of Terrestrial
Ecosystems consortium the National Museum of Natural History in Washington, D.C., USA.
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Chapter 12
The End-Cretaceous Extinction
and Ecosystem Change
Conrad C. Labandeira, Francisco J. Rodríguez-Tovar, and Alfred Uchman
12.1 Introduction
One of the most phenomenal events in the history of Phanerozoic life was the end-
Cretaceous (K-Pg) mass extinction, occurring 66.04 Ma ago (Vandenberghe et al.
2012; Husson et al. 2014 for recent calibrations), an event that has been important
for the subsequent evolutionary and ecological history of the continental and
marine biota. Interest in this event was reignited during the early 1980s, when evi-
dence for extraterrestrial causation was proposed based on several lines of evidence
(Alvarez et al. 1980, 1984), later identified with an impact site (Hildebrand et al.
1991). Currently there is near consensus that the K-Pg event was caused by a bolide
hitting the carbonate platform in Yucatan, Mexico (but see Schoene et al. 2014),
which resulted in a significantly altered global environment, including altered sedi-
mentation patterns (D’Hondt 2005), wildfires (Wolbach et al. 1985), and elevated
atmospheric temperatures, and pCO2 and pO2 concentrations (Gale et al. 2001;
C.C. Labandeira, (*)

Department of Paleobiology, MRC-121, National Museum of Natural History, Smithsonian
Institution, P.O. Box 37012, Washington, DC 200013-7012, USA
Department of Geology, Rhodes University, Grahamstown 6140, South Africa
College Park, MD 21740, USA
College of Life Sciences, Capital Normal University, Beijing 100048, China
F.J. Rodríguez-Tovar
Department of Stratigraphy and Paleontology, University of Granada, Granada 18007, Spain
A. Uchman
Institute of Geological Sciences, Jagiellonian University, Kraków 30-063, Poland

DOI 10.1007/978-94-017-9597-5_12
266 C.C. Labandeira et al.
Beerling et al. 2002; Schulte et al. 2010). However, the most consequential, long-
lasting effects were on organisms, not only immediately after the event, but also
throughout the intervening time between the event and today.
In the continental realm, the most obvious group that became extinct were major
lineages of non-avian dinosaurs (Sereno 1999), which suffered a sudden fate
(Sheehan et al. 2000); the previously declining pterosaurs (Penny and Phillips
2004); and several lineages of birds (Hou et al. 1996, but see Cooper and Penney
1997) that underwent major diversity declines (Gelfo and Pascual 2001). For birds,
nine lineages survived that became the progenitors to the 42 extant orders of birds
(Jarvis et al. 2014). There was extinction of at least one and probably several major
clades of mammals (MacLeod 2005), although 39 lineages continued across the
boundary, giving rise to the 25 orders of mammals of today (Bininda-Emonds et al.
2007). The similar style of survivorship of bird versus mammal lineages across the
K-Pg boundary is noteworthy. Both major vertebrate lineages seem to support a
slow-fuse model (Bininda-Emonds et al. 2007; Jarvis et al. 2014), in which over-
whelmingly fewer, nondiverse lineages were present deep in the Cretaceous or ear-
lier. These comparatively few lineages were “triggered” soon after the end-Cretaceous
ecological crisis to provide a stunningly rapid cascade of evolutionary diversifica-
tion within 10–15 Ma into the numerous lineages of birds and mammals that pres-
ently occupy virtually every significant continental and marine habitat on the planet.
Other organisms in terrestrial and freshwater habitats suffered considerable
losses, including land plants (Johnson 2002; Wilf and Johnson 2004; McElwain and
Punyasena 2007), with important regional differences (Askin and Jacobson 1996);
bivalves (Hartman 1998; Hartman et al. 2009); crocodyliforms (Brochu 2004); and
many avian dinosaurs (Sereno 1999; Retallack 2004). Groups that were minimally
or not affected were amphibians (Archibald and Bryant 1990); and non-archosaurian
reptiles such as turtles, lizards, rhynchocephalians, amphisbaenians, and
choristoderes (MacLeod et al. 1997; Novacek 1999). Major insect lineages were
unaffected at the family level (Labandeira and Sepkoski 1993), although ecological
relationships with plants were significantly reduced (Labandeira et al. 2002a, b;
Wilf et al. 2006; Donovan et al. 2014). Spiders, predators overwhelmingly
consuming insects, also were not reduced in family-level diversity (Penney et al.
2003), and appear to track the family-level diversity of insects before and after the
event (Penney 2003).
For the marine realm, as in the continental realm, some clades experienced
significant or outright extinctions, whereas others did not (D’Hondt 2005). The
principle clades that suffered major extinctions at the K-Pg boundary were diatoms
(MacLeod 1998); calcareous nannoplankton (Bown 2005); other phytoplankton
(MacLeod et al. 1997); benthic and planktic foraminifera (Kuhnt and Collins 1996;
Arenillas et al. 2000; Molina 2015); scleractinian corals, particularly colonial taxa
inhabiting warmer waters (Rosen and Turnšek 1989); echinoderms (Jeffrey 2001);
mollusks such as belemnoids, ammonites (Marshall and Ward 1996) and inoceramid
and rudistid clams (Ward et al. 1991; Raup and Jablonski 1993; Lockwood 2003).
Vertebrate extinctions included many lineages of teleost fish (Friedman 2009,
2010), mosasaurs (MacLeod et al. 1997), and plesiosaurs (O’Keefe 2001).
Apparently, there were minimal levels of extinctions, if any, of radiolaria (MacLeod
12 The End-Cretaceous Extinction and Ecosystem Change 267
et al. 1997), dinoflagellates (Gedl 2004); probably ostracods (Brouwers and De

Deckker 1993); and brachiopods, which experienced a diversification event during
the early Paleocene (MacLeod et al. 1997).
It is evident that these data are overwhelmingly based on identifications and tal-
lies of body fossils, inventoried as either specimens or taxa, which represent diversi-
ties or abundances through time. A neglected but crucial aspect toward understanding
the K-Pg event and its recovery phase is the collection, analysis, and interpretation
of ecological data. When appropriately linked to various types of diversity data and
to assessments of phylogeny based on paleobiological occurrence data and molecu-
lar phylogenies, ecological data can yield insights into mechanisms and patterns
that may not be evident solely from a focus on taxonomic affiliations or occurrence
tallies in particular habitats. In this contribution, we attempt to fill this void by pro-
viding two approaches toward understanding the ecology of the K-Pg extinction in
both the continental and marine realms. For the continental realm, we will employ
the system of analyzing damage types (DTs), that record distinctive plant–insect
interactions throughout the K-Pg event to the late Paleogene (Labandeira et al.
2007). This approach has resulted in a fuller understanding of the patterns and
processes during this tumultuous interval in terrestrial earth history. From a differ-
ent branch of paleoichnology, we employ sedimentologically based tools for under-
standing marine organism–sediment relationships and behavioral correlates,
mainly based on detailed ichnofabrics from studies of polished sections and exam-
ined burrow fills, including isotopic composition studies, which yields a better
understanding of organism response to changes in the marine realm.
12.2 Evolutionary and Ecological Dynamics of the K-Pg

Event and Its Recovery
There are two basic aspects to the generation of new lineages and their associated
ecologies during a major mass-extinction crisis. The initial phase involves lineage-
sorting processes inherent in the extinction, whereas the subsequent phase molds
the selected lineages within the context of newly created ecospace.
12.2.1 Selectivity and Sorting During the Mass Extinction
The effect of the K-Pg event on the evolutionary trajectory of life has been exam-
ined extensively, particularly for marine mollusks (Jablonski 1989, 2001, 2005).
The K-Pg mass extinction resulted in dramatic decreases in standing diversity
(Jablonski 2005), although there were other significant patterns that have been
recorded related to lineage selectivity and its consequences on affected taxa
(Jablonski and Raup 1995). One of the more important conclusions of these studies
has been the central role that mass extinction selectivity plays in favoring certain
taxa-specific attributes over others (Jablonski 2001). The targeting of attributes
during mass extinctions differs substantively from times of ambient, background

extinction rates (Kitchell et al. 1986). In addition to providing dramatic decreases in
standing diversity, another pattern resulted in more widely distributed taxa becom-
ing considerably extinction averse compared to taxa with much narrower biogeo-
graphic ranges (Jablonski 2005). This extinction differential tends to favor
taxonomically higher-ranked taxa for survival over their lower-ranked subordinates.
Consequently, there was extinction of species with narrower geographic ranges,
many of which were members of more broadly distributed (and temporally persis-
tent) genera.
Perhaps the most important factor for selectivity during mass extinction was the
relationship between life habits and feeding biology (Jablonski 2005). Most buffered
of all were deposit feeders that occurred in the benthos or on sediment substrates
that subsisted on detritus or dead organisms (Arthur and Zachos 1987; Twichett
2006). Deposit-feeding clades suffered much lower extinction levels than those
groups occurring in the water column or having more direct trophic links to
photosynthetic organisms (Sheehan and Hansen 1986). Consequently, in the marine
realm, benthic deposit feeders and scavengers were favored over suspension and
filter feeders existing in the water column that were embedded in food webs
connected to photosynthetic organisms. For the continental realm, small
insectivorous animals and aquatic invertebrates in streams were favored over most
large-bodied vertebrates such as dinosaurs (Sheehan and Hansen 1986). Nevertheless,
Levinton (1996) suggested that deposit feeders should have suffered extinction
levels comparable to organisms more directly linked to photosynthetic organisms.
This alternative pattern would be attributable to all dead organic material ultimately
originating from more inclusive food chains of primary producers and their
herbivores, although the effects would have been separated by a temporal lag of 3–6
months after impact. Contrary evidence is the presence of a local detritivorous
earthworm fauna within several thousand years of the K-Pg event that consumed
organic material preserved regionally as lignites (Chin et al. 2013). Another life-
history source of immunity from extinction was marine organisms that possessed
planktic life stages, versus those that do not (Kitchell et al. 1986). Those organisms
with planktic life stages, such as many crustaceans with planktotrophic larvae,
preferentially survived the K-Pg extinction. A considerably longer effect, resulting
from a decrease of organic flux to the sea, were the negative consequences on
calcium carbonate production by marine plankton, which took a few million years
for the open-ocean ecosystem to be restored to full operational capacity, well into
the mid Paleocene (D’Hondt 2005).
12.2.2 Establishing New Lineages and Ecologies

During the Recovery
The recovery phase also had interesting evolutionary and ecological dynamics that
were separate from the much shorter, preceding extinction phase. One secondary
effect, based on a combination of theoretical models (Erwin 2001) and observations
(Jablonski 2001), was the probable lack of an association between the severity of
the extinction and the length of recovery. In part, this would have been attributable
to the collapse of ecosystems and thus the removal of ecospace that would disallow
a rapid, orderly, logistic recolonization of pre-existing habitats. The destruction of
ecospace was rebuilt, figuratively and trophically, from the ground up, and the
idiosyncratic nature of lineage survivorship suggests that the recovery pattern was
not attributable to global trophic principles, but rather represent a summation of
lineages resulting from disparate patterns in time and space (McKenna and Farrell
2006), with no single, unifying ecological explanation (Jablonski 2005).
The vagaries of organismic occupation of ecospace during the recovery phase
also indicate that origination rates of major lineages are reset at mass extinction
episodes, rather than at intervals of background extinction (Jablonski 1989). It is
during the recovery phase that entirely novel ecologies are created for surviving
lineages (Solé et al. 2002); indicating that mass extinction and subsequent recoveries
ultimately are the key intervals for establishing major evolutionary trajectories
(Erwin 1998; Krug and Jablonski 2012). Indeed, the effects of the K-Pg event are
reflected in current disruptions in the biogeographical distributions and ages of
marine bivalve genera (Krug et al. 2009), and in the times of origin of particular
extant plant–insect interactions (Labandeira 2005). These patterns indicate an
increase in origination rates that followed the K-Pg mass extinctions and eventually
reached a peak about 10 my after the event (Kirchner and Weil 2000). Effects of
these origination-rate increases persist to the present day. An associated, but
opposite, phenomenon is the survival of some lineages that survived the mass
extinction event in radically decreased diversity, only to be finally extirpated early
within a short-lived recovery (Erwin 2001). Such clades are termed “dead clade
walking” (Jablonski 2002), and their early, post-event demise probably involved
mismatches among biotic interactions.
12.3 The Continental Perspective of Ecological Disruption

and Its Consequences
In this section, the effects of the K-Pg crisis on aquatic and terrestrial communities
will be discussed, although the focus will be insects and their associations with
plants from floras spanning the K-Pg boundary in the Western Interior of North
America (Labandeira et al. 2002a, b; Wilf et al. 2000, 2006; Winkler et al. 2010;
Labandeira and Currano 2013; Donovan et al. 2014). The analytical techniques used
in these studies were developed during the late 1990’s (Wilf and Labandeira 1999;
Wilf et al. 2001; Labandeira et al. 2007), and have expanded in scope to allow
examination of plant–insect herbivore dynamics immediately preceding, during,
and following the Paleocene–Eocene Thermal Maximum (PETM) floras that occur
in the same or nearby basins (Wilf and Labandeira 1999; Wilf et al. 2001; Currano
et al. 2008, 2010). Examination of plant-host and insect herbivore dynamics of the
PETM studies can be seen as an extension of the K-Pg work, the latter detailed
below, particularly as it documents the final stage of recovery from the major effects
of the end-Cretaceous ecological crisis.
Parenthetically, it should be noted that study of plant–insect interactions in the
fossil record never has been a major theme in paleoichnology. Mainstream
paleoichnology has been principally driven by varied studies of the organismic
alteration of sediment substrates in the fossil record from a wide variety of
continental and marine depositional environments (Bromley et al. 2007). By
contrast, much of the recent study of plant–insect interactions, principally herbivory,
has targeted well-preserved, angiosperm-dominated fossil floras from several well-
studied areas worldwide, including western North America (Wilf et al. 2001, 2006;
Labandeira et al. 2002a), western Europe (Wappler et al. 2009, 2012), east Africa
(Currano et al. 2011), and southern Patagonia and northern Colombia in South
America (Wilf et al. 2005; Wing et al. 2009). The two approaches—sedimentologi-
cal and plant–insect associational―have allowed for differing collection proce-
dures, statistical protocols, interpretations, research-driven questions, and reference
to overarching theory.
12.3.1 Aquatic Communities
The fate of invertebrates in freshwater aquatic communities during the K-Pg event
is minimally understood, but is best demonstrated for bivalves, which experienced
a major contraction of taxa that survived into the Paleocene (Hartman 1998). Major
insect lineages such as mayflies, dragonflies, stoneflies, caddisflies, nematocerous
flies, and beetles, which have actively feeding aquatic immatures, did not suffer
extinction at the family level above that of the background level (Labandeira 2005),
and likely were preferentially buffered against extinction (Sheehan and Hansen
1986). There is virtually no data on survivorship of major freshwater malacostracan
lineages across the boundary.
12.3.2 Terrestrial Communities
Terrestrial communities house the bulk of biodiversity in the modern world, and
undoubtedly did so during the latest Cretaceous. Most of this biomass occurs as
land plants and arthropods, the two hyperdiverse groups that provide macroscopic
structure to terrestrial ecosystems. Because of the ecological importance of these
two groups and the absence of body-fossil insects near the K-Pg boundary (e.g.,
Larsson 1975; Pike 1994), a plant–insect associational analysis was conducted
across the boundary in the Williston Basin of North Dakota, where abundant,
diverse, and well-preserved floras are widespread (Johnson 2002; Labandeira et al.
2002a, b; Wilf et al. 2006; Donovan et al. 2014). When these ecological studies of
K-Pg plant–insect interactions (Labandeira et al. 2002a, b) were linked to those of
the PETM (Wilf et al. 2006; Currano et al. 2008, 2010; Donovan et al. 2014), the
results (discussed below) parallel those found in the evolutionary studies of birds
(Jarvis et al. 2014) and mammals (Bininda-Emonds et al. 2007), also occurring
within 10 to 15 my after the boundary. Additionally, examinations of K-Pg to PETM
floras have been extended to western Europe (Wappler et al. 2009, 2012; Dunne
et al. 2012), and the results are suggestive of the pattern in North America. However,
a valid test of Williston Basin data in Europe would require a regional, continuous
section of well-preserved floras traversing the K-Pg boundary, currently absent, for
further progress toward understanding the broader biogeographical extent of this
global event.
12.3.2.1 Plants
The change in plant diversity across the K-Pg boundary was once thought to have
been gradual (Hickey 1981) or stepwise (Frederiksen 1989), based on megafloral
and palynological data, respectively. However, subsequent evidence from more
intensely collected sections from the Williston Basin have clearly established a
catastrophic extinction pattern (Johnson 2002), equivalent to a 57 % maximum
estimate for extinction based on megafloral species-level data (Wilf and Johnson
2004), and a 30 % minimum estimate based on palynological genus-level data
(Nichols 2002). These more recent data indicate a major turnover from more highly
diverse and warm-adapted late Maastrichtian floras occupying a variety of habitats,
to depauperate, cool-adapted early Paleocene floras that largely occupied mire
habitats (Johnson 2002).
12.3.2.2 Insect Herbivory
With the possible exception of the Denver Basin to the south, the Williston Basin
floral sequence represents the best combination of conditions for an analysis of
insect herbivore patterns on floras straddling the K-Pg boundary. These floras are
stratigraphically linked to a composite section of 183 m of strata that represent 2.2
my, of which the lower 1.4 my interval is of latest Cretaceous age and the upper 0.8
my interval is of earliest Paleocene age (Fig. 12.1). Within this system of floras,
13,441 specimens representing 380 plant-organ morphotypes were examined from
143 localities derived from 106 discrete stratigraphic levels (Fig. 12.1). Typical of
most floras for this time interval, the Williston Basin floras are dominated by dicoty-
ledonous angiosperms (86.3 %), but also include monocotyledonous angiosperms,
conifers, cycads, ginkgos, ferns, a horsetail, and bryophytes (Fig. 12.2). The total
assemblage represents a variety of fluvially dominated environments, including
abandoned channels, sand bars, overbank deposits, ponds, and swamps, which
formed a mosaic of distinctive habitats. Additional details are provided in the two
reports from this study (Labandeira et al. 2002a, b).
Fig. 12.1 Stratigraphic and sampling data for the 51 insect-mediated damage types from a
183 meter composite section straddling the K-Pg boundary (orange bar), from the Williston Basin
of southwestern North Dakota. The estimated time duration for this interval is 2.2 million years
(my) before present, of which 0.8 my are assigned to the postboundary interval and 1.4 my are
allocated to the pre-boundary interval. The Cretaceous strata are within the Hell Creek and Fort
Union formations, and the Paleocene strata are entirely within the Fort Union Formation. Placement
of the zero datum is at the K-Pg boundary, not the Hell Creek/Fort Union formational contact,
because the contact is diachronous with respect to the K-Pg boundary timeline. Depicted are the
raw presence/absence data for damage types, categorized by functional feeding group and DT
subgroup at bottom, and representing 106 discrete horizons. Eight damage types have single
stratigraphic occurrences (dots); the ranges of those with multiple stratigraphic occurrences are
shown as vertical shaded bars, categorized by host specificity from the color scheme at upper left
(see text). Of the 14 specialized damage types with multiple stratigraphic occurrences that
disappear at or below the K-Pg boundary, six (42.9 %) reappear during the latest Paleocene to
middle Eocene of Wyoming and Utah. All associations are extant today. Four confidence intervals
that exceed our sampling range are shown without end bars. Total specimen frequencies are given
at right, including all plant organs; note logarithmic scale. The gray horizontal lines in the main
figure and corresponding gray squares at right indicate the 14 horizons with more than 200
specimens of identified dicot leaves; meter levels for these floras are: −75.0, −65.0, −56.8, −47.8,
−36.9, −31.4, −15.0, −3.6, +0.2, +1.3, +30.9, +42.4, +50.0. Abbreviations: mar margin feeding,
skeleton. skeletonization, spl. surface feeding, p. piercing-and-sucking, o. oviposition. Limitations
on graphical presentation cause the flora at +0.2 m, which are Paleocene, to appear within the
orange line representing the K-Pg boundary. From Labandeira et al. (2002b), reprinted with
permission by the National Academy of Sciences
Fig. 12.2 A spectrum of generalized (c, g, i, j) and specialized (a, b, d, e, f, h, i, k) plant–insect
associations from the Williston Basin of southwestern North Dakota. Associations range from the
earliest Paleocene at upper left, 14.4 m above the K/T boundary, and continue to the older
associations of the latest Cretaceous at lower right, 85.5 m below the boundary. All material is from
the Denver Museum of Nature and Science (DMNH) or the Yale Peabody Museum (YPM).
Following each plant host are, respectively, morphotype number (indicated by the prefixes HC or
FU), specimen number, NMNH locality number (loc.), and ± meter distances from the K/T
boundary. Damage types are indicated by the prefix DT. Scale bars: solid, 1 cm; backslashed,
0.1 cm. (a) Two linear mines with oviposition sites (arrows), following secondary and then primary
venation, terminating in a large pupation chamber (DT59) on the dicot Paranymphaea crassifolia
(FU1), DMNH-20055, loc. 563, +14.4 m. (b) Single gall (DT33) on primary vein of Cercidiphyllum
genetrix (Cercidiphyllaceae, FU5), DMNH-20042, loc. 562, +8.4 m. (c) Free feeding (DT26) on
Platanus raynoldsi (Platanaceae, FU16), DMNH-20035, loc. 2217, +1.3 m. (d) Skeletonization
(DT51) on a probable lauralean leaf (HC32), DMNH-19984, loc. 2097, −31.4 m. (e) Multiple galls
(DT33) on Trochodendroides nebrascensis (Cercidiphyllaceae, HC103), DMNH-19976, loc.
1489, −33.7 m. (f) Initial phase of a serpentine mine (DT45) on Marmarthia pearsoni (Lauraceae,
HC162), DMNH-7228, loc. 2087, −36.9 m. (g) Cuspate margin feeding (DT12, arrow) on
Metasequoia sp. (Cupressaceae, HC35), DMNH-13108, loc. 567, −56.8 m. (h) Serpentine leaf
mine (DT43) assigned to the Nepticulidae (Lepidoptera) on unidentified Rosaceae (HC80), YPM-
6367a, loc. 567, −56.8 m. (i) Hole feeding pattern (DT57) on an unknown genus of Urticales
(HC81), DMNH-19539, loc. 2203, −56.8 m. (j) General skeletonization (DT16) on Erlingdorfia
montana (Platanaceae, HC57), DMNH-11013, loc. 571, −61.7 m. (k) Large scale-insect
impressions (DT53) centered on primary veins of E. montana, DMNH-18829b, loc. 571, −61.7 m.
(l) Slot hole feeding (DT08) on an unidentified genus of Platanaceae (HC109), DMNH-18658, loc.
434, −88.5 m. See Labandeira et al. (2007) for descriptions of damage types. From Labandeira
et al. (2002b), reprinted with permission by the National Academy of Sciences
Of the examined specimens, 9292 (69 %) were late Maastrichtian and 4149
(31 %) were early Paleocene in age. This breakdown approximately was associated
with the Hell Creek Formation, and the Fort Union Formation, respectively, but
since the Hell Creek–Fort Union formational contact is diachronous with respect to
the K-Pg boundary (the zero datum of Fig. 12.1), some earliest Paleocene floras
were within the uppermost Hell Creek Formation. Sampling did not favor either the
Paleocene or the Maastrichtian portions of this study, as specimen coverage was
approximately equal for any 5-m interval above and below the boundary. In addition,
confidence intervals were placed on the tops of ranges for each damage type (or DT)
to correct for sampling intensity and to provide estimates for the likely time of
extinction for each relevant DT (Labandeira et al. 2002a).
The presence–absence matrix represented 51 DTs from eight functional feeding
groups (FFGs): hole feeding, margin feeding, skeletonization, surface feeding, gall-
ing, leaf mining, piercing-and-sucking, and oviposition (Figs. 12.1, 12.2, and 12.3).
Each DT was categorized as to whether it was a generalized, intermediate, or spe-
cialized interaction (Figs. 12.1 and 12.2), but with particular attention to document-
ing unique host-specialist associations (Fig. 12.3), based on a variety of criteria.
Three major patterns resulted from an analysis of the plant–insect interactions data-
set. First, generalized DTs are better represented than intermediate and specialized
DTs in the Paleocene side, with all generalized DT’s crossing the boundary whereas
10 of 16 of the intermediate and 6 of 20 of the specialized cross the boundary.
Second, of the 14 Maastrichtian DTs that are represented by large sample size, 10
have a last appearance just below the boundary, indicating extirpation at or just
below the K-Pg boundary, a finding buttressed by confidence intervals. Third, after
the decrease of the early Paleocene, herbivory increases in both frequency and rich-
ness during the later Paleocene. Most of this herbivory is generalized, as the more
specialized DTs remain consistently low in frequency and richness. The frequency
and richness of insect damage was analyzed throughout the section, and confined to
the 14 discrete horizons (Fig. 12.1) and to the most abundant, identifiably dicot
leaves to avoid biases in DT sampling on less abundant and poorly preserved non-
dicot leaves. These analyses show a decrease in herbivory at the boundary, with the
most significant decrease attributed to the intermediate and specialized DTs (data
Fig. 12.3 (continued) (i) Detail of mine in h, showing median frass trail at upper left. (j) Three
aborted leaf mines, two of which have coiled initial phase (loc. 900, DMNH-7325). (k) Complete
leaf mine with extensive terminal chamber illustrating trail (loc. 900, DMNH-7264). (l)
Enlargement of terminal mine chamber in k, showing path of undulatory frass trail, indicated by
white arrows. (m) Two aborted leaf mines with coiled initial phases (loc. 428, DMNH-7498). (n)
Enlargement of leaf mine at upper left in m, showing coiled (darkened) initial phase. (o)
Enlargement of leaf mine at lower right in m, revealing coiled leaf mine at lower right in k, indi-
cating coiled (darkened) initial phase, and subsequent curvilinear phase along median primary and
branching secondary veins (arrow). (p) Two adjacent leaf mines aborted early in development (loc.
900, DMNH-7313). Abbreviations: DMNH Denver Museum of Nature and Science, YPM Yale
Peabody Museum. Scale bars: 1.0 cm, solid; 0.1 cm, striped. From Labandeira et al. (2002a),
reprinted with permission by the Geological Society of America
Fig. 12.3 Host-specific association between Marmarthia pearsoni (Laurales) and a gracillariid
leaf miner (Lepidoptera), from the uppermost Hell Creek Formation. This highly stereotyped dam-
age type, DT45, represents a specialized serpentine miner typically with an initial coiled phase, a
subsequent curvilinear trajectory, and modestly expanded terminal chamber. It occurs exclusively
on plant host HC162 at YPM localities 900 and 428. (a) Near-complete leaf mine following the
primary venation (loc. 900, DMNH-7313). (b) Enlargement of a, with dark colored, medial frass
trail detectable at upper left corner, along a primary vein. (c) Complete leaf mine with terminal
chamber at upper right (loc. 900, DMNH-7263). (d) Enlargement of latter serpentine phase and
terminal chamber of leaf mine in c. (e) Close-up of terminal chamber displaying dark frass trail
(top arrow) and chamber edge (bottom arrow). (f) Fragment of leaf with a portion of leaf mine; note
coiled early phase (loc. 900, DMNH-7199). (g) Close-up of mine in f. (h) Complete leaf mine bounded
by median and lateral primary veins of plant host, typical for this species (loc. 900, DMNH-20023).
Fig. 12.4 Plant and insect-feeding diversity for bulk floras, standardized to sample sizes of 400
leaf specimens each. Orange-yellow data points are Cretaceous floras; blue data points are
Paleocene floras. Ma million years ago. Plant richness (a) was standardized by means of rarefaction,
with error bars indicating 95 % confidence intervals. Insect damage was standardized by means of
random resampling without replacement, with ±1σ error bars around the mean of 5000 iterations,
both for (b) all damage morphotypes and (c) mine morphotypes only. There is a strong negative
correlation of plant and insect damage richness for Mexican Hat (Mex. Hat) and Castle Rock
(C. Rock). A separate analysis (not shown in the figure) excluded most external feeding and other
generalized damage morphotypes, yielding results nearly identical to (b). Abbreviations: K-T
Cretaceous–Paleogene boundary, P-E Paleocene–Eocene boundary. Reprinted from Wilf et al.
(2006), with permission from the American Association for the Advancement of Science
not shown; Labandeira et al. 2002a, Fig. 3). The sustained decrease of herbivory
above the boundary, and the failure of intermediate and specialized DTs to recover
do record a major event affecting insect herbivores, documented in particular for
leaf-mining taxa (Donovan et al. 2014).
These results were tested in a subsequent study (Wilf et al. 2006), in which insect
feeding damage from 14,999 dicot leaves were examined from 14 latest Cretaceous,
Paleocene, and early Eocene sites to understand post-extinction patterns of herbiv-
ory, including generalized and specialized forms of interactions (Fig. 12.4).
Expectedly, most of the Paleocene sites displayed low richness of plants and insect
damage, with two unexpected exceptions in the early Paleocene, both within 1.7 my
of the end-Cretaceous extinction. One site from the Denver Basin, Castle Rock,
showed exceptionally high plant diversity but virtually no specialized feeding;
another site from the Williston Basin, Mexican Hat, conversely exhibited a typically
depauperate Paleocene flora but high levels of specialized herbivory. These dispa-
rate results indicate that, for about 2 my after the end-Cretaceous extinction, local
community structure of plants and insect herbivores remained significantly unbal-
anced, and did not regain latest Cretaceous levels of herbivory and specialization
until much later, during the late Paleocene.
12.3.2.3 Implications for Macroevolutionary Patterns of Specialist

Insect Herbivores
The above studies are based on assessments of insect herbivore functional feeding
groups and their damage types in floras across the K-Pg boundary, through the
Paleocene, and well into the early Eocene. These data may be important for detect-
ing the immediate and longer-term ecological processes of plant–insect interactions
after a major environmental perturbation. Given the long lag times toward increased
levels of herbivory and host-plant specialization following the K-Pg event, it would
appear to support the gradual rather than instantaneous colonization of plant hosts
after a major ecological crisis. This pattern also is detected from examination of
specialized leaf-mining lineages and their delayed colonization of available plant
hosts (Lopez-Vaamonde et al. 2006; Donovan et al. 2014), consistent with the long
lead-times to diversification predicted by the resource abundance–dependent diver-
sification hypothesis (Nyman et al. 2012). The post-event fossil pattern also is
inconsistent with geologically rapid, synchronous patterns of insects that co-radiate
onto their host plants, documented for certain chrysomelid beetles and their bursera-
ceous hosts (Becerra 2003). Alternatively, these data could address the issue of
quick, local adaptation to allopatric populations of novel host plants from a broadly
distributed specialist species (Rosenzweig 1995; Zvereva et al. 2010).
If host specialists were disproportionately extirpated at the K-Pg mass extinction
event, then specialization of insect herbivores could be an evolutionary dead end. A
tendency toward lineage phylogenetic stasis has been documented for recent clades
of leaf miners (Connor and Taverner 1997; Lopez-Vaamonde et al. 2003), those
gallers with more limited host ranges (Hardy and Cook 2010), and some
Dendroctonus bark beetles (Kelley and Farrell 1998). This especially would be true
if the lineages of host-plant specialists are clustered in particular clades vulnerable
to a mass extinction (Roy et al. 2009). However, there are good reasons to indicate
that some specialized relationships are not evolutionary cul-de-sacs (Colles et al.
2009); counterexamples include doniciine beetles on aquatic reeds (Kölsch and
Pedersen 2008) and perhaps other Dendroctonus bark beetle taxa (Kelley and Farrell
1998). While host-plant specialization may lead to extinction at times of major eco-
system crises, such as the K-Pg, during other, much more prolonged intervals of
background extinction, specialist and generalist herbivore lineages may experience
stasis and have bidirectional acquisition of host-plant feeding preferences
(DiMichele et al. 2004; Forister et al. 2012; Thompson 2013).
12.3.2.4 Insect Pollination
The data and methods of assessing patterns of insect pollination across the K-Pg
boundary are considerably more difficult than that those that evaluate insect her-
bivory. The palynological record, however, can reveal broad trends in the frequency
of zoophilous (insect vectored) versus anemophilous (wind dispersed) pollen.
Several examinations have documented a significant reduction of zoophilous pollen
at the K-Pg boundary (Frederiksen 1989; Sweet and Braman 2001), indicating a
disproportionate extinction of pollen vectored by insects at the boundary. In one

study documenting a lineage of dominantly Late Cretaceous zoophilous pollen
across the K-Pg boundary, a sole anemophilous palynospecies evidently survived
the event in a local section (McIver et al. 1991). Other lines of evidence also indi-
cate the emergence of pollinators immediately after the K-Pg event. One is a fortu-
itous occurrence of an entrapped stingless bee associated in a flower of its orchid
host found in Dominican amber (Ramírez et al. 2007). A molecularly based recon-
struction of orchid phylogeny based mostly on this fossil indicated that the time of
origin of the insect-pollinated orchid lineage was immediately after the K-Pg event.
This result parallels the same time of origin, using a similar method of phylogenetic
reconstruction, for nymphalid butterflies (Wahlberg et al. 2009), another major pol-
linator clade likely originating in the wake of the K-Pg crisis.
12.3.2.5 Blood Feeding on Dinosaur Hosts
Some associations are known from the Late Cretaceous involve live and dead dino-
saurs. Perhaps the most intriguing association involves an example of blood feeding
between Culicoides, the most diverse genus of extant blood-feeding of the mos-
quito-like dipteran family Ceratopogonidae (biting midges, no-see-ums, punkies)
and its inferred live host, the dinosaur Corythosaurus. Female ceratopogonids cur-
rently are major vectors of arboviruses, parasitic protozoa, and filarial worms that
cause diseases such as African Horsesickness, Bovine Ephemeral Fever, Bluetongue
Virus, and occasional filariasis, which are associated with acute dermatitis and skin
lesions (Lehane 1991). (Male ceratopogonids are nectar feeders and often are pol-
linators.) Early appearing ceratopogonid lineages occur in Early Cretaceous ambers
and include taxa whose modern representatives, such as Leptoconops and certain
basal species of Forcipomyia and Culicoides feed on reptiles such as turtles, iguanas
and lizards (Wirth and Hubert 1962; Auezova et al. 1990). In particular, these cera-
topogonids attack hosts with vulnerable skin regions lacking scales such as eye
membranes, the anal vent area, and underbelly of individuals, or alternatively in
heavily vascularized regions of the skin that have narrow spaces of exposed skin
between thickened scutes (Auezova et al. 1990; Borkent 1995).
An association has been between certain Late Cretaceous species of Culicoides—
particularly C. canadensis and C. bullus of Campanian Grassy Lake Amber of
Canada and possibly C. filapalpis of Coniacian Taimyr Amber of Russia—and dino-
saurs (Borkent 1995). This interaction is based on the mouthpart structure of fossil
and modern Culicoides (Borkent 1995). Those Culicoides species with a combina-
tion of finely toothed mandibular stylets and coarse, retorsely toothed maxillary
stylets indicate feeding on vertebrates, rather than insects (McIver et al. 1991). In
addition, the number CO2 detecting capitate sensillae on the maxillary palps of
Culicoides species is directly associated with vertebrate host size: those with a
greater number of sensillae (n = 29–74) feed on small hosts such as small birds and
small mammals, whereas those species with fewer sensillae (n = 11–17) feed on
large mammals, with some species possessing an intermediate number of sensillae
(n = 29–36) that feed on birds and mammals of intermediate sizes (Rowley and
Cornford 1972; Braverman and Hulley 1979). An examination of the fossil record
from both the Late Cretaceous of north-central North America northern Russia indi-
cate that mammals and birds were comparatively small and that the only large ver-
tebrate candidates as hosts were large dinosaurs, such as Corythosaurus which
possessed exposed integumental surfaces for blood-feeding ceratopogonids that co-
occurred with Grassy Lake species of C. canadensis and C. bullus (Borkent 1995)
Both of these and the Russian species of Culicoides have several features of mouth-
part structure and anatomy that would strongly indicate blood feeding on vulnerable
integumental areas of large dinosaurs.
12.3.2.6 Other Interactions
Based on scant evidence, there are other types of interactions that likely were extir-
pated at the K-Pg boundary. Most of these associations involve plant pathogens, and
those involving various relationships with large vertebrates, particularly dinosaurs.
The documented fossil record of plant pathogens is almost nonexistent, with the
exception of epiphyllous fungi (Labandeira and Prevec 2014). Mid-Cretaceous
floras such as the Dakota Formation display a significant epiphyllous mycota (DT58
on page 15 of Labandeira et al. 2007), and foliar fungi are known from floras
spanning the K-Pg boundary of the Williston Basin (Labandeira personal
observation), though they have not been characterized other than assignment to
DT58. The other major plant-parasitic pathogen groups of viruses, bacteria, and
nematodes may have instances of tissue damage in the fossil record (Labandeira and
Prevec 2014), but have not been documented formally.
Sediments from approximately the same age as the example of the biting midge
and dinosaur parasitism reveal an association between scarab beetles and dinosaur
dung rich in conifer fragments (Chin and Gill 1996). Evidence of carrion
communities on dinosaur carcasses includes beetle borings in and on bone material
from the Campanian of southern Utah and northwestern Montana, and from the
Maastrichtian of northwestern Madagascar (Rogers 1992; Roberts et al. 2007).
Wasp cocoons associated with decomposing dinosaur eggs were described from
middle Campanian to lower Maastrichtian sediments of northern Patagonia, in
Argentina (Genise and Sarzetti 2011). These Late Cretaceous associations occur
within several million years of the K-Pg boundary likely were extirpated by the
demise of their dinosaur hosts.
12.4 The Marine Perspective of Ecological Disruption

and Its Consequences
Although the end-Cretaceous mass extinction is one of the best documented events
of the Phanerozoic, until recently, there have been few detailed ichnological analy-
ses focusing on this extinction in marine environments (Ekdale and Bromley 1984a;
Savrda 1993; Stinnesbeck et al. 1993, 1996; Keller et al. 1994; Ekdale and
Stinnesbeck 1998). This absence of research is a consequence of difficulties inherent
in examining discrete trace fossils from lower Danian sediments. However, in Spain
and France, the K-Pg boundary transition usually is marked by a 2–3 mm-thick red,
iron-rich, boundary layer at the base of several-centimeter-thick interval of dark
clay. In most cases, material infilling the earliest Danian trace fossil assemblage is
similar in color to that of host Danian strata, precluding any direct, visual, differen-
tiation. Typically, both the latest Maastrichtian and the earliest Danian ichnoassem-
blages are observed below the K-Pg boundary, and contrast with the light color of
Maastrichtian strata. The latest Maastrichtian ichnoassemblage consists of struc-
tures filled with sediments that are only slightly darker than the host material, while
the earliest Danian forms are filled with a dark-colored matrix similar to marly,
lowermost Danian strata that contrast strongly with the light color of the
Maastrichtian host rock. In order to analyze the ichnology of the K-Pg transition in
Spain and France, one recent, fruitful methodology has been applied in four bound-
ary sections. These methods have focused on detailed ichnofabric analysis based on
the study of polished sections and on analyses, including isotopic studies, of mate-
rial infilling various sedimentary trace fossils (Rodríguez-Tovar et al. 2002, 2004,
2006; Rodríguez-Tovar and Uchman 2004a, b, 2006, 2008).
12.4.1 A Selective Impact Favoring the Deposit Feeding

Community
Marine ichnoassemblages recorded in pelagic and hemipelagic, non-turbiditic

facies of the K-Pg boundary transition globally are very similar in composition.
At one distal, continental shelf deposit in Denmark, the Maastrichtian assemblage
consists mostly of Thalassinoides, Zoophycos, and Chondrites, whereas basal
Danian sediments are comprised of Planolites, Thalassinoides, and “small
Chondrites-like forms” that now are recognized as Phycosiphon (Ekdale and
Bromley 1984a). In very proximal, continental shelf deposits from Alabama, Savrda
(1993) observed that Thalassinoides, Ophiomorpha, and Planolites penetrated
estuarine sandy deposits of the lowermost Danian. In shallow neritic settings
examined in Mexico, trace-fossil assemblages principally consisted of Chondrites,
Ophiomorpha, Planolites, and Zoophycos (Ekdale and Stinnesbeck 1998). In sec-
tions studied from the south (Agost and Caravaca) and north (Sopelana) of Spain
and in southwestern France (Bidart), there is correspondence to open, deep-sea
pelagic sedimentation (Fig. 12.5). At these Spanish and French sites, a well-
developed, oldest Danian, endobenthic community is recognized, composed typi-
cally of Chondrites, Zoophycos, Planolites, Thalassinoides, and Alcyonidiopsis
(Rodríguez-Tovar and Uchman 2004a, b; Rodríguez-Tovar et al. 2011) (Fig. 12.6).
Ichnoassemblages at the K-Pg boundary transition principally consist of Chond-
rites, Zoophycos, Planolites, Thalassinoides, Ophiomorpha, and Alcyonidiopsis as
Fig. 12.5 The range of black-filled trace fossils are displayed in stratigraphic sections from Bidart,
southwestern France, and Caravaca, southeastern Spain. The trace-fossil infilling is derived from
the K-Pg boundary layer. The Bidart lithologic column is after Rodríguez-Tovar et al. (2011).
Abbreviation: cl base colonization levels in lowermost Danian dark sediments. The planktic
foraminiferal zonation and stratigraphic correlation of the Caravaca section is based on Arz et al.
(2000) and Arenillas et al. (2004)
the most common ichnotaxa (Figs. 12.5 and 12.6), and are similar to those from
other Late Cretaceous, fine-grained, marly sediments (Ekdale and Bromley 1984b).
These assemblages reveal the dominance of feeding traces that consist of a variety
of behaviors―including domichnia, fodinichnia, pascichnia, and chemich-
nia―and were produced predominantly by deposit feeders obtaining food from
the sediment. This spectrum of feeding ecologies agrees with favorable conditions
for detritus- and deposit-feeding tracemakers, associated with the availability of
abundant food that was established immediately after the impact event (Morrow and
Hasiotis 2007). As mentioned above, deposit-feeding clades were less affected by
the extinction than by groups inhabiting the water column or narrowly linked
Fig. 12.6 Black-filled, lowermost Danian trace fossils are shown from the uppermost Maastrichtian
in stratigraphic sections of Bidart, southwestern France, and Caravaca, southeastern Spain. (a)
Chondrites targionii occurring in a polished section of a horizontal slab, Caravaca section. (b)
Thalassinoides isp. (Th) and Planolites isp. (Pl) occurring in an oblique parting surface, Bidart
section. (c) Thalassinoides isp. (Th) and Planolites isp. (Pl) occurring in a horizontal parting
surfaces, Bidart section. (d) Thalassinoides isp. (Th), Chondrites isp (Ch), and Planolites isp. (Pl)
occurring in a polished section of a vertical slab, Bidart section. (e) A lobe of Zoophycos isp. in a
horizontal parting surface, Caravaca section. (f) Zoophycos isp. in a vertical section of a parting
surface, Caravaca section
trophically to photosynthetic organisms (Arthur and Zachos 1987; Jablonski 2005;

Twichett 2006). However, along K-Pg sites along the Gulf Coastal plain, apparently
the producer of Thalassinoides experienced a significant bout of dwarfism after the
boundary, as burrow diameters underwent a significant decrease in earliest Paleocene
sediments (Wiest et al. 2015).
Ichnological studies of the K-Pg boundary interval from deep sea turbidite
facies are less well known than those from non-turbiditic sediments. In the Uzgruň
section of Moravia, in the Czech Republic, Chondrites intricatus, Ch. targionii,
Ophiomorpha annulata, O. rudis, Palaeophycus tubularis, Planolites isp.,
Phycosiphon incertum, Thalassinoides isp., and Trichichnus isp. occur through the
boundary interval (Uchman et al. 2005). They occur in the sediments underlying
and overlying the boundary, and lack distinct morphological changes. In addition,
the trace-fossil diversity pattern at the ichnogenus level does not display signifi-
cant change through this interval (Uchman 2004, 2007). Interestingly, after a
Cretaceous peak, the number of new graphoglyptid ichnogenera in the Paleocene
decreased considerably, while the contribution of graphoglyptid taxa in ichnoas-
semblages of turbiditic sediments increased in general (Uchman 2003).
Stratigraphic resolution of these changes is poor, making it very difficult to link
these changes precisely to the K-Pg event. However, it is possible that the coinci-
dental the drop of ambient, deep-sea, water temperature during the Paleocene
(Barron and Peterson 1991) may have affected the infaunal farming activity of
graphoglyptid tracemakers.
12.4.2 Minor Disruption in the Macrobenthic Tracemaker

Community
As noted above, in the Spanish and French sections there is a differentiation between
pre-event latest Maastrichtian ichnoassemblages, consisting of trace fossils with
lighter-hued infillings of Chondrites, Zoophycos, and Planolites, versus post-event,
earliest Danian ichnoassemblages of darkly filled structures that contain principally
Chondrites, Zoophycos, Planolites, Thalassinoides, and Alcyonidiopsis (Rodríguez-
Tovar and Uchman 2004a, b; Rodríguez-Tovar et al. 2011). These two ichnoassem-
blages are differentiated by matrix color, nature of the burrow infill material, and
isotopic composition. Minor variations between sections, such as the presence or
abundance of Chondrites and Zoophycos, probably have local importance that can
be related to differences in feeding strategies of the trace-fossil tracemakers related
to food content or site bathymetry.
Similarly, in other K-Pg boundary sections, Thalassinoides, Zoophycos, and
Chondrites are representative of the upper Maastrichtian assemblage, while
Planolites and Thalassinoides are dominant in the lower Danian (Ekdale and
Bromley 1984a; Savrda 1993). Thus, trace-fossil assemblages do not change sig-
nificantly across the K-Pg boundary, appearing in lower Danian ichnoassemblages,
and by comparison are less abundant and diverse than Maastrichtian ichnoassem-
blages. Environmental changes associated with K-Pg boundary phenomena appar-
ently did not have a significant impact on macrobenthic tracemakers, and had a
minimal effect on the marine macrobenthic tracemaker community at the K-Pg
boundary.
12.4.3 An Unfavorable Habitat for Macrobenthic

Colonization?
Historically, the rusty, red boundary layer in various K-Pg boundary sections of
Spain and France has been considered non-bioturbated, presumably revealing unfa-
vorable environmental conditions that were inhospitable for colonization by organ-
isms. Nevertheless, a detailed analysis does show that the K-Pg boundary layer at
the Caravaca section exhibits a highly bioturbated fabric that includes Zoophycos
and Chondrites vertically crossing the boundary layer (Rodríguez-Tovar and
Uchman 2008). This boundary interval also is penetrated horizontally by Chondrites
ramifications. The “unfavorable” conditions displayed by the iron-rich boundary
layer evidently did not impede colonization by tracemakers. This important coloni-
zation event was related to a repertoire of producer behaviors that was not depen-
dent on substrate features. Zoophycos and Chondrites tracemakers constructed
open, or at least partly ventilated, burrows that accommodated various substrate
features, and this was followed by further infaunal colonization of sediment poor in
oxygenated pore waters and food (Rodríguez-Tovar and Uchman 2008).
By way of analogy, a recent environmental disaster occurred at Doñana National
Park in southern Spain that was caused by the failure of a tailings pond adjacent a
pyrite mine at Aznalcóllar, near Sevilla (Rodríguez-Tovar and Martín-Peinado
2009). The sedimentary wedge resulting from the outflow of sedimentary mine
waste from the tailings pond, replete with elevated concentrations of pollutants and
toxic elements, was colonized within 10 years by the ghost ant, Tapinoma nigerrima.
Tapinoma nigerrima is characterized by aggressive life habits and opportunistic
behavior. Notably, the colonized mine-waste substrate was enriched in various
pollutants that included mercury, arsenic, lead, thallium, antimony, and iridium, and
was characterized by locally elevated heavy elemental concentrations. A comparison
of the leaked element abundances from Aznalcóllar with K-Pg boundary sections
revealed that in several cases, such as iridium, the values obtained in the polluted
soil of Aznalcóllar are higher than those recorded for the K-Pg rusty brown boundary
layer (Rodríguez-Tovar and Martín-Peinado 2009). Nests of T. nigerrima occurred
throughout the tailings layers, and an ant-fashioned biofabric was created from
particles within the polluted soil, providing evidence for the irrelevance of substrate
structure in determining the ant-generated biofabric in the polluted substrate. The
particular response of T. nigerrima to the Aznalcóllar disaster is relevant to
interpretation of ichnofaunal colonization of the K-Pg boundary event, at least for
sections from Spain and France. This relevance is based on similarities between
Aznalcóllar soils and the K-Pg boundary layer, such as the presence of strongly
anomalous, life-destroying chemical elements.
A similar example recently has been studied in the contaminated marsh area of
the Tinto River near Huelva, in southwestern Spain, This marsh is characterized by
high soil concentrations of toxic elements, such as copper, zinc, and arsenic
(Rodríguez-Tovar and Martín-Peinado 2014). An ichnological analysis revealed the
presence of biogenic structures produced by the activity of the earthworm Lumbricus
terrestris and the beetle Platystethus. Colonization of this polluted substrate is
possible due to the particular features of the tracemakers: Lumbricus terrestris

shows a great resistance to elevated concentrations of a number of contaminating
elements, whereas Platystethus produces traces that are relatively independent of
substrate features. These patterns indicate that substrate colonization at Huelva
could be comparable to that of Planolites and Thalassinoides tracemakers immedi-
ately after the K-Pg boundary event (Rodríguez-Tovar and Martín-Peinado 2014).
12.4.4 A Relatively Rapid Recovery
Based on observations on microfaunal assemblages, calibrations of the initial recov-

ery of the marine biota associated with the K-Pg boundary were estimated in the
range of thousands of years (kyr). The initial recovery of planktic foraminifera was
estimated at about 230 kyr (Keller and Barrera 1990), and the early pioneer, calcare-
ous nannoflora appeared approximately 25 kyr after the K-Pg mass extinction event
(Lamolda et al. 2005). Approximately ten kyr is proposed for the time involved in
restoration of food webs and restructuring of marine ecosystems, a process occur-
ring after the oceans were repopulated by planktonic species with high turnover
rates. After an initial, comparatively low occurrence of post-event benthic foramin-
ifera, a subsequent and rapid recovery was found, as complex trophic webs reap-
peared approximately seven kyr after the K-Pg boundary. During the early phase of
the recovery interval, the presence of an epifauna tolerant of low oxygen occurred
from 600 to 1200 years after the event (Coccioni and Galeotti 1994).
Two significant observations are important for understanding the response of the
macrobenthic tracemaker community to the K-Pg boundary event and their subse-
quent, comparatively rapid recovery. First is the presence of iron oxide spherules in
Thalassinoides burrow infillings (Rodríguez-Tovar 2005). A second consideration
is physical disturbance resulting from bioturbation of the K-Pg boundary layer
(Rodríguez-Tovar and Uchman 2008). Stereomicroscopic and field-emission SEM
analyses of Thalassinoides at the K-Pg boundary layer from Agost display numer-
ous iron oxide spherules in the infilling material. In addition, the composition, inter-
nal texture, morphology, and size of the infilling were similar to the ichnofabric that
was confined to other sections of the 2–3 mm-thick, rusty, K-Pg boundary layer
(Martínez-Ruiz et al. 1997, 1999). These observations, in conjunction with the
homogeneity of the infilling material and the absence of erosional surfaces capping
Thalassinoides burrows, were interpreted as evidence of rapid colonization by
Thalassinoides tracemakers, occurring almost contemporaneously with the spher-
ule layer deposit (Rodríguez-Tovar 2005) and by a smaller-bodied species revealed
by data from the Gulf of Mexico (Wiest et al. 2015). Subsequently, a detailed analy-
sis of the dark boundary layer from the Caravaca section revealed discrete bioturba-
tion that commenced about 14 mm from the rusty-boundary layer, immediately
above the first laminated interval (Rodríguez-Tovar and Uchman 2008). Recently,
high-resolution geochemical analyses from the K-Pg boundary at the Caravaca sec-
tion support the conclusion that the recovery to pre-impact levels of oxygenation
was almost instantaneous, with absolute values in the order of 102 yr (Sosa-Montes
de Oca et al. 2013).
12.4.5 Iterative and Continuous Colonization after the Mass

Extinction
The absence of a clear color differentiation between burrow infillings of the earliest
Danian ichnoassemblage and the dark, marly, lower Danian host sediments from
the K-Pg boundary sections of Spain and France prevented, at least initially, con-
clusive identification of the colonized stratal horizons and possible assessment of
the recovery’s initiation. In this context, the analysis of carbon isotope composition
of the infill from passively filled burrows became a useful tool (Rodríguez-Tovar
et al. 2002, 2004, 2006). A comparison of the δ13C data from infilled Danian trace
fossils in the uppermost Cretaceous sediments in the Agost section allowed for
identification and assessment of the relative timing of the macrobenthic coloniza-
tion phases. This examination distinguished Cretaceous from Danian trace fossils
based on isotopic composition, and revealed that different isotopic values from the
dark-infilling material could be correlated with those obtained in particular hori-
zons within the Danian marly interval. This analysis provided evidence for different
phases of colonization, deployed in succession across the K-Pg boundary interval
(Rodríguez-Tovar et al. 2002, 2004, 2006).
A subsequent, detailed ichnofabric analysis of the 7–10 cm-thick, dark Danian
boundary layer at the Caravaca section allowed for identification of two bioturbated
horizons, separated by two laminated, unbioturbated layers (Rodríguez-Tovar and
Uchman 2006). The first laminated layer, 14 mm thick, rests just above the rusty
boundary layer. It is overlain by a 26 mm thick, bioturbated horizon, which is
covered by a 36 mm thick layer that exhibits convolute lamination. Above, the
sediment is again bioturbated (Rodríguez-Tovar and Uchman 2008). From the
bioturbated horizons in the dark boundary layer, trace fossils pipe downward
continuously to the uppermost horizons of lighter-hued, Maastrichtian sediments.
These ichnofossils cross-cut the rusty boundary layer, penetrating up to 90 cm
below into Maastrichtian marls. Zoophycos and Chondrites penetrate up to 90 cm
and 35 cm, respectively, below the rusty boundary layer (Rodríguez-Tovar and
Uchman 2006, 2008). These data suggest multiple, post-event colonization events.
12.5 Can Trace-Fossil Records Address Biologic Effects

of the K-Pg Event?
Although several advantages of trace-fossil data over body-fossil data previously

have been discussed (Labandeira 2007), the following eight issues, derived from
studies in this contribution, represent utilitarian, recent approaches toward under-
standing ecological and environmental issues in the deep-time fossil record. These
approaches can be applied to both continental and marine trace-fossil records.
12.5.1 Previously Unapplied Analytic Techniques
Trace-fossil data are as eminently amenable to quantification and analysis by a vari-

ety of statistical techniques, as are body-fossil data. The quantification of abundant
trace-fossil data can be seen as a departure from studies of single, or at most a few
specimens of ichnological taxa in previous studies. As well, carbon isotopic analy-
sis, heavy element analysis, and SEM field emission studies provide detailed docu-
mentation at local to regional scales of organism–environment relationships before,
at, and after the K-Pg boundary. The analytic techniques developed for character-
izing plant–insect interactions of entire floras and for understanding the substrate
relationships of organisms expand the applicability of trace-fossil approaches to
new areas of inquiry.
12.5.2 A Multitude of Data
Plant–insect associational studies of bulk floras or biotas require hundreds to (tens

of) thousands of specimens. Such studies also capture data from a multitudinous
array of specimens, plant morphotypes, insect damage types, localities, stratal lev-
els, and habitats. As a result, data from plant–insect interactions become ideally
suited for examinations of time series originating from bulk-collected floras from
multiple stratal horizons. Such data are concordant with modern ecological tech-
niques for examining trends in space, but importantly, the fossil data uniquely offer
the opportunity for examining temporal patterns. For example, the effects of the
K-Pg event in macroinfaunal habitats of the offshore marine realm in Spain and
France took approximately 102–103 yr, according to recent high-resolution geo-
chemical information (Sosa-Montes de Oca et al. 2013). By contrast, in terrestrial,
angiosperm-dominated communities of North Dakota the recovery time was 107 yr
(Wilf et al. 2006), a difference of approximately 4–5 orders of magnitude.
12.5.3 Unique Ecological Data
The analyses of plant–insect interaction trace fossils are the most successful way of
capturing large datasets of trophic data in the fossil record. Because of the absence
of interpretable insect body-fossil data in most deposits, plant–insect interactional
data can provide trophic data that otherwise would be unavailable. These paleoeco-
logical data also can be used as raw input in other approaches recently used in the
fossil record, such as the construction of site-specific food webs (Wilf 2008; Dunne
et al. 2012), and studies of niche conservatism through time (Solé et al. 2002).
Similarly, unique ecological data such as detailed ichnofabric analysis from pol-
ished microscopic sections can reveal a variety of specific organism–substrate rela-
tionships that are unavailable from more traditional, more macroscopically based
ichnological approaches.
12.5.4 Ability to Test and Generate Hypotheses
Because of typically large datasets, trace-fossil data can provide the type and
amount of data that are available to test hypotheses established by modern plant–
insect interaction theory. Alternatively, trace-fossil data also can generate hypothe-
ses from deep-time trends of plant–insect interactions that are testable using modern
data. Specific examples of fossil plant–insect associational data, such as lag times
involved in post-event occupation of ecospace, increasingly are becoming important
for evaluating hypotheses and concepts that are derived from modern theories.
Likewise, in the shallow marine realm, hypotheses regarding the phases of post-
event organism colonization of sediment and hardrock substrates can provide evi-
dence for or against the geochronologically instantaneous establishment of
burrowing or whether a more prolonged colonization process is involved.
12.5.5 Employing Data from Both the Preserved

and “Nonpreserved” Sedimentary Record
An analysis of the material in passively infilled, earliest Danian burrows supports

other significant types of data that have been gathered from the sedimentary record
for the post-K-Pg recovery event. Collectively, these data provide inferences indi-
cating the prevalent environmental conditions at that time. In some cases, there is
recognition of a laterally extensive horizon of colonization if the original layering in
sediment was preserved. However, if the initial sediment is not preserved, burrows
store the lost sedimentary record, and provide some portion of information. Such
information can be used to characterize different phases of colonization and provide
an evaluation of the relative time to recovery.
In the terrestrial plant–insect associational record, relevant data can be found in
more distant basins. Such basins can provide quantifiable insect damage data that
may be closer to the major event boundary of interest. In both the marine and con-
tinental records, regionally extensive strata should be intensively explored for col-
lection of data that are unavailable at historically more intensively explored but very
local stratigraphic sections.
12.5.6 Data with Enhanced Biostratigraphic Resolution
A consequence of the impact of the K-Pg boundary event on microplankton is the

near-complete absence of these groups in the first few centimeters immediately
above the K-Pg boundary. The K-Pg event strongly affected diatoms, calcareous
nannoplankton, phytoplankton, and planktic foraminifera. Depauperate micro-
planktic biotas impede a high-resolution biostratigraphy, resulting in doubts about
completeness of the sedimentary record or the existence of hiatuses that are

biostratigraphically unresolvable. However, in the Agost section, there is the initial
appearance of Paleogene planktonic foraminifera located a few centimeters above
the K-Pg boundary (Arenillas et al. 2004), calibrated to ~5.7–6.7 kyr later in the
K-Pg boundary. The high-resolution (intrasubzone), stable isotope data that were
obtained from infill material of trace fossils in the Agost section show a close cor-
respondence with those from the upper Maastrichtian and lower Danian sediments.
These data provide evidence for completeness, and the absence of hiatuses within
biostratigraphic resolution (intrasubzone) (Rodríguez-Tovar et al. 2006). On land, a
similar exploration of the thin and locally present FU0 layer in the Williston Basin,
for example, can reveal illuminating patterns of plant–insect interactions at the
K-Pg boundary that could be different from the subjacent latest Maastrichtian and
superjacent earliest Paleocene strata (Labandeira et al. 2002b).
12.5.7 A Major Role in the Debate on Catastrophic vs. Gradual

Extinction
The perennial debate about how catastrophic was the end-Cretaceous mass extinc-
tion, include two, highly differentiated and opposite positions of catastrophic versus
gradual perspectives (e.g., Smit 1990 vs. Keller et al. 1995, respectively).
Catastrophism versus gradualism perspectives still represent an unsolved flash point
regarding the biologic consequences of the end-Cretaceous event. This issue, in
part, can be related to the presence/absence of several species of microfossils,
principally planktic foraminiferans and calcareous nannofossils, immediately below
and above the K-Pg boundary, data that has been perceived to support both positions.
Maastrichtian taxa found in Danian sediments are considered as totally or partially
reworked (e.g., Smit 1990; Pospichal 1994; Henriksson 1996; Molina et al. 1998;
Tantawy 2003; Gallala et al. 2009), or as taxa that survived the disaster, but rapidly
disappear during the earliest Danian (e.g., Perch-Nielsen et al. 1982; Keller 1988;
Keller et al. 1995; Gardin and Monechi 1998; Gardin 2002; Bown 2005).
Within this debate, the sedimentological context of trace fossils is important. For
example, the redistribution of microfossils by tracemakers, even immediately below
the K-Pg boundary, remains a possibility that has been considered minimally and
currently lacks deeper analyses by micropaleontologists and paleoecologists
(Thierstein and Okada 1979; Thierstein 1981; Smit and Romein 1985; Pospichal
and Wise 1990; Pospichal et al. 1990; Henriksson 1996; Pospichal 1996; Romein
et al. 1996; Mai et al. 2003; Bown 2005; Lamolda et al. 2005). Recently, detailed
analyses focusing on calcareous nannofossils from the burrow fillings of Zoophycos,
Thalassinoides, Chondrites, and their surrounding sediments across the K-Pg
boundary transition at the Bidart (Rodríguez-Tovar et al. 2010), and Caravaca
(Kęsdzierski et al. 2011) sections, revealed Danian calcareous nannofossils in
lightly-hued Maastrichtian sediments. In these boundary sedimentary sequences, it
was only the dark infillings of Danian burrows that piped down across the K-Pg
boundary, and conversely, Cretaceous nannoplankton occurred above the boundary
layer that was conveyed up onto the seafloor by earliest Danian tracemakers such as
Thalassinoides through burrow excavation (Rodríguez-Tovar et al. 2010; Kędzierski
et al. 2011).
The response of generalized to specialized insect herbivores to their plant-host
spectrum after the end-Cretaceous event can provide a terrestrial perspective.
Although the terrestrial stratigraphic record is poorer than the marine record at this
time, the considerable delay in the recovery of associational diversity and special-
ization levels to that of the latest Maastrichtian is significant. The presence of lag
times from 104 to 105 orders of magnitude between the rapid response of the marine
realm, versus the much more prolonged terrestrial recovery, may hint at differences
in ecological structure. The ecological recovery potential and flexibility of the shal-
low marine realm evidently is considerably greater than that of the terrestrial realm.
12.5.8 Understanding Ecologic and Evolutionary Response

to Future Environmental Crises
Uncertainties regarding the effect of the K-Pg mass extinction event on evolution-
ary and ecological aspects of the biota can be addressed by analogy to the response
of modern organisms soon after recent disasters. Although a comparison of
paleoenvironmental events with recent examples of environmental crises is rarely
applied, nevertheless it is a useful tool (Kuhnt et al. 2005) to understand biotic
response to dramatic past, recent, and future environmental change. The response
of tracemakers after the K-Pg boundary event as well as those recorded after a
drastic environmental disaster occurring at Doñana National Park in southern Spain
(Rodríguez-Tovar and Martín-Peinado 2009; Martín-Peinado and Rodríguez-Tovar
2010) – and a parallel crisis associated with the Tinto river marsh at Huelva
(Rodríguez-Tovar and Martín-Peinado 2014), in southwestern Spain—indicate that
high concentrations of heavy metals accumulated in soils and sediments can be
dispersed at geologically ephemeral time scales (Rodríguez-Tovar and Martín-
Peinado 2011). These two, recent incidents reveal that a better understanding of
significant paleoenvironmental change, either abrupt or extended in time, can have
profound consequences for the biota, their lag recovery times, and colonization of
previously “uninhabitable” habitats.
Though not discussed in this contribution, related studies relevant to the recovery
of plant–insect interactions have been done in real time in defaunation experiments
of small, mangrove islands in Florida Bay. These studies have recorded the lag
times of recolonization of major plant and insect groups and their associations
following complete island defaunation (Simberloff and Wilson 1969). The pattern
of the re-establishment of major trophic groups (Simberloff 1976), including gener-
alized and specialized associations of a local biota is relevant to issues such as the
return of plant–insect associations after the K-Pg event.
12.6 Conclusions
Based on the results of studies from the terrestrial and marine realm that are detailed
in this contribution, there are several major applications of ichnological data, some
of which cannot be replicated solely by examination of body-fossil data. For a better
understanding of the consequences of major crises in the history of life, such as the
K-Pg ecological crisis, the combination of ichnological data with body-fossil and
physical data is essential, such as analyses of the diversity and abundance of insect
damage on plant hosts in conjunction with various sedimentological and geochemi-
cal studies. Integration of such data provides a realistic assessment of organismic
response that takes into account all of the major, relevant parameters involved.
These features are: (1) the environment in the broadest sense of the term; (2) the
organisms themselves; (3) organismic behavior and organism interactions with one
another and their environment; and (4) and the temporal dimension that includes
event durations and times to recovery. Application of all four aspects of the data
provides a more integrative and complete approach toward understanding environ-
ment–organism interactions during crucial ecologic crises in the fossil and associ-
ated sedimentological record. Moreover, understanding organism response to such
events can be put to practical use in approaching similar catastrophes for the future.
Acknowledgments We extend our gratitude to Gabriela Mángano and Luis Buatois for inviting
us to provide this contribution to the volume. Reviewers Karen Chin and Charles Savrda provided
critical remarks that improved the paper. We thank Finnegan Marsh for producing Figs. 12.1, 12.2,
12.3, and 12.4. Research by Rodríguez-Tovar was carried out with financial support of the Research
Projects CGL2008-03007, and CGL2012-33281 (Ministerio de Economía y Competitividad),
P08-RNM-03715 and the Research Group RNM-178 (Junta de Andalucía). A. Uchman received
additional support from the Jagiellonian University (DS funds). This is contribution 260 of the
Evolution of Terrestrial Ecosystems consortium at the National Museum of Natural History, in
Washington, D.C.
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Chapter 13
The Phanerozoic Four Revolutions
and Evolution of Paleosol Ichnofacies
Jorge F. Genise, Emilio Bedatou, Eduardo S. Bellosi, Laura C. Sarzetti,

M. Victoria Sánchez, and J. Marcelo Krause
13.1 Introduction
Which is the appropriate scale to analyze how paleosol ichnology reflects major
evolutionary events in the history of life? Surely, the scale of the ichnofacies model,
which arises from the most elementary aspect of scientific reasoning: to recognize
recurrent patterns. Repetitive ichnoassemblages are typical of different periods in
the history of life and that they can be grouped in major categories: Seilacherian
ichnofacies. Ichnofacies are recurrent associations of trace fossils in time and space,
and it is their recurrence that grants the essential, broad scale, data for this evolu-
tionary analysis. Mesozoic ichnoassemblages include or are dominated by crayfish
and earthworm trace fossils, whereas insect trace fossils dominate most Cenozoic
assemblages (Genise 2016). To ignore these patterns would result minimally in an
unacceptable loss of scientific information.
Life on Earth has been evolving since its beginning during the Archean.
Accordingly, paleosol ichnofacies, as a reflection of behaviors and ecologic prefer-
ences of soil-inhabiting organisms, should also have evolved. As pointed out by
MacEachern et al. (2007, 2012), the Coprinisphaera Ichnofacies cannot be older
J.F. Genise (*) • E.S. Bellosi • L.C. Sarzetti • M.V. Sánchez

CONICET, División Icnología, Museo Argentino de Ciencias Naturales,
Av. Ángel Gallardo 470, 1405 Buenos Aires, Argentina
E. Bedatou
CONICET, INCITAP, Facultad de Ciencias Exactas y Naturales, Universidad
Nacional de La Pampa, Av. Uruguay 151, 6300 Santa Rosa, La Pampa, Argentina
J.M. Krause
CONICET, Museo Paleontológico Egidio Feruglio, Av. Fontana 140,
9100 Trelew, Chubut, Argentina

DOI 10.1007/978-94-017-9597-5_13
302 J.F. Genise et al.
than the Paleogene when grass-dominated communities appear. What happened,

then, with paleosol ichnofacies prior to the Cenozoic? How did ichnofacies evolve?
It is impossible to think in static ichnoassemblages from the oldest paleosols-
bearing rhizoliths recorded in the Early Devonian to the youngest ones in the
Holocene. Alternatively, tracking ichnofacies evolution, an uncommon approach in
theoretical ichnology, should allow us a better understanding of evolution of life
itself and it would represent a significant contribution of ichnology for its delayed
integration with other paleontological disciplines. The idea of major evolutionary
changes characterizing the history of life has been designed mostly from data and
interpretations provided by paleontological disciplines other than ichnology. Can
ichnology, and particularly paleosol ichnology, with its own major evolutionary
steps, offer a parallel exploration to the history of life?
On the other hand, which is the influence of soil types on the evolution of paleo-
sol ichnofacies? Much of the major events in life history, such as the end-Permian
crisis, the advent of angiosperms during the mid-Cretaceous, or the end-Cretaceous
mass extinction, produced no significant changes in the nature of soils. In contrast,
it is clear that the increasing complexity of terrestrial ecosystems, from early micro-
bial habitats to the development of complex plant communities, triggered the
appearance of the different soil types. Retallack (1992a) proposed that the principal
changes of land environments were deployed in a stepwise mode following the ori-
gin of different plant formations, from microbial earths to grasslands. Is it possible
to use paleosol ichnofacies to test this hypothesis?
In this chapter, we will travel along a two-way road looking at the consequences,
if any, that major evolutionary changes had on paleosol ichnofaunas, and vice versa.
As a result, we expect to unveil the big picture of the evolution of paleosol ichnofa-
cies accompanying major evolutionary changes in the history of life.
13.2 The Rhizolith Revolution: The Paleozoic Shy Beginning

of Paleosol Trace Fossils
The Paleozoic was a very innovative period for history of life (see Chaps. 3–7). It
includes the appearance of the first soils bearing subaerial trace fossils (Tables 13.1
and 13.2; Figs. 13.1 and 13.3). Before that, different paleosol features produced
mostly by chemical and physical weathering have been recognized in Archean and
Proterozoic rocks (Retallack 2001a). These characters would include microbial
borings, soil horizons, peds, illuviation argillans, cryogenic cracks, slickensides,
nodules, and sepic-plasmic clay microfabrics (Retallack 1992b). These paleosols
older than 1 Ga would include Entisols, Inceptisols, Aridisols, Gelisols, Andisols,
Vertisols, and Oxisols, whereas “green clays” would have formed in a reducing
atmosphere older than 2.2 Ga (Retallack 2001a). Microbial earths would have devel-
oped also in terrestrial environments during the Precambrian (Driese et al. 1997).
For various reasons, including the absence of most groups of soil invertebrates
that produce preservable traces, paleosol trace fossils are scarce during the Paleozoic.
Table 13.1 Recorded cases of root and invertebrate trace fossils in paleosols
13
Locality Formation Age Trace fossils References

Yemen, Ma’rib Unnamed Holocene Root traces Pietsch and Kühn (2012)
South Africa, Clanwillam Unnamed Holocene Termite earth mounds Moore and Picker (1991)
South Africa, Western cape Unnamed Holocene Root casts Cramer and Hawkins (2009)
Sultanate of Oman, Wahiba Unnamed Holocene Celliforma Radies et al. (2005)
Sand Sea Coprinisphaera
Rosellichnus
Termite nests
USA, San Miguel Island Unnamed Holocene Rhizoconcretions Stewart and Thorson (1994)
New Zealand, Karikari, Unnamed Quaternary (Holocene and Cladichnus Gregory et al. (2004)
and Aupouri Peninsulas Pleistocene) Planolites
Skolithos
Taenidium
Insect burrows
Root traces
Spain, Canary Islands Unnamed Quaternary (Holocene and Rebuffoichnus guanche Alonso-Zarza and Silva (2002),
Pleistocene) Rebuffoichnus casamiquelai Mikúlaš and Genise (2003), Genise
Tombownichnus parabolicus and Edwards (2003), Alonso-Zarza
Tombownichnus plenus et al. (2008), Meco et al. (2010,
Palmiraichnus castellanosi 2011), La Roche et al. (2014), Genise
“Acridian ootheca” et al. (2013a)
Rhizoliths
Australia Unnamed Quaternary (Holocene and Rebuffoichnus casamiquelai Lea (1925)
Pleistocene)
The Phanerozoic Four Revolutions and Evolution of Paleosol Ichnofacies
Argentina, Quequén Unnamed Late Pleistocene-Holocene Root traces Tonello et al. (2002)
Uruguay Sopas Late Pleistocene Castrichnus incolumis Ubilla (1996), Verde et al. (2007)
Coprinisphaera
Taenidium serpentinum
Meniscate burrows
Rhizoliths
303
(continued)
304

Argentina, Santa Fé Tezanos Pinto Late Pleistocene Coprinisphaera Iriondo and Krohling (1996),
Ant nests Kröhling (1999)
Root traces
Argentina, Buenos Aires Buenos Aires Late Pleistocene Barberichnus bonaerensis Laza (1995, 2006a), Genise et al.
Coprinisphaera (2000)
Termite nests
Argentina, Buenos Aires Luján Late Pleistocene Coprinisphaera Laza (1995, 1997), Genise et al.
and Santa Fé Ant nests (2000)
Namibia Homeb Silt Late Pleistocene Taenidium Smith et al. (1993)
Termitichnus
Pelletal burrows and chambers
Namibia Sossus Sand Late Pleistocene Taenidium Smith and Mason (1998)
(Khommabes Termitichnus
Carbonates) Digitichnus
Argentina, Tucumán Tafí del Valle Late Pleistocene Coprinisphaera Fontaine et al. (1995)
South Africa, Still Bay Waenhuiskrans Middle and Late Pleistocene Rhizoliths Roberts et al. (2008)
Italy, Santo Stefano Island Villa Giuli Pleistocene cf. Rebuffoichnus or cf. Sacchi and Petti (2008)
Fictovichnus
Canada, Ontario Pleistocene sands Pleistocene Rhizoconcretions Kindle (1923)
USA, Sapelo Island Unnamed Pleistocene Taenidium Gregory et al. (2004)
Root traces
New Zealand, Northland Unnamed Pleistocene Ant burrows Gregory and Campbell (2003),
Bee burrows Gregory et al. (2009)
Bee cells
“Phoebichnus look-alike” root
system
Australia Bridgewater Pleistocene Palmiraichnus Zeuner and Manning (1976),
Houston (1987)
J.F. Genise et al.
13
USA, Washington Washtucna Soil Pleistocene Cicada burrows O’Geen and Busacca (2001),
Root traces Blinnikov et al. (2002)
Ecuador, Quito Cangahua Pleistocene Coprinisphaera Sauer (1955), Clapperton and Vera
(1986), Laza (2006b), Sánchez et al
(2013)
Brazil, Osorio Chuí Pleistocene Celliforma Netto et al. (2007)
Coprinisphaera
Krausichnus
?Monesichnus
Palmiraichnus
Taenidium barretti
Termitichnus
Vondrichnus
Rhizoliths
Wasp cocoons
British West Indies, Ironshore Pleistocene Rhizoliths Jones and Ng (1988)
Cayman Brac
Spain, Mallorca Unnamed Pleistocene Rhizoconcretions Calvet Rovira et al. (1975), Klappa
(1980)
Spain, Ibiza Unnamed Pleistocene Rhizoliths Klappa (1980)
Kenya, East Turkana Koobi Fora Pleistocene Root traces Cohen (1982), Mount and Cohen
Root casts (1984)
USA, Louisiana Citronelle Early Pleistocene Tree casts Mossa and Schumacher (1993)
South Africa, Unnamed Early Pleistocene Planolites Miller and Mason (2000)
Namaqualand Skolithos
Taenidium
Termitichnus namibiensis
Vertical rhizoconcretions
Tanzania, Olduvai Gorge Olduvai Early Pleistocene Meniscate burrows Ashley and Driese (2000)
Rhizoliths
Termite and ant traces
305
(continued)
306

Argentina, Entre Ríos and Ensenada Early Pleistocene Coprinisphaera Frenguelli (1938)
Santa Fé
Spain, Mallorca Unamed Pliocene-Early Pleistocene Rebuffoichnus Mas and Ripoll (2010)
India, Punjab Boulder Pliocene-Pleistocene Termitichnus Tandon and Naug (1984)
Meniscate burrows
Argentina, Buenos Aires San Andrés Late Pliocene-Early Barberichnus bonaerensis Laza (1995, 2006a), Cantil et al.
Pleistocene Coprinisphaera (2013)
Kenya and Tanzania Laetoli Late Pliocene Coprinisphaera Sands (1987), Ritchie (1987),
Celliforma Darlington (2005), Krell and
Lazaichnus amplus Schawaller (2011), Darlington (2011)
Wasp cocoons
Termite nests
Burrows
Root traces
Argentina, Buenos Aires Chapadmalal Late Pliocene Coprinisphaera Laza (1995)
Ant nests
Termite nests
Argentina, Buenos Aires Barranca de los Late Pliocene Tacuruichnus farinai Genise (1997)
Lobos
Argentina, Jujuy Maimará Early Pliocene Coprinisphaera Laza (2006b)
Argentina, Salta and Jujuy Piquete Early Pliocene Coprinisphaera Alonso et al. (1982)
United Arab Emirates, Abu Unnamed Early Pliocene Rosellichnus arabicus Genise and Bown (1996)
Dhabi Ant nests
Argentina, Buenos Aires Monte Hermoso Late Miocene-Early Pliocene Coprinisphaera Laza (1986b)
United Arab Emirates, Abu Baynunah Late Miocene Rosellichnus arabicus Bown and Genise (1993)
Dhabi Termite nests
Pakistan Dhok Pathan Late Miocene Coprinisphaera-like traces Retallack (1991a)
J.F. Genise et al.
Argentina, Neuquén, Río Collón-Curá Late Miocene Celliforma Frenguelli (1939), Genise and Bown
13
Negro, Chubut and Santa Coprinisphaera (1996), Laza (1986b), Villafañe et al.
Cruz Lazaichnus fistulosus (2008), Bedatou (2010), Sarzetti et al
Fictovichnus sciuttoi (2014)
Rosellichnus patagonicus
Teisseirei barattinia
Burrows
Argentina, San Luis Paso de Las Carretas Late Miocene Celliforma Pascual and Bondesio (1981)
Coprinisphaera
Argentina, San Juan Las Flores Late Miocene Coprinisphaera Contreras (1996)
Argentina, Catamarca Andalhualá Late Miocene Coprinisphaera Genise et al. (2000)
Argentina, La Pampa and Cerro Azul Late Miocene Attaichnus kuenzelii Laza (1982, 2006b), Genise et al.
Buenos Aires Coprinisphaera (2013b)
Quirogaichnus coniunctnus
Chad Chad Basin Late Miocene Coatonichnus globosus Duringer et al. (2006, 2007)
Microfavichnus alveolatus
Quirogaichnus
Termitichnus schneideri
Vondrichnus planoglobus
Root traces
USA, Oregon Rattlesnake Late Miocene Insect burrows Retallack et al. (2002a)
Root traces
Root systems
Perú, Loreto Pebas Late Miocene Planolites Rebata et al. (2006)
Taenidium
Root traces
Spain, Teruel Unit II Late Miocene Cf. Alonso-Zarza et al. (2012)
Cellifroma Root traces
USA, Kansas Ogallala Middle Miocene-Early Daimoniobarax nephroides Martin and Bennett (1977), Gobetz
Pliocene Daimoniobarax tschinkeli and Martin (2006), Yelinek and Chin
Possible dung beetle burrows (2007), Smith et al. (2011)
307
Root traces
(continued)
308

Kenya Maboko Middle Miocene Root traces Retallack et al. (2002b)
USA, Colorado Pawnee Creek Miocene Beaconites kytosichnus Hembree and Hasiotis (2008)
B. knestosichnus
Parowanichnus perirhizaterion
Rhizoliths
Spain “Gypsum and grey Miocene Celliforma Uchman and Alvaro (2000)
shales” Cellicalichnus aff. habari
Rosellichnus cf. arabicus
Spongeliomorpha
Labyrintichnus terrerensis
Taenidium barretti
Beaconites filiformis
Polykladichnus aragonensis
Kenya Hiwegi (Kibanga Miocene Cellicalichnus habari Thackray (1994), Genise (2000)
Mb.) Root traces
Burrows
Wasp cocoons
Kenya Hiwegi (Fossil Bed Miocene Root traces Retallack et al. (1995)
Mb.) Cocoons
Honduras, Tegucigalpa El Periodista Miocene Palmiraichnus Domínguez-Alonso and Coca-Abia
(Member) (1998)
Argentina, Santa Cruz Pinturas Early Miocene Coprinisphaera Bown et al. (1988), Bown and Laza
Palmiraichnus (1990), Genise and Bown (1994a),
Skolithos Genise and Hazeldine (1998a),
Syntermesichnus fontanai Kramarz and Bellosi (2005),
Taenidium barretti Cosarinsky et al. (2005), Laza
Burrows (2006b), Bedatou et al. (2007,
Earthworm diffuse boxwork 2008b), Genise et al. (2008b),
Rhizoliths Bedatou (2010)
J.F. Genise et al.
Root traces
13
Argentina, Santa Cruz Santa Cruz Early Miocene Celliforma Genise and Bown (1994a), Tauber
Coprinisphaera (1996)
Ant nests
USA, Florida Tampa Early Miocene Celliforma nuda Brown (1935), Genise (2000)
Ethiopia Bakate Early Miocene Termite nests Bown and Genise (1993)
Germany Unnamed Early Miocene Celliforma Sauer and Schremmer (1969)
Argentina, Neuquén Puesto Burgos Late Oligocene-Early Coprinisphaera Pazos (2011)
Miocene Bee cells
USA, Oregon John Day Late Oligocene-Early Pallichnus dakotensis Retallack et al. (2000), Retallack
Miocene Edaphichnium lumbricatum (2004)
Taenidium
Scaphichnium
Termitichnus
Root traces
Spain, Aragón Sariñena and Oligocene–Early Miocene Coprinisphaera Hamer et al. (2007)
Uncastillo Scolicia
Ant nests
Bee cells
Dung beetle burrows
Dung beetle nests and boli
Horizontal crawling traces of
arthropods
Rhizoliths
Spider burrows
Spain, Ebro Solsona Oligocene Taenidium de Gibert and Sáez (2009)
Vertical ornamented burrows

Germany Unnamed Oligocene Celliforma Schütze (1907)
Argentina, Chubut Deseado Oligocene Coprinisphaera Frenguelli (1938), Laza (1986a)
Teisseirei
(continued)
309
310

USA, South Dakota Brule Oligocene Cellicalichnus ficoides Retallack (1983, 1984), Genise
Celliforma (2000)
Pallichnus dakotensis
Root traces
Argentina, Mendoza Rodados lustrosos Oligocene Coprinisphaera Genise et al. (2000)
Egypt, El Fayum Jebel Qatrani Oligocene Fleaglellius pagodus Bown (1982), Genise and Bown
Krausichnus trompitus (1994b)
Macanopsis
Masrichnus issawii
Scaphichnium hamatum
Termitichnus qatrani
Termitichnus simplicidens
Vondrichnus obovatus
Crayfish? burrows
Rhizoliths
Czech Republic Doupov Mountains Early Oligocene Cf. Celliforma and/or Fejfar and Kaiser (2005)
Rebuffoichnus
USA, Colorado White River Early Oligocene Cellicalichnus ficoides Retallack (1983), Hembree and
Edaphichnium Hasiotis (2007)
Fictovichnus isp.
Macanopsis
Pallichnus dakotensis
Parowanichnus
Planolites
Backfilled burrows
Rhizoliths
Wasp cocoons
J.F. Genise et al.
13
USA, Montana Renova Late Eocene-Early Steinichnus Sheldon and Tabor (2009), Sheldon
Oligocene Taenidium and Hamer (2010)
Root traces
Wasp cocoons
USA, Wyoming Bridger Late Eocene Celliforma spirifer Brown (1934), Genise (2000)
Antarctica, Seymour Island La Meseta Late Eocene Coprinisphaera Laza and Reguero (1990), Laza
(2006b)
France Unnamed Late Eocene Celliforma arvernensis Ducreux et al. (1988)
England, Isle of Wight Bembridge Late Eocene Fictovichnus sciuttoi Edwards et al. (1998), Armenteros
Limestone and Daley (1998), Genise et al.
(2007)
Argentina, Salta Lumbrera Middle–Late Eocene Root traces Del Papa (2006)
Uruguay Queguay Middle Eocene–Middle Celliforma germanica Martínez et al. (1997, 2001), Verde
Oligocene C. spirifer and Genise (2007), Alonso-Zarza
C. rosellii et al. (2011)
Fictovichnus gobiensis
Fictovichnus sciuttoi
Rosellichnus isp.
Bee nests
“Agapostemonini cluster”
Rhizoliths
(continued)
311
312

Argentina, Chubut Sarmiento Middle Eocene–Early Chubutolithes gaimanensis Andreis (1972), Laza (1986a), Bown
Miocene Coprinisphaera lazai and Ratcliffe (1988), Genise and
Coprinisphaera murguiai Bown (1990), Bellosi et al. (2001,
Coprinisphaera kraglievichi 2002a, 2010), Mikúlaš and Genise
Coprinisphaera kheprii (2003), Bellosi and Genise (2004),
Coprinisphaera tonnii Genise and Cladera (2004), Genise
Feoichnus challa et al. (2004, 2008b), Krause and
Lazaichnus fistulosus Genise (2004), Cosarinsky et al.
Pallichnus (2005), Laza (2006b), Bedatou et al.
Taenidium (2007, 2008b). Krause et al. (2008,
Teisseirei barattinia 2010), Sánchez and Genise (2009),
Tombownichnus pepei Sánchez et al. (2010b), Bellosi and
Tombownichnus parabolicus González (2010), Bedatou (2010)
Burrows
Earthworm diffuse boxwork
Root marks
Meniscate burrows
Pedotubules
Rhizoliths
Rhizoconcretions
France, Bouxwiller Unnamed Eocene Cf. Fictovichnus gobiensis Kuntz (2010, 2012)
Cf. Fictovichnus sciuttoi
USA, Utah Uinta Eocene Edaphichnium Sandau (2005)
Ant nests
Meniscate burrows
Wasp cocoons
Argentina, Chubut Koluel Kaike Early–Middle Eocene Coprinisphaera Laza (2006b), Krause and Bellosi
Feoichnus challa (2006), Krause et al. (2008)
Root traces
J.F. Genise et al.
13
Uruguay, Nueva Palmira Asencio Early Eocene Celliforma Roselli (1987), Genise and Laza
Coprinisphaera murguiai (1998), Genise et al. (1998),
Coprinisphaera kraglievichi González (1999), Genise and Verde
Corimbatichnus fernandezi (2000), Genise and Zelich (2001),
Elipsoideichnus meyeri Mikúlaš and Genise (2003), Genise
Krausichnus et al. (2004, 2008b), Cosarinsky et al.
Guerraichnus poligibbus (2005), Laza (2006b), Verde and
Lazaichnus fistulosus Genise (2010), Tófalo and Pazos
Monesichnus ameghinoi (2010), Bellosi et al. (2016)
Palmiraichnus castellanosi
Rebuffoichnus barattinia
Teisseirei
Tombownichnus plenus
Tombownichnus parabolicus
Uruguay auroranormae
Uruguay rivasi
Meniscate burrows
Root traces
Termite nests
USA, Wyoming Wasatch Early Eocene Camborygma eumekenomos Zonneveld et al. (2006), Bohacs et al.
Celliforma (2007), Hasiotis and Honey (2000)
Eatonichnus
Planolites
Skolithos
Root traces
(continued)
313
314

Argentina, La Pampa Gran Salitral Early Eocene Celliforma germanica Melchor (2002), Melchor et al.
Celliforma rosellii (2002)
Rosellichnus isp.
Rebuffoichnus
Skolithos linearis
Taenidium barretti
Teisseirei barattinia
Ornamented burrow fillings
Plant traces
Rhizoliths
Argentina, Jujuy Maíz Gordo Late Paleocene–Early Krausichnus trompitus DeCelles et al. (2011)
Eocene Root traces
USA, Wyoming Willwood Late Paleocene–Early Camborygma litonomos Bown and Kraus (1983, 1987),
Eocene Cylindricum Hasiotis et al. (1993a), Kraus and
Edaphichnium lumbricatum Hasiotis (2006), Smith et al. (2008a,
Ichnogyrus b, c, 2009)
Macanopsis
Naktodemasis bowni
Planolites
Scaphichnium hamatum
Steinichnus
Cocoon traces
Meniscate burrows
Pedogenically modified
Camborygma litonomos
Rhizoliths
Root traces
J.F. Genise et al.
13
USA, Utah Claron Paleocene-Eocene Camborygma eumekenomos Bown et al. (1997), Hasiotis and
Camborygma litonomos Bown (1996)
Celliforma
Eatonichnus uthaensis
Eatonichnus claronensis
Fictovichnus
Parowanichnus formicoides
Wasp cocoons
USA, Wyoming Hanna Late Paleocene Camborygma eumekenomos Hasiotis and Honey (1995, 2000)
Root traces
Argentina, Rocas Coloradas Peñas Coloradas Late Paleocene Eatonichnus Genise et al. (2001a, 2008b)
USA, Wyoming Unnamed Upper Late Paleocene Camborygma Hasiotis and Honey (1995)
Paleocene unit Root traces
USA, Wyoming Fort Union Late Paleocene Camborygma symplokonomos Hasiotis and Honey (2000), Kraus
Camborygma eumekenomos and Hasiotis (2006), Adams et al
Camborygma litonomos (2011)
Root traces
Rhizoconcretions
Slovenia Trstelj Late Paleocene Rhizoconcretions Kosir (2004)
USA, North Dakota Fort Union, Sentinel Early Paleocene Undetermined burrows Fastovsky and McSweeney (1991)
Butte Member Root traces
Stumps
USA, Montana Hell Creek Late Cretaceous Root traces Fastovsky et al. (1989)
Brazil, Bauru Basin Adamantina Late Cretaceous Coprinisphaera Souza Carvalho et al. (2009)
USA, Montana Two Medicine Late Cretaceous Celliforma Martin and Varricchio (2011)
Fictovichnus sciuttoi
Skolithos
Teisseirei
Tombownichnus
(continued)
315
316

Romania, Southern Sânpetru Late Cretaceous Burrows Therrien et al. (2009)
Carpathian Mountains Crayfish? burrows
Root traces
Romania, Tuştea Densuş-Ciula Late Cretaceous Burrows Therrien (2005)
Root traces
Romania, Vurpăr Red Continental Late Cretaceous Burrows Therrien (2005)
Strata Root traces
Argentina, Rio Negro Allen Late Cretaceous Fictovichnus sciuttoi Genise and Sarzetti (2011)
Argentina, Chubut Laguna Palacios Late Cretaceous Beaconites Sciutto (1995), Sciutto and Martínez
Cellicalichnus meniscatus (1996), González (1999), Genise
Cellicalichnus chubutensis (2000), Bellosi and Sciutto (2002),
Loloichnus baqueroensis Bellosi et al. (2002b), Genise et al.
Rebuffoichnus casamiquelai (2002a, 2004, 2007, 2008a, b),
Skolithos Mikúlaš and Genise (2003); Genise
Taenidium and Bellosi (2004), Bedatou et al.
Tombownichnus plenus (2005, 2007, 2008a, b), Bedatou
Earthworm diffuse boxwork (2010)
Insect nests
Root traces
Argentina, Chubut Bajo Barreal Late Cretaceous Pallichnus dakotensis Sciutto and Martínez (1996), Bellosi
Fictovichnus sciuttoi et al. (2002b), Genise et al. (2007,
Earthworms Ichnofabrics 2008b), Umazano et al. (2008),
Insect traces Bedatou et al. (2008b), Bellosi et al.
Meniscate burrows (2010), Bedatou (2010)
Rhizoconcretions
Root moulds
J.F. Genise et al.
13
USA, Alaska Prince Creek Late Cretaceous Root traces Spicer and Parrish (1987)
France, Albas Aquitaine Basin Late Cretaceous Root traces Wright et al. (1995)
Gobi Desert, Mongolia Djadokhta Late Cretaceous Fictovichnus gobiensis Johnston et al. (1996), Loope and
Rhizoliths Dingus (1999)
USA, Utah Kaiparowits Late Cretaceous Socialites tumulus Roberts and Tapanila (2006)
Burrows
Root traces
USA, Alabama Tuscaloosa Late Cretaceous Taenidium serpentinum Savrda et al. (2000)
Root traces
USA, Arizona Dakota Late Cretaceous Cellicalichnus dakotensis Elliott and Nations (1998), Genise
(2000)
Uruguay Mercedes Late Cretaceous Celliforma spirifer Veroslavsky and Martinez (1996),
Celliforma germanica Veroslavsky et al. (1997), Tófalo and
Fictovichnus gobiensis Pazos (2010), Alonso-Zarza et al.
Bee nests (2011)
Rhizoliths
Wasp cocoons
Argentina, Santa Cruz Mata Amarilla Late Cretaceous Rhizoliths Varela et al. (2012)
Stumps
South Korea, Jinju Cretaceous Rhizoconcretions Kim et al. (2002)
Kyongsangnamdo
Argentina, Chubut Castillo Early Cretaceous Root traces Sciutto (1981); Bellosi et al. (2002b)
Argentina, Chubut Cerro Barcino Early Cretaceous Rhizolith balls Genise et al. (2010a)
Argentina, Chubut Matasiete Early Cretaceous Palaeophycus Bellosi et al. (2002b), Paredes et al.
Planolites (2003, 2007)

Taenidium
Root traces
(continued)
317
318

Argentina, Santa Cruz Bajo Tigre Early Cretaceous Beaconites Genise (2001), Cladera et al. (2002),
Castrichnus incolumis Bedatou et al. (2006, 2007, 2008a, b,
Dagnichnus titoi 2009), Genise et al. (2008a, b),
Loloichnus baqueroensis Bedatou (2010)
Taenidium
Root traces
Argentina, Santa Cruz Punta del Barco Early Cretaceous Beaconites Cladera et al. (2002), Bedatou et al.
Castrichnus incolumis (2006, 2007, 2008a, b, 2009), Genise
Cellicalichnus meniscatus et al. (2008a, b), Bedatou (2010)
Dagnichnus titoi
Loloichnus baqueroensis
Taenidium
Root traces
Spain, Soria Tierra de Lara Group Late Jurassic-Early Root traces Wright et al. (1995)
Cretaceous
Argentina, Santa Cruz Bajo Grande Late Jurassic Beaconites Cladera et al. (2002), Bedatou et al.
Loloichnus baqueroensis (2007, 2008a, b, 2009), Bedatou
Taenidium (2010)
Root traces
J.F. Genise et al.
13
USA, Colorado Morrison Late Jurassic cf. Ancorichnus isp. Hasiotis (1993, 1999, 2004), Hasiotis
Camborygma litonomos and Demko (1996, 1998), Hasiotis
Camborygma eumekenomos and Kirkland (1997), Hasiotis et al.
Camborygma airioklados (1998, 2004), Fiorillo (1999), Demko
Celliforma et al. (2004), Parrish et al. (2004),
Coprinisphaera Bromley et al. (2007)
cf. Cylindrichum
cf. Rosellichnus isp.
Naktodemasis bowni
Scoyenia isp.
Steinichnus
“Ant nests”
“Bee nests”
“Beetle burrows”
Cocoons
Crayfish burrows
Horizontal striated burrow
Plant and root traces
Quasi-vertical striated burrow
Rhizoliths
“Termite nests”
Vertical burrows
Vertical striated burrows
Iran, Binalud Mountains Aghounj Early-Middle Jurassic Cellicalichnus antiquus Fürsich et al. (2010)
South Africa Elliot/Clarens Early Jurassic Taenidium Smith and Kitching (1997), Bordy
“Termite nests” (2008), Bordy et al. (2004, 2005,
Root traces 2009), Genise et al. (2005)

(continued)
319
320

Poland, Holy Cross Unnamed Early Jurassic Planolites Pieńkowski and Niedźwiezki (2008)
Mountains Scoyenia
Arthropod burrows
Chambered insect nests
Earthworm burrows
Radial chambers
Canada McCoy Brook Early Jurassic Planolites Tanner (1996)
Burrows
Root traces
USA East Berlin Early Jurassic Palaeophycus striatus Gierlowski-Kordesch (1991)
Poland Zajage Early Jurassic Scoyenia Gierlinski et al (2004)
Spongeliomorpha
Root traces
USA, Arizona Owl Rock Late Triassic Crayfish burrows Tanner (2000)
Root traces
USA, Arizona Chinle Late Triassic Archeoentomichnus Hasiotis and Mitchell (1993),
metapolypholeos Hasiotis (1993), Hasiotis et al.
Camborygma symplokonomos (1993b, 2004), Hasiotis and Dubiel
Camborygma eumekenomos (1993a, b, 1994, 1995), Kowaleski
Camborygma litonomos et al. (1998), Therrien and Fastovsky
Camborygma araioklados (2000), Cleveland et al. (2008),
Taenidium Lucas et al. (2010), Tanner and Lucas
Taenidium serpentinum (2012)
Skolithos
Naktodemasis bowni
“Bee cells”
“Beetle burrows”
Inclined burrows
J-shaped burrows
Rhizoconcretions
J.F. Genise et al.
Root traces
13
Argentina, La Rioja Ischigualasto Late Triassic Taenidium Melchor et al. (2001), Genise et al.
Skolithos (2001b, 2004)
Rhizoliths
Antarctica Lashly Middle Triassic Fossil roots Retallack and Alonso-Zarza (1998)
Insect? burrows
Antarctica, Kitching Ridge Fremeouw Early Triassic Burrows Miller and Collinson (1994), Miller
Crustacean burrows and Smail (1996), Retallack et al.
Root traces (1996a), Babcock et al. (1998),
Miller et al (2001)
Australia, New South Narrabeen Group Early Triassic Skolithos Retallack (1976, 1997a)
Wales Crayfish burrows
Earthworm burrows
Insect burrows
Root traces
South Africa Katberg Late Permian–Early Triassic Katbergia carltonichnus Retallack et al. (2003), Gastaldo and
Macanopsis Rolerson (2008)
Scoyenia
Skolithos
Root traces
China, Xinjiang Uyghur Turpan Basin Late Permian–Early Triassic Root traces Thomas et al. (2011)
Rhizoliths
Russia, Arkhangelsk Salarevo Late Permian Root traces Yakimenko et al. (2000)
Oblast
Russia, Tetyushi Urzhumskii, Late Permian Root traces Inozemtsev et al. (2011)
Severodvinskii, and
Vyatskii horizons
Antarctica, Transantarctic Buckley Late Permian Root traces Retallack et al. (1996a)
Mountains
(continued)
321
322

USA, New Mexico Abo Late Permian Scoyenia Kessler et al. (2001), Mack et al.
Adhesive meniscate burrows (2003)
Rhizoconcretions
Antarctica, Victoria Land Beacon Supergroup Permian-Triassic Skolithos Retallack et al. (1995, 1997)
Root traces
Antarctica, Prydz Bay Unnamed Permian-Triassic Root traces Turner (1993)
USA, Kansas Speiser Shale Early Permian Rhizoliths Hembree et al. (2005)
China, Ghizou Liangshan Early Permian Root traces Wang et al. (2013)
USA, Nebraska Lansing-Kansas City Late Carboniferous Root traces Prather (1985)
Groups
Russia Moscow and Mezen Late Carboniferous Root traces Kabanov et al. (2010)
Continental-Platform
Basins
USA, Pennsylvania Pottsville Late Carboniferous Root traces Gill and Yemane (1996),Wnuk and
Pfefferkorn (1987)
USA, Missouri Fort Scott Late Carboniferous Root mats Retallack and Germán-Heins (1994)
Root traces
USA, Ohio Glenshaw and Late Carboniferous Millipede burrows Hembree (2009), Hembree and
Casselman Burrows with chambers Nadon (2011)
Root traces
Australia, New South Seaham Middle Carboniferous Skolithos Retallack (1999a)
wales Root traces
England, Bristol Clifton Down Early Carboniferous Root traces Vanstone (1991)
Mudstone
USA, New York Walton Late Devonian Burrows Retallack (1985)
Root traces
J.F. Genise et al.
13
USA, Pennsylvania and Catskill Middle–Late Devonian Plant and root traces Driese et al. (1997)
New York
USA, New York Oneonta Middle Devonian Meniscate burrows Dunagan and Driese (1999)
Rhizoliths
UK, Wales Old Red Sandstone Middle Devonian Beaconites antarcticus Morrissey and Braddy (2004),
Taenidium barretti Morrissey et al. (2012), Hillier et al.
Root traces (2008)
Antarctica, Victoria Land Aztec Siltstone Middle Devonian Rhizoconcretions Retallack (1997b)
Canada, Quebec Malbaie Middle Devonian Plant and root traces Driese and Mora (2001)
USA, Gilboa Unnamed Middle Devonian Plant and root traces Driese and Mora (2001)
Norway Wood Bay Early Devonian Skolithos helicoidalis Blomeier et al. (2003), Volohonsky
(Spitsbergen Is.) Burrows et al. (2008)
Canada, Gaspé Bay Battery Point Early Devonian Plant and root traces Driese and Mora (2001), Elick et al.
(1998)
Ichnotaxonomy and affinities are included as published, regardless of its reliability. In a few cases, ichnotaxonomy was updated according to recent revisions.
Invertebrate cases in the table intended to be all recorded, whereas in the case of root traces is not an exhaustive list
323
Table 13.2 Paleosol ichnofacies showing trace fossil composition, typical sedimentary environments, and associated plant formations
324
Dominant trace
Ichnofacies Description fossils Sedimentary environments Vegetation
Scoyenia Dominated by meniscate structures (feeding Scoyenia, Taenidium, Continental low energy deposits Little or no vegetation
and locomotion) both trackways and Beaconites, with sediments ranging from sandy cover. Semi-aquatic
continuous trails. There is a mix of Diplichnites, to argillaceous. Both slightly vegetation may be
invertebrate, vertebrate, and plant traces. Diplopodichnus, submerged sediments with present.
Invertebrates are mainly detritus-feeders, Spongeliomorpha, periodical exposition to air and
deposit-feeders, or predators. Vertebrates are Skolithos, subaerial sediments periodically
predators or herbivorous. Vertebrate traces in Fuersichnus, submerged can be represented.
this ichnofacies are mainly mammalian and vertebrate tracks. Typically transitions between fluvial
bird tracks. Ichnodiversity of invertebrate and lacustrine environments,
traces is usually low, but individual traces can including floodplains, ponds, lake
be abundant. Vertebrate footprints may be margins, ephemeral lakes, and
diverse and abundant. humid interdunes.
Coprinisphaera Dominant breeding structures of beetles and Coprinisphaera, Paleosols associated to ecosystems Open herbaceous
bees, but traces of other insects such as ants, Celliforma, of open herbaceous communities, communities. Mostly
termites, cicadas, and moths may be present. Teisseirei, from dry and cold to humid and grass-dominated
Meniscate tubes, mammal caves, and Monesichnus, warm climate. habitats.
rhizoliths can be present. Most common Uruguay, Paleosols formed in different
vertebrate structures are rodent burrows. Palmiraichnus, sedimentary environments including
Moderate to relatively high ichnodiversity Feoichnus, alluvial plains, desiccated
and abundance Tombownichnus, floodplains, abandoned fluvial bars,
Lazaichnus, crevasse splays, levees, and
rhizoliths, vertebrate vegetated aeolian deposits.
burrows
Celliforma Dominance of chambers and chambered trace Celliforma, Calcretes formed under semiarid Scrubs and woodlands
fossils assigned to bees and wasps. Other Rebuffoichnus, (and seasonal) to arid climate. from arid to semiarid
associated components are rhizoliths, Fictovichnus, Palustrine carbonates. environments.
vertebrate coprolites, and large vertebrate Pallichnus, Palustrine vegetation or
burrows. Moderate ichnodiversity, high rhizoliths. bare soils due to
abundance. frequent flooding.
J.F. Genise et al.
13
Termitichnus Dominance of chambers and chambered trace Termitichnus, Paleosols of closed forests Closed forests of warm
fossils attributed to termites. Other Krausichnus, developed under warm and humid and humid climates.
components are J-shaped burrows, rhizoliths, Vondrichnus, conditions. Moderately developed
large vertebrate burrows, and pelleted burrow Fleaglellius, paleosols on channel-belt deposits.
fillings. Low to moderately ichnodiversity rhizoliths, large
and abundance. vertebrate burrows.
Camborygma Dominated by ichnogenera attributed to Camborygma, Paleosols with evidence for a high Local marshes, bogs,
crayfishes and earthworms. The former are Loloichnus, and fluctuating water table, mostly swamps, or wetlands as
vertical to subvertical burrows, commonly Dagnichnus, under warm climates. Paleosols can a subset of forest,
Y-branched, in some cases connected by Cellicalichnus be formed on different deposits scrub, and open
horizontal burrows, with or without meniscatus, (channel, floodplain, overbank, herbaceous
chambers. Rarely breeding structures are Edaphichnium, levee-crevasse splay, gravity flows, communities
present. Earthworm trace fossils are diffuse diffuse boxworks, loessic, ponds, etc.) and on almost
boxworks, in some cases meniscate, or rhizoliths. any lithology.
connected to chambers. Cross-cuttings are
very common. Low ichnodiversity, high
abundance.
Rhizoliths Associations composed almost exclusively of Root traces, Different types of paleosols formed Diverse. Not related to
plant traces, including stumps and any kind of rhizoliths, on any kind of sedimentary deposit. any particular plant
rhizoliths. paleorhizospheres, Indicative of subaerial exposure. formation
Usually low ichnodiversity and abundance stump casts.
commonly high.
Modified from Genise et al. (2000), Buatois and Mángano (2007, 2011) and Melchor et al. (2012)
325
Fig. 13.1 Stratigraphic ranges of crustacean, earthworm, plant, and undetermined trace fossils in
paleosols. Figure starts in the Devonian when the first rizholiths are recognized in paleosols. Only
reliable attributions to producers in each group are included, whereas trace fossils with nonreliable
attributions are included in the group of trace fossils with undetermined affinities. Numbers in
unnamed trace fossils with reliable attributions indicate consulted references: (1) Bown (1982),
Fiorillo (1999), Hasiotis (1993, 1999, 2004), Hasiotis and Demko (1996, 1998), Hasiotis and
Kirkland (1997), Retallack (1997a), Tanner (2000), Therrien et al. (2009); (2) Pieńkowski and
Niedźwiezki (2008); (3) Bedatou et al. (2005, 2006, 2007, 2008a, b), Bedatou (2010); (4) Pieńkowski
and Niedźwiezki (2008), Retallack (1976, 1997a); (5) Smith et al. (1993)
13 The Phanerozoic Four Revolutions and Evolution of Paleosol Ichnofacies 327
A possible oldest evidence of subaerial trace fossils are drab haloes attributed to
filaments of biological soil crusts produced by microorganisms and lichens from the
Early Cambrian Flinders Ranges, Australia (Retallack 2008). The Cambrian-
Ordovician Grindstone Range Sandstone of Australia shows trackways and resting
traces of arthropods (Diplichnites, Selenichnites), which are interpreted as being
produced on the surface of thin and weakly developed paleosols (Retallack 2009).
Since Middle–Late Ordovician, soils included liverwort plants, representing low-
biomass, high soil-respiration ecosystems (Retallack 1997b). Examples of these
conditions would be the red, well-drained, weakly calcareous paleosol of Nova
Scotia, which displays small reduction haloes assigned to mold colonies (Boucot
et al. 1974), and the Aridisol of the Late Ordovician Juniata Formation from
Pennsylvania (Retallack and Feakes 1987; Feakes and Retallack 1988; Retallack
1993, 2001b). However, this evidence was recently criticized based on the deposi-
tional environment, evidence for liverworts, and the nature of S. beerboweri and its
producer, among other issues (Davies et al. 2010) (see Chap. 5).
Undisputed subaerial meniscate trace fossils are younger. They come from the
upper Silurian-Middle Devonian Old Red Sandstone of Wales (Morrissey and
Braddy 2004; Hillier et al. 2008; Morrissey et al. 2012), the Middle Devonian
Oneonta Formation of the USA (Dunagan and Driese 1999), and the Late Permian
Abo Formation of the USA (Mack et al. 2003). Also recorded are Skolithos from
the Early Devonian Wood Bay Formation of Norway (Volohonsky et al. 2008) and
the Middle Carboniferous Seaham Formation of Australia (Retallack 1999a).
Absence of true assemblages and the paucity of data hamper a more definite inclu-
sion of these assemblages in the ichnofacies model. However, they would be con-
sidered herein tentatively as the oldest examples of the Scoyenia Ichnofacies.
Usually this ichnofacies, indicative of alternative subaerial and subaquatic condi-
tions (Fig. 13.4a), is not recognized as a paleosol one. However, during subaerial
exposure the substrate may develop a temporal plant cover producing roots and
traces of soil organisms (Genise et al. 2010b; Melchor et al. 2012). Its mention
herein responds to its record as the only Paleozoic one indicative of subaerial expo-
sure bearing invertebrate trace fossils. However, for practical purposes, the
Scoyenia Ichnofacies will not be treated further in this chapter (neither included in
Table 13.1), as it has a very different background and approach from the other
ichnofacies that occur in post-Paleozoic paleosols (see Chaps. 5, 6 and 12).
Apart from the 9 undisputed cases of Paleozoic paleosols bearing invertebrate
trace fossils, other 17 examples show different types of rhizoliths as exclusive
trace fossils (Table 13.1, Fig. 13.1). The first assemblage composed only of rhizo-
liths is Early Devonian (Driese and Mora 2001), in correspondence with the first
plant origination event, marked by the advent of lycophytes and sphenophytes,
and the first large adaptive radiation of plants (Cascales et al. 2010). This assem-
blage is recognized herein as the first ichnologic revolution in paleosols (Fig. 13.3).
These exclusive rhizolith assemblages, assuming the diversity of different types of
rhizoliths is underestimated, are considered herein for practical purposes as cases of
a Rhizolith Ichnofacies, a proposal similar to that of Melchor et al. (2012). Its name
refers to the initial appearance of rhizoliths, the “Cinderella” of paleosol trace fossils,
considering that root traces are largely understudied and unnamed in comparison
with animal traces. After future and badly needed research on different root sys-
tems and rhizolith branching patterns, the paleoenvironmental meaning of this
ichnofacies could be extended greatly. For instance, rhizoliths and root systems
constitute indicators of subaerial exposure or vegetation type, but also of drainage
conditions in soils and paleosols (Sarjeant 1975; Jenik 1978; Retallack 1988;
2001a). The study of iron oxide-depleted zones around root trace fossils also
assists to define several specific categories of drainage conditions in paleosols
(Kraus and Hasiotis 2006). In addition, further analyses on the reasons for the
absence of associated invertebrate trace fossils in these assemblages will also
increase its value. There are about 38 cases of the Rhizolith Ichnofacies occurring
along different intervals of the Phanerozoic, even though Table 13.1 is not an
exhaustive compendium of all cases recorded from the literature.
Late Silurian and Devonian soils vegetated by the first herbaceous and rhizomatous
plants (xeromorphic rhyniophytes), called “brakeland” formations, were repre-
sented by stronger developed, well-drained paleosols showing complicated biotur-
bation patterns (Retallack 1992a, 2001a). By the Early Devonian, root systems of
primitive vascular plants were large and stout, reaching 1 m deep (Elick et al. 1998).
Rhizome trace fossils in these Entisols consist of dichotomous and fibrous casts and
molds presumably produced by trimerophytes. In addition, expansion of swampy
vegetation in permanently waterlogged terrains gave way to peaty soils or Histosols
(Rice et al. 1995). An old Inceptisol showing large root trace fossils, leaf litter, and
spodic attributes was recognized in the Late Devonian of the USA (Retallack 1985).
The advent of forested soils represented a change in weathering of soil minerals,
and also a renewed step in stability and reduced erosion of landscapes due to deeper
penetration of tree roots. Decaying vegetation of these more complex ecosystems
probably favored the recycling of nutrients, thus originating new subsurface soil
horizons that resulted in the illuviated clay accumulations of Alfisols and Ultisols,
or in the iron and organic matter concentrations of Spodosols (Retallack 2001a).
The most ancient forest ecosystems are recorded in Middle Devonian reddish
Alfisols of Antarctica. These well-drained paleosols bear calcareous rhizoconcre-
tions, ferruginized concretions, and deep and robust root trace fossils surrounded by
large drab-haloes, attributed to highly seasonal, warm subhumid environments
(Retallack 1997b). Diversity in vegetation communities since the Middle Paleozoic
increased weathering rates, which consequently reduced the stability of minerals
through the exportation of ions from soil waters and acidizing the rhizosphere
(Knoll and James 1987). Root structures diversified anatomically in the Mississipian
with the expansion of various forest types (Pfefferkorn and Fuchs 1991). Gill and
Yemane (1996) described an ancient Ultisol from the lower Pennsylvanian of the
USA, containing deep carbonaceous root trace fossils. A more ancient sandy soil
enriched in iron and organic matter (Spodosol) exhibits stout woody root trace fossils
probably of conifers (Vanstone 1991). Pennsylvanian calcic Vertisols, Inceptisols,
and Alfisols formed in alluvial floodplains of the Appalachian basin bear different
root trace fossils (rhizohaloes, rhizocretions, rhizotubules) that vary in depth, repre-
senting fluctuating water tables (Hembree and Nadon 2011). Tropical rain forests
dominated by large-leaved seed ferns (pteridosperms), are recorded since the
Pennsylvanian (USA), and are represented by kaolinitic Oxisols showing large root
trace fossils (Retallack and Germán-Heins 1994).
In sum, the most important soil types were already developed by the Devonian–
Carboniferous, as well as most types of plant formations from different environ-
ments, from arid herbaceous communities to swampy or forested habitats developed
under humid climates. Thus, the paucity of data on paleosol ichnofaunas should
respond to other causes, such as the still relatively scarce record of Paleozoic
paleosols or to the scarcity of soil inhabiting organisms capable of leaving preserv-
able traces. The latter will be a recurrent issue along this chapter. Most of the rec-
ognizable tracemakers of Mesozoic and Cenozoic ichnoassemblages, such as
earthworms and crayfishes, have no body fossil record for the Paleozoic, whereas
the first holometabolous insects appear at the Mississippian–Pennsylvanian transi-
tion (Nel et al. 2007; Béthoux 2008; Wiegmann et al. 2009; Labandeira 2011). In
turn, most Paleozoic invertebrate trace fossils in paleosols have been attributed to
millipedes (Retallack 1999a, 2001b; Morrissey and Braddy 2004; Hembree 2009).
Voigt (2007) analyzed the possible occurrence of insect trace fossils in Permo-
Carboniferous basins of North America and Europe.
The Paleozoic records the appearance of the first paleosols bearing subaerial trace
fossils. Paleozoic assemblages are composed only of rhizoliths (the Rhizolith
Ichnofacies) or in some cases dominated by invertebrate burrows and trackways (the
Scoyenia Ichnofacies) (Table 13.1). The largest floral extinction, occurred at the end
of the Carboniferous (Cascales et al. 2010) is not reflected in changes in paleosols or
trace fossil assemblages. This stasis is only interrupted after the end-Permian mass
extinction event. The assemblages containing only rhizoliths, which dominate the
Paleozoic, are grouped in the Rhizolith Ichnofacies that is recorded extensively from
the Devonian along the rest of the Phanerozoic (Table 13.1, Figs. 13.1, and 13.3).
The Rhizolith Ichnofacies is a practical concept, to be explored considering three
points: (1) the recurrency of paleosols showing only root traces, (2) the dominance of
these assemblages in Paleozoic paleosols, and (3) the potential paleoenvironmental
significance of different rhizolith morphologies.
13.3 The Camborygma Revolution: The Triassic Appearance

of Crayfish and Earthworm Trace Fossils
The Late Permian catastrophic loss of plant biodiversity (Ward et al. 2000;
Michaelson 2002; Arche and López-Gómez 2005) had an uncommonly long period
of recovery in the Triassic. The modification in terrestrial ecosystems was represented
by a global change in vegetation corresponding to the appearance and expansion of
seed plants and the culmination of widespread coal accumulation (Faure et al. 1995;
Retallack et al. 1996b).
In the Southern Hemisphere, this biotic crisis on land is documented in few
paleosols from high latitudes that cross the Paleozoic-Mesozoic boundary (Smith
1995; Retallack 1999b; Retallack and Krull 1999). According to Retallack
(2001a), Late Permian–Early Triassic paleosols from Antarctica, South Africa,

and Australia record abrupt environmental (e.g., acidification, biological produc-
tivity) and paleontological changes, also reflected in the replacement of paleosol
types (e.g., Histosols to ferruginized paleosols). In several localities of Antarctica,
Late Permian carbonaceous root traces (Vertebraria) were replaced by silica or
clayey infilled roots (Retallack et al. 2005). Mid-latitude paleosols from China
show similar climate controlled, long-term changes, from gleyed, organic-matter
rich paleosols indicative of humid conditions, to paleosols with calcic and gypsic
concentrations formed in unstable, semiarid environments, through the Permian-
Triassic boundary (Thomas et al. 2011). Rhizoliths, root haloes, and root moulds
were the only trace fossils observed in these paleosols.
Advanced cone-bearing and seed-producing gymnosperms, along with free-sporing
lycopsids and ferns, became the dominant groups in early Mesozoic floras (Niklas et al.
1985; Visscher et al. 2004; Retallack et al. 2011). The major radiation of conifer fami-
lies (Cephalotaxaceae, Pinnaceae, Taxaceae, Araucariaceae, Podocarpaceae, and
Cheirolepidiaceae) occurred during the Triassic in a global scenario of increasing tem-
perature and seasonality (Archibold 1995; Willis and McElwain 2002). By the early
Jurassic, the expansion of Cycadales, Bennettitales, and Ginkgoales resulted in a new
floristic change at global scale. Eighty percent of plant species were Gymnosperms by
the Middle Jurassic (Hallan 1994; Brenchley and Harper 1998).
In spite of the mentioned large-scale vegetational changes, new or particular types
of paleosols were not observed in Mesozoic strata. The flora preserved in the Upper
Jurassic Morrison Formation illustrates an environment that supported giant sauro-
pods under warm, semiarid to subhumid, seasonal conditions. Floodplains had an
herbaceous groundcover mixed with low-growing woody shrubs (ferns, seed ferns,
ginkgos, horsetails) adapted to severe droughts (Parrish et al. 2004). Watercourses
and lake margins show riparian open-forests of conifers with an herbaceous and
shrubby understory (Turner and Peterson 2004; Engelmann et al. 2004). Calcic
Vertisols, Aridisols, and Alfisols developed in such scenario with fluctuating water
tables (Retallack 1997c; Demko et al. 2004).
The Early Triassic shows the appearance of the first assemblages of paleosol
trace fossils that display a moderate diversity of ichnotaxa and trace makers
(Table 13.1; Figs. 13.1 and 13.3). Crayfish, earthworm, and other invertebrate bur-
rows, probably some of them produced by insects made their appearance in the
geologic record. Nevertheless, the attribution of Early Triassic trace fossils to par-
ticular groups of animals was mostly tentative or weakly supported in the different
study cases. Retallack (1976, 1997a) mentioned crayfish, insect, and earthworm
burrows from the Narrabeen Group of Australia. Among them, earthworm traces,
containing fecal pellets, are the most reliable identified ones (Retallack 1976).
The body-fossil record of earthworms, as of other soft-bodied organisms, is fragmentary
and probably very incomplete to be used as control for the trace-fossil record. Some
tentative body fossils of oligochaetes were recorded earlier during the Middle
Ordovician (Conway Morris et al. 1982). However, these identifications are uncer-
tain and have been disputed (Humphreys 2003). Unquestionable evidence was pre-
sented by Pierce et al. (1990), who described a fossil earthworm embryo
(Oligochaeta: Lumbricidae) preserved with part of its cocoon from beneath a

Holocene midden at Potterne, UK. The earliest evidence is not body fossils but
rather clitellate cocoons, parataxonomically named Burejospermum, Dictyothylakos,
and Pilothylakos, attributed either to Hirudinea or Oligochaeta. These cocoons are
diverse and were recorded worldwide from the Late Triassic to the Neogene (Manum
et al. 1991; Jansson et al. 2008; Tosolini and Pole 2010). The earliest specimens
were described from the Upper Triassic of Greenland (Harris and Rest 1966) and
Sweden (Manum et al. 1991). Other records were documented in the Jurassic of
Norway (Manum et al. 1991), England (Harris 1961), and Australia (Jansson et al.
2008). Cretaceous cocoons were described from Greenland (Manum et al. 1991),
Germany (Manum et al. 1991), Australia, and New Zealand (Tosolini and Pole
2010). Younger material comes from Holocene (post-glacial) deposits from Ontario,
Canada (Schwert 1979).
The body-fossil record of crayfishes is more complete. Miller et al. (2001) dis-
cussed the difficulties in separating burrows of crayfish from those of small tetrapods
in the Lower Triassic Fremeuow Formation of Antarctica. By contrast, Retallack
et al. (2003) and Gastaldo and Rolerson (2008) attributed very tentatively, Macanopsis
and Katbergia respectively, from the upper Permian–Lower Triassic interval of the
Karoo Basin from South Africa, to crustaceans. The oldest body fossil of Parastacidae
is Palaeochinastacus australianus (Martin et al. 2008), recorded from the Lower
Cretaceous of the Otway Group in Australia. Other body fossil records from the
Southern Hemisphere are younger (Sokol 1987; Aguirre-Urreta 1992; Feldmann and
Pole 1994). In a recent molecular phylogeny of the Parastacidae calibrated by body
fossils, Toon et al. (2010) estimated that the Parastacidae originated around 185 Ma
during the Early Jurassic. The oldest Astacidae comes from the Lower Cretaceous of
Spain (Garassino 1997). The oldest body-fossil record of possible Cambaridae
comes from the Upper Triassic Chinle Formation of USA (Hasiotis and Mitchell
1993; Crandall et al. 2000). Although this particular attribution has been disputed
(Rode and Babcock 2003), well documented, reliable crayfish trace fossils included
in the ichnogenus Camborygma, are recorded from this formation (Hasiotis and
Mitchell 1993; Hasiotis et al. 1993b). These trace fossils undoubtedly share with
modern crayfish burrows the typical Y branching, longitudinal connecting tunnels,
chambers, wall surface texture, and dependence on the water table (Hasiotis and
Mitchell 1993; Hasiotis et al. 1993b).
In contrast, the attribution of Late Triassic and Late Jurassic trace fossils from
the Chinle and Morrison formations of the USA (Hasiotis 2000, 2003, 2004) to
particular groups of insects, such as bees, dung beetles, ants, and termites, among
others, and of Early Jurassic trace fossils from the Elliot and Clarens formations of
South Africa to termites (Bordy et al. 2004), has been disputed and mostly unac-
cepted (Engel 2001; Genise et al. 2004, 2005; Grimaldi and Engel 2005; Bromley
et al. 2007; Alonso-Zarza et al. 2008; Lucas et al. 2010; Tapanila and Roberts 2012).
Undoubtedly, insects would have been conspicuous inhabitants of the Triassic and
Jurassic soils and probably many of the simple vertical, inclined, J-shaped, menis-
cate, or chambered burrows of the Chinle or Morrison Formations belong to them.
However, it is hard to unequivocally attribute any trace fossil in a paleosol from
these units to a particular group of insects because most of these simple trace fossils
lack diagnostic characters. The Chinle and Morrison formations are exceptional in
that preserve diverse ichnofaunas, which would contribute significantly to our
understanding of ichnofacies evolution and the ichnofacies model. Unfortunately,
the lack of ichnotaxonomic treatment of these trace fossils and the brief and poorly
documented descriptions reflected in the poor understanding of their affinities and
paleoenvironmental significance allow us only a tentative inclusion of these asso-
ciations in the ichnofacies paradigm (Bromley et al. 2007). The end-Triassic mass
extinction event seems to have had little effect on diversity of paleosol ichnofaunas
judging by their increase in diversity between the Early and Late Triassic, represented
in the Chinle Formation, and the comparable ichnodiversity between the Chinle and
Morrison formations (Hasiotis 2000, 2004).
Genise et al. (2008a) proposed that terrestrial crayfishes could be capable of con-
structing nests, which basically showed two architectures: necked cells attached to
parental burrows (Cellicalichnus meniscatus) and central chambers surrounded by
radiating, short, meniscate tunnels (Dagnichnus titoi). The attribution of these struc-
tures from the Upper Jurassic and Lower Cretaceous of Argentina to crayfishes
were based on their occurrence in the same beds, with abundant crayfish trace
fossils (Loloichnus baqueroensis) (Bedatou et al. 2008a), and in the case of
Cellicalichnus meniscatus because its cells are connected to tunnels indistinguish-
able from the former. In addition, Dagnichnus resembles the bauplan of thalassini-
dean calichnia (i.e., small cells or tunnels connected with large chambers or tunnels).
There are two outstanding and well described trace fossils from Lower Jurassic
paleosols of Poland and Iran that deserve special comment (Pieńkowski and
Niedźwiezki 2008; Fürsich et al. 2010). Are these trace fossils the pioneer expres-
sions of larval parental care? Do they represent a major step in the evolution of
paleosol ichnofaunas? Pieńkowski and Niedźwiezki (2008) described a trace fossil
composed of a central chamber surrounded by radiating cells, resembling the bau-
plan of Dagnichnus, from the Lower Jurassic of Sołtyków in Poland. Fürsich et al.
(2010) described Cellicalichnus antiquus, a trace fossil composed of horizontal tun-
nels bearing opposite pairs of side branches that are considered breeding cells, from
the Lower Jurassic Aghounj Formation of Iran. These authors made a well sup-
ported analysis of the affinities of these trace fossils, evaluating the possibilities of
bees or bee ancestors, and crayfishes (Genise et al. 2008a; Genise and Verde 2010)
as putative producers. They concluded that an insect origin was more likely because
its morphology was more similar to that of insect nests than of extant crayfish traces.
These cases, which probably have counterparts in the Chinle or Morrison forma-
tions, may represent a major acquisition in soil invertebrate behavior, the construc-
tion of cells in soils for larval development, regardless their crustacean or insect
affinities (Genise 2016; Genise et al. 2008a; Genise and Verde 2010).
The Upper Jurassic and Lower Cretaceous formations of Patagonia, where the
most diverse ichnoassemblages are recorded, are clearly dominated by earthworm
and crayfish trace fossils (Bedatou et al. 2008a; Bedatou 2010), whereas other
ichnoassemblages elsewhere are less diverse. At present, burrowing crayfishes
are found in a diversity of soils and vegetation supporting periodical waterlogging.
In South America, crayfishes inhabit swamp, marshes, and mossy bogs occurring in
evergreen forests and grasslands (Rudolph 1997; Rudolph and Crandall 2005;
Buckup 2003; Noro 2007) (Fig. 13.4b). In North America they live in wet prairies
with emergent sedges and grasses, marshes, and swamps (Grow 1981; Huner and
Barr 1991; Jordan et al. 2000), whereas in Oceania they inhabit marshes, swamp,
peatlands, and wetlands, occurring in grasslands, low shrublands, scrubs, and for-
ests (Lake and Newcombe 1975; Suter and Richardson 1977; Horwitz and Knott
1983, 1991; Richardson 1983; Horwitz et al. 1985; Growns and Richardson 1988;
Hamr and Richardson 1994; Richardson and Wong 1995; Whitmore et al. 2000).
Accordingly, the Camborygma Ichnofacies would be indicative of local marshes,
bogs, swamps, or wetlands as a subset of forest, scrub, and open herbaceous com-
munities (Mueller-Dombois and Ellenberg 1974).
In sum, Triassic, Jurassic, and most Cretaceous trace-fossil associations recorded
from paleosols of Australia, USA, South America, and Europe include or are domi-
nated by earthworm, crayfish, root, and/or undetermined trace fossils, which in
some cases could be produced by unidentified groups of insects (Tables 13.1 and
13.2; Figs. 13.1, 13.2, and 13.3). The Mesozoic record of trace fossils in paleosols
is greater than that of the Paleozoic, but it contains fewer and less studied occur-
rences than those of the Cenozoic. The lack of formally defined Mesozoic paleosol
ichnofacies until now reflects this scenario. A new ichnofacies dominated by cray-
fish and earthworm trace fossils, indicative of paleosols with fluctuating, high water
tables and distinctive of wetlands and swamps, may be defined at least tentatively
with the database presented herein. This ichnofacies is called the Camborygma
Ichnofacies, honoring the first ichnogenus named after crayfishes and also the oldest
record of crayfishes (Hasiotis and Mitchell 1993; Hasiotis et al. 1993b). The
Camborygma and Rhizolith Ichnofacies are the dominant paleosol ichnofacies from
the Early Triassic to the Early Cretaceous, and represent stasis until the advent of
new ichnotaxa and ichnofacies by the Late Cretaceous, the next revolution.
13.4 The Celliforma Revolution: The Late Cretaceous

Advent of Recognizable Insect Trace Fossils
in Paleosols and New Ichnofacies
By the Cretaceous, the Triassic–Jurassic scenario began to change slowly because

of the appearance and early diversification of different groups of holometabolous
insects capable of constructing preservable and distinct traces in soils (Genise and
Bown 1994a). This already “old” observation, put forward almost 20 years ago, is
still in force. Ants, bees, some groups of beetles, wasps, and termites that construct
preservable and recognizable traces in soils, probably favored by the diversification
of flowering plants, appeared by the Cretaceous when their first trace fossils are
also recorded (Genise 2016).
The body-fossil record of these distantly related groups of insects began in the
Cretaceous, even though there are abundant and impressive Triassic and Jurassic
334
Fig. 13.2 Stratigraphic ranges of insect trace fossils in paleosols. Only reliable attributions to
insects are included. Numbers in unnamed trace fossils with reliable attributions indicate consulted
references: (1) Gregory et al. (2009); (2) Genise and Bown (1994a, 1996), Hamer et al. (2007),
Iriondo and Krohling (1996), Kröhling (1999), Laza (1982, 1995, 1997), Sandau (2005), Tauber
(1996); (3) Gregory et al. (2009); (4) Hamer et al. (2007), Gregory et al. (2009); (5) Genise and
Verde (unpubl.), Verde and Genise (2007), Veroslavsky and Martínez (1996), Veroslavsky et al.
(1997); (6) Hamer et al. (2007), Yelinek and Chin (2007); (7) O’Geen and Busacca (2001); (8)
Hasiotis and Bown (1996), Hembree and Hasiotis (2007), Netto et al. (2007), Retallack et al.
(1995), Sandau (2005), Sheldon and Hamer (2010), Thackray (1994); (9) González (1999), Sciutto
and Martínez (1996); (10) Sciutto and Martínez (1996); (11) Moore and Picker (1991); (12) Bown
and Genise (1993), Darlington (2005, 2011), Genise et al. (2000), Krell and Schawaller (2011),
Laza (1995, 2006a), Radies et al. (2005)
localities bearing fossil insects worldwide (Grimaldi and Engel 2005). Here, it is
provided a brief summary of the body-fossil record of common insect producers of
trace fossils in Cretaceous–Cenozoic paleosols. The records of Dynastinae
(Scarabaeidae) are from the Cenozoic of the USA and Germany, the oldest one of
which is Oryctoantiquus borealis from the Middle Eocene of the USA (Ratcliffe
et al. 2005). There are records of Melolonthinae (Scarabaeidae) from the Cenozoic
of Russia, Germany, Czech Republic, and the USA, but the oldest records are from
the Lower Cretaceous of Russia: Cretoserica latitibialis Nikolajev (1998) and three
species of Lithanomala Nikolajev (1993) (Krell 2007). The oldest weevils, mostly
from the families Nemonychidae and Belidae, come from Late Jurassic deposits of
Karatau in South Kazakhstan (Gratshev and Zherikhin 2003; Oberprieler et al.
2007; McKenna et al. 2009; Legalov 2010). The oldest Curculionidae is
Arariperhinus monnei from the Santana Formation, Brazil (Lower Cretaceous,
Aptian-Albian) (Fernandes de Aquino Santos et al. 2011). Other Early Cretaceous
weevil fossils come from Sierra del Montsec (Spain), Yixian (China), Bon-Tsagan,
(Mongolia), Khetana and Ulya River (Russia), and Santana (Brazil); most of these
are members of the Nemonychidae (Gromov et al. 1993; Oberprieler et al. 2007;
Fernandes de Aquino Santos et al. 2011). Most families of extant weevils arose by
the end of the Cretaceous (McKenna et al. 2009). The oldest termite body fossils are
Baissatermes lapideus Rasnitsyn (2008) from the Lower Cretaceous (Barremian) of
Baissa, Siberia (Engel et al. 2007, 2009) and Morazatermes krishnai Engel and
Delclòs (2010) from the Lower Cretaceous (Albian) of Spain. Other termites, also
recorded from the same deposits but presently known only from the forewings are
Cantabritermes simplex Engel and Delclòs (2010) and Aragonitermes teruelensis
Engel and Delclòs (2010) in amber from Teruel, Spain. The oldest undisputed bee
is Cretotrigona prisca from New Jersey amber, which is closely related to the extant
stingless bees (Michener and Grimaldi 1988a; Engel 2000). The precise age of C.
prisca is still unclear since it was considered initially to be 80 My (Michener and
Grimaldi 1988a, b) but later estimated in 70 My (Grimaldi 1999), and still later in
65 My (Engel 2000). Poinar and Danforth (2006) described Melittosphex burmensis
from the Lower Cretaceous Burmese amber (Myanmar), which was originally con-
sidered as a transitional form between crabronid wasps and extant bees. More
recently, Danforth and Poinar (2011) proposed that M. burmensis was a pollen-
collector and accordingly the oldest bee.
The oldest ants are recorded from the Early Cretaceous (Dlussky 1996; Nel et al.
2004; Ward 2007; Perrichot et al. 2008), and are unexpectedly diversified with several
distinct genera assigned minimally to two subfamilies (Perrichot et al. 2008).
The oldest body-fossil records are Haidomyrmodes mammuthus (Sphecomyrminae)
(Perrichot et al. 2007) and Gerontoformica cretacica (uncertain subfamily) (Nel et al.
2004) from the Lower Cretaceous (Upper Albian) of Charente-Maritime locality
(France), and Haidomyrmex cerberus (Dlussky 1996), Sphecomyrmodes orientalis,
Sphecomyrma sp. (both Sphecomyrminae), Burmomyrma rossi, and Myanmyrma
gracilis (both uncertain subfamily) from the Lower Cretaceous (Upper Albian) of
the Myanmar locality in Burma (France) (Dlussky 1996, Engel and Grimaldi 2006,
Ward 2007; Perrichot et al. 2008). Finally, the oldest Aculeata is from the Upper
Jurassic of Karatau in Kazakahstan (Central Asia) and was placed in the extinct
family Bethylonymidae (Rasnitsyn 1975, 2002). This family probably represents
the closest relative and gave rise to all modern aculeates (Rasnitsyn 2002; Brady
et al. 2009). Many modern vespoid families diverged in the Late Jurassic and
throughout the Early Cretaceous, although few families, including Pompilidae,
Mutiliidae, and Sapygidae, may have originated considerably more recently
(Brady et al. 2009). The earliest specimens of Sphecidae (Angarosphecidae)
appeared during the Early Cretaceous and these were recorded from many localities
Fig. 13.3 Ichnodiversity of trace fossils in paleosols through the Phanerozoic. To emphasize
changes in ichnodiversity, all trace fossils in Table 13.1 are included, independently of the reliability
of their ichnotaxonomy and affinities. The Scoyenia Ichnofacies is depicted only for the Paleozoic
to emphasize that its examples are one of the oldest ichnologic evidence of subaerial exposure and
principally in Eurasia (Spain, Burma, Mongolia, China, Russia, UK, and

Transbaikalia) (Evans 1969; Rasnitsyn 1975, 1980, 2000; Darling and Sharkey
1990; Rasnitsyn et al. 1998, 1999; Rasnitsyn and Ansorge 2000; Rasnitsyn and
Martinez-Delclòs 2000). The oldest Pompilidae is Bryopompilus interfector (tribe
Bryopompilini) from the Lower Cretaceous (Albian) amber of Myanmar (Burma)
(Engel and Grimaldi 2006).
These phylogenetically unrelated groups of insects, the most common producers
of trace fossils in paleosol since the Cretaceous, share a similar behavioral trait.
Representatives of these taxa line or construct different parts of their nests, cocoons,
or pupation chambers, which are not mere excavations, increasing significantly the
potential of preservation (Genise and Bown 1994a). Analogously, insects reinvented
several millions of years later, two behavioral acquisitions of Jurassic crayfishes:
linings for reinforcing walls and chambers for rearing larvae (Genise et al. 2008a).
The need of Jurassic and Cretaceous crayfishes to maintain water for breathing
inside their burrows probably favored the acquisition of pelletal linings to increase
isolation from soil and retain water within the burrows (Bedatou et al. 2008a).
Similar behavioral acquisitions, such as pelletal and fluidized linings, and other
features exclusive of insects, such as free-standing walls and organic linings, occur
for the first time in Late Cretaceous paleosol trace fossils, such as cocoons, pupa-
tion chambers, and nests, which can be attributable to insects. These behavioral
traits are recorded for the first time in a few deposits worldwide (Table 13.1;
Figs. 13.2 and 13.3).
The advent of flowering plants by the Early Cretaceous was probably one of
the greatest innovations, together with the radiation of Triassic seed plant lin-
eages, for Mesozoic continental environments. Examples of paleosols related to
primitive angiosperms are known from North and South America. Paleosols of
Barremian age are related to the earliest angiosperms in the eastern USA and cor-
respond to Entisols, pink clayey Inceptisols, and coaly Histosols covered by coni-
fers (Retallack and Dilcher 1986; Retallack 2001a). In southern Patagonia
Fig. 13.3 (continued) because they constitute about half of the Paleozoic cases. The dotted line
indicates that it is recorded up to the Holocene, but these post-Paleozoic cases are not treated herein.
The first case of a paleosol bearing only rhizoliths, the rhizolith revolution, took place during the
Early Devonian. The Rhizolith Ichnofacies is later extensively recorded along the Phanerozoic. The
Camborygma revolution, during the Early Triassic, indicates the advent of assemblages dominated
or including earthworm and crayfish trace fossils (Camborygma Ichnofacies). The last record of this
ichnofacies is from the Eocene, but dotted line indicates that extant equivalent examples exist. The
Celliforma revolution, during the Late Cretaceous, is indicated by a rise in the ichnodiversity, which
includes records of the first recognizable insect trace fossils in paleosols and the oldest records of
the Celliforma Ichnofacies. The Coprinisphaera revolution, by the early Eocene, is envinced by
another rise in the ichnodiversity triggered by the EECO, the appearance and spreading of grass-
dominated habitats, and of the oldest cases of the Coprinisphaera Ichnofacies. The Termitichnus
Ichnofacies appeared by the Oligocene in closed forest paleoenvironments. The Celliforma and
Termitichnus ichnofacies could have been originated by the Early Cretaceous due the diversification
of its producers by that time. Accordingly, its possible origin is prolonged by a dotted line, although
there is no record of trace fossils attributable to soil termites, bees, or wasps
(Argentina), the first Barremian–Aptian angiosperms occupied wetlands that were

affected by recurrent volcanic ashfalls. They grew on andic and kaolinitic Entisols
that supported diverse conifers (Cheirolepidaceae, Podocarpaceae and subordinate
Araucareaceae, Cupressaceae), along with Bennettitales, Cycadales, some
Ginkgoales, ferns, lycopods, and bryophytes (Cladera et al. 2002; Del Fueyo et al.
2007). Similar, poorly-drained, Aptian environments of the central USA were
also occupied by angiosperms, growing on Entisols and Histosols of lowlands and
coastal-marine settings (Retallack and Dilcher 1981). Apparently, Early and mid
Cretaceous angiosperms were early-successional colonizers (Retallack and
Dilcher 1986). Detailed sedimentologic and ichnologic information from Upper
Jurassic–Lower Cretaceous continental successions of southern Patagonia
(Argentina) indicates that terrestrial environments, paleosols and associated trace
fossils show no changes with the appearance of angiosperms. Lithofacies associa-
tion and paleosol types of the pre-angiosperm Late Jurassic Bajo Grande
Formation are similar to those of the post-angiosperm Aptian Bajo Tigre Formation
(Cladera et al. 2002; Bedatou et al. 2009).
According to Retallack (1986, 1991b) the significant modification on the weath-
ering pattern produced by the expansion of angiosperms since the Cretaceous (Knoll
and James 1987) had no consequences on soils since Jurassic paleosols are similar
to those of the early Paleogene. However, the diversification of termites, related to
the radiation of angiosperms, may have caused a change in Oxisols (Schaefer 2001)
through the incorporation of oval pellets rich in gibbsite, Fe-oxides, and charcoal,
along with elongate burrows filled with these microaggregates (Eschenbrenner
1986; Schaefer 2001). Thus, it is probable that proliferation of termites intro-
duced a modification of soils in tropical ecosystems. Nevertheless, most of middle
and Late Cretaceous examples of new insect trace fossils suggest no comparable
changes in paleosols in temperate regions. In any case, the appearance of flower-
ing plants had little effect on soil types, which remained mostly the same since the
Carboniferous.
The record of the third revolution, mostly Late Cretaceous in timing (Table 13.1;
Figs. 13.2 and 13.3), includes Fictovichnus gobiensis from the Djadokhta Formation
of Mongolia (Johnston et al. 1996); Cellicalichnus dakotensis from the Dakota
Formation of the USA (Elliot and Nations 1998); Pallichnus dakotensis and
Fictovichnus sciuttoi from the Bajo Barreal Formation of Argentina (Genise et al.
2007); Cellicalichnus chubutensis and Rebuffoichnus casamiquelai from the
Laguna Palacios Formation of Argentina (Genise et al. 2002a), and Fictovichnus
sciuttoi from the Allen Formation of Argentina (Genise and Sarzetti 2011).
Particularly interesting are the first records of assemblages of the Celliforma
Ichnofacies in the Mercedes Formation of Uruguay (Alonso-Zarza et al. 2011) and
the Two Medicine Formation of the USA (Martin and Varricchio 2011). The for-
mer assemblage is composed of Celliforma spirifer, C. germanica, Fictovichnus
gobiensis, and rhizoliths, whereas the latter is composed of Fictovichnus sciuttoi,
Rebuffoichnus isp, and Skolithos isp. It is possible that other cases mentioned pre-
viously also may represent examples of the Celliforma Ichnofacies, such as that of
the Allen Formation (Genise and Sarzetti 2011). However the lack of recorded
assemblages composed of different ichnotaxa or carbonate-rich paleosols preclude

their inclusion until more evidence is recovered to understand the complete
paleoenvironmental significance of this ichnofacies. Even when some of these ich-
nogenera, such as Fictovichnus, Pallichnus, or Rebuffoichnus, are also representa-
tives of the Coprinisphaera or Celliforma Ichnofacies (Genise et al. 2000, 2010b),
when they do occur in associations dominated by earthworm or crayfish traces
(i.e., Laguna Palacios Formation), they can be considered more likely as secondary
participants of the Camborygma Ichnofacies, rather than indicators of Cretaceous
examples of the Coprinisphaera or Celliforma Ichnofacies. The same is true for
North American formations, like Claron and Willwood, in which insect trace fos-
sils like Parowanichnus, Eatonichnus, Celliforma, Naktodemasis, and wasp
cocoons are associated with crayfish and earthworm traces (Bown and Kraus 1983,
1987; Bown et al. 1997; Hasiotis and Bown 1996). The Celliforma ichnofacies is
indicative of calcretes developed under arid or semiarid conditions and palustrine
carbonates (Fig. 13.4c, d).
The presence of the Termitichnus Ichnofacies during the Cretaceous is a pos-
sibility considering that termites were already diversified by this period. However,
there are no cases recorded of associations of termite nests for the Mesozoic, with
the exception of those structures described by Bordy et al. (2004) from the Jurassic
of the Karoo Basin, which probably deserve a different interpretation (Genise
et al. 2005; Alonso-Zarza et al. 2008). Two cases of trace fossils attributed to
social insects, including termites, require a brief analysis. Genise et al. (2010a)
described rhizolith balls from the Lower Cretaceous Cerro Barcino Formation of
Argentina, and discussed their affinities, such as self-induced and localized over-
growth of secondary and tertiary rootlets, crayfish feeding chambers, and termite
or ant chambers for agricultural purposes. Tentatively, they were assigned to ant
fungus gardens, the oldest evidence of insect agriculture, which would represent
an important element of the Cretaceous revolution. The other case, Socialites
tumulus, described by Roberts and Tapanila (2006) from the Upper Cretaceous
Kaiparowits Formation of the USA, was interpreted as an ant or termite nest.
However, the morphology may also resemble bioturbated stump casts or megarhi-
zoliths. In any case, both records do not qualify as Cretaceous examples of the
Termitichnus Ichnofacies.
In sum, bee cells and nests, putative ant and termite nests, coleopteran pupation
chambers, and wasp cocoons are the Late Cretaceous pioneer trace fossils that sig-
nal another great change that would undergo paleosol ichnofaunas during the
Eocene, the fourth revolution (Tables 13.1 and 13.2; Figs 13.2 and 13.3). The
increase of ichnodiversity; the appearance of the first recognizable, constructed or
lined, insect trace fossils in paleosols included as secondary components of the
Camborygma Ichnofacies; and the first record of the Celliforma Ichnofacies, also
are keystones of the third revolution in paleosol ichnofaunas. The possibility of the
oldest physical evidence of sociality and insect agriculture is equally significant
from an evolutionary viewpoint. According to the body-fossil record, Early–mid
Cretaceous records of the Celliforma Ichnofacies could be expectable. The lack of
them could be an artifact or it could correspond to the absence of some behaviors
related to wall construction in the earliest representatives of the involved groups.
Fig. 13.4 Modern analogous environments of the six paleosol ichnofacies. (a) Scoyenia
Ichnofacies. Drying ephemeral lake deprived of vegetation in Bajo de los Huesos, Chubut,
Argentina. Note several vertebrate trackways in the foreground, and some dessiccation cracks in
the lower right corner; (b) Camborygma Ichnofacies. Wetland in a glade of an evergreen forest at
Rucapihuel, Chile. Note the soil flooded by the rise of water table in the center of the figure. (c)
and (d) Celliforma ichnofacies at Las Tablas de Daimiel, Spain. Bare soil in a palustrine environ-
ment rich in carbonate (c), and carbonate rich soil supporting sparse vegetation. The calcrete was
broken and exposed to the surface by plowing. (e) Coprinisphaera Ichnofacies. Savanna domi-
nated by grasses at the Pilanesberg National Park, South Africa. Bare soil produced by trampling
of the rhino (at the center) and other large mammals is optimal for bee nesting. Large size herbi-
vores like this provide dung for coleopteran brood balls. (f) Termitichnus Ichnofacies. Closed for-
est of warm and humid climate in Misiones, Argentina. Tall trees, with interlocking crowns and
understory of tree ferns
13.5 The Coprinisphaera Revolution: The Paleogene

Explosion of Insect Trace Fossils Related
To Grass-Dominated Habitats
The end Cretaceous mass extinction event seems to have had little effect on paleosol
ichnofaunas (Fig. 13.3), although the Paleocene record is comparatively scarce for
an accurate evaluation. One of the most significant floristic changes related to this
biotic crisis was the extinction of several species of evergreen angiosperms and the
subsequent predominance of conifers and deciduous angiosperms in the early
Paleogene vegetation (Retallack 2001a).
In South America, particularly in Central Patagonia, the Late Cretaceous succes-
sions end with a clear dominance of earthworm and crayfish trace fossils, as in most
Mesozoic units, and with the appearance of a few insect trace fossils (Genise et al.
2002a, 2004; Bedatou et al. 2008a). For the same region, the Paleocene–Eocene Rio
Chico Group shows scarce trace fossils, including the appearance of the first trace
fossil attributable to cicadas, Feoichnus challa, in the Koluel Kaike Formation of
Patagonia (Krause et al. 2008, 2010). The earliest body fossils of true cicadas
(Cicadoidea) are from the Triassic of Russia, France, and Australia (Shcherbakov
2008). These specimens are included in the family Tettigarctidae (hairy cicadas).
However, the distinct fossorial forelegs of Cicadoidea nymphs apparently were
present only by the mid Cretaceous, in amber from New Jersey (Grimaldi and Engel
2005). In South America, body fossils of hairy cicadas also are known from the
Lower Cretaceous of Brazil (Lefebvre et al. 1998; Shcherbakov 2008). The earliest
record of the family Cicadidae is from the Paleocene (ca. 60 Ma) of North America
(Cooper 1941). Another ichnogenus, Eatonichnus, attributable to dung beetles
(Bown et al. 1997; Krause et al. 2007; Sánchez et al. 2010a), is recorded for the first
time from the Peñas Coloradas Formation in Patagonia (Genise et al. 2001a) and the
Claron and Colter formations in the USA (Bown et al. 1997). Few recognizable
earthworm burrows and doubtful crayfish trace fossils are recorded from the Rio
Chico Group (Krause et al. 2007; unpub. data).
In contrast, in North America, crayfish burrows (Camborygma) are still abundant
in Upper Paleocene–Lower Eocene units of Wyoming (Hasiotis and Honey 1995,
2000; Smith et al. 2008b) and Utah (Hasiotis and Bown 1996). The most diverse
Paleocene assemblage is from the Willwood Formation (Bown and Kraus 1983,
1987; Hasiotis et al. 1993a; Smith et al. 2008a, b, c, 2009) and from the Paleocene–
Eocene Claron Formation (Hasiotis and Bown 1996; Bown et al. 1997), both of
which are Cenozoic cases of the Camborygma Ichnofacies. The evidence that sup-
ports this assumption is: (1) the abundance of Camborygma and Edaphichnium in
the former deposits, (2) the abundance of Camborygma in the latter deposits, and
(3) the paleosols exhibiting fluctuating water tables. In those assemblages, recog-
nizable insect trace fossils are associated with crayfish and earthworm ones, as in
the Upper Cretaceous deposits of Patagonia. The Paleocene–Eocene Thermal
Maximum (PETM) produced the reduction in size and an increase in diversity of
trace fossils in the Willwood Formation of USA (Smith et al. 2009).
In both Patagonia and the Western Interior of the USA, crayfish trace fossils are
mostly absent in younger deposits. In South America, soil crayfishes are at present
restricted to humid environments of Chile and Brazil (Bedatou et al. 2008a).
Earthworm trace fossils, although less recorded, are similarly present in Cenozoic
trace-fossil assemblages from younger units. Earthworms and crayfishes inhabit
modern South and North American soils, indicating that at a global scale the
end Cretaceous mass extinction had little direct effect on these organisms (Figs.13.1
and 13.3), even though regionally they displayed changes in their geographic
distribution (Bedatou et al. 2008a). In sum, the end Cretaceous mass extinction
(see Chap. 14), a major event in the history of life, was less important for paleosol
ichnofaunas than previous events, as the rising and diversification of angiosperms
during the Early Cretaceous, and later events, such as the origin and diversification
of grass-dominated habitats during the Middle-Late Eocene cooling trend after the
EECO. The fossorial habit of invertebrate soil organisms could have favored its
survival in critical episodes, such as proposed for fossorial vertebrates (Archibald
and Bryant 1990; Sheehan and Fastovsky 1992; MacLeod et al. 1997; Robertson
et al. 2004; Longrich et al. 2012).
The last major step in the evolution of plant communities and soils took place
during the middle Paleogene–early Neogene with the expansion of new, fast-
growing monocot plants: the grasses. Grass-dominated habitats (Fig. 13.4e) are
high-productivity ecosystems associated with dry or nonhumid habitats (subhumid
to semiarid) of different continents, currently occupying 40 % of global land surface
(Anderson 2006). However, grasses can also grow in less favorable conditions and
environments, such as highly seasonal and tropical, cool temperate, high-mountain
prairies, salt marshes, and are successional after periodic fires. Grass-dominated
habitat expansion promoted significant changes in biota, favoring development of
large vertebrates (grazers) on ground and diverse soil invertebrates, despite of the
development as a defensive strategy of chemically harmful effects on mammals,
insects, mites, and fungi (Retallack 2001a). As a consequence of changes in their
rooting system and soil biota, grassland soils acquired particular characteristics in
their soil aggregates, such as coarse granular or near-mollic to very fine-granular or
crumb ped structure. Other diagnostic character of these new soils is a dark surface
horizon rich in nutrients (mollic epipedon), displaying evidence of intense activity
of burrowing invertebrates, such as fecal pellets of earthworms (Retallack 2001a).
These features defined a new type of soil: the Mollisol. The appearance of a granu-
lar surface horizon in Oligocene-Miocene paleosols of the USA was considered a
proxy for grasslands, which replaced and displaced wooded shrublands and dry
woodlands (Retallack 1990). Fossil Mollisols present abundant, short and fine root
traces. Desert grasslands, bunch grasslands, and rangelands developed in the latest
Eocene–early Oligocene in NW and central USA (Retallack 2001a, 2009). The first
short (sod) grassland Mollisols with fine crumb peds and dense and fine rhizoliths
appeared by the early Miocene (19 Ma) in the USA and Kenya. Both examples
occurred in dry climates based on the presence of shallow, calcic horizons (Retallack
2004; Retallack et al. 1995).
Twenty millions years before, a set of particular factors converged in Central

Patagonia (Argentina) to trigger the formation of more ancient grass-dominated
habitats. Main factors were probably a wetter climate and soils formed in fresh vol-
canic ashes. In addition, the effect of herbivorous mammals and associated dung
beetles would also have contributed to the spread of these habitats. Paleosols of the
Sarmiento Formation formed in pyroclastic mudstones composed of dacitic–rhyo-
litic glass shards (61–99 %) and andesine plagioclase (Bellosi 2010). The distinctive
properties of this acid, low bulk-density volcaniclastic material benefited plant
growth and rooting by providing high fertility, rapid weathering, a high moisture-
holding capacity, and elevated macroporosity resulting in an appropriate medium
for the extension of root hairs in a well-aerated medium (Nanzyo 2002). The appear-
ance of the oldest grass-dominated habitats and grasslands is matter of debate and
controversial evidence (Genise 2016). The first documented grass-dominated habi-
tats for South America are middle Eocene (39 Ma) and occur at Gran Barranca in
the middle Eocene-lower Miocene Sarmiento Formation of Patagonia. This pro-
posal is supported by paleosols and loessic deposits (Bellosi and González 2010),
phytoliths (Zucol et al. 2010; Sánchez et al. 2010c) and density and diversity of
dung beetle brood balls (Sánchez et al. 2010b). The only thorough and comparative
phytolith analysis of Coprinisphaera and bearing paleosols (Sánchez et al. 2010c)
showed that grasses were the second dominant group after palms, as expectable for
grassy savannahs with palms or palm groves (Cabrera 1971). The oldest dung bee-
tles attributable to ball-making species are Eocene (Krell 2007). Morphology and
behavior of dung beetles reflect a strong adaptation to exploit mammal excrement in
open-grass habitats (Halffter and Edmonds 1982; Hanski and Cambefort 1991).
African savannahs show the greatest diversity of dung beetles, whereas South
American forests show a large diversity because of the Quaternary extinction of
large mammals (Halffter 1991). Trace fossils of the Sarmiento Formation are clear
examples of the Coprinisphaera Ichnofacies, indicating open-herbaceous commu-
nities (Genise et al. 2000). High density (40–100 balls/m2), abundance and diversity
in sizes and ichnospecies of Coprinisphaera, indicating a large number and diver-
sity of dung beetles (Sánchez et al. 2010b), reflect the middle Eocene appearance of
open-grass habitats (Bellosi et al. 2010), coincident with the increase of diversity
and size of mammals (Woodburne et al. 2014). Probably the appearance of grass-
dominated habitats was diachonous in different continents through the middle
Paleogene–early Neogene. Those of Gran Barranca were probably similar to the
present Chaco landscapes of northern Argentina (Cabrera 1971), represented by a
mosaic vegetation composed of grassy savannahs with palms and palm groves, and
forest in patches and riverbanks. Such a paleolandscape is concordant with all sedi-
mentologic, pedologic, paleontologic, and ichnologic evidence for Gran Barranca.
A possible still older evidence for a wooded grass-dominated habitat, developed
at lower latitudes, is represented in the renowned lower Eocene Asencio Formation
of Uruguay. The precise age of this unit is unknown because of the lack of datable
organisms or rocks. However, the lateritic character of the soils triggered by the
Early Eocene Climatic Optimum, the presence of abundant large Coprinisphaera
that can be only related with large mammals, and the age of the underlying and
overlying formations, indicate that this formation is early Eocene in age (Genise
et al. 2004; Bellosi et al. 2004, 2016). The Asencio Formation represents not only
the oldest example of the Coprinisphaera Ichnofacies, but also one of its best exam-
ples because of the diversity of the ichnoassemblage. The presence of a grass-
dominated habitat is indicated by the extraordinary development of the
Coprinisphaera Ichnofacies and the record of Ultisols (González 1999), developed
under a warm and seasonal-humid climate. When analyzed with more resolution,
the Asencio Formation comprises two alternating ichnoassemblages which corre-
spond to the different ichnofabrics recognized either in the duricrusts or in nodular
beds Bellosi et al. (2016). The duricrusts, representing drier periods and shallower
layers of the soil, bear ichnoassemblages dominated by Palmiraichnus, Teisseirei,
and secondarily Rebuffoichnus. In turn, the nodular beds, representing wetter peri-
ods and deeper layers of the soil, bear more diverse ichnoassemblages including all
ichnotaxa recorded for the formation. Previous and current research shows that the
Asencio ichnofauna is composed of traces of cicadas (Monesichnus), dung beetles
(Coprinisphaera), bees (Palmiraichnus, Elipsoideichnus, Uruguay, Corimbatichnus,
Celliforma), sphinx moths (Teisseirei), termites (Krausichnus), beetles
(Rebuffoichnus), crayfish (Loloichnus), and cleptoparasites and detritivores
(Tombownichnus, Lazaichnus) (Genise and Bown 1996; Genise and Hazeldine
1998a, b; Genise and Laza 1998; Genise 2000; Genise et al. 2002b, 2013c; Genise
and Verde 2000; Mikúlaš and Genise 2003; Sánchez and Genise 2009; Verde and
Genise 2010). The extraordinary diversity of this trace-fossil assemblage reveals in
turn a higher diversity of insects inhabiting lateritic soils that may be the direct con-
sequence of the high temperatures and precipitation related to the EECO in this
grass-dominated habitat. This and the contemporaneous assemblage from the Gran
Salitral Formation of La Pampa (Melchor et al. 2002) record the first appearance of
Teisseirei barattinia, the only paleosol trace fossil attributed to sphinx moths Genise
et al (2013c). The oldest records of Sphingidae recognized by Sohn et al. (2012) are
an adult from the middle Eocene Baltic Amber (Kusnezov 1941), and a caterpillar
from the early Miocene of Baden-Wüttemberg, Esslinger, Germany (Zeuner 1927).
Other early body fossil Sphingidae is Mioclanis shanwangiana from the middle
Miocene (~15–17 My) of Shanwang, Shandong, China (Zhang et al. 1994). Other
species named Sphinx snelleni was described by Weyenbergh (1869) as a sphingid
from the Upper Jurassic Solenhofen Limestone deposits in Bavaria, Germany.
However, it has later proved to be a wood wasp of the hymenopteran family Siricidae
(Kitching and Sadler 2011). Skalski (1990) recorded also a Sphingidae from the
Baltic amber, but Ross (1996) considered this claim to be unsubstantiated.
Other outstanding assemblages belonging to the Coprinisphaera Ichnofacies are
those from different stratigraphic levels of the middle Eocene–lower Miocene
Sarmiento Formation of Argentina (Genise 2016; Genise et al. 2004; Bellosi et al.
2010; Sánchez et al. 2010b). Previous and current research demonstrates that the
Sarmiento ichnofauna is composed of trace fossils of cicadas (Feoichnus), dung
beetles (Coprinisphaera, Chubutolithes, Eatonichnus, Pallichnus), sphinx moths
(Teisseirei), bees (Celliforma), earthworms (Lazaichnus, Castrichnus, diffuse box-
works), and cleptoparasites (Tombownichnus) (Krause and Genise 2004; Krause

et al. 2008; Bellosi et al. 2010; Sánchez and Genise 2009; Sánchez et al. 2010b). In
this case, several paleolatitudinal degrees southwards and about 10 million years
later, the extraordinary diversity of dung beetle trace fossils (Sánchez et al. 2010b)
cannot be attributed directly to the EECO, but to the establishment of extended
grass-dominated habitats, bearing abundant large mammals, which provided the
dung for beetle brood masses. The high-resolution ichnostratigraphy of
Coprinisphaera at the locality of Gran Barranca records important paleoenviron-
mental changes in those former grass-dominated habitats, including an increase in
mammal size by the late Oligocene (Sánchez et al. 2010b).
The major evolutionary event during the Eocene, which represents the appear-
ance and establishment of grass-dominated habitats, produced the appearance of
larger mammals and abundant dung, which in turn promoted the evolution of
coprophagy in dung beetles. Ichnology can reconstruct partially this history (Genise
2016). The first ichnologic evidence of dung-beetle coprophagy is probably recorded
by trace fossils found in dinosaur coprolites from the Upper Cretaceous Two
Medicine Formation of Montana (Chin and Gill 1996) (but see Arillo and Ortuño
2008). Theoretically, in the early stages of coprophage evolution, dung beetles pro-
duced meniscate burrows inside the dung pads and underneath in the soil substrate.
In more derived behavioral stages, dung beetles construct, and bury in the soil,
brood balls for rearing their larvae, which are preserved as trace fossils and recog-
nized as specimens of Coprinisphaera. Which is the oldest record of Coprinisphaera
and accordingly of brood ball construction? Halffter and Edmonds (1982) and Krell
(2006) hypothesized about ball-making dung beetles utilizing the abundant dung of
dinosaurs, whereas Arillo and Ortuño (2008) disregarded dinosaurs as dung provid-
ers. Arillo and Ortuño (2008) based their conclusions first, on the mixture of reptil-
ian feces with excretory products; and second, on the poor attraction of extant dung
beetles to those feces. With the only exception of a single putative specimen of
Coprinisphaera found in the Cretaceous of Brazil (Souza Carvalho et al. 2009), there
are no reliable records older than Eocene. Which is the ichnologic record of the evolu-
tion of coprophagy between the Cretaceous burrows of the Two Medicine Formation
and the Eocene brood balls of South America? Sánchez et al. (2010a) described a
pupation chamber of extant dung beetles, which matches the morphology of
Eatonichnus (and partially Chubutolithes), trace fossils that are found in Paleocene–
early Eocene deposits of Argentina and the USA (Bown et al. 1997; Genise et al.
2001a; Krause and Genise 2004; Krause et al. 2007), predating the appearance of
Coprinisphaera. These Paleocene trace fossils probably reflect the life habit of
Paleocene dung beetles that exhibit similar behaviors to extant species of Eucranini
(Scarabaeinae), which provision their nests with fecal pellets of small mammals, as
those recorded for the Paleocene. Mature larvae construct a helical pupation chamber
composed of pellets similar to Eatonichnus (Sánchez et al. 2010a).
Can the body-fossil record of dung beetles corroborate this evolutionary history
proposed by ichnology? Regrettably, the body-fossil record of American dung
beetles is young. The oldest body fossils identified as Scarabaeinae are Prionocephale
deplanate Lin (1980) from the Upper Cretaceous of China and Cretonitis copripes
Nikolajev (2007) from the Lower Cretaceous of Russia (Krell 2007). However,
among those genera whose species are recognized ball makers, the oldest is
Gymnopleurus eocaenicus Meunier (1921) (Gymnopleurini) from the Eocene of
Germany (Krell 2007). The fossil record of other ball makers is still younger. The
oldest Dichotomiini are Heliocopris antiquus Fujiyama (1968) from the Miocene of
Noto, Japan, and Anachalcos mfwangani Paulian (1976) from the Miocene of Lake
Victoria, Kenya. The oldest Phanaeini are Phanaeus antiquus Horn (1876) from the
Pleistocene of the Port Kennedy caves, in the USA, and Palaeocopris labreae Pierce
(1946) from the Pleistocene of Rancho La Brea, USA. Finally, the oldest Canthonini
are Copris leakeyorum Paulian (1976) and Metacatharsius rusingae Paulian (1976)
from the Miocene of Lake Victoria, Kenya.
By the Eocene, there are also diverse assemblages belonging to the Celliforma
Ichnofacies from palustrine carbonates. The Gran Salitral Formation of Argentina
(early Eocene) probably was influenced by the elevated temperatures of the EECO,
but in drier conditions than those recorded for the Asencio Formation. The assem-
blage is represented by Celliforma ispp., Teisseirei, Rosellichnus, Skolithos,
Taenidium, and Fictovichnus (Melchor et al. 2002). Another assemblage of possible
late Eocene age occurs in the Queguay Formation of Uruguay, and is represented by
Celliforma spirifer, C. germanica, C. rosellii, Fictovichnus gobiensis, Fictovichnus
sciuttoi, and rhizoliths (Alonso-Zarza et al. 2011).
After the Eocene, few diverse ichnoassemblages in paleosols can be recognized.
The youngest paleosol ichnofacies appeared by the Oligocene in the Jebel Qatrani
Formation of Egypt. Paleosols of closed-forest paleoenvironments (Fig. 13.4f) pre-
served in this unit include the first assemblage dominated by termite and ant trace
fossils (Termitichnus, Fleaglellius, Vondrichnus, Krausichnus) (Genise and Bown
1994b), composing the Termitichnus Ichnofacies (Tables 13.1 and 13.2; Figs. 13.2
and 13.3). This incipient ichnofacies has another potential example in some paleo-
sols of the Chui Formation in Brazil (Netto et al. 2007). Even when diverse ichnoas-
semblages may be expected in closed-forest soils, the geologic record of these
associations is very scarce. The lateritization process involves the destruction and
incorporation of old termite nests to the soils (e.g., Eschenbrenner 1986; Schaefer
2001), probably reducing the probability of tropical termite nests to cross the tapho-
nomic barrier.
Other diverse trace-fossil assemblages in paleosols are recorded in the lower
Oligocene White River Formation of the USA (Hembree and Hasiotis 2007); the
upper Oligocene–lower Miocene John Day Formation of the USA (Retallack 2004);
the lower Miocene Pinturas Formation of Argentina (Genise and Bown 1994a;
Bedatou 2010); Miocene gypsum and gray shales of Spain (Uchman and Alvaro
2000); the Miocene Pawnee Creek Formation of the USA (Hembree and Hasiotis
2008); the upper Miocene Collon Cura Formation of Argentina (Bedatou 2010);
and the upper Miocene of the northern African Chad Basin (Duringer et al. 2007).
The remaining Miocene, Pleistocene, and Holocene record is composed mostly of
descriptions or citations of single or a few trace fossils of the above mentioned
groups of insects from different formations, which can be included in the
Coprinisphaera and Celliforma Ichnofacies. The peaks shown by Fig. 13.3 for the
Neogene, even when comparable with older ones, correspond to no particular

evolutionary events, but rather to more extensive research mostly carried out in
Pliocene and Pleistocene deposits of Argentina and Africa.
13.6 Conclusions
Our analysis of a database composed of 166 cases of invertebrate and plant trace
fossils in paleosols resulted in the recognition of four major evolutionary steps from
the first paleosols-bearing rhizoliths preserved in Early Devonian rocks to those of
the Holocene. Each step constitutes a revolution for paleosol ichnofaunas, which
triggered the appearance of a new ichnofacies. These events reflect some of the
major changes in the history of life. Other historical events, such as the Cretaceous–
Paleocene event, have no reflection in paleosol ichnofaunas or alternatively the data
is too scarce for detection.
The first revolution is associated with the appearance and expansion of vascular
plants, and took place in the Early Devonian with the first paleosols showing only
ichnoassemblages composed of rhizoliths, which constitute half the cases of the
Paleozoic and are recorded up to the Holocene. The appearance of vascular plants
by the late Silurian resulted in the Early Devonian appearance of several types of
paleosols (i.e., Histosols, Spodosols, Alfisols, and Ultisols) linked to the develop-
ment of new ecosystems. These rhizolith assemblages compose an archetypal ich-
nofacies, the Rhizolith Ichnofacies, which would be indicative of subaerial exposure
and the presence of different types of vegetation. Depending on needed and pending
studies on rooting patterns, in the future this ichnofacies can yield more precise and
significative paleoenvironmental data. Also, the absence of invertebrate trace fossils
may be a clue to interpret better the paleoenvironment. Other Paleozoic ichnofau-
nas, composed of scarce invertebrate trace fossils, some of them meniscate, are
considered herein as representative of the Scoyenia Ichnofacies, reflecting the peri-
odical subaerial exposure of the deposits.
These ichnoassemblages are the only recorded for the rest of the Paleozoic until
the second revolution, after the end-Permian mass extinction occured, with the
appearance of trace-fossil assemblages dominated by earthworm (Edaphichnium,
Castrichnus, diffuse boxworks) and crayfish (Camborygma, Loloichnus,
Dagnichnus, Cellicalichnus, Katbergia) trace fossils. These ichnoassemblages,
indicative of paleosols with high fluctuating water tables and representative of local
marshes, bogs, swamps, and wetlands, may be grouped into a new archetypal ich-
nofacies: the Camborygma Ichnofacies, which should be defined more completely
in the future, when better-documented cases are available.
The third revolution occurred in the Late Cretaceous. By that time, the diversi-
fication of flowering plants triggered the diversification of certain soil-inhabiting
insects, such as ants, termites, bees, wasps, and some beetles, which were capable
of constructing linings and free-standing walls for their chambers and nests that
consequently acquired a high preservation potential. By the Late Cretaceous, the
first recognizable insect trace fossils in paleosols occurred, either as isolated exam-
ples, as part of the Camborygma Ichnofacies, or in a new one: the Celliforma
Ichnofacies. The end Cretaceous mass extinction shows no global changes in
paleosol ichnofaunas, although the Paleocene record is scarce for a thorough evalu-
ation. In southern South America, the Camborygma Ichnofacies, which was domi-
nant since the Late Jurassic, dissapeared. In North America the Camborygma
Ichnofacies remains dominant in Paleocene–Early Eocene deposits, bearing recog-
nizable insect trace fossils, resembling those assemblages of the South American
Late Cretaceous.
The fourth and most important revolution for ichnodiversity took place in the
middle Eocene, influenced by the advent and expansion of grass-dominated habitats.
Other favorable conditions that promoted this ichnofacies were the early Eocene
Climatic Optimum, and the establishment of modern insect groups. These conditions
also lead to the appearance of a new soil type, the Mollisols, displaying granular or
crumb peds. Trace fossils of cicadas (Feoichnus, Monesichnus, Naktodemasis), dung
beetles (Coprinisphaera, Eatonichnus, Chubutolithes), bees (Palmiraichnus,
Elipsoideichnus, Uruguay, Corimbatichnus, Rosellichnus), sphinx moths (Teisseirei),
ants (Attaichnus), termites (Krausichnus), and cleptoparasites and detritivores
(Tombownichnus, Lazaichnus) appeared or diversified during this revolution that led
to the establishment of the Coprinisphaera Ichnofacies. By the Oligocene, in
closed-forest environments, also appears the first assemblage dominated by termite
and ant trace fossils (Termitichnus, Fleaglellius, Vondrichnus, Krausichnus), com-
prising the Termitichnus Ichnofacies. The Coprinisphaera revolution is followed
during the Neogene by stasis that was interrupted by the occasional appearance of
new trace fossils of the same, previously mentioned groups of insects. In contrast
with older ichnofacies (Scoyenia and Camborygma), which are recorded in flooded
or high water table paleosols, the younger ichnofacies (Celliforma, Coprinisphaera,
and Termitichnus) reflect paleosols with lower water tables. It is as if the terrestrial
ichnofaunas would have shift to drier environments, favored by morphological and
behavioral adaptations of their producers.
The ichnofacies model for paleosols is an ongoing approach that still has weak
points, all of which arise in the present incompleteness of the trace-fossil record.
For developing a stronger foundation, additional documentation will be required,
and surely it will be a future task of ichnologists to complete and strengthen the
models. There are about 166 case studies (25 for the Paleozoic, 39 for the Mesozoic,
and 102 for the Cenozoic), many of which are understudied or involve few trace
fossils that presently are insufficient to resolve the entire scenario. Despite this, the
model is a very promising one, and currently allows us to recognize within the fog
of incompleteness, the silhouettes of five paleosol ichnofacies, each one including a
number of ichnoassemblages that show sharply or broadly defined paleoenviron-
mental significance. After the novel approach presented in this chapter, the paleosol
ichnofacies model also provides an evolutionary scenario to track the major changes
in the history of terrestrial life.
Acknowledgements We thank Conrad Labandeira and Nicholas Minter for improving the origi-
nal manuscript, and Gabriela Mángano and Luis Buatois for fruitful comments and also for invit-
ing us to contribute to this book. The present research was supported by grant PICT 07/1972 and
2012/022 of the FONCYT of Argentina to J.F.G.
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Chapter 14
The Cenozoic Radiation of Mammals
Verónica Krapovickas and Sergio Vizcaíno
14.1 Introduction
Mammals are at present among the most successful vertebrates on Earth. On land
they inhabit almost all the habitats from the desert to the poles and have diverse
locomotor and dietary niches and size ranges. The Mesozoic record of mammals
spans about 155 Ma (the earliest mammals are known from the Late Triassic), more
than twice the duration of the entire Cenozoic Era (Luo 2007). Even though their
fossil record is relatively scarce, Mesozoic mammals were diverse and had assorted
biologic roles (Luo 2007). The end-Cretaceous mass extinction event that elimi-
nated non-avian dinosaurs and many other mostly marine taxa has been proposed
to having triggered the evolutionary radiation of Cenozoic mammals (Archibald
and Deutschman 2001). Following e.g. Alroy (1999), Benton (1995), and Foote
et al. (1999), among others, most mammalian orders originated and diversified in a
short period of time (10 Mya) soon after the Cretaceous/Tertiary (K/T) boundary.
Studies based on molecular data introduced two other models for the diversifica-
tion on mammals: the long-fuse and the short-fuse models. The first model postu-
lated an extended lag for the radiation of mammals as orders originated in the
Cretaceous but diversified after the K/T transition (e.g. Springer et al. 2003). The
second model proposed that the origin of orders and their diversification occurred
V. Krapovickas (*)
IDEAN-CONICET, Departamento de Ciencias Geológicas, FCEN, Universidad de
Buenos Aires, Ciudad Universitaria, Pabellón 2, Ciudad Autónoma de Buenos Aires
C1428EHA, Argentina
S. Vizcaíno
CONICET, División Paleontología Vertebrados, Museo de La Plata, Unidades de
Investigación Anexo Museo, FCNyM-UNLP, 60 y 122, B1900FWA LA Plata, Argentina

DOI 10.1007/978-94-017-9597-5_14
372 V. Krapovickas and S. Vizcaíno
well back into the Cretaceous (e.g. Springer 1997; Kumar and Hedges 1998).
Recent studies integrating morphologic and molecular data proposed that the ori-
gin of extant orders of mammals occurred during the Late Cretaceous and major
diversification occurred during and after the early Eocene, remaining the diversifi-
cation rates unchanged after the K/T transition (Bininda-Emonds et al. 2007). Up
to now, fossil evidence supporting the previous models was restricted to body fos-
sils. However, although even more sparse, the Mesozoic footprint record of early
mammals may provide further evidence.
In this contribution we briefly summarize the Mesozoic and Cenozoic record of
footprints attributed to mammals worldwide. Also, we address in more detail the
evolutionary implications of the Cenozoic ichnologic record of South American
mammals. Regarding the latter, we will first analyze chronologic and geographic
changes in the composition of the mammalian ichnofaunas and compare them with
those of body fossils. Second, we will consider the paleoenvironmental distribution
of mammalian footprints and their paleoecological significance.
14.1.1 Mammalian Ichnology Before the K/T Transition
The Mesozoic record of mammals is mostly composed of isolated teeth; postcranial

skeletons and skulls are rare (Kielan–Jaworowska et al. 2004). This fact complicates
in-depth evaluation of morphologic variations of footprints produced by distinct
early mammalian lineages, making the assignment to putative producers of
footprints vague. Distinguishing footprints assignable to mammals from other taxa
of the mammalian evolutionary line as nonmammalian cynodonts and even basal
synapsids is problematic (e.g. Schultz-Pittman et al. 1996; Lockley et al. 2004).
One of the most remarkable footprints known from the Mesozoic is Ameghinichnus
patagonicus, from the Middle Jurassic La Matilde Formation of Santa Cruz
Province, Argentina. They are one of the unquestioned examples of footprints
attributed to early mammals. Casamiquela (1964) proposed a mammalian
trackmaker due to the morphology of the manus and pes prints and the galloping
and hopping gaits evidenced on the trackways. This author inferred that the
trackmaker was a quadrupedal animal with similarly-shaped, pentadactyl manus
and pes, with the manus being slightly smaller than the pes. The impressions of
manual and pedal digits II–IV are of similar length, and those of digits I and V are
slightly shorter. The metacarpal/tarsal and phalangeal pads are well preserved, and
there are no claw marks (Casamiquela 1964). The trackways record a walking gait,
with the manus-pes sets in an alternate position in relation to the midline. In contrast,
hopping trackways have manus-pes sets grouped at relatively the same position in
relation to the midline (Casamiquela 1964). Most authors supported Casamiquela’s
(1964) identification of the Ameghinichnus trackmaker as an early mammal (e.g.
Leonardi and de Oliveira 1990; Leonardi 1994; Rainforth and Lockley 1996; Martin
and Rauhut 2005; Rougier et al. 2007a, b). Casamiquela (1964) proposed
“pantotheres” as the mammalian producer, whereas Kielan-Jaworowska and
Gambaryan (1994) attributed Ameghinichnus to multituberculates.
14 The Cenozoic Radiation of Mammals 373
Similar tracks from other ichnofossiliferous localities worldwide have been

referred to Ameghinichnus, as it is the best defined ichnotaxa assigned to an early
mammal. Olsen (1980) described a series of ichnofossils from the Lower Jurassic
Newark Supergroup, later recognized as representing several species of
Ameghinichnus, which he ascribed to nonmammalian cynodonts, possibly
tritheledontids (Olsen et al. 2002). Footprints with hair impressions have been
recorded in the Triassic Stomberg group in Lesotho and inferred to have been
produced by nonmammalian cynodonts or early mammals (Ellenberger 1972, 1974,
1975). De Valais (2009) recognized the ichnotaxa Eopentapodiscus to be the only
synonymous with Ameghinichnus, after reviewing numerous icnotaxa
(Acropentapodiscus, Amphibiopodiscus, Aristopentapodiscus, Dinopentapodiscus,
Eoameghinichnus, Grypopentapodiscus, and Pseudameghinichnus) once attributed
to the ichnogenus from the Upper Triassic-Lower Jurassic Elliot Formation of
Lesotho (Ellenberger 1970), and accepted the assignment to Ameghinichnus given
by other authors to materials from the Towaco Formation (Lower Jurassic) of the
Newark basin, USA (Olsen and Rainforth 2001), and the Lower Jurassic of
Sołtyków, Poland (Gierliński et al. 2004).
Another ichnotaxon, Brasilichnium elusivum, attributed to Mesozoic early
mammals is recorded in numerous localities. This ichnotaxon was originally
described from the Botucatu Formation (Upper Jurassic-Lower Cretaceous) of
Brazil and interpreted as corresponding to small-sized mammals (Leonardi 1980,
1981; Fernandes and Carvalho 2008). Later, it was recorded in classical Lower
Jurassic outcrops of USA, including the Navajo Sandstone and the Moenave
Formation, and reinterpreted as possibly produced by advanced therapsids or early
mammals (Lockley and Hunt 1995; Lockley et al. 1998; Rainforth and Lockley
1996; Loope 2006). Other Cretaceous footprints of mammals have been reposted
from the Gate Formation (Late Cretaceous) of Alberta, Canada, and ascribed to
several mammalian trackmakers (Sarjeant 2000; McCrea and Sarjeant 2001).
At present, there is just one example of footprints attributed to marsupial
mammals, Duquettichnus kooli, described by Sarjeant and Thulborn (1986) from
the Lower Cretaceous of the Peace River Canyon, British Columbia. The footprints
record syndactyly of pedal digits II and III, supporting a marsupial origin.
Some of the most significant and better preserved ichnologic records of Mesozoic
mammals are from Gondwana, despite their relatively scarce body-fossil record
(e.g., Kota Formation, India; Yadagiri 1984). This differs from the body-fossil
record in boreal landmasses, where Mesozoic mammalian diversity and abundance
is much higher (Bonaparte 1986, 1990, 1995; Bonaparte and Kielan-Jaworowska
1987; Luo et al. 2002; Kielan–Jaworowska et al. 2004).
14.1.2 Mammalian Ichnology After the K/T Transition
The global ichnologic record of mammals is poor for the Paleogene, most likely
representing less than 17 % of the record for the entire Cenozoic (McDonald et al.
2007). Paleocene footprints are extremely rare, with just a few tracksites reported
worldwide (Hunt and Lucas 2007), all of them in North America (e.g. Peabody
1954; Johnson 1986; Lockley and Meyer 2000).
The only confirmed tracksite that contain Paleocene mammal footprints is from
Alberta, Canada. It corresponds to a trackway preserved on a fallen block from
strata belonging to the Porcupine Hills Formation at Calgary (McCrea et al. 2004).
The footprints were ascribed to Sarjeantipes whitea and interpreted as produced by
creodont mammals. In 1928, Rutherford and Russell unveiled Paleocene footprints
found near Red Deer, Alberta, attributed to a mammalian track maker (Rutherford
and Russell 1928; Russell 1930). McCrea et al. (2004) re-studied the material
suggesting a crocodilian track maker. It is notable that, even apparently depauperate,
the Paleocene tetrapod footprints have scarce mammals represented and is mostly
composed by amphibian, reptiles, and birds, in contrast with mammal footprints
represented during the rest of the Cenozoic (Hunt and Lucas 2008) that became
more frequent through the Eocene and Oligocene.
The most diverse and well-studied Eocene mammalian ichnofauna is from the
late Eocene of Texas (Sarjeant and Langston 1994). It records footprints attributed
to insectivoran, creodont, carnivore, mesonychian, rodent, perissodactyl, and
artiodactyl trackmakers. The trans-Peco ichnofauna is one of the best recorded
examples of a Paleogene ichnofauna globally, even though it could be enlightening
to reevaluate this ichnassemblage in the light of new discoveries. Another remarkable
ichnofauna is from the Chuckanut Formation of Northern Washington, USA
(Mustoe 2002). This ichnofauna is mainly composed of multiple bird tracks and
also record remarkable mammal footprints attributed to extinct lineages, such as
Pantodonta or Dinocerata and early equids or tapiroids. Other records are from
Europe (e.g. Desnoyers 1859; Ellenberger 1980), Asia (West et al. 1983; Abbassi
and Lockley 2004; Ataabadi and Khazaee 2004), North America (e.g. Greben and
Lockley 1992; Lockley and Hunt 1995; Moussa 1968), and South America (e.g.
Noblet et al. 1995).
One of the best preserved and abundant mammalian footprints corresponds to the
lower Oligocene of southeastern France (Demathieu et al. 1984; Costeur et al.
2009). It records abundant perissodactyl (early Rhinocerotids) and artiodactyl
(Lophiomerycids and/or Entelodonts) track and trackways and a footprint attributed
to a mustelid-like carnivore. Some other examples are recorded from additional
localities of Europe (e.g. Tobein 1952; Casanovas-Cladellas and Santafé-Llopis
1982; Astibia et al. 1994; Prats and López 1995; Murelaga et al. 2000; Uchman
et al. 2004), North America (e.g. Nixon 1991; Lockley and Hunt 1995; Terry and
Wells 1995), and South America (e.g. Noblet et al. 1995).
For the Neogene the number of recorded tracksites grows exponentially (for
more detail, see McDonald et al. 2007). Herein, we briefly highlight some of the
classic outcrops documenting relevant mammal footprints. The research performed
in these localities has documented a wide variety of mammal footprints and
established the most used ichnotaxonomy of Cenozoic tracks and trackways. This is
the case of the work of Vyalov (1965, 1966) who reported numerous bird and
mammal footprints from the Miocene of Ukraine. The mammalian footprints were
ascribed to numerous ichnotaxa assigned to canids (Bestiopeda gracilis), felids
(Bestiopeda bestia, Bestiopeda sanguinolenta), artiodactyls (Pecoripeda amalphaea,

Pecoripeda dicrocervoides, Pecoripeda djali, Pecoripeda gazelle, Pecoripeda
satyri), and perissodactyls (Hippipeda aurelianis). The Miocene of the Romanian
Carpathians records also a diverse and excellent preserved avian and mammalian
ichnofauna (Panin and Avram 1962; Panin 1965). Mammals are represented by
canids (Canipeda longigriffa), felids (Felipeda felis, Felipeda lynxi), artiodactyl
(Pecoripeda gazelle), and proboscidean (Proboscipeda enigmatica) footprints.
Other significant records are known from Europe (e.g. Pérez-Lorente et al. 1999;
Astibia et al. 2007; Fornós et al. 2002), North America (e.g. Robertson and Sternberg
1942; Scrivner and Bottjer 1986; Lockley and Hunt 1995; McNeil et al. 1999; Lucas
2007), Oceania (e.g Carey et al. 2011), Africa (e.g. Leakey and Harris 1987; Robert
et al. 2008; Scott et al. 2009), and South America (e.g. Aramayo and Manera de
Bianco 1987a, b). Globally, the Neogene mammalian record is dominated by
ungulate footprints as perissodactyls, artiodactyls, and proboscideans, followed by
carnivores as felids and canids.
Reviews of Cenozoic vertebrate ichnology are scarce. The contributions of
Leonardi (1994), McDonald et al. (2007), Hunt and Lucas (2007), and Lucas (2007)
are the starting point to deeper ichnologic analyses, not only compiling bibliography,
but also evaluating the biologic, ethologic, and ecologic information provided by
footprints in the framework of the evolutionary history of their producers. Hopefully,
identifying the problems and promises of the mammalian paleoichnologic record of
each continent will help to develop a research program that will permit global
comparisons.
14.2 The South American Case Study
The geologic history of the main continental land masses has influenced the evolu-
tion and dispersal of their mammal faunas, resulting in radical differences among
them. Thus, from many aspects, direct comparisons of mammalian communities
between continents are a difficult task. For instance, the magnitude of the disparity
achieved is so that the faunas are not used for global age correlations and particular
land-mammal-age systems are used for different continental land masses (see
Woodburne 2006 for North America; Lindsay 1997 for Eurasia; Megirian et al.
2010 for Australia; and Flynn and Swisher 1995 for South America).
The long-term isolation of the South American continent for the greater part of
the Cenozoic produced a highly peculiar terrestrial biota, of which one of the best-
known components is its endemic mammalian fauna; probably the best fossil record
of mammals among southern hemisphere continents (Croft 2012). Several of the
main groups discussed below became partially or fully extinct and/or developed
morphologies many times outlying the shape ranges of their living closest relatives
(e.g., carnivorous marsupials, glyptodonts, and sloths among xenarthrans, native
ungulate groups). As a consequence, inferences about their paleobiology cannot
rely on simplistic actualistic reference to the biology of their living closest relatives.
In the last two decades, some effort has been made to overcome this situation
through the application of morphofunctional, biomechanical, and ecomorphologic
approaches. Overviews on this issue with respect to carnivorous marsupials are
available in Prevosti et al. (2012), on xenarthrans in Vizcaíno et al. (2008), Bargo
et al. (2012) and Vizcaíno et al. (2012b), and on ungulates in Cassini et al. (2012).
In a strict sense, the inferences resulting from those approaches are in many
instances difficult to verify. However, occasionally direct evidence comes to our aid.
This is the case when trace fossils, such as footprints attributable to these groups,
offer a tool to test functionally based hypotheses about locomotion and/or use of the
substrate, and to generate new hypotheses providing valuable paleobiologic
information on gait style and particular behavior (e.g., digging, scratching), which
may serve as proxies for the size of the putative trackmaker.
In most cases, footprints and bones are not associated in the same stratigraphic
units or facies, so then tracks become the primary source for recording the presence
of the producer at those levels. As they are normally preserved in situ in the facies
where they were produced, fossil footprints involve small temporal and spatial error
spans for the analysis of faunal taxonomic composition and distribution and imply
a close relationship between the footprints and the environments inhabited by the
producers. Thus, fossil footprints emerge as useful tools for paleocommunity
analysis and paleoenvironmental interpretations.
Fossil footprints in South America have been reported from different localities
and ages, including ichnologic associations preserved on a single surface or several
stratigraphically closely related surfaces. Leonardi (1994) summarized the
ichnologic fossil record in South America known two decades ago, including that of
Cenozoic mammals. In the present state of knowledge, the already mentioned
peculiarities of the South American mammalian fauna preclude general comparisons
of its paleoichnologic record with that from other continents.
In the second part of this chapter we endeavor to identify the problems and
promises of the mammalian paleoichnology record of South America. In doing so,
we update the information provided by Leonardi (1994) with new findings,
reconsidering the ages according to new chronologic information and reviewing the
potential producers. We also analyze chronologic and geographic changes in the
composition of the mammalian faunas expressed by the ichnologic record in com-
parison with that of body fossils. Finally, we evaluate the paleoenvironmental distri-
bution of mammalian footprints and their paleoecologic significance.
14.3 Faunistic Content Through the Cenozoic
Based mostly on the fossil record from Argentine lowland, high-latitude sites, dif-
ferent authors, such as Ameghino, Simpson, Reig, Ortiz-Jaureguizar, and Pascual,
among others (for a complete account, see Goin et al. 2012), have stressed the
episodic nature of the Cenozoic evolution of South American mammals, probably
as a consequence of several important temporal gaps in the knowledge of these
successions. The increasing amount of information collected in the last decades

from new high-latitude sites and in many other areas of South America, especially
the notable faunas uncovered from the Andes, produced at least two different and
complementary effects. At a broader regional scale, as the biochronologic sequence
of Neogene mammalian associations of southern South America became better
known, cycles or phases characterized by faunal turnovers and a distinctive
taxonomic composition were recognized, and paleoclimate and tectonics were
identified as major driving forces in addition to the intrinsic evolutionary patterns of
each mammalian lineage (Goin et al. 2012). At a continental scale, the actual
patterns are far more complex, with Cenozoic mammal faunas responding also to
biogeographic, sea level, ecologic, and environmental changes (Flynn et al. 2012).
Three of the five successive phases in South American mammalian evolution
recognized by Goin et al. (2012) involved the Cenozoic: the Early South American
(?latest Cretaceous-latest Eocene), characterized by major radiations within
Metatheria and Eutheria; the Late South American (early Oligocene-middle
Pliocene), with standardization of relatively few lineages among metatherians and
marked radiation of hypsodont types among South American native ungulates; and
the Inter-American (late Pliocene-Recent), produced by the mixture of North and
South American therian lineages, with progressive decline of native faunas.
According to these authors, the events that triggered these phases include global
cooling, full development of the Circumpolar Antarctic Current, and arrival of
platyrrhine primates and caviomorph rodents between the Early and Late South
American phases. The transition from the Late South American and the Inter-
American phases is marked by the Panamanian connection between the Americas
and the beginning of the Great American Biotic Interchange (GABI).
During the Cenozoic part of the Early South American phase, marsupials, xenar-
thrans, and native ungulates underwent their first Cenozoic main radiation.
Marsupials exploited a variety of adaptive zones and diets, including several small
to medium granivorous, insectivorous, and insectivorous–frugivorous groups (poly-
dolopimorphian; didelphimorphian and paucituberculates; Abello et al. 2012) and
medium to large carnivorous and carnivorous–omnivorous (i.e., sparassodonts).
Xenarthrans include the Cingulata and the Pilosa. The Cingulata are exemplified
by armadillos and glyptodonts, noted for the armor covering the head, body, and
tail. The Pilosa are composed of the Vermilingua—the anteaters—and the Tardigrada
or Folivora—living tree sloths and a much larger diversity of fossil sloths, including
the gigantic ground sloths. The scarce early record of xenarthrans suggests that most
of them inhabited regions not yet yielding fossils, an explanation widely accepted
by paleontologists, and consistent with the high diversity of xenarthrans in the
modern tropics and the correspondingly poor early Cenozoic record of that region
(Flynn et al. 2012).
According to Cifelli (1985, 1993), the ungulate groups endemic to South America
and the modern Ungulata (Perissodactyla and Cetartiodactyla) share a most recent
common ancestor or even form a single clade themselves (Cifelli 1985, 1993).
However, a recent phenomic approach split endemic South American ungulates
between Pan-Euungulata and Afrotheria (O’Leary et al. 2013). Endemic South
American ungulates fall into five groups: the astrapotheres, pyrotheres, notoungulates,
litopterns, and xenungulates (Simpson 1980; Marshall and Muizon 1988).
Astrapotheres (including trigonostylopids) were rhinoceros-like mammals (Cifelli
1985) found in deposits of Paleocene to Miocene age. Pyrotheres were elephant-
like, with tusks and bilophodont cheek teeth, and were never as diverse nor did they
cover as great a span of time as the astrapotheres. They are known only from the
middle Eocene through the late Oligocene. Notoungulates are by far the most
diverse and abundant lineage of South American ungulates (nearly 140 species in 13
families; Croft 1999), and include animals similar to rhinoceroses, hippopotamuses,
rabbits, and rodents; others do not closely resemble any living mammal. Litopterns
were the second most successful group of South American ungulates in terms of
diversity and longevity, spanning from the late Paleocene to the late Pleistocene
(Marshall and Cifelli 1990). They include forms similar to antelopes, horses, and
camels. Xenungulates are primitive, poorly known, tapir-like mammals, restricted
to the Paleocene deposits of Brazil and Argentina (Gelfo et al. 2008).
The taxonomic and ecologic shift that marked the passage to the Late South
American phase (Goin et al. 2012) included the arrival of caviomorph rodents and
platyrrhine primates in South America, the last records of marsupials sensitive to
low temperatures, the beginning of a rapid diversification of the “shrew opossums”
(Paucituberculata), the radiation of large modern borhyaenoids (Sparassodonta),
and the origins of the Argyrolagoidea. Among native ungulates, the diversity of
hypsodont notoungulates increased after the early Oligocene, and low-crowned,
bunodont ungulate types became restricted to the lower latitudes; Astrapotheria and
Pyrotheria developed their larger forms during the Oligocene-Miocene (Vizcaíno
et al. 2012a), disappeared by the middle late Miocene, and, by the Miocene, some
lineages developed limb specialization convergent on that of equids (e.g., in proter-
otheriid Litopterna).
According to the most recent findings (Antoine et al. 2012), the oldest records of
caviomorph rodent come from the late middle Eocene of Perú. Caviomorphs are
among the most abundant taxa in every post-Oligocene fauna and soon after their
arrival they radiated in a series of lineages that persist up to the present day, evolving
some fossil forms that weigh in excess of a hundred kilograms (Vizcaíno et al.
2012a). The oldest record of a platyrrhine primate in South America is Branisella
from the late Oligocene of Salla, Bolivia (Kay et al. 1998). Fossil South American
primates are rare arboreal forms, mostly no larger than 10 kg, but at least one taxon
reached 20 kg (Halenar 2011).
Goin et al. (2012) described the Inter-American phase as a series of pulses of the
so called Great American Biotic Interchange (Stehli and Webb 1985), possibly
reflecting glacial versus interglacial conditions in the Northern Hemisphere. The
first North American forms to arrive in South America included cricetid rodents,
procyonid, and mustelid carnivorans, and tayassuid artiodactyls. During the early
Pleistocene gomphotheriids (proboscideans), camelids and cervids (artiodactyls),
equids and tapirids (perissodactyls) are recorded in South America for the first time,
in addition to several carnivoran lineages (felids, canids, and ursids).
As mentioned above, there is a significant biogeographic differentiation at a

continental scale that was considered recently by Goin et al. (2012) and Flynn et al.
(2012). Both contributions stressed the significance of the evidence provided by
sites of different ages along the Andes, especially during Goin’s et al. earlier phases
of the Cenozoic. For instance, the latest Eocene-early Oligocene faunas of central
Chile and Patagonia differ in taxonomic representation: the Chilean Tinguirirican
fauna lacks hegetotheres but possesses rodents, and a number of species are distinct
between the two areas (Croft et al. 2008; Flynn et al. 2003; Flynn et al. 2012). Also,
the middle Miocene fauna of Quebrada Honda in southernmost Bolivia is more
similar to the slightly older high-latitude fauna of Collón-Curá in Patagonia than to
the contemporaneous low-latitude fauna of La Venta of Colombia (Croft 2007; Goin
et al. 2012). However, there is also evidence of some late biogeographic differences
during the Inter-American phase, such as the differential distributions within ground
sloths (e.g. Eremotherium and Megatherium; Cartelle and De Iuliis 1995) and
proboscideans (Cuvieronius and Notiomastodon; Mothé et al. 2012 and references
therein), among others.
14.4 Methods
To analyze the composition of the Cenozoic South American mammal paleocommu-

nities, we compiled incidence (= presence/absence) data for ichnologic assemblages
containing diverse footprints and dismissed those represented only by one ichnologic
morphotype. We conducted a cluster analysis to explore trace-fossil assemblage
groupings (Q.mode) and used the Jaccard similarity index for binary data. The anal-
ysis was performed in the statistical package PAST (Hammer et al. 2001).
In the second analysis we compiled a dataset of environments of footprint pres-
ervation by age, for all assemblages with information available in the literature. The
environmental data were classified in seven categories: (1) floodplain, (2) exposed
sandbar, (3) marginal lacustrine, (4) wet interdune, (5) dune, (6) salt flat, and (7)
marginal marine. The number of occurrences of each category per age was plotted
in histograms.
14.5 The Cenozoic Track Record of South American

Mammals
14.5.1 The Footprints of the San Jeronimo Group, Peru
The San Jerónimo Group (early Eocene-early Oligocene) of Peru (Carlotto 2006)
holds relevant ichnologic assemblages misinterpreted for nearly a century as
dinosaurian in origin, and consequently their stratigraphic levels were taken as
Cretaceous in age.
Fig. 14.1 Map indicating mammal-bearing fossil footprint localities in South America
Originally, Gregory (1916) described fossil footprints from the Cuzco valley,
Peru, preserved near the base of the “Couches Rouges” (San Jerónimo Group).
Later, Noblet et al. (1987, 1995) described other footprint assemblages from three
localities, two near Cuzco city (Kayra and Paruro) and the third at Langui near the
city of Sicuani (Fig. 14.1). During the past decade, the age of the San Jerónimo
Group was established as early Eocene to early Oligocene by stratigraphic studies
and analysis of volcanic tuff and fission tracks (Carlotto 1998, 2002, 2006; Carlotto
et al. 1995, 2005; Fornari et al. 2002). Subsequently, the producers of the footprints
described by Noblet et al. (1987, 1995) were reinterpreted by Lockley et al. (1999)
as notoungulates or litopterns and by Carlotto (2011) as birds, although the material
was never examined in detail. The material first studied by Noblet was illustrated by
Leonardi (1994) and is that discussed herein.
The San Jerónimo Group is divided in two units, both preserving fossil footprints.
The footprints from Paruro occur in the lower-middle Eocene Kayra Formation
(lower unit). The upper Eocene-lower Oligocene Soncco Formation (upper unit)
records two different assemblages, one at Kayra and the other at Langui, near
Sicuani (Fig. 14.1).
14.5.1.1 Paruro Footprints
The footprint assemblage comprises three morphotypes preserved on a single sur-

face. The best preserved morphotype consists on homopod tridactyl footprints of a
rodent-like mammal of small size (approximately 6 cm long and 3 cm wide), with
long pointed digits and a plantar pad of at least one third of the total length of the
footprint (Fig. 14.2c) (see plate XXXIV, Fig. 4 in Leonardi 1994). The second foot-
print morphotype has a poorly defined morphology and is not well illustrated. It
consists of at least a tetradactyl footprint of an apparently homopod quadruped (see
plate XXXIV, Fig. 2 in Leonardi 1994). The footprints are approximately 7 cm long
and 7 cm wide. The material corresponds to three trackways and an isolated
footprint. Leonardi (1994) described it as footprints of the size of a cat or a small
dog. One poorly defined trackway of a bipedal and at least tridactyl animal repre-
sents the third morphotype preserved at Paruro (Fig. 14.2a and b) (see plate XXXIV,
Figs. 1 and 3 in Leonardi 1994). It has marked toe drags that gave the impression to
Noblet and Leonardi that it was produced by a sick or very old individual.
14.5.1.2 Kayra Footprints
The ichnologic record of Kayra involves an isolated tridactyl footprint, 13 cm long.

It represents the impression of a mesaxonic foot with three digits directed forward.
The central digit is wider and longer than the lateral digits, though it is incomplete,
and lateral digits are pointed (Fig. 14.2e). The plantar pad is wide and is as long as
the central digit. The footprints are interpreted here as produced by a toxodontid
notoungulate.
14.5.1.3 Langui Footprints
The footprints of Langui are preserved on a single sub-vertical surface with at least
three different morphologic types. An unclassifiable fourth morphotype was
mentioned by Leonardi (1994). Unfortunately, there is no illustration of the footprint
and for this reason we won’t consider it here. The largest morphotype is represented
by one trackway with two tridactyl footprints, approximately 20 cm long with a
central digit longer than the lateral digits, possibly attributable to phorusrhacids (see
plate XXXV, Figs. 4 and 1 in Leonardi 1994). The second morphotype consists of
one trackway with seven tridactyl footprints, 10–13 cm long, of a quadruped homo-
pod mammal with hoof-like toes (Fig. 14.2d) (see plate XXXV, Figs. 1, 2 and 6 in
Leonardi 1994). The footprints are assigned to Macrauchenichnus isp. and
Fig. 14.2 Footprints of the San Jeronimo Group, Peru. Paruro tracksite photographed by Noblet
(a) and the tracksite today (b); (c) homopod tridactyl footprints, Paruro. Scale bar = 3 cm; (d)
Tridactyl quadruped homopod footprint, Langui; (e) isolated tridactyl footprint, Kayra
interpreted as possibly produced by toxodontid notoungulates or macraucheniid

litopterns. The morphotype III consists of a trackway of a quadrupedal, rodent-like
mammal. The manus footprints are overprinted by the well-defined pes footprints.
Pes footprints are tridactyl with elongated digits and a plantar pad (see plate XXXV,
Figs. 1 and 3 in Leonardi 1994). The footprints are approximately 7 cm long, and
the morphology of the pes footprint and the trackway pattern are highly comparable
with those of the small heteropod footprints described by Krapovickas et al. (2009a,
b, c; see below), suggesting that they were produced by a rodent-like typothere
notoungulate or a caviid rodent.
14.5.2 The Footprints of the Vinchina Basin, Argentina
The Vinchina Basin of La Rioja Province, northwestern Argentina (Fig. 14.1), is a

thick continental succession deposited under a wide variety of depositional systems
(i.e., fluvial, lacustrine, and eolian) that bears diverse and exceptionally well-
preserved trace-fossil assemblages. Recent interpretations of the stratigraphy of the
basin (Ciccioli et al. 2010, 2014; Limarino et al. 2010) divided the sedimentary
succession into five formations, from bottom to top, Puesto La Flecha (Eocene),
Vallecito (Oligocene-early Miocene), Vinchina (early Miocene), Toro Negro (late
Miocene-Pliocene), and El Corral formations (Pliocene). The body-fossil record for
these units (e.g., Turner 1964; Ramos 1970; De Iuliis et al. 2004; Rodríguez Brizuela
and Tauber 2006) is fragmentary and scarce. Conversely, fluvial and lacustrine
facies of the basin host a rich invertebrate and vertebrate ichnofauna (Frenguelli
1950; Bonaparte 1965; Tripaldi et al. 2001; Krapovickas et al. 2009a; Melchor et al.
2010, 2012; Krapovickas and Nasif 2011). The most remarkable assemblages of
mammal footprints are recorded in the Vinchina Formation, at Quebrada de la Troya
(lower portion) and Quebrada del Yeso, and Toro Negro Formation at Quebrada de
la Troya (upper portion) (Fig. 14.1).
14.5.2.1 Lower Quebrada de la Troya Footprints
The footprints of Quebrada de la Troya, nearby the town of San José de Vinchina,
are preserved in floodplain deposits of meandering fluvial systems of the Vinchina
Formation. One of the most peculiar footprints is Tacheria troyana, approximately
10 cm long, produced by tetradactyl homopod quadrupeds with long digits and well
developed plantar pads (Fig. 14.3f). The trackmaker was interpreted by Krapovickas
and Nasif (2011) as a dinomyid caviomorph rodent or a close relative. There are
also preserved small tridactyl footprints of a homopod rodent-like mammal, possi-
bly a caviomorph rodent or typothere notoungulate (Fig. 14.3h); tridactyl homopod
footprints with short broad digits and blunt tips assigned to Macrauchenichnus isp.
and interpreted as toxodontid notoungulates or macraucheniid litopterns (Fig. 14.3b
and d); large tridactyl footprints of rheiformes; footprints of shorebirds assigned to
Gruipeda isp.; and a small tridactyl to tetradactyl footprint with pronounced scratch
marks (Krapovickas et al. 2009b, c)
14.5.2.2 Quebrada del Yeso Footprints
In 1950 Frenguelli mentioned a fossil footprint assemblage preserved in a continen-

tal succession at Quebrada del Yeso, between the towns Villa Castelli and Jagüe, La
Rioja Province, Argentina. He described Venatoripes riojanus, a trackway of large
manus-pes sets interpreted originally as produced by a large-size reptile, since the
bearing levels were considered early Permian in age (see Fig. 2 in Frenguelli 1950).
Later, the succession was interpreted as deposited during the Miocene–Pliocene and
the tracks were reinterpreted as produced by xenarthran megatheroids (Turner 1960;
Bonaparte 1965). Recently, the outcrops were relocated as close to the top of the
Vinchina Formation, suggesting an early-middle Miocene age for the footprint
assemblage (Limarino, C.O. pers. comm.). Other elements of the assemblage were
documented by Bonaparte (1965), interpreting them as produced by several avian
and mammalian taxa, such as rheiformes, charadriiformes, ralliformes, and
macraucheniid litopterns (see Fig. 1 in Bonaparte 1965).
Fig. 14.3 Footprints of the Vinchina Basin, Argentina. (a) General view of the main tracking
surface of Toro Negro; (b) General view of the main tracking surface of Lower Vinchina; (c) Small
heteropod footprints, Toro Negro; (d) Macrauchenichnus isp. Scale bar = 3 cm, Lower Vinchina;
(e) Oval impressions, Toro Negro; (f) Tacheria troyana, Lower Vinchina. Scale bar = 3 cm; (g)
Macrauchenichnus rector, Toro Negro; (h) Small tridactyl footprints, Lower Vinchina. Scale
bar = 3 cm
14.5.2.3 Upper Quebrada de la Troya Footprints
The Miocene deposits of the Toro Negro Formation at Quebrada de la Troya, La

Rioja Province, Argentina, host a rich vertebrate and invertebrate ichnofauna docu-
mented by Krapovickas et al. (2009a, b, c). The trace fossils are recorded from the
lower part of the Lower Member of the Toro Negro Formation. The vertebrate ich-
nofauna is the most diverse of the three ichnologic assemblages preserved in the
Vinchina basin and includes avian and mammalian footprints (Fig. 14.3a). The
mammalian footprints ascribed to Macrauchenichnus rector are interpreted as pro-
duced by medium sized macraucheniid litopterns (Fig. 14.3a and g), cf. Venatoripes
riojanus as large tardigrade mammals (Fig. 14.3a), kidney-like footprints as medium
to small tardigrades, oval impressions as proterotheriid litopterns (Fig. 14.3e), and
small heteropod footprint originally interpreted as hegetotheriid typotheres or cav-
iomorph rodents and herein as caviid caviomorph rodents (Fig. 14.3c). The avian
footprints (e.g., Fuscinapeda sirin) are interpreted as produced by shorebirds,
incumbent footprint as a large cursorial bird like rheiformes or phororhacids, and
slender anisodactyl footprints as perching birds. The tetrapod footprints and rare
Palaeophycus and Helminthopsis are preserved on the top of exposed sandbars of
anastomosing fluvial systems developed under a semi-arid climate. Crevasse-splay
deposits preserve abundant meniscate trace fossils and dwelling tubes as Taenidium,
Scoyenia, and Palaeophycus ascribed to the Scoyenia Ichnofacies.
14.5.3 The Footprints of the Santa Cruz Formation, Argentina
The Santa Cruz Formation (late early Miocene) is a continental succession exposed
from the Andean Precordillera to the Atlantic coast of southern Patagonia (Matheos
and Raigemborn 2012). Along the Atlantic coast, between the Río Coyle and Río
Gallegos, Tauber (1997) defined two members, from base to top, the Estancia La
Costa and the Estancia La Angelina members. The basal portion of the Estancia la
Angelina Member records undetermined mammalian fossils footprints in the pres-
ent-day intertidal zone (Figs. 14.1 and 14.4d) (Raigemborn et al. 2012). These beds
represent distal-floodplain environments with the development of moderately well-
drained paleosols due to seasonal shifts between subhumid and semiarid climatic
conditions (Krapovickas 2012).
14.5.4 The Footprints of the Mariño Formation, Argentina
The Mariño Formation (Miocene) records a trackway exposed on the margins of the
Provincial Road 17 at Cacheuta, Mendoza Province, western Argentina (Fig. 14.1).
The footprints consist of a single trackway with oval footprints, approximately 3 cm
long, preserved in eolian strata deposited under arid to semi-arid climatic conditions
Fig. 14.4 Footprints of the Mariño and Santa Cruz formations, Argentina. (a) Tridactyl footprints
on fluvial wave-rippled fine-grained sandstone strata, Salagasta; (b) detail of the tridactyl
footprints; (c) Oval footprints, Cacheuta; (d) undetermined mammalian footprints, Santa Cruz
(Zavattieri et al. 2001). The footprints correspond to a quadruped monodactyl

homopod trackmaker, most-likely a proteroteriid litoptern (Fig. 14.4c). Other expo-
sures of the Mariño Formation at Salagasta, Mendoza Province (Fig. 14.1), record
tridactyl footprints on fluvial wave-rippled fine-grained sandstone strata (Ahumada
2004). The footprints were most-likely produced by toxodontiid notoungulates or
macraucheniid litopterns (Fig. 14.4a and b).
14.5.5 The Footprints of Las Flores Formation, Argentina
At Puchuzun at the Calingasta valley, San Juan Province, western Argentina (Fig. 14.1),
the Las Flores Formation of late Miocene age holds a diverse ichnologic assem-
blage representing mostly footprints of aquatic and terrestrial birds and a few mor-
photypes of medium and large size mammals. They consist on bird footprints of
Anseriformes, Charadriformes, and Phoenicopteriformes, in addition to footprints
of hegetotheriid notoungulates and a large ground sloth preserved in marginal
lacustrine facies (Contreras 1996, 2006; Contreras and Peralta 2011).
14.5.6 The Footprint of Quebrada del Jarillal Formation,

Argentina
The Cenozoic succession exposed at Sierra de Mogna, San Juan Province, Argentina
(Fig. 14.1) records a set of manus-pes footprints preserved on levels of the Quebrada
del Jarillal Formation (middle-upper Miocene). The footprints were originally
described and illustrated by Casamiquela in Cuerda et al. (1984) (see Plate XV,
Fig. 2 and Plate XVI, Figs. 1 and 2 in Cuerda et al. 1984) and later also documented
by Leonardi (1994, plate XIX, Fig. 11). Krapovickas and Nasif (2011) assigned the
material to Tacheria troyana and interpreted it as produced by a dinomyid
caviomorph rodent or a close relative.
14.5.7 The Footprints of the Tertiary of the Puna, Argentina
Near the border between the provinces of Salta and Jujuy, Argentina, at Catua
(Fig. 14.1), in 1980 R.N. Alonso found mammalian footprints, possibly rodents,
on eolian dune cross-beds of possible Miocene age (Leonardi 1994). In addition,
R.N. Alonso also discovered a small isolated footprint cast of a mammalian track-
maker, probably a rodent, preserved on salt flats deposits at Salar del Rincón, Salta
Province, Argentina (Fig. 14.1). The bearing levels possibly correspond to the
Pliocene Trinchera Formation (Leonardi 1994).
14.5.8 The Footprints from the Río Negro Formation
Extensive sea cliffs of northeast Patagonia, Río Negro Province, expose an upper
Miocene-lower Pliocene succession corresponding to the Río Negro Formation
(Fig. 14.1). The succession corresponds to a complete suite of eolian and eolian-
related sub-environments (Zavala and Freije 2001). The Río Negro ichnologic
record from the margins of the Río Negro River at Carmen de Patagones was origi-
nally described by Casamiquela (1974), who identified novel ground sloth bipedal
footprints assigned to Megatherichnum oportoi and interpreted as produced by a
megatheriid close to Megatherium. Later, Angulo and Casamiquela (1982) studied
the footprint record of the strata cropping out at the cliffts of the San Matías Gulf
at several localities. The most diverse ichnologic assemblage is the one recorded at
Balneario El Cóndor, including footprints of megatheriid ground sloths of large and
medium size assigned to Megatherichnum oportoi and Falsatorichnum calceocan-
nabius, respectively (see Plate II in Angulo and Casamiquela 1982); Porcellusignum
consulcator, corresponding to heteropod footprints with a tetradactyl manus and
tridactyl pes most-likely produced by capybaras (see Plate II in Angulo and
Casamiquela 1982); Macrauchenichnus rector that consist of tridactyl blunt
Fig. 14.5 Footprints of the Río Negro Formation. (a) Megatherichnum oportoi; (b) Large slab
containing footprints of marsupial saber-toothed tiger; (c) Macrauchenichnus isp.; (d) Schematic
drawing of Macrauchenichnus rector, scale bar = 10 cm
footprints produced by macraucheniid litopterns (Fig. 14.5d) (see Plate VIII, IX,
and X in Angulo and Casamiquela 1982); and Caballichnus impersonalis, subcir-
cular footpritns of monodactyl trackmakers interpreted as equids (see Plate V, VI,
VII in Angulo and Casamiquela 1982). These authors also mentioned the presence
of Macrauchenichnus rector in other localities in the studied area (Balneario La
Lobería, Balneario La Boca, and Pozo Salado). Because of the poor quality of pres-
ervation of the material used to erect Falsatorichnum calceocannabius and
Caballichnus impersonalis, the validity of both ichnotaxa has been questioned
(e.g., Melchor 2009). Moreover, the extremely simple morphology (subcircular
outline) together with the lack of preservational details expressed on C.
impersonalis make its assignation to equids questionable. This would imply extend-
ing the record of equids in South America to the late Miocene prior to the GABI.
Aramayo et al. (2004, 2007) recorded additional material from the area. These
authors reported footprints of ground sloths of large size assigned to Megatherichnum
oportoi (Fig. 14.5a) and of medium size assigned to cf. Milodontidichnum isp.,
similar in size and shape to Falsatorichnum calceocannabius, but interpreted as a
mylodonthid . Other material corresponds to capybara footprints (cf. Porcellusignum
isp.), two ungulate footprints of proteroteriid litopterns and undetermined ungu-
lates, pentadactyl footprints with claw marks most-likely produced by the marsu-
pial saber-toothed tiger (Fig. 14.5b), tridactyl footprints of large size produced by
phorusrhacids, and two types of tridactyl bird footprints, ones of shorebirds and the
others are webbed footprints of flamingos. Aditional material has been recently
documented by Carmona (pers. com., 2012; Fig. 5c). The tridactyl footprints highly
resemble Macrauchenichnus isp. from the Vinchina Formation (Fig. 14.3d).
14.5.9 The Footprints of Antofagasta, Chile
Plio-Pleistocene marine strata that bear fossil footprints preserved in coastal dunes
facies are exposed near Antofagasta, Chile, 3 km from the recent marine coast
(Fig. 14.1) (Casamiquela and Chong Diaz 1975). The footprints correspond to a
quadruped monodactyl mammal of medium size. The impression of forefoot and
hind foot are sub-equal and oval and are more deeply impressed on the anterior
margin. Casamiquela and Chong Diaz (1975) interpreted that the footprints were
made by equids, although the morphology of the footprints does not correspond
exactly with recent equids.
14.5.10 The Footprints of Claromecó, Argentina
In the present-day intertidal zone of the Caracolero beach at Claromecó, Buenos

Aires, Argentina (Fig. 14.1), deposits of late Pleistocene age record fossil footprints
most likely produced by a large sized mylodontid ground sloth (Azcuy et al. 2011).
The footprints have a very different outline and differ from other footprints
interpreted as produced by large ground sloth (e.g. Venatoripes riojanus,
Megatherichnum oportoi, Neomegatherichnum pehuencoensis). They consist of a
plantigrade pes track with a general oval outline, with four digits and a prominent
sole print. The tracks are approximately 60 cm long and 40 cm wide. The digit III
impression is the longest and has a prominent claw mark, the lateral digits (II and
IV) are subequal in length and show a sharp ending, and laterally to digit IV is the
small impression of digit V.
14.5.11 The Footprints of Monte Hermoso, Argentina
Casamiquela (1983) described ground sloth footprints from fluvial plain deposits of
late Pleistocene age on the Atlantic coast at Monte Hermoso, Buenos Aires.
Argentina (Fig. 14.1). The footprints correspond to Iribarnichnum megamericanum
and Acunaichnus dorregoensis, interpreted as megatheriids and mylodontid ground
sloths, respectively.
14.5.12 The Footprints of Pehuen-Có, Argentina
The continental succession exposed on the Atlantic coast of southern Buenos Aires
Province, Pehuen-Có (Fig. 14.1), records the most diverse association of Cenozoic
mammalian footprints from South America (Aramayo and Manera de Bianco
1987a; b; 1996; 2009). The footprints are preserved in the present-day intertidal
zone in strata of late Pleistocene age and represent a wide variety of mammals, such
as large ground sloths (Fig. 14.6a, b and e) (the megatheriid Neomegatherichnum
pehuencoensis interpreted as produced by Megatherium and the mylodontid
Milodontichnum rosalensis), macraucheniid litopterns (Fig. 14.6c) (Eumacrau-
chenichnus patachonicus), gomphotheres (Stegomastodonichnum australis), car-
nivorans (Pumaeichnum biancoi, Pehuencoichnus gracilis, Mustelidichnum
enigmaticum), camelids (Fig. 14.6d) (Lamaichnum guanicoe and Megalamaichnum
tulipensis), cervids (Odocoileinichnum commune), equids (Hippipeda isp.), glypto-
donts (Fig. 14.6f), bears, and caviomorph rodents, along with birds, such as phoe-
nicopteriforms (Phoenicopterichnum rector), tinamiforms, anseriforms, rheiforms,
and shorebirds. The trackway-bearing levels correspond to floodplain deposits
where most of the footprints are preserved on mudstone intervals with desiccation
cracks and ripple marks (Manera de Bianco and Aramayo 2004).
14.6 Faunistic and Biogeographic Similarity of Footprints

Assemblages
The cluster analysis of the mammalian taxa represented by the fossil footprint
record in South America shows a clearly identifiable change in diversity since the
Eocene to the Pleistocene. The main dichotomy separates the Eocene to Miocene
ichnologic assemblages from the Pliocene-Pleistocene ones. These branches
separate the assemblages in which the producers belong to lineages that under-
went diversification and ecologic specialization in isolation from other conti-
nents, from other assemblages including also representatives of North American
lineages (Fig. 14.7a).
Fig. 14.6 The footprints of Pehuen-Có, Argentina. (a) Neomegatherichnum pehuencoensis track-
ways; (b) Milodontichnum rosalensis trackways; (c) Tracking surface with numerous impressions
of Eumacrauchenichnus patachonicus; (d) Lamaichnum guanicoe; (e) Detail of the pes of
Neomegatherichnum pehuencoensis; (f) Isolated footprint of glyptodont
a b
Lower Troya
Lower Troya
Upper Troya
Upper Troya
Pehuen-Co
Pehuen-Co
Cond.-Lob.
Cond.-Lob.
Qda. Yeso
Qda. Yeso
Puchuzum
Puchuzum
El Condor
El Condor
Paruro
Paruro
Langui
Langui
0,96
0,84
0,72
0,6
Similarity
0,48
0,36
0,24
0,12
Fig. 14.7 Dendogram with the results of the cluster analysis, including the principal Cenozoic
mammalian ichnofaunas
The Eocene-early Miocene ichnologic assemblages mostly represent forms of

uncertain affinity. This is the case of rodent-like footprints identified as produced by
small caviomorph rodents or typothere notoungulates and footprints assigned to
small toxodontid notoungulates or macraucheniid litopterns. Also represented are
footprints of large dinomyids caviomorph rodents, large undetermined notoungu-
lates, and medium-size footprints of undetermined tetradactyl mammals (Fig. 14.8).
The rodent-like footprints of the Paruro assemblage may be highly significant.
This assemblage comes from the Kayra Formation. The base of this formation is
placed near the Paleocene-Eocene boundary and its limit with the unconformably
overlying Soncco Formation is at about 43 Ma (Carlotto 2006), constraining the
chronologic span of the formation to the early-middle Eocene. The recent discovery
of rodents in the middle Eocene of the Peruvian Amazonia (Antoine et al. 2012)
renders plausible a caviomorph affinity of the Paruro rodent-like footprints and
could reinforce the recent hypothesis of Antoine et al. (2012) about the southward
expansion of the earliest caviomorphs during the late middle Eocene–early
Oligocene period, from low (Peruvian Amazonia) to middle (central Chile) and then
to high latitudes (Patagonian Argentina). However, the stratigraphic information
provided in the report of the footprints (Noblet et al. 1995) indicates that they come
from the lower levels of the section, suggesting an early Eocene age. If this were
true, and further careful analysis confirms its caviomorph affinity, these footprints
would expand the early record of rodents in South America to the early Eocene.
There is a marked resemblance between the early Oligocene Langui ichnoas-
semblage of Peru and the early Miocene Lower Vinchina ichnoassemblage of north-
western Argentina (Fig. 14.7). The fact that the Lower Vinchina record is more
Fig. 14.8 Paruro footprints: (1) caviomorph rodents or typothere notoungulates, (2) undetermined
tetradactyl mammals. Kayra footprints, (3) toxodontid notoungulates. Langui footprints: (4)
toxodontid notoungulates or macraucheniid litopterns, (5) caviomorph rodents or typothere
notoungulates. Lower Quebrada de la Troya footprints, (6) Tacheria troyana, (7) caviomorph
rodents or typothere notoungulates, (8) Macrauchenichnus isp. Quebrada del Yeso footprints, (9)
Venatoripes riojanus, (10) macraucheniid litopterns. Upper Quebrada de la Troya footprints, (11)
caviomorph rodents, (12) cf. Venatoripes riojanus, (13) Macrauchenichnus rector, (14)
proterotheriid litopterns. Cacheuta footprints, (15) proterotheriid litopterns. Salagasta footprints,
(16) toxodontiid notoungulates or macraucheniid litopterns. Sierra de Mogna, (17) Tacheria
troyana. Catua footprints, (18) caviomorph rodents or typothere notoungulates. Salar del Rincón
footprints, (19) caviomorph rodents or typothere notoungulates. Puchuzun footprints (20)
hegetotheriid notoungulates, (21) ground sloth. Río Negro footprints, (22) Porcellusignum
consulcator, cf. Porcellusignum isp., (23) Megatherichnum oportoi, Falsatorichnum
calceocannabius, cf. Milodontidichnum isp., (24) Macrauchenichnus rector, (25) proteroteriid
litopterns, (26) marsupial saber-toothed tiger, (27) Caballichnus impersonalis. Antofagasta
footprints, (28) equids. Claromecó footprints, (29) ground sloth. Monte Hermoso footprints: (30)
Iribarnichnum megamericanum and Acunaichnus dorregoensis. Pehuen-Có footprints, (31)
Neomegatherichnum pehuencoensis, Milodontichnum rosalensis, (32) Eumacrauchenichnus
patachonicus, (33) glyptodonts, (34) Stegomastodonichnum australis, (35) Pumaeichnum biancoi,
Pehuencoichnus gracilis, Mustelidichnum enigmaticum, Bear footprints, (36) Lamaichnum
guanicoe, Megalamaichnum tulipensis, Odocoileinichnum commune, (37) caviomorph rodents,
(38) Hippipeda isp.
similar to an older assemblage located to the north (Langui), rather than younger
assemblages (middle-late Miocene) located in the same geographic area (Upper
Vinchina and Toro Negro), suggests that at least during the earliest Miocene these
areas were biogeographically connected and an isolation mechanism potentially
operated from the early Miocene onwards. These could be the result of similar cli-
matic conditions and the prevalence of similar faunas. However, it is not possible
yet to contrast the similarities of the Lower Vinchina ichnoassemblage with any
other ichnologic assemblage of the same age elsewhere.
Among the Miocene assemblages footprints assigned to ground sloth, macrauche-
niiid and proterotheriid litopterns, caviomorph rodents and rodent-like caviomorph
rodents or typothere notoungulates are well represented; footprints assigned to tox-
odontid notoungulates or macraucheniid litopterns were also present (Fig. 14.8).
Not surprisingly, the major similitudes between ichnoassemblages are among those
from the same age (middle-late Miocene) and geographic area (northwestern Argentina),
as is the case of the Upper Vinchina and Toro Negro ichnoassemblages (Fig. 14.7).
During this period, footprints of large mammals, such as ground sloths and
macraucheniids, are present for the first time, though they did not achieve the major
sizes reached during the Plio-Pleistocene. Proteroteriid footprints are also present
for the first time (Fig. 14.8). The middle Miocene ichnofaunas, as well as the osteo-
dentary faunas, mark the peak of known diversification achieved by mammals after
the arrival of primates and rodents but before the arrival of North American immi-
grants (Marshall and Cifelli 1990).
The late Miocene-Pliocene and late Pleistocene assemblages include footprints
of both native South American mammals and North American taxa arrived during
the GABI, recording a variety of ground sloths (Megatheriidae and Milodontidae),
macraucheniid and proteroteriid litopterns, hydrochoerid caviomorph rodents and
North American equids in both assemblages (Fig. 14.8). Footprints of other taxa are
recorded in a single assemblage; a carnivorous marsupial (Thylacosmilidae) is
recorded in outcrops of late Miocene-early Pliocene age of Río Negro, Argentina,
and glyptodonts, gomphotheriids (proboscideans), camelids, and cervids (artiodac-
tyls), and several carnivoran lineages (felids, ursids, mustelids, and bears) are repre-
sented in the late Pleistocene ichnoassemblages of Pehuen-có, Buenos Aires,
Argentina (Fig. 14.8).
Comparing our results with the faunal evolutionary phases proposed by Goin
et al. (2012) described above, the two major clusters roughly separate the faunas
corresponding to the Early and Late South American phases from the Interamerican
phase (Fig. 14.7). The dichotomy recorded within the first cluster described does
not fit with the separations between the Early and Late South American phases, but
there are several factors that may be affecting the results. First, only one lower
Eocene locality may represent the Early South American phase; however, this
Eocene locality clusters with the Oligocene and Miocene localities of the Late
South American phase. The main problem is the ambiguous affinity of the rodent-
like footprints (typothere/caviomorph), because the incorporation of caviomorph
rodents to the South American record was taken as one of the events that triggered
the Late South American Phase. When this footprint is removed from the analysis
(Fig. 14.7b), the Eocene locality remains separated from a cluster constituted by the
Oligocene and Miocene localities (Late South American phase). However, if the
potential affinity of the Paruro footprints is established by a powerful method of
trackmaker identification to caviomorphs, it would bring the beginning of the Late
South American phase backward to the early Eocene, even earlier than the earliest
record of body fossils (Antoine et al. 2012).
14.7 The Paleoenvironmental Distribution of Mammalian

Ichnofossils
South American fossil footprints are mostly recorded in fully continental sedimen-
tary systems, with just one example to date, recorded from marginal marine systems
corresponding to Plio-Pleistocene outcrops of Antofagasta, Chile (Fig. 14.9).
Fig. 14.9 Environmental distribution of Cenozoic mammalian footprints
Among the continental sedimentary systems preserving fossil footprints there are
fluvial, lacustrine, and eolian deposits. Fossil footprints are mostly preserved in
fluvial deposits, particularly on floodplains with just one example preserved on
exposed fluvial sandbars (Fig. 14.9). There are scarce data about fluvial styles pre-
serving fossil footprints in meandering fluvial systems from the lower Miocene
Vinchina Formation (Lower Vinchina ichnoassemblage) and anastomosing fluvial
systems from the upper Miocene Toro Negro Formation (Toro Negro ichnoassem-
blage). Lacustrine deposits preserving fossils footprints involve uniquely marginal-
lacustrine facies and are only mentioned for the upper Miocene Puchuzun
ichnoassemblages (Fig. 14.9). Fossil footprints in eolian deposits are common in
Miocene and Pliocene outcrops, preserved mostly on dune and wet interdune facies
(footprints of Cacheuta, Catua, and Río Negro). Footprints are also recorded on
Pliocene salt flats of the Argentinean Puna (Fig. 14.9).
There is a broad relationship between the facies bearing footprints and biotic and
abiotic conditions prevailing at different moments during the Cenozoic in South
America. During the Eocene-Oligocene, relatively homogenous conditions were
present across the entire continent. Forested areas were widespread together with
tropical climate conditions that reached the summit during the early Eocene Climatic
Optimun (Zachos et al. 2001; Flynn et al 2012). During this time, mammal foot-
prints are recorded only in floodplain facies (Fig. 14.9).
Large-scale faunal provinciality reflecting latitudinal and altitudinal environ-
mental gradients appear to be present by the early middle Miocene or late Oligocene
of South America (Flynn et al. 2012). There are also environmental mosaics at spe-
cific areas within these gradients, as reflected by the lower Miocene of Santa Cruz
in Patagonia (Kay et al. 2012), the upper Miocene of Toro Negro in northwestern
Argentina (Krapovickas et al. 2009a, b, c) and the middle Miocene of La Venta in
Colombia (Kay and Madden 1997).
Miocene deposits involving mammalian footprints are recorded in subhumid,

semiarid, and arid climates (Fig. 14.9). Examples are distal overbank areas of high-
latitude settings (Santa Cruz) with moderately well-drained paleosols due to sea-
sonal shifts between subhumid and semiarid climatic; exposed sandbars of
anastomosing fluvial systems under semi-arid climates of mid-latitude areas at
northwestern Argentina (Toro Negro) and in arid climates from mid-latitude areas
of central western Argentina (Mariño) and northwestern Argentina (Puna). Among
these, Toro Negro is the richest assemblage. In this unit, inter-channel areas of anas-
tomosed fluvial systems, under a semi-arid climatic regime, have a high preserva-
tion potential for tracks due to seasonal flooding events, which rapidly bury and
protect the footprints, preventing rapid degradation of exposed tracks (Krapovickas
et al. 2009a, b, c). In comparison, the distal overbank areas of low-sinuosity rivers
(Raigemborn et al. 2012) with low rates of deposition and high subaerial exposure
result in lower preservation potential for footprints.
Glacial and interglacial oscillations occurred during the Pliocene and Pleistocene,
involving increased aridity. Late Miocene-Pliocene deposits bearing mammalian
footprints are mostly developed in arid climates of high- (Rio Negro; late Miocene-
Pliocene) and mid-latitude (Puna; Pliocene) settings. For most of the middle and
late Pleistocene, dry and cold climate caused open areas to predominate in South
America (Cione et al. 2009). Pleistocene floodplain facies of the Pampean region
bear one of the most impressive footprint assemblages worldwide.
14.8 South American Mammal Tracks and Implications

for Paleoecology
In ichnology, as well as in other paleontologic disciplines, paleoecology may be

seen from two different complementary perspectives (Kay et al. 2012). The most
frequent approach is to reconstruct environmental parameters, such as substrate
consistency, climate, food supply, topography (e.g., coastlines, exposed fluvial
bars), water saturation, water energy, salinity, oxygenation, rate of sedimentation,
light, and temperature, among others. The other approach is to understand aspects
of the paleoecology of individual species based upon their adaptive characteristics
such as body size, diet, locomotion, and substrate preference (paleoautecology), and
to build from this the overall niche characteristics of the fauna as a whole
(paleosynecology) (e.g., Gastaldo et al. 1996; Falcon-Lang et al. 2007; Krapovickas
et al. 2009a, b, c; Minter and Braddy 2009; Wilson et al. 2009; Kubo 2011). Both
approaches have been scarcely applied to the study of South American mammalian
ichnoassemblages.
Krapovickas et al. (2009a, b, c) used fossil footprints to infer characteristics of
ancient environments. The tetrapod fauna of the lower interval of the Toro Negro
Formation revealed by footprints suggested an open environment, possibly related
with a seasonal arid or semiarid climate, associated with nearby forested areas
(Fig. 14.10).
Fig. 14.10 Reconstruction of the Miocene Vinchina anastomosed fluvial system with exposed
sandbars and the putative vertebrate and invertebrate producers
Manera de Bianco and Aramayo (2004) studied the taphonomy of the Pehuen- Có
tracksite, considering footprint association, preservation quality, producers, and
substrate. They found that mudstones without desiccation cracks or ripple marks
display the best preservation quality, reflecting a differential preservation potential
related to the proximity to the floodplain water body.
Most of the paloeautoecologic studies are qualitative or biomechanical analysis
of the locomotion style or use of the substrate by the producers. Casamiquela (1974)
analyzed Megatherichnum oportoi, concluding that the absence of hand prints
reflects a bipedal locomotion, in accordance to many features of the skeleton of the
megatheres. This claim resulted controversial and has been reevaluated in detail by
numerous authors. Casinos (1996) studied the biomechanics of locomotion of
Pleistocene Megatherium americanum by means of the estimation of body mass and
using measurements of several trackways originally assigned to megatheres
(Frenguelli 1950; Bonaparte 1965; Casamiquela 1974, 1983; Aramayo and Manera
de Bianco 1987a, b). Casinos (1996) calculated different mechanical parameters such
as speed, Froude number, indicators of athletic abilities, and bending and resistance
movements of the vertebrate column calculated in both bipedal and quadrupedal
conditions. He concluded that it was not possible to determine whether Megatherium
was better adapted to bipedal or quadrupedal locomotion. Either bipedal or quadru-
pedal, in all cases the results indicated a walking mode of locomotion. Studying
Neomegatherichnum pehuencoensis, Blanco and Czerwonogora (2003) reassessed
the locomotory abilities of the Pleistocene Megatherium americanum. In doing so,

they developed a geometric model to calculate the percentage of body weight sup-
ported by each pair of limbs to estimate the pressure they should have exerted on the
ground and calculated the speed of locomotion after Alexander (1976). Their con-
clusion was that Megatherium walked roughly as a strategy to minimize the costs of
transport and to better withstand big lateral bending moments that appear in bipedal
walk. McDonald (2007) inferred the locomotion of Paramylodon harlani integrat-
ing morphologic and ichnologic data of ground sloth tracks of the Nevada state
Prison, Carson City, USA, assigned to Paramylodon. He concluded that the known
studies of the anatomy of ground sloth and their tracks have not been conclusive
about their capacity of bipedal locomotion. He also concluded that the evidence
suggests a quadrupedal primary mode of locomotion and that the support for bipedal
locomotion has been based on the general absence of manus impressions.
In terms of paleosinecology, Krapovickas et al. (2009a, b, c) analyzed the pres-
ervation potential of mammal tracks to estimate the magnitude of time passed on the
formation of the Toro Negro tracking surface in order to analyze the footprint
assemblage as a partial sample of the original tetrapod community that inhabited
western Argentina during the early Miocene. To characterize the paleocommunity
structure, these authors identified the number of trackmakers and the relative abun-
dance of footprints. The latter was interpreted as a measure of the activity of the
producers and not necessarily as an indicator of the number of individuals involved.
Krapovickas et al. (2009a, b, c) measured the relative abundance of footprints by
counting the number of footprints and developed an index that records the percent-
age of the tracking surface which is bioturbated by each ichnotaxa, the Relative
Bioturbated Area index.
The study of ichnofossils of the Toro Negro Formation expanded the knowledge
of its tetrapod paleocommunity in three main ways: by reinforcing the presence of
certain taxa as indicated by body fossils (large and medium-size ground sloths,
proterotheriid litopterns, caviid rodents, and birds), by refining the level of
identification of certain taxa (shorebirds, perching birds, and large cursorial birds),
and by adding taxa that would otherwise be unknown by body fossils (macrauche-
niid litopterns) (Fig. 14.11).
14.9 Future Directions
Despite the fact that first studies began almost a hundred years ago, the ichnologic
knowledge of South American mammals is still in its infancy. As mentioned above,
in this chapter we attempt to to identify problems and promises of the South
American record hoping to contribute to develop a major research program in mam-
malian paleoichnology that will permit global comparisons.
To begin with, there are extensive areas of South America where the Cenozoic
ichnologic content is still unknown, revealing the necessity of intensive field work:
there are no specific reports for Bolivia, Brazil, Colombia, Ecuador, French Guiana,
Guiana, Paraguay, Surinam, Uruguay, and Venezuela.
Fig. 14.11 Comparison of the tetrapod ichnological and osteological evidence from the Toro
Negro Formation at Quebrada de La Troya, Argentina. Black outline silhouette: indeterminate
bones. Black silhouette: material represented by both evidences. Light gray silhouette: material
represented by osteological evidence. Blue silhouette (medium gray): material represented by the
ichnological evidence. Modified from Krapovickas et al. (2009a, b, c)
Most ichnologic studies on Cenozoic South American mammals do not include

detailed analyses of the paleoenvironmental context and the bearing facies, and only
general aspects are mentioned. Much more analyses on ancient environments and
its prevalent climates need to be done to understand under which conditions
inhabited the mammalian trackmakers.
From a systematic point of view, at present, there are numerous ichnotaxa erected
to name very similar forms produced most likely by related trackmakers. There is a
strong need of a comprehensive and updated ichnotaxonomic revision of the existing
ichnotaxa in accordance with morphologic and paleobiologic characteristics of the
trackmakers. Moreover, in order to provide novel evolutionary, locomotory, and
distributional information, the attribution of footprints to body fossils groups needs
to be independent from the known body fossil spatial and temporal distribution.
Currently, ichnologists analyze putative trackmakers based on anatomic features of
the postcranial skeleton that are transcribed to the footprint morphology and
trackway pattern. In order to use footprints as an independent source of evidence,
when analyzing footprint producers, it is preferable to establish osteologic
synapomorphies that could be identified in the fossil footprints (Carrano and Wilson
2001). Unfortunately, until now, phylogenetic studies of South American mammals
do not include characters of the distal autopodium in their analyses. The study of
trackmakers on the light of the cladistic method of trackmaker identification is still
a pending task on the ichnologic agenda.
In terms of trackmaker function, there is scarce information about the range of
functional capabilities revealed in South American mammal trackways; it is
necessary to conduct deep analysis of the locomotory modes expressed in trackways
and analyze how this range compares to that of living mammals. The estimation of
the body mass of the footprint producers is another pending issue.
Finally, mammalian fossil footprints are barely used for habitat reconstruction
analysis. Both taxon dependent and taxon-free analysis based on the footprint
record could result in useful tools for habitat reconstruction.
These are the major problems and promises we were able to identify for South
America so far. We understand that reviewing all the available evidence from other
continents would identify particular and common problems and promises, contrib-
uting to develop a global paleoichnologic mammalian research program.
14.10 Conclusions
Globally, some of the most significant and better preserved ichnologic records of
Mesozoic mammals are from Gondwana, including Ameghinichnus, Brasilichnium,
and Duquettichnus, in contrast to its meager body-fossil record. In boreal land-
masses, the diversity and abundance of the Mesozoic mammalian body-fossil record
are much higher, and their footprint record is not as well known.
The Paleocene tetrapod footprint record, albeit apparently depauperate, has
scarce mammals represented and is mostly composed by amphibian, reptiles, and
birds. In contrast, mammal footprints become more common during the Eocene and
Oligocene. The Neogene mammalian record is dominated by ungulate footprints,
such as perissodactyls, artiodactyls, and proboscideans, followed in abundance by
carnivores as felids and canids.
Presently there are more than 20 known localities that record footprints of South
American mammals. Two main ichnologic assemblages are distinguished: (1) those
in which the producers belong to lineages that underwent diversification and eco-
logic specialization in isolation from other continents, and (2) those including also
representatives of North American lineages.
The Eocene and Oligocene ichnologic assemblages mostly represent forms of
uncertain affinity up to now, including small caviomorph rodents or typothere
notoungulates, large undetermined notoungulates, medium-size undetermined tet-
radactyl mammals, Macrauchenichnus isp. assigned to small toxodontid notoungu-
lates or macraucheniid litopterns. Among the Miocene assemblages there are
footprints assigned to ground sloths (e.g. Venatoripes riojanus, cf. Venatoripes rio-
janus, kidney-like footprints), macraucheniiids (e.g. Macrauchenichnus rector) and
proterotheriid litopterns, rodent-like caviomorph rodents or typothere notoungu-
lates, hegetotheriid notoungulates, and toxodontid notoungulates or macraucheniid
litopterns (e.g. Macrauchenichnus isp.), and Tacheria troyana assigned to a large
dinomyid caviomorph rodent.
The late Miocene-Pliocene and late Pleistocene assemblages include footprints
of both native South American mammals and North American taxa that arrived dur-
ing the GABI. The native South American lineages consist of a variety of ground
sloths (e.g., Megatherichnum oportoi, Falsatorichnum calceocannabius) including
megatheriids (e.g., Iribarnichnum megamericanum, Neomegatherichnum pehuen-
coensis) and mylodontids (e.g., Acunaichnus dorregoensis, Milodontichnum
rosalensis), glyptodonts, macraucheniid (e.g., Macrauchenichnus rector,

Eumacrauchenichnus patachonicus) and proterotheriid litopterns, hydrochoerid
caviomorph rodents (e.g., Porcellusignum consulcator), undetermined ungulates,
and carnivorous marsupials. The North American examples are equids (e.g.,
Caballichnus impersonalis, Ichnhippus cotaposi, Hippipeda isp.), gomphotheriids
(e.g., Stegomastodonichnum australis), camelids (e.g., Lamaichnum guanicoe,
Megalamaichnum tulipensis), cervids (e.g., Odocoileinichnum commune), and sev-
eral carnivoran lineages as felids and mustelids (e.g., Pumaeichnum biancoi,
Pehuencoichnus gracilis, Mustelidichnum enigmaticum).
The cluster analysis of the mammalian taxa represented by the fossil footprint
record of South America, compared with the faunal evolutionary phases proposed
by Goin et al. (2012), shows two major clusters that roughly separate the faunas
corresponding to the Early and Late South American phases from the Interamerican
phase.
The early Eocene rodent-like footprints of Paruro are of great importance for the
understanding of the early evolution of caviomorph rodents. If future careful analy-
sis confirms the caviomorph affinity of the Paruro footprints, they could reinforce
the recent hypothesis of Antoine et al. (2012) about the southward expansion of the
earliest caviomorphs during the late middle Eocene–early Oligocene period.
Moreover, the footprints would expand the early record of rodents in South America
to the early Eocene.
Footprints were uniquely preserved on floodplain facies during the Eocene,
Oligocene, and Pleistocene. The record of mammalian fossil footprints in deserts of
South America occur in a variety of sub-environments (dunes, wet interdunes, and
salt flats), involving semi-arid and arid climates, during the Miocene and Pliocene.
Both approaches, the reconstruction of environmental parameters and the study
of the paleoecology of individual species and communities, have been scarcely
applied to South American mammalian ichnoassemblages.
Acknowledgments We thank D. Croft and S. Lucas for reviewing the paper. E. Ahumada,
S. Aramayo, S. Bargo, V. Carlotto, E. Cerdeño, V. Contreras, T. Manera de Bianco, and A. Zavattieri
for the photographs and information that gently shared with us. Funding for this research was
provided by UBACyT—X728 and PICT-2011-2334 (V.K.). This is the contribution R-100 of the
Instituto de Estudios Andinos Don Pablo Groeber.
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Chapter 15
Major Events in Hominin Evolution
Martin Lockley, Jeff Meldrum, and Jeong Yul Kim
15.1 Introduction
Until recently (reviewed in Kim et al. 2008a, b; Lockley et al. 2007a, b, 2008a, b;
Lockley and Roberts 2004), the study of fossil hominin tracks had been largely
neglected by ichnologists, paleontologists, and anthropologists, with the exception
of the famous Laetoli hominin trackways in east Africa. While it may be speculative
to consider why this has been so, the study of hominin footprints, like the study of
hominins in general, is of potential mutual interest to several disciplines (paleontol-
ogy, ichnology, anthropology, archeology), and so has not been claimed as the
exclusive province of any one of these. The distinction between hominids and homi-
nins is sometimes confusing to nonspecialists. Here we use the term hominin, in
reference to the tribe Hominini, to include representatives of the “australopiths” and
the “hominans”, as outlined by Wood (2005), and Wood and Richmond (2000).
With a few notable exceptions (Hay and Leakey 1982; Leakey and Harris 1987;
Pales 1976) there were almost no hominin footprint-bearing sites that had been
subjected to any sort of comprehensive analysis, prior to 2000. Even though a spate
of publications in 2004 through 2009 (Meldrum 2004a, b; Lockley et al. 2007a, b,
2008a, b; Meldrum et al. 2010, 2011) marked a re-awakening of interest, most of the
few dozen publications to emerge at this time could best be classified as preliminary
M. Lockley (*)
University of Colorado Denver, CB172, PO Box 173364, Denver, CO 80217-3364, USA
J. Meldrum
Department of Biological Sciences, Idaho State University,
921 S. 8th Ave., Stop 8007, Pocatello, ID 83209-8007, USA
J.Y. Kim
Department of Earth Science Education, Korea National University of Education,
Cheongwon, Chungbuk 361-892, Korea

DOI 10.1007/978-94-017-9597-5_15
412 M. Lockley et al.
site reports, some dealing with sites that had been known for decades but never
studied. Nevertheless, since the publication of two special volumes in Ichnos (vol.
15 (3–4) for 2008, and vol. 16 (1–2) for 2009), under the title Hominid Ichnology,
interest in the subject has been sustained, and a number of papers have emerged
dealing both with the oldest African sites, and with other younger more widely dis-
tributed sites (e.g., Bennet and Morse 2014). In accordance with the theme of this
volume, we herein review the hominin track record from the perspective of the role
of trace fossils (primarily fossil footprints) in highlighting major events in hominin
evolution, including emphasis on new studies and perspectives not discussed in pre-
vious summaries (Lockley et al. 2007a, b, 2008a, b).
While the focus of this contribution is on fossil footprints and their interpretation, the
renaissance in “Hominid Ichnology”, represented in part by the Ichnos volumes, has
been accompanied by independent, but simultaneous, interest in redefining the scope of
hominid ichnology (Hasiotis et al. 2007; Baucon et al. 2008; Kim et al. 2008a, b). In
short, the authors of these articles, as well as those of the present article (Kim et al. 2004,
2008a, b), simultaneously proposed that hominid ichnology should include the study of
a wide variety of traces, in addition to footprints. These traces include, but are not lim-
ited to, tool marks and artifacts made by modifying (flaking, engraving, sculpting, exca-
vating), of wood, bone, rock, and earth (soil) substrates, as well as various forms of
painting and writing, also on a wide range of substrates. While recently made popular in
Crime Scene Investigation (CSI) dramas, involving detection of all manner of traces on
diverse substrates (Lockley 1999), such facets of hominid ichnology were already evi-
dent during the so called Late Paleolithic Cultural Revolution (discussed below) and
raise fundamental questions about the importance of ichnology in understanding and
reflecting complex cultural behavior during one of the most debated “major events in
evolution”—the emergence of modern humans. Hominid ichnology bridges the gap
between tetrapod ichnology and what Baucon et al. (2008) appropriately identified as
the important subdiscipline of ichnoarcheology. The etymology of words like ichnos
and trace, discussed below, underscore the deep relationships between trace fossils and
writing. Without such ‘ichnologic’ representations of the behavior of modern humans
and their ancestors, this volume and all its predecessors would not be possible.
15.2 Early Hominin Bipedalism: Laetoli Revisited
Even the non-paleontologist and non-anthropologist is likely aware of the famous

footprints from Laetoli in Tanzania discovered by the Mary Leakey team in 1976
(Leakey and Hay 1979; Hay and Leakey 1982; Leakey and Harris 1987). They
ostensibly indicate a group, possibly a family unit, of three individuals, one of which
was slightly smaller (creating trackway G-1) walking to the left-hand side of the
other two (creating trackways G2/3), which appeared to be two individuals stepping
in each other’s footprints (Fig. 15.1). Trackmaker height estimates are about 1.10–
1.15 m for G-1 and 1.32–1.52 m for G-2/3 (White and Suwa 1987). As noted below,
despite this group-of-three scenario, and the many analyses of the trackways that
15 Major Events in Hominin Evolution 413
Fig. 15.1 Pliocene

hominin trackway from
Laetoli (left) include the
G-1 trackway (left) and the
overprinted, “double”
G-2/3 trackways, possibly
open to interpretation as a
twice-overprinted, “treble”
G-2/3/4 trackway.
Photogrammetric image
(right) after Kim et al.
(2008b) shows three
consecutive tracks (a
stride) and highlights the
multiple hallux (big toe)
traces in the G-2/3
trackways
have been published since 1976 (e.g., Charteris et al. 1981; Tuttle 1990; Tuttle et al.
1990; White and Suwa 1987; Meldrum 2004a, b; 2007a; Meldrum et al. 2011), this
interpretation has recently been challenged, and it has been claimed that there are in
fact three trackmakers represented by the overlapping series not two (Musiba et al.
2011; Matthews et al. 2011). This observation implies a G-1 + G-2/3/4 scenario.
Clearly, the most significant implication of the Laetoli trackways and the one on
which most researchers agree, is that they provide direct evidence of a ~3.6 Ma old
hominin that was capable of walking upright. As almost every anthropologic text
proclaims, the transition to a fully upright gait was a major event in hominin evolution,
perhaps the seminal adaptation of this radiation. Moreover, given the fragmentary
state of many body fossils of this age, trackways arguably provide the best evidence
that hominin bipedalism was established by 3.6 Ma. Mary Leakey proclaimed this
“the most remarkable find I have made in my whole career” (Lewin 1982, p. 220).
A few other facts may be inferred from the Laetoli tracks and their broader geologic
and paleontologic context. First, they form part of a huge trackway assemblage domi-
nated by the tracks of non-hominin mammals and birds, including some that evidently
represent extinct species (Leakey and Harris 1987). We may also assert that the body
fossil record shows that representatives of genus Australopithecus were extant at the
time when the tracks were made. Lastly, we may note that the tracks were recently
given the name Praehominipes laetoliensis (Meldrum et al. 2011). The formal name
and diagnosis distinguish them from the previously erected ichnotaxon Hominipes
modernus (Kim et al. 2009), the latter being the formal name applied to fossil footprints
unequivocally attributed to Homo sapiens (and possibly H. neanderthalensis).
Beyond these facts there is far less agreement about the interpretation of the
trackway evidence, though Kim et al. (2007) discussed their paleontologic,
stratigraphic, and sedimentologic significance. As we have seen, even the num-
ber of individuals producing a trackway has recently been disputed (Musiba
et al. 2011). In addition, there is disagreement as to whether the tracks were
made by a trackmaker with a foot architecture manifestly distinct, in its mosaic
and/or intermediate nature (Stern and Susman 1983; Susman et al. 1984;
Deloison 1991, 1992; Clarke 1999; Meldrum 2000, 2002; 2004; Berge et al.
2006; Meldrum and Chapman 2007; Bennett et al. 2009; Hatala 2014) or an as
yet unrecognized relatively derived hominin (e.g., Day and Wickens 1980;
Charteris et al. 1981; Alexander 1984; Suwa 1984; Lovejoy 1988; Tuttle 1985,
1996; Tuttle et al. 1990, 1991; Schmid 2004; Sellers et al. 2005; Harcourt-Smith
and Hilton 2005; Kimbel and Delezene 2009; Raichlen et al. 2010; Tuttle 2014).
Closely intertwined with this debate is the question of whether the footprints are
essentially indistinguishable from arched modern hominin tracks (Tuttle 1990;
Tuttle et al. 1990; Crompton et al. 2012) or whether they show evidence of
primitive features, including a midtarsal break (sensu Meldrum et al. 2011),
which would imply that the trackmaker’s foot morphology, and dynamic foot-
print registration differed from that known for modern humans.
It is perhaps surprising that such basic evidence as the number of individual track-
ways is in dispute. However, it should be remembered that after the initial excavation,
molding, and analysis, which included some 3D imaging using 1980s technology, the
tracks were reburied and most researchers had to rely on studying planimetric figures
or fiberglass replicas representing only a short segment of the G1 and G2/3 trackways
(Lockley and Matthews 2007; Meldrum 2007a). Even after the tracks were exhumed
by the Getty Conservation Institute (Demas and Agnew 1996) in order to assess dis-
turbances resulting from the reburial, they were again reburied in order to conserve
them. It was only in 2011 that they were exhumed for a second time and subjected to
high resolution photogrammetric analysis (Musiba et al. 2011; Matthews et al. 2011).
These results suggest an assemblage of four, rather than three trackways, in which a
group of three rather than two have overlapping footsteps. While popular inference
has been tempted to interpret the “group-of-three” scenario as possible evidence for a
“family” unit consisting of two larger adults and a smaller juvenile offspring, the new
interpretation casts doubt on such a “nuclear family” scenario. Meldrum (2007a),
after a careful analysis of two sets of stereophotographs of the G2/3 trackway, deter-
mined that the lengths of the G2/3 footprints were approximately 25 cm (G2-25) and
20 cm (G3-9), respectively. The more recent evidence (Musiba et al. 2011; Matthews
et al. 2011) calls for a careful reanalysis of the size of the overlapping footprints,
which may revise these size estimates. What might the number and size composition
imply about the trackmaker demographics and behavior? What induced three track-
makers to follow closely in line, with the two followers stepping repeatedly in the
footprints of the leading individual? An unfamiliar substrate? Given the uncertainty
surrounding the overstepped footprints (G2/3 or G2/3/4), we can only note that the G1
trail has footprints ~18 cm long and ~8 cm wide with a step of 41–44 cm, indicating
a trackmaker with a stature of ~1.32 m (cf. White and Suwa 1987).
As noted by Lockley et al. (2008a, p. 107), the non-hominin Laetoli track

assemblages allows a census that ostensibly attributes most trackmakers to
“extant species, or species that are indistinguishable from modern species (or
generic, and higher categories) on the basis of tracks. The only exceptions are
the identification of extinct chalicothere and Hipparion tracks, the latter studied
by Renders (1984). The census clearly shows that rabbits (or other lagomorphs)
were abundant in the area, numerically constituting 88.8 % of the non-hominin
vertebrate tracks.” Thus, based on rabbit abundance, Laetoli was “the Watership
Down” of the ichnologic world (Lockley 1999, p. 241)! However, as this census
is based on individual prints—not trackways—it is not a reliable measure of
biomass, although potentially useful as a general indicator of ecology and the
activity of faunal components.
Lastly, we may note that the Laetoli tracks are preserved in reworked volcanicla-
stic sediments, which allow for inferences about runoff and seasonal climatic
regimes (Hay and Leakey 1982; Houck et al. 2009). In conclusion, therefore, the
Laetoli tracks offer us valuable evidence of a major evolutionary event; the emer-
gence and nature of early hominin bipedalism. They also form part of a rich assem-
blage that has attracted wide scientific interest. However, disparate interpretations
of the trackmakers and their behavior continue to be debated, and access to the
entire trackways, rather than partial replicas, remains restricted by the need to keep
them covered in order to preserve them.
15.3 Walking Erect Phase II: In and Out of Africa
15.3.1 On the Shores of Lake Turkana
There is a huge temporal gap in the hominin track record between the ~3.6 Ma
Laetoli occurrence and two ~1.5 Ma track sites recorded in the Koobi Fora Formation
on the eastern shore of Lake Turkana. As noted by Behrensmeyer and Laporte (1981,
p. 3), the first discovery of Koobi Fora footprints (from a site known as GaJi10) con-
tributed “a reference point on hominin foot morphology, locomotion behavior, and
ecology 2 Myr younger than the Laetoli occurrences and ~1 Myr older than the Late
Pleistocene human footprints in Europe.” Unlike the Laetoli tracks, which are
unequivocally Pliocene in age, the Koobi Fora tracks can be assigned to the Lower
Pleistocene, which spans the interval between ~1.8 and ~0.8 Ma.
At the time of the first discovery, Behrensmeyer and Laporte (1981) inferred that
the Koobi Fora tracks may have been produced by a Homo erectus (H. ergaster)
individual ~1.5–1.6 Ma. This date is more or less confirmed by Bennett et al. (2009),
who stated that the track layer is just below a tuff dated at 1.435 Ma. The decided
lack of details of morphology in the tracks provides no evidence of significant
changes in the hominin foot since the Laetoli trackway.
Behrensmeyer and Laporte (1981) reported tracks visible in a single trackway at
site GaJi120, but Bennett et al. (2009) reported that two more tracks were excavated
at the site. According to Behrensmeyer and Laporte (1981), the tracks range in
length from 25 to 32 cm and the mean foot size dimensions are estimated at 26 cm
long and 10 cm wide, suggesting an individual 1.6–1.8 m in height. In addition to
some isolated tracks at the Ileret site, Bennett et al. (2009) reported at least four
trackway segments, one from the lower level, and three from the upper level, and
they estimated the height of the track makers as between 1.75 and 1.78 m (±0.26 m),
with a possible subadult print from the lower level representing an individual only
0.92 m (±0.13 m).
Bennett et al. (2009) reported another site at Ileret, Kenya, about 45 km north of
the GaJi10 site that reveals two hominin track-bearing layers, ~5 m apart strati-
graphically, dated at ~1.53 Ma. The Ileret site reveals tracks which have better pres-
ervation than the GaJi10 site footprints as they reveal individual digital pad
impressions. Dingwall et al. (2013) concluded that some or all of the footprints
found on three levels could be either Homo erectus (H. ergaster) or Paranthropus
boisei. Bennett et al. (2009) concluded that the footprints provide “the oldest evi-
dence of an essentially modern human-like foot anatomy, with a relatively adducted
hallux, medial longitudinal arch, and medial weight transfer before push-off.” We
find support for only one of these three points (i.e. a relatively adducted hallux). We
find no evidence to support the consistent presence of a longitudinal arch or for a
medial weight transfer. The suggestions of a medial longitudinal arch in the clearest
examples (such as depicted on the cover of Science; Bennett et al. 2009) are the
result of distortions caused by secondary impressions with extrusion left by passing
ungulates, which obscure the medial margin of the hominin footprints (Fig. 15.2).
In other instances, the preservation is so poor that the actual topography of the
medial contact surface is indiscernible. A number of the published examples show
a clear lack of medial weight transfer. Although differences of substrate properties
may influence the general appearance of individual footprints as demonstrated by
Morse et al. (2013), we observe consistent distinctions to the specific appearance of
modern human footprints, regardless of substrate. Therefore, we maintain that
Homo erectus (H. ergaster) had neither a fixed longitudinal arch (sensu Mauch et al.
2008) nor a modern toe-off mechanism, in concordance with analysis of the hallucal
metatarsal KNM BK-63 (Meldrum et al. 2010), or that of the Dmanisi hominin post
crania (contra Lordkipanidze et al. 2007; Pontzer et al. 2010).
15.3.2 Out of Africa
The phrase “Out of Africa” has become synonymous with the now entrenched view
that our hominin ancestors originated in Africa, an opinion that can be traced back at
least to Darwin’s Descent of Man (Darwin 1872). Certainly there is little or no evi-
dence to refute the idea that australopithecines and early Homo had their origins in
Africa and may never have left that continent (although interpretations of Homo flo-
resiensis from Indonesia, as potentially a relict late australopith/early Homo sp.
raises the possibility of earlier dispersal out of Africa). Likewise, it has traditionally
Fig. 15.2 Tracks attributed

to Homo erectus (H.
ergaster) at Ileret, Kenya
(after Bennett et al. 2009)
been assumed that Homo erectus (H. ergaster), a species that existed from ~2.0 to
0.2 Ma, with little pronounced morphologic change, originated in Africa, even
though primitive representatives of this species dispersed across the old world at an
early date. For example, remains are reported from Dmanisi, Georgia as early as
~1.8 Ma, raising the possibility of an Asian origin for H. erectus (Ferring et al. 2011).
Following the hominin footprint trail from the early Pleistocene sites east of
Lake Turkana, the next youngest reports are from three European sites, one dated to
the Early Pleistocene, between 1.0 and 0.78 Ma, and the other two dated to the
Middle Pleistocene. The oldest site, recently reported by Ashton et al. (2014), is
associated with estuarine sediments of the Cromer Forest-bed Formation at the
Happisburgh site in east Anglia, England. The footprints identified at this site occur
in parallel-laminated silts, but are not very well preserved. In fact, only one foot-
print shows toe impressions and none preserve clear evidence of a medial arch.
However, although the footprints are susceptible to destruction by marine erosion
soon after they are exhumed, about 50 elongate footprints (length 14–25 cm, width
6–11 cm) were nevertheless identified in an area of about 12 m2. Ashton et al. (2005,
p. 7) argued that “the shape of the footprints suggests that they were most likely
made by hominins and none of the prints are consistent with those formed by other
mammals.” Moreover, Parfitt et al. (2005) have established that flint tools made by
hominins occur in this same formation.
Prior to the reports from Happisburgh, the oldest, best-dated, and best-studied
footprint site was the Middle Pleistocene Roccamonfina Volcano site in Italy,
reported by Mietto et al. (2003) and Avanzini et al. (2004, 2008), dated between
~385,000 and ~325,000 yBP. The trackways occur in a volcanic ash deposited on
the slopes of the Rocamonfina volcanic complex and, due to the slope, one trackway
of 27 footprints has a zig-zag configuration, indicating an individual switch-backing.
Another trackway of 19 prints follows a gently curved course, with a few hand- or
palm-prints where the trackmaker put his/her hand on the ground. A third trackway
of ten prints forms a straight line. The footprint details are quite indistinct and not
very well-preserved. Most lack clear toe impressions, even though faint traces of the
hallux are recognized in some cases, even indicating a gap between digits I and II
typical of habitually unshod individuals. The tracks are generally flat, but show pos-
sible inconsistent traces of a transient medial longitudinal arch in some cases. They
are described as short and broad (~20–24 cm long and ~10–12 cm wide), suggesting
a stature for the trackmaker of ~1.56 m. Based on age, these tracks likely represent
a pre-sapiens or pre-neanderthalensis species such as H. heidlebergensis, or possi-
bly late Homo erectus (H. ergaster) (Scaillet et al. 2008).
A second mid-Pleistocene footprint from Terra Amata, in southern France, has
been tentatively dated at ~300,000–400,000 YYBP (De Lumley 1966, 1967;
Miskovski 1967; Meldrum 2006, p. 246; De Lumley et al. 2011). The site only
reveals one track, but it clearly shows a diagnostic hominin big toe (hallux) trace.
However, like the Rocamonfina tracks, there is little evidence of a well-defined
medial longitudinal arch (Meldrum 2004a, b). It is interesting that the lack of a well-
defined arch is noted in both the Italian and French tracks. This is presumably open
to several possible interpretations. For example: (1) the lack of a discernible arch
could be due to poor footprint preservation, resulting from suboptimal substrate
conditions at the time of registration, or post-track-making or post-exhumation
weathering of the footprints, (2) the trackmaker may have been an individual or a
member of a species in which arches were not well-defined, or inconspicuous as the
result of having robust fleshy feet, or (3) hominins exhibit a mosaic pattern in evolu-
tion of features of the foot and stability of the foot’s medial column was still lacking
at this stage of hominin evolution.
Barnaby (1975) noted that hominin footprints from a volcanic ash near
Demirköprü, Turkey were assigned an age of 250,000 yBP, suggesting they repre-
sent a pre-sapiens species (Ozansoy 1969). However, subsequent thermolumines-
cence dating of associated tuffs in the area gave much younger dates on the order
of 65 ± 7 ka and 49 ± 9 ka (Westaway et al. 2003, 2004, 2006). These dates have
been amended, yet again, to suggest dates as young as 12 ka (Tekkaya 1976; see
Lockley et al. 2008a, b for summary). Some 50 pairs of footprints have been
removed and are stored at the MTA Museum (Ocakoglu, personal communica-
tion). They measure ~29 cm in length and 11 cm wide, and show a robust hallux
impression and well-developed ball, heel pad, and medial longitudinal arch, sug-
gesting the footprint of a fully modern foot.
15.4 Tracking the Dawn of H. sapiens in Africa
The fossil footprint record in Southern Africa is important for understanding

Middle-Late Pleistocene transitions, which were more or less temporally coincident
with the emergence of modern Homo sapiens. Roberts (2008) reported the two
important tracksites: one at Nahoon Point, near East London (Deacon 1966;
Mountain 1966), now dated at about 127,000 ± 8000, and another in the Langebaan
Lagoon area, 130 km south of Cape Town, dated to about 117,000 YYBP (Gore
1997; Roberts and Berger 1997; Roberts 2008). Thus, both track sites represent the
last interglacial period that coincides with the sub-Series boundary between Middle
and Late Pleistocene (Gibbard 2003). The Nahoon Point tracks are better preserved,
showing well-preserved toe impressions and a footprint length of ~19 cm, and are
associated with poorly preserved mammal and bird tracks. The Langebaan lagoon
tracks are larger (~23 cm long), but less well-defined. They are associated with
probable hyena tracks (Roberts 2008). Hatala et al. (2011), Richmond et al. (2011)
and Zimmer et al. (2012) reported an assemblage of about 350 tracks comprising 18
trackways from Lake Natron, Tanzania dated at about 120,000 YYBP (Wong 2011).
Charles Helm (personal communication 2012) reports a number of other mammal
and bird track sites along the South African coast.
15.5 The Late Pleistocene: H. sapiens Produces a New Type

of Ichnologic Record
The ichnologic record of Homo sapiens in the Late Pleistocene contains evidence of
an evolutionary event that can only be characterized as “revolutionary.” Simply put,
this is because humans changed the fundamental nature of the ichnologic record.
Whereas they had previously created footprints, and a few butcher marks on bone,
only inadvertently, by as early as 30,000–35,000 years ago they were deliberately
producing artifacts and artwork, which most anthropologists acknowledge as an
unprecedented “creative revolution.” Before proceeding any further we need to
explore the justification for including human creations, such as art work and other
traces produced by the growth of civilization, under the broad umbrella of verte-
brate ichnology. The word ichnos derives from the Greek meaning footprint or
trace. In this regard the English word “trace” has deep etymologic roots connecting
it with the words, draw, drag, and trace. So, for example, a draw horse, drags or
draws a plow leaving a furrow or trace (see Smith and Hall 1914 or Harper 2012, for
Latin translation of “to drag” as traho).
On the one hand, one might argue that it would be simpler and more consistent
with our account of the pre-Late Pleistocene record, to confine our discussion of
hominin ichnology to footprints left “inadvertently” by humans, or human ances-
tors, and therefore to “avoid” discussion of a plethora of traces created deliberately
by humans, leaving it to be dealt with in the fields of archeology and anthropology
(Kim et al. 2008a; Baucon et al. 2008). On the other hand, it can be persuasively
argued that it is entirely arbitrary to draw a line between human traces produced
inadvertently and those produced deliberately. A bird builds a nest deliberately, not
inadvertently, and the same goes for animals that dig burrows. One might also argue
that avoiding the problem is the lazy way out, because it fails to address the question
of what is legitimately classified as a hominin trace fossil. The separation or avoid-
ance approach would in effect treat humans as a “special” species, somehow outside
the evolutionary continuum, creating a charge of unwarranted bias, contrary to
much contemporary scientific philosophy. Moreover it is difficult if not impossible
to draw an arbitrary dividing line between the cultural artifacts and traces produced
by H. sapiens and non-H. sapiens species. Pre- or non-sapiens species were already
producing trace fossils, such as butcher marks on bone and knapping marks on flint
long before the first sapiens cultural revolution was underway in the Late Paleolithic.
As noted in Sect. 15.1, there is also scientific precedent, for including many diverse
sapiens artifacts as an integral part of hominin ichnology (Hasiotis et al. 2007; Baucon
et al. 2008). Kim et al. (2008a) presented some of the aforementioned rationale in argu-
ing for four categories of hominin trace fossils, including (1) Pliocene through Holocene
footprints, (2) Pleistocene through Holocene butcher marks (feeding traces), (3) Early
and Mid Pleistocene stone tools, and (4) Late Pleistocene multimedia technology
(including art, dwelling traces, etc.). Examples of traces in the latter two categories can
be found through to the present time. In a similar, but more detailed evolutionary scheme,
Rothschild and Lister (2003), in a standard textbook on evolution, listed ten major events
in hominin evolution, occurring in the last 5 million years. Among these, a majority
leave a trace fossil record, notably (1) bipedal locomotion, (2) utilization of new food
resources, (3) stone tool manufacture, (4) control of fire, (6) symbolic communication,
(9), and complex cultural and technologic diversification (10). Hasiotis et al. (2007)
proposed a classification of hominid trace fossils that is somewhat different from the
fourfold classification of Kim et al. (2008a). Their classification includes “features, bio-
facts/ecofacts and artifacts” which are further divided into multiple categories, including
almost all manufactured items or creations including structural remains, lithic, ceramic,
and metal items. However, in general, the two classifications are similar in so far as they
recognize almost all human creations that leave physical traces as some category of trace
fossil. As noted above, this discussion has been enlarged by Baucon et al. (2008) in their
definitions of the broad scope of ichnoarcheology.
For convenience, in the sections that follow, we discriminate between artifacts
themselves and the traces left by their creation. For example, stones are not trace
fossils, but worked stone tools, and especially flake scars are trace fossils. Likewise,
a fire is an artifact in a different category from the charred hearth left behind as the
trace of the fire. In the same way a shelter or building is not in the same category as
the post holes or foundation ditches created during their construction.
In any event, from a philosophic and methodologic perspective, we arrive at the

following conclusions. There is no scientific justification for confining discussion of
hominin ichnology only to footprints (produced inadvertently or otherwise). Thus,
we may include such diverse traces as flaked stone tools, fire traces, cave painting,
ditches, and foundations. While these trace fossil categories are commonly different
from those associated with animal activity during most of the Phanerozoic, they are
nevertheless unequivocal evidence of behavior and activity. Like other trace fossils,
evidence of activity of organisms (Mcllroy 2004) or work of organisms (ICZN
1999), hominin traces are significant in paleontology, stratigraphy, and sedimentol-
ogy, but unlike other trace fossils, however, hominin traces can uniquely contribute
to understanding the work or activities of our ancestors (Kim et al. 2007). Since it is
we humans who attempt to evaluate “differences” between traces produced by dif-
ferent species, some subjectivity is perhaps inevitable, and the question arises as to
whether these are “differences of kind or differences of degree.” These issues are
further discussed below. It is also worth noting that while the creation of a diverse
suite of new hominin trace fossils in the Late Pleistocene is a considered a revolu-
tionary leap forward in cultural terms (Diamond 1999; Mellars 2006) appearing as
a major ichnologic event in the trace fossil record (Hasiotis et al. 2007; Kim et al.
2008a), it is not necessarily an anomalous evolutionary event. Rather it can be
viewed as an example of a “punctuated evolution” scenario, or a change in the
tempo of cultural evolution following a long Early–Middle Paleolithic period of
relative stasis.
15.5.1 Breaking New Ground: Prelude to the Late Pleistocene

Cultural Revolution
The geologically defined Late Pleistocene age extends from ~130,000 to 10,000
yBP. However, in comparison with the rapid developments of human cultural devel-
opment in the later part of the Late Pleistocene, which falls within the range of 14C
dating techniques, the early part of this age (before ~50,000) has, until recently, pro-
duced comparatively little evidence of any major evolutionary events. There are no
Late Pleistocene hominin tracksites dated with confidence in the first half of this
interval (between ~130,000 and ~65,000). By contrast the latter part of the Late
Pleistocene saw the worldwide spread of Homo sapiens. Unequivocal evidence,
including footprints, establishes the presence of modern humans (H. sapiens) in both
Australia and the Americas, as well as in ostensibly remote regions such as the
Tibetan Plateau and Jeju island, Korea (Zhang and Li 2002; Kim et al. 2009). Along
the way there may have been interactions with, even gene flow between, other late-
Homo species—Neanderthals, Denisovans, and Red Deer Cave hominins (Meldrum
2012b). Tracks left by any of these species would likely be fully modern in appear-
ance, albeit robust in proportions. In contrast, the persistence of more archaic homi-
nins into this period—Homo heidlebergensis, even Homo erectus (Swisher et al.
1996) and the enigmatic and as yet very restricted species H. floresiensis (Morwood
et al. 2005) raises the possibility of additional footprint morphotypes. Given the
perennial interest in the relationship of Neanderthals to modern humans, their
inferred extinction as recently as ~25–30,000 yBP is generally considered a major
event in hominin evolutionary history (Delson and Harvati 2006 and refs. therein).
However, despite a footprint record spanning this time interval, it is not sufficient to
shed useful light on the timing of this or other similar events.
However, before accepting that the so called Late Paleolithic cultural revolution
or “great leap forward” into modernity (Diamond 1999; Mellars 2006) occurred
quite abruptly ~50,000 yBP, perhaps coincident with, or in some way related to, the
extinction of the Neanderthals it is important to note that this is a rather Eurocentric
view that has been challenged by McBrearty and Brooks (2000). These authors
show that many technologic advances, considered typical of European cultures
existing between ~10,000 and ~50,000 yBP, have been identified significantly ear-
lier in Africa on the order of >100,000 yBP.
Lockley et al. (2009) reported a total sample of 19 Late Pleistocene hominin
tracksites from diverse localities around the world. Such geographically widespread
occurrences demonstrate that the hominin track record is consistent with that
obtained from archeologic evidence. In comparison with the important Pliocene
through earliest Late Pleistocene track record in Africa, the record from most of the
Late Pleistocene of Africa is sparse. Scott et al. (2008a, b) reported an isolated and
poorly preserved track from the Lake Bogoria area of the Kenyan Rift valley. The
track is ~20 cm long and 8.5 cm wide, and occurs in association with the footprints
of bovids, suids, and birds.
Turning to Europe, we find some of the earliest of the Late Pleistocene track
records associated with caves. Based on published dates, the oldest Late Pleistocene
human footprints are those reported from Vârtop Cave, Romania (Onac et al. 2005)
and assigned an age of ~62,000 yBP. Three footprints, one with a well-defined out-
line (22 cm long and 10.6 cm wide), are preserved. However, details are obscured by
infill of soda straws and moonmilk. Of note was a distinctive gap (1.6 cm) between
the first and second digits (Fig. 15.3). According to these authors, the Vârtop Cave
tracks may be the only footprint sample unequivocally attributable to Neanderthals.
Human footprints were reported from a “deep Middle Paleolithic layer” at
Theopetra Cave in Thessaly, Central Greece (Facorellis et al. 2001). This cave was
evidently occupied from the Middle Paleolithic until the post Neolithic, with mini-
mal 14C dates of ~48,000 yBP obtained for the oldest layers with which the foot-
prints are associated.
15.5.2 The Hominin Track Record Goes Underground
Many Late Pleistocene hominin footprint sites are associated with Late Paleolithic
cultural evidence found in subterranean cave sites in southern France, Spain, and
Italy. The region is perhaps most famous for cave paintings, but though less often
reported in detail, many sites contain tracks and other evidence of Late Paleolithic
Fig. 15.3 Purported

Neanderthal footprints,
Vârtop Cave, Romania.
Note growth of cave
calcite, obscuring track
morphology
activity. Tracks are documented from Lascaux (Berriere and Sahly 1964) and the
Niaux cave system (Pales 1976). Tracks from the Niaux caves have been described
and illustrated in detail by Pales (1976), and apparently include patterned arrange-
ments of footprints that have been interpreted as deliberate activity of children at
play (Lockley and Meyer 2000) (Fig. 15.4). Here the issue of intention is raised,
reminding us that by the Late Paleolithic not all human tracks were made as the
result of purely unintentional passage of individuals through a particular area.
Indeed, it is generally assumed, if not explicitly stated, that humans entered under-
ground caves deliberately, even if the motives, other than creating artwork, are not
clear. In a similar vein, human tracks from Grotte de Cabrerets or “Pech Merle”
cave famously reveal traces indicating an individual using a walking stick (Begouen
1927; Vallois 1927, 1931).
In other cases, the interpretations put on tracks are more ambiguous. Tracks from
Grotte Aldène assigned an age of ~15,000 yBP by Casteret (1948), but assigned and
age of ~8000 yBP by Ambert et al. (2000), are associated with charcoal, hyena
tracks, and cave bear nests, but the behavior of the track maker, in relation to other
animal and human traces is unclear. In Fontanet Cave, footprints suggest a child
may have followed a puppy or fox into a cave, but this inference can hardly be con-
firmed or denied on the basis of available analysis (Bahn and Vertut 1988). As noted
Fig. 15.4 (a) Tracks in a patterned arrangement from Niaux cave, suggest children at play (after
Pales 1976 and Lockley and Meyer 2000, Fig. 10.12). (b) Track from Peche Merle cave
elsewhere (Lockley et al. 2007a; 2009), the footprints of children appear as common
components of the track record of Paleolithic caves. For example, in Chauvet Cave
near Vallon-Pont-d’Arc in southern France, a trail of footprints claimed to represent
those of a young boy, about 8 years old and about 1.5 m tall, are possibly the oldest
European footprints of Homo sapiens, perhaps as old as 35,000 yBP. At Ariège
three trackways of children are recorded (Bahn and Vertut 1988). The footprints
extend for about 50 m across the cave floor and may be between 20,000 and
30,000 years old (Harrington 1999; García 1999, 2001). Many other Late Paleolithic
caves with hominin tracks have been mentioned or illustrated without detailed
descriptions (Kuhn 1955; Marshack 1972; Vialou 1986; Bahn and Vertut 1988). As
already implied, in many cases these sites reveal evidence that other animals, evi-
dently all mammalian carnivores, shared these cave habitats, at least on some occa-
sions (Lockley et al. 2007a, b; 2009).
Tana della Basura cave near Toirano, Northern Italy is here mentioned separately
due to the intriguing but controversial suggestion that the “human tracks” reported
by Chiapella (1952) are attributable to Neanderthals (Pales 1954, 1960). At present,
despite the skeletal evidence that may be available from various sites to help distin-
guish H. sapiens from H. neaderthalensis, we have no reliable criteria for identify-
ing isolated Neanderthal footprints. As pointed out by Molleson et al. (1972) and by
Onac et al. (2005), the date of this site may be as young as 12,000 yBP, in which
case the Neanderthal claim is questionable. Its morphology certainly does not dis-
tinguish it from a H. sapiens (Fig. 15.5).
The abundance of human tracks at cave sites is perhaps surprising, especially
given the lack of detailed description of many sites. For example, in Ojo Guareña, a
cave near Burgos, Spain, hundreds of footprints are reported, but the site is illus-
trated by only two photographs (Marcos 2001). Dates of 15,600 yBP, suggesting a
Late Pleistocene age, were obtained from carbonized wood. As noted by Lockley
et al. (2008a, b), a “light patina of carbonate” gives the tracks “a very ancient
appearance” (translation of phrases from Marcos 2001, p. 35).
This type of discrepancy between abundance and documentation is understand-
able when considering the problems of access, and the dangers of disturbing the
footprints by walking on them, or otherwise causing damage. Likewise, dating of
the cave site tracks is often difficult due to the lack of suitable materials in the
Fig. 15.5 Footprint from

Tana della Basura cave
near Toirano, Northern
Italy
substrate, the likely differences in age between substrate and tracks, and the preference
for dating other materials such as charcoal which may or may not be contemporane-
ous with the tracks. For example, footprints from Tempranas Cave, near Niembro in
the Llanes region of Asturias Spain (Noval Fonseca 2007) remain undated at the
present time.
Despite the problems involved in obtaining accurate dates, tracks in caves have
good preservation potential and, as noted above, tracks are apparently more abun-
dant, at least at some sites, than the literature might suggest. However, another fac-
tor must be considered—that is the attraction of cave paintings. On the positive side,
efforts to date cave paintings make it possible to infer the age of footprints, assum-
ing there is no strong evidence to suggest that paintings and footprints represent
different phases of activity. On the negative side, paintings distract attention from
other features, such as footprints and charcoal. Obviously, paintings indicate that
caves were frequented by people on foot, even if footprints are not found. Footprints
may occur in caves without paintings or other evidence of human habitation (see
below), but it is impossible to speculate on how common tracks are in caves lacking
other human-produced evidence.
For all the problems and ambiguities that surround the discovery, documenta-
tion, dating, and interpretation of cave site footprints, cave tracks indicate a certain
type of behavior and ecology. For whatever reason, modern humans (H. sapiens)
and perhaps close relatives (e.g. H. neanderthalensis) began to frequent caves
sometime in the Late Pleistocene. This can be considered a significant event in
hominin evolution and, in the most general terms, the evidence seems to suggest a
human impulse to colonize new habitats. This was just one example of the geo-
graphic spread of humans to colonize new regions, such as Australia and the
Americas. In ecologic terms, as explicitly noted by Lockley et al. (2007a, b; 2008a, b),
the nonhuman footprints reported from Late Pleistocene caves are almost exclu-
sively those of carnivores (including bear, hyena, and fox), and stand in contrast to
open-air hominin track sites, where tracks of ungulates and birds are typically
dominant. The aforementioned cave-dwelling carnivores had evidently inhabited
caves long before their habitations were invaded by modern humans. Thus, human
cave-colonization behavior precipitated new “ecologic” interactions between
humans and cave dwelling mammals. While the co-occurrence of footprints of
both groups is tangible evidence of such cohabitation, even more evocative evi-
dence of interaction is found in the archeologic record of paintings and supposed
shrines indicative of “cave bear cults” (a once-popular notion, especially for the
Middle Paleolithic, but long discredited by taphonomic studies [Bahn 2012]).
15.5.3 Vertebrate Ichnology Investigates Cave Paintings
Lockley and Meyer (2000) and Kim et al. (2008a) noted that cave art (or the
more neutral term “painting”) is itself a type of hominin ichnology, as is any
type of engraving. Likewise, as noted by Lockley et al. (2008a, p. 113), “[b]
ecause tracks are a type of symbol or signature of the trackmaker, their artistic
renderings have sometimes been the subject of debate.” Indeed, as noted above,
many human creations can be considered as ichnologic phenomena, including
most forms of sculpture and writing. However, in order to constrain the discus-
sion to the realm of prehistory, we confine our discussion to the implications of
Late Pleistocene evidence, which sheds light on evolutionary events. For exam-
ple, just as footprints indicate the co-occurrence of modern humans and Late
Pleistocene carnivores in caves, so cave art provides direct evidence of the inter-
action of humans with the Pleistocene megafauna. The interest for vertebrate
ichnology is compounded when Paleolithic hominins also depicted the foot-
prints of the animals they were tracking, observing, or hunting (Mithen 1988).
In this regard, it could be inferred that just as tracks are potentially useful as a
census of animals in a particular area (Lockley 1991), so cave art is also poten-
tially a census of animals in a given area in the past. For example, at Closquier
cave, in southern France, there are depictions of what is interpreted as the now-
extinct great auk, “a seabird that could only live in a cold biotope” (Clottes and
Courtir 1996, p. 128). Just as these authors at first found it difficult to interpret
the depiction of the great auk, so too footprints depicted by Late Paleolithic art-
ists have proved difficult to interpret. As noted by Lockley et al. (2009, p. 113),
a particularly “interesting episode in the annals of anthropology was a debate
over whether certain symbols represented tracks or were representations of
female genitalia (Bahn 1986).”
European Cave art, especially from France and Spain, has been the subject of
endless debate. For example, do animal depictions indicate hunting activity,
sympathetic magic, or other shamanistic symbolism? Likewise, one can argue
endlessly as to whether animal and track depictions are realistic or stylized, the
result of superior or inferior artistic ability. Possibly the most significant depic-
tions from an ichnologic view point are these that show the co-occurrence of
animals and the tracks they made. One famous example is the bovids depicted
at Altimira, Spain, where the animals are seen in profile, but their feet are shown
in “plan view” appearing as cloven hoofed tracks stuck on the end of the legs.
While it is obvious from such examples that Late Paleolithic humans could
correlate between animals and their tracks, this is still a highly significant record
from the “dawn of vertebrate ichnology” essentially no different from that pro-
duced by contemporary vertebrate ichnologists who correlate between tracks
and track makers. Thus, our Paleolithic ancestors deserve full credit for being
the authors of the first the vertebrate track field guides (Seilacher 2007).
Whether, we can infer that they only interpreted the track-track maker correlation
in causal terms, the way we do, remains an open question, and we have to
consider the possibility that tracks had other significance in various Paleolithic
cultures (Lockley 1999); see Baucon et al. (2008) for discussion of podomorphs
(footprint representations) and “ichnohierophanies” (traces of religious or
spiritual significance).
15.6 New Intercontinental Travel Frontiers
Archeology and anthropology regard the colonization of Australia and the Americas
by Homo sapiens (if not earlier Homo species), as evidence of the ability of modern
humans to permanently expand their ranges into previously “uninhabited” territory,
by undertaking what we can describe as intercontinental travel. Exactly when and
where the first emigrants broke out of the “old world” to set foot in Australia and
America is unknown. However, these expansions of range seem to have been part of a
pattern that was first manifest with the spread of human ancestors “out of Africa” into
Europe and Asia, including inaccessible regions, such as present day Tibet (Zhang
and Li. 2002) and various islands of the Indonesia archipelago (Morwood et al. 2005).
15.6.1 Into Australia
The colonization of Australia is considered (Bowler et al. 2003) a major event in

hominin evolution, reflecting an ever-increasing ability of humans to colonize new
continents. Humans probably first colonized Australia around 50,000 yBP, showing
their ability to cross the famous Wallace line which separates the marsupial-
dominated faunas of Australia from the placental-dominated faunas of southeast
Asia (Oppenheimer 2009). Webb et al. (2006) and Webb (2007) reported tracks that
have been optically dated to between ~19,000 and 23,000 yBP in the Willandra
Lakes region of southeastern Australia. This is evidently one of the world’s largest
collection of Pleistocene human footprints, with at least 123 footprints, and the site
is part of a property nominated for World Heritage status. The largest tracks mea-
sure 29 cm in length by 10 cm wide. The tracks for which close-up photos have been
published (Fig. 15.6) present an unusual and less than fully modern morphology:
quite flat, exceptionally broad heel, deep and long toe impressions (up to 7 cm) and
a large robust hallux, but lack any clear medial arch and ball impressions (Webb
et al. 2006, Fig. 4).
15.6.2 The Trail to the New World
According to controversial footprint evidence, modern humans may have first set
foot in the New World as early as ~40,000 yBP (Gonzalez et al. 2006a; Huddart
et al. 2008). According to the initial claims of these authors, footprints from a Late
Pleistocene site from Toluquilla Quarry, at Valsequillo, near Puebla Mexico, indi-
cate this reliable 40,000 yBP date for the presence of Homo sapiens in the New
World. However, while the date may be credible, the features interpreted as foot-
prints are controversial, and several of the original authors now admit that these
purported tracks are of questionable origin (Morse et al. 2010). They lack clear
Fig. 15.6 Homo sapiens tracks from the New World and Australia. (a–c) tracks from the
Acahualinca site Nicaragua (after Lockley et al. 2009, Fig. 10), (d) track from Cuatrocienegas,
Mexico (after Gonzalez et al. 2009), (e) track from Willandra Lakes site, Australia
evidence of the big toe (hallux) impression, arch, or differentiated ball and heel
traces. In short they are elongate depressions or traces that may be artifacts pro-
duced by quarry equipment. Renne et al. (2005) referred to these features as
“alleged” footprints and also questioned the dates, instead inferring much older
dates of 1.3 Ma, corroborated by paleomagnetic studies. However, Gonzalez et al.
(2006b) and Huddart et al. (2008) held to their claim arguing that the dating of
Renne et al. (2005) is incorrect.
According to evidence other than footprints humans are not proven to have
been resident in the Americas prior to ~20,000 yBP (Nemecek 2000). Footprints
on the western coast of Canada, dating to 12,500 yBP, may indicate a coastal
route for colonization of the Americas (De Pastino 2015). Tracks from Buenos
Aires Province Argentina (Aramayo and Manera de Bianco 2009), have pro-
duced dates in the range of ~12,000–16,000 yBP. Hominin tracksites reported
from Monte Verde, Chile are associated with dates of 11,500–12,500 yBP
(Dillehay 1999). The rediscovery of a tracksite associated with tufa deposits at
Cuatrociénegas, Coahuila, Mexico has produced tentative dates of ~10,000 yBP

(Gonzalez et al. 2006c, d, 2007, 2009). Two sets of tracks from Cuatrociénegas
have recently yielded U-series dates of 10.55 ± 0.03 ka and 7.24 ± 0.13 ka (Morse
et al. 2010). Given the doubts about the age of the Valsequillo footprints referred
to above, Morse et al. (2014) claimed that the former date represents the oldest
known for any footprints from Mexico. Other tracksites reported from Mexico
(Ordoñez 1945; Aveleyra Arroyo de Anda 1950; Rodríguez-de la Rosa et al.
2004) are poorly known, but likely include mostly Holocene rather than
Pleistocene tracks.
Among the Holocene track sites listed by Lockley et al. (2007a, b; 2008a, b),
two sites from La Olla and Monte Hermoso Argentina dated at ~7000 yBP
(Bayon and Politis 1996, 1998; Aramayo and Manera de Bianco 2009; Bayón
et al. 2011) are significant, and comparable in age to a site from Laguna La María
(near Villa Cañás) dated to ~8000 yBP. Abundant well-preserved tracks from the
footprint museum at Acahualinca, (Huellas de Acahualinca) Managua, Nicaragua
(Flint 1883; Brinton 1887) have been 14C-dated at 5945 ± 145 yBP (Bryan 1973),
at 6500 yBP by Bice (1979), and between 2000 and 6000 yBP (Schmincke et al.,
2005, 2007, 2008, 2009). The site is now the type locality for Hominipes moder-
nus (Kim et al. 2008b) and preserves the trackways of at least 15 individuals, an
ungulate, a possum and a bird (Lockley et al. 2007a, b; 2008a, b). Another nearby
site, known as El Recreo (Williams 1952), has yielded bison and tapir tracks.
Plant remains are also reported (Brown 1947). The Oro Grande Site near
Victorville, southern California (Rector 1979, 1983, 1999), has given a 14C date
of 5070 ± 120 yBP (Rector 1983) for tracks of at least four individuals, where
tracks of raccoon, coyote, and ungulates were also documented.
Haberland and Grebe (1957) reported a tracksite from El Salvador tenta-
tively dated between ~1200 and 1800 yBP. Undated footprints were also
reported from a volcanic deposit near Guaimaca, Honduras (Veliz 1978 ).
Similarly sparse information comes from a report ( Anon undated) of a cave
site from Naj Tunich, Guatemala, with footprints attributed to indigenous
Mayan inhabitants tentatively dated on the basis of artifacts, not footprints,
between ~1450 and 1950 yBP.
To complete the New World track record, we may refer to Willey et al.
(2009), who noted footprints reported from Pocket Cave in Arizona, and dated
at ~1450–1500 yBP based on dendrochronology. Finally, footprints from east-
ern North America are associated with underground caverns, such as Jaguar
Cave, Tennessee, Unknown Cave, Kentucky, third Unnamed Cave, Tennessee,
Fisher Ridge Cave, Kentucky, Mud Glyph Cave, Tennessee, Sequoyah Caverns,
Alabama, Footprint Cave, Virginia, and Lon Odell Memorial Cave, Missouri
(Watson et al. 2005; Willey et al. 2005, 2009). Collectively, these cave sites date
from between 4695 ± 85 and ~400 yBP.
15.6.3 Other Exploratory Trails
As summarized by Lockley et al. (2008a, b), Zhang and Li (2002) and Zhang et al.
(2003) reported a series of hand and footprints associated with calcareous tufa
deposits at an elevation of 4200 m on the Tibetan Plateau optically dated at about
20,000 yBP. This unusual combination of hand and footprints may be related to the
site being a hot springs with a hearth, therefore presumably used as a campsite,
rather than an area simply passed through.
Late Pleistocene footprints dated at about 19,000–25,000 yBP from Jeju Island,
Korea (Kim and Kim 2004a, b; Kim et al. 2004, 2009, 2010) indicate that humans
were exploring remote islands at about the same time that they were exploring the
Tibetan Plateau. As a coastal site, Jeju provides evidence of a variety of bird and
mammal tracks diverse invertebrate traces and body fossils, at multiple levels, sug-
gesting an ecology obviously different from that found in Tibet. Nevertheless,
despite the geographic and ecologic differences between these two Asian sites, they
both provide striking evidence of the ability of Late Paleolithic humans to explore
habitats that had not previously been frequented by hominins.
15.7 The Prelude to History
15.7.1 Life on the Sea Shore
Lockley et al. (2007a, b, 2008a, b) have already listed known hominin track sites
that bridge the gap between unequivocally Pleistocene and prehistoric Holocene
sites, to those that yield comparatively recent dates that bring us into historic time.
It is not necessary to repeat detailed accounts of these sites which are already widely
distributed. However, a brief summary allows us to pick out features that highlight
the utility of footprints in interpreting “events” in historic time.
Among the earliest Holocene tracks to fit in our post-Pleistocene category, we
can cite abundant human footprints from near the Sebkra el Azrag, Mauritania
(Mafart 2006). These date from ~9000 yBP and are associated with footprints of
elephants, hippopotamus, giant eland, and bovids.
Younger Holocene footprint sites are known from near Clare Bay, in South
Australia (Belperio and Fotheringham 1990), estimated to be ~5000
yBP. Undocumented tracks are also reported from near Broome in western Australia
(Long 1998; Baldwin personal communication 2011). Both are coastal sites. Other
significant coastal hominin tracksites are known in Britain (Aldhouse-Green et al.
1995; Cowell et al. 1993; Roberts et al. 1996; Roberts 2009; Doyle 2007; Bennett
et al. 2010a, b). An assemblage of exceptionally preserved footprints of between six
and nine children (ranging in length from 114 to 206 mm) has been described from
Walvis Bay, Namibia (Bennett and Morse 2014). They date to 1500 yBP and are
preceded by the tracks of an apparent flock of domesticated sheep/goats.
Human footprints reported from Rawthey Cave, Cumbria, England (Chamberlain

et al. 1997) could be as recent as the 14th century AD. Some coastal sites (Patton
1993) have only the tracks of domestic animals. Human tracks have also been
reported from Holocene beach rock in Greece (Bromley et al. 2009).
The coastal situation of many Holocene track sites seems to reflect two factors.
(1) Suitable environments for track preservation (e.g., estuarine and beach sub-
strates) and (2) preferred habitats or foraging sites of humans. We know from other
lines of evidence, such as shell middens, that foraging was an important activity at
such sites.
15.7.2 In the Shadow of Volcanoes
Human tracks are reported from numerous agricultural sites (rice fields) in Japan,
(Harada and Noto 1984) that date from ~720 to 1600 yBP. At many of these Japanese
sites, volcanic ash played an important role in track preservation. Human tracks also
occur at various stratigraphic levels in volcanic ashes dated at 1400 AD on Motutapu
Island, New Zealand. Nichol (1982) suggested that track makers were evidently not
deterred by the continuing ash falling over the area. Traces of digging sticks suggest
that individuals were digging gardens.
Footprints from Hawaii Volcanoes National Park site are from two distinct
footprint-bearing horizons, the younger precisely dated to a 1790 Kilauea eruption
and ashfall (Meldrum 2004a, b; Moniz Nakamura 2009). The record of modern
pedestrian trackways in ash of remarkably similar physical and chemical character-
istics as the Laetoli ashfall, provides in the Hawaiian footprints a meaningful com-
parison and contrast to the Laetoli hominin footprints. This contrast highlights the
diagnostic distinctions between Praehominipes laetoliensis and Hominipes moder-
nus (Meldrum 2004a, b; Meldrum et al. 2011). Because of the drama associated
with volcanic eruptions, various scenarios regarding the relationship between ash
fall and track-making activity have been proposed, not only for the Hawaiian sites
(Jaggar 1921, 1934; Meldrum 2004a, b; Mayor and Sarjeant 2001), but also for the
aforementioned Nicaraguan site (Schmincke et al. 2005, 2007, 2008, 2009) and the
New Zealand levels (Nichol 1982).
Regarding the Hawaiian sites, Moniz Nakamura (2009) suggested footprints were
made in the area both before and after the eruption, thus indicating that individuals
were not deterred by the ash fall and not fleeing or about to perish as a direct result of
the eruption. Were indigenous populations accustomed to repeat volcanic activity, and
undeterred by smaller eruptions? Possibly, but sometimes large eruptions did create
truly dramatic, life-threatening scenarios as in the case of the famous 79 AD eruption
of Vesuvius. Unlike this eruption, which is not associated with footprints, footprints
are found in association with deposits created by the 3780 yBP eruption of Vesuvius,
known as the Avellino plinian eruption, that fell on Nola, a Bronze Age village near
Naples, Italy (Mastrolorenzo et al. 2006). These devastating ash falls created a remark-
able footprint record NNW of Vesuvius where “thousands of footprints [are] directed
NNW away from the volcano.” These “testify to an en masse exodus from the devas-
tated zone” (Mastrolorenzo et al. 2006, p. 4368). Footprints registered on “all horizons
in the ash bed” indicating that “the evacuation occurred during the settling of the surge
cloud.” “Flood and lahar deposits overlying the surge bed also include footprints and
local raindrop imprints as well, thus testifying that the ongoing exodus occurred dur-
ing both the ash fall and the post-eruption rainstorms and floods” (Mastrolorenzo et al.
2006, p. 4368).
From these accounts of human tracks in volcanic ashes, we can infer that, at the
times indicated above, humans lived in volcanic terrains, as they do in some areas
today, risking the dangers inherent in such environments. Although we do not know
exactly how they reacted to localized or small eruptions, the evidence at some sites
(New Zealand, Nicaragua, Hawaii) suggests they were not sufficiently deterred by
the threats, to vacate these areas. Thus, on occasion they did not evacuate, or move
very far from small eruptions that created small ash falls. However, as the Nola
eruption indicates, humans reacted to large eruptions and ash falls by fleeing,
although in this case escape was not possible and individuals literally died in their
tracks.
15.8 Vertebrate Ichnology Transcends Planet Earth
One of the major events in human history was the landing of humans at six sites on
the Moon (Fig. 15.7). Arguably, this is a major evolutionary event akin to the migra-
tion of a species onto a new continent on which that species had not previously set
foot. Just as the fossil record, including the track record, provides evidence of such
migrations, so the artifacts and tracks and traces left by humans on the Moon also
provide unequivocal evidence that the range of living humans has extended from one
celestial body (planet Earth) to another (its moon). In the case of the machines that
humans have landed on Mars, the artifact and trace fossil evidence is slightly differ-
ent, and it is intriguing to speculate as to how it might be interpreted and compared
with the lunar evidence, by someone unfamiliar with human history (Lockley 1999).
While most humans regard the Moon landing as a momentous event in human
history, it is perhaps too recent an event to have had its evolutionary significance
evaluated in the context of deep time. In any event, the track making activity of
humans on the Moon, while creating visually spectacular traces, is so distinctive
as to have been treated as a special event, unrelated to the long history of hominin
track making on Earth. Nevertheless, while it is legitimate to regard lunar and ter-
restrial tracks records as two quite separate ichnologic records, in obviously dif-
ferent substrates, environments, and geographic locations, they are still
indisputable parts of the continuum of an expanding hominin track record
(Fig. 15.6). So, what are the implications?
Humans are the only large vertebrate species to have set foot on the Moon. In this
regard, they crossed a new frontier in much the same way as they did when they
cross the Wallace line between Asia and Australia, or the Bering Straits between
Fig. 15.7 The dispersal of hominins from the old world to the new world and Moon as demon-
strated by the footprint record, is in broad agreement with the evidence obtained from the record
of body fossils and artifacts
Asia and North America. The principle is succinctly stated by Morwood and
Oosterzee (2007, p. 182): “Modern humans were the only large Asian animals that
made the west-to-east crossing to Greater Australia on their own account.” Here,
these authors evidently imply that the migration into Australia was not accidental,
as in the oft-cited, but speculative, explanation of dispersal of animals, rafted by
accident from one land mass to another. Lockley (1999) described some of the con-
figurations of human traces on the Moon. First, of course they are limited to a very
small area, and secondly they loop out from the lunar module and back. Secondly
the total census of individual trackways is known to represent only 12 individuals.
An intelligent species analyzing the footprint evidence might correctly infer that
very few individual track makers, of a distinctive bipedal species, had visited the
Moon, but had been unable to explore very far. The track makers were not adapted
to colonize extensively.
It is perhaps unduly speculative to consider how another intelligent species
might interpret the trails of the lunar rover, the actual rover itself, or the trails and
machines on Mars (Lockley 1999). We know that the Martian traces were made
without humans being present on the planet, but would this be deduced by an intel-
ligent species unaware of the details of an Earth-based space program? It is perhaps
debatable as to whether traces left by robotic machines legitimately fall in the cat-
egory of biogenic sedimentary structures. Clearly, feet and shod foot traces repre-
sent human functional anatomy and behavior, but lunar and Martian rovers represent
a different type of function, only indirectly related to human movement. Since these
robotic machines are tools, we argue that in principle the traces they leave are simi-
lar to those produced by other human-manufactured tools that register traces on a
variety of substrates.
In conclusion, as noted below, a broad-based interpretation of hominin ichnol-
ogy indicates that there have been at least four major threshold events that starkly
punctuate the ichnologic record. The first was bipedalism, the second was the spread
of hominins to Australia and the New World, the third was painting, sculpture, and
tool making, associated with extensive cave exploration, and the fourth was the
ability to set foot on other celestial bodies.
15.9 Discussion
At least 65 hominin tracksites are reported in the literature (Lockley et al. 2007a, b,
2008a, b) of which about one third (~24) are Late Pleistocene or older. Only a few
(4) sites represent pre-sapiens track makers. The advent of erect posture and gait
was clearly a major event in hominin evolution. However, the question of whether
such features as midfoot morphology (flexibility vs. a fully modern arch) and sepa-
ration of the big toe (digit I) from traces of digits II–V, and their different inferred
lengths, constitute evidence of major evolutionary changes remains open to ques-
tion, and may not be fully resolved without additions to the body and trace fossil
records. Debates about the affinity of the 3.6-million-year-old Laetoli track maker
are made partly on the basis of the age of the sites and contemporary body fossils.
However, they are also at least partially based on footprint morphology, when cor-
related pedal skeletal fossils are known. Tuttle et al. (1990) inferred that the Laetoli
track maker may have been an as-yet-unknown hominin indistinguishable from
modern H. sapiens, and therefore not an australopithecine as inferred by Stern and
Susman (1983; see also Suwa 1984). In contrast, Meldrum (2006, 2007a) pointed to
evidence of mid-tarsal pressure ridges and extrusion fronts, in the GI trail indicating
ape-like midfoot flexibility largely if not altogether absent in modern humans (but
see DeSilva and Gill 2013). The distinctions in the Laetoli tracks when compared to
modern unshod pedestrians are expressed in the introduction of the new ichnotaxon
Praehominipes laetoliensis (Meldrum et al. 2011) in contrast to footprints of fully
modern humans, designated as Hominipes modernus (Kim et al. 2008a, b). However,
if we exclude the overprinted trackways (G2 and G3), and possibly a fourth (G4)
according to Musiba et al. (2011), the Praehominipes sample of un-obscured foot-
prints remains small consisting of only the G1 trackway (Fig. 15.1).
The inference of an H. erectus (H. ergaster) trackmaker, in the case of the poorly
preserved Koobi Fora footprints (Behrensmeyer and Laporte 1981), is based on age.
The Ileret footprints also exhibit insufficient detail to infer modern footprint mor-
phology with certainty. This inference is contra Bennett et al. (2010a), who con-
cluded that these footprints provide “the oldest evidence of an essentially modern
human-like foot anatomy, with a relatively adducted hallux, medial longitudinal
arch, and medial weight transfer before push-off.” We find support for only one of
these three points (i.e. a relatively adducted hallux). We find no evidence to support
the consistent presence of a longitudinal arch or for a medial weight transfer.
Preservation is poor, and the actual topography of the contact surface is indiscern-
ible. Even in footprints as young as 300,000–400,000 years old, such as the Terra
Amata footprint (De Lumley 1966, 1967; De Lumley et al. 2011), evidence of a
modern arch is lacking (Meldrum 2004a, b; 2006). Likewise, tracks from the Middle
Pleistocene Roccamonfina Volcano site in Italy (Mietto et al. 2003; Avanzini et al.
2004, 2008), which falls in the same age bracket as the Terra Amata footprint, are
not sufficiently well-preserved to determine if a fully modern arch is present.
Thus, given the small size and quality of the pre-Late Pleistocene footprint
sample, few universally accepted conclusions can be drawn regarding the possi-
bility of diagnostic differences between tracks made by various hominin species
of early to middle Homo. However, the strong possibility remains that much of the
hominin history of bipedalism took place on flat flexible feet in contrast to the
modern human foot form of relatively recent vintage, as characterized by the ich-
nospecies Hominipes modernus (Kim et al. 2009). The scarce fossil record of the
distal hallucal metatarsal, a key element in the modern longitudinal arch, clearly
points to this interpretation.
Distinguishing between the tracks of Homo sapiens and the comparatively
unknown track record of H. neanderthalensis on the basis of footprint morphology
is evidently not reliable at present, despite a few claims to the contrary (Onac et al.
2005). It stands to reason that the robusticity of the neanderthal skeleton would
produce footprints with relatively larger breadth to length ratios. However, the sam-
ple of Neanderthal footprints is too meager to test this prediction.
As noted by Lockley et al. (2008a, b), although the majority of known sites have
assigned ages, in many cases the dates are uncertain, and the literature indicates
frequent age date revisions, such as in the case of the Turkish and Nicaraguan sites.
These authors also discussed the respective proportion of outdoor or open-air sites
versus cave sites as about 65 % versus 35 %. Likewise, the proportion of sites in
well-documented volcaniclastic/pyroclastic rather than non-volcaniclastic sub-
strates is discussed by Houck et al. (2009).
The study of hominid traces raises philosophic issues that deal with the contro-
versial question of the “exclusivity” of the human species in comparison with other
vertebrates. Many scientists and philosophers argue for and against the idea that
humans are fundamentally different from other species (see Guldberg 2010 for dis-
cussion of both sides of this debate). The traditional argument in favor of human
exclusivity is that we have language, self-awareness, and culture, not to mention the
power to change the environment in ways that other species cannot. This is not to
say that other microbial, plant, and animal species have not changed environments
dramatically, but in different ways. We also recognize that the exclusivity argument,
with respect to tool use, language, and self-awareness, has been challenged in cases
were such attributes have been reported for other species (Galef 2003, 2009; Rendell
and Whitehead 2001). However, detailed discussion of these debates is beyond the
scope of this paper. As the foregoing discussions emphasize, an objective look at the
hominin track record indicates that the ichnologic record of H. sapiens is unlike that
of any other species, with respect to many features including, geographic distribu-
tion, diversity of traces registered, substrates on which traces are registered, and
behavioral implications of traces (e.g., Hasiotis et al. 2007; Baucon et al. 2008; Kim
et al. 2008a). Again, in the context of this review of footprints and other traces cre-
ated by sapiens and pre-sapiens hominins, it is clear that, even though foot and hand
prints may be similar among all hominins, the diversity and behavioral implications
of most other sapiens-produced traces is different from those of pre-sapiens homi-
nins and other vertebrates both in absolute and relative terms (i.e. they are both
differences of “degree and kind”). Thus, the use and creation of stone, bone and
wood tools, and artifacts, while not entirely unknown in pre-sapiens hominins or
other vertebrate species, reaches a degree of complexity indicating behaviors and
cultural shifts that truly represent major evolutionary advances during the sapiens
phase of hominin prehistory. This shift in turn heralded entirely novel and ulti-
mately historically documented additions to the anthropologic and trace fossil
records, such as ceramics, metal-work, and writing which have no pre-sapiens pre-
cursors (Hasiotis et al. 2007; Baucon et al. 2008; Kim et al. 2008a). In short, as
hominins have evolved over the last ~4 million years, their ichnologic record has
become increasingly well differentiated from that of their ancestors, a process that
shows its most dramatic acceleration or shift after about 30,000 yBP.
While prior to that date multiple hominin species, a half dozen or more, coex-
isted across the landscape at any given time it is generally inferred that since
~30 kyBP, H. sapiens has been the only or “exclusive” extant hominin species
However, there is a growing appreciation for the bushiness of the hominin (even
hominoid) tree, as well as accumulating examples of quite recent persistence of
individual branches of said tree, as evidenced in the fossil record (Meldrum 2012a,
b). Is it justified to simply assume we are “the last hominin standing”? Or is there a
case to be made for the possible existence of “relict hominoids” today? The discov-
ery of the extraordinarily recent remains of Homo floresiensis, combined with the
acknowledgement of historical anecdotes of encounters with little hairy “people” in
the mountain forests of Flores should have made that point clearly enough (e.g.,
Forth 2012).
There is considerable ichnologic evidence for the existence of relict hominoids.
By far the most extensive footprint evidence is attributed to the so-called “Bigfoot”
(or sasquatch) of North America, as well as reports of similar tracks in Asia
(Meldrum and Gouxing 2012), where the comparatively recent existence of the
giant hominoid Gigantopithecus is unequivocally accepted (Meldrum 2004b, 2006,
2007b and references therein). Meldrum (2007b) formally named and diagnosed the
alleged sasquatch tracks as Anthropoidipes ameriborealis.
While it is outside the scope of the present review to explore this intriguing but
controversial issue further, it is fair to say that the possible existence of relict homi-
noids represents a significant dimension in hominid evolution. While most anthro-
pologists ignore or refute the existence of relict hominoids without detailed analysis
of the evidence at hand, there is nevertheless an extensive literature on the subject
including a number of books by bona fide scientists who have taken the evidence
seriously (see Lockley 1999 and Meldrum 2006 for reviews). Did Gigantopithecus
or some similar large bipedal relict hominoid (such as a form of paranthropine)
cross from Asia to North America, along with Homo sapiens, during the Pleistocene,
along with 75 % of the mammals now considered endemic to this continent? And if
so, what sort of ichnologic evidence might we expect to find? Are contemporary
reports of the tracks of a giant hominid in North America’s remote mountain forests,
or those of a diminutive “hobbit” in the jungles of southeast Asia, any more surpris-
ing than H. sapiens tracks on the Moon, or 25,000 year old tracks alongside pictures
of Great Auks in a deep cave in the South of France, or 20,000 year old tracks on
the Tibetan plateau at the height of the Ice Age?
Finally, it is relevant to consider the extent to which Homo sapiens’ intentional
behavior is affecting the track record as a whole. While large scale megalopolis
construction and landscape alteration will surely leave an ichnologic footprint that
dwarfs such relatively local traces as Japanese rice paddies, it will at the same time
remove or diminish the track record of many other species, by erasing or modifying
substrates. This process could be considered similar, or analogous, to other natural,
non-human-induced, processes such as erosion, that lead to the differential
preservation or destruction of trace-bearing deposits. While such human-generated
traces may vastly modify natural cycles of track registration, preservation, exhuma-
tion, and destruction, humans also intentionally preserve the ancient track record.
This is done not just by preserving fossil footprints in museums, but in the creation
of a symbolic, documentary record (a type of ichnologic record: e.g., this book) of
extinct species and lost cultures that would not otherwise exist without human
intentionality. Thus, Homo sapiens has created a wide array of highly distinctive
and “exclusively” human traces.
15.10 Conclusions
We conclude that the track record has significant implications for our understanding
of major events in hominin evolution and can be summarized in two categories:
major and minor events. Major events include the following:
1. The advent of bipedalism, confirmed by the Laetoli site, and dated not later than
~3.6 Myr.
2. The arrival of modern humans in Australia and the New World (the Americas),
confirmed by footprints and other archeologic evidence 15,000–20,000 yBP. A
date of ~40,000 yBP has been suggested for colonization of Australia, but in the
case of the Americas such an early date, although claimed in one case, is contro-
versial and dubious.
3. The ichnology manifest in the creation of art (painting, sculpture, etc.,) and tools
in the Late Paleolithic, especially between ~30,000 and 10,000 yBP, indicates
that humans underwent a “cultural revolution” at this time. They made extensive
use of underground cave sites, leaving 2D and 3D art and artifacts and, in one
case, a set of footprints indicating a game or ritual.
4. The extraordinary Holocene diversification of culturally- and technologically
produced large and small scale traces culminated in the arrival of track making
Homo sapiens on the Moon, and the generation of traces on Mars by human-
manipulated machines.
A common theme of events 2–4 is geographical expansion of the Homo sapiens,
and a steady increase in the diversity and size range (both large and small) of trace
fossils produced.
Other minor but significant events recorded in the hominin track record can be
listed as follows:
1. Inferred significant modifications in the morphology of the hominin foot and
corresponding footprints between 3.6 million and ~50,000 yBP. The extent to
which these modifications are evident in the track record is debatable, but there
are two distinct polar morphologies (Praehominipes and Hominipes) now docu-
mented in the ichnologic literature.
2. Morphologic distinctions between inferred H. sapiens and H. neanderthalensis
footprints could be of significance, if unequivocally established, but at present,
claims of inferred differences are poorly documented.
3. The movements to, or colonization of, remote sites, such as the Tibetan Plateau
and remote archipelagos, such as Jeju Island, Korea, or New Zealand, are dem-
onstrated by footprints which are as old as, or older than, other archeologic evi-
dence. As such, these represent significant events in the history of prehistoric
human exploration, at least on the regional scale.
4. The colonization and use of caves and rock shelters as art galleries, often depicting
animals, represents a significant development in human–animal interactions.
5. An extraordinary diversification and complexification of human-produced trace
fossils since the Late Pleistocene and Early Holocene has allowed paleontolo-
gists to expand the academic definition of vertebrate ichnology into realms tradi-
tionally considered the province of archeology and anthropology.
6. This recent complexification of the trace fossil record by Homo sapiens
activity has profound implications for how recent, present, and future traces
will be registered as dominant components of the deep time ichnological
record. In this regard humans are presently creating an ichnological revolu-
tion with significant implications for debate about the global impact and
exclusivity of our species.
Acknowledgements We thank Gabriela Mángano and Luis Buatois, University of

Saskatchewan, for their invitation to contribute this article and their help and patience during the
editing process. We also thank Neffra Matthews for the photogrammetric work done on the
Laetoli tracks illustrated in Fig. 15.1, and first published in Kim et al. (2008b, Fig. 7). We par-
ticularly thank Andrea Baucon, Geology and Paleontology Office, Centro Cultural Raiano,
Idanha-a-Nova, Portugal and Gustavo Politis, Facultad de Ciencias Sociales, UNCPBA,
Olavarria, Argentina for their helpful reviews.
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Soc of America abstracts of 108th annual meeting 44. p 13
Chapter 16
Recurrent Patterns and Processes:
The Significance of Ichnology
in Evolutionary Paleoecology
Luis A. Buatois and M. Gabriela Mángano
16.1 Introduction
Gould (1980) established a distinction between idiographic and nomothetic paleobi-

ology. Whereas the former deals with individual evolutionary events, the latter
attempts to find regularities in the history of life. This book is rooted in an idio-
graphic approach because most chapters review a single major evolutionary event.
However, after gathering such a wealth of information on specific events, it is worth
to close the book with a brief attempt at exploring the nomothetic perspective by try-
ing to find recurrent patterns and processes in evolutionary paleoecology using ich-
nologic data. In fact, as noted by Gould (1980) himself regarding J. J. Sepkoski’s
(1978, 1979, 1984) kinetic model of diversification, the approach employed here
attempts to reach a balance between nomothetics and idiographics by producing a
nomothetic model which is empirically grounded (see D. Sepkoski 2012 for an
insightful review). Therefore, this chapter compares several evolutionary events,
searching for common themes. In order to do so, we will compare organism-substrate
interactions during evolutionary radiations, benthic fauna response to mass extinc-
tions, patterns of ecospace colonization, and environmental shifts through time.


DOI 10.1007/978-94-017-9597-5_16
450 L.A. Buatois and M.G. Mángano
16.2 Evolutionary Radiations
Evolutionary radiations are dramatic proliferations of taxa within clades (Erwin 1992;
Simões et al. 2016). From an ichnologic perspective, three major events in marine
settings are compared in this book and are, therefore, addressed in this chapter, the
Cambrian Explosion (see Chap. 3), the Great Ordovician Biodiversification Event
(see Chap. 4) and the Mesozoic Marine Revolution (see Chap. 9). Figures 16.1 and 16.2
summarize changes in ichnodiversity and ichnodisparity through time, respectively.
Only invertebrate trace fossils were considered. Ichnogeneric occurrences were
compiled on a case-by-case basis, therefore summarizing actual occurrences.
However, the curves were plotted as “range-through” data, and therefore they are
based on lower and upper appearances for each ichnogenus/category of architec-
tural design and then extrapolating their presence through any intervening gap in the
continuity of its record. This allows eliminating the noise generated by small-scale
fluctuations in ichnodiversity and ichnodisparity that may result from the uneven
number of studies for each time span. However, a disadvantage of this approach is
that it is not particularly useful to detect potential drops in ichnodiversity and ichno-
disparity as a result of mass extinctions (see Sect. 16.3).
Compilation of ichnogeneric diversity through geologic time indicates that the
Cambrian Explosion, the Great Ordovician Biodiversification Event and the
Mesozoic Marine Revolution, all evolutionary radiations established based on body
fossils, are associated with increases in ichnodiversity (Fig. 16.1). This pattern
shows a remarkable consistency with our knowledge derived from the analysis of
diversity based on the body-fossil record. In fact, similarities between the curve of
Fig. 16.1 (continued) were compiled at the ichnogenus level because the taxonomy is more firmly
established than for ichnospecies. Total number of ichnogenera is 523 ichnogenera (417 for biotur-
bation structures and 106 for bioerosion structures). Synonymies have been checked to make ichno-
taxonomy consistent. Individual curves were constructed for continental, shallow-marine and
deep-marine bioturbation ichnotaxa, and marine and continental bioerosion. In turn, separate ichno-
diversity curves were compiled for all marine bioturbation ichnogenera, all marine ichnogenera
(bioerosion plus bioturbation), and all continental ichnogenera (bioerosion plus bioturbation). In
order to differentiate between shallow marine, deep marine and continental occurrences, original
paleoenvironmental interpretations in the literature, as well as personal data, were considered.
Curves plotted as “range-through” data, and therefore based on lower and upper appearances for
each ichnogenus, extrapolating their presence through any intervening gap in the continuity of its
record. Temporal resolution is at Epoch rank. Because there are no formal subdivisions for the
Ediacaran Period, this bin has been further divided using the informal subdivision of Vendian (560–
550 Ma) and Namibian (550–541 Ma) (sensu Jensen et al. 2006). Biogenic structures recorded in
modern sediments have been included in the Holocene. Although it occurs in both hardgrounds and
firmgrounds, Gastrochaenolites was considered under bioerosion. The record of mirobioerosion
ichnotaxa extends further into the Precambrian with Granulohyalichnus and Tubulohyalichnus rang-
ing into 3.5 Ga (Furnes et al. 2004, 2007; Banerjee et al. 2006, 2007; McLoughlin et al. 2009), rep-
resenting in fact the oldest trace fossils. Because these two ichnogenera are produced by microbial
activity and our analysis is centered on invertebrates, they have not been considered in the discussion
presented in the text
16
Recurrent Patterns and Processes…
Fig. 16.1 Ichnodiversity changes through geologic time. Compilation based on literature and personal data. The main literature sources used to compile the ich-
nogeneric list were the Treatise volume on trace fossils (Häntzschel 1975) and the more recent revision by Knaust (2012). In addition, we have checked each
ichnogenus by consulting the original ichnologic literature and pertinent reviews in order to revise its validity and potential synonyms. The ichnodiversity curves
451
16 Recurrent Patterns and Processes… 453
marine ichnodiversity and that of marine animal genera produced by Sepkoski

(1997) are striking with regards to the Cambrian Explosion and the Great Ordovician
Biodiversification Event (Buatois et al. 2016a). A 433 % increase in ichnodiversity
took place in marine environments during the Terraneuvian (12 and 64 ichnogenera
in the Ediacaran and Terraneuvian, respectively), whereas a 77 % occurred as a
result of the Great Ordovician Biodiversification Event (82 ichnogenera in the
Furongian and 145 ichnogenera by the Late Ordovician). Whereas the Cambrian
Explosion is essentially restricted to bioturbation structures, the Great Ordovician
Biodiversification Event is expressed by bioerosion structures as well, an event that
has been referred to as the Ordovician bioerosion revolution (Wilson and Palmer
2006). Ichnogeneric compilations show that the explosion in bioerosion took place
approximately 80 my after the Cambrian Explosion in bioturbation (Buatois et al.
2016a). The explosive diversification of the early Cambrian supports an exponential
model for the early phases of diversification as advocated originally by Sepkoski
(1978) based on shelly fossils. Despite these similarities, two main differences are
apparent. First, trace-fossil data indicate that the rapid diversification took place in
the early Cambrian, rather than in the late early Cambrian as indicated by shelly
fossils, suggesting that the Fortunian may be regarded as part of the phylogenetic
fuse (Mángano and Buatois 2014; see Chap. 3). Second, marine animal genera show
minor diversity fluctuations during the middle to late Cambrian, but ichnodiversity
reached a plateau that spanned from the middle Cambrian to the onset of the Great
Ordovician Biodiversification Event (Buatois et al. 2016a).
The ichnodiversity curve for marine trace fossils shows another, but more modest,
increase in the Early Jurassic (8 %; 165 and 178 ichnogenera in the Late Triassic and
Early Jurassic, respectively) and a more important one in the Late Cretaceous (19 %;
187 and 223 ichnogenera in the Early and Late Cretaceous, respectively) (Fig. 16.1).
A more limited, but constant, increase took place between the Early Jurassic and
Early Cretaceous. These changes in ichnodiversity are expressed by both bioerosion
and bioturbation structures. This increase is undoubtedly a reflection of the Mesozoic
Marine Revolution. Comparing ichnodiversity levels between the Late Triassic
(164 ichnogenera) and the Late Cretaceous (223 ichnogenera) reveals a total 35 %
ichnodiversity increase that is attributed to this evolutionary radiation. Ichnologic
data indicate that the Early Jurassic experienced a change in ichnotaxonomic com-
position and the complexity of infaunal tiering, which is consistent with the timing
of the Mesozoic Marine Revolution as revealed by our compilations (see Chap. 9).
Fig. 16.2 Ichnodisparity changes through geologic time. Compilation based on literature and per-
sonal data. Total number of categories of architectural designs is 79 (58 for bioturbation structures
and 21 for bioerosion structures). Curves were constructed for ichnodisparity following the same
environmental subdivision used for ichnodiversity. Curves plotted as “range-through” data, and
therefore based on lower and upper appearances for each category of architectural design, extrapo-
lating their presence through any intervening gap in the continuity of its record. Ichnodisparity was
compiled following the approach of Buatois and Mángano (2013; see also Chap. 1) and categories
of architectural designs summarized by Buatois et al. (2016b). Table 16.1 summarizes the different
categories and the ichnogenera included. For further information, see caption of Fig. 16.1
Table 16.1 Categories of architectural designs and their ichnotaxa in alphabetical order (after
Buatois et al. 2016b)
Architectural designs Ichnogenera
1-Simple horizontal trails Archaeonassa, Circulichnis, Cochlichnus,
Gordia, Helminthoidichnites, Helminthopsis,
Herpystezoum, Mermia, Talitrichnus
2-Trilobate flattened trails Curvolithus, Trisulcus
3-Chevronate trails Protovirgularia, Rhadhostium
4-Trails with undulating transverse bars and Climactichnites, Steinsfjordichnus
furrows
5-Bilobate trails and paired grooves Carpatichnis, Cruziana, Davichnia,
Didymaulichnus, Diplopodichnus,
Taphrhelminthoides, Tumblagoodichnus
6-Trackways and scratch imprints Acanthichnus, Acripes, Allocotichnus,
Angulichnus, Arachnomorphichnus, Arcichnus,
Asaphoidichnus, Biformites, Bifurcatichnus,
Bifurculapes, Climacodichnus, Coenobichnus,
Conopsoides, Copeza, Danstairia,
Dendroidichnites, Dimorphichnus, Diplichnites,
Etterwindichnus, Euproopichnus,
Foersterichnus, Glasbachichnium,
Glaciichnium, Hamipes, Harpepus,
Harpichnus, Heftebergichnus,
Heteropodichnus, Heterotripodichnus,
Hexapodichnus, Homopodichnus, Irichnus,
Ixalichnus, Kalnaichnus, Keircalia,
Kivanichnus, Konbergichnium, Kouphichnium,
Lineatichnus, Lithographus, Lusatichnium,
Maculichna, Megapodichnus,
Merostomichnites, Mirandaichnium,
Mitchellichnus, Monomorphichnus,
Multipodichnus, Octopodichnus,
Oklahomaichnus, Oniscoidichnus,
Orchesteropus, Paleohelcura, Palmichnium,
Parahamipes, Pentapodichnus, Petalichnus,
Pirandikus, Protichnites, Pterichnus,
Punctichnium, Siskemia, Stiallia, Stiaria,
Striatichnium, Tambia, Tarichnus, Tarsichnus,
Taslerella, Tasmanadia, Telsonichnus,
Teratichnus, Terricolichnus, Tortilichnus,
Trachomatichnus, Umfolozia, Vadichnites,
Warvichnium
7-Bilaterally symmetrical short, scratched Avolatichnium, Arborichnus, Chagrinichnites,
imprints Cheliceratichnus, Gluckstadella,
Huilmuichnus, Kingella, Orbiculichnus,
Pollichianum, Ramosichnus, Rotterodichnium,
Solusichnium, Surculichnus, Tonganoxichnus,
Tripartichnus
(continued)

8-Bilaterally symmetrical short, scratched Aglaspidichnus, Alph, Cardioichnus,
impressions and burrows Cheiichnus, Craticulichnum, Crescentichnus,
Faciemichnus, Limulicubichnus,
Raaschichnus, Rusophycus, Selenichnites,
Svalbardichnus
9-Fan-shaped to radiating scratched imprints Kimberichnus, Radichnus
10-Passively filled horizontal burrows Didymaulyponomos, Palaeophycus
11-Simple actively filled (massive) horizontal Furculosus, Macaronichnus, Nenoxites,
to oblique structures Planolites, Sericichnus, Torrowangea,
12-Simple actively filled (meniscate) Ancorichnus, Beaconites, Compaginatichnus,
horizontal to oblique structures Entradichnus, Imponoglyphus,
Jamesonichnites, Scoyenia, Squamichnus,
Taenidium
13-Simple, actively filled (pelletoidal) Alcyonidiopsis, Castrichnus, Edaphichnium,
horizontal burrows Quebecichnus, Sphaerapus, Tubotomaculum,
Tubularina
14-Complex actively filled horizontal Bichordites, Bolonia, Nereites,
structures Paramargaritichnus, Parataenidium,
Psammichnites, Rutichnus, Scolecocoprus,
Scolicia, Tylichnus
15-Armored burrows Crininicaminus, Diopatrichnus, Ereipichnus,
Ichnospongiella, Lepidenteron, Nummipera
16-Horizontal branching burrow systems Agrichnium, Arachnostega, Korymbichnus,
Labyrintichnus, Multina, Paracanthorhaphe,
Pilichnus, Saportia, Shanwangichnus,
Taotieichnus, Vagorichnus, Virgaichnus
17-Horizontal burrows with horizontal to Arthrophycus, Caugichnus, Ctenopholeus,
vertical branches Intexalvichnus, Microspherichnus, Phycodes,
Phymatoderma, Polykampton, Saerichnites,
Streptichnus, Treptichnus, Tuberculichnus,
Vitichnus
18-Surface-coverage branching burrows Oldhamia
19-Radial to rosetted structures Arenituba, Asterichnites, Asterichnus,
Bifasciculus, Capodistria, Cladichnus,
Clematischnia, Cycloichnus, Dactyloidites,
Dactylophycus, Guanshanichnus,
Gyrophyllites, Haentzschelinia, Hartsellea,
Heliochone, Monocraterion,
Parahaentzschelinia, Phoebichnus, Radiichnus,
Rotamedusa, Scotolithus, Sphaerichnus,
Stelloglyphus, Taxichnites, Volkichnium
20-Sheaf burrow of thickly lined tubes Bornichnus, Schaubcylindrichnus
21-Horizontal burrows with serial chambers Halimedides, Omanichnus, Sidichnus,
Solanichnium, Strobilorhaphe
22-Horizontal burrows with simple vertically Halopoa, Teichichnus, Trichophycus
oriented spreiten
23-Horizontal spiral burrows Multilaqueichnus, Pramollichnus, Spiralites,
Spirodesmos, Spirophycus
24-Horizontal helicoidal burrows Augerinoichnus, Avetoichnus, Helicodromites
(continued)

25-Burrows with complex vertically oriented Caridolites, Daedalus, Dictyodora, Euflabella,
spreiten Gyrochorte, Heimdallia, Paradictyodora,
Stellavelum, Syringomorpha, Tursia
26-Burrows with horizontal spreiten Criophycus, Falcichnites, Fuersichnus,
Hydrancylus, Lamellaeichnus, Lobichnus,
Lophoctenium, Multilamella, Phycosiphon,
Rhizocorallium
27-Burrows with helicoidal spreiten Echinospira, Spirophyton, Zoophycos
28-Basal axial tubes with feather-like and Hillichnus
spreite-like structures
29-Mound-shaped structures Amanitichnus, Chomatichnus, Trusheimichnus
30-Isolated and serial oval to almond-shaped Calceoformites, Lockeia, Oravaichnium,
burrows Ptychoplasma
31-Pentameral-shaped imprints and burrows Asteriacites, Pentichnus
32-Oval-shaped imprints Astacimorphichnus, Corpusculichnus,
Epibaion, Ichnocumulus, Musculopodus
33-Dumbbell- and arrow-shaped burrows Arthraria, Bifungites, Monofungites
34-Vertical plug-shaped burrows Amphorichnus, Astropolichnus, Bergaueria,
Conichnus, Conostichus, Lithoplaision,
Mammillichnis, Metaichna, Solicyclus
35-Vertical unbranched burrows Altichnus, Bathichnus, Caletichnus,
Cylindricum, Digitichnus, Felderichnus,
Funalichnus, Guerraichnus, Laevicyclus,
Lingulichnus, Lunatubichnus, Oikobesalon,
Pustulichnus, Scalichnus, Siphonichnus,
Skolithos
36-Vertical single U- and Y-shaped burrows Arenicolites, Balanoglossites, Catenichnus,
Diplocraterion, Glyphichnus, Liholites,
Loloichnus, Paratisoa, Parmaichnus, Pholeus,
Polarichnus, Psilonichnus, Solemyatuba, Tisoa
37-Vertical multiple U- and Y-shaped burrows Kauriichnus, Lanicoidichna, Polykladichnus
38-Vertical simple J-shaped burrows Artichnus, Keilorites, Naviculichnium
39-Burrows with vertical tubes and stacked Tasselia
discs
40-Vertical helicoidal burrows Gyrolithes, Lapispira
41-Burrows with shaft or bunch with Chondrites, Fascifodina, Lennea, Pragichnus,
downwards radiating probes Saronichnus, Skolichnus, Trichichnus
42-Vertical concentrically filled burrows Cylindrichnus, Rosselia
43-Horizontal, branched concentrically filled Asterosoma, Patagonichnus
burrows
44-Spiral graphoglyptids Spirorhaphe
45-Guided meandering graphoglyptids Cosmorhaphe, Helminthorhaphe,
Spirocosmorhaphe
46-Uniramous meandering graphoglyptids Belocosmorhaphe, Belorhaphe,
Dendrotichnium, Helicolithus, Helicorhaphe,
Punctorhaphe, Ubinia, Urohelminthoida
(continued)

47-Radial graphoglyptids Arabesca, Chondrorhaphe, Dendrorhaphe,
Estrellichnus, Fascisichnium, Glockerichnus,
Lorenzinia, Persichnus, Tuapseichnium,
Yakutatia
48-Biramous meandering graphoglyptids Desmograpton, Oscillorhaphe, Paleomeandron
49-Regular to irregular network Acanthorhaphe, Megagrapton, Paleodictyon,
graphoglyptids Protopaleodictyon
50-Maze and boxwork burrows Ardelia, Ophiomorpha, Sinusichnus,
Spongeliomorpha, Thalassinoides
51-Vertical to oblique simple ornamented Capayanichnus, Liticuniculatus, Lunulichnus,
burrows Spirographites
52-Simple to complex burrows with terminal Camborygma, Egbellichnus, Katbergia,
chambers Macanopsis, Platicytes
53-Chambers surrounded by burrows Maiakarichnus, Dagnichnus
54-Isolated, clustered or interconnected cells Cellicalichnus, Celliforma, Corimbatichnus,
Elipsoideichnus, Palmiraichnus, Rosellichnus,
Uruguay
55-Chambers with discrete thick linings Chubutolithes, Coprinisphaera, Eatonichnus,
Feoichnus, Monesichnus, Quirogaichnus,
Rebuffoichnus, Teisseirei
56-Excavated chambers with thin linings Fictovichnus, Pallichnus, Scaphichnium
undetachable from rock matrix
57-Interconnected chambers and boxworks Attaichnus, Barberichnus, Coatonichnus,
Daimoniobarax, Fleaglellius, Krausichnus,
Masrichnus, Microfavichnus, Parowanichnus,
Socialites, Syntermesichnus, Tacuruichnus,
Termitichnus, Vondrichnus
58-Holes, pits and galleries in walls and Lazaichnus, Tombownichnus
fillings
59-Cylindrical vertical to oblique borings Carporichnus, Flagrichnus, Linkichnus,
Pecinolites, Stipitichnus, Trypanites,
Tubulohyalichnus
60-Borings with elliptical to sub-rectangular Osprioneides, Xylonichnus
cross sections
61-Winding borings Lapispecus, Maeandropolydora
62-U-shaped borings Canaliparva, Caulostrepsis, Diorygma,
Pseudopolydorites, Sertaterebrites
63- Circular holes and pit-shaped borings Circolites, Curvichnus, Dipatulichnus,
Lamniporichnus, Oichnus, Planavolites,
Polydorichnus, Stellatichnus, Tremichnus
64-Globular to spherical borings Granulohyalichnus, Planobola
65-Pouch borings Asthenopodichnium, Aurimorpha, Bascomella,
Cuenulites, Cubiculum, Petroxestes, Rogerella,
Umbichnus
66-Clavate-shaped borings Cavernula, Cylindricavus, Gastrochaenolites,
Palaeosabella, Phrixichnus, Ramosulcichnus,
Sanctum, Teredolites, Xylokrypta
(continued)

67-Fracture-shaped bioerosion traces Belichnus, Bicrescomanducator, Caedichnus,
Mandibulichnus
68-Branched tubular borings Calciroda, Clionoides, Cunctichnus,
Dekosichnus, Ichnoreticulina, Paleobuprestis,
Paleoipidus, Paleoscolytus, Runia, Scolecia,
Talpina
69-Non-camerate network borings Anobichnium, Filuroda, Orthogonum,
Rodocanalis
70-Camerate network borings Feldmannia, Haimeina, Ichnogutta, Iramena,
Pennatichnus, Pinaceocladichnus, Rhopalia,
Saccomorpha
71-Non-camerate boxwork borings Chaetophorites, Cycalichnus, Eurygonum
72-Camerate boxwork borings Entobia, Unellichnus
73-Spiral borings Helicotaphrichnus, Spirichnus
74-Radial borings Fascichnus, Polyactina
75-Dendritic and rosetted borings Abeliella, Calcideletrix, Clionolithes,
Dendrina, Dictyoporus, Megascolytinus,
Nododendrina, Pyrodendrina,
Scolytolarvariumichnus
76-Single circular to tear-shaped attachment Anellusichnus, Centrichnus, Kardopomorphos,
bioerosion traces Lacrimichnus, Ophthalmichnus
77-Multiple attachment bioerosion traces Finichnus, Flosculichnus, Podichnus
78-Elongate or branched attachment Camarichnus, Canalichnus, Renichnus,
bioerosion traces Stellichnus, Sulcichnus
79-Groove bioerosion traces Ericichnus, Gnathichnus, Osteocallis,
Radulichnus
Analysis of ichnodiversity changes from the Ediacaran to the Holocene supports

the three-phase kinetic model of Sepkoski (1984) based on analysis of marine body
fossils. According to this model, the three main evolutionary radiations in the marine
biosphere display an early exponential growth followed by subsequent slowing of
growth. In the case of the Cambrian and Ordovician radiations there was very rapid
growth of diversity in their initial stages until a plateau was reached later, whereas
diversity rose slowly during the onset of the Mesozoic Marine Revolution and con-
tinues to do so. Controversies regarding this model have essentially revolved around
potential geologic and paleontologic biases, such as increased exposure of sedimen-
tary rocks, uneven number of studies per interval, and increased ease of fossil
extraction (Raup 1976; Peters and Foote 2001; Alroy et al. 2008; Alroy 2010). For
our interest here, the most serious controversy is the one on the diversity levels
attained in the post-Paleozoic world. It has been argued that standardized curves
show a final diversity only slightly higher than the Paleozoic maximum (e.g., Alroy
et al. 2008; Alroy 2010, 2014) in sharp contrast with the much higher diversity levels
shown by Sepkoski’s curves (see also Bambach et al. 2004). However, new stan-
dardized curves show that Cenozoic diversity doubles Paleozoic values, supporting
sustained marine diversification during the Mesozoic–Cenozoic (Bush and Bambach
2015), a pattern more consistent with the original Sepkoski’s curves.
In particular, one of the potential biases affecting estimation of post-Paleozoic

diversity levels is the so-called Pull of the Recent (Raup 1979). This effect operates
in two ways. First, there is an increase in the volume of Cenozoic (and to a lesser
extent Mesozoic) outcrops. Second, because the modern is much better sampled
that the rest of the fossil record, taxa that occur in just one stratigraphic interval and
have a modern representative will be plotted as occurring through the whole inter-
val, resulting in an inflation of diversity levels in post-Paleozoic strata. Interestingly,
the trace-fossil record is not biased in the same way. Certainly, neoichnologic stud-
ies are instrumental in recognizing some biogenic structures in the modern, helping
to extend the stratigraphic range of some ichnotaxa. However, at the same time,
modern counterparts of some trace fossils are lacking not because these behaviors
have gone extinct, but because identification of modern traces is typically much
more difficult than with fossil material, particularly in the case of bioturbation struc-
tures (Bromley 1996; Buatois and Mángano 2011a). Trace fossils tend to be
enhanced by diagenetic processes that assist in their recognition (Magwood 1992)
and many biogenic structures are cumulative structures, which consist of both abandoned
and active components (Bromley and Frey 1974); casts of modern representatives of
cumulative structures would only reflect the morphology of the open components
occupied by the producer, resulting in a simpler morphology than the actual overall
architecture, making identification of the ichnotaxon virtually impossible (Frey
1975; Frey and Seilacher 1980; Magwood 1992). The practical result of this would
be an artificial drop in ichnodiversity in the modern. This more complex pattern
exhibited by the trace-fossil record is referred herein as the Push and Pull of the
Recent. To avoid this problem, we have plotted in the final curves together the
Holocene and the modern.
Similar to Sepkoski’s curves, ichnologic data show a phase of exponential growth
in ichnodiversity followed by an equilibrium stage during the Cambrian Explosion
and the Great Ordovician Biodiversification Event. Identical to body fossils, the
Mesozoic Marine Revolution is associated with a slower increase in ichnodiversity.
The overall coincidence between Sepkoski’s curves and those presented in this
chapter indicates similar diversity trajectories for animal diversity and their behav-
iors. However, in the case of post-Paleozoic faunas, whereas the curve for marine
animals does not show any evidence of a plateau, ichnologic information seems to
suggest that a plateau may have been reached for both marine bioturbation and
bioerosion.
In contrast, trends in ichnodisparity reveal a more complicated picture (Fig. 16.2).
Whereas the Cambrian Explosion is associated with an increase in both ichnodis-
parity and ichnodiversity (Mángano and Buatois 2014; see Chap. 3), the Great
Ordovician Biodiversification Event and the Mesozoic Marine Revolution display
different trends. A 263 % increase in ichnodisparity took place during the
Terraneuvian as a result of the Cambrian Explosion (8 and 29 categories of archi-
tectural designs in the Ediacaran and Terraneuvian, respectively), but only a 45 % is
associated with the Great Ordovician Biodiversification Event (38 categories of
architectural design in the Furongian and 55 by the Late Ordovician). In addition,
if bioturbation and bioerosion structures are considered separately, it is clear that
bioerosion is responsible for the overall increase (17 % increase in ichnodisparity

for bioturbation structures and 367 % for bioerosion structures). Whereas the
Cambrian Explosion was the main event in the Phanerozoic for large-scale innova-
tions in animal-sediment interactions, it is the Great Ordovician Biodiversification
Event that played this role regarding bioerosion (Buatois et al. 2016a). If bioturba-
tion structures are considered separately for shallow- and deep-marine environ-
ments, it is apparent that ichnodisparity in shallow-marine settings remained very
close to Cambrian levels, whereas an increase took place in the deep sea (88 %; 17
categories of architectural design in the Furongian and 32 by the Late Ordovician).
In the same fashion that broad innovations in bioerosion lagged behind those in
bioturbation, the establishment of the main types of animal-sediment interactions in
the deep sea took place later than in shallow-marine settings, as originally envis-
aged in models of deep-sea colonization (e.g., Uchman 2004). Because approxi-
mately half of the architectural categories that typify Ordovician deep-sea
ichnofaunas first occurred in shallow-marine settings during the Cambrian, the
overall impact of Ordovician deep-sea categories in global marine ichnodisparity
levels is limited (Buatois et al. 2016a).
In turn, although an increase in ichnodisparity is apparent as a result of the
Mesozoic Marine Revolution, distinguishing between bioturbation and bioerosion
structures shows that the increase is restricted to the latter (19 % increase between
the Late Triassic and the Late Cretaceous). Ichnodisparity of bioturbation structures
in both shallow- and deep-marine settings does not display any significant increase
in connection to the Mesozoic Marine Revolution.
A main conclusion of our analysis of secular changes in ichnodiversity and ich-
nodisparity during the Cambrian Explosion, the Great Ordovician Biodiversification
Event and the Mesozoic Marine Revolution is that, although increases in ichnodi-
versity are invariably linked to evolutionary radiations, these are necessary but not
sufficient conditions for ichnodisparity increases. In order to further explore the
links between ichnodiversity and ichnodisparity and the possible underlying causes
for increases in the latter, it is now useful to analyze the results of our compilations
regarding continental environments.
Ichnodiversity compilations for continental environments show a more protracted
process of colonization through the whole Phanerozoic as revealed by bioturbation
structures (Fig. 16.1). Although the process of invasion of the continents seems to
have been slow and gradual, an initial rapid increase took place by the Silurian-
Devonian transition (see Chap. 6), with an impressive 967 % increase in ichnodiver-
sity between the Wenlock and the Early Devonian (3 and 32 ichnogenera in the
Wenlock and Early Devonian, respectively). As in the case of the Cambrian Explosion,
this rapid diversification in continental settings is consistent with Sepkoski’s (1978)
exponential model for early diversification. Subsequent to that increase a plateau until
the Early Mississippian is apparent. However, this may be in part simply an artifact
due to the scarcity of continental outcrops of Late Devonian age (see Chap. 6).
Subsequently, a slow but constant increase in ichnodiversity took place until the early
Permian (Cisuralian), which displays an unusual peak (65 %, 91 ichnogenera). This
peak has been noted in earlier studies and deemed controversial (Buatois et al. 1998).
It reflects the presence of a wide variety of arthropod trackways in overbank deposits,

which at least in part results from a marked tendency towards oversplitting in track-
way taxonomy. In particular, many arthropod are only known from the Permian of
Germany (e.g., Holub and Kozur 1981; Walter 1983); many of these are considered
doubtful (see Chap. 6). After a plateau that lasted the whole Triassic, another more
modest peak is apparent in the Early Jurassic. This is, at least in part, a monographic
effect (Raup 1976) resulting from the high number of new arthropod trackway genera
introduced by Hitchcock (1858, 1865) based on his analysis of continental trace fos-
sils from the Connecticut Valley. Many of these have never been recorded elsewhere
and are now regarded as doubtful (Rainforth 2005), so they have been omitted in our
compilation. However, even leaving aside these doubtful ichnotaxa and only retain-
ing those regarded as valid, the Early Jurassic ichnodiversity peak still persists
(21 %; 61 and 74 ichnogenera in the Late Triassic and Early Jurassic, respectively).
Since the Middle Jurassic onwards, ichnodiversity experienced a relatively constant
increase.
Bioerosion in continental environments experienced a much later develop-
ment than bioturbation. Although wood borings were mentioned for the early
Permian (von Kušta 1880), the earliest convincing evidence of bioerosion in conti-
nental environments is from the Late Triassic, with the recording of six ichnotaxa
representing wood bioerosion (Tapanila and Roberts 2012). Notably, this record
predates the earliest record of marine bioerosion in wood, which is Early Jurassic
(Villegas-Martín et al. 2012). In turn, the earliest record of bioerosion in bone is
from the Early Jurassic (undetermined ichnotaxon), with the ichnogenus Cubiculum
being recorded for the first time in the Late Jurassic (Xing et al. 2015). This was
followed by a plateau until the Late Cretaceous with the addition of ten new ichno-
genera. Since then, ichnodiversity of bioerosion structures reached a plateau that
continues until the Holocene.
After the initial rapid increase in ichnodiversity of bioturbation structures by the
Silurian-Devonian transition, colonization of freshwater settings was apparently
quite gradual. It has been suggested that this slow pace may have resulted from the
need to develop innovative styles of reproduction and dispersal, as well as complex
osmoregulatory systems (Miller and Labandeira 2002). The fact that no plateau
indicating an equilibrium stage in diversification is apparent for bioturbation struc-
tures may suggest that the invasion of terrestrial environments is a still ongoing
process. This is consistent with the fact that the utilization of freshwater infaunal
ecospace seems to have been less complete than in their marine counterparts (Miller
and Labandeira 2002). Continental bioerosion ichnogenera seem to have reached a
plateau by the end of the Mesozoic, but their ichnotaxonomy is still in its infancy
and it is not unexpected that this trend will change with further studies.
As with ichnodiversity, ichnodisparity in continental environments reflects the
process of protracted colonization (Fig. 16.2). Ichnodisparity curves of bioturbation
structures reveal a rapid increase with the onset of widespread colonization by the
Silurian-Devonian transition (see Chap. 6), with a 267 % increase in ichnodisparity
between the Wenlock and the Early Devonian (3 and 11 categories of architectural
designs in the Wenlock and Early Devonian, respectively). This was followed by a
very slow increase until the Late Cretaceous that records a hike (29 %; 28 categories
of architectural design in the Early Cretaceous and 36 in the Late Cretaceous), albeit
limited, reaching ichnodisparity levels that persist until the Recent. The Late
Cretaceous ichnodisparity peak is also expressed by bioerosion structures.
However, it should be noted that all categories of architectural designs in freshwa-
ter subaqueous environments and transitional terrestrial to subaqueous settings
originated in marine environments and only subsequently occurred in continental
settings. In contrast, there are six categories that are exclusive of terrestrial settings
(see Chap. 11). In other words, the main innovations required to invade freshwater
were already present in the marine realm, but the only true behavioral innovations
in continental environments were those required to colonize subaerially exposed
settings. These are mostly the architectural designs developed by insects nesting in
paleosols (see Chap. 13).
To summarize, our analysis shows that the key factor in building up ichnodisparity
is not the existence of an evolutionary radiation per se, but the colonization of empty
ecospace. The Cambrian Explosion records the colonization of empty ecospace and,
therefore, displayed an increase in both ichnodisparity and ichnodiversity. A similar
situation is expressed by bioerosion during the Great Ordovician Biodiversification
Event in connection with the colonization of hard substrates. A third increase in
ichnodisparity is revealed by the colonization of paleosols which experienced
dramatic evolutionary innovations by the end of the Mesozoic. Finally, the cradle of
evolutionary innovations is in shallow-marine settings (see Sect. 16.5). This is
revealed by the appearance in shallow seas of all the architectural designs that sub-
sequently expanded into freshwater and a substantial number of those that later
bloomed in the deep sea. In almost all these cases, the ichnodisparity increase is
followed by a subsequent further increase in ichnodiversity, as it is illustrated, by
the diversification of insect nesting structures during the Cenozoic, following the
establishment of the basic architectural designs in the Late Cretaceous. This pattern
supports analysis based on body fossils that indicates a “first disparity, then diver-
sity” scenario (Foote 1993, 1997; Erwin 2007; see Chap. 6).
Finally, the limitations of ichnodiversity analysis to unravel evolutionary trends
need to be addressed (see Buatois and Mángano 2013 for a detailed analysis). As
stated many times, the fact that ichnodiversity cannot be equated with the actual
diversity of organisms represents a cautionary note (Ekdale 1985). Behavioral con-
vergence resulting in different organisms being able to produce the same ichno-
taxon may mask evolutionary trends. The opposite case, the one of several ichnotaxa
resulting from the work of a single producer, also undoubtedly introduced a bias in
ichnodiversity analysis through geologic time. Because of this, the different chap-
ters exploring the ichnologic blueprints of evolutionary radiations use a plurality of
conceptual and methodological tools rather than being restricted to an analysis of
the number of ichnogenera or architectural designs. For example, it is clear that
burrows produced by decapod crustaceans represent one of the most, if not the
most, archetypal biogenic structures of the Mesozoic Marine Revolution. However,
because some of its most characteristic structures, namely burrow systems included
in Thalassinoides, are produced not only by decapod crustaceans, but by other

organisms with a stratigraphic range that extends well before the Mesozoic, the
importance of crustacean burrows remains undetected if only ichnodiversity is con-
sidered. In other words, the long stratigraphic range of Thalassinoides is misleading
with respect to its key role during the Mesozoic Marine Revolution. A clearer pic-
ture emerges when, for example, abundance is considered as well (Carmona et al.
2004; see Chap. 9).
Also, in part as a result of behavioral convergence, ichnogeneric extinction rates
are remarkably low. It has been noted that increases in diversity within clades may be
produced from a drop in extinction rate rather that diversification per se, resulting in
the so-called “pseudoradiations” (Simões et al. 2016). In the case of trace-fossil data,
extinction rates can be regarded not only as low but also as constant, the latter at least
since the Ordovician. Because of the impossibility of equating biological diversity
and ichnodiversity, the trace-fossil expression of evolutionary radiations is not pri-
marily of a phylogenetic nature, but rather reflects behavioral innovations.
Other problems are evident while dealing with ichnodiversity changes in specific
environments through the Phanerozoic. Because there is a trend to increased infaunaliza-
tion through time, the colonization of deeper tiers is conducive to obliteration of the more
shallowly emplaced structures, which in many cases represent a substantial contribution to
ichnodiversity. As a result, a decrease in ichnodiversity through time rather than an
increase, has been noted in the case of tidal flats (Mángano et al. 2002; Mángano and
Buatois 2015) and lakes (see Chap. 11). These cases underscore the importance of care-
ful evaluation of taphonomic overprints.
Also, using ichnodiversity at ichnogeneric level may preclude detection of other
patterns. Ichnospecies reflect minor behavioral variations and, therefore, they may
help to detect more subtle trends in behavioral innovations (see Chap. 3). A large
number of ichnogenera that have their first occurrence early in the Paleozoic seem
to have displayed an increase in the number of ichnospecies through the Phanerozoic
(see Chaps. 3 and 9). Exploring patterns of ichnospecies distribution through time
is a promising line of research, although this would imply in-depth critical reevalu-
ation of trace-fossil taxonomy at this rank, something beyond the scope of the
present study.
Regardless of potential shortcomings, our review indicates that ichnologic
information (including both ichnodiversity and ichnodisparity) is highly useful as
an independent line of evidence to understand paleobiologic trends through geologic
time.
16.3 Benthic Fauna Response to Mass Extinctions
Ichnologic information typically reveals a set of common responses to the three

mass extinctions analyzed in this book, the end-Permian (see Chap. 7), the end-
Triassic (see Chap. 8), and the end-Cretaceous (see Chap. 12). Most of these extinc-
tion events are characterized by a reduction in ichnodiversity, decrease in degree of
bioturbation, reduction of bioturbation depth, size reduction, dominance of simple

trace fossils in post-extinction strata, preferential survival of ichnotaxa produced by
deposit feeders, and selective extinction in shallow-marine ecosystems. How pro-
nounced were these effects seems to depend on the severity of the mass extinction
event, with the end-Permian mass extinction clearly showing the most severe impact
from an ichnologic standpoint, showing consistency with body-fossil data. In turn,
this is reflected in the pattern of recovery after the extinction, with longer recovery
times involved in the end-Permian mass extinction in comparison with the end-
Cretaceous event. Ichnologic information on the other two major events, the end-
Ordovician and late Devonian extinctions is patchy, with only a few studies available
(McCann 1990; Herringshaw and Davies 2008; Buatois et al. 2013) and, therefore,
any generalization remains premature.
Although these extinctions are associated to a decrease in ichnodiversity, actual
extinction of invertebrate ichnotaxa is quite rare. On the contrary, ichnotaxa that
disappeared after the mass extinctions for the most part reappeared during the
recovery time, representing Lazarus ichnotaxa (Gibert 2003). As noted previously,
trace-fossil extinction rates tend to be very low, a fact that, at least in the case of
relatively simple behaviors, results from behavioral convergence. Some ichnotaxa
(e.g., the undermat miner Oldhamia, the large trail Climactichnites), however, seem
to reflect behaviors that have apparently disappeared by the end of the Cambrian,
together with their producers.
Because the curves illustrated in Figs. 16.1 and 16.2 have been compiled as range
through data, they are of limited use to detect changes in ichnodiversity as a result
of mass extinctions. For example, whereas a significant drop in ichnodiversity is
associated with the end-Permian mass extinction (see Chap. 7), this is not reflected
by the ichnodiversity curves. Similarly, the end-Triassic mass extinction resulted in
a decrease in ichnodiversity, albeit more modest (see Chap. 8), which is not recorded
in our ichnodiversity curves. The absence of an ichnodiversity drop in these curves
is simply an artifact resulting from the methodology employed in their construction.
It is therefore surprising that a 7 % decrease in total marine ichnodiversity (5 % for
bioturbation structures and 15 % for bioerosion structures) is revealed by these
curves in connection with the end-Cretaceous mass extinction. The Late Cretaceous
peak and subsequent drop in ichnodiversity are also present in the curves that were
plotted eliminating the occurrence of singletons.
Decrease in the degree of bioturbation has been noted in connection with many
mass extinction events. Although comparative analysis of mass extinctions from a
trace-fossil perspective are still in their infancy, available information seems to
suggest that the extent of this decrease in intensity of bioturbation tends to reflect
how profound was the impact of the event. This is illustrated by the end-Permian
mass extinction, which shows a collapse of the mixed layer (see Chap. 7 and
Sect. 16.4; Buatois and Mángano 2011b; Hofmann et al. 2015). A similar situation
has not been recorded so far associated with any of the other mass extinction
events. In fact, intense bioturbation has been noted in Danian post-extinction
deposits (see Chap. 12).
Reduction in the size of trace fossils has been detected in connection with many
mass extinction events, most notably the end-Permian (see Chap. 7) and end-
Triassic (see Chap. 8). In other cases, such as the end-Cretaceous, this trend is not
apparent (see Chap. 12, although see Wiest et al. 2015 for documentation of trace-
fossil size reduction in the aftermath of the end-Cretaceous mass extinction). The
so-called Lilliput effect has been noted in the paleobiologic literature to explain the
sharp decrease in body size, based on both the body fossil and trace-fossil records,
in the aftermath of mass extinctions (Urbanek 1993; Twitchett 2007). Explanations
for this size reduction are still unclear, with both externalist and internalist causes
being involved. Limited oxygen and food have been invoked as likely candidates for
the former (Twitchett 2007), whereas survival of small taxa; the dwarfing of larger
lineages and the evolutionary miniaturization from larger ancestral stocks fall
among the later (Harries and Knorr 2009).
Dominance of simple trace fossils is another typical feature in post-extinction
strata. For example, simple trace fossils attributed to Planolites are the first in reap-
pearing after the end-Permian mass extinction (see Chap. 7). Simple forms, such as
Planolites and Palaeophycus, seem to be common in Lower Jurassic post-extinction
deposits as well (see Chap. 8). The prevalence of these simple morphologies overall
reflect very simple feeding behaviors, most likely reflecting opportunistic population
strategies in the aftermath of mass extinctions (Fraiser and Bottjer 2009).
It has long been noted that organisms are differentially affected during extinctions
according to trophic type (Jablonski and Raup 1995; Smith and Jeffrey 1998; Hansen
et al. 2004; Twitchett 2006). In particular, suspension feeders tend to have been more
affected than deposit feeders during mass extinctions. In this regard, it has been con-
cluded that both selective deposit feeding and omnivory significantly enhance sur-
vivability during mass extinction events (Twitchett 2006). This is clearly illustrated
by the overwhelming dominance of deposit-feeding traces in Danian post-extinction
deposits (see Chap. 12). Also, post-extinction late Devonian ichnofaunas are domi-
nated by structures produced by selective deposit feeders, whereas suspension-feed-
ing burrows are absent (Buatois et al. 2013).
Ichnofaunas from shallow-marine environments tend to be more strongly affected
than those in marginal-marine and deep-marine settings. Marginal-marine, brackish-
water faunas typically consist of opportunistic animals that flourish under extreme
conditions. Therefore, they are able to rapidly colonize environments after a major
disturbance, such as mass extinctions. Of the five colonization phases recognized
for brackish-water settings by Buatois et al. (2005), only the end of the first
(Ediacaran–Ordovician) and the third (Permian–Triassic) coincide with mass
extinctions (the Late Ordovician and Late Triassic mass extinctions, respectively).
The remaining three of the “Big Five”, the late Devonian, end-Permian, and end-
Cretaceous mass extinctions, does not show any impact in marginal-marine biotas.
In the same vein, ichnofaunas from deep-water settings do not seem to have been
strongly affected by mass extinctions (Uchman 2004). No major crisis has been
associated with any of the “Big Five” in the deep sea, other than a reduction in the
diversity and abundance of graphoglyptids in the case of the end-Ordovician and
end-Cretaceous mass extinctions (Uchman 2003).
16.4 Patterns of Ecospace Colonization
Having established that there are some recurrent ichnologic patterns involving evo-
lutionary radiations and mass extinctions, we may now explore if recurrent motifs
can be established in connection with colonization of empty or underutilized eco-
space. In other words, is it possible to detect trace-fossil assemblages that recur
through geologic time not as a response to environmental constrains (as is the case
of ichnofacies), but driven by large-scale evolutionary controls instead? We have
attempted to address this question in a previous paper, referring to this particular
recurrence of ichnoassemblages revealing common strategies in ecospace coloniza-
tion as “the Déjà vu effect” (Buatois and Mángano 2011b). In this chapter, we argue
that three main types of trace-fossil assemblages can be recognized in connection
with exploitation of empty or underutilized ecospace.
The first of these assemblages consists of very small grazing trails (e.g., Helminthopsis,
Helminthoidichnites, Gordia) produced by epifaunal or very shallow infaunal vermiform
animals, very shallow-tier feeding structures (e.g., Treptichnus, Oldhamia, Pilichnus) of
infaunal vermiform organisms, and (in some cases) trackways (e.g., Diplichnites,
Stiaria) of epifaunal arthropods (Buatois and Mángano 2011b). Almost invariably trace
fossils occur in association with structures suggestive of microbial stabilization, such as
wrinkle marks, palimpsest ripples, and old elephant skin (Noffke 2010; Buatois and
Mángano 2012a). The associated primary sedimentary fabric and bedding-plane physi-
cal sedimentary structures are superbly preserved (e.g., Buatois et al. 1997). Ichnofaunas
reveal superficial and shallow-tier grazing and mining of microbial mats in the absence
of substantial infaunal bioturbation. The presence of microbial mats allows preserva-
tion of minute trace fossils, displaying delicate fine morphologic features (Buatois and
Mángano 2011b). These trace-fossil assemblages occur across a wide range of envi-
ronmental settings, such as Ediacaran-lower Cambrian shallow-marine (e.g., Gehling
1999; Buatois et al. 2014), Cambrian deep-marine (e.g., Buatois and Mángano 2003),
and late Paleozoic continental to fluvio-estuarine (Buatois et al. 1997; Mángano et al.
1997) settings, invariably in connection with empty or underutilized ecospace.
The second assemblage is characterized by shallow-tier burrows
(Spongeliomorpha, Halopoa), trails (Cruziana), and resting traces (Rusophycus)
with well-developed ornamentation, typically bioglyphs. Very shallow-tier burrows
and trackways may be present as well. The occurrence of well-preserved bioglyphs
and the sharp burrow boundaries suggest emplacement in sediment that was firm at
or close to the sea bottom. These trace-fossil assemblages are associated with wide-
spread firm substrates and the virtual absence of a mixed layer (Droser et al. 2004;
Jensen et al. 2005; Buatois and Mángano 2011b; Mángano et al. 2013; Hofmann
et al. 2015). This is illustrated by two apparently unrelated ichnofaunas: those in
Cambrian–Ordovician and Lower Triassic shallow-marine environments. Both
deposits are characterized by preferential preservation of epifaunal and very
shallow-tier infaunal trace fossils; paucity of mid- and deep-tier trace fossils;
absence of mottled bioturbation textures; and dominance of cohesive substrates in
open-marine environments (Hofmann et al. 2015). In the case of lower Paleozoic strata,
the mixed layer was still incipient and poorly developed (Droser et al. 2004; Jensen
et al. 2005; Callow and Brasier 2009; Mángano et al. 2013), whereas in the Lower
Triassic the mixed layer may have collapsed as a result of the end-Permian extinction
(Buatois and Mángano 2011b; Hofmann et al. 2015; see Chap. 9). The ichnologic
responses in both situations were essentially identical.
The third assemblage is characterized by the presence of horizontal branched
burrow systems (e.g., Multina, Vagorichnus) at the base of event sandstone beds
(Buatois and Mángano 2011b). These burrow systems are preserved along litho-
logic interfaces creating virtually no disturbance in the primary sedimentary fabric.
Multina is common at the base of lower Paleozoic marine tempestites and turbidites,
where it records post-event colonization (e.g., Buatois and Mángano 2004, 2012b;
Buatois et al. 2009). Vagorichnus has been recorded at the base of Jurassic lacus-
trine turbidites (Buatois et al. 1995, 1996). Although these occurrences are from
disparate environments and from rocks of different ages, they are all associated with
an increase in burrowing depth reflecting initial exploitation of the infaunal ecospace
(Buatois and Mángano 2011b). From an evolutionary standpoint, this incipient
colonization of the infaunal ecospace preceded the onset of more intense bioturbation
and the establishment of a well-developed mixed layer.
The recurrent presence of these three types of trace-fossil assemblages in rocks of
different ages and formed under a wide variety of environmental settings can only be
understood under the light of the evolutionary prism. These assemblages reveal a
limited repertoire of behavioral strategies that allow benthic faunas to colonize empty
or underutilized ecospace. Fossilization of these behavioral strategies in the form of
biogenic structures is mediated by a set of preservational conditions resulting from
the recurrent opening of a taphonomic window. This window was responsible for the
preservation of surface and shallow-tier structures which typically are obliterated by
deeper-tier and more intense bioturbation (Bromley 1996).
16.5 Environmental Shifts Through Time
A number of studies published for the most part in the eighties and nineties revealed
that the fossil record can be characterized by a recurrent pattern of onshore origina-
tion of evolutionary innovations and their subsequent expansion to deeper water,
resulting in the proposal of the so-called “onshore–offshore model” (e.g., Jablonski
et al. 1983; Sepkoski and Miller 1985; Jablonski and Bottjer 1990; Sepkoski 1991;
Sepkoski and Sheehan 1983). In fact, it has been argued that an onshore–offshore
pattern is displayed by Sepkoski’s evolutionary faunas themselves, as illustrated by
the replacement of trilobite-rich communities by mollusk-rich communities in shal-
low-water niches throughout the Paleozoic (Sepkoski and Miller 1985).
The trace-fossil record provides further empirical support to this model across a
broad range of scales. First, an onshore–offshore pattern is evident from individual
ichnotaxa, showing either evidence of expansion or retreat. In the first case, an
ichnogenus occurring for the first time in shallow water subsequently extends its
environmental range into deeper-marine settings without loss of onshore represen-
tatives. In the second case, migration into deeper water occurs together with loss of
onshore representatives (Bottjer et al. 1988; Stanley and Pickerill 1993). Examples
of expansion include Ophiomorpha (Bottjer et al. 1988; Tchoumatchenco and

Uchman 2001), Scolicia (Seilacher 1986) and Rhizocorallium commune (Knaust
2013). An example of retreat is Zoophycos (Bottjer et al. 1988).
Second, entire groups of trace fossils may display the onshore–offshore pattern.
The best example is graphoglyptids, which in the early Cambrian occur in shallow
water, but subsequently migrated to the deep sea (Crimes and Anderson 1985; Jensen
and Mens 1999), where they became the core component of the Nereites Ichnofacies.
Third, the onshore–offshore pattern is also displayed by ichnofabrics. For example,
it has been argued that in carbonate settings intense bioturbation first took place in
shallow-water settings and only later seaward (Droser and Bottjer 1989). This pat-
tern may reflect the fact that niche partitioning within the substrate first developed
in shallow water and subsequently expanded into deeper water.
Fourth, and at a larger scale, the onshore–offshore pattern is displayed in connection
with the colonization of empty or underutilized ecospace. This is shown by trace-fossil
evidence of original colonization of shallow-marine settings and the progressive
subsequent expansion into the deep sea, marginal-marine environments and conti-
nental settings. Also, this pattern is apparent within continental environments them-
selves, as demonstrated by the initial colonization of lake-margin settings and later
expansion into subaqueous lacustrine settings (Buatois and Mangano 1993; Buatois
et al. 1998; see Chaps. 5 and 6). This is certainly analogous to the migration of plants
from upland areas to the lowlands during the Carboniferous-Permian transition
(DiMichele and Aronson 1992) and the origination of aquatic insects in running
water and subsequent expansion into lacustrine habitats (Wooton 1988).
16.6 Conclusions
Comparative analysis of the ichnologic record of major evolutionary events allows

detecting recurrent patterns that may help to further increase our understanding of
the underlying evolutionary dynamics of animal-substrate interactions through geo-
logic time. In particular, we have detected recurrent patterns of organism-substrate
interactions during evolutionary radiations, benthic fauna response to mass extinc-
tions, strategies of ecospace colonization, and environmental shifts through time.
Evolutionary radiations are invariably associated with an ichnodiversity increase,
whereas increases in ichnodisparity are linked to the colonization of empty eco-
space. Evaluation of mass extinction events suggests common responses from the
perspective of animal-substrate interactions, namely a reduction in ichnodiversity,
decrease in degree of bioturbation, reduction of bioturbation depth, size reduction,
dominance of simple trace fossils in post-extinction strata, preferential survival of
ichnotaxa produced by deposit feeders, and selective extinction in shallow-marine
ecosystems. The recurrent presence of certain trace-fossil assemblages in rocks of
different ages and formed under a wide variety of environmental settings seems to
suggest a limited repertoire of behavioral strategies for colonization of empty or
underutilized ecospace. The trace-fossil record provides further empirical support
to the onshore–offshore model across a broad range of scales.
Acknowledgments We thank José Carlos Garcia Ramos, Jorge Genise, Verónica Krapovickas,
Conrad Labandeira, Marisa Lopez-Cabrera, Radek Mikuláš, Nic Minter, Eduardo Olivero, Laura
Piñuela, Cecilia Pirrone, Eric Roberts, Alfred Uchman, Mark Wilson, and Max Wisshak for gener-
ously providing highly useful advice and information at different stages of the compilation.
Eduardo Olivero and Roy Plotnick reviewed the chapter providing highly useful feedback. Kai
Zhou assisted us with plotting the information in tables and charts.
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Index
A Arenicolites isp., 21, 46, 64, 70, 85, 88

Abarenicola pacifica, 105 Aristopentapodiscus, 373
Acanthichnus, 454 Artichnus, 21, 43, 93
Acropentapodiscus, 373 Artichnus pholeoides, 94
Acunaichnus dorregoensis, 390 Asaphoidichnus, 454
Ademosynidae, 195 Asencio Formation, 344
Adephagan beetles, 201 Asiatoceratodus, 238
Agua de la Peña Group, 192 Asteriacites, 9, 21, 43, 44, 66
Alcyonidiopsis, 280, 283 Asteriacites lumbricalis, 42, 67
Allocotichnus, 454 Asterosoma, 21, 46, 47, 52, 63, 65, 68, 69, 76,
Ameghinichnus, 372, 373 78, 85, 89, 95
Ameghinichnus patagonicus, 372 Asterosoma coxii, 52
Amphibiopodiscus, 373 Asterosoma ludwigae, 52
Anachalcos mfwangani, 346 Asterosoma radiciformis, 52
Anchisauripus, 6, 8 Asterosoma striata, 52
Ancorichnus, 45 Asterosoma surlyki, 52
Angulata Zone, 7, 11 Atreipus, 6, 8, 147
Angulichnus, 454 Australopithecus, 413
Anomoepus, 6–8 Australopiths, 411
Anthropoidipes ameriborealis, 437 Avolatichnium, 192
Anyao Formation, 198 Azolla ferns, 202
Apatopus, 5, 6, 146
Arachnomorphichnus, 454
Arachnostega, 21, 45, 46, 49, 61 B
Aragonitermes teruelensis, 335 Baissatermes lapideus, 335
Arariperhinus monnei, 335 Baissoferidae, 211, 216
Archaeonassa, 34, 62, 223, 454 Balanoglossites, 46
Archaeonassa fossulata, 192 Barberichnus bonaerensis, 304
Archeoentomichnus metapolypholeos, 320 Barrancapus, 5
Archosaur trackways, 6 Bathichnus paramoudrae, 80
Archosaurs, 137, 138, 140, 143, 145–147, 160 Batrachopus, 6, 7
Arcichnus, 21, 43, 44, 454 Bauplan, 332
Arenicolites, 10–12, 46, 189, 194, 244 B. coronus, 209

DOI 10.1007/978-94-017-9597-5
476 Index
Beaconites, 316, 324 Camborygma, 181, 189, 194, 215, 325, 331,
Beaconites antarcticus, 209 341, 347
Beaconites filiformis, 308 Camborygma araioklados, 320
Behavioral convergence, 241–244 Camborygma eumekenomos, 315, 320
Belostomatidae, 195 Camborygma Ichnofacies, 330, 331, 333, 340
Bestiopeda bestia, 375 Camborygma litonomos, 315
Bestiopeda sanguinolenta, 375 Camborygma symplokonomos, 315, 320
Bichordites, 41 Cambrian Evolutionary Fauna (CEF), 20
Bichordites kuzunensis, 42 Canipeda longigriffa, 375
Biformites, 454 Cantabritermes simplex, 335
Bifurcatichnus, 454 Cardioichnus, 41
Bifurculapes, 192, 454 Caririchnium, 206
Biostratigraphic resolution, 288–289 Carnivorous archosaur trackways (Apatopus), 6
Biotal revolutions, 231–233 Carpatichnis, 454
Bipedalism Castrichnus, 242, 344
3D imaging, 414 Castrichnus incolumis, 303
foot architecture, 414 Catastrophic vs. gradual extinction, 289–290
footprints, 412 Caulostrepsis, 53, 57, 60, 61, 92
group-of-three scenario, 414 Cave bear cults, 426
Laetoli trackways, 413 Cave paintings, 426–427
non-hominin Laetoli track, 415 Cellicalichnus, 347
paleontologic context, 413 Cellicalichnus antiquus, 332
sedimentologic significance, 414 Cellicalichnus chubutensis, 316, 338
trackmaker height, 412 Cellicalichnus dakotensis, 338
volcaniclastic sediments, 415 Cellicalichnus ficoides, 310
Bivalve-produced ichnogenera, 35–40, Cellicalichnus meniscatus, 316, 325, 332
55–56 Celliforma, 242, 309, 324, 339, 344
Bivalves Celliforma germanica, 338, 346
epifaunal and semi-infaunal taxa, 35 Celliforma ichnofacies, 340
evolutionary diversification, 35 Celliforma nuda, 309
seagrasses and mangroves, 40 Celliforma rosellii, 346
tellinaceans, 40 Celliforma spirifer, 338, 346
Blood feeding, dinosaur hosts, 278–279 Cenomanian–Turonian transition in Europe, 79
Blue Lias Formation, 11 Cenozoic diversity, 458
Bornichnus, 46, 52 Cenozoic main radiation, 377
Brachiosauridae, 148 Cenozoic mammalian footprints, 373–376,
Brachychirotherium, 6, 138, 146, 147, 188 379–398
Brakeland formations, 328 Cenozoic radiation, mammals
Branchiopods, 202 Antofagasta, 389
Brasilichnium, 6, 188 biogeographic differentiation, 379
Brasilichnium elusivum, 373 Claromecó, 389
Brontopodus, 147 Faunistic Content, 376–379
Bryopompilus interfector, 337 footprints, 373
Bucklandi Zone, 11 assemblages, 390–394
Burejospermum, 331 and bones, 376
Burmomyrma rossi, 337 fossil, 376
Kayra, 381
Langui, 381–382
C Paruro, 381
Caballichnus impersonalis, 388–389 Quebrada de la Troya, 383, 385
Calceocannabius, 388 Quebrada del Yeso, 383
Calceoformites, 5 record, 372
Camarasaurs, 148 inhabit, 371
Camarichnus, 458 K/T transition, 371
Index 477
mammal-age systems, 375 Cretonitis copripes, 345

Monte Hermoso, 390 Cretoserica latitibialis, 335
Pehuen-Có, 390 Cretotaenia pallipes, 214
South American, 375 Cretotrigona prisca, 335
Vinchina Basin, 382–385 Crime Scene Investigation (CSI), 412
Cenozoic South American mammal, 379–390 Crocodilians tracks, 6
Cenozoic succession, 387 Crustacean-produced ichnotaxa, 24–32, 59
Cenozoic Track Record, South American Callianassinae, 33
Mammals, 379–390 Cretaceous, 32
Central Atlantic Magmatic Province (CAMP), decapod burrows, 33
2, 6, 12 Decapod excavations, 32
Chaoboridae, 202 detritus-feeding shrimp, 33
Characeae, 229 Jurassic, 32
Characichnos, 188 macroevolutionary analysis, 24
Chironomidae, 202 Mesozoic, 32, 33
Chirotherium, 138 Neogene, 33
Chondrites, 9–12, 21, 46, 47, 62, 64, 65, 68, ophiomorphids, 24
70, 71, 76, 78, 80–82, 90, 96, 280, Order Decapoda, 24
283, 284, 289 paleoecological data, 24
Chondrites ichnoguild, 81 Paleogene, 33
Chondrites targionii, 283 Phanerozoic, 32
Chresmodidae, 201, 210, 216 post-Paleozoic shallow-marine
Chubutolithes, 344 ichnofaunas, 32
Chuí Formation, 33 Protocallianassinae, 33
Cicatricula, 54 thalassinidean, 24
Circolites, 53, 56, 83 Triassic, 32
Circulichnis, 454 Cruziana, 5, 9, 10, 187, 188, 242, 454, 466
Cladichnus, 303 Cruziana problematica, 192
Climactichnites, 34, 454 Cubiculum, 461
Clionoides, 54, 57 Culicoides, 278, 279
Clionolithes, 54, 73 Culicoides bullus, 278, 279
Cochlichnus, 5, 189, 223, 454 Culicoides canadensis, 278, 279
Cochlichnus anguineus, 189, 192, 198, 209 Culicoides filapalpis, 278
Coenagronidae, 228 Cunctichnus, 58
Coenobichnus, 21, 24, 31, 32, 454 Curvolithus, 47, 48, 454
Coleoptera, 182 Curvolithus simplex, 77
Colymbothetidae, 195 Cylindrichnus, 9, 21, 46, 47, 52, 63, 68, 70,
Conopsoides, 454 76, 90
Copeza, 454 Cylindrichnus concentricus (Cy), 65
Coprinisphaera, 242, 303, 324, 343–345 Cylindrichnus-Rosselia ichnoguild, 64
Coprinisphaera Ichnofacies, 301, 337, 340,
343, 344, 348
Copris leakeyorum, 346 D
Coptoclava longipoda, 214 Dagnichnus, 325, 332, 347
Coptoclavidae, 195, 211 Dagnichnus titoi, 332
Corduliidae, 228 Dasyleptid bristletails, 195
Corythosaurus, 278, 279 Davichnia, 454
Couches Rouges, 380 Deep-tier Chondrites isp., 65, 78
Cretaceous–Paleocene boundary, 86 Deep-tier Phycosiphon incertum (Ph), 65
Cretaceous–Paleogene phase, 87 Dendroctonus, 277
Cretaceous–Paleolake Baissa, 240 Dendroichnites, 225
Cretaceous Western Interior Seaway of North Dictyothylakos, 331
America, 77 Dicynodontipus, 188
Cretochaulus lacustris, 214 Didymaulichnus, 454
478 Index
Didymaulichnus lyelli, 192 Egbellichnus jordidegiberti, 225

Digitichnus, 304 Electrobatinae, 230
Dilophosaurus, 8 Empididae, 212
Dimorphichnus, 454 End-Cretaceous extinction and ecosystem
Dinopentapodiscus, 373 change
Dinosauromorphs, 139–142 biogeographic ranges, 268
Dinosaurs, 136, 137, 139–141, 149–154, continental realm, 266, 267
160, 161 ecological disruption, 269–286
early diversification, 144–145 ecospace, 269
origin and history, 142–147 extraterrestrial causation, 265
replacement dynamics, 145–147 feeding clades, 268
skeletal, 145 insect lineages, 266
timing of appearance, 143–144 K-Pg boundary, 266
Diplichnites, 5, 187, 192, 324, 327, 454, 466 K-Pg event, 267–269
Diplocraterion, 4, 5, 7, 9–12, 21, 48, 64, 68, macrobenthic colonization, 284–285
70, 76, 85, 88 macrobenthic tracemaker community, 283
Diplocraterion habichi, 68 marine realm, 266
Diplocraterion parallelum, 64, 69, 71, 77 mass extinction, 267–268
Diplopodichnus, 187, 324, 454 recovery phase, 268
Diplopodichnus biformis, 192 terrestrial and freshwater habitats, 266
Diptera, 182, 196 vertebrate lineages, 266
Dromaeosauripus, 206 End-Triassic mass extinction, 1–14, 62, 464
Dry Biome, 185–186 Entobia, 54
Duquettichnus kooli, 373 Environmental shifts, 467–468
Dysoneuridae, 211, 216 Eoameghinichnus, 373
Dytiscidae, 195 Eopentapodiscus, 373
Eosauropus, 6
Epeorominidae, 200
E Ephemeroptera, 196
Eatonichnus, 242, 339, 341, 344, 345 Ericichnus, 458
Eatonichnus claronensis, 315 Etterwindichnus, 454
Eatonichnus uthaensis, 315 Eubrontes, 6–8, 145
Echinocardium cordatum, 41 Eubrontes giganteus, 8
Echinocardium group, 41, 105 Eucommiidite, 206
Echinoderm-produced ichnotaxa, 41, 56 Euflabella, 21, 48, 52, 84, 102
Atelostomata, 43 Eumacrauchenichnus patachonicus, 390
Cretaceous and Cenozoic, 43 Euproopichnus, 454
deep-burrowing irregular echinoids, 40 Evazoum, 6, 188
diverse marine groups, 40
early Paleozoic, 42
fascioles, 43 F
holothurids, 43 Falsatorichnum, 388, 389
irregular echinoids, 40 Falsatorichnum calceocannabius, 387
Modern Evolutionary Fauna, 44 Feeding community, 280–283
ophiuroids, 43 Felipeda felis, 375
Paleozoic and post-Paleozoic echinoids, 42 Felipeda lynxi, 375
spatangoids, 40 Feoichnus, 324, 344
Spatangus group, 41 Feoichnus challa, 341
sulfide-rich and anoxic sediment, 43 Fictovichnus, 315, 324, 339, 346
Ecospace occupation, 60, 103–104, 226, Fictovichnus gobiensis, 338, 346
234–237, 241, 245, 267, 269, 288, Fleaglellius, 325, 346
461–462, 466–467 Fleming Fjord Formation, 189
Edaphichnium, 325, 341, 347 Foersterichnus, 24, 32, 454
Edaphichnium lumbricatum, 309 Footprints of Antofagasta, 389
Index 479
Footprints of Claromecó, 389 Halopoa isp., 77

Forcipomyia, 278 Halovelia electrodominica, 230
Fossil record across the T–J Boundary, 3–12 Harpepus, 454
marine record Harpichnus, 454
deep-sea ichnofauna, 9 Hawaii Volcanoes National Park site, 432
shallow-marine ichnofauna, 9–12 Helicodromites, 48, 89
terrestrial record, 5–8 Helicopsychidae, 211
Fuersichnus, 324 Helicotaphrichnus, 53, 57, 58
Fuersichnus communis, 190 Heliocopris antiquus, 346
Fuscinapeda sirin, 385 Hell Creek Formation, 274
Helminthoida, 12
Helminthoidichnites, 5, 189, 454, 466
G Helminthoidichnites tenuis, 192, 193, 198
G2/3 footprints, 414 Helminthopsis, 5, 189, 385, 454, 466
Gastrochaenolites, 21, 35, 36, 38, 53–56, 61, Helminthopsis abeli, 192, 198
62, 73, 79, 81, 83, 86, 92, 450–451 Helminthopsis hieroglyphica, 198, 209
Gerontoformica cretacica, 337 Hemeroscopidae, 210
Gerridae, 230 Hemeroscopus baissicus, 214
Getty Conservation Institute, 414 Hemiptera, 196, 210
Gigantopithecus, 437, 438 Herpystezoum, 454
Glacial and interglacial oscillations, 396 Heteropodichnus, 454
Glaciichnium, 225, 454 Heterotripodichnus, 454
Global carbon system, 2 Hettangian, 7
Global ichnodiversity, 61–63, 70, 84–85, 89, Hexapodichnus, 454
93, 95, 103, 199, 233–235, Hillichnus, 22, 35, 36, 39, 40, 48, 106
336–337, 339, 348, 450–453, Hillichnus lobosensis, 38
458–463 Hipparion, 415
Global Stratotype Section and Point (GSSP), 1 Hippipeda, 390
Glyphichnus, 21, 26, 31 Hippipeda aurelianis, 375
Gnathichnus, 53, 56, 62, 73, 92, 458 Holocene
Gordia, 454, 466 footprints, 420
Gordia indianaensis, 192 track sites, 430
Gordia marina, 192 Holometabolan taxa, 201
Grallator, 6–8, 145, 147, 188 Hominin evolution
Granulohyalichnus, 450–451 body and trace fossil records, 435
Great Ordovician Biodiversification Event, etymology, 412
453, 459, 462 footprint-bearing sites, 411
Green River biota, 229 hominid traces, 436
Gregariousness, 152–155 Ichnos, 412
Grypopentapodiscus, 373 Lake Turkana, 415–416
Gwyneddichnium, 6, 188, 191 non-volcaniclastic substrates, 436
Gymnopleurus eocaenicus, 346 Out of Africa, 416–419
Gyrochorte, 48 pre-Late Pleistocene, 436
Gyrochorte comosa, 66, 77 pre-sapiens track, 435
Gyrolithes, 22, 26, 29, 30, 32, 80, 88 sea shore, 431
shadow of volcanoes, 432–433
trace fossils, 412
H track record, 422–426, 437
Haenamichnus, 159, 206 Hominipes modernus, 413, 430, 432, 435, 436
Haentzschelinia, 22, 48, 52, 66, 75 Homo erectus (H. ergaster), 415–418, 421, 435
Haentzschelinia ottoi, 66, 67 Homo floresiensis, 416, 437
Haidomyrmex cerberus, 337 Homo heidlebergensis, 418, 421
Haidomyrmodes mammuthus, 337 Homo neaderthalensis, 425
Halopoa, 466 Homo neanderthalensis, 413, 426, 436, 439
480 Index
Homopodichnus, 454 Kouphichnium, 5, 9, 10, 454

Homo sapiens, 413, 424, 428, 429, 435–439 K-Pg mass extinctions, 84, 87, 218, 227,
ichnologic record, 419–427 265–291, 341, 372–373, 463–465
Middle-Late Pleistocene, 419 K-Pg plant–insect interactions, 270
Nahoon Point tracks, 419 Krausichnus, 325, 344, 346
Human footprints Krausichnus pagodus, 310
Pleistocene, 415 Krausichnus trompitus, 310
Sebkra el Azrag, 431
Theopetra Cave, 422
Vârtop Cave, 422 L
Human-like foot anatomy, 435 Labyrintichnus terrerensis, 308
Hydrophilopsia baissensis, 214 Lacustrine ecosystems, 182–184, 238–241, 244
Hydropsychidae, 228, 229 bioturbation, 183
Hydroptilidae, 229 dispersal adaptations, 183, 184
Hymenoptera, 196 evolutionary novelties and innovations, 184
exploration, 182
formation, isolation and destruction, 183
I long-lived lakes, 184
Ichnofabrics, 10, 47, 68, 70, 80, 82, 86, 90, taphonomic megabiases, 182
99–101, 103–105, 187, 209, 236, Lacustrine trophic webs
280, 285–287, 344, 468 bottom–up processes, 238
Ichnoguilds, 60, 64, 67–68, 75–76, 80–81, ecosystem structure and function, 238
89–90, 92, 104, 234, 236 energy-flow diagrams, 239
Ichnos, 412 evolutionary history, 238
Ichnotaxonomic problems, 241 herbivore-driven food web, 240
Ignotomis, 206 Paleolake Messel, 241
Ignotornis, 155 Paleozoic lakes, 239
Insect herbivory, 271–277 physiochemical conditions, 241
Insect pollination, 277–278 Laetoli hominin trackways, 411
Intercontinental travel frontiers, 428–431 Lagerpeton, 140
Intertidal carbonate-sand flat, 25 Lagerstätten, 182
Intra-Valanginian Discontinuity, 79 Lajas Formation, 67
Intricatus, 283 Lamaichnum guanicoe, 390
Iribarnichnum megamericanum, 390 Langui footprints, 381–382
Irichnus, 454 Langui ichnoassemblage, 392
Isochirotherium, 138 Lapispecus, 57
Ixalichnus, 454 Lapispira, 22, 49, 52, 75
Las Flores Formation, 386
Las Hoyas fossil site, 240
J Late Permian tetrapods, 137
Jamesonichnites, 45, 48 Lazaichnus, 324, 344
Jindongornipes, 206 Lepidostomatidae, 211
Jurassic-Cretaceous boundary, 155 Leptoconops, 278
Lestidae, 228
Libellulidae, 228
K Liliensternus, 145
Katbergia, 331, 347 Lilstock Formation, 11
Katbergia carltonichnus, 321 Limnephilidae, 229
Kayentapus, 8 Limoniidae, 196, 202
Kayra footprints, 381 Linckichnus, 5
Keircalia, 454 Lithanomala, 335
Kimberichnus, 454 Lithographus, 454
Koranornis, 206 Lockeia, 5, 9, 10, 22, 35–37, 39, 61, 62, 88, 187
Korymbichnus, 49 Lockeia amygdaloides, 66, 190
Index 481
Lockeia siliquaria, 38, 189, 191 ichnologic record, 20, 106

Loloichnus, 325, 344, 347 ichnotaxa, 20
Loloichnus baqueroensis, 316, 332 ichnotaxonomic treatment, 52
Lonchidion, 239 infaunalization, 60–93
Lophoctenium, 22, 35, 37 Jurassic, 63–75
Lumbricus terrestris, 284, 285 marine paleocommunities, 20
Lusatichnium, 225, 454 mixed layer, 99–101
Lycoptera, 214 modern-marine ecosystems, 19
Neogene, 88–93
neoichnological observations, 20
M paleoenvironmental breadth, 93–96
Macanopsis, 22, 26, 30, 67, 88, 321, 331 Paleogene, 84–87
Macanopsis plataniformis, 29 paleogeography and climate, 101–103
Macaronichnus, 22, 44, 49, 52, 76, 84, 85, 88, post-Paleozoic bioturbators, 106
95, 102 post-Paleozoic shallow-marine
Macaronichnus segregatis, 77 environments, 20
Macoma balthica, 105 in shallow-water settings, 106
Macrauchenichnus, 383, 389 sponges (see Sponges)
Macrauchenichnus rector, 385, 387 tiering structure, 60–93
Maculichna, 454 Triassic, 61–63
Madygen deposits, 194 worms, 57–58
Madygenerpton, 239 Metacatharsius rusingae, 346
Maeandropolydora, 57, 61, 92 Microvory, 202
Maiakarichnus, 22, 26, 29, 31, 33 Milodontidichnum, 389
Manus, 372 Minisauripus, 206
Mariño Formation, 385–386 Mioclanis shanwangiana, 344
Marmarthia pearsoni, 274–275 Mirandaichnium, 454
Megalamaichnum tulipensis, 390 Mitchellichnus, 454
Megaloptera, 182 Mixed layer depth, 99
Megalosauripus, 8 Mixed layer traces, 100
Megatherichnum oportoi, 387, 389, 397 Modern Evolutionary Fauna (MEF), 19, 20,
Megatherium, 387 32, 74, 75, 84–87, 103
Megatherium americanum, 398 Monesichnus, 324, 344
Melittosphex burmensis, 335 Monocraterion, 5, 9
Mermia, 181, 189, 454 Monomorphichnus, 10, 454
Merostomichnites, 9, 10, 454 Monomorphichnus lineatus, 198
Mesoleuctra, 195 Monte Hermoso, 390
Mesoleuctra–Mesoneta assemblage, 203 Morazatermes krishnai, 335
Mesonetidae, 200 Morrison Formation, 203, 330
Mesozoic mammals, 371 Moyenisauropus, 8
Mesozoic marine revolution (MMR), 24–40, Multina, 467
54–55, 450 Multipodichnus, 454
algae, 59 Myanmyrma gracilis, 337
behavioral evolution, 96–99 Myriapodites, 454
bioeroders, 52
bioturbation and ichnofaunas, 103–106
bryozoans, 58 N
coevolutionary developments, 19 Naktodemasis, 339
cretaceous, 75–84 Naktodemasis bowni, 320
crustaceans, 58–59 Naviculichnium, 44, 93
echinoids, 56 Naviculichnium marginatum, 94
foraminifera, 59–60 Necrotaulidae, 195, 216
fungi, 59 Neomegatherichnum pehuencoensis, 389, 397
ichnofaunal composition, 60–93 Nereites, 49, 468
482 Index
Neuroptera, 182 Paleodictyon, 242

Nododendrina, 60 Paleogene insects, 227
Notonectidae, 195 Paleosol ichnofacies
Nummipera, 86 Paleosol trace fossils, 302–329
Nyasasaurus, 144, 145 Paleo-thermocline, 192
Paleozoic Evolutionary Fauna (PEF), 20
Pallichnus, 324, 339, 344
O Pallichnus dakotensis, 309, 316, 338
Ochteridae, 196 Palmichnium, 454
Octopodichnus, 454 Palmiraichnus, 324, 344
Ocypode quadrata, 25, 90 Palmiraichnus castellanosi, 303
Odocoileinichnum commune, 390 Parabrontopodus, 8, 147
Oecanthoperlidae, 216 Paracanthorhaphe togwunia, 198
Oichnus, 53, 55, 73, 74, 86, 92 Paradictyodora, 39
Oldhamia, 464, 466 Paramylodon, 398
Oligichnos limnos, 224 Paranthropus boisei, 416
Oneonta Formation, 327 Parasynaptichnium, 138
Oniscoidichnus, 10, 454 Pareiasaurs, 137
Onshore origination, 105–106, 244, 467–468 Parmaichnus, 31
Ophioichnus, 22, 43, 44, 454 Parowanichnus, 339
Ophiomorpha, 22, 26, 28, 30–33, 64, 66–68, Parowanichnus formicoides, 315
70, 74, 76, 80, 84, 85, 88–91, 93, Paruro footprints, 381, 393
102, 280, 468 Passaic Formation, 191
Ophiomorpha annulata, 94, 283 Patagonichnus, 22, 49, 52, 85, 88
Ophiomorpha irregulaire, 77 Paucituberculata, 378
Ophiomorpha nodosa (Op), 90–91 Pecoripeda amalphaea, 375
Ophiomorpha rudis, 283 Pecoripeda dicrocervoides, 375
Opisthocoelicaudia, 148 Pecoripeda djali, 375
Oravaichnium, 35 Pecoripeda gazelle, 375
Orchesteropus, 454 Pecoripeda satyri, 375
Ornithischian tracks, 6, 7 Pehuen-Có, Argentina, 390, 391
Ornithopodichnus, 206 Pelycosaurs, 137
Oro Grande Site, 430 Perleididae, 238
Orthogonum, 59 Permichnium, 454
Oryctoantiquus borealis, 335 Petalichnus, 454
Osteocallis, 458 Petroxestes, 53, 56
Otozoum, 6, 7 Phanaeus antiquus, 346
Ovipositional lesions, 201 Philopotamidae, 196
Phoebichnus, 24, 49, 50, 52, 67, 68, 75, 76
Phoebichnus trochoides, 66, 69
P Phoenicopterichnum rector, 390
Palaeochinastacus australianus, 331 Pholeus, 23, 26, 30, 32, 61
Palaeocopris labreae, 346 Phryganeina, 201
Palaeogyrinus, 230 Phycodes, 9
Palaeoniscidae, 238 Phycosiphon, 23, 47, 50, 64, 66–68, 76, 80,
Palaeophycus, 5, 9–11, 22, 49, 52, 62–64, 67, 90, 280
68, 70, 76, 85, 90, 188, 385, 465 Phycosiphon incertum (Ph), 65, 67
Palaeophycus heberti, 65 Phycosiphon-Chondrites ichnoguild, 64
Palaeophycus striatus, 192 Pilichnus, 466
Palaeophycus tubularis, 192, 283 Pilothylakos, 331
Palaeosabella, 53, 57, 61 Planolites, 5, 9–12, 23, 50, 52, 62, 63, 68, 70,
Paleocene tetrapod footprints, 374 76, 80, 82, 85, 90–91, 100, 188,
Paleocene–Eocene Thermal Maximum 280, 283, 285, 303, 465
(PETM), 269, 341 Planolites annularis, 209
Index 483
Planolites beverleyensis, 64 Q
Planolites ichnoguild, 80 Quebrada de la Troya footprints, 383, 385
Planolites montanus (Pl), 65, 189 Quebrada del Jarillal Formation, 387
Planorbis Zone, 11 Quebrada del Yeso footprints, 383
Plant and insect-feeding diversity, 276 Quirogaichnus, 242
Plant diversity, 271
Plant–insect associational
studies, 287 R
Plateosaurus, 8 Radulichnus, 53, 55, 73, 92, 458
Platyperlidae, 200 Ramosulcichnus, 57
Platystethus, 284 Rebuffoichnus, 324, 339, 344
Plecoptera, 195 Rebuffoichnus casamiquelai, 303, 338
Plecopteran nymphs, 200 Rebuffoichnus guanche, 303
Plectrotarsidae, 211 Rebuffoichnus isp., 338
Pleistocene–Holocene Bahamian-type Rebuffoichnus sciuttoi, 315, 316, 338, 346
carbonates, 90–91 Renichnus, 53, 55, 92
Plesiornis, 8 Rhadostium, 454
Plover Formation, 65 Rhizocorallium, 5, 9–12, 24
Polykladichnus, 23, 50, 62, 189, 194 Rhizocorallium commune, 468
Polykladichnus aragonensis, 308 Rhizocorallium irregulare, 64
Porcellusignum, 389 Rhizocorallium isp., 23, 61, 62, 66–68, 70, 72,
Porcellusignum consulcator, 387 74–76, 84, 85, 88, 99, 105
Portlandia arctica, 105 Rhizocorallium jenense jenense, 71
Praehominipes, 435 Rhizocorallium jenense spinosus, 71
Praehominipes laetoliensis, 413, 432, 435 Rhizolith ichnofacies, 327–329, 333, 337, 347
Pre-Planorbis Beds, 11 Rhynchosauroides, 5–8, 147, 188, 192
Prionocephale deplanate, 345 Rio Chico Group, 341
Prorhyacophilidae, 195 Río Negro Formation, 387–389
Prorotodactylus, 140 Rogerella, 53, 58, 60, 73, 83, 92
Prosauropod tracks, 6, 7 Ropalonaria, 53, 58
Prostomium, 45 Rosellichnus, 303, 346
Protichnites, 192, 454 Rosselia, 50
Protocallianassa, 33 Rotodactylus, 140, 141
Protochirotherium, 138 Rusophycus, 5, 10, 187, 188, 242, 466
Protovirgularia, 9, 10, 23, 35–37, 39, 61, 62, Rusophycus stromnessi, 192
75, 76, 88, 95, 454
Protovirgularia dichotoma, 77
Psammichnites, 34 S
Pseudameghinichnus, 373 Salinity barrier, 241, 243, 244
Pseudotetrasauropus, 6 San Gerónimo Group, 379–382
Psilonichnus, 30 Santa Cruz Formation, 385
Psiloceras spelae, 10 Sarjeantipes whitea, 374
Psilonichnus, 23, 26, 67, 84, Saronichnus, 23, 35–37, 39, 40, 88, 106
88, 90 Saronichnus abeli, 38
Psilonichnus upsilon, 29, 90–91 Sauropodomorph tracks, 6
Psychodidae, 202 Sauropods, 148, 149, 154
Pteraichnus, 158, 206 Sauropod tracks, 8
Pteraichnus saltwashensis, 158 Scalichnus, 5, 23, 35, 36, 39, 88
Pterichnus, 454 Scalichnus phiale, 38
Pterosaur locomotion, 157–160 Scaphichnium, 309
Pterosaurs, 139, 157–160 Schaubcylindrichnus, 23, 47, 50, 52, 63, 67,
Ptychoplasma, 35 68, 76, 79, 85, 90
Pumaeichnum biancoi, 390 Schaubcylindrichnus coronus, 77
Punctichnium, 454 Schizophoridae, 195, 201
484 Index
Sciomyzidae, 212 Survivability, 465

Scolicia, 41, 42, 89, 309, 468 Synapsids tracks, 6
Scoyenia, 5, 181, 186, 189, 191, 321, 324 Synaptichnium, 138
Scoyenia beerboweri, 327 Syntermesichnus fontanai, 308
Scoyenia gracilis, 189–191
Scoyenia Ichnofacies, 327, 336, 340, 347, 385
Secundumichnus, 454 T
Seilacherian ichnofacies, 301 Tacheria troyana, 383
Selenichnites, 10, 327 Taenidium, 5, 10, 51, 188, 303, 304, 309, 320,
Semidendrina, 59 321, 324, 346, 385
Sepkoski’s curves, 459 Taenidium barretti, 209, 308
Sericostomatidae, 216, 229 Taenidium-Phycosiphon ichnoguild, 80
Shandongornipes muxiai, 157 Taenidium serpentinum, 64, 303, 320
Shrew opossums, 378 Talpina, 53, 57, 60, 73
Sialis strausi, 230 Taphrhelminthoides, 454
Siberioperla, 195 Tapinoma nigerrima, 284
Siberioperlidae, 200 Tasmanitids, 185
Sieblosiidae, 227 Taxonomic revisions, 242
Simuliidae, 202 Teichichnus, 5, 9, 10, 23, 47, 51, 52, 64, 68,
Sinusichnus, 23, 26, 31, 32 70, 76, 78, 80, 82, 85, 90
Sinusichnus sinuosus, 29 Teichichnus-Chondrites ichnoguild, 76
Siphlonuridae, 196, 200 Teichichnus flexuosus, 52
Siphonichnus, 23, 35, 36, 39, 64, 68, 88 Teichichnus ichnoguild, 64
Siphonichnus eccacensis, 38 Teichichnus nodosus, 52
Skolithos, 5, 9, 10, 12, 23, 50, 62, 68, 76, 85, Teichichnus patens, 52
88, 90, 189, 192, 194, 303, 308, Teichichnus rectus, 52, 65, 69
320, 321, 324, 327, 338, 346 Teichichnus sigmoidalis, 52
Skolithos linearis, 90–91 Teichichnus spiralis, 52
Socialites tumulus, 339 Teichichnus zigzag, 52
Solemyatuba, 37 Teisseirei, 315, 324, 344, 346
Solemyatuba ypsilon, 38 Teisseirei barattinia, 344
Sonidae, 210 Temnospondyl amphibians, 137
Spatangus group, 41 Teredolites, 83
Sphecomyrma sp., 337 Termitichnus, 242, 309, 325, 346
Sphecomyrmodes orientalis, 337 Termitichnus Ichnofacies, 336, 339, 340
Sphenodontian track, 6 Terra Amata footprint, 436
Sphingopus, 140 Tetrapods, limb postures, 137
Sphinx snelleni, 344 Tetrasauropus, 6, 188
Spirichnus, 53, 58 Thalassinidean, 24
Spongeliomorpha, 5, 23, 26, 29, 32, 74, 88, Thalassinoides, 9–12, 23, 24, 26, 27, 30–32,
188, 191, 324, 466 61–67, 70, 71, 74, 76, 78–80,
Spongeliomorpha milfordensis, 189, 191 82–84, 88, 89, 93, 100–102, 192,
Sponges, 52–54, 62, 73, 81, 83, 84, 86, 92, 95 280, 283, 285, 289, 290, 463
Stegomastodonichnum australis, 390 Thalassinoides ichnoguild, 80, 81
Steinsfjordichnus, 454 Thalassinoides paradoxicus, 81
Stiallia, 187 Thalassinoides suevicus, 77, 94
Stiaria, 466 Theropods, 8, 142, 145, 147, 150–152, 154,
Stomatopod crustaceans, 87 160, 161
Stratiomyiida, 212 Theropod tracks, 6
Striatichnium, 454 Titanosauriformes, 148
Stychopterus, 214 Tombownichnus, 315, 324, 344, 345
Subtidal callianassid mound, 25 Tombownichnus parabolicus, 303
Sundance Biota, 207 Tombownichnus plenus, 303
Sundance Formation, 203 Treptichnus, 5, 189, 466
Index 485
Treptichnus pedum, 45 Westbury Formation, 10

Treptichnus pollardi, 189, 191, 192 Wet Biome, 185, 186
Triaplidae, 195 Worm-produced ichnotaxa, 45–51,
Triassocoridae, 195 57–58
Trichichnus, 283 bioturbators, 44
Trichoptera, 182, 195, 216, 228 echiurans, 45
Trisulcus, 454 enteropneusts, 45
Trypanites isp., 53, 57, 61, 62, 73, 83, 86, 92 evolutionary history, 51
Tuberculichnus vagans, 198 nemerteans, 44
Tubotomaculum, 31 ophelids (scolecids), 44
Tubulohyalichnus, 450–451 polychaetes, 44, 51
Tumblagoodichnus, 454 post-Paleozoic rocks, 52
priapulids, 45
scolecida and palpata, 44
U sipunculids, 45
U- and Y-shaped burrows, 243 sophisticated spreiten, 45
Uhangrichnus, 206
Undichna, 189, 208
Undichna bina, 192 X
Undichna britannica, 192 Xenarthrans, 377
Uruguay, 324 Xylonichnus, 5
V Y
Vagorichnus, 467 Younger Holocene footprint
Vagorichnus anyao, 198 sites, 431
Venatoripes riojanus, 383, 385, 389
Vinchina Basin, Argentina, 382–385
Vinchina Formation, 383 Z
Vitimotaulidae, 211, 216 Zacallatidae, 227
Vondrichnus, 242, 325, 346 Zoophilous pollen, 277
Zoophycos, 51, 61, 76, 80, 96, 99, 101, 105,
280, 283, 284, 289
W Zoophycos-Chondrites ichnoguild, 81
Warvichnium, 225 Zoophycos evolution, 97, 98
Wasp cocoons, 279 Zoophycos-Taenidium ichnoguild, 81

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Topics in Geobiology 40

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8 The Late Triassic Mass Extinction Event ............................................. 1

16 Recurrent Patterns and Processes: The Significance

Index ................................................................................................................. 475

“A beginning has been made and the discipline has been

paleobiologic literature shows that there is an increased awareness of the impor-

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

Colin Barras and Richard J. Twitchett

© Springer Science+Business Media Dordrecht 2016 1

excursions from a number of localities worldwide, including among others example

8.2 Trace-Fossil Record Across the T–J Boundary

8.2.1 Terrestrial Record

8.2.2 Marine Record

8.2.2.1 Deep-sea Ichnofauna Across the T–J Boundary

Nearly 30 ichnogenera inhabited the deep-sea in the Carboniferous, before a halv-

8.2.2.2 Shallow-Marine Ichnofauna Across the T–J Boundary

dysaerobic conditions contain common Chondrites, with rare Rhizocorallium

However, while there is evidence of (ichno)taxonomic extinction in the terres-

Luis A. Buatois, Noelia B. Carmona, H. Allen Curran, Renata G. Netto,

Modern-marine ecosystems are the result of numerous evolutionary innovations

L.A. Buatois (*) • M.G. Mángano

© Springer Science+Business Media Dordrecht 2016 19

9.2 The Cast of Characters

9.2.1 The Main Groups of Burrowers

The MEF is dominated by bivalves, gastropods, echinoids, crustaceans, and marine

Phoebichnus Jurassic-Pleistocene Worm-like organisms, Fodinichnia Deposit-feeding Gallery biodiffusive

Crustaceans of the Order Decapoda (Superclass Crustacea, Class Malacostraca) are

Ophiomorpha refers to simple to complex, branching burrows with distinctive,

Fig. 9.3 Examples of Thalassinoides in Mesozoic–Cenozoic shallow-marine deposits. (a) Vaca

Fig. 9.4 Examples of Ophiomorpha in Mesozoic–Cenozoic shallow-marine deposits. (a)

Tubotomaculum consists of spindle-shaped burrows, displaying a spreite and

Gastropods and Polyplacophorans

Locomotion and feeding structures produced by gastropods and chitons can be

exhibited by several other major groups (e.g., echinoids), is widely recognized

Fig. 9.6 Schematic reconstructions of bivalve-produced ichnogenera. (a) Reconstruction of the

Fig. 9.7 Preservational variants of Protovirgularia in Mesozoic–Cenozoic shallow-marine depos-

some specimens can represent semipermanent domiciles (Seilacher 1953; Rindsberg

feeding structure produced as a result of a chemosymbiotic life strategy. At present,

due to sparse occurrence. Hillichnus reflects a level of behavioral complexity

polychaete larval infestation in recently sediment-filled bivalve borings.

reported that Cylindrichnus concentricus ichnofabrics occur commonly in Mesozoic

This ichnogenus is interpreted as the locomotion trace of infaunal predators, such as

range of shallow- to deep-marine and even continental environments, being most

9.2.2 The Main Groups of Bioeroders

Sponge borings typically consist of anastomosing channel networks generally form-

of Entobia. Therefore, the ichnospecies Entobia devonica is an early representative

Gastropods typically produced predatory borings, such as Oichnus (Wisshak et al.

Gastrochaenolites comprises clavate or drop-like trace fossils that are circular to

Helicotaphrichnus is a helicoidal boring made in the columella of gastropod