Atlas of Trace Fossils in Well Core

Dirk Knaust
Atlas of
Trace Fossils
in Well Core
Appearance, Taxonomy
and Interpretation
Atlas of Trace Fossils in Well Core
Dirk Knaust
Atlas of Trace Fossils in

Well Core
Appearance, Taxonomy and Interpretation
123
Dirk Knaust
Statoil ASA
Stavanger
Norway
ISBN 978-3-319-49836-2 ISBN 978-3-319-49837-9 (eBook)

DOI 10.1007/978-3-319-49837-9
Library of Congress Control Number: 2016958461
© Springer International Publishing AG 2017

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Cover image: Sectioned well-core sample of a cross-bedded sandstone with mud drapes and floating granulae,
comprising the cross section of a large spreite burrow assigned to Teichichnus zigzag. Middle Jurassic (Bathonian)
Tarbert Formation (tidally influenced delta front), Oseberg Sør Field, Norwegian North Sea (well 30/9-4S, ca.
3347.2 m). Image width ca. 9 cm. Photograph taken by the author.
Printed on acid-free paper
This Springer imprint is published by Springer Nature

The registered company is Springer International Publishing AG
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For my son and daughter,
Per Ragnar and Solveig Alida
Foreword
It is not easy to tell which of the graduate students will become masters in their field. When I
met Dirk Knaust in 1997 at the Fourth International Ichnofabric Workshop on San Salvador, he
was just another graduate student, obviously very intelligent, but there were others with longer
strings of publications. But as the years progressed, Dirk’s work in ichnology accelerated, and it
became clear that he is a perfectionist whose body of work is ripening at last. Building on his
doctoral study of the Muschelkalk, he expanded his range to the North Sea and many other
regions, and as we all know, the best ichnologist is the one who has seen the most trace fossils.
In his maturity, he has produced a series of papers revising Asteriacites, Rhizocorallium,
Balanoglossites, Pholeus, Oichnus, and other ichnogenera (Knaust 2002, 2008, 2013; Wisshak
et al. 2015; Knaust and Neumann 2016), brought the world’s attention to extraordinarily
preserved tracemakers in the Muschelkalk (Knaust 2010a, b), and elucidated the connection
between bioturbation and reservoir quality (Knaust 2009a, b; Knaust et al. 2014).
Dirk’s most important effort in recent years has been, together with Richard G. Bromley, to
organize and edit Trace Fossils as Indicators of Sedimentary Environments, a thick volume
that brings ichnology to sedimentologists and other geologists—and sedimentology back to
ichnologists (Knaust and Bromley 2012). He carried an advance copy of the book to Ichnia
2012, where it was a big hit among ichnologists on the long drives between field trip stops on
the Avalon Peninsula of Newfoundland. The copy passed slowly from hand to hand to the rear
of the bus and back to the front again. I think he must have sold to forty people on the bus
within a few hours.
Currently, Dirk is at work with me on leading a revision of the trace-fossil volume of the
Treatise on Invertebrate Paleontology. As he pointed out in a keynote address at Ichnia 2016,
the number of named invertebrate ichnogenera has more than doubled since Walter Häntz-
schel’s last revision in 1975. After Bertling et al. (2006) reached a consensus on which criteria
were best for differentiating ichnotaxa, Dirk applied these ichnotaxobases systematically to
the entire corpus of invertebrate trace fossils, using them to define categories within a key
(Knaust 2012)—an effort that required consulting the diagnoses and major revisions of every
ichnogenus. He reasoned that ichnogenera falling within the same category are potential
synonyms, and indeed some have been found by this method. The key is effectively a massive
test of the consensus criteria, generating predictions that might otherwise not be investigated.
It is also a way to guide ichnologists toward a viable classification of trace fossils that utilizes
ichnofamilies within the context of tracemaker behavior and paleoenvironments. In a recent
paper, Dirk erected a new ichnofamily, the Siphonichnidae, to accommodate bivalvian activity
as diverse as those recorded in Parahaentzschelinia, Scalichnus, and Hillichnus (Knaust
2015). He has also drawn attention to the meiobenthos as significant agents of bioturbation,
even identifying fossil tracemakers in some cases (Knaust 2010a, b).
The book at hand, Atlas of Trace Fossils in Well Core—Appearance, Taxonomy and
Interpretation, was developed over a period of years from the series of in-house workshops
that Dirk has led from 2011 onwards. It is clear that this work comes from long gestation of
ideas and presentation. It includes two sections: First, a brief introduction acquaints the reader
with the fundamentals of ichnology, with special regard to their use in petroleum geology.
vii
viii Foreword
Second, in the main part of the book, 39 genera of trace fossils and associated features are
discussed individually. For each ichnogenus, sections are given on their morphology, fill and
size, ichnotaxonomy, substrate, appearance in core, similar trace fossils, producers, ethology
(behavior), depositional environment, ichnofacies, age, and finally reservoir quality, accom-
panied by a generous number of illustrations. The treatment is condensed but draws on a very
broad knowledge of the literature as well as extensive personal experience. Many of the
illustrations are new; others are borrowed from the best in the literature. For many ichno-
genera, the criteria for identification of the major ichnospecies are given, and close attention is
paid to possible tracemakers and their behavior. Is such detail necessary? Not for every
project, but the studies that pay attention to ichnospecies do yield finer details about depo-
sitional environments than those which identify trace fossils only to ichnogenus, or ignore
ichnotaxonomy altogether.
The trace fossils that have been chosen for treatment in this volume include all the
ichnogenera that are ordinarily found in cores, whether taken for academic or economic
purposes, plus ichnogenera that are common in geological contexts with which Dirk is
familiar. The result is that here is not only the most up-to-date compendium of information on
trace fossils in cores, but also an advancement of science on trace fossils such as Phoebichnus
that many ichnologists have not recognized in their own material.
Anyone who wishes to use trace fossils in core or outcrop can benefit from reading this
book, but those who want information on how trace fossils determine the porosity and per-
meability of rocks will find it particularly useful (Knaust 2014). Dirk gives specific infor-
mation about the properties that each ichnogenus may lend reservoir rocks, whether aiding or
hindering the flow of pore fluids.
Need I remind readers that what controls petroleum flow also applies to groundwater?
Hydrogeologists and environmental geologists may be pleasantly surprised to find that they
can better understand the properties of aquifers and aquitards by reference to ichnology. This
is a barely touched area of science, but one that will becoming increasingly important as water
tables fall and engineering projects respond to environmental changes. The permeability of the
limestone underlying Miami, for instance, is determined partly by its trace fossils. It will not
be surprising if the stratigraphic distribution of trace fossils turns out to determine the fate of
different areas of Florida as sea level rises.
You have in your hands an authoritative book, one that will have an influence for years to
come throughout the field of invertebrate ichnology as well as aspects of sedimentology,
petroleum geology, and environmental geology. It has built on decades of work by the
ichnological community and Knaust’s own, and is a fitting companion for the scientist who
works with trace fossils.
July 2016 Andrew K. Rindsberg

University of West Alabama
Livingston
References
Bertling M, Braddy SJ, Bromley RG et al (2006) Names for trace fossils: a uniform approach. Lethaia 39:
265–286
Knaust D (2002) Ichnogenus Pholeus Fiege, 1944, revisited. J Paleontol 76:882–891
Knaust D (2008) Balanoglossites Mägdefrau, 1932 from the Middle Triassic of Germany: part of a complex trace
fossil probably produced by boring and burrowing polychaetes. Paläontologische Zeitschrift 82:347–372
Knaust D (2009a) Ichnology as a tool in carbonate reservoir characterization: a case study from the Permian–
Triassic Khuff Formation in the Middle East. GeoArabia 14:17–38
Knaust D (2009b) Characterisation of a Campanian deep-sea fan system in the Norwegian Sea by means of
ichnofabrics. Mar Pet Geol 26:1199–1211
Knaust D (2010a) Remarkably preserved benthic organisms and their traces from a Middle Triassic (Muschel-
kalk) mud flat. Lethaia 43:344–356
Foreword ix
Knaust D (2010b) Meiobenthic trace fossils comprising a miniature ichnofabric from Late Permian carbonates
of the Oman Mountains. Palaeogeogr Palaeoclimatol Palaeoecol 286:81–87
Knaust D (2012) Trace-fossil systematics. In: Knaust D, Bromley RG (eds) Trace fossils as indicators of
sedimentary environments. Developments in Sedimentology, vol 64, pp 79–101
Knaust D (2013) The ichnogenus Rhizocorallium: classification, trace makers, palaeoenvironments and
evolution. Earth Sci Rev 126:1–47
Knaust D (2014) Classification of bioturbation-related reservoir quality in the Khuff Formation (Middle East):
towards a genetic approach. In: Pöppelreiter MC (ed) Permo-Triassic Sequence of the Arabian Plate. EAGE,
pp 247–267
Knaust D (2015) Siphonichnidae (new ichnofamily) attributed to the burrowing activity of bivalves: ichno-
taxonomy, behaviour and palaeoenvironmental implications. Earth Sci Rev 150:497–519
Knaust D, Bromley RG (2012) Trace fossils as indicators of sedimentary environments. Developments in
Sedimentology, vol 64. Elsevier, Oxford, xxx+960 pp
Knaust D, Neumann C (2016) Asteriacites von Schlotheim, 1820—the oldest valid ichnogenus name—and
other asterozoan-produced trace fossils. Earth Sci Rev 157:111–120
Knaust D, Warchoł M, Kane IA (2014) Ichnodiversity and ichnoabundance: revealing depositional trends in a
confined turbidite system. Sedimentology 62:2218–2267
Wisshak M, Kroh A, Bertling M et al (2015) In defence of an iconic ichnogenus—Oichnus Bromley, 1981.
Ann Soc Geol Pol 85:445–451
Preface
This book provides the reader with a blend of high-quality photographs, figures, and accom-
panied text for the identification of trace fossils in well core and outcrop. Ichnological data are
becoming more and more crucial in sedimentological and paleoenvironmental interpretation,
not only in the exploration and exploitation of hydrocarbon but also in the characterization of
aquifers and in scientific drilling. Key features include the identification and interpretation of
trace fossils in core and outcrop, integrated sedimentological–ichnological core logging, and
hydrocarbon reservoir characterization. It has been prepared for an audience in the fields of
sedimentology, paleontology, and petroleum geoscience from academia (graduate students and
professionals) and industry (reservoir geologists).
After an introduction to the study of trace fossils in well core and an outlining of ichno-
logical basics, principles and concepts, this book offers a detailed description and interpre-
tation of 39 commonly occurring ichnogenera together with recurrently associated features
such as diffuse bioturbate texture, plant roots and their traces, borings and pseudo-trace fossils.
The trace fossils are highlighted by their expression in well core, illustrated with numerous
original photographs and supplemented with carefully selected schematic drawings from the
literature. This unique information is complemented by examples of trace fossils in outcrop, as
well as relevant key figures from existing work.
Each chapter is treated in a consistent manner, stating the ichnogenus name and author in
the title, followed by sections on morphology, fill and size; ichnotaxonomy; substrate;
appearance in core; similar trace fossils; producers; ethology; depositional environment;
ichnofacies; age as well as reservoir quality. This book is rounded off with an extensive list of
references for further reading. The material for the book originated from the author’s con-
tinuous work with trace fossils, chiefly in core, over the last two decades.
The well-core examples selected for this book mainly originate from the Norwegian
Continental Shelf, which has been subject to extensive exploration and exploitation for oil and
gas over the last half-century, although data from other regions of the world have been added.
Based on this, siliciclastic rocks are overrepresented as compared to carbonates, and the
majority of material comes from Mesozoic strata; however, all major paleoenvironments are
covered. The presented trace fossils and associated features are thus just examples of possible
occurrences in core, and other regions or stratigraphical units may return other interesting
ichnological data. It is my hope that this book will promote further studies in this field.
Many colleagues and friends have shared their ideas over the last years, as well as
specimens and literature; these include, in alphabetical order:
Andrea Baucon (Milano), Zain Belaústegui (Barcelona), Markus Bertling (Münster),
Richard G. Bromley (Copenhagen), Luis A. Buatois (Saskatoon), Richard H.T. Callow
(Stavanger), Kevin J. Cunningham (Miami), H. Allen Curran (Northampton), Andrei V.
Dronov (Moscow), Allan A. Ekdale (Salt Lake City), Christian C. Emig (Marseille), Christian
Gaillard (Lyon), Jorge F. Genise (Buenos Aires), Jean Gérard (Madrid), Jordi de Gibert
(Barcelona, deceased), Murray K. Gingras (Edmonton), Roland Goldring (Reading, deceased),
Murray R. Gregory (Auckland), Hans Hagdorn (Ingelfingen), Geir Helgesen (Stavanger),
William Helland-Hansen (Bergen), Günther Hertweck (Wilhelmshaven), Sören Jensen
xi
xii Preface
(Badajoz), Jostein Myking Kjærefjord (Bergen), Christian Klug (Zurich), Kantimati Kulkarni
(Pune), James A. MacEachern (Burnaby), M. Gabriela Mángano (Saskatoon),
Anthony J. Martin (Atlanta), Allard Martinius (Trondheim), Duncan McIlroy (St. John’s),
Renata Meneguolo (Stavanger), Radek Mikuláš (Praha), Masakazu Nara (Kochi), Carlos Neto
de Carvalho (Idanha-a-Nova), Renata G. Netto (São Leopoldo), Christian Neumann (Berlin),
Jan Kresten Nielsen (Oslo), Eduardo B. Olivero (Ushuaia), Ørjan Berge Øygard (Bergen),
S. George Pemberton (Edmonton), John E. Pollard (Manchester), Lars Rennan (Trondheim),
Andrew K. Rindsberg (Livingston), Francisco J. Rodríguez-Tovar (Granada), Jennifer J. Scott
(Calgary), Koji Seike (Tokyo), Adolf Seilacher (Tübingen, deceased), Andrew M. Taylor
(Northwich), Roger T.K. Thomas (Lancaster), Alfred Uchman (Krakow), Lothar Vallon
(Faxe), Michał Warchoł (Bergen), Andreas Wetzel (Basel), Max Wisshak (Wilhelmshaven),
Beate Witzel (Berlin) and Lijun Zhang (Jiaozuo).
Several ideas originated during the International Congresses on Ichnology and International
Ichnofabric Workshops and also during in-house core workshops, special projects, and
teaching trace-fossil analysis. Statoil ASA, in particular Sture Leiknes (Bergen), Frode
Hadler-Jacobsen (Trondheim), Kjell Sunde (Bergen), and Ole Jacob Martinsen (Bergen), is
thanked for providing me with the opportunity to study trace fossils in so many cores and to
enable the publication of selected parts of this knowledge. This book has greatly benefited
from the thorough reviews provided by Andrew K. Rindsberg (Livingston) and Andreas
Wetzel (Basel), whose timely suggestions are much appreciated.
Stavanger, Norway Dirk Knaust

August 2016
Contents
1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Ichnological Basics, Principles and Concepts . . . . . . . . . . . . . . . . . . . . . . . . 5

2.1 Terminology and Definitions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2.2 Some Principles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
3 Applications of Trace-Fossil Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

3.1 Facies Interpretation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.2 Stratigraphy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
3.3 Reservoir Quality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
4 Methodology in Ichnological Core Logging . . . . . . . . . . . . . . . . . . . . . . . . . 21

4.1 Identification of Bounding Surfaces and Quantification of Bioturbation. . . 21
4.2 Identification and Documentation of Key Trace Fossils . . . . . . . . . . . . . . 22
4.3 Analysis of Burrow Size and Tiering Patterns . . . . . . . . . . . . . . . . . . . . 22
4.4 Quantification of Ichnodiversity and Ichnoabundance . . . . . . . . . . . . . . . 24
4.5 Advanced Techniques and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
4.6 Neoichnological Approach and Analog Studies . . . . . . . . . . . . . . . . . . . 26
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
5 Selected Trace Fossils in Core and Outcrop. . . . . . . . . . . . . . . . . . . . . . . . . 27

5.1 Classification of Burrows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
5.2 Arenicolites Salter, 1857 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
5.3 Artichnus Zhang et al., 2008 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
5.4 Asterosoma von Otto, 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
5.5 Bergaueria Prantl, 1946 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
5.6 Bornichnus Bromley and Uchman, 2003 . . . . . . . . . . . . . . . . . . . . . . . . 44
5.7 Camborygma Hasiotis and Mitchell, 1993 . . . . . . . . . . . . . . . . . . . . . . . 48
5.8 Chondrites von Sternberg, 1833 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
5.9 Conichnus Männil, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
5.10 Cylindrichnus Toots in Howard, 1966. . . . . . . . . . . . . . . . . . . . . . . . . . 65
5.11 Diplocraterion Torell, 1870 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
5.12 Hillichnus Bromley et al., 2003 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
5.13 Lingulichnus Hakes, 1976 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
5.14 Macaronichnus Clifton and Thompson, 1978. . . . . . . . . . . . . . . . . . . . . 85
5.15 Nereites MacLeay in Murchison, 1839 . . . . . . . . . . . . . . . . . . . . . . . . . 90
5.16 Ophiomorpha Lundgren, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
5.17 Palaeophycus Hall, 1847. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
5.18 Paradictyodora Olivero et al., 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
5.19 Parahaentzschelinia Chamberlain, 1971 . . . . . . . . . . . . . . . . . . . . . . . . 106
xiii
xiv Contents
5.20 Phoebichnus Bromley and Asgaard, 1972 . . . . . . . . . . . . . . . . . . . . . . . 109

5.21 Phycosiphon Fischer-Ooster, 1858 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
5.22 Planolites Nicholson, 1873 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
5.23 Rhizocorallium Zenker, 1836 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
5.24 Rosselia Dahmer, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
5.25 Schaubcylindrichnus Frey and Howard, 1981. . . . . . . . . . . . . . . . . . . . . 129
5.26 Scolicia de Quatrefages, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
5.27 Scoyenia White, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
5.28 Siphonichnus Stanistreet et al., 1980 . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
5.29 Skolithos Haldeman, 1840 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
5.30 Taenidium Heer, 1877 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
5.31 Teichichnus Seilacher, 1955. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
5.32 Thalassinoides Ehrenberg, 1944 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
5.33 Tisoa de Serres, 1840 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
5.34 Trichichnus Frey, 1970b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165
5.35 Virgaichnus Knaust, 2010a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
5.36 Zoophycos Massalongo, 1855 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172
5.37 Diffuse Bioturbate Texture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
5.38 Plant Roots and Their Traces. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179
5.39 Borings. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
5.40 Pseudo-trace Fossils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
About the Author
Dirk Knaust is Specialist in Sedimentology in Statoil’s Research and Technology unit in

Stavanger, Norway. After working as a geologist in an underground mine and graduating in
Germany with a Ph.D. in Geology (Carbonate Sedimentology, Stratigraphy, and Paleontology
in the Triassic), in 1997 he followed an offer from the Norwegian oil and gas industry working
in Exploration and Field Development. Since then, he has been frequently exposed to a vast
amount of well core for investigation. The author has studied various aspects of trace fossils
and ichnology, which is documented in about 60 publications in scientific journals and edited
volumes. He is co-editor of the book Trace Fossils as Indicators of Sedimentary Environments
(Elsevier) and associate editor of the journal Ichnos, amongst others. Dirk is leading author of
a revised version of the Trace Fossils volume to be published in the Treatise of Invertebrate
Paleontology.
The author inspecting dinosaur footprints on a Jurassic carbonate mud flat in Portugal
xv
Introduction
1
Driven by the need of a successful hydrocarbon exploration shoreline, shelf, slope or deep sea; e.g. Knaust and Bromley
and exploitation, the value of ichnology has been recognized 2012) and very well suit cases where data coverage is scant,
since an early phase of the oil and gas industry. It is therefore such as new exploration areas. The ichnofacies approach is
no wonder that trace-fossil analysis of well core samples and then complemented by ichnofabric analysis, a concept based
relevant outcrop analogs has played a key role in the geo- on the detailed interweaves between sedimentary features
logical interpretation of subsurface data. Trace fossils are and ichnological signals (Ekdale et al. 2012). It therefore
increasingly regarded as important for different kinds of becomes important in the meticulous description and inter-
sedimentological interpretations, and a wealth of publica- pretation of bioturbated sediments and sedimentary rocks,
tions has been dedicated to this subject as summarized in a particularly in core.
number of volumes (e.g. Crimes and Harper 1970, 1977; Extensive scientific drilling began with the Deep Sea
Frey 1975; Basan 1978; Ekdale et al. 1984; Curran 1985; Drilling Project (DSDP, operated from 1968 to 1983) and its
Bromley 1990, 1996; Maples and West 1992; Pemberton successors, the Ocean Drilling Program (ODP, operated
1992; Donovan 1994; Pemberton et al. 2001; McIlroy 2004, from 1985 to 2004), and the Integrated Ocean Drilling
2015; Seilacher 2007; Miller 2007; Bromley et al. 2007; Program (IODP, operated from 2003). These programs have
MacEachern et al. 2007; Avanzini and Petti 2008; Hasiotis provided another important impulse and comprehensive
2010; Buatois and Mángano 2011; Knaust and Bromley dataset for ichnological analysis based on core. Several
2012). hundred kilometers of core were obtained through these
From the beginning of industrial core sampling, for campaigns and resulted in dozens of reports with ichnolog-
instance in the exploration for coal in Germany, trace fossils ical information (e.g. Warme et al. 1973; Chamberlain 1975;
were commonly regarded together with other fossils from a Ekdale 1977, 1980; Fütterer 1984; Wetzel 1987). Related to
biostratigraphical perspective (e.g. Gothan 1932, Jessen this exceptional dataset, Chamberlain (1978) delivered an
1950; Fiebig 1956). About the same time, the boxcore overview of trace fossils in core, still building the funda-
sampling method was developed for shallow-marine mentals for modern trace-fossil analysis in core.
deposits of the German North Sea by researchers from the Building upon the eminent work done by Chamberlain
Senckenberg Institute (e.g. Schäfer 1952, 1956; Reineck (1978), the study of trace fossils in core soon emerged as a
1958, 1963), which enabled the observation of burrows special field in the oil and gas industry. As an example, the
along with their producers. This successful approach was Norwegian oil and gas company Statoil has a long history of
later applied to the coastal systems (Sapelo Island, Georgia; trace-fossil analysis based on core. Comprehensive well core
e.g. Howard and Dörjes 1972; Howard and Frey 1973, 1985) material from the giant gas and oil field Troll, offshore
and elsewhere (e.g. Howard and Reineck 1981; Howard and Norway, provided enough material for completing probably
Scott 1983), which was closely tied to research in similar the first extensive trace-fossil atlas by Bockelie and Howard
paleoenvironments in the U.S. Western Interior Basin by (1984), which at that time was prepared for the Research
Weimer and Hoyt (1964), then by Howard and Frey (1973). Centre in Norsk Hydro. This volume gives a good intro-
Several sources of data, including from core, facilitated duction to the study of trace fossils in core and treats 35
the development of the seminal ichnofacies concept by ichnotaxa with respect to morphology and paleoenviron-
Seilacher (1967). Ichnofacies interpretations are generally ment, nicely illustrated with numerous line drawings and
done on a broader scale with wide facies belts (e.g. core photographs. For more than three decades, this volume
© Springer International Publishing AG 2017 1

D. Knaust, Atlas of Trace Fossils in Well Core,
DOI 10.1007/978-3-319-49837-9_1
2 1 Introduction
became an in-house standard and has been the basis for For all these reasons, identification of trace fossils with
many core descriptions, although only a small fraction of confidence has become more crucial than ever but also faces
this study has reached the public domain (Bockelie 1991, old challenges. Previous compilations of trace fossils in core
1994). In the 1990s, Frederic Bockelie and later also Jean have addressed these issues and presented a solid overview
Gérard from Elf (now Total) used to work together with of the commonest forms (e.g. Chamberlain 1975, 1978;
Richard Bromley on Jurassic core material from the North Bockelie and Howard 1984; Taylor 1995; Pemberton et al.
Sea, which influenced the contents of Bromley’s (1990) 2001; Gerard and Bromley 2008). From a current perspec-
Trace Fossils, and was subsequently published in an tive, however, beside easily recognized burrows are many
ichnofabric atlas (Gerard and Bromley 2008). Similar work other forms out of several hundred burrow ichnogenera
was done in the UK sector of the North Sea by Roland (Knaust 2012), which of course also occur in cored rock
Goldring, John Pollard and their students (Duncan McIlroy, samples. In addition to the more common ichnogenera
Andrew Taylor, Stuart Gowland, and Stuart Buck). In 1991, (many of which are not covered by the outdated Treatise on
the First International Ichnofabric Workshop, based on core, Invertebrate Paleontology volume on Trace Fossils;
was launched in Bergen and Oslo, an ongoing series of Häntzschel 1975), the rare forms may bear important
workshops organized every two years around the world information and aid in the reconstruction of paleoenviron-
(Ekdale et al. 2012). mental conditions.
Since the 1980s, modern ichnology has entered the oil
and gas business as an integrated part of core descriptions
and facies interpretations in exploration and production. In
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Bromley RG, Buatois LA, Mángano G et al (eds) (2007) Sediment—
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Cambridge, 347 pp
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fold but three main uses may be highlighted: (1) Historically, J Paleontol 49:1074–1096
probably the most relevant application is the utilization of Chamberlain CK (1978) Recognition of trace fossils in cores. In:
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identification and characterization of key bounding surfaces Issue 9, 351 pp
Cunningham KJ, Sukop MC (2011) Multiple technologies applied to
in the stratigraphical column and their use in sequence-
characterization of the porosity and permeability of the Biscayne
stratigraphical subdivisions and paleoenvironmental recon- Aquifer, Florida. USGS, Open-File Report 2011–1037, 8 pp
structions. (3) Currently evolving is the understanding of the Cunningham KJ, Sukop MC (2012) Megaporosity and permeability of
impact of trace fossils and bioturbate texture on rock proper- Thalassinoides-dominated ichnofabrics in the Cretaceous
karst-carbonate Edwards-trinity aquifer system, Texas. U.S. Geo-
ties (such as porosity and permeability) and fluid behavior
logical Survey, Open-file report 2012–1021, 4 pp
(connectivity), similar to the related and already established Cunningham KJ, Sukop MC, Huang H et al (2009) Prominence of
knowledge of bioturbation and sediment properties in bio- ichnologically influenced macroporosity in the karst Biscayne
logical and ecological science. aquifer: stratiform “super-K” zones. GSA Bull 121:164–186
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sedimentary environments. Developments in Sedimentology, vol Deutschen Geologischen Gesellschaft 101:23–43
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Ekdale AA, Bromley RG, Pemberton SG (1984) Ichnology: the use of ichnology. SEPM Short Course Notes, vol 52, 380 pp
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Ichnological Basics, Principles and Concepts
2
2.1 Terminology and Definitions There are many other categories of traces with less
importance for the purpose of this book, of which coprolites
Ichnology is the study of traces produced by organisms (i.e. fossil feces) are probably the most important. An
(animals, plants and microbes) on or within a substrate. It overview of accepted groups of traces is given by Bertling
deals with all aspects related to modern (neoichnology) and et al. (2006).
fossil traces (paleoichnology), bioturbation and bioerosion, Bioturbation is the process by which the primary struc-
and is interdisciplinary in combining sedimentological, ture and properties of a sediment are modified by the activity
paleontological, biological and ecological methods (Bromley of organisms living within it, which may result in sediment
1996). It complements and constrains sedimentological mixing (Bromley 1996). The latter expression is often
interpretations and serves as a powerful tool in reservoir loosely applied to the product of this process, which is better
characterization. defined by the term bioturbate texture (Frey 1973). Bioero-
The subjects of paleoichnology are trace fossils (also sion, in contrast, comprises processes of mechanical or
called ichnofossils), which are fossilized structures produced biochemical destruction of hard substrates by organisms.
in substrates ranging from unlithified sediment to sedimen- From a sedimentological point of view, bioturbation, bio-
tary rock or organic matter (including shell, bone, wood and erosion, biodeposition and biostratification structures can be
peat) by the activity of organisms. Traces of organisms can grouped together as biogenic sedimentary structures (Frey
be grouped into categories, depending on the type of sub- 1973).
strate and manner of origin (Fig. 2.1): Ichnofacies as concept was established by Seilacher
(1967) based on his and others, earlier work. Trace-fossil
• Burrows: Most common trace-fossil category, compris- communities (ichnocoenoses) were linked to an overall
ing galleries, tunnels, shafts, chambers, etc., excavated ocean profile, mainly related to the behavioral response of
by animals within an unconsolidated substrate. the tracemakers to a bathymetric gradient in food supply.
• Bioerosion trace fossils: If the excavation takes place in a Ichnofacies represents a powerful tool when working on a
consolidated and lithified substrate, the resulting trace is larger scale (e.g. basin scale) and screening new areas, where
a bioerosional trace fossil such as a boring or a scratch. a rough interpretation of the paleoenvironment in terms
• Trails: Trails are surface features, in which the producer of broad facies belts can be given. The ichnofacies concept
leaves a continuous path behind it while moving. has been continuously updated, refined and extended into
• Trackways: In contrast to trails, trackways are discon- the continental realm. Current overviews and discussions are
tinuous paths which originate from walking animals. provided by Buatois and Mángano (2011), MacEachern et al.
Individual imprints of the trackway are called tracks. (2012) and Melchor et al. (2012, for continental ichnofacies).
• Plant-root traces: Most traces are related to the activity The ichnofabric concept regards all aspects of the texture
of animals, although plants can also leave their traces by and internal structure of a sediment that result from biotur-
means of their roots. bation at all scales (Ekdale and Bromley 1983, 1991; Bromley

DOI 10.1007/978-3-319-49837-9_2
6 2 Ichnological Basics, Principles and Concepts
Burrow Trail Trackway
Imprint Boring Fecal pellet (coprolite)
Fig. 2.1 Major categories of traces that may become trace fossils if fossilized. Bedding-surface views except for burrow (upper left), which is
vertical section. Imprint image courtesy of H. Allen Curran (Northampton)
and Ekdale 1986). It has been developed on sectioned rock resting, scraping hard substrate (such as wood) while
faces, where different cross-cutting relations can be related to feeding, building chambers while breeding, and leaving
successive colonization or tiering, and changing degree of their excrement in form of fecal pellets. Other examples
bioturbation can be analyzed. Compared with the ichnofacies include many species of crustaceans and molluscs, able
concept, which puts emphasis on the recognition of recurrent to produce different traces and burrows with contrasting
ichnocoenoses and facies belts, the purpose of the ichnofabric characteristics (Fig. 2.2, see also Fig. 5.85).
concept is mainly the analysis of different stages contained in • Many different organisms can produce the same trace:
a particular piece of bioturbated rock. Therefore it is a valu- Simple vertical shafts without branching (e.g. Skolithos)
able tool in the detailed interpretation of rock samples from would be a good example, because they can be produced by
core (Taylor et al. 2003; Ekdale et al. 2012). many different organisms such as priapulids, holothurians,
polychaetes, phoronids, crustaceans, anthozoans, insects,
spiders and even plant roots (Fig. 2.3).
2.2 Some Principles • The tracemaker is rarely known: Particularly true for
many trace fossils, the tracemaker is only preserved
The study of trace fossils is related to various challenges of under rare circumstances (e.g. exceptionally preserved
which the following are highlighted. fossils or fossillagerstätten, Fig. 2.4; see also Figs. 5.103
and 5.133). In most cases, the producer can be inferred at
• One type of organism can produce many different traces: a higher taxonomic rank with some uncertainty, for
For example, given a particular insect which is able instance by analyzing particular features of the trace (e.g.
digging a burrow into the substrate, leaving a trackway architecture, scratches and fecal pellets) or reconstruction
on the surface due to locomotion, or an imprint while of its functional morphology.
2.2 Some Principles 7
Trail Burrow
Chiton
(Polyplacophora)
Bioerosion Fecal pellets

Fig. 2.2 Various traces produced by chiton (Polyplacophora)
Holothurians
Priapulids Polychaetes
Skolithos verticalis
Phoronids Crustaceans
Anthozoans Aplacophorans
Arachnids Plant roots
Insects
Fig. 2.3 Skolithos, a very simple trace, can potentially be produced by a wide range of organisms of different phyla and environments
(a) (b)
Fig. 2.4 Examples of trace fossils (mainly trails and shallow burrows) see Knaust (2007, 2010, 2015). a Bedding surface with many trails and
that preserve their producers at the termination of the trace. The very burrows, most of which preserve their producers at the termination in
fine-grained (micritic) sediment and favored taphonomic and diagenetic form of weathered sulfide aggregates (e.g. arthropods, nemerteans,
circumstances (e.g. microbial growth, lowered oxygenation) prevented nematodes) or calcite crystals (e.g. involutinidae foraminifers, turbel-
a total loss of the organic material and promoted exceptional fossil larian platyhelminthes). b Undulating bedding surface with pustules
preservation, which allows a determination of higher taxonomic due to microbial modification with a trail occurring together with its
categories. Middle Triassic (Anisian-Ladinian) Meissner Formation supposed nemertean (ribbon worm) producer which is preserved as
(Muschelkalk), Thuringia, Germany. Scale bars = 1 mm. For details limonite aggregate. Note the slightly sinuous fecal string
2.2 Some Principles 9
Rhizocorallium commune Rhizocorallium jenense
softground firmground
deposit-feeding suspension-feeding
Cruziana Ichnofacies Glossifungites Ichnofacies
(a) (b)
Fig. 2.5 The only two valid ichnospecies of Rhizocorallium, with pouch-shaped and inclined burrow with passive fill and a dense pattern
different morphological features mainly due to contrasting substrate of scratches on the margin of burrow. After Knaust (2013), republished
conditions. Scale bars = 1 cm. a R. commune produced in softground, a with permission of Elsevier; permission conveyed through Copyright
wide horizontal burrow with actively created spreite, fecal pellets and Clearance Center, Inc. See also Fig. 5.117
occasional branching. b R. jenense produced in firmground, a narrow
• The same trace becomes preserved differently in various Spongeliomorpha (Fig. 5.85). Composite trace fossils
substrates: Different categories of substrate (e.g. soft- originate from the interpenetration of ichnotaxonomically
ground, firmground and hardground) preserve the traces different parts, which can be identified as such by their
of the same producer in different ways. The ichnogenus cross-cutting relationship. Complex trace fossils are
Rhizocorallium is only one example (Fig. 2.5), where a morphologically complex structures, including com-
probable polychaete makes extensive horizontal spreite pound trace fossils, which are characterized by their high
burrows with occasional branching and active fill degree of organization, for instance Zoophycos and
(R. commune, probably resulting from deposit feeding), Hillichnus.
while in firmground the burrows are shorter and inclined, • Multiple colonization phases and surfaces, tiers and
unbranched and open or passively filled (R. jenense, cross-cutting relationship lead to complex ichnofabrics:
probably resulting from suspension feeding). Traces are rarely single, and interaction among different
• Compound, composite and complex trace fossils: generations of traces with contrasting features is the norm
Trace-fossil architecture can be complicated by the (Fig. 2.6). This can result in partly or completely bio-
interaction of different tracemakers, or producers with turbated substrate, which may preserve discrete traces on
contrasting behavior. Compound trace fossils are those top of a diffuse (bioturbated) background. Interaction of
comprising intergradational forms of ichnotaxonomically benthic communities may also lead to reburrowing of
different parts, such as Thalassinoides-Ophiomorpha- existing burrows by subsequent producers.
(a)
(b) (c)
M
References 11
b Fig. 2.6 Ichnofabrics with multiple colonization surfaces in outcrop it due to repeated bioturbation. The mudstone layer above it contains
and in core. Scale bars = 5 cm (a) and 1 cm (b, c). a Outcrop large burrows actively filled with sand (Thalassinoides and Ophiomor-
photograph of a limestone bedding plane showing hardground features pha) as part of deep-tier burrowing through the overlying sand
with dense occurrence of Gastrochaenolites isp., produced by boring (turbidite). Upper Cretaceous (Maastrichtian) Springar Formation
bivalves (in places preserved) in a shallow-marine environment. (deepmarine), Norwegian Sea (well 6604/10-1, ca. 3647.5 m).
A second colonization of the same surface is documented by a network c Ripple-laminated fine-grained sandstone with intervals containing
of calichified root traces, which belong to the overlying eolian Macaronichnus segregatis (M) in vertical core section. The displayed
sandstone. This surface is a regional angular unconformity between sandstone shows multiple colonization surfaces (arrow heads) from
Cretaceous platform carbonates and Pliocene to Pleistocene eolian dune which the muddy spreite burrows Teichichnus zigzag penetrate the
deposits (brownish patches of sand). Cliff section south of Taghazout, underlying sediment, indicating rapid deposition (sandy tidal-flat
western Morocco. b Parts of two sandy turbidite layers interbedded deposit). The succession is interrupted by a medium-grained sandstone
with hemipelagic mudstone in vertical core section. The top surface of layer in its lower part, probably a storm deposit (tempestite). Middle
the lower turbidite served as a colonization surface (arrow head), which Jurassic (Bathonian) Tarbert Formation (sandy tidal flat), Oseberg Sør
resulted in a mixed layer with incorporated green clay minerals beneath Field, Norwegian North Sea (well 30/9-F-26, ca. 4466.8 m)
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Frey RW (1973) Concepts in the study of biogenic sedimentary Taylor A, Goldring R, Gowland S (2003) Analysis and application of
structures. J Sediment Petrol 43:6–19 ichnofabrics. Earth Sci Rev 60:227–259
Applications of Trace-Fossil Analysis
3
With respect to reservoir prediction and characterization, other methods, especially sedimentology (Fig. 3.1). For
ichnology can contribute with three major themes, which are instance, in cases where a new basin is entered or a new
facies interpretation, stratigraphy, and reservoir quality. petroleum play is explored, only a limited set of data is
typically available, of which trace-fossil analysis can be used
to delineate the overall setting (such as deep marine versus
3.1 Facies Interpretation shelf). Smaller areas with better data coverage can be dif-
ferentiated with respect to their paleoenvironments and
The interpretation of facies and depositional environments is summarized in paleogeographical maps. Great value can be
probably the best recognized application for trace fossils and gained by the identification of limiting factors such as
bioturbate textures. Depending on available data, the size of oxygen depletion, salinity fluctuation and reduction, light
the study area and subject to be investigated, the ichnofacies availability, hydraulic energy, substrate consistency, avail-
and ichnofabric concepts can be utilized in combination with ability of organic matter and so on.
(a) Ichnofacies (Seilacher, 1967) (b) Ichnofabric (Bromley and Ekdale, 1986)
Fig. 3.1 Ichnofacies (a) and ichnofabric concept (b), two brilliant, ichnofabric concept integrates various ichnological information of a
often used, and still debated approaches in the utilization of sectioned unit and thus provides a wide range of information.
ichnological information. The ichnofacies concept was originally Reprinted from Seilacher (1967) (a), republished with permission of
designed for trace fossils outlining broad facies belts and thus becomes Elsevier, permission conveyed through Copyright Clearance Center,
applicable in areas where relative little data is available, whereas the Inc.; and Bromley (1990) (b), reprinted with permission of Springer

DOI 10.1007/978-3-319-49837-9_3
14 3 Applications of Trace-Fossil Analysis
3.2 Stratigraphy role in the development of reservoir quality, including rock

composition, sedimentary structures, cementation and dis-
In general, trace fossils are poor stratigraphical markers. solution, and fracturing (Fig. 3.3). In addition, bioturbation
However, some periods (e.g. in the lower Paleozoic) have (organism/sediment interactions) and the resulting textures
been successfully dated by means of trace fossils (e.g. ich- and trace fossils has proved to be an important agent in the
nospecies of Cruziana). This procedure might be helpful in postdepositional modification of sediments (Fig. 3.4).
exploration activities of large basins with relatively few data The process of bioturbation results in the origin of highly
as demonstrated in North Africa. More common is the use of variable textures and structures, which may increase or
trace fossils and bioturbate textures in sequence stratigraphy, decrease the rock’s porosity and thus reservoir quality
where particular intervals and key stratigraphical surfaces (Fig. 3.5). In many cases, intense bioturbation leads to the
can be identified and correlated (Fig. 3.2). In addition to homogenization of alternating sediments or even an increase
many other methods, ichnological analysis is important in of mud due to the introduction of clay minerals, which under
constraining sequence-stratigraphical interpretations and to certain circumstances may help protecting the existing
solve a wide range of problems in the following areas: porosity. More often, however, biogenically introduced mud
reduces the porosity and diminishes the reservoir quality.
• Delineation of sequence-stratigraphic surfaces by means The reorganized rock fabric and discrete trace fossils can act
of their ichnological signal, which can be studied in as preferred fluid paths for soluble mineral phases and pro-
outcrop and in subsurface datasets (e.g. cores and bore- mote early diagenetic cementation leading to reduced
hole images). porosity. In contrast, open and sand-filled shafts and gal-
• Facies reconstruction and characterization of juxtaposed leries of burrow systems can improve porosity significantly
facies types to recognize changes in deposition and and to turn a fluid barrier into a reservoir. Finally, bioturbate
deviations from continuous facies transitions (e.g. Wal- textures and trace fossils are an important constituent of
ther’s Law). mudstone and shale and may lead to an increased hetero-
• Vertical facies trends, reflecting shallowing- or geneity due to the incorporation of silt and sand. This latter
deepening-upward in response to autocyclic or allocyclic process plays an important role in the producibility and
processes, often recognizable by changing degree of performance of shale gas reservoirs.
bioturbation and trace-fossil content. (This is particularly A wide range of infaunal organisms with contrasting
crucial in relatively uniform lithologies such as in car- burrowing styles is known and some create complex bio-
bonate systems.) turbate textures and ichnofabrics. Some organisms (e.g.
• General aspects, such as degree of bioturbation, crustaceans) process the sediment and destroy the primary
trace-fossil associations, ichnodiversity etc. in comparison bedding, which leads to a uniform fabric with an isotropic
to one another, which can contribute to the delineation of character. Other animals (e.g. some kinds of polychaetes) are
particular systems tracts. more selective and repeatedly rework the sediment whilst
they selectively feed on the deposit. The result is a more
anisotropic rock fabric with connectivity preferred in a
particular direction (Fig. 3.6). More advanced architects
3.3 Reservoir Quality (e.g. some worms, bivalves and shrimp) are responsible for
rather complex burrow systems with three-dimensional
The flow behavior within a reservoir is dependent on its branching, wall lining and active fill. Consequently, these
reservoir quality, which in turn is a function of porosity, bioturbators import a high degree of heterogeneity to the
permeability and connectivity. A number of factors play a sediment.
3.3 Reservoir Quality 15
Subaerial Maximum regressive Ravinement Regressive surface of Maximum flooding

unconformity (SU) surface (MRS) surface (RS) marine erosion (RSME) surface (MFS)
Fluvial to Carbonate shoal Shoreface Marginal-marine

marginal-marine
Offshore
Sandy tidal flat

Lower shoreface
The surface near the top of
the image separates A laminated siltstone with
bioturbated sandstone of a sparse Phycosiphon is
sandy tidal flat from sharply eroded at its top
deeper-marine deposits and overlain by coarse-
Supratidal flat above it, and is grained sandstone with rip-
demarcated by the up clasts. This erosion Interval of totally
occurrence of a sideritic surface occurs regionally bioturbated, very fine-
spreiten burrow and is associated with grained sandstone
Diplocraterion and deeply penetrating, sand- comprising Phycosiphon,
numerous reworked quartz filled, uncompacted Cylindrichnus, Planolites,
pebbles (lag deposit) partly burrows indicative of a Schaubcylindrichnus and
Alluvial plain incorporated into the
burrow and the adjacent
firmground omission Siphonichnus. Skarv Field,
surface (Glossifungites Norwegian Sea (well
host rock due to surface). Lower Jurassic 6507/5-1, ca. 3538.6 m).
Boundary between Hegre A dense root system bioturbation. Middle Cook Formation,
Group (Upper Triassic) and (probably resulting from Jurassic Hugin Formation, Norwegian North Sea (well
Statfjord Group (Lower Cordaitales) is deeply Sleipner Field, Norwegian 35/10-1, ca. 3655.8 m).
Jurassic). The Hegre penetrating from an North Sea (well 15/9-1, ca.
Group in the lower part of erosive, blackened surface 3660.35 m).
the image consists of a and is filled with organic-
mixed siliciclastic- rich sediment, dolomite,
carbonate rock with caliche and gypsum. It is sharply
features, fractures and overlain by an oolitic
complex burrow systems in limestone. Upper Permian
the firmground, as well as carbonate platform of the
Taenidium (not figured). It Khuff Formation, South
results from pedogenesis Pars Field, offshore Iran
in an alluvial environment. (well SP10, 2937.5 m).
The overlaying coarse-
grained sandstone of the
Statfjord Group is weakly
bedded, contains a basal
lag deposit with polymict
clasts and granulae, and is
without bioturbate textures.
It results from fluvial
processes. Johan
Sverdrup Field, Norwegian
North Sea (well 16/2-15,
ca. 1972.7 m).
Fig. 3.2 Interpreted core examples of key stratigraphical surfaces Catuneanu (2013, upper part, republished with permission of Elsevier;
(indicated by arrows, middle and lower parts) and their relationship permission conveyed through Copyright Clearance Center, Inc.). Width
within the sequence-stratigraphic framework provided by Zesshin and of core images is approximately 10 cm
Fig. 3.3 Important processes 1. Sedimentary 2. Ichnological

and their products, which result in
small-scale heterogeneities of
reservoirs in sedimentary rocks.
During the evolution of a
reservoir, each process in
isolation or combination with
others may have an impact on
reservoir quality. Modified after
Knaust (2013)
- Lithological composition - Trace fossils

- Sedimentary structures - Bioturbate texture
3. Diagenetic 4. Structural
- Cementation and dissolution - Fractures

- Grain coating - Faults
(a) (b)
<5% bioturbation 70% bioturbation Ophiomorpha rudis
23,8% / 6,69 mD 20,9% / 1,29 mD
5 cm 1 cm
(c) (e)
(d)
Well 6604/10-1
100.000
10.000
Average: 23,9; 4,532

Permeability [mD]
1.000
Average: 18,2; 0,492
0.100
0.010
0.0 5.0 10.0 15.0 20.0 25.0 30.0
Porosity [%]
Sandstone (no bioturbation) Sandstone (bioturbated)

b Fig. 3.4 Impact of bioturbation on gas-reservoir quality based on permission of GEUS. d Permeability and porosity cross plot showing the
examples from Upper Cretaceous deep-marine turbidite deposits of the properties of bioturbated versus unbioturbated sandstone, as shown in
Vøring Basin, Norwegian Sea. a Core examples of almost unbioturbated (a) (based on horizontal core plug measurements). The bioturbated
versus intensely bioturbated sandstone and their corresponding porosity sandstone is clearly reduced in porosity and permeability, the latter of
and permeability measurements (based on horizontal core plug data), which is about one order of magnitude lower than measured for
which indicate reduced properties for bioturbated sandstone. Springar unbioturbated sandstone. e Thick unit of turbiditic sandstone with
Formation (Maastrichtian, well 6604/10-1). b Sandstone in thin section upward-increasing bioturbation (Ophiomorpha rudis) towards a sharp
with homogeneous texture and incorporated clay minerals, some of bedding surface at its top (ca. 4476.45 m), which goes hand in hand with
which outline the wall of a discrete Ophiomorpha rudis burrow. Same a reduction of porosity and permeability due to incorporation of mud.
well as in (a). c Callianassid shrimp in its lined burrow similar to The overlying sand unit is unbioturbated and has reasonably high
Ophiomorpha. From Bromley and Asgaard (1972), republished with properties. Kvitnos Formation (Santonian, well 6707/10-2A)
Fig. 3.5 Architectural elements

and characteristics of burrows,
Orientation Branching
which may have an impact on
reservoir quality, such as burrow
orientation with respect to
bedding (e.g. horizontal, vertical,
oblique), branching (e.g.
unbranched, branched,
dichotomous, network, boxwork),
presence or absence of lining or
mantle, and burrow fill (e.g.
active, passive) with respect to
host rock (e.g. sand, mud).
Modified after Knaust (2013)
Lining/mantle Fill
(a)
(b)
Fig. 3.6 Schematic illustration presenting stages of the fluid flow molecules travel to the fracture network or permeable flow path
within bioturbated, gas-charged tight mudstone. From Bednarz and through diffusion. b Millimetric scale. Efficiency of the flux of the
McIlroy (2015), republished with permission of Elsevier; permission diffusively migrating molecules into the fracture network is dependent
conveyed through Copyright Clearance Center, Inc. a Micro-scale. As on the distance from oil- or gas-charged pore to the closest fracture or
pressure drops, hydrocarbon molecules enter the organo-porosity permeable flow-path. Dense ichnofabric network minimises the
through desorption from pore walls and kerogen material. If the pore distance of the diffusive flow through partitioning the hosting rock
is connected to the (micro-) fracture or micro-channel, molecules travel with permeable and brittle silty tubes and may also improve fracture
through the conductive flow paths to the well bore. If there is no spacing and/or complexity
fracture or micro-channel connected to the organo-porosity, the
References East Greenland. Grønlands Geologiske Undersøgelse, Rapport 49:

7–13
Bromley RG, Ekdale AA (1986) Composite ichnofabrics and tiering of
Bednarz M, McIlroy D (2015) Organism–sediment interactions in shale- burrows. Geol Mag 123:59–65
hydrocarbon reservoir facies—Three-dimensional reconstruction of Knaust D (2013) Bioturbation and reservoir quality: towards a genetic
complex ichnofabric geometries and pore-networks. Int J Coal Geol approach. AAPG Search and Discovery Article #50900
150–151:238–251 Seilacher A (1967) Bathymetry of trace fossils. Mar Geol 5:413–428
Bromley RG (1990) Trace fossils: biology and taphonomy. Unwin Zesshin M, Catuneanu O (2013) High-resolution sequence stratigraphy
Hyman, London, 293 pp of clastic shelves I: units and bounding surfaces. Mar Pet Geol
Bromley RG, Asgaard U (1972) Notes on Greenland trace fossils, 1. 39:1–25
Freshwater Cruziana from the Upper Triassic of Jameson Land,
Methodology in Ichnological Core Logging
4
Well coring is an expensive but valuable method to obtain associations below and above a particular surface, and
information about the geological situation in the subsurface, identifying firm- and hardgrounds on the basis of passively
and a versatile analysis including ichnological study may filled burrows (Glossifungites and Trypanites ichnofacies),
lead to a robust interpretation. The amount of investigation are just two examples that help to subdivide the inspected
and recommended workflow are dependent on the goal, the core interval into genetically related units (see Sect. 3.2).
core (e.g. preservation, length, preparation, facies, etc.), the One of the very first steps performed by the logging
task at hand (e.g. what kind of information and in which sedimentologist is the quantification of bioturbation, which
detail should be extracted), and the skills of the investigator. is an indicator of the conditions affecting the infaunal
In an optimal case, ichnological data is integrated with other community. Different scales and methods are available, such
relevant data and information with respect to regional geol- as the semiquantitative field classification of ichnofabrics
ogy, sedimentology, petrophysics and others. Based on my (Droser and Bottjer 1986), and bioturbation on bedding
own experience, the following overall steps can be performed planes (Miller and Smail 1997). Widely applied and thus
and account for different tasks but may build on each other. comparable from study to study is the bioturbation scale of
Reineck (1963; see also Taylor and Goldring 1993), which
1. Identification of bounding surfaces and quantification of ranges from grade 0 (no bioturbation) to 6 (bedding com-
bioturbation. pletely destroyed). These seven grades of bioturbation (0–6)
2. Identification and documentation of key trace fossils. contain very different values ranging from grade 0 (0%),
3. Analysis of burrow size and tiering patterns. grade 1 (1–4%), grade 2 (5–30%), grade 3 (31–60%), grade
4. Quantification of ichnodiversity and ichnoabundance. 4 (61–90%), grade 5 (91–99%) to grade 6 (100%) (Fig. 4.1).
5. Advanced techniques and methods. Because this scale was developed for the description of
6. Neoichnological approach and analog studies. modern bioturbation encountered in boxcores and is expli-
citely limited to narrow intervals of the section (e.g. the
Steps 1 and 2 are most relevant for (broad) ichnofacies decimeter range in boxcores), it is at best applicable to
analysis, whereas steps 3 and 4 have great potential in individual (small-scale) ichnofabrics which were not subject
(narrow) ichnofabric studies. Steps 5 and 6 add some to repeated bioturbation, non-depositon, erosion and com-
information for enhanced interpretation. paction. From a geological and paleoichnological perspec-
tive, however, the Reineck (1963) scale is limited in its
application and a linear distribution of the different grades of
4.1 Identification of Bounding Surfaces and bioturbation is recommended (Knaust 2012). The subdivi-
Quantification of Bioturbation sion of such a linear scale into equally distributed categories
depends on the logging scale and the length of the interval.
Deposition of sediment is a discontinuous process frequently Intervals with an increment of 10 or 20% of bioturbation
interrupted by periods of nondeposition or erosion. structures would be common (e.g. in a scale of 1:10–1:50),
These and other surfaces can be characterized with differ- whereas larger logging scales (e.g. 1:200) might well be
ent methods, wherein the ichnological content can be served by a threefold subdivision. The chief advantage of a
investigated in conjunction with other features for their linear scale is its suitability for quantitative and statistical
identification and characterization. Contrasting trace-fossil analysis of different units (e.g. Martin 1993; Knaust 2010).

DOI 10.1007/978-3-319-49837-9_4
22 4 Methodology in Ichnological Core Logging
4.2 Identification and Documentation of fractionally displayed and may be misidentified as other,
Key Trace Fossils simpler burrows. Even so, morphological details, for
instance such related to the burrows fill, wall structure, cross
A second important step is the identification and documen- section and pellets, may provide important clues for
tation of trace fossils. This challenging task often sets a trace-fossil identification. The size limit imposed by cores is
hurdle to the responsible worker and in the worst case this an important constraint when different size classes of trace
information is neglected. The outline of selected trace fossils fossils are addressed. Cores are commonly taken in a sub-
in the following section will lower this hurdle and encourage vertical direction (except in inclined or horizontal drilling
the investigator in charge of to perform this task to a certain and inclined bedding) and thus overrepresent this orienta-
degree. tion. In most cases, identification of trace fossils is restricted
Of course, the number of described and named trace to the ichnogenus level if not more, specific diagnostic
fossils is high: several hundred invertebrate ichnogenera, of features are exposed. This fact has an impact on subsequent
which only a small fraction can be recognized with confi- facies interpretation, which of course turns out to be less
dence in sectioned core, whereas the majority of traces detailed than at the ichnospecies level. And, lest we forget,
escapes the conventional core logging process due to their ichnological observations achieved from core are always
occurrence along bedding planes, size issues and featureless limited in comparison with those obtained from outcrop.
appearance. Therefore it is important to be aware of the From a practical point of view, core logs (including trace
selectivity and bias of the collected trace-fossil data before fossils) can be drawn by hand, subsequently digitized using
utilizing them in advanced interpretations. It must be also drawing software, or recorded directly in an application
stressed that the list of trace fossils may change from region specifically designed for that task (Fig. 4.2). Displaying
to region and differ within stratigraphical units. Finally, the burrows goes hand in hand with the log of the bioturbation
subjectivity and experience of the core logger and interpreter intensity (see above). While some workers prefer the com-
play major roles on the outcome of the trace fossils recorded. mon convention of displaying different symbols for each
The exposed surface area displaying burrows for inves- kind of trace fossil, others apply acronyms based on the
tigation is important, such as full core or sectioned core. If ichnotaxon’s name. Given that burrows in core are com-
sectioned, it may be worthwhile studying the counterpart as monly fragmentary and often allow for different assign-
well, as it commonly reveals a slightly different section ments, a good description and sketch should accompany the
figure and thus facilitates three-dimensional reconstruction. log entry. For more advanced studies, drawing entire ich-
Trace fossils having a complex morphology are only nofabrics on a transparent sheet directly over the core section
not only reveals the different burrows and their features, but
also their cross-cutting relationships. Finally, high-resolution
photography is a standard way of documentation, although it
Degree of bioturbation
may not be as persuade as a detailed drawing in the
6 trace-fossil analysis.
100 % 5
5
90 %
80 % 4 4.3 Analysis of Burrow Size and

4 Tiering Patterns
70 %
60 %
Amount
3 After logging key trace fossils, additional information can be

50 % 3
obtained from the core. Important information includes the
40 %
2 analysis of trace fossils and bioturbate texture in relation to
30 % key stratigraphical surfaces, the succession of colonization,
20 % 2
as well as cross-cutting relationships. The ichnofabric
1
10 % approach has become very useful in such investigations and
1
0% is widely applied in core and in outcrop (e.g. Knaust 1998;
Reineck (1963) Knaust (2012)
Fig. 4.3).
It has turned out that burrow size may depend on envi-
Fig. 4.1 Quantification of bioturbation. The widely used bioturbation
scale of Reineck (1963) with dissimilar proportions of categories ronmental factors and thus can be crucial in interpretations,
compared with a linear distribution as proposed by Knaust (2012). particularly in cases where other proxies are lacking. Any
Modified after Knaust (2012) particular trace fossil (e.g. Chondrites) can occur in a
4.3 Analysis of Burrow Size and Tiering Patterns 23
(a)
(b)
Fig. 4.2 Examples of core logs including ichnological information. a Hand drawn and subsequently redrawn core section. b Core section logged
directly in a core-logging application
(a) (b) (c)

Planolites-Nereites ichnofabric (totally bioturbated)
Grain size range Cl Si VFS FS MS CS VCS Gr Pe
% area covered 0.1 1 10 100
60 Horizontal, unbranched
and sand-filled burrows,
Planolites
up to 1 cm in diameter
Order of deposition and

10 Vertical spreiten burrows with
bioturbation (tiering)
concave-up meniscate sand-fill,
Teichichnus up to 6 cm deep and 10 cm wide
Cylindrical, unbranched and

Nereites 18 horizontal burrows with irregular
course and a dark, silty core,
0.1-0.3 cm in diameter
10 Horizontal to oblique, bow-shaped
Cylindrichnus burrows, concentrically filled with
sand and clay, partly compacted, up to
1 cm in diameter and 7 cm in length
2 Subhorizontal, gently curved burrows with a thick

Schaubcylindrichnus light-colored wall and a passive burrow fill,
unbranched, elliptical cross section (compacted),
about 0.5 cm in diameter
Fig. 4.3 Example of an ichnofabric analysis based on a core section (a), scale bar = 1 cm, from which major burrows were drawn with different
colors on an overlaid transparent sheet (b), and finally displayed in an ichnofabric constituent diagram (c)
“standard” size range, which needs to be qualified by cali- subdivide the total number of recognized trace fossils into
bration with “standard” conditions (e.g. open-marine envi- grades, commonly resulting in three to six grades of ichno-
ronment), or largely reduced in burrow diameter because of diversity. Unfortunately, the resulting numbers only
environmental influences (e.g. poor oxygenation, lowered indicate the presence of particular trace fossils but do not
salinity). Changes in size classes may be abrupt or gradual, provide information about their abundances. Therefore, the
and in each case it is important to get it documented. ichnoabundance can be calculated for particular core inter-
A relatively complete list of trace fossils in core is often vals, and is a combination of the ichnodiversity (e.g. number
beneficial for a particular project. In addition to a few key of trace fossils) and their abundance. According to the
trace fossils which are easily identified, it can be worthwhile abundance of a logged trace fossil, different grades can be
also to identify less common forms. In this way, the spec- subdivided following an exponential growth factor (for
trum is increased and comparison of trace-fossil associations details and case study, see Knaust et al. 2014; Fig. 4.4).
between different core intervals can be achieved with good
confidence.
4.5 Advanced Techniques and Methods
4.4 Quantification of Ichnodiversity and The outlined procedure of ichnological core logging can be
Ichnoabundance extended in cases where the core data is challenging or in
studies where enhanced interpretation becomes necessary.
After capturing the degree of bioturbation and identifying Various techniques and methods are available for the dif-
trace fossils, the next step could reveal information about the ferent tasks and only a few common ones are mentioned here
number and abundance of recognized trace fossils. Although (see also Taylor et al. 2003; Gingras et al. 2011).
ichnodiversity must not be confused with biodiversity, the A semiquantitative estimation of the degree of bioturba-
number of contemporaneous burrows is a good indication of tion can be achieved by either putting a transparent sheet
prevailing conditions at the sedimentary surface during the with a grid on top of the sectioned core surface or point-
time of deposition. Ichnodiversity has been frequently counting the number of hits where bioturbation is encoun-
evaluated in case studies and the conventional approach is to tered (Marenco and Bottjer 2010; Knaust 2012). Grid size
4.5 Advanced Techniques and Methods 25
(a) Ichnodiversity (b) Ichnoabundance
ID=n IA=2nVR+4nR+8nC+16nVC+32nA+64nVA
n = number of ichnotaxa
VR = very rare (n=1)
R = rare (n=2-6)
C = common (n=7-9)
VC = very common (n=10-22)
A = abundant (n=23-41)
VA = very abundant (n 42)
Fig. 4.4 Logged ichnodiversity (a) versus calculated ichnoabundance benthic organisms) multiplied with the abundance (frequency) of each
(b) of a dataset from the confined deep-marine turbidite system of the grade (ranging from very rare to very abundant). Comparison of the
Eocene Grès dʼAnnot Formation, southeastern France. The ichno- results of the two methods reveals significant trends for ichnoabun-
diversity is simply a count of ichnotaxa (richness) for each locality dance, such as an increase from a proximal to a distal position within
without regard for the abundance of each ichnotaxon (ichnogenera in the depositional system, which are hardly reflected in the ichnodiversity
this case). In contrast, the ichnoabundance is calculated by the sum of plot. Adapted from Knaust et al. (2014), republished with permission of
an exponential growth factor (reflecting the population dynamics of Wiley; permission conveyed through Copyright Clearance Center, Inc.
(a) (b) (c)
Fig. 4.5 Core example of a completely bioturbated sandstone from the Berge Øygard (Bergen). a Overall CT-scan revealing numerous discrete
Paleocene Grumantbyen Formation (shelf) near Longyearbyen, Sval- burrows within a bioturbate texture due to density contrasts. b Ortho-
bard, treated with CT-scan and processed subsequently. Core diameter gonally sliced CT-scan overlain by discrete burrow parts (in blue
ca. 6 cm. Images courtesy of Lars Rennan (Trondheim) and Ørjan color). c Segmented burrows after removing small spots
(e.g. with centimeter scale) is somewhat dependent on the interval. Another method of calculation is by means of
average size of the burrows present and the number of digital treatment in image analysis software (Dorador and
counts (e.g. 300) must be representative for the investigated Rodriguez-Tovar 2015).
(a) (b)
Fig. 4.6 Example of a neoichnological dataset for comparison. Maps of diversity (a) and sediment disruption resulting from infaunal activity (b).
a modern sandy tidal flat (delta front) of the Fraser River Delta (Boundary From Dashtgard (2011), republished with permission of Elsevier;
Bay), British Columbia, western Canada, showing the ichnological permission conveyed through Copyright Clearance Center, Inc.
Displaying trace fossils and bioturbate texture might be Droser ML, Bottjer DJ (1986) A semiquantitative field classification of
difficult under certain circumstances, e.g. low contrast in color, ichnofabric. J Sediment Petrol 56:558–559
Garton M, McIlroy D (2006) Large thin slicing: a new method for the
grain size, lithology, and so on. Ultraviolet light instead of study of fabrics in lithified sediments. J Sediment Res 76:1252–
visible light might be a good way to do that, as well as 1256
CT-Scans (Fig. 4.5). At broader scale, borehole image anal- Gingras MK, Baniak G, Gordon J et al (2012) Porosity and
ysis may reveal particular bioturbation patterns (Knaust 2012); permeability in bioturbated sediments. In: Knaust D, Bromley RG
(eds) Trace fossils as indicators of sedimentary environments.
while at a narrower scale, microfabric analysis based on thin Developments in Sedimentology, vol 64, pp 837–868
sections could show disturbances in bedding and reorientation Gingras MK, MacEachern JA, Dashtgard JE (2011) Process ichnology
of grains related to bioturbation (Garton and McIlroy 2006). and the elucidation of physico-chemical stress. Sediment Geol
Finally, dedicated minipermeameter measurements have 237:115–134
Knaust D (1998) Trace fossils and ichnofabrics on the Lower Muschel-
shown to be helpful in the evaluation of the reservoir properties kalk carbonate ramp (Triassic) of Germany: tool for high-resolution
of burrows versus matrix (Gingras et al. 2012). sequence stratigraphy. Geol Rundsch 87:21–31
Knaust D (2010) The end-Permian mass extinction and its aftermath on
an equatorial carbonate platform: insights from ichnology. Terra
Nova 22:195–202
4.6 Neoichnological Approach and Knaust D (2012) Methodology and techniques. In: Knaust D, Bromley
Analog Studies RG (eds) Trace fossils as indicators of sedimentary environments.
Developments in Sedimentology, vol 64, pp 245–271
As a final step, links between the collected data and the Knaust D, Warchoł M, Kane IA (2014) Ichnodiversity and ichnoabun-
dance: revealing depositional trends in a confined turbidite system.
subsequent interpretation can be made by comparing the Sedimentology 62:2218–2267
ichnological findings in core with neoichnological evidence Marenco KN, Bottjer DJ (2010) The intersection grid technique for
and similar cases described in literature, as well as applying quantifying the extent of bioturbation on bedding planes. Palaios
analog studies with a focus on ichnology integrated with 25:457–462
Martin AJ (1993) Semiquantitative and statistical analysis of bioturbate
sedimentology (Fig. 4.6). textures, Sequatchie Formation (Upper Ordovician), Georgia and
Tennessee, USA. Ichnos 2:117–136
Miller MF, Smail SE (1997) A semiquantitative field method for
References evaluating bioturbation on bedding planes. Palaios 12:391–396
Reineck H-E (1963) Sedimentgefüge im Bereich der südlichen
Nordsee. Abhandlungen der Senckenbergischen Naturforschenden
Dashtgard SE (2011) Neoichnology of the lower delta plain: Fraser Gesellschaft 505:1–138
River Delta, British Columbia, Canada: implications for the Taylor AM, Goldring R (1993) Description and analysis of bioturbation
ichnology of deltas. Palaeogeogr Palaeoclimatol Palaeoecol 307: and ichnofabric. J Geol Soc, London 150:141–148
98–108 Taylor A, Goldring R, Gowland S (2003) Analysis and application of
Dorador J, Rodriguez-Tovar F (2015) Ichnofabric characterization in ichnofabrics. Earth Sci Rev 60:227–259
cores: a method of digital image treatment. Ann Soc Geol Pol
85:465–471
Selected Trace Fossils in Core and Outcrop
5
This section contains many well-known trace fossils but sections through the burrows may very precisely reveal
also several poorly understood ichnotaxa, some of which morphological details such as wall structure, fill, cross sec-
have just emerged during the last 10–15 years. It attempts tion, pellets, etc. These circumstances call for a nomencla-
to give a comprehensive overview of each treated trace ture adapted for ichnotaxa identified in core, and in many
fossil in a standardized way, starting with its morphology, cases, identification is limited to the ichnogenus level, which
fill and size; appearance in outcrop; an evaluation of ichno- of course influences the precision of the paleoenvironmental
taxonomical status; and preferred substrate. This first part interpretation. It therefore becomes obvious that ichnological
is supplemented with key illustrations from outcrop, col- observations achieved from core are not fully comparable
lections and literature. The main part of each section deals with those obtained from outcrop. For instance, the trace
with the appearance of particular trace fossils in core, fossils characteristic of subenvironments of deep-sea fan
including a description and core image examples, as well as systems are well known from various outcrop studies,
comparison with similar trace fossils. This part is mainly whereas vertical core sections from such systems display a
based on material from offshore Norway, but samples from very different suite of trace fossils. This is also related to the
elsewhere have been involved as well. The last part of each fact that in distal deep-sea deposits so-called graphoglyptids
section outlines additional information, including the are preferably preserved at the base of turbidite beds, which
interpretation of the tracemaker, its behavior (ethology) are hard to study in core.
and, quite importantly, notes on the depositional environ- For hands-on identification of trace fossils in core, an
ment based on personal observations and on a review of experienced investigator may rely on his or her ability of
published evidence. It rounds up with comments on ichno- pattern recognition and in this way may be able to identify
facies, age range and impact of the trace fossil on reservoir the most common ichnotaxa. Another approach is to follow
quality. Figure 5.1 gives a summary of all ichnogenera a determination key (Fig. 5.2), in which diagnostic criteria
dealt with herein, including their abundance, number of (e.g. ichnotaxobases) are arranged in a hierarchical order and
ichnospecies, similar trace fossils, inferred producers and thus allow trace-fossil identification based on major mor-
paleoenvironment. phological features. This key follows the morphological
classification of trace fossils as proposed by Knaust (2012a)
and contains all burrows that are listed alphabetically and
5.1 Classification of Burrows further explained in detail below. In addition to these ich-
notaxa, the hierarchically higher trace-fossil categories bio-
Trace-fossil identification is not straightforward in turbate texture, plant roots and their traces, as well as
two-dimensional core slabs, and often only limited infor- pseudo-trace fossils are added subsequently. Trace-fossil
mation about the whole morphology, particularly of complex associations and their relationship to particular ichnofacies
trace fossils, can be achieved. In contrast, different planes of on a basin profil are displayed in Fig. 5.3.

DOI 10.1007/978-3-319-49837-9_5
28 5 Selected Trace Fossils in Core and Outcrop
Paleoenvironment
Acro- Number of
nyms Ichnogenera ichnospecies Similar trace fossils Inferred producers Continental Paralic Shallow marine Slope Deep marine
Art Artichnus 1 Tei Holothurians
Ast Asterosoma ±5 Cyl, Ros, Art, Euflabella Polychaetes, crustaceans, holothurians, bivalves
Cho Chondrites 4+ Pilichnus, Skolichnus, Pragichnus, Phymatoderma, Vir Polychaetes, sipunculids, bivalves
Cyl Cylindrichnus 7 Ast, Ros, Laevicyclus, Pah, root traces, Glyphichnus, Catenarichnus Polychaetes, holothurians
Mac Macaronichnus 1 Pla, Gordia, Ner Polychaetes
Ner Nereites 6+ Phy, Mac, Cho Enteropneusts
Oph Ophiomorpha 8+ Tha, Spongeliomorpha, Psilonichnus, Pholeus, Sce Thalassinidean shrimp (e.g. callianassids)
Pal Palaeophycus ±20 Pla, Mac, Oph, Sip Annelids, arthropods
Phy Phycosiphon 1 Ner, Cho Worms
Pla Planolites ±10 Pal, Mac Worms, arthropods, bivalves
Ros Rosselia 1 Pah, Ast, Cyl, Lin, Sko Polychaetes, sea anemones
Sch Schaubcylindrichnus 1 Pal Enteropneusts
Sip Siphonichnus 1 Sko, Pal, Mac, Lin, Dip, Trypanites Bivalves
Sko Skolithos ±20 Cylindricum, Capayanichnus, Tri, Sip, Tha, Oph Worms, crustaceans, sea anemones, holothurians, insects, plant roots
Tae Taenidium ±7 Sce, Ancorichnus, Zoo, Rhi Insects, arthropods, worms
Tei Teichichnus ±12 Dip, Rhi, Phycodes, Art, Pad, Scal Annelids, arthropods, bivalves, holothurians
Tha Thalassinoides ±15 Spongeliomorpha, Psilonichnus, Parmaichnus, Cam Thalassinidean shrimp (e.g. callianassids), arthropods, sea anemones, worms
Zoo Zoophycos ±6 Rhi, Spirophyton, Echinospira, Tae Polychaetes, echiurans, sipunculans
Are Arenicolites ±15 Dip, Catenarichnus, Polykladichnus, Sko, Pal Polychaetes, crustaceans, holothurians, insect
Ber Bergaueria ±13 Con, Piscichnus Sea anemones
Bor Bornichnus 1 Root traces,Cho, Pilichnus, Vir Polychaetes, crustaceans
Cam Camborygma 4 Loloichnus, Oph, Tha, Spongeliomorpha, Psilonichnus, Pholeus Crayfish
Con Conichnus 3 Conostichus, Dip, Sca, foot prints Sea anemones, bivalves
Dip Diplocraterion 8 Rhi, Tei, Catenichnus, Tis, Sca, Oph Polychaetes, crustaceans, holothurians, echiurans
Hil Hillichnus 2 Ast, Lophoctenium, Pal, Sko, Pah, Tei Bivalves
Lin Lingulichnus 3 Sip, Con, Sko Lingulid brachiopods
Pad Paradictyodora 2 Dictyodora, Tei, Heimdallia, Stellavelum, Zavitokichnus, Euflabella Bivalves, polychaetes
Pah Parahaentzschelinia 2 Ros, Pad Bivalves
Pho Phoebichnus 4 Stelloglyphus, Oph, Mac, Sip, root traces Arthropods (e.g. crustaceans), worms
Rhi Rhizocorallium 2 Dip, Zoo, Tis, Tae, Sco Polychaetes, crustaceans, insects (e.g. mayflies)
Sco Scolicia ±5 Tae, Zoo Echinoids
Sce Scoyenia 2 Tae Millipeds, insects
Tis Tisoa 4 Dip, Rhi, Are, Paratisoa, Bathichnus Polychaetes
Tri Trichichnus 1 Polykladichnus, Sko Sipunculids
Vir Virgaichnus 1 Tha, Cho, Bor, root traces Nemerteans
More common Dominant Dominant

Less common Subordinate Subordinate
Fig. 5.1 Overview of the abundance of trace fossils observed in core and as described in this book, including some of their features
Burrows
Orientation Subhorizontal Subvertical
Branching Unbranched Branched Unbranched Branched

club-shaped
Tunnel with
meandering
Cylindrical
Bifurcated
U-shaped
Cylindrical
Helicoidal
to conical
U-shaped
Boxwork
Prismatic
J-shaped
U-, bow-
Conical,
Winding/
shaped
Lobate
shaped
shaped
Funnel-
Radial
shafts
shaped
Plug-
Wall-
Shape
Passive
Passive
Passive
Passive
Passive
Passive
Passive
Passive
Passive
Active
Active
Active
Active
Active
Active
Active
Active
Active
Active
Active
Active
Active
Fill
Schaubcylindrichnus
Macaronichnus
Palaeophycus
Ophiomorpha
Siphonichnus
Cylindrichnus
Lining / mantle
Phoebichnus
Phycosiphon
Lingulichnus
Asterosoma
Bornichnus
Hillichnus
Artichnus
Rosselia
Nereites
Parahaentzschelinia
Arenicolites
Trichichnus
Conichnus
Skolithos
Tisoa
No lining / mantle
Paradictyodora
Thalassinoides
Rhizocorallium
Diplocraterion
Camborygma
Teichichnus
Virgaichnus
Zoophycos
Bergaueria
Chondrites
Taenidium
Planolites
Scoyenia
Scolicia
Ichnotaxa in red font may be branched or unbranched

Underlined ichnotaxa include ornamented ichnospecies
Fig. 5.2 Morphological classification of burrows observed in core, following hierarchically arranged diagnostic criteria (modified after Knaust
2012a). The burrows being described and figured in the following sections are listed alphabetically
5.2 Arenicolites Salter, 1857 29
Coastal dunes
Continental ichnofacies Mean sea level
Backshore
Foreshore
(undifferentiated) Upper
Camborygma Shoreface Fairweather wave base
Lower
Palaeophycus Storm wave base
Offshore
Planolites
transition Offshore
Scoyenia
Taenidium Beach
Slope
Bioturbate texture C o n t i n e n t a l s h e l f
Plant roots Basin floor
Teredolites Trypanites Glossifungites

Ichnofacies Ichnofacies Ichnofacies
Psilonichnus Teredolites Gastrochaenolites Diplocraterion
Ichnofacies Trypanites Rhizocorallium
Plant roots Thalassinoides
Entobia Tisoa
Palaeosabella
Rogerella
Skolithos Talpina
Ichnofacies
Arenicolites
Bergaueria
Bornichnus Cruziana
Conichnus Ichnofacies
Cylindrichnus Artichnus
Diplocraterion Asterosoma
Hillichnus Bornichnus Zoophycos
Lingulichnus Cylindrichnus Ichnofacies
Macaronichnus Hillichnus Phycosiphon
Ophiomorpha Palaeophycus Teichichnus
Parahaentzschelinia Paradictyodora Trichichnus Nereites
Schaubcylindrichnus Phoebichnus Zoophycos Ichnofacies
Siphonichnus Phycosiphon
Skolithos Chondrites
Planolites Asterosoma Hillichnus
Bioturbate texture Rhizocorallium Bornichnus Nereites
Rosselia Chondrites Ophiomorpha
Asterosoma Schaubcylindrichnus Nereites
Chondrites Parahaentzschelinia
Scolicia Tisoa Phycosiphon
Palaeophycus Taenidium
Rosselia Planolites
Teichichnus Scolicia
Thalassinoides Trichichnus
Trichichnus Bioturbate texture
Virgaichnus
Bioturbate texture Asterosoma
Tisoa
Arenicolites Zoophycos
Chondrites
Lingulichnus
Nereites
Ophiomorpha
Siphonichnus
Predominant ichnotaxon Skolithos
Accessory ichnotaxon Zoophycos
Fig. 5.3 Trace-fossil associations as dealt with in this book and their occurrence in particular ichnofacies
5.2 Arenicolites Salter, 1857 (e.g. A. carbonarius), while vertical adjustments of the
tracemaker can lead to the development of a thin spreite-like
Morphology, Fill and Size: Arenicolites refers to unbranched structure. Collapsed Arenicolites can also lead to such
U-shaped burrows having a subvertical orientation, with or spreite-like features and resembles Diplocraterion. Tube
without lining and passive fill (Rindsberg and Kopaska- diameter typically is in the range of a few millimeters, and
Merkel 2005; Bradshaw 2010; Figs. 5.4 and 5.5; see also distance between the tubes as well as burrow depth ranges
Fig. 5.138c, d). Secondary successive branching may occur from a few to several centimeters.
Fig. 5.4 Morphological

variability shown on Arenicolites
variabilis (a–f, scale bar = 5 cm)
and oolithic burrow wall (g, scale
bar = 1 cm). From Fürsich
(1974a)
Ichnotaxonomy: About 15 ichnospecies of Arenicolites Skolithos (vertical) or Palaeophycus (horizontal). Com-

have been described in the literature, many of them pending pacted Arenicolites may resemble Palaeophycus.
a critical review. Differentiation criteria include secondary Producers: Polychaete worms and amphipod crustaceans
branching (A. carbonarius) and orientation of the limbs are the most common producers of modern Arenicolites-like
(vertical in A. sparsus, the most characteristic ichnospecies; burrows in marine settings (e.g. Gingras et al. 2008; Brad-
inclined in A. curvatus; variable in A. variabilis; subhori- shaw 2010; Baucon et al. 2014; Fig. 5.7), although other
zontal in A. longistriatus), and presence or absence of lining organisms such as holothurians, sipunculans and echiurids
(Rindsberg and Kopaska-Merkel 2005). may produce Arenicolites too (Smilek and Hembree 2012;
Substrate: Arenicolites is a characteristic burrow in sandy Baucon and Felletti 2013; Baucon et al. 2014). Insects may
(siliciclastic and carbonate) substrate but also occurs in produce similar traces in continental environments (Rinds-
muddy sediment. berg and Kopaska-Merkel 2005).
Appearance in Core: Its simple U-shaped morphology Ethology: Most Arenicolites result from the dwelling
makes Arenicolites easy to recognize in core sections (domichnial) activity of a suspension-feeding organism.
(Fig. 5.6, see also Fig. 5.146d). Although complete burrows Depositional Environment: Arenicolites is known from a
are rarely exposed, individual burrow parts with vertical, wide range of continental to deep-marine environments
J-shaped, elliptical and circular sections become visible in and often occurs in association with other trace fossils
dependence on the section plane orientation with respect to (Fig. 5.9b). Arenicolites is commonly associated with
the burrows. The burrows are passively filled and the high-energy deposition, for instance stormdeposition (lower
occurrence of mud lining may enhance their appearance. to middle shoreface) and migrating dunes and barforms.
Similar Trace Fossils: The U-shaped morphology of Mass occurrences of Arenicolites in low diversity is
Arenicolites is similar to that of Diplocraterion, which can indicative of stressed environments, such as reduced and
be distinguished by its spreite; although some collapsed fluctuating salinity or increased organic productivity, and
Arenicolites can resemble Diplocraterion very confusingly. reflects opportunistic colonization (e.g. Price and McCann
Another U-shaped burrow, Catenarichnus, is a broad 1990; Bradshaw 2010).
arc-like burrow (Bradshaw 2002). Partly preserved Poly- Ichnofacies: Arenicolites belongs to the Skolithos Ichno-
kladichnus (upper parts) could be mistaken for Arenicolites, facies and, subordinately, is part of the Cruziana Ichnofacies.
while incomplete core sections of Arenicolites may resemble Opportunistic colonization of event beds (e.g., storm deposits)
5.2 Arenicolites Salter, 1857 31
(a) (b)
(c) (d)
(e) (f)
Fig. 5.5 Arenicolites in outcrop. Scale bars = 1 cm. a Vertical section sandstone displaying most of an A. sparsus burrow (post-turbidite).
of a sandstone-mudstone alternation containing numerous Arenicolites. Eocene Grès dʼAnnot Formation (deep marine, confined turbidite
Lower Triassic Buntsandstein Group (fluvial), Kahla, Thuringia, system), Braux, southeastern France. After Knaust et al. (2014),
Germany. b Roof bed (lower bedding surface) of section in (a), republished with permission of Wiley; permission conveyed through
showing numerous burrow apertures filled with sand. c Slab of bedding Copyright Clearance Center, Inc. f Sandstone bedding plane preserving
surface with A. cf. variabilis burrow apertures and weathered horizontal the paired plugs of the filled openings of A. cf. sparsus (pre-turbidite).
burrows, probably belonging to Palaeophycus isp. Upper Triassic Eocene Grès dʼAnnot Formation (deep marine, confined turbidite
Kågeröd Formation (fluvial), Bornholm, Denmark. From Knaust system), Chalufy, southeastern France. After Knaust et al. (2014),
(2015b). d Same slab as in (c) (ca. 2–3 cm thick), lower surface with republished with permission of Wiley; permission conveyed through
A. cf. variabilis (vertical burrows) and Palaeophycus cf. alternatus Copyright Clearance Center, Inc.
(horizontal burrows). From Knaust (2015b). e Vertical section of a
Fig. 5.6 Arenicolites in sectioned core. Scale bars = 1 cm. a, area, Norwegian Barents Sea (well 7220/7-2S, ca. 1156.5 m). d Sand-
b Cross-bedded sandstone with reactivation surfaces, from which stone-mudstone alternation with mud-filled Arenicolites partly pre-
mud-lined, passively filled, U-shaped Arenicolites (only partly seen) served and penetrating the sandstone layers from their top. Upper
penetrate the substrate. They are accompanied by bioturbate texture and Jurassic (Kimmeridgian) Heather Formation (offshore), Fram Field,
small equilibrium traces. Upper Triassic (Norian) Fruholmen Formation Norwegian North Sea (well 35/11-11, ca. 2577.5 m). e Dolomitic
(fluvial to deltaic floodplain), Skavl Discovery, Johan Castberg area, mudstone and wackestone having a horizon with mud-lined U-shaped
Norwegian Barents Sea (well 7220/7-2S, ca. 1385.9 and 1395.9 m). burrows. Lower Triassic (Olenekian) Khuff Formation (carbonate
c Heterolithic sandstone with ripple lamination and silty layers, platform with restricted lagoon), South Pars Field, Persian Gulf, Iran
displaying U-shaped burrows with funnel-shaped apertures. Upper (well SP-9, ca. 2866.5 m). From Knaust (2009a), republished with
Triassic (Rhaetian) to Lower Jurassic (Hettangian) Tubåen Formation permission of GulfPetroLink
(marginal marine, tidally influenced), Skavl Discovery, Johan Castberg
was originally referred to the “Arenicolites Ichnofacies” by Reservoir Quality: Given its passive fill, often consisting
Bromley and Asgaard (1991) but such ichnocoenoses are now of sand, a greater abundance of Arenicolites may increase
included in the Skolithos Ichnofacies. the reservoir quality. However, the occurrence of a mud
Age: Arenicolites is known from the Early Cambrian (e.g. lining along the burrow margin may affect negatively
Crimes et al. 1977) to Holocene (e.g. Baucon and Felletti reservoir quality.
2013).
5.3 Artichnus Zhang et al., 2008 33
(a) (b)
(c)
Fig. 5.7 Examples of organisms producing Arenicolites-like burrows (2012a), republished with permission of Elsevier; permission conveyed
in modern sediments. a The polychaete lugworm Abarenicola pacifica through Copyright Clearance Center, Inc. c Resin cast from a boxcore
in its U-shaped burrow. Note the fecal mound above the sediment from the southern North Sea with Arenicolites-like burrows (upside
surface (arrow head). From Dashtgard and Gingras (2012), republished down, up to 3 cm long) produced by C. volutator. Original from
with permission of Elsevier; permission conveyed through Copyright Hans-Erich Reineck, Senckenberg, Wilhemshaven, Germany. From
Clearance Center, Inc. b Monospecific assemblage of Arenicolites-like Knaust (2012b), republished with permission of Elsevier; permission
burrows together with their producer, the amphipod crustacean conveyed through Copyright Clearance Center, Inc.
Corophium volutator. Bay of Fundy, Canada. From Gingras et al.
5.3 Artichnus Zhang et al., 2008
Morphology, Fill and Size: Artichnus is a wide, J-shaped

burrow with a narrow, upward tapering shaft and the distal
end tapering to a blind termination (Figs. 5.8 and 5.9). The
burrow lumen is thickly laminated and accompanied by a
spreite, which is best developed in the lower part of the
burrow. The size of Artichnus in the type material is quite
variable and ranges from 5 to 15 cm in length and 2 to 5 cm
in width (Zhang et al. 2008).
Ichnotaxonomy: A. pholeoides and A. giberti are the only
described ichnospecies. Fig. 5.8 Schematic reconstruction of Artichnus based on observations
Substrate: Artichnus is common in mud-dominated sub- of material from the Eocene Grès d’Annot Formation, southeastern
strates such as heterolithic deposits. France (see Fig. 5.9)
(a) (b)
Fig. 5.9 Artichnus in the Eocene Grès d’Annot Formation (deep with permission of Wiley; permission conveyed through Copyright
marine, turbiditic), southeastern France. Scale bars = 1 cm. a Longitu- Clearance Center, Inc. b Three specimens with a spindle-shaped
dinal burrow part (lower left) and two shafts (centre) displaying the appearance (arrow heads) co-occurring with Scolicia (S) and Arenico-
thick lamination of the burrows. From Knaust et al. (2014), republished lites (A)
Appearance in Core: Although the J-shaped morphology Depositional Environment: Infaunal holothurians, such as
of Artichnus is hard to prove in two-dimensional core sec- Apodida, burrow in littoral to deep-sea environments. So far,
tions, these spreite burrows are typically expressed by their related trace fossils have been described from deep-marine
retrusive spreite element (often dominant) and the passively deposits (Zhang et al. 2008; Knaust et al. 2014). Ayranci and
filled lumen (causative tube) included within it (Fig. 5.10; Dashtgard (2013) and Ayranci et al. (2014) document
see also Ayranci and Dashtgard 2013; Ayranci et al. 2014). Artichnus-shaped traces from modern delta-front and pro-
Longitudinal sections may hint at the J-shaped morphology, delta deposits and hypothesize that in the rock record this
while in cross and oblique sections the lumen appears to be trace could be used as evidence of stable euhaline conditions
circular and elliptical, respectively. The lumen is only a few and, in particular, of deposition and colonization below
millimeters in diameter and reaches a length of a few cen- storm-wave base.
timeters. The entire burrow lies typically in the centimeter Ichnofacies: Artichnus fits nicely within the Cruziana
range. Ichnofacies.
Similar Trace Fossils: Artichnus strongly resembles the Age: In addition to the above-mentioned Jurassic and
similar spreite burrow Teichichnus and has undoubtedly Cretaceous occurrences, Artichnus has been recorded from
been mistaken as such in many cases. Distinction of both Eocene and modern deposits. As this ichnogenus was only
ichnotaxa in core is not straightforward and depends on the recently named, it seems likely that its range will be
recognition of the overall morphology (J-shaped versus extended in the near future.
wall-shaped) as well as on the occurrence of a comparatively Reservoir Quality: A general reduction of reservoir
thick lumen surrounded by a few laminae. Moreover, quality can be assumed of Artichnus-bearing sedimentary
Artichnus has generally a smaller vertical extent compared rocks due to the predominance of the mud-filled laminae and
with Teichichnus. spreite, especially where occurring in high densities.
Producers: Artichnus is interpreted to be produced by
burrowing holothurians (sea cucumbers), probably belong-
ing to the order Apodida (Zhang et al. 2008; Ayranci and 5.4 Asterosoma von Otto, 1854
Dashtgard 2013; Fig. 5.11).
Ethology: The J-shaped morphology of Artichnus with its Morphology, Fill and Size: Asterosoma is a morphologically
thickly laminated rim suggests that it is a dwelling trace variable trace fossil. It consists of a number of arm-like,
(domichnion) of a suspension- or detritus-feeding tracemaker. often bulbous burrows radiating outwards from a central axis
5.4 Asterosoma von Otto, 1854 35
Fig. 5.10 Artichnus in sectioned (a)

core of heterolithic sandstone.
Scale bars = 1 cm. a Dense
ichnofabric with numerous
burrows displaying the lumen in
longitudinal and cross section,
with the dominance of retrusive
spreite elements below it. Lower
Jurassic (Pliensbachian-Toarcian)
Cook Formation (shallow
marine), Norwegian North Sea
(well 34/5-1S, ca. 3658.5 m).
b Thickly laminated burrows
(upper part) accompanied with
spreiten (lower right). Lower
Jurassic (Toarcian-Aalenian) Stø
Formation (offshore), Snøhvit
Field, Norwegian Barents Sea
(well 7120/8-3, ca. 2211.75 m).
c Sandstone bed with numerous
small, mud-dominated specimens.
Lower Jurassic
(Hettangian-Sinemurian) Nansen
Formation (marginal marine),
Oseberg Sør Field, Norwegian
North Sea (well 30/9-16, ca. (b) (d)
3464.9 m). d Dense ichnofabric
displaying individual burrows
with thickly laminated lumina in
longitudinal, oblique and cross
sections. Upper Cretaceous San
Antonio/San Juan Formation, La
Vieja, Macal Tacata, Venezuela
(c)
variable in dimension, number, and orientation and may

show tension fractures as well as faint wall ornamentation
(Fig. 5.13).
Ichnotaxonomy: About five ichnospecies of Asterosoma
have been described so far, of which A. radiciforme and
A. ludwigae are probably the most common (Figs. 5.14 and
5.15).
Substrate: Asterosoma occurs in siliciclastics and car-
bonates alike, although the tracemakers seem to prefer sandy
substrates.
Appearance in Core: Only fractions of the entire burrow
system are commonly exposed in core sections (Fig. 5.16).
Those burrow parts typically appear in form of clusters of
“bulbs”, in which mud laminae surround a passively, often
sand-filled lumen. Parts of the shaft and the ascending tubes
may be associated with the bulbous burrow parts.
Similar Trace Fossils: Ichnospecies of Asterosoma in
core may be confused with other burrows containing a
concentrically laminated fill, such as Cylindrichnus (a
bow-shaped burrow, commonly unbranched), Rosselia (a
dominantly vertical burrow), Artichnus (J-shaped burrow),
Hillichnus (sand-filled tubes, Fig. 5.64a) and Euflabella
(palmate spreite burrow). Transitions between those forms
may occur and may be related to the activity of the same or a
similar producer.
Producers: As with Cylindrichnus and Rosselia, poly-
Fig. 5.11 Schematic diagram of a holothurian of the order Apodida as
potential producer of Artichnus. From Zhang et al. (2008). a Develop- chaete and other worms are potential producers of Astero-
ment of initial burrow. b Well-developed burrow showing the central soma (Chamberlain 1971; Bromley 1996; Pemberton et al.
lumen surrounded by a thick rim of laminae. c Abandonment of burrow 2001; Dashtgard and Gingras 2012; Monaco 2014). Based
with slight effect of compaction
on the occurrence of superficial striae, other authors attribute
Asterosoma to the activity of crustaceans (Müller 1971;
and tapering towards blind extremities, which may be Gregory 1985; Schlirf 2000; Neto de Carvalho and Rodri-
extended into very fine galleries (Schlirf 2000; Bromley and gues 2007). This interpretation is less conclusive because
Uchman 2003; Bradshaw 2010; Fig. 5.12). The burrows are other organisms than crustaceans (including worms) are also
Fig. 5.12 Historical figure of the type ichnospecies of Asterosoma, A. radiciforme, in its original description by von Otto (1854)
Fig. 5.13 Reconstructions of Asterosoma. a Schematic diagram of the (Sp), several of which show the central lumen emerging as an ascending
construction of Asterosoma. After Pemberton et al. (2001), republished tube (At). Total length is about 35 cm. Reconstruction after Bromley
with permission of the Geological Association of Canada. b Asterosoma and Uchman (2003), republished with permission of the Geological
from the Lower to Middle Jurassic of Bornholm, Denmark. A shaft (Sh) Society of Denmark
leads up to a fan-like group of intersecting, steeply inclined spindles
capable producing scratches (see Knaust 2008), and such ranging from paralic to deep-marine settings. In many cases,
features are rather tension fractures than striae (Monaco the ichnotaxonomical determination does not go below the
2014). In addition, other groups of organisms may be con- ichnogenus level and therefore, little is yet known about the
sidered as tracemakers of Asterosoma. For instance, Ayranci environmental significance of individual ichnospecies.
and Dashtgard (2013) describe multiple diminutive burrows Asterosoma is a common constituent of the lower shoreface
with a thick mud lining (Artichnus) produced by holothuri- to offshore transition (or distal ramp setting in carbonate
ans (sea cucumbers) from modern delta deposits; in sec- systems), but also occurs in other parts of the shelf (e.g.
tioned core these burrows mimic Asterosoma (Dashtgard and Farrow 1966; Howard 1972; Gowland 1996; MacEachern
Gingras 2012). In contrast, Percival (1981) interpreted and Bann 2008; Joseph et al. 2012; Pemberton et al. 2012).
star-shaped burrows similar to Asterosoma as the result of Likewise, it is common in deltaic successions, where it
the feeding activity of tellinid bivalves. occurs most frequently in delta front and prodelta deposits
Ethology: Asterosoma is commonly regarded as the (e.g. McIlroy 2004; MacEachern et al. 2005; Gani et al.
feeding trace (fodinichnion) of a deposit-feeding (Schlirf 2007; Carmona et al. 2008, 2009; Dafoe et al. 2010; Tonkin
2003; Bradshaw 2010) or suspension-feeding animal (Neto 2012). Some Asterosoma producers tolerate reduced and
de Carvalho and Rodrigues 2007). fluctuating salinities and occur in environments with
Depositional Environment: Asterosoma has been fre- brackish conditions, including estuaries, bayhead deltas and
quently reported from a wide range of marine environments other paralic deposits (e.g. Greb and Chesnut 1994; Hubbard
(a) (b)
(c) (d)
(e) (f)
Fig. 5.14 Asterosoma in outcrop. Scale bars = 22 cm (a) and 1 cm Wiley; permission conveyed through Copyright Clearance Center, Inc.
(b–f). a Bedding plane with A. radiciforme. Upper Miocene (shallow d Lower bedding plane with A.radiciforme. Eocene, Grès dʼAnnot
marine), East Cape, North Island, New Zealand. b Vertical section Formation (deep marine, turbiditic), southeastern France. From Knaust
showing a cluster of Asterosoma arms with a thick mud lining et al. (2014), republished with permission of Wiley; permission
surrounding a thin, passively filled tube. Lower Jurassic (Hettangian) conveyed through Copyright Clearance Center, Inc. e, f A. ludwigae on
Höganäs Formation (nearshore), Helsingborg, southern Sweden. bedding plane of glauconitic sandstone. Upper Cretaceous
c Lower bedding plane with A. ludwigae (between arrow heads). (Coniacian-Santonian) Bavnodde Greensand (shelf), near Rønne,
Eocene, Grès dʼAnnot Formation (deep marine, turbiditic), southeastern Bornholm, Denmark
France. From Knaust et al. (2014), republished with permission of
Fig. 5.15 Different ichnospecies and preservational variants of Asterosoma. From Seilacher (2007), republished with permission of Springer
et al. 2004; MacEachern and Gingras 2007; Bradshaw 2010; Ichnofacies: Asterosoma is mainly a component of the
Leszczyński 2010; Gingras et al. 2012a; Joeckel and Korus Cruziana Ichnofacies, although it occurs in the Skolithos,
2012; Pearson et al. 2013). Asterosoma even occurs on tidal Zoophycos and Nereites ichnofacies as well.
flats (e.g. Miller and Knox 1985; Knaust 2009a; Knaust et al. Age: Asterosoma has been described from strata of the
2012). Occurrences of Asterosoma in deep-marine environ- entire Phanerozoic from Early Cambrian (e.g. Desai et al.
ments are related to slope-, fan- and basin-floor deposits (e.g. 2010) to Holocene (e.g. Dashtgard et al. 2008).
Pickerill 1980; Powichrowski 1989; Heard and Pickering Reservoir Quality: Asterosoma is a burrow with an active
2008; Uchman and Wetzel 2011; Hubbard et al. 2012; (mud-dominated) fill, which is less favorable for reservoir
Knaust et al. 2014; Monaco 2014). In general, Asterosoma is quality and fluid migration (La Croix et al. 2013; Knaust
related to well-oxygenated environments, although exceptions 2014a).
exist (e.g. Neto de Carvalho and Rodrigues 2007).
(a) (b)
(c)
(e)
(d)
5.5 Bergaueria Prantl 1946 41
b Fig. 5.16 Asterosoma in sectioned core. Scale bars = 1 cm. a Sand- sandstone showing parts of an Asterosoma system. Lower Jurassic
stone with basal pebble-lag deposit and a cluster of mud-lined, bulbous (Pliensbachian-Toarcian) Tofte Formation (fan delta), Åsgard Field,
Asterosoma. Lower Jurassic (Pliensbachian) Nordmela Formation (tidal Norwegian Sea (well 6506/12-I-2H, 4867.6 m). d Sandstone with
flat), Iskrystall Discovery, Norwegian Barents Sea (well 7219/8-2, ca. clustered Asterosoma sectioned in oblique direction. Lower to Middle
3019.1 m). b Sandstone with admixed granulae and bioclasts (trans- Jurassic (Toarcian-Aalenian) Ile Formation (tidal-influenced delta),
gressive unit), partly calcite cemented and displaying an Asterosoma Norwegian Sea (well 6406/8-1, ca. 4388.2 m). e Dolomitic limestone
ichnofabric with horizontal and oblique burrow segments (some of with a dense Asterosoma ichnofabric. Upper Permian Khuff Formation
which are indicated by arrow heads). Middle Jurassic (Bathonian- (muddy tidal flat to grain-shoal transition), South Pars Field, Persian
Oxfordian) Hugin Formation (shoreface), Ivar Aasen Field, Norwegian Gulf, Iran (well SP9, ca. 3025.15 m). From Knaust (2009a), repub-
North Sea (well 16/1-16, ca. 2398.5 m). c Heterolithic and glauconitic lished with permission of GulfPetroLink
5.5 Bergaueria Prantl, 1946 Substrate: Bergaueria is common in siliciclastic deposits

of softground origin and is most easily recognized at the
Morphology, Fill and Size: Bergaueria is defined as a base of sandstone beds. It also occurs in carbonates.
hemispherical to shallow cylindrical, vertical trace fossil Appearance in Core: In core samples, Bergaueria typi-
with a rounded base (Alpert 1973; Pemberton et al. 1988; cally appears as plug-shaped depressions along the base of
Fig. 5.17). Its diameter is generally greater than or equal to sandstone layers (Fig. 5.19). If exposed in an axial position,
its length. Burrow walls are smooth and unornamented; a central peak at the base of the burrow may become visible
although a lining may be present, and the base may contain a (Fig. 5.19c). Bergaueria occurs solitarily or in colonies.
shallow central depression and radial or biradial ridges Similar Trace Fossils: Other plug-shaped burrows might
(Alpert 1973; Pemberton et al. 1988). Burrow fill is gener- be confused with Bergaueria. Especially Conichnus can be
ally structureless and most commonly attached to and similar to Bergaueria but is conical with downward tapering
genetically related to sediment from the overlying bed, laminae, is commonly larger than Bergaueria, and probably
forming convex hyporelief preservation (Pemberton et al. results from a similar maker and mode of construction.
1988; Mata et al. 2012). The size of Bergaueria ranges from Another burrow similar in shape to Bergaueria is Piscich-
less than a centimeter to more than a decimeter in burrow nus, which results from the nesting and yet-feeding activity
diameter and depth. of rays and thus is larger than Bergaueria. In addition to
Ichnotaxonomy: About a dozen ichnospecies of Berg- burrows, there are also sedimentary erosion features such as
aueria have been erected based on morphological differ- potholes (or pot casts), which may resemble Bergaueria.
ences (Figs. 5.17 and 5.18). However, pot casts are not only larger than Bergaueria but
Fig. 5.17 Examples of ichnospecies of Bergaueria. From Seilacher (2007), republished with permission of Springer
(a) (b)
(c) (d)
(e) (f)
Fig. 5.18 Bergaueria preserved in positive hyporelief from outcrops. Cross Mountains, southern Poland. c–f B. perata (c), B. hemispherica
Scale bars = 1 cm. a Holotype of B. perata as originally designed by (d) and B. elliptica (e, f) from Middle Triassic (Anisian-Ladinian)
Prantl (1946). Upper Ordovician Letná Formation, Zdice at Beroun, carbonates of the Meissner Formation (Muschelkalk), Thuringia,
Czech Republic. Original in the National Museum in Praha. After Germany. Note slight affection by pressure solution, particularly in
Mikuláš (2006), republished with permission of the author. b B. sucta on (e) and (f). From Knaust (2007b), republished with permission of SEPM
a sandstone bed from the Uper Cambrian of Wiśniówka Duża, Holy
5.5 Bergaueria Prantl 1946 43
(a) (b)
(c) (d)
Fig. 5.19 Bergaueria in sectioned core. Scale bars = 1 cm. 3195.5 m). c Silty mudstone with Phycosiphon, which is truncated with
a Heterolithic sandstone with ripple lamination and a shifted depression Bergaueria and overlain by cross-bedded sandstone (also with
along a bedding plane, which results from the vertical adjustment of the Phycosiphon). Note the central depression at the base of the burrow.
Bergaueria tracemaker. Lower Jurassic (Sinemurian-Pliensbachian) Lower Jurassic (Hettangian-Pliensbachian) Amundsen Formation
Tilje Formation (sandy tidal flat), Heidrun Field, Norwegian Sea (well (lower shoreface), Fram area, Norwegian North Sea (well 35/10-1,
6507/7-A-38, ca. 2789.5 m). b Cross-bedded, fine-grained sandstone ca. 3655 m). d Ripple-laminated sandstone with mud drapes and a
overlain by medium-grained sandstone and Bergaueria along the Bergaueria, that is accompanied by synaeresis cracks. Middle Jurassic
interface. Lower Jurassic (Sinemurian-Pliensbachian) Tilje Formation (Bajocian) Ness Formation (delta plain), Valemon Field, Norwegian
(sandy tidal flat), Heidrun Field, Norwegian Sea (well 6507/7-A-27, ca. North Sea (well 34/10-23, ca. 4197.8 m)
(a) (b)
Fig. 5.20 Dwelling sea anemones (Actiniaria) as potential producers b Two specimens accumulating debris in a ring-like structure due to the
of Bergaueria on a sandy tidal flat on the Pacific coast of California. activity of the tentacles
a Single specimen with only tentacles emerging on the sandy surface.
also differ from it by their tendency to be enlarged at their 5.6 Bornichnus Bromley and Uchman, 2003
base and having a more irregular shape.
Producers: Sea anemones (Actiniaria) produce Morphology, Fill and Size: Small burrows composed of a
Bergaueria-like traces in modern sediments (Schäfer 1962; crowded tangle of millimetric lined tubes in that are clo-
Bromley 1996). Cerianthus, for instance, can deeply burrow sely and tortuously branched (Fig. 5.22). The whole trace
below the sediment-water interface while extending its ten- fossil occupies an ovoid region of sediment a few cen-
tacles and mouth above the sediment surface to feed (Frey timeters in size (Bromley and Uchman 2003), while the
1970a; Dashtgard and Gingras 2012; Fig. 5.20). In case of tube diameter typically lies within the size range of
disturbance (e.g. due to storm), the animal is able to retract 1–2 mm. In addition, more loosely arranged branched
most of its body down into the sediment for shelter burrows with less tortuous but more linear tubes are also
(Fig. 5.21). Varying burrowing strategies are utilized by the included in Bornichnus.
anemones, including vertical adjustment, which leads to Ichnotaxonomy: B. tortuosus is the only described
collapsed, chevron-like traces, and attachment to a buried ichnospecies. Loosely organized burrows with similar
hard substrate (such as gravel) with only limited vertical character and size occur too and could probably be assigned
shifting (Dashtgard and Gingras 2012). to a new ichnospecies.
Ethology: Bergaueria is interpreted as the dwelling or Substrate: Bornichnus is known from sandy substrate.
resting trace (domichnion or cubichnion) of sea anemones Appearance in Core: These small trace fossils with a tube
(e.g. Pacześna 2010). diameter of only 1–3 mm are easily overlooked but appear
Depositional Environment: Bergaueria is commonly rela- in core as clusters (ovoid areas) or loosely concentrated areas
ted to high-energy nearshore environments such as beaches containing dark-gray burrows (Figs. 5.23 and 5.118a). Thick
and sandy tidal flats, but does also occur in other marine mud lining may result in entirely mud-filled burrows,
environments down to the deep sea (e.g. Książkiewicz 1977). although in some instances the passive fill becomes clearly
Ichnofacies: Bergaueria typically occurs in the Skolithos visible. Morphology and branching pattern is quite variable
Ichnofacies. and includes loosely winding burrow elements with
Age: Bergaueria is known from the Lower Cambrian T-shaped branching points and bifurcation. Association with
(e.g. Pacześna 2010; Mata et al. 2012) to the Holocene (Frey Ophiomorpha burrows is common.
1970a). Some dubious forms are reported from Ediacaran Similar Trace Fossils: Bornichnus resembles other small
deposits (e.g. Fedonkin 1981; Narbonne and Hofmann 1987; trace fossils with which it could be confused, such as rootlets
Seilacher 2007). (plant-root traces), Chondrites (dichotomously branched
Reservoir Quality: Because of its passive sand fill, the burrows with active fill and no lining), Pilichnus (horizontal
occurrence of Bergaueria generally has a slight positive burrows with dichotomous branching), and Virgaichnus
effect on reservoir quality. (boxwork with pinching and swelling burrow diameter).
5.6 Bornichnus Bromley and Uchman, 2003 45
Fig. 5.21 Environmental and taphonomic history of Bergaueria (=“Alpertia”) as a result of burrowing sea anemones. Based on observations on
Middle Devonian strata of Poland. After Orłowski and Radwański (1986)
(a) (b)
(c) (d)
(e) (f)
Fig. 5.22 Bornichnus in outcrop. Scale bars = 1 cm. a B. tortuosus in Madsegrav near Rønne, Bornholm, Denmark. See Nielsen et al.
vertical section from its type locality. Two ovoid clusters with loosely (1996). d Tangled burrows with thick lining. Same locality as in (c).
tangled burrows within glauconitic, cross-bedded sandstone with e B. tortuosus in vertical section occurring in isolation or in connection
limonitic mud flasers (sandy tidal flat). Lower Jurassic with crustacean burrows (e.g. Thalassinoides, Gyrolithes and Sponge-
(Pliensbachian-Toarcian) Sorthat Formation (marginal marine), near liomorpha). The thickly lined burrows are impregnated with ferrugi-
Rønne, Bornholm, Denmark. See Bromley and Uchman (2003) for nous material and thus readily visible. Upper Miocene near Lepe,
details. b As in (a), close-up view of an ovoid cluster consisting of Huelva, southwestern Spain. See Belaústegui et al. (2016b) for
numerous tiny burrows enhanced by limonite. c Vertical section of sand geological setting. f Crowded tangle of thickly lined tubes clustered
with Ophiomorpha nodosa shafts and associated B. tortuosus. Lower around a crustacean tunnel (middle). Same locality as in (e)
Cretaceous (Berriasian) Robbedale Formation (shallow marine),
5.6 Bornichnus Bromley and Uchman, 2003 47
(a) (b)
(c)
Fig. 5.23 Bornichnus in sectioned core. Scale bars = 1 cm. a Bornich- some of which show passive sand fill surrounded by a thick mud lining.
nus ichnofabric with clusters of thickly mud-lined, partially mud-filled Middle Jurassic (Bajocian-Oxfordian) Hugin Formation (shallow mar-
tiny burrows. Middle Jurassic (Bajocian-Oxfordian) Hugin Formation ine), Sleipner Vest Field, Norwegian North Sea (well 15/9-5, ca.
(shallow marine), Norwegian North Sea (well 25/10-12ST2, ca. 2161 m). 3592.5 m). c Cluster of Bornichnus with thick mud lining in turbiditic
b B. tortuosus with an ovoid region with tangled and branched burrows, sandstone. Paleocene (deep marine, lobe complex), off Tanzania
Producers: Various species of modern polychaetes are Ethology: Deposit-feeding (fodinichnial) behavior of the
known to produce Bornichnus-like traces, such as Capito- Bornichnus tracemaker can be inferred.
mastus cf. aciculatus, Scoloplos armiger and Heteromastus Depositional Environment: B. tortuosus was originally
filiformis (e.g. Schäfer 1962; Hertweck 1972; Hertweck et al. described from tidal-flat deposits (Bromley and Uchman
2007; Fig. 5.24). However, co-occurrence of Bornichnus 2003). The examples presented in Fig. 5.23 extend this
and Ophiomorpha may suggest a close relationship between range into the shoreface, shelf and slope, where the producer
these trace fossils. For instance, small burrows similar to has colonized sandy event deposits (e.g. tempestites and
Bornichnus were described originating from brooding turbidites).
chambers connected with large Ophiomorpha burrows and Ichnofacies: Too little data is available for an unequivo-
were produced by juvenile shrimp (Forbes 1973; Bromley cal ichnofacies assignment of Bornichnus, but existing
and Frey 1974; Curran 1976; Verde and Martinez 2004; findings indicate the preferred occurrence within the Sko-
Fig. 5.25). Alternatively, such relationship between small lithos and Cruziana ichnofacies, subordinately also in the
and large burrows could also be the result of commensalism, Zoophycos Ichnofacies.
e.g. between polychaetes and crustaceans as suggested by de Age: The new records have extended the original Lower
Gibert et al. (2006). Jurassic (Pliensbachian to Toarcian) age to the Paleocene.
chamber, or laterally to downward-branched systems

(Hasiotis 2010; Fig. 5.27). Depending on the complexity of
the burrow architecture, their length can range from 10 to
more than 400 cm, with a burrow diameter from 1 to 14 cm
(Hasiotis 2010). The burrow wall can have bioglyphs (e.g.
scratches, striae). In exceptional cases, a tower (chimney)
consisting of pelletoidal sediment can be preserved above
the paleosurface.
Ichnotaxonomy: Based on their burrow architecture, four
ichnospecies of Camborygma are discriminated (Hasiotis
and Mitchell 1993; Fig. 5.26).
Substrate: Crayfish burrows (Camborygma and similar
ichnogenera) are typically related with indurated and car-
bonate-rich paleosols where they occur in a variety of clastic
substrates (e.g. sandstone, mudstone, pyroclastic deposits,
etc.).
Appearance in Core: Crayfish burrows such as Cam-
borygma have been broadly overlooked in core and are
only sporadically recognized as such (e.g. Hasiotis 2010)
because only fractions of the complex burrow systems are
typically exposed in slabbed core (Fig. 5.28). The burrows
are characterized by a large diameter (1 cm or more),
often a sharp margin (because of the firm paleosol sub-
strate), and passive fill (debris) contrasting from the sur-
rounding host rock. Circular to slightly elliptical tunnel
cross sections are common, occasionally accompanied by
vertical shafts and, in rare cases, chamber-like extensions.
Because of its association with seasonally wet soils,
Camborygma typically is associated with root traces and
Taenidium, and some burrow parts may be affected by
calichification.
Similar Trace Fossils: Because of its complex burrow
Fig. 5.24 Vertical section with a fine sand-silt intercalation and
architecture, Camborygma may be confused with morpho-
burrows of the modern polychaete Capitomastus cf. aciculatus from the
nearshore (2–10 m water depth) of the Georgia coastal region, Sapelo logically similar crustacean burrows, making a clear dis-
Island, USA. From Hertweck (1972), republished with permission of tinction in core impossible. Loloichnus is another crayfish
Schweizerbart (www.schweizerbart.de/home/senckenberg) burrow from continental deposits which differs from Cam-
borygma by the absence of chambers, common tunnels and
multiple shafts (Bedatou et al. 2008). Other decapod crus-
Reservoir Quality: Due to the high amount of mud within tacean burrows (e.g. Ophiomorpha, Thalassinoides, Spon-
the burrow, Bornichnus has the potential of decreasing the geliomorpha, Psilonichnus, Pholeus and others) may partly
reservoir quality of otherwise clean sandstone. resemble crayfish burrows (e.g. Martin et al. 2008) but are
restricted to marine environments.
Producers: Camborygma is the burrow of crayfish
5.7 Camborygma Hasiotis and Mitchell, 1993 (continental decapod crustaceans), which have modern
counterparts (Fig. 5.29).
Morphology, Fill and Size: Camborygma includes simple to Ethology: Camborygma is a trace serving for dwelling
complex burrow systems with subvertical shafts, subhori- and reproducing (domichnion; Hasiotis 2010).
zontal tunnels, and chambers (Hasiotis and Mitchell 1993; Depositional Environment: Camborygma is a continental
Fig. 5.26). They can be simple vertical shafts with a basal trace fossil typically associated with weakly- to well-
5.7 Camborygma Hasiotis and Mitchell, 1993 49
Fig. 5.25 Resin cast of the burrow of the modern shrimp Upogebia burrows (ca. 1 mm in diameter) emerge (inset). Scale bar = 10 cm.
affinis (similar to fossil Ophiomorpha and Thalassinoides) sampled in After Bromley and Frey (1974), republished with permission of the
tidal creeks on Sapelo Island, Georgia. Some burrow terminations are Geological Society of Denmark
swollen chambers with a rough surface, from which numerous minute
Fig. 5.26 Architectural morphologies of the different ichnospecies of with multiple shafts and chambers or long burrows that branch at depth
Camborygma with respect to position on the floodplain and depth to the indicate low water tables, decreasing access to surface waters, and long-
water table. C. litonomos composed of simple shafts with little branching term burrow occupation. Modified from Hasiotis and Honey (2000),
and few chambers imply high water table, periodic connectivity to open Smith (2007)
water sources, and shorter-term burrow occupation. Complex burrows
(a) (b)
(c) (d)
Fig. 5.27 Camborygma in caliche pebble-bearing sandstone to con- associated with horizontal tunnels in longitudinal section (left) and
glomerate (alluvial deposits on a floodplain) of the Upper Triassic cross section (right). b Vertical shaft. c Oblique shaft with Y-shaped
(Carnian) Kågeröd Formation at Risebæk, Bornholm, Denmark. Scale branching (upper part) and chamber-like extension (lower part).
bars = 5 cm. After Knaust (2015b). The burrows are passively filled d Burrow cross section
with green mud that is cracked due to desiccation. a Oblique shaft
developed paleosols formed in proximal to distal alluvial and hydrophilic burrows reflect the depth and fluctuation of the
marginal-lacustrine environments (channel, levee and over- ancient water table (Hasiotis and Mitchell 1993). Burrow
bank, floodplain). A high abundance of burrowing crayfish density and size variations are other aspects in the recon-
occurs in humid to hot, seasonal wet climates (Hasiotis struction of paleoenvironments. Lateral variation in burrow
2010), although continental and semiarid occurrences are density may reflect spatial heterogeneity in water table and
reported too (Knaust 2015b; Fiorillo et al. 2016). Starting soil moisture levels, while crayfish size increases from the
from a discontinuity surface at the top, the burrow shafts fluvial channel towards the overbank (Kowalewski et al.
penetrate the vadose zone and typically branch below the 1998). This size segregation along an environmental gradi-
water table with the chamber situated in the phreatic zone ent may directly be used in reservoir prediction as long as
(Figs. 5.26 and 5.30). Architecture and depth of the the data density is sufficient for that.
5.7 Camborygma Hasiotis and Mitchell, 1993 51
(a) (b) (e)

t
t
t
s
t
t
(c)
(d)
s
t
t
c
Fig. 5.28 Camborygma in sectioned core from the Upper Triassic 2736.5 m). c Sandstone paleosol with passively filled shaft (s) and
(Norian-Rhaetian) Lunde Formation (fluvial) in the Snorre Field (a– chamber-like extended tunnel (t) (well 34/7A-9H, ca. 2595.5 m).
d) and the Upper Triassic to Lower Jurassic Hegre Group (alluvial) in d Mudstone paleosol with root traces and calichified tunnel (t) and shaft
the Johan Sverdrup Field (e). Scale bars = 1 cm. a Fine-grained (s) with basal chamber (c) (well 34/7-1, ca. 2532.5 m). e Complex
sandstone containing calichified tunnel elements (t) and a sand-filled burrow system following a fractured limestone, displaying a long
shaft (s) (well 34/7A-4H, ca. 2872.5 m). b Mudstone paleosol with root vertical shaft with horizontal branches filled with green mudstone
traces and sand-filled shaft (s) and tunnels (t) (well 34/7A-9H, ca. within a caliche deposit (well 16/2-17S, ca. 2028.5 m)
Fig. 5.29 Cambarus (Puncticambarus) georgiae Hobbs, 1981, a modern burrowing crayfish from Georgia, USA (from Hobbs 1981). Entire
length ca. 7–8 cm
Fig. 5.30 Generalized crayfish burrows with respect to the water table (from Hobbs 1981)
Ichnofacies: Camborygma is a common constituent of reservoir quality and connectivity (e.g. Fig. 5.28b, c).
continental ichnofacies, including Scoyenia, Coprinisphaera The burrows may increase lateral connectivity of channel-
and Celliforma ichnofacies (Buatois and Mángano 2011; margin deposits (e.g. levee and overbank deposits) with
Melchor et al. 2012). sand-filled channel deposits. However, due to the process of
Age: Crayfish evolved in the Permian and crayfish bur- pedogenization, such burrows may also act as conduits for
rows such as Camborygma are recorded from continental carbonate-rich solutions that become precipitated in form of
deposits of Permian to Holocene age (Hasiotis 2010). pedogenic carbonate (e.g. Fig. 5.28a, d). In that case,
Reservoir Quality: Given their large dimensions and pas- calichified burrows would reduce reservoir quality and
sive fill, Camborygma generally contributes to an improved connectivity.
5.8 Chondrites von Sternberg, 1833 53
5.8 Chondrites von Sternberg, 1833 to the surface, which ramifies with depth under acute angle
to form a dendritic or root-like system (Osgood 1970; Fu
Morphology, Fill and Size: Chondrites is one of the most 1991; Fig. 5.32). Most of the burrows show an active fill,
common and widely distributed trace fossils; due to its sometimes with portions preserving a meniscate structure.
rootlike appearance it was originally interpreted as a plant The burrows are unlined. The tunnel diameter remains
fossil (Fig. 5.31). It consists of tunnel systems possessing a constant in different parts of the burrow and typically is in
single or a small number of master shafts, presumably open the range of less than 1 mm to a few millimeters. The lateral
Fig. 5.31 Historical figures of specimens of Chondrites originally granulata), Lower Jurassic, Germany. b C. intricatus, Oligocene,
interpreted as plant fossils (algae, fucoids). From Steinmann (1907, Switzerland. c C. targionii, Oligocene, Switzerland. d C. bollensis
a–c) and Saporta (1873, d). Burrow diameter is typically in the size (now P. granulata), Jurassic, France
range of few millimeters. a C. bollensis (now Phymatoderma
Fig. 5.32 Different morphologies of Chondrites. a From Tauber (1949), republished with permission of the Geological Survey of Austria. b From
Simpson (1956). c From Staub (1899)
Fig. 5.33 Characteristic ichnospecies of Chondrites, including the four that Fu (1991) regarded as valid. C. bollensis is now assigned to
Phymatoderma granulata. From Seilacher (2007), republished with permission of Springer
extent of individual burrow systems ranges from a few C. recurvus; Figs. 5.33 and 5.34), although this view is still
centimeters to over a decimeter, although continuations debated (e.g. Uchman 1999) and additional ichnospecies
between different burrow systems appear in densely biotur- were subsequently introduced.
bated beds. The burrow systems may reach several cen- Substrate: Chondrites was preferably produced in
timeters in depth, although the true vertical extent often fine-grained softgrounds, although the occasional occurrence
remains uncertain due to incomplete preservation and sub- of uncompacted specimens indicates relatively cohesive
sequent compaction. substrates. It occurs in siliciclastic deposits (such as mud-
Ichnotaxonomy: Chondrites appears with a wide range of stone, marlstone and sandstone) and carbonates (including
morphologies which has given reason for establishing about calcilutite and chalk).
150 ichnospecies. After a revision of the ichnogenus Appearance in Core: In core sections, Chondrites appears
Chondrites, Fu (1991) concluded that only four ichnospecies as clusters of burrows of a similar diameter. Some burrows
were valid (C. targionii, C. intricatus, C. patulus and show the typical branching at an acute angle. Depending on
(a) (b)
(c) (d)
r i
(e) (f)
i
i
t
i
Fig. 5.34 Chondrites ichnospecies in outcrop. Scale bars = 1 cm. (i) and C. recurvus (r). Same locality as in (b). e C. intricatus (i) and C.
a C. intricatus (i) and C. targionii (t). Triassic part of the Hamrat Duri targionii (t). Cretaceous marlstone (flysch), Salzburg Alps (Muntigl),
Group (deep marine), Hajar Mountains, Oman. b C. intricatus. Austria. Coll. Stadtmuseum Berlin. Image courtesy of Beate Witzel
Cretaceous marlstone (flysch), Ligurian Alps (east of Albenga), (Berlin). f C. intricatus. Middle Triassic (Anisian) limestone (shallow
northern Italy. c C. patulus. Same locality as in (b). d C. intricatus marine), Udelfangen Formation near Trier, western Germany
b Fig. 5.35 Chondrites in sectioned core. Scale bars = 1 cm. a Lami- Åsgard Formation (carbonate shelf), Johan Sverdrup Field, Norwegian
nated, dark-gray mudstone with sand-filled Chondrites as the only trace North Sea (well 16/5-2S, ca. 1946 m). d Marly limestone, thoroughly
fossil, plus pyrite nodules. Note the branching of some burrows at an bioturbated, showing indistinct Zoophycos spreite burrows, some of
acute angle. Middle Jurassic (Callovian) Vestland Group (shallow which are intensively bioturbated with Chondrites of different size
marine, restricted basin), Johan Sverdrup Field, Norwegian North Sea classes. Note the dark-gray mud fill of Chondrites. Lower Cretaceous
(well 16/2-16AT2, ca. 2341.5 m). b Marly limestone with Chondrites (Berriasian) Åsgard Formation (carbonate shelf), Johan Sverdrup Field,
of different sizes cross-cutting a Zoophycos spreite (lower part). Lower Norwegian North Sea (well 16/5-2S, ca. 1948.5 m). e Chalk with a
Cretaceous (Berriasian) Åsgard Formation (carbonate shelf), Johan Zoophycos-Chondrites ichnofabric, consisting of a colonization sur-
Sverdrup Field, Norwegian North Sea (well 16/2-11A, ca. 2169.5 m). face, stripe-like Zoophycos spreite burrows and spotty Chondrites.
c Marly limestone (slightly faulted) with Chondrites of different size Upper Cretaceous Shetland Group (carbonate shelf, allochthonous),
classes, some of which are restricted to the sediment fill of large Oseberg Field, Norwegian North Sea (well 30/9-B-46A, ca. 3362.5 m).
Thalassinoides and Zoophycos burrows. Lower Cretaceous (Berriasian) f As in (e), detailed view
(a) (b) (c)
1cm 10 cm
Fig. 5.36 Schematic diagrams of ichnogenera included in the Chon- through Copyright Clearance Center, Inc. c Pragichnus fascis, from
drites group (other than Chondrites). a Pilichnus dichotomus (plan Mikuláš (1997), republished with permission of Schweizerbart (www.
view), from Uchman (1999). b Skolichnus hoernesii, from Uchman schweizerbart.de/journals/njgpa)
(2010), republished with permission of Elsevier; permission conveyed
Fig. 5.37 Phymatoderma granulata, originally assigned to “Chon- Jurassic (Toarcian) black shale (bedding plane) of Holzmaden, southern
drites bollensis”, has a morphology consistent with that in Chondrites Germany. Coll. Palaeontological Institute and Museum, University of
but differs from it by its internal composition of fecel pellets. Lower Zurich
Fig. 5.38 Different interpretations of Chondrites behavior and pro- republished with permission of the Geological Survey of Austria.
ducer. a Deposit-feeding system (fodinichnion) within the sediment by a c Chemosymbiosis of the bivalve Thyasira flexuosa by means of its
vermiform animal. From Richter (1931), republished with permission of extensible vermiform foot. From Dando and Southward (1986), © The
Schweizerbart (www.schweizerbart.de/home/senckenberg). b Dwelling Marine Biological Association, published by Cambridge University
(domichnion) of a suspension-feeding annelid. From Tauber (1949), Press, reproduced with permission
the direction of the section with individual burrows, circular lining. Rutichnus DʼAlessandro et al., 1987 branches in a
(transverse), elliptical (oblique) and elongate (longitudinal) similar manner as Chondrites but is thickly walled and has a
figures become visible (Fig. 5.35; see also Figs. 5.50a, meniscate fill and external rugosity. Planolites is a common
5.121a and 5.161f). A subhorizontal burrow orientation is trace fossil with a horizontal course, active fill and no lining,
most common. In composite ichnofabrics, Chondrites occu- and thus may be confused with Chondrites in core sections,
pies the deepest tier while cross-cutting older traces such as from which it differs by its unbranched nature. Finally, the
Zoophycos, Phycosiphon and Thalassinoides (Ekdale and complex, three-dimensional, irregular burrow system
Bromley 1991; Fig. 3.1b), although in modern sediments Virgaichnus has similarities with Chondrites but differs from
Thalassinoides and Zoophycos can penetrate deeper than it by a higher degree of irregularity, varying burrow diameter
Chondrites. Because of this relationship, Chondrites often and passive fill (Knaust 2010a).
occurs within preexisting burrows (e.g. Thalassinoides, Producers: Chondrites is a heterogeneous and polygenetic
Zoophycos, Planolites), which the tracemaker has trace-fossil group with many ichnospecies, reflecting its wide
reburrowed. morphological variation. Still the burrow architecture itself
Similar Trace Fossils: The Chondrites group contains remains relatively simple comprising only a few significant
some ichnotaxa that were previously regarded as part of the features. Therefore, together with its wide range in age and
ichnogenus Chondrites but are now excluded from it environments, it is very likely that animals of different groups
because of their different morphology. In core, those ich- were producers of Chondrites. Annelids (e.g. polychaetes;
notaxa may be easily confused with Chondrites sensu stricto. Tauber 1949; Hertweck et al. 2007) and sipunculans (Simp-
They include irregularly winding and dichotomously bran- son 1956) are likely candidates. In homology to the peristaltic
ched Pilichnus Uchman, 1999; the radial trace fossil movement of a vermiform organism, the extended and
Skolichnus Uchman, 2010; and the root-like trace fossil branched foot of some bivalves which thrive with
Pragichnus Chlupáč, 1987 (Mikuláš 1997; Fig. 5.36). Hor- chemosymbiosis (e.g. thyasirid and lucinid bivalves) are
izontally branched tunnels filled with fecal pellets were known to produce Chondrites-like burrows in sediments (e.g.
occasionally included in Chondrites (e.g. Kotake 1991) but Dando and Southward 1986; Seilacher 1990, 2007; Fu 1991;
belong to the ichnogenus Phymatoderma (Fu 1991; Miller Dufour and Feldbeck 2003).
2011; Izumi 2012; Fig. 5.37). The dendritic trace fossil Ethology: Due to its wide morphological variability,
Hartsellea Rindsberg, 1994 is another candidate for confu- different behavior models have been derived for Chondrites
sion but differs from Chondrites by branching upward and (Fig. 5.38). Most commonly applied is the interpretation of
5.9 Conichnus Männil, 1966 59
Fig. 5.39 Chondrites in Devonian shale of North America. Scale shale bed from where Chondrites and Zoophycos penetrate the
bars = 1 cm. a Chondrites in black shale (outcrop, bedding plane) underlying substrate. The large bright spot in the lower right corner
filled with contrasting material from the overlying gray shale bed. of the image is a pyrite nodule. Well core 57, New Albany Shale, Upper
Huron Shale, Upper Devonian (Famennian), Clay City, Kentucky. Devonian, Illinois Basin
b Laminated black shale in vertical core section, overlain by a gray
subsurface deposit-feeding behavior (e.g. Osgood 1970), Ichnofacies: Although Chondrites must be regarded as a
although other models include suspension-feeding facies-crossing trace fossil, its widest distribution is docu-
(e.g. Tauber 1949), detritus-feeding on the sea floor (Simp- mented in deep-sea (flysch) deposits belonging to the
son 1956), and chemosymbiosis (Seilacher 1990, 2007; Fu Nereites Ichnofacies.
1991). Age: Chondrites is a common trace fossil from the
Depositional Environment: Many case studies have Cambrian (Webby 1984) to Holocene (Wetzel 1981, 2008)
shown that Chondrites can be regarded as a good indicator and has a wide distribution in the Cretaceous to Paleogene
of dysoxic (between anaerobic and aerobic) settings, where alpine flysch, from which it was originally described as a
the dissolved oxygen in bottom and pore waters of sedi- plant fossil (Fu 1991).
mentary basins is between 0.2 and 1 ml/l (Bromley and Reservoir Quality: In their case study from the Creta-
Ekdale 1984; Savrda and Bottjer 1991; Martin 2004). In ceous Ben Nevis Formation reservoir off Newfoundland,
existing models, such as the one developed by Savrda and Tonkin et al. (2010) found that Chondrites in mudstone-rich
Bottjer 1991), decreasing oxygenation goes hand in hand facies had a net effect of permeability reduction. In other
with reduction of ichnodiversity, and Chondrites appears to cases, however, a positive impact on porosity and perme-
be the last trace fossil before anaerobic conditions set in as ability distribution due to the presence of Chondrites may be
shown by lamination, high organic matter and sulfide min- noticed, chiefly because of the introduction of sandy material
eralization coinciding with decreasing burrow size and into mud-dominated facies and thus an increased hetero-
decreasing depth of bioturbation. Such conditions can often geneity. This may be particularly important for the perfor-
be found in the deep-marine environments (e.g. basin plains, mance of hydrocarbon reservoirs in shale (e.g. Schieber
submarine fans), from which Chondrites is frequently 1999, 2003; Bednarz and McIlroy 2015; Fig. 5.39).
reported. Stagnant conditions, however, also occur on the
shelf (e.g. minibasins) and in nearshore restricted basins (e.g.
estuaries, lagoons, embayments etc.). The fact that the 5.9 Conichnus Männil, 1966
Chondrites-producer tolerates low oxygen content within the
sediment results in its frequent occurrence in transgressive Morphology, Fill and Size: Conichnus is a relatively large,
deposits and particularly in association with maximum conical burrow with subcircular cross section and subvertical
flooding intervals. orientation. Its internal fill is largely passive and a thin lining
3
5
2
1 4
Fig. 5.40 Vertical sections of Conichnus conicus type specimens with 6 Fragments of skeletons of different organisms. From Männil (1966),
passive fill from the Ordovician of Estonia. 1 Fine clastic material, republished with permission of the Borissiak Paleontological Institute
predominantly organogenic. 2 Same, relatively coarse. 3 Fine-grained Moscow. Scale bar = ca. 1 cm
limestone. 4 Marl with thin shaly layers. 5 “Bituminous” limestone.
along the burrow wall may be present (Frey and Howard 1981; and a characteristic apical, papilla-like protuberance (Männil
Pemberton et al. 1988; Figs. 5.40, 5.41 and 5.42). Conichnus 1966; Dronov et al. 2005).
can be related to internal, convex-down (chevron-like) Substrate: Conichnus is a characteristic trace fossil in sandy
structures (due to the adjustment of the producer), or it can be substrate (looseground) of siliciclastic and carbonate origin.
located above such a structure (equilibrium trace). Burrow Appearance in Core: Complete specimens of Conichnus
depth typically ranges between a few centimeters and several are rarely displayed in core samples because of their rela-
decimeters. tively large size. The subvertical burrows show a typically
Ichnotaxonomy: The type ichnospecies, C. conicus, is the plug-shaped morphology (Fig. 5.43). The length/diameter
most common ichnospecies of Conichnus. Nielsen et al. ratio of the burrows is relatively high (about 3:1 to 10:1 or
(1996) described C. conosinus from the Cretaceous of more) and is related to the upward movement (adjustment)
Bornholm, Denmark, which contains an upper dish-shaped of the producing animal during sediment aggradation.
depression above the cone-shaped lower burrow. In addition, Internally, Conichnus burrows often show downward-
Chen et al. (2005) introduced the questionable C. wudan- deflected laminae which lead to a chevron-like appearance.
gensis from the Devonian of China. The inclusion of Such a structure can also be present below the actual
Amphorichnus papillatus in C. conicus (Frey and Howard Conichnus burrow as a response to the vertical adjustment of
1981) remains controversial, because the former is distin- the producer. In addition, some parts of the laterally adjacent
guished from C. conicus by its amphora-like morphology sediment may be impacted by the activity of the producer
and thus show downward-deflected laminae, listric faults or

collapsed sediment. The fill of Conichnus can be reburrowed
to various extents.
Similar Trace Fossils: As pointed out by Buck and
Goldring (2003), Conichnus may be confused with a number
of conical biogenic structures and sedimentary structures of
different natures, such as sand collapse into a cavity, escape
and equilibrium traces of organisms, and biodeformational
excavations by organisms (Fig. 5.44). In addition, fluid-
escape structures can also resemble Conichnus. Conostichus
resembles Conichnus, but differs from it by having an
ornamented wall, which unfortunately can hardly be seen in
core slabs. Slabbed specimens of Conichnus with internal
downward-deformed laminae may resemble Diplocraterion,
which can be roughly in the same size range. Diplocraterion,
however, consists of a spreite and marginal burrow, occa-
sionally accompanied by fecal pellets. The lower, retrusive
burrow part of large spreite burrows (e.g. Teichichnus) may
look like Conichnus in cases where the upper burrow part
has been eroded. Even the footprints of tetrapods (e.g.
dinosaurs) may produce imprints superficially similar to
deformed Conichnus.
Producers: Based on modern analogs, Conichnus can be
regarded as the product of the activity of sea anemones
(Actiniaria) (Schäfer 1962; Shinn 1968; Frey and Howard
1981; et al. 2008; Figs. 5.45, 5.46 and 5.47) and, to a lesser
degree, bivalves (Gingras et al. 2008).
Ethology: The domichnion Conichnus usually results
from the dwelling activity of anemones, which often is
accompanied by an adjustment of the animal’s position in
order to keep pace with shifting substrate thickness
(Fig. 5.41). The latter process may result in equilibrium
traces associated with Conichnus.
Depositional Environment: Conichnus is a common
element of nearshore to shallow-marine environments with
high-energy sedimentary processes, high sediment supply and
frequently shifting substrate (Abad et al. 2006). Conichnus also
occurs in intertidal to shallow subtidal environments (e.g. tidal
flats, estuaries; Curran and Frey 1977) and associated sand
waves, dunes and megaripples (Shinn 1968; Savrda 2002)
as well as flood-tidal deltas and lagoons (Mata et al. 2012).
Ichnofacies: Conichnus is a constituent of the Skolithos
Ichnofacies.
Age: Anthozoa-produced trace fossils such as Conichnus
occur from the Early Cambrian (Pacześna 2010; Mata et al.
2012) to Holocene (e.g. Gingras et al. 2008).
Fig. 5.41 Vertically extensive Conichnus specimen occurring in Upper Reservoir Quality: Given their relatively large size and
Cretaceous sandstone of Alabama, USA. Bold lines and letters delineate passive (sand) fill, deep penetration of Conichnus may con-
burrow segments. Burrow width is ca. 15 cm. From Savrda (2002),
reprinted by permission of the publisher (Taylor & Francis Ltd., http:// tribute to an increased net-reservoir distribution and vertical
www.tandfonline.com) connectivity.
Fig. 5.42 Conichnus conicus in shallow-marine (nearshore) sand- each other due to small adjustments of the producer. C. conicus is
stone. Scale bars = 1 cm (a, b) and 22 cm (c, d). a, b Lower reburrowed with Macaronichnus segregatis. This trace resembles
Cretaceous (Berriasian) Robbedale Formation, Arnager Bay, Bornholm, the feeding trace of rays similar to Piscichnus waitemata and was
Denmark. Note the deformed coarse sand layer related to the lowermost tentatively interpreted as such by Carmona et al. (2008). d Upper
burrow segment. c Lower Miocene Chenque Formation, Santa Cruz, Cretaceous (Campanian) Neslen Formation, East Canyon, Book Cliffs,
Patagonia, Argentina. Note the overlap of several specimens on top of Colorado, USA
Fig. 5.43 Conichnus in sectioned core. Scale bars = 1 cm. a–c Lower delta plain), East Canyon, Book Cliffs, Colorado, USA. Well HCR#1,
to Middle Jurassic (Toarcian-Aalenian) Ile Formation (tidal-influenced ca. 75.6 ft. e, f Lower Cretaceous (Berriasian) Åsgard Formation
delta), Njord Field area, Norwegian Sea (well 6407/10-2). All (carbonate shelf), Johan Sverdrup Field, Norwegian North Sea. The
specimens show significant soft-sediment deformation related to the relatively small burrows are located within a thin layer of biolaminite.
process of burrow generation. a Ca. 3456.8 m. b Ca. 3455.4 m. c Ca. e Well 16/2-11A, ca. 2175.5 m. f Well 16/2-21, ca. 1930.5 m
3685.7 m. d Upper Cretaceous (Campanian) Neslen Formation (lower
(a) (b) (c)
Fig. 5.44 Conichnus-like sedimentary structures and traces. Scale 3481.3 m). c Lower parts of burrows showing retrusive spreite
bars = 1 cm. a, b Sedimentary collapse structures above Ophiomorpha structures. Middle Jurassic sandstone, coastal outcrop in the Brora
tunnels in sectioned core. Middle Jurassic Hugin Formation (marginal area, Scotland, UK
marine, tidally influenced), well 15/9-8 (Sleipner Field, ca. 3483.5 m,
Fig. 5.45 Schematic illustration of modern anemones within their from a muddy sand flat. b Haloclava producta. c The sea onion
burrows from the southeastern Atlantic coast of the United States. From Paranthus rapiformis
Ruppert and Fox (1988). a Extended specimen of Edwardsia elegans
5.10 Cylindrichnus Toots in Howard, 1966 65
Fig. 5.46 Development of Conostichus, a trace fossil similar to contracted physa driven deep. d Anemone in place with expanded
Conichus. a Burrowing anemone on surface. b Physa driving into physa as holdfast. From Chamberlain (1971), © Paleontological
substrate using its body weight, then drawing its body into the Society, published by Cambridge University Press, reproduced with
substrate. c Body column (scapus) expanded against sediment, and permission
5.10 Cylindrichnus Toots in Howard, 1966 regarded as valid ichnogenus (Ekdale and Harding 2015). C.
concentricus is the type ichnospecies (Fig. 5.49). C. japoni-
Morphology, Fill and Size: Cylindrichnus, originally descri- cus Shuto and Shiraishi, 1979, C. pustulosus Frey and
bed from the petroleum-rich Western Interior Basin (USA), Bromley, 1985, C. errans DʼAlessandro and Bromley, 1986,
includes bow-shaped to broadly U-shaped burrows with a C. operosus Orłowski, 1989, C. candelabrus Głuszek, 1998,
passive fill and a concentric lining (Fig. 5.48). They are and C. helix de Gibert et al., 2006 were introduced subse-
commonly unbranched, although branching has been docu- quently, but C. hollowus Nilsen and Kerr, 1978 remains a
mented (e.g. in C. candelabrus Głuszek, 1998). Burrow nomen nudum.
apertures may be slightly enlarged to form funnel-shaped Substrate: C. concentricus is a common element in sili-
entrances, or may be constricted. The diameter of Cylindr- ciclastic sedimentary rocks, where it occurs in a deep-tier
ichnus is in the range of a centimeter or less, and the burrows position and is typically associated with sandy substrates
may reach a length of several centimeters to decimeters, with (Goldring 1996; Fig. 5.50a–c, f–h). It also occurs in car-
a vertical extent of several centimeters. bonates, including chalk (e.g. Frey and Bromley 1985;
Ichnotaxonomy: After some controversy over the ichno- Goldring et al. 2005; Knaust 2009a; Fig. 5.50d, e). Cylindr-
taxonomic status of Cylindrichnus (Goldring 1996), it is now ichnus was built in softground sediment but can show
Fig. 5.47 Modern anemone Cerianthus producing Conichnus-like adjusting its burrow while moving towards the sediment surface. From
burrows. Left Dwelling structure with mucus-lined wall and sediment Schäfer (1962), republished with permission of Schweizerbart (www.
fill below. Second animal burrowing from the sediment surface into the schweizerbart.de/home/senckenberg)
sediment. Right Equilibrium trace of a sediment-buried animal
transitions to such firmground burrows as Glyphichnus finer-grained sediment that thickens upwards. Most frequent
(Goldring et al. 2002). are random cross sections through the burrow system, dis-
Appearance in Core: Because of the limited size of core playing the diagnostic thick lining enclosing a thin central to
samples, only parts of Cylindrichnus burrows are exposed eccentric tube with passive fill. Clustering of burrow sec-
(Fig. 5.50; see also Figs. 4.3 and 5.129a, e). Funnel-shaped tions may occur in dense Cylindrichnus ichnofabrics.
openings of the overall bow-shaped burrows are common Similar Trace Fossils: Cylindrichnus shares its internal
features of Cylindrichnus in core, which continue with composition, such as the thick lining enclosing a passively
tapering and inclining tubes in their deeper parts. Vertical filled tube, with few other trace fossils, particularly Astero-
sections of such burrow parts appear with a V-shaped soma, Rosselia and Artichnus. So far, no consensus has been
structure consisting of a steeply inclined central tube with reached whether or not Cylindrichnus must be regarded as
passive fill, which is surrounded by a wall of predominantly a transitional form to the last mentioned ichnogenera.
Fig. 5.48 Cylindrichnus concentricus (bow-shaped burrows) and their apertural neck (enlarged) leading down to main section of burrow.
infill (from Goldring 1996). a Typical cross sections of five burrows. c Reconstruction of a complete burrow. d Ichnofabric constituent
b Reconstruction as a shallow, bow-like burrow with constricted diagram constructed from the interval with C. concentricus
(a) (b)
(c) (d)
(e) (f)
(g) (h)
Fig. 5.49 Cylindrichnus concentricus in outcrop. Scale bars = 1 cm. in the middle. Weakly developed scratches in the latter indicate partly
a Cross-laminated sandstone with a bow-shaped burrow in longitudinal firmground conditions and suggest transition to the ichnogenus
section and many oblique and cross sections. Miocene Mount Glyphichnus. Same locality as in (c). e Heterolithic sandstone with a
Messenger Formation (deep marine, overbank), sea cliffs of the thickly mud-lined and branched burrow. Campanian Neslen Formation
Taranaki Peninsula, North Island, New Zealand. b Cross-laminated (marginal marine), Jim Canyon, Book Cliffs, Utah, USA.
sandstone with numerous burrow cross sections. Same locality as in (a). f Cross-bedded sandstone with an elongate and thickly sand-lined
c Cross-bedded sandstone with numerous bow-shaped burrows burrow. Campanian Sego Formation (tide-dominated river delta), San
displayed in various sections and enhanced by brownish iron mineral Arroyo, Book Cliffs, Colorado, USA. g Sandstone bedding plane
staining. Eocene Battfjellet Formation (deltaic), Brongniart Fjellet, showing the lower part of a bow-shaped burrow. Campanian Sego
Svalbard. d Sandstone beds with burrow penetration from the top Formation (tide-dominated river delta), Book Cliffs, Colorado, USA.
surface (mainly oblique and cross sections), and a longitudinal section h Sandstone with an oblique burrow section. Same locality as in (g)
(a) (b) (c)
(d)
(e)
(h)
(f) (g)
b Fig. 5.50 Cylindrichnus in sectioned core. Scale bars = 1 cm. carbonate ramp), South Pars Field, Persian Gulf, Iran (well SP9, ca.
a Heterolithic sandstone with high degree of bioturbation, resulting in 3173.9 m). From Knaust (2009a), republished with permission of
an ichnofabric composed of mud-lined Cylindrichnus overprinting GulfPetroLink. e As in (d), thin section showing Cylindrichnus in cross
Chondrites. Middle Jurassic (Bathonian-Oxfordian) Hugin Formation section and significant mud lining. f Heterolithic sandstone with
(restricted lower shoreface), Gina Krog Field, Norwegian North Sea Cylindrichnus burrows (partly with funnel-shaped aperture) on several
(well 15/5-7, ca. 3908.5 m). b Silty sandstone with several bow-shaped colonization surfaces. Upper Jurassic (Oxfordian) Heather Formation
burrows in longitudinal sections. Early Jurassic (Pliensbachian- (shelf turbidites), Fram Field, Norwegian North Sea (well 35/11-11, ca.
Toarcian) Ror Formation (offshore), Lavrans Discovery, Norwegian 2715.5 m). g Sandstone with funnel-shaped burrow aperture. Eocene
Sea (well 6406/2-1, ca. 4853.1 m). c Ichnofabric with multiple burrows Battfjellet Formation (deltaic), Sysselmannbreen well, Svalbard (well
overlying each other. Paleocene Grumantbyen Formation (lower BH 10-2008). h Sandstone with numerous reworked parts of Cylindr-
shoreface to offshore transition), Sysselmannbreen well, Svalbard (well ichnus. These ichnoclasts became resistant due to early diagenetic
BH 10-2008, ca. 825.5 m). d Cylindrichnus in cross-bedded oolitic cementation by siderite along an omission surface prior to reworking.
limestone (grainstone and wackestone). Upper Permian Khuff Forma- Middle Jurassic (Bathonian-Oxfordian) Hugin Formation (shoreface),
tion (storm-reworked sand shoals and sand waves, inner to outer Sleipner Vest Field, Norwegian North Sea (well 15/9-5, ca. 3554 m)
In general, the bow-shaped bauplan of Cylindrichnus differs deposits, sand dunes and shoals, where vertical and steeply
from the one in Asterosoma (with multiply branched or inclined forms predominate (e.g. Howard 1966; McCarthy
radiating horizontal parts), Rosselia (vertical and conical 1979; Pemberton and Frey 1984; Frey and Howard 1985;
termination) and Artichnus (thickly lined, predominantly Frey 1990; Olariu et al. 2012). Cylindrichnus is a common
horizontal lumen). Subvertical parts of Cylindrichnus, dis- constituent of marginal-marine and estuarine environments
playing a funnel-shaped aperture, may lead to confusion with brackish conditions (Jurassic and younger; Netto and
with other vertical trace fossils such as Laevicyclus (formerly Rossetti 2003; MacEachern and Gingras 2007; Buatois and
Monocraterion sensu lato; Stanley and Pickerill 1998; Mángano 2011; Gingras and MacEachern 2012) and occurs
Knaust 2015a), while cylindrical parts resemble Skolithos in association with delta-front and prodelta deposits (e.g.
(Frey and Howard 1985; Fig. 5.51), or have been mistaken Tonkin 2012). Monoichnogeneric occurrences in such set-
as root traces (e.g. Frébourg et al. 2010; see Fig. 5.50d). tings are a good indicator for stressed environments with
Cylindrichnus can be associated with omission (hiatus) reduced salinities. Cylindrichnus is occasionally reported
surfaces and may show intergradations with the firmground from deep-marine deposits (e.g. Nilsen and Kerr 1978;
burrow Glyphichnus (Goldring et al. 2002). Catenarichnus Fig. 5.49a).
Bradshaw, 2002 shares its overall bow-shaped morphology Ichnofacies: Cylindrichnus preferably occurs in the distal
and passive fill with Cylindrichnus but only sometimes has a Skolithos Ichnofacies and is typical within the Cruziana
thin lining. Ichnofacies.
Producers: Polychaete worms (such as terebellids) are Age: Cylindrichnus is common in Mesozoic and Ceno-
good candidates for producing Cylindrichnus-like burrows zoic deposits (Goldring 1996) but also occurs in Paleozoic
(Dashtgard et al. 2008; Belaústegui and de Gibert 2013; strata (e.g. Głuszek 1998; Buatois et al. 2002). Orłowski
Fig. 5.52). In estuarine environments, maldanid polychaetes (1989), Gámez Vintaned et al. (2006) and Desai et al. (2010)
have been observed in Cylindrichnus-like tubes that are used described Cylindrichnus from Early Cambrian deposits.
for head-down mining as well as head-up suspension and Modern examples are given by Dashtgard et al. (2008).
interface deposit-feeding (MacEachern and Gingras 2007). Reservoir Quality: Suspension-feeding animals such as
Funnel-feeding holothurians (sea cucumbers) also produce the producers of Cylindrichnus introduce a certain amount of
bow-shaped burrows with a thick lining around a central mud when colonizing the top surface of sandy waves or
lumen (e.g. Ayranci and Dashtgard 2013) similar to shoals. In this way, dwelling burrows commonly reduce the
Artichnus. sand/mud ratio within the affected interval. As a result,
Ethology: A suspension-feeding behavior of polychaetes horizons with a considerable decrease in porosity and per-
living within the sediment (domichnion) is inferred for meability can occur within highly permeable units and act as
Cylindrichnus. relatively thin baffles or barriers (Knaust 2009a). Those
Depositional Environment: Cylindrichnus preferably horizons with reduced porosity and permeability can either
occurs in shelf settings up to the lower shoreface with occur locally or follow flooding surfaces at the base of
moderate- to low-energy regime, where more horizontal shallowing-upward cycles and then are widespread. In
burrow components and forms slightly oblique to bedding contrast to this, La Croix et al. (2013) noticed that vertical
occur (e.g. Fürsich 1974a). In addition, it is a characteristic connections were generated by Cylindrichnus within fine-
trace fossil in high-energy deposits, including storm grained sandstone of a gas reservoir.
5.11 Diplocraterion Torell, 1870 71
Fig. 5.52 Diagram showing the reconstruction of Cylindrichnus

concentricus as produced by a terebellid polychaete, and the ichno-
fabric resulting from its activity. Scale bar = 1 cm. Reproduced from
Belaústegui et al. (2011). Copyright © 2011 Elsevier Masson SAS. All
rights reserved
5.11 Diplocraterion Torell, 1870
Morphology, Fill and Size: The ichnogenus Diplocraterion

consists of vertical U-shaped spreite burrows (Fürsich 1974b;
Fig. 5.53). The spreite can be retrusive or protrusive, or both
(e.g. Goldring 1962, 1964; Fig. 5.54). It ranges in size from a
few millimeters in length to several decimeters (Fig. 5.55), the
larger size group being more obvious in cores.
Ichnotaxonomy: Morphological differences in Diplocra-
terion have resulted in the erection of several ichnospecies,
of which the type, D. parallelum, remains most important. In
addition to the five ichnospecies distinguished by Fürsich
(1974b), three ichnospecies have subsequently been intro-
duced, resulting in a total of eight ichnospecies. Schlirf
(2011) also includes burrows with an oblique orientation
into Diplocraterion, a procedure which is not followed here
(cf. Knaust 2013).
Substrate: Diplocraterion is produced in softgrounds and
firmgrounds in siliciclastic and carbonate settings.
Appearance in Core: The spreite bounded by the
U-shaped tube (“limbs”) is conspicuous and relatively easy
to recognize in core. It can be sectioned at various angles and
thus produces different projections (Fig. 5.56; see also
Chakraborty and Bhattacharya 2013; Fig. 5.57). Longitudi-
nal sections along the plane of the spreite burrow produce
the most complete expression, while oblique sections across
the spreite are more common and result in a thin but elongate
spreite combined with the passively filled tube. Depending
on the protrusive or retrusive character of the burrow, the
tube may be located either at the base of the spreite or higher
up within it. Oblique sections are also common and show a
“feathering-out” of the spreite. The marginal tube can be
Fig. 5.51 Cylindrichnus concentricus (left) and Skolithos linearis
(right). Scale bar = 1 cm. After Howard and Frey (1984), republished filled with sand or mud, in the latter case often containing
with permission of Canadian Science Publishing; permission conveyed high content of organic material or, more rarely, plant debris.
through Copyright Clearance Center, Inc. The tube can be also laterally enlarged and display features
Fig. 5.54 Retrusive (left) and protrusive (right) Diplocraterion. From

Seilacher (1967), republished with permission of Elsevier; permission
conveyed through Copyright Clearance Center, Inc.
Fig. 5.53 Idealized sketch of Diplocraterion with its elements, in be confused with the spreite burrow Teichichnus (Fig. 5.58).
which the spreite (=septum) between the tube results from the
protrusive and retrusive movement of the tracemaker. One free tube Catenichnus McCarthy, 1979 is a bow-shaped spreite bur-
is shown opening to a normal aperture; the other tube is shown as row similar to Diplocraterion and Teichichnus. Some
having been plugged before erosion and sedimentation took place. specimens of the slender and very narrow U-shaped burrow
From Goldring (1962), republished with permission of Springer D. habichi resemble Tisoa siphonalis and may better be
attributed to it, particularly when lacking a spreite. Collapsed
U-burrows (e.g. Arenicolites) can resemble Diplocraterion,
of lateral accretion, resulting from the adjustment of the the collapsed structure mimicking a spreite. Large
producer in response to loose and shifting substrate. The Ophiomorpha shafts with a thick mud lining and active fill
spreite may incorporate granulae and pebbles and may show can actually look alike small Diplocraterion (Fig. 5.59).
diagenetic modifications such as sideritic cementation. Producers: Two main groups are generally regarded as
A conspicuous feature of some burrows is the concentration producers of Diplocraterion: crustaceans and polychaete
of millimetric elliptical mud pellets (fecal pellets, Coprulus worms. The amphipod Corophium, for instance, is known to
oblongus) within the spreite. produce incipient Diplocraterion (e.g. Dashtgard and
Similar Trace Fossils: Diplocraterion is part of the ich- Gingras 2012), although fossorial crustaceans seem to be
nofamily Rhizocoralliidae and shows close affinity to the absent in the Paleozoic (e.g. Carmona et al. 2004) and thus
ichnogenus Rhizocorallium, which includes horizontal to alternative interpretations become necessary. Boxcores taken
oblique spreite burrows (Schlirf 2011; Knaust 2013). Both in firm substrate of the Baltic Sea have revealed oblique
ichnogenera may partly originate from the same kind of spreite burrows akin to Diplocraterion and Rhizocorallium
producer. Incomplete core sections of Diplocraterion may (Winn 2006; Knaust 2013), where they were produced by
(a) (b)
(c) (d)
(e) (f)
Fig. 5.55 Diplocraterion parallelum in outcrop. Scale bars = 1 cm. a, (shallow marine), Due Odde, Bornholm, Denmark. d Sandstone bedding
b Sandstone with a dense D. parallelum assemblage seen as slit-like surface with slit-like burrow apertures. Lower Cambrian erratic boulder,
traces on the bedding plane (a) and as vertical spreite burrow in vertical northeastern Germany. Coll. Natural History Museum Berlin. e,
section (b). Lower Cambrian Hardeberga Formation (shallow marine), f Vertical spreite burrows in massive (micritic) limestone. Triassic
Snogebæk, Bornholm, Denmark. See Clausen and Vilhjálmsson (1986). (Rhaetian)/Jurassic (Hettangian) boundary, Western Bergamasc Alps,
c Vertical spreite burrow. Lower Cambrian Hardeberga Formation Lombardy, Italy
(a) Ethology: Suspension-feeding behavior within their

(a)
dwelling (domichnion) can be assumed for the producers of
(b) most Diplocraterion (e.g. Goldring 1962; Fürsich 1974b),
although deposit-feeding has been also considered (e.g.
Leaman and McIlroy, 2016).
Depositional Environment: Ichnospecies of Diplocrate-
(b) rion have been commonly recorded in marginal-marine
environments such as tidal settings and estuaries (e.g. bars
and sand flats; Buatois and Mángano 2011; Gingras et al.
2012a; Desjardins et al. 2012; Higgs and Higgs 2015). This
(c) implies that their producers are able to tolerate reduction in
salinity (brackish conditions) and are often subject to
high-energy sedimentation with repeated erosion and depo-
sition (e.g. storm deposition; Figs. 5.61 and 5.62). Mea-
surements of high-density occurrences of Diplocraterion
have shown that the burrow planes may show a preferred
orientation along the bedding plane as a reaction to waves
(d) and currents (e.g. Clausen and Vilhjálmsson 1986; Buckman
1992; Gaillard and Racheboeuf 2006; Rodríguez-Tovar and
Pérez-Valera 2013). Brackish conditions with fluctuating
salinity are also common in the glaciomarine and deltaic
settings from which Diplocraterion has been recorded (Netto
et al. 2012; Tonkin 2012), as well as hyperpycnal flow
(c) deposits (Buatois et al. 2011). Diplocraterion may also
occur in shoreface and offshore environments, where it is
often related to transgressive surfaces (ravinement surfaces,
(d) omission surfaces) due to the action of waves or tides (e.g.
Mason and Christie 1986; Dam 1990; Taylor and Gawthorpe
Fig. 5.56 Schematic diagram showing different sections through 1993; Goldring et al. 1998; Rodríguez-Tovar et al. 2007).
Diplocraterion parallelum with retrusive (lower part) and protrusive
Such surfaces can be utilized for stratigraphical correlation
(upper part) spreite burrows as they would appear in sectioned core.
a Longitudinal section perpendicular to burrow plane. b Oblique and serve as markers (e.g. sequence boundaries; see Olóriz
section through lower part of marginal tube and protrusive spreite and Rodríguez-Tovar 2000). Monoichnospecific Diplocra-
part. c Cross section through retrusive spreite burrow. d Cross section terion produced in laminated rocks may indicate oxygen-
through protrusive spreite burrow
deficient sediment (Leszczynski et al. 1996). Furthermore,
Diplocraterion is one of those ichnotaxa that first appear
terebellid polychaetes. The polychaete Polydora is another after mass extinctions such as at the end Permian (e.g.
candidate for producing Diplocraterion in firmgrounds (Sei- Knaust 2010b; Chen et al. 2011). Isolated occurrences of
lacher 1967; Gingras et al. 2001, 2012a; Fig. 5.60). The fecal Diplocraterion in continental (e.g. Kim and Paik 1997; Xing
pellets Coprulus oblongus are similar to those produced on et al. 2016) and deep-marine deposits (e.g. Crimes et al.
modern tidal flats by the polychaete Heteromastus, which 1981) are sporadically reported, although Martin et al. (2016)
makes polychaetes likely producers for Diplocraterion. More have shown that shallow-marine Diplocraterion may occur
subordinately, holothurians (sea cucumbers) and echiurans within continental deposits. Nevertheless, mayflies are
are known to produce U-shaped burrows with minimal spreite known to produce spreite-like burrows into continental fir-
(Bromley 1996; Dashtgard and Gingras 2012; Smilek and mgrounds (see Knaust 2013), which could resemble
Hembree 2012) similar to Diplocraterion and Teichichnus. Diplocraterion.
(a) (b) (c)
(e)
(d)
b Fig. 5.57 Diplocraterion parallelum in sectioned core. Scale bars = 1 cm. ravinement surface, from which a muddy spreite burrow with a thick
a Highly bioturbated cross-bedded sandstone with large mud- and U-shaped tube penetrates the underlying sediment. The spreite contains
sand-filled burrows (partly retrusive) sectioned roughly longitudinal to millimetric elliptical fecal pellets (Coprulus oblongus) and crystalline
the burrow plane. Lower Jurassic (Sinemurian-Pliensbachian) Tilje granulae incorporated from the ravinement (omission) surface. Pre-
Formation (nearshore, tidal-influenced), Skarv Field, Norwegian North ferred cementation by iron-bearing minerals (siderite?) has led to
Sea (well 6507/5-1, ca. 3580.8 m). b Heterolithic sandstone with wavy differential compaction between burrow and surrounding sediment.
bedding and large D. parallelum (retrusive) sectioned mostly oblique to Middle Jurassic (Bajocian) Hugin Formation (transition between sandy
the burrow plane. Lower Jurassic (Pliensbachian) Tilje Formation tidal flat below the ravinement surface and deeper-marine environment
(mixed tidal flat), Njord Field area, Norwegian North Sea (well above it), Sleipner Field, Norwegian North Sea (well 15/9-1,
6407/10-1, ca. 2990.85 m). c Heterolithic, ripple-laminated sandstone 3660.35 m). e Medium-grained sandstone with D. parallelum devel-
with a D. parallelum ichnofabric. The spreite burrows (partly retrusive) oped below an erosion surface. The fill of the spreite burrow is
were sectioned at various angles (transverse, oblique and longitudinal to heterolithic (sand and mud) and includes sideritic mud clasts (ocher
the burrow plane). Lower Jurassic (Sinemurian-Pliensbachian) Tilje color). Middle Jurassic (Callovian) Fensfjord Formation (upper shore-
Formation (mixed tidal flat), Njord Field area, Norwegian North Sea face), Gjøa Field, Norwegian North Sea (well 36/7-1, ca. 2341.8 m)
(well 6407/10-1, ca. 2995.1 m). d Heterolithic sandstone with a
High
Depth/width ratio of spreite
Diplocraterion
Teichichnus
Low
High Low
Basal curvature of spreite
Fig. 5.58 Various forms of vertical spreite trace fossils showing the of the spreite. Variable scale. Different sources, modified after Corner
transition field between Diplocraterion and Teichichnus. The forms are and Fjalstad (1993), reprinted by permission of the publisher (Taylor &
arranged roughly according to the depth/width ratio and basal curvature Francis Ltd., http://www.tandfonline.com)
Fig. 5.59 Dense Ophiomorpha ichnofabric in Upper Cretaceous lined and actively filled vertical shafts (e.g. upper left) resemble
(Maastrichtian) Springar Formation (deep marine, turbidites) from Diplocraterion and may be confused with it. Scale bar = 1 cm
the Gro Discovery (well 6604/10-1, ca. 3545.5 m). Some of the thickly
Ichnofacies: Diplocraterion is a major constituent of

the Skolithos Ichnofacies (softgrounds), but also occurs
within the substrate-controlled Glossifungites Ichnofacies
(firmgrounds).
Age: Diplocraterion is a long-ranging trace fossil known
from the Cambrian (e.g. Cornish 1986; Bromley and Hanken
1991) to Holocene (e.g. Corner and Fjalstad 1993; Dasht-
gard and Gingras 2012).
Reservoir Quality: Diplocraterion often occurs at a
relatively large size and high concentration along omis-
sion surfaces (e.g. ravinement surfaces, sequence bound-
aries) and therefore may have an influence on the
producibility of a reservoir. For instance, high-density
occurrences of Diplocraterion with a passive sand fill may
promote vertical connectivity, while the incorporation of
mud may act as a baffle. Improved reservoir quality due to
sediment cleaning and the occurrence of highly permeable
Fig. 5.60 Diagram illustrating a Polydora (polychaete worm) associ-
vertical conduits has been documented by Leaman and
ation and the resulting Diplocraterion-like traces from modern firm-
ground at Willapa Bay, Washington. Scale bar = ca. 1 cm. Illustration by McIlroy (2016).
Tom Saunders in Gingras et al. (2001), republished with permission of
Elsevier; permission conveyed through Copyright Clearance Center, Inc.
Fig. 5.61 Movement pattern of Diplocraterion parallelum in response all have been truncated to a common erosion surface. It is considered
to the amount of sedimentation or erosion as indicated by the that repeated phases of erosion and sedimentation led to the develop-
adjustment to depth and mode of preservation. Upper Devonian Baggy ment of the various types. Stage (a), development of burrow (1). With
Beds, England, UK. After Goldring (1964), republished with permis- degradation of the surface, this tube migrates downwards, and at
sion of Elsevier; permission conveyed through Copyright Clearance intervals, new tubes (2 and 3) are constructed (e and c). Sedimentation
Center, Inc. Heights of solid arrows show amount of sedimentation or follows (d and e) but some of the tubes are abandoned. Stage (f), all
erosion. D. parallelum occurs in the various types shown in (f), where tubes are abandoned and erosion reduces them to a common base
5.12 Hillichnus Bromley et al., 2003 79
Fig. 5.62 Different models of Diplocraterion parallelum, lacking the erosion. d Erosion—stability—erosion. e Stability followed by depo-
retrusive vector. These represent variation in three factors: ontogenetic sition. f Rapid erosion followed by deposition. After Bromley and
growth, rate of deposition or erosion, and amount of deposition or Hanken (1991), reprinted by permission of the publisher (Taylor &
erosion. a Stable sedimentary surface. b Slow, steady erosion. c Rapid Francis Ltd., http://www.tandfonline.com)
5.12 Hillichnus Bromley et al., 2003 Ichnotaxonomy: Beside the type ichnospecies H. lobo-
sensis, H. agrioensis was introduced by Pazos and Fernán-
Morphology, Fill and Size: Hillichnus is a highly complex dez (2010) for more regular forms with lateral and vertical
trace fossil comprising various elements occurring at dif- spreite-like components. Due to the occurrence of rising
ferent levels, which could be assigned to different ichnotaxa tubes, Hillichnus can be regarded as part of the ichnofamily
if occurring in isolation. A basal axial tube complex is Siphonichnidae (Knaust 2015a; Fig. 5.141).
accompanied on either side by feather- and spreite-like Substrate: Hillichnus preferably occurs in thin-bedded
structures. The upper part contains an array of upward- sandstone and mudstone facies, or ripple-laminated sand-
curving, linear to undulating tubes that can be mantled stone with a slightly heterogeneous appearance due to the
(Bromley et al. 2003; Fig. 5.63). The entire trace fossil has a admixture of clay material and organic matter.
rather large size, ranging from 10 to 20 cm in width and Appearance in Core: The complex nature of Hillichnus
vertical extension. This disparity in diameter within the same makes this trace fossil difficult to recognize, particularly in
trace fossil is one of its most diagnostic features. core. The individual elements of this trace fossils resemble
(a) however, these structures become very extended, feather-like

and curving and become more slender distally”, called lateral
tubules. In the upper part, rising tubes (inclined, arcuate) are
developed, typically having a distinct mud lining.
Similar Trace Fossils: Due to the nature of core material,
only a small and perhaps unrepresentative part of Hillichnus
can be ordinarily be exposed, which in turn may result in its
identification as different trace fossils. Lateral lamellae of the
basal part can be clustered and thus be reminiscent of
Asterosoma, although no passively filled central tube is
present. The curved lamellae and lateral tubules (feathered
(b) serpent structures) seem to be the trickiest part of Hillichnus,
because they may be easily confused with Lophoctenium or
crowded Palaeophycus. Individual rising tubes, although
inclined and arcuate, may resemble Skolithos. In addition,
funnel-shaped construction of rising tubes probably results
from stationary feeding and conforms to Parahaentzschel-
inia. Finally, spreite structures are common elements of
both, H. lobosensis and H. agrioensis, and may result in
Teichichnus-like burrows.
Producers: Functional interpretation of individual burrow
parts and comparison with modern analogs allowed Bromley
et al. (2003) to interpret Hillichnus as the product of
deposit-feeding bivalves (Paleotaxodonta or Protobranchia).
Ethology: A subsurface deposit-feeding (fodinichnial)
Fig. 5.63 Hillichnus architecture. a The structure of H. lobosensis mode of life of bivalves (e.g. tellinacean) is inferred for
dissected as a five-level model. Structures show a range of variation, in Hillichnus. The bivalve moved through the sediment and
particular those at level C. From Bromley et al. (2003), republished
utilized its palpal tentacles to exploit adjacent sediment for
with permission of Elsevier; permission conveyed through Copyright
Clearance Center, Inc. b Sandstone bedding plane with numerous food (Figs. 5.65 and 5.66). Siphonal excursions to the sea
burrow systems aff. Hillichnus isp. from the Barremian to Aptian floor result in the preservation of an array of upward-curving
Almargem Formation (fluvial to marginal marine?) at Quinta do Grajal, tubes. Association with pyrite may indicate chemosymbiosis
Belas, Portugal. The twig-like, palmate branching pattern is accompa-
with the farming of sulfide-oxidizing bacteria (Bromley et al.
nied by numerous short ascending tubes which represent the probing
traces of the siphon. The dense occurrence of those probing traces led to 2003).
the interpretation of this structure as algae or meniscate backfill of a Depositional Environment: H. lobosensis was originally
different ichnotaxon and recently described as Cladichnus lusitanicum described from deep-marine inner fan environments of sub-
(Neto de Carvalho et al. 2016). Original specimen in the Geological
marine canyons, where it occurs in overbank/levee deposits
Museum of Lisbon, Portugal. Scale bar = 5 cm
(Bromley et al. 2003). Pazos and Fernández (2010) described
H. agrioensis from marginal-marine deposits with tidal
influence. Similar marginal-marine deposits with sandy tidal
different ichnotaxa and could be assigned to those if found in flats can also be assumed for the above presented data from
isolation. Thus, the recognition of Hillichnus depends on the core as well as for the example from Portugal (Fig. 5.63b).
combined occurrence of various diagnostic elements Given the wide distribution of its producers, Hillichnus
(Fig. 5.64). In addition, association with other bivalve-pro- should be present with a very broad distribution.
duced trace fossils facilitates the identification of Hillichnus Ichnofacies: The few records of Hillichnus belong to the
in core. Following the terminology proposed by Bromley deep-marine Nereites Ichnofacies as well as the marginal-
et al. (2003; Fig. 5.63a), the basal segmented structure marine Skolithos and Cruziana ichnofacies.
“appears as a series of discontinuous arcs arranged in line so Age: Only recently, a decent understanding of this com-
as to suggest a chain of barrels”. Slightly higher up in the plex trace fossil has been achieved, which explains its poor
section, the basal tube is a thinly mud-lined and segmented record so far, including Early Jurassic (Ekdale et al. 2012),
structure. Lateral to this basal tube, alternating structures Early Cretaceous (Pazos and Fernández 2010; Neto de
“are now seen to expand in sand into distinct bunches of Carvalho et al. 2016) and Paleocene (Bromley et al. 2003)
curved lamellae, black, somewhat resembling spreiten” examples. Further research will probably extend the strati-
(referred to as lateral lamellae). “Within mud laminae, graphical range of Hillichnus.
5.12 Hillichnus Bromley et al., 2003 81
(a) (b)
t
t
Fig. 5.64 Various burrow elements potentially belonging to Hillich- shoreface), Snøhvit Field, Norwegian Barents Sea (well 7120/8-2, ca.
nus lobosensis in sectioned core. Scale bars = 1 cm. a Clusters of 2109.8 m). b Feathered serpent structures, which consist of clustered
wrinkled structures composed of dark lining material, which are sand-filled tubes surrounded by a thick mud lining (resembling
interpreted as lateral lamellae (resembling Asterosoma, arrow heads) Palaeophycus or Lophoctenium). Lower Jurassic (Pliensbachian) Tilje
and associated with tube sections, which are interpreted as parts of the Formation (marginal marine), Norwegian Sea (well 6607/12-3, ca.
basal tube (t). Lower Jurassic (Toarcian) Stø Formation (lower 4222.85 m)
Fig. 5.65 Successive activities of a deposit-feeding tellinacean b Having withdrawn its siphons, the bivalve digs a short distance
bivalve that could have produced the trace fossil Hillichnus lobosensis. forwards. c The siphons extend along a new path to the surface for
From Bromley et al. (2003), republished with permission of Elsevier; respiration. d A new bunch of probes is undertaken by the inhalant
permission conveyed through Copyright Clearance Center, Inc. a The siphon at the feeding level
inhalant siphon probes successively at a horizon of exploitation.
Lingulichnus preferably occurs in very fine- to medium-

grained sandstone but has been reported rarely from arenitic
limestone with initial firmground conditions (Knaust et al.
2012; Fig. 5.68).
Appearance in Core: The overall morphology of the bur-
rows (vertical, inclined, J- or U-shaped) gives a first indication
on Lingulichnus in core (Fig. 5.69). The pedicle trace is a
relatively small (few millimeters), elongate burrow with cir-
cular cross section and passive fill (Fig. 5.69a). In L. verticalis,
the pedicle trace is commonly located in the lower part of the
burrow, but often it cross-cuts the upper, laminated portion of
the burrow. The valve trace is larger and may have an extended
spade shape (Fig. 5.69b). It may be passively filled or show
internal lamination (active fill) that can be extended as a long
Fig. 5.66 The benthic bivalve Yoldia sp., demonstrating deposit- funnel (Fig. 5.69c, d). Proximal parts of Lingulichnus are
feeding by means of palp proboscides. From Ward and Shumway typically elliptical in cross section (Fig. 5.69e–g).
(2004), republished with permission of Elsevier; permission conveyed Similar Trace Fossils: Siphonichnus is probably the most
through Copyright Clearance Center, Inc.
similar trace fossil to Lingulichnus. It differs from Lingul-
ichnus by its circular instead of elliptical cross section, and
the internal lamination is penetrated by the passively filled
Reservoir Quality: Overall, a slight reduction of reservoir core in a more consistent manner. The internal trace (core) is
properties can be assumed because of intermingling of generally wider than the pedicle trace in Lingulichnus, and
mud-rich sediment into sandy facies by deposit-feeding of only preserves a relatively small portion of the outer lami-
the Hillichnus producer. nated burrow. Some of the adjusted Lingulichnus resemble
Rosselia but are more irregularly laminated and lack the
passively filled terminal burrow in their upper part. They
5.13 Lingulichnus Hakes, 1976 could also be mistaken as Conichnus, which may occur as
stacked funnel-shaped burrows. However, Conichnus does
Morphology, Fill and Size: Lingulichnus consists of verti- not have a passively filled burrow continuing to the apex of
cally or obliquely oriented burrows having a straight, sinu- the funnel-shaped burrow. In horizontal section, Lingulich-
ous or J- to U-shaped morphology (Zonneveld and nus can resemble Lockeia, which differs by its almond-
Pemberton 2003; Fig. 5.67). Cross sections vary with shaped cross section and the lack of a pedicle trace
respect to their position within the burrow and range from (Rindsberg 1994). Finally, the pedicle trace could be mis-
elliptical to subcircular. Portions of the burrows are filled taken as a root trace or as Skolithos, if occurring in isolation
with concentric laminae with a spreite-like appearance, (e.g. due to erosion).
which may be penetrated by a passively filled central lumen. Producers: On the basis of modern analogs (Emig et al.
The size of Lingulichnus is typically in the range of a few 1978) and direct evidence in fossil Lingulichnus (Zonneveld
centimeters with burrow diameters around 1 cm. et al. 2007), lingulide brachiopods (e.g. Lingula and Glot-
Ichnotaxonomy: Depending on their shape and orientation tidia) can be assumed as the producers of this trace fossil
within the substrate, three ichnospecies are differentiated (Figs. 5.69 and 5.70).
(Zonneveld and Pemberton 2003): L. verticalis (straight, Ethology: Lingulichnus mainly represents a dwelling
vertical; Fig. 5.68), L. inclinatus (straight, inclined), and trace (domichnion) of lingulide brachiopods, which have a
L. hamatus (J- and U-shaped, vertical). suspension-feeding behavior. The tracemaker is able to cope
Substrate: The distribution of lingulide brachiopods (the with increased sedimentation rates and adjusts its burrow
producers of Lingulichnus) is controlled by the grain size of accordingly (equilibrichnion; see Zonneveld et al. 2007).
the substrate (Zonneveld et al. 2007). Accordingly, the Depositional Environment: Lingulichnus occurs in many
abundance of Lingulichnus continuously decreases with marine environments but commonly dominates in shallow-
increasing mud content, and is lacking in muddy substrates. and marginal-marine successions (Zonneveld et al. 2007).
5.13 Lingulichnus Hakes, 1976 83
Fig. 5.67 Lingulide infaunal behavior and resulting ichnospecies of (L. inclinatus). b A lingulide in process of arching its pedicle to initiate
Lingulichnus in response to exhumation and sudden burial. Cross reburrowing after exhumation and the formation of a U-shaped burrow
sections showing preservation in horizontal aspect are displayed at (L. hamatus). The brachiopod props itself up with its pedicle, orienting
base. After Thayer and Steele-Petrović (1975), from Zonneveld and its shell downward (T1). The brachiopod burrows using scissorlike
Pemberton (2003), reprinted by permission of the publisher (Taylor & motion of valves. The brachiopod typically burrows deeply enough
Francis Ltd., http://www.tandfonline.com). a Lingulide brachiopod in (T2) to return to the surface in a vertical orientation (T3)
normal dwelling position (L. verticalis) and in oblique orientation
(a) (b)
Fig. 5.68 Lingulichnus verticalis in grainy packstone (arenite) with incipient firmground conditions. Middle Triassic (Anisian) Jena Formation
(Muschelkalk), Thuringia, Germany. Scale bars = 1 cm. a Slightly abraded bedding plane. b Vertical section
(a) (b) (c)
(d) (e) (g)
(f)
Fig. 5.69 Lingulichnus verticalis in sectioned core. Scale bars = 1 cm. bearing L. verticalis. Lower Jurassic (Pliensbachian) Åre Formation
a Burrow in homogeneous sandstone, with a long pedicle trace (arrow (delta plain, tidal flat), Skuld Field, Norwegian North Sea (well
head) and a mud-laminated upper part. Note the adjustment of the 6608/10-14S, ca. 2629.35 m). e Same interval as in (d), but with
burrow in response to rapid deposition. Upper Jurassic (Oxfordian) increased bioturbation resulting in a dense L. verticalis ichnofabric.
Sognefjord Formation (shallow marine), Vega Field, Norwegian North Individual burrows (in addition to many pedicle-trace cross-sections)
Sea (well 35/11-6, ca. 3187.65 m). b Individual specimens showing the can be recognized in the upper left of the image. f Dense ichnofabric of
extended pedicle trace in the lower part of the burrow, and the L. verticalis due to total bioturbation. Some discrete pedicle traces and
spade-shaped laminated trace in the upper part of the burrows. Middle laminated burrow parts are still recognizable. Middle Jurassic (Aale-
Jurassic (Bathonian) Hugin Formation (marginal marine, tidal flat), nian) Ile Formation (delta plain, tidal flat), Skuld Field, Norwegian
Gudrun Field, Norwegian North Sea (well 15/3-9T2, ca. 4503.5 m). North Sea (well 6608/10-14S, ca. 2550.65 m). g Well-preserved
c Ripple-laminated sandstone with several burrows showing the burrow with a pedicle trace in the lower part and a spade-shaped
extended pedicle traces in the lower part (arrow heads) and a wide burrow above, the latter strongly inclined to the left, reflecting
funnel with lamination in the upper part. Lower Jurassic adjustment of the tracemaker to continuous sedimentation. Several
(Pliensbachian-Toarcian) Tofte Formation (fan delta, tidal flat), Skuld casts of linguloids are preserved at the burrows aperture (arrows).
Field, Norwegian North Sea (well 6608/10-14S, ca. 2591.95 m). Lower Jurassic (Toarcian-Pliensbachian) Cook Formation (marginal
d Ripple-laminated sandstone with several colonization surfaces marine), Norwegian North Sea (well 35/10-1, ca. 3653.0 m)
5.14 Macaronichnus Clifton and Thompson, 1978 85
substrates, Lingulichnus is often associated with sandy event

beds such as tempestites. Rapid or sudden burial is consid-
ered as a requirement to preserve Lingulichnus, whereas
specimens in soft substrates with extensive bioturbation only
have a low preservation potential (Kowalewski and Demko
1997). Because lingulide brachiopods can tolerate fluctua-
tions and temporarily lowered salinity, Lingulichnus can be
found in marginal-marine settings dominated by brackish
conditions (e.g. estuaries). In such environments, burrow size
is commonly reduced (Buatois et al. 2005; Gingras et al.
2012a).
Ichnofacies: Lingulichnus belongs to the Skolithos Ichno-
facies but may occur in transitions to the Cruziana
Ichnofacies.
Age: Lingulichnus is recorded from the Early Cambrian
to the Holocene. It is most commonly reported from Paleo-
zoic strata, which is probably related to a preservation bias
(Zonneveld and Pemberton 2003).
Reservoir Quality: The influence of Lingulichnus on
reservoir quality has not been studied so far. Judging from its
morphological features such as subvertical orientation and the
nature of its (partly) passive fill, Lingulichnus is capable of
slightly increasing reservoir quality and vertical connectivity.
5.14 Macaronichnus Clifton and Thompson,

1978
Morphology, Fill and Size: Macaronichnus includes pre-

dominantly horizontal, cylindrical burrows of indefinite
length with a straight, winding, meandering or spiral-like
course (Fig. 5.71), although oblique and vertical burrows
can occur too (e.g. Uchman et al. 2016; see Fig. 5.42c). The
burrows are characterized by an active fill of pale sand, and
an outer mantle composed of dark mineral grains (Bromley
et al. 2009; Figs. 5.72 and 5.73; see also 5.178c). The bur-
rows are unbranched (although occasional branching has
been described by Rodríguez-Tovar and Aguirre 2014) and
Fig. 5.70 Sedimentary structures constructed by modern lingulide
commonly occur in high density. Their diameter ranges
brachiopods that resulted in Lingulichnus verticalis. After Emig et al.
(1978), from Zonneveld and Pemberton (2003), reprinted by permis- between 0.2 and 2.0 cm, and their length can be at least
sion of the publisher (Taylor & Francis Ltd., http://www.tandfonline. several centimeters (Savrda and Uddin 2005).
com). a Longitudinal section of a lingulide within its dwelling structure Ichnotaxonomy: M. segregatis is the only ichnospecies of
(L. verticalis). P pedicle; PM pedicle mass; R retraction position;
Macaronichnus. It includes M. segregatis segregatis and
i inhalant pseudosiphon; e exhalant pseudosiphon projecting above the
sediment-water interface. b Retraction of a lingulide brachiopod into its three other morphological forms described as ichnosub-
burrow. c Formation of lingulide equilibrium trace (L. verticalis). Cross species by Bromley et al. (2009), M. s. lineiformis, M. s.
sections through different levels of a lingulide equilibrium trace are maeandriformis and M. s. spiriformis (Fig. 5.71). Rodrí-
shown at right
guez-Tovar and Aguirre (2014) added a fourth, ichnosub-
species M. s. degiberti, for occasionally branched and partly
It can be related to storm deposits in proximal offshore to oblique to vertically oriented burrows.
lower shoreface settings and is common in intertidal envi- Substrate: Macaronichnus is typically associated with
ronments. Due to the tracemaker’s preference for grainy sandy substrate (looseground).
Macaronichnus segregatis crowded Macaronichnus but the latter having no bifurcation.

This also applies to Nereites, in cases where the actively
filled central core is surrounded by a poorly lobed mantle.
Producers: By neoichnological comparison, opheliid
polychaetes have been identified as producers of modern
Macaronichnus, such as Ophelia limacina (Clifton and
Thompson 1978), Euzonus sp. (Seike 2007, 2008; Dafoe
et al. 2008a, b) and Travisia japonica (Seike et al. 2011)
(Fig. 5.75).
Ethology: M. segregatis is produced by intrastratal
linear forms meandering forms spiralled forms deposit-feeding of opheliid polychaetes (Clifton and
M. s. lineiformis M. s. maeandriformis M. s. spiriformis Thompson 1978; Seike 2008). The polychaetes feed on the
microbes on the surface of the (quartz) grains and process
Fig. 5.71 Morphological variability of Macaronichnus segregatis.
these grains through their gut, while segregating dark-
From Bromley et al. (2009), reprinted by permission of the publisher
(Taylor & Francis Ltd., http://www.tandfonline.com) colored grains with their bristles around their bodies.
Depositional Environment: M. segregatis is a shallow-
marine trace fossil most common in foreshore, shoreface and
Appearance in Core: In core, Macaronichnus typically delta-front deposits (Nara and Seike 2004; Seike 2007;
appears as crowds of more or less horizontal cylindrical Bromley et al. 2009; Quiroz et al. 2010), as well as intertidal
burrows with a winding course, which can be recognized as and shallow subtidal deposits (Clifton and Thompson 1978;
elongate to elliptical and circular burrow sections (Fig. 5.74, Seike 2008). Rodríguez-Tovar and Aguirre (2014) described
see also Fig. 2.6c). Burrow fill contrasts from the sur- M. segregatis from shelf deposits. Under certain circum-
rounding sediment by its pale color and the diagnostically stances, it also can occur in upper slope environments (e.g.
dark mantle (consisting of mica or heavy minerals) sur- Figure 5.74d) where longshore or upwelling currents pro-
rounding that core. vide favorable conditions for the tracemaker. The polychaete
Similar Trace Fossils: Macaronichnus is a simple hori- Euzonus responds with its burrows to beach morphody-
zontal burrow that shares similarities with several other trace namics (Seike 2008). The worms burrow horizontally in
fossils. The active fill of Planolites differs from Maca- various directions in relatively stable conditions, whereas
ronichnus by lacking a mantle, which also applies to Gordia. they move preferentially landward after heavy erosion of the
Chondrites may also create a dense ichnofabric similar to beach face by a storm (Fig. 5.76).
(a) (b)
Fig. 5.72 Burrow architecture of Macaronichnus segregatis-like 2008). Note the pale core surrounded by a mantle with dark mineral
traces produced by the polychaete worm Euzonus in modern beach grains. Scale bars = 1 cm. a Bedding-plane view. b Vertical section
sand of the Hasaki coast, central Japan (original resin peels from Seike
(a)
(b) (c)
(d) (e)
Fig. 5.73 Macaronichnus segregatis in outcrop. Scale bars = 1 cm. Svalbard. d Sandstone with a dense M. s. lineiformis ichnofabric on
a M. s. maeandriformis on a sandstone bedding. Eocene Battfjellet the bedding plane. Lower Cambrian Hardeberga Formation (sandy tidal
Formation (lower delta plain), Brongniartfjellet (Van Keulenfjorden), flat), Snogebæk, Bornholm, Denmark. e Sectioned vertical surface of a
Svalbard. b Sandstone with a dense M. s. maeandriformis ichnofabric on glauconite-rich cross-bedded sand within a fault zone (overturned
the bedding plane. Paleocene Firkanten Formation (shallow marine), section) displaying a cluster of M. segregatis. Lower Cretaceous Arnager
Longyearbyen, Svalbard. c M. s. maeandriformis in bioturbated siltstone. Greensand Formation (storm-dominated shoreface), near Rønne, Born-
Paleocene Grumantbyen Formation (shelf), near Longyearbyen, holm, Denmark
(a) (b) (d)
(c)
Fig. 5.74 Macaronichnus segregatis in sectioned core. Scale bars = 1 cm. equilibrium trace fossils (upper layer) are consistent with a high-energy
a Heterolithic (silty) sandstone with ripple lamination and discrete environment and rapid sedimentation. Middle Jurassic (Bathonian)
burrows consisting of a pale core surrounded by a thin dark mantle. Tarbert Formation (sandy tidal flat), Oseberg Sør Field, Norwegian
Middle Jurassic (Callovian) Fensfjord Formation (shallow marine, North Sea (well 30/9-14, ca. 3133.35 m). d Dense Macaronichnus
lower shoreface), Gjøa Field, Norwegian North Sea (well 36/7-1, ca. ichnofabric on top of a thick turbiditic sandstone bed (light quartzitic
2384.5 m). b Heterolithic (silty) sandstone with ripple lamination and sandstone). The overlain dark silty material is partly incorporated in
small burrows. Middle Jurassic (Bathonian-Callovian) Hugin Forma- some burrows of the shallower tier. The burrows consist of an actively
tion (shallow marine), Gina Krog Field, Norwegian North Sea (well filled core (quartz sand) and a reworked sandy mantle (darker sand).
15/5-7, ca. 3894.0 m). c Cross-bedded sandstone with an inten- Lower Cretaceous (Albian, deep-marine channel system), off Tanzania
sively burrowed layer chiefly containing Macaronichnus. Associated
(a) (b) (c)
Fig. 5.75 Opheliid polychaetes as the producers of modern Maca- (2004). c Travisia japonica. From Seike et al. (2011), republished with
ronichnus-like traces. Scale bars = 1 cm. a Ophelia limacine. From permission of Elsevier; permission conveyed through Copyright
www.marinespecies.org. b Euzonus mucronata. From Nara and Seike Clearance Center, Inc.
(a) (b)
(c) (d)
Fig. 5.76 Resin peels of vertical sections of modern beach deposits in various directions (a and b), whereas under storm conditions, the
from the Hasaki coast, central Japan (a and c) together with schematic tracemakers are forced landward, which results in that preferred
diagrams showing the corresponding burrowing behavior inferred from orientation (c and d). Original resin peels and drawings from Seike
the traces (b and d). Under fair-weather conditions, the worms burrow (2008), republished with permission of Springer. Scale bars = 1 cm
Ichnofacies: Macaronichnus assemblages are common backfilled core can show a meniscate fill, consisting of either
constituents of the Skolithos Ichnofacies (e.g. Pemberton muddy substrate (e.g. fecal material) or sandy sediment.
et al. 2012). Nereites ranges in size from few millimeters to over 1 cm in
Age: M. segregatis has frequently been reported from width/diameter.
Mesozoic and Cenozoic deposits (e.g. Clifton and Thomp- Ichnotaxonomy: About 30 ichnospecies of Nereites have
son 1978; Quiroz et al. 2010), occasionally also from the been described, many of them now regarded as preserva-
Paleozoic (e.g. Bromley 1996; Knaust 2004a) as old as Early tional variants of a few ichnospecies (Uchman 1995; Man-
Cambrian (Fig. 5.73d). gano et al. 2000; Fig. 5.78). Helminthoida, a junior synonym
Reservoir Quality: Several studies have demonstrated the of Nereites (Uchman 1995), can still sometimes be found in
subtle but positive effect of Macaronichnus ichnofabrics on the literature (e.g. Pemberton et al. 2001), as it is the case for
enhanced reservoir quality, which is related to the sorting Scalarituba, another junior synonym of Nereites (Uchman
and cleaning effect due to sediment feeding by the worms 1995; Mangano et al. 2000).
(Gingras et al. 2002; Pemberton and Gingras 2005; Pem- Substrate: Nereites preferably occurs in silty to fine-
berton et al. 2008; Dafoe et al. 2008b; Knaust 2009a, 2014a; grained sandy substrate with a certain amount of mud
Gordon et al. 2010). admixture. In siltstone, the Scalarituba preservational aspect
is more usual.
Appearance in Core: Nereites typically appears in mass
5.15 Nereites MacLeay in Murchison, 1839 occurrences with burrows displaying an actively filled tunnel
enveloped by a halo of reworked sediment (Uchman 1995).
Morphology, Fill and Size: Nereites is defined as a pre- Nereites with a muddy core and sandy halo are common,
dominantly horizontal, unbranched, meandering to winding although sandy burrows also occur (Fig. 5.79; see also
burrow or trail, consisting of an actively filled central core Figs. 4.3, 5.121b, 5.129b, 5.132d and 5.156d). Depending
and a thick lobed mantle (Uchman 1995; Fig. 5.77). The on the degree of winding and meandering of the burrows, a
wide range of sections can occur in core, including semi-
circular cross sections and more or less elongate longitudinal
sections.
Similar Trace Fossils: Particularly in core, Nereites
occurs with Phycosiphon, and both trace fossils can be quite
similar to each other, which weakens their distinction.
However, Phycosiphon is a winding burrow with a muddy or
silty core and a sandy spreite around it. As a result, Phy-
cosiphon burrows often appear as paired cross sections.
Nereites is generally larger than Phycosiphon and has a
concentric or bilobate halo around the mud-rich core (Cal-
low et al. 2013). In contrast to Phycosiphon, vertical loops
and twists are not developed. Nereites shares similarities
with the trace fossil Macaronichnus, which consists of an
actively sand-filled core with surrounding mantle. Nereites
of low contrast could be also mistaken for Chondrites,
which, however, shows branching in three dimensions.
Producers: The producer of Nereites remains uncertain,
but a kind of vermiform animal, probably an enteropneust, is
most likely (Mangano et al. 2000). Rindsberg and Martin
(2003) and Martin and Rindsberg (2007) also consider
arthropods as the producer of Nereites.
Ethology: Nereites could be classified as the trace of a
deposit-feeder (fodinichnion), although the combined loco-
motion and feeding activities would be consistent with an
Fig. 5.77 Nereites from the Lower Devonian of Germany in plan
view (top) and oblique view (bottom). From Seilacher (2007), interpretation as grazing trace (pascichnion; Mangano et al.
republished with permission of Springer 2000).
5.15 Nereites MacLeay in Murchison, 1839 91
(a) (b)
(c) (d)
Fig. 5.78 Nereites from outcrop. Scale bars = 1 cm. a Tidly mean- specimen from the Middle Devonian of Thuringia, Germany (coll.
dering N. irregularis in Eocene shale (flysch), Bregenzerwald, Austria. University of Greifswald). d Nereites (formerly Helminthoida) in
Senckenberg coll., Frankfurt Main (original of Richer 1928). b Detail Cretaceous helminthoid flysch of the Ligurian Alps east of Albenga,
of Nereites isp. from the Middle Devonian of Thuringia, Germany. Italy
Senckenberg coll., Frankfurt Main (original of Richer 1928). c Another
Depositional Environment: Nereites is a typical element Ichnofacies: Nereites is the namesake of the Nereites
of deep-sea deposits, where it preferably occurs in sediments Ichnofacies (Seilacher 1967) and is typical of basin-floor
originating from deposition under moderate energy (Wetzel (flysch) deposits (Uchman 1995). It also occurs in other
2002, and references therein). It also occurs in slope deposits ichnofacies, for instance in the Zoophycos and Cruziana
(Zoophycos Ichnofacies, e.g. channel-levee deposits; e.g. ichnofacies.
Callow et al. 2013), and is common in shelf deposits Age: Nereites is reported from the Cambrian (e.g. Aceño-
(Cruziana Ichnofacies). Finally, Nereites has been reported laza and Alonso 2001) to Holocene (Wetzel 2002).
from sandy estuarine deposits and tidal flats (Martin and Reservoir Quality: The impact of Nereites on reservoir
Rindsberg 2007; Neto de Carvalho and Baucon 2010) and quality depends on the composition of the burrows and their
even from lacustrine (Hu et al. 1998) and glacial (Netto et al. contrast with the hosting sediment. Sand-dominated burrows
2012) deposits. Nereites is a shallow-tier component and occurring in high density, for instance, have the potential to
occurs as postdepositional trace fossil just below the surface increase connectivity in a reservoir (Bednarz and McIlroy
layer within oxygenated sediment (Fig. 5.80). 2015; Fig. 3.6).
(a) (b)
(c)
(d) (e)
Fig. 5.79 Nereites in sectioned core (a–d) and full core (e). Scale original stratification seems to remain intact because of reworking at a
bars = 1 cm. a Heterolithic (silty) sandstone with Nereites consisting of small scale (small burrow size, cryptic bioturbate texture). Upper
a dark (silty) inner zone and a light (sandy) mantle. In silty (dark) layers, Jurassic (Oxfordian) Heather Formation (offshore), Fram Field, Norwe-
burrow density is higher than in sandy (light) layers. Lower Jurassic gian North Sea (well 35/11-9, ca. 2650.25 m). d Completely bioturbated
(Pliensbachian) Amundsen Formation (shallow marine, shelf), Norwe- silty sandstone with a dense Nereites ichnofabric and discrete Schaub-
gian North Sea (well 35/10-1, ca. 3657.2 m). b Cluster of large burrows cylindrichnus. Upper Jurassic (Oxfordian) Heather Formation (off-
with dark muddy core and light sandy mantle in cross section. Upper shore), Fram Field, Norwegian North Sea (well 35/11-9, ca. 2645.3 m).
Cretaceous (Campanian) Neslen Formation (lower delta plain), East e Large Nereites with complex muddy core surrounded by sandy mantle.
Canyon, Book Cliffs, Colorado, USA. Well HCR#1, ca. 266 ft. c Silty Paleocene Grumantbyen Formation (lower shoreface to offshore tran-
sandstone with complete bioturbation due to Nereites, in which the sition), Svalbard (well BH 9-2006, ca. 389 m)
5.16 Ophiomorpha Lundgren, 1891 93
• O. irregulaire—contorted, sand-cored mud pellets

(Fig. 5.84a–c)
• O. annulata—elongated sand pellets perpendicular to
burrow axis
• O. recta—small mud pellets
• O. rudis—irregularly distributed, knobby sand pellets
(Figs. 5.83e, f and 5.84e–g)
• O. puerilis—cylindrical, rod-shaped pellets with rounded
ends
• O. ashiyaensis—granular ornamentation.
O. isabeli Mayoral (1986) is based on the occurrence of a

pseudopelletal structure due to diagenetic processes (e.g.
compaction or differential dissolution) and thus does not
warrant its own ichnotaxonomical name. In contrast to all
other Ophiomorpha ichnospecies, O. puerilis de Gibert et al.
Fig. 5.80 Nereites as shallow-tier trace fossil in deep-marine sedi- 2006 contains fecal pellets assignable to Coprulus oblongus,
ments. From Wetzel (2002) which today is produced by polychaetes (Knaust 2008).
Substrate: Ophiomorpha is a typical constituent of clean
sandstone, both homogeneous and cross-bedded. Due to the
burrowing activity of its producer, a relatively large amount
5.16 Ophiomorpha Lundgren, 1891
of organic matter and fine-grained sediment can be intro-
duced into the burrow and may lead to a heterogenization of
Morphology, Fill and Size: Ophiomorpha consists of box-
the host sediment. The Ophiomorpha tracemaker is capable
works constituting a horizontal maze with vertical shafts
of penetrating sandy beds 1 m thick or more (e.g. storm or
(Fig. 5.81). The burrows are circular to elliptical in cross
turbidite deposits) before turning horizontally at the
section. Branching is Y- and T-shaped, typically with
mud/sand interface. Thus, thin mudstone interlayers can be
enlargement of the junctions. Passive fill is common,
completely penetrated due to the producer’s ability of
although some burrow segments may have active meniscate
searching for food (Fig. 2.6b). Ophiomorpha also occurs in
fill (Figs. 2.6b and 5.84e–g). Burrow lining is diagnostic of
chalk.
Ophiomorpha, consisting of pellets of sand and/or mud
Appearance in Core: Given their diagnostic appearance
along the wall. The burrow diameter varies from one ich-
with the pelleted wall, identification of Ophiomorpha in core
nospecies to another and ranges between 3 and 30 mm,
is comparatively straightforward (Fig. 5.84). Burrows are
while complete burrow systems are reported to be several
commonly larger than accompanying trace fossils and the
meters in extent and more than 1 m in depth.
pelleted wall is quite distinct, although a more homogeneous
Ichnotaxonomy: Several ichnospecies are distinguished
mud lining can give reason for confusion with Palaeophycus.
on the basis of overall morphology and the shape and
Horizontal burrow parts typically dominate (Fig. 5.146e) but
composition of the pellets (Uchman 2009):
vertical shafts are sometimes exposed in core sections
(Fig. 5.146f). It must be stressed that Ophiomorpha can be
• O. nodosa—regularly distributed, knobby sand pellets
passively or actively filled. Active fill typically is meniscate
(Figs. 5.22c, 5.82, 5.83c, d, 5.84d and 5.178d)
(Figs. 5.84e, g, 5.146f and 5.161d).
• O. borneensis—regularly distributed, bilobate sand pel-
Similar Trace Fossils: The local discontinuation of pel-
lets (Fig. 5.83a, b)
lets may give reason for confusion with other (supposed)
Fig. 5.81 Morphological

variability of Ophiomorpha and Shafts Regular maze Regular boxwork
modern analogs. After
Chamberlain and Baer (1973),
Frey et al. (1978), adapted from
Anderson and Droser (1998),
republished with permission of
Wiley; permission conveyed
through Copyright Clearance
Center, Inc.
Irregular maze Irregular boxwork
Tiered maze Meander maze
crustacean burrows such as Thalassinoides, Spongeliomor- tracemakers may resemble Chondrites but differ from it by
pha, Psilonichnus, Pholeus and Scoyenia (Fig. 5.85). Bur- their boxwork architecture and the occurrence of pellets.
rows with compact linings may resemble Palaeophycus and Open Ophiomorpha tunnel elements may be subject to
gradational transitions to other crustacean burrows can be collaps, which then may lead to dawnwards deflected lami-
found in dependence on the substrate properties. However, nae in the overburden sediment, mimicking Conichnus
Palaeophycus differs from Ophiomorpha by preferred hori- (Fig. 5.44a, b).
zontal orientation and its usually unbranched nature. Producers: By comparison with modern analogs, there is
Actively filled Ophiomorpha tunnels are superficially similar good confidence that the producers of Ophiomorpha belong to
to the backfilled burrows Taenidium and Scolicia, which are thalassinidean shrimp, particularly callianassids (Figs. 5.25,
unbranched and lack pellets or significant vertical compo- 5.86 and 5.87). In fact, several hundred fossorial species of the
nents, while actively filled shafts may resemble Diplocra- extant thalassinideans (families Callianassidae and Upogebi-
terion (Fig. 5.58). Small Ophiomorpha produced by juvenile idae) are known (Knaust et al. 2012).
Fig. 5.82 General characteristics

of O. nodosa. After Pollard et al.
(1993). Scale bars = 1 cm except
(f) = 10 cm. a Dissected burrow
with sandy wall pellets, smooth
inner surface to lining and muddy
sand restriction. b Taper of
restricted burrow. c Two shafts (c)
with taper into muddy layer,
assumed to be colonization
surface. d Shaft truncation. e Base
of restricted shaft leading into T-
junction with another gallery. (d)
f Lined shaft passing down into
humic rich sand without lining.
Humic sand truncated and
burrowed by Thalassinoides cf.
suevicus
(b)
(a)
(e)
(f)
Ethology: Ophiomorpha-producing shrimp can be regarded as a significant component of shallow-marine

deposit- and suspension-feeders (domichnial or fodinichnial), facies (Frey et al. 1978; Pollard et al. 1993), several ich-
depending on the involved species but often also within the nospecies of Ophiomorpha are characteristic of deep-marine
same species (Nickell and Atkinson 1995). Interconnected, deposits (Tchoumatchenco and Uchman 2001; Uchman
highly organized and systematic burrow elements (such as 2009), and its cross-facies relationship is discussed by
polygons and sinusoidal segments), particularly along the Monaco et al. (2009). Few reports exist where Ophiomorpha
sandstone/mudstone interface, indicate more advanced is documented from continental settings, although those
behavior and have been interpreted as agrichnia (Cummings burrows can be also produced by other means and assigned
and Hodgson 2011). The pellets of Ophiomorpha often to different ichnotaxa (Goldring and Pollard 1995; cf. Bau-
originated from the mixture of mucus and sediment. Because con et al. 2014). Although not exclusively, Ophiomorpha is
of their occurrence along the interface between open, oxy- a typical component related to high-energy environments.
genized burrow and anoxic host sediment, these pellets have Estimations of the time available for colonization and bur-
a high potential for diagenetic modification. row construction, referred to the colonization window,
Depositional Environment: Ophiomorpha is one of allowed Pollard et al. (1993) to differentiate sandy shoreline
the commonest trace fossils and occurs in a wide range sedimentary environments such as shoreface, offshore tidal
of paleoenvironments (Leaman et al. 2015). Originally shelf sand-wave facies, and estuarine facies (Fig. 5.88). On
(a) (b)
(c) (d)
(e) (f)
Fig. 5.83 Ophiomorpha in outcrop. Scale bars = 1 cm except bedding plane of a sandstone. Eocene Grès d’Annot Formation (deep
(f) = 10 cm. a O. borneensis in sandstone. Lower Cretaceous (Berri- marine, turbiditic), southeastern France. From Knaust et al. (2014),
asian) Robbedale Formation (shallow marine, nearshore), Arnager Bay, republished with permission of Wiley; permission conveyed through
Bornholm, Denmark. b Part of O. borneensis. Upper Cretaceous Copyright Clearance Center, Inc. f A cluster of O. rudis on the bedding
(Campanian) Bearpaw-Horseshoe Canyon Formation (marginal marine), plane of a sandstone. Miocene Mount Messenger Formation (deep
near Drumheller, Alberta, Canada. c, d O. nodosa in limestone. Note the marine, channel-levee system), sea cliff of the Taranaki Peninsula, North
adjacent occurrence of passive and active burrow fill. Erratic boulder, Island, New Zealand
Cretaceous, Greifswalder Oie, northeastern Germany. e O. rudis on the
(a) (b) (c) (g)
(f)
(d) (e)
Fig. 5.84 Ophiomorpha in sectioned core. Scale bars = 1 cm. 35/11-11, ca. 2724.5 m). e O. rudis in thin-bedded heterolithics. Upper
a O. irregulaire shaft. Lower Jurassic (Sinemurian-Pliensbachian) Cretaceous (Maastrichtian) Springar Formation (deep marine), Norwe-
Tilje Formation (nearshore, tidal-influenced), Skarv Field, Norwegian gian Sea (well 6604/10-1, ca. 3648.5 m). f O. rudis shaft and tunnels
North Sea (well 6507/5-1, 3581.2 m). b O. irregulaire shaft. Middle filled with glauconitic sediment. Lower Cretaceous (Albian, deep
Jurassic (Bajocian) Ile Formation (shoreface, tidal-influenced), Norwe- marine, channel system), off Tanzania. g O. rudis ichnofabric with total
gian Sea (well 6406/8-1, ca. 4421.5 m). c O. irregulaire shaft. Middle bioturbation in the upper part and discrete, actively filled burrows in the
Jurassic (Bathonian-Callovian) Hugin Formation (shoreface, tidal- lower part. Upper Cretaceous (Santonian) Kvitnos Formation (deep
influenced), Sleipner Vest Field, Norwegian North Sea (well 15/9-5, marine, turbiditic), Aasta Hansteen Field (well 6707/10-2A, 4477.0-
ca. 3627.2 m). d O. cf. nodosa ichnofabric in sandstone. Upper Jurassic 4477.5 m). See Fig. 3.4e for porosity/permeability changes due to
(Callovian) Heather Formation (offshore), Fram Field area (well bioturbation
loose soft firm

Ophiomorpha nodosa Thalassinoides suevicus Spongeliomorpha iberica
(pelleted) (smooth) (scratched)
Fig. 5.85 Burrow part showing three different components, which features, three different names are applied on the ichnogenus level.
result from the activity of the same kind of organism (a supposed Modified after Schlirf (2000)
thalassinidean crustacean) in different substrate. Due to this contrasting
Fig. 5.86 Three examples of

burrowing crustaceans. From
Bromley and Asgaard (1972),
GEUS. a Nephrops norvegicus,
b Callianassa major, and
c Glyphea rosenkrantzi
Fig. 5.87 Mounded topography

created by the deep-burrowing
callianassid Glypturus
acanthochirus on a wide tidal flat
in the Bahamas. From Knaust
et al. (2012), republished with
permission of Elsevier;
permission conveyed through
Copyright Clearance Center, Inc.
Fig. 5.88 Diagram to illustrate colonization window. From Pollard limited to periods of mud deposition. c Shoreface environment where
et al. (1993). a Sand-wave migration and opportunity for colonization physical restraints on colonization probably were limited to infrequent
of the trough area. b Schematic diagram to illustrate depositional storms
processes on estuarine point bar where opportunity for colonization was
Fig. 5.89 Dense Ophiomorpha-

Thalassinoides boxwork in
hummocky cross-stratified
sandstone with upwards-
increasing degree of cementation
with iron-stained minerals
(limonite?). Middle Miocene
Tatsukushi Formation (Misaki
Group, shoreface), southeast
coast of Shikoku Island, southern
Japan
the basis of a sequence-stratigraphic analysis in the Upper end-Permian mass extinction. From the Late Jurassic onward,
Cretaceous of Utah, Anderson and Droser (1998) argued that Ophiomorpha can be also found in deep-marine deposits
Ophiomorpha-related bioturbation is more pervasive in (Tchoumatchenco and Uchman 2001; Uchman 2009).
lowstand systems tracts compared to transgressive systems Reservoir Quality: Given their complex nature and mul-
tracts, because of the predominance of marginal- and tiple kinds of producers involved, Ophiomorpha can tend
nearshore-marine, sand-dominated settings as favorable both improve and diminish the quality of reservoir rock.
habitats for colonization by Ophiomorpha producers. Because of its preferred suspension-feeding mode of life,
Ichnofacies: Within the shallow-marine realm, shrimp introduce a considerable amount of mud into their
Ophiomorpha with its significant vertical shafts belongs to sand-hosted dwellings, which can lead to a drastic reduction
the Skolithos Ichnofacies, but overlaps with the Cruziana of porosity and permeability as known from the Vøring case
Ichnofacies, where the horizontal burrow elements are study (Fig. 3.4). In the same area, thin gas-bearing sandstone
more pronounced. In the deep sea, the O. rudis ichnosub- beds within shaly matrix are vertically connected by long
facies was established by Uchman (2009) as part of the Ophiomorpha shafts (Fig. 5.84e). The best documented
Nereites Ichnofacies. There, O. rudis characterizes high- example of an Ophiomorpha- and Thalassinoides-dominated
energetic proximal and axial turbidite deposits (e.g. Knaust reservoir is the Biscayne aquifer in Florida, which relies on the
2009b). burrowing activity of callianassid shrimp (Cunningham et al.
Age: Burrowing shrimp appear in the Permian, from which 2009, 2012). In the shallow-marine Cretaceous Ben Nevis
Ophiomorpha is known (e.g. Chamberlain and Baer 1973), to Formation off Newfoundland, Ophiomorpha-dominated ich-
the present (e.g. Leaman et al. 2015). Baucon et al. (2014) nofabrics are documented as important agents of the net-pay
reported Ophiomorpha from Permian fluvial deposits in Italy intervals (Tonkin et al. 2010). The relatively large caliber of
and suggested that their ghost shrimps producers invaded Ophiomorpha burrows promotes enhanced fluid flow and
marine environments during the recovery following the thus precipitation of early diagenetic minerals (Fig. 5.89).
5.17 Palaeophycus Hall, 1847 101
5.17 Palaeophycus Hall, 1847
Morphology, Fill and Size: Palaeophycus refers to subhori-

zontal, essentially cylindrical, straight or slightly curved
burrows with a lining and passive fill (Figs. 5.90 and 5.91).
The burrows are typically unbranched or have unsystematic
branching (Keighley and Pickerill 1995). The size of Palaeo-
phycus may range from 1 mm or less to more than 1 cm in
diameter.
Ichnotaxonomy: The ichnogenus Palaeophycus includes
about 20 valid ichnospecies of which thin-walled P. tubu-
laris and thick-walled P. heberti are most relevant in core
samples. Other ichnospecies are very thinly lined and differ
from each other by varying kinds of striae (Pemberton and
Frey 1982).
Substrate: Palaeophycus occurs in diverse grain sizes
mainly in soft, but also in firm substrate of both, siliciclastic
Fig. 5.90 Palaeophycus heberti in a reconstruction by Howard and
and carbonate deposits. Frey (1984), republished with permission of Canadian Science
Appearance in Core: The more or less horizontally ori- Publishing; permission conveyed through Copyright Clearance Center,
ented burrows with their lined wall and the passive fill make Inc. Scale bar = 1 cm
(a) (b)
(c) (d)
(e)
Fig. 5.91 Palaeophycus in outcrop. Scale bars = 1 cm. a P. heberti in burrows. c Large, elongate P. cf. heberti burrows on bedding plane of
the rim of chert nodules (silicified limestone, vertical section). Middle silty sandstone. Paleocene Grumantbyen Formation (proximal shelf),
Permian Kapp Starostin Formation (mixed siliciclastic carbonate ramp), near Longyearbyen, Svalbard. d Same as in (c), close-up view. e P. cf.
Akseløya, Svalbard. b Same as in (a), detail view showing crowded alternatus, burrow with weak annulation. Same locality as in (c)
plain and delta front; Fig. 5.93c). These occurrences indicate

that the tracemaker was euryhaline. Palaeophycus is also
common in shoreface and offshore deposits, where it occurs
at much higher ichnodiversities. It has also been reported
from continental slopes (Hubbard et al. 2012) and deep-sea
fans (Uchman and Wetzel 2012; Fig. 5.93d).
Ichnofacies: In the marine realm, Palaeophycus belongs
to the Cruziana Ichnofacies and, subordinately, to the Sko-
lithos, Zoophycos and Nereites ichnofacies, while continental
Palaeophycus occurs within a wide range of defined ichno-
facies (MacEachern et al. 2012; Melchor et al. 2012), of
which the Mermia and Scoyenia ichnofacies are probably
most relevant.
Age: Aside from dubious reports of Palaeophycus from
Fig. 5.92 General morphology of Palaeophycus and its appearance in the Upper Proterozoic, reliable occurrences are known
slabbed core throughout the entire Phanerozoic.
Reservoir Quality: Under optimal circumstances and if
Palaeophycus distinctive in core. They occur as circular, occurring in high densities, passively sand-filled Palaeo-
elliptical and elongated sections (Figs. 5.92, see also phycus burrows may contribute to an improved reservoir
Fig. 5.111b). Their appearance (e.g. wall thickness, size, quality.
density, etc.) is related to the encountered ichnospecies,
facies and limiting environmental factors (Fig. 5.93).
Palaeophycus may occur as individual burrows or in high 5.18 Paradictyodora Olivero et al., 2004
densities.
Similar Trace Fossils: Palaeophycus is a relatively sim- Morphology, Fill and Size: Paradictyodora is a complex
ple trace fossil with correspondingly high potential for vertical spreite burrow that widens upwards, displaying a
confusion with similar ichnogenera. Planolites and Maca- prismatic to conical shape (Figs. 5.94, 5.95 and 5.96). It
ronichnus resemble Palaeophycus in morphology but differ consists of subvertical folded laminae produced by the lat-
from it by their active fill. Other lined burrows, such as the eral migration of a subvertical J-shaped tube (Olivero et al.
horizontal parts of Ophiomorpha and Siphonichnus, may be 2004). Paradictyodora is a relatively large burrow reaching
similar to Palaeophycus, as is the case for parts of more several centimeters in vertical and lateral extent.
complex trace fossils such as Hillichnus (Fig. 5.64b). A de- Ichnotaxonomy: Only two ichnospecies of Paradictyo-
tailed study of many specimens commonly reveals the true dora have been described so far: P. antarctica and
nature of Palaeophycus. P. flabelliformis. Tursia DʼAlessandro and Fürsich, 2005
Producers: Vermiform animals (e.g. annelids) are the was erected approximately at the same time as Paradictyo-
most likely producers of Palaeophycus, although other dora and is a junior synonym of it (Serpagli et al. 2008).
groups of organisms (e.g. arthropods) cannot be ruled out. Substrate: All few records of Paradictyodora to date are
Modern traces similar to Palaeophycus are produced by from sandy substrates.
nereidid polychaetes (Dashtgard and Gingras 2012; Gingras Appearance in Core: In horizontal view from the top
et al. 2012). (bedding-parallel), the laminae of Paradictyodora merge
Ethology: Palaeophycus is commonly interpreted as the into a regularly to irregularly meandering or spiraling band
dwelling (domichnion) of a predaceous or suspension- with backfilled meniscate structure. In vertical view, laminae
feeding animal (Pemberton and Frey 1982). appear as series of regularly imbricate, oblique to subvertical
Depositional Environment: Palaeophycus occurs in a bands (Olivero et al. 2004; Fig. 5.97). The few specimens
wide range of paleoenvironments in marine and continental observed in core show a winding and undulating trace ca.
settings. In the continental realm, Palaeophycus is common 12–15 cm long in vertical section. Individual spreite bur-
in (but not restricted to) fluvial and lacustrine deposits rows are filled with alternating sand and mud, while a
(Fig. 5.5c, d). Low-diversity assemblages with diminutive band-like trace appears in horizontal section.
burrows occur in marginal-marine environments with Similar Trace Fossils: Paradictyodora may be confused
brackish conditions (e.g. estuarine, intertidal and subtidal; with similar vertical spreite burrows such as Dictyodora,
Fig. 5.93a, b), while dense Palaeophycus assemblages with Teichichnus, Heimdallia, Stellavelum, Zavitokichnus and
large burrows are reported from delta deposits (lower delta Euflabella (Olivero et al. 2004; DʼAlessandro and Fürsich
5.18 Paradictyodora Olivero et al., 2004 103
(a) (b)
(c) (d)
b Fig. 5.93 Palaeophycus in sectioned core. Scale bars = 1 cm. a Sand- (well 15/9-1, ca. 3532.3 m). c Sandstone with dense ichnofabric almost
stone with small specimens in oblique and cross section. Middle entirely consisting of large Palaeophycus. Lower Jurassic (Pliens-
Jurassic (Bathonian) Hugin Formation (marginal marine), Gudrun bachian) Cook Formation (marginal marine, deltaic?), Norwegian
Field, Norwegian North Sea (well 15/3-9T2, ca. 4490.4 m). b Argilla- North Sea (well 34/2-2, ca. 3670.0 m). d Silty sandstone with
ceous sandstone with complete bioturbation, consisting of a dense individual Palaeophycus. Upper Cretaceous (Campanian) Nise Forma-
ichnofabric of small Palaeophycus. Middle Jurassic (Callovian) Hugin tion (deep marine), Aasta Hansteen Field, Norwegian Sea (well
Formation (shallow marine), Sleipner Field, Norwegian North Sea 6707/10-1, ca. 3021.25 m)
Fig. 5.95 Variant of Paradictyodora antarctica (schematic recon-

struction). From Olivero et al. (2004), © Paleontological Society,
published by Cambridge University Press, reproduced with permission
Fig. 5.94 Reconstruction of Paradictyodora antarctica. From Olivero
et al. (2004), © Paleontological Society, published by Cambridge
University Press, reproduced with permission. a Three-dimensional inner shelf to lower shoreface settings (Bourgeois 1980;
view with the causative burrow in black and successive curved Serpagli et al. 2008) and in protected shoreface deposits
segments along the meandering band (1–6). b Plan view and vertical
section
(DʼAlessandro and Fürsich 2005).
Ichnofacies: Paradictyodora seems to be part of the
Cruziana Ichnofacies.
2005; Michalík and Šimo 2010; Olivero and López Cabrera
2013).
Producers: Deposit-feeding activity by the lateral shift of
the inhalant siphon of tellinid bivalves and by the lateral
displacement of the feeding shaft of the polychaete Areni-
cola were assumed as potential tracemaking processes by
DʼAlessandro and Fürsich (2005) (Fig. 5.98). Serpagli et al.
(2008) argue that the preservation of the inhalant siphon
trace (e.g. Fig. 5.97a, b) supports the tellinid bivalve model
(Fig. 5.98).
Ethology: Deposit-feeding (fodinichnial) behavior can be
inferred from the winding spreite burrows (Olivero et al.
2004; DʼAlessandro and Fürsich 2005). Fig. 5.96 Plan view of course of lateral displacement of Paradictyo-
Depositional Environment: P. antarctica originally is dora (=Tursia) flabelliformis. Arrows indicate direction of spreite
displacement. After DʼAlessandro and Fürsich (2005), reprinted by
described from fine-grained deposits of deep-marine fan
permission of the publisher (Taylor & Francis Ltd., http://www.
systems (Olivero et al. 2004). P. flabelliformis occurs in tandfonline.com)
5.18 Paradictyodora Olivero et al., 2004 105
(a) (b) (d)
(c)
(e) (f)
b Fig. 5.97 Paradictyodora in sectioned core from the Middle Jurassic arrow heads). c Same specimen as in (a) and (b), horizontal section at
(Bathonian-Callovian) Hugin Formation (shallow marine, a–f) and the the top (well 15/6-4, ca. 10637′). d Sandstone with an alternating
Lower Jurassic (Pliensbachian) Amundsen Formation (shallow marine, mud-sand spreite burrow with undulating course in vertical section.
g), Norwegian North Sea. Scale bars = 1 cm. a, b Completely Sleipner Vest Field (well 15/9-1, ca. 3544.0 m). e, f Intensely
bioturbated sandstone with a vertical section (part and counterpart) of bioturbated sandstone with a muddy spreite burrow in vertical and
an undulating and vertical spreite burrow, predominantly filled with horizontal section (well 25/7-2, ca. 4465.3 m)
mud and showing an internal, sand-filled tubular burrow (between
Fig. 5.98 Alternative ethological models of Paradictyodora (=Tursia) DʼAlessandro and Fürsich (2005), reprinted by permission of the
flabelliformis. a Interpretation as produced by lateral shift of the siphon publisher (Taylor & Francis Ltd., http://www.tandfonline.com). c Inter-
of a deep infaunal deposit-feeding bivalve (tellinid model). b Interpre- pretation as produced by the wandering position of the inhalant siphon
tation as expression of the lateral shift of one shaft of a vertical of a tellinid bivalve, while the trace of the exhalant siphon (left) remains
U-shaped burrow in connection with the feeding activity of a finally preserved. From Serpagli et al. (2008)
deposit-feeding worm-like organism (Arenicola model). a and b from
Age: Paradictyodora has been described from the Late (Chamberlain 1971), which leads to an overall funnel-
Cretaceous (Bourgeois 1980; Olivero et al. 2004) to the shaped burrow. A conical depression may be left on the
Pleistocene (DʼAlessandro and Fürsich 2005). The core upper surface of a bed where mud-filled tubes have been
material presented herein extends the occurrence of Para- extensively developed. The type ichnospecies P. ardelia is
dictyodora to the Early Jurassic. relatively small (1.5 mm in tunnel diameter, 15–20 mm deep
Reservoir Quality: No evaluation of the reservoir quality and 15–60 mm in surface diameter) and has an inclined
influenced by Paradictyodora is known so far. Given its main tunnel at its apex (Fig. 5.99). In contrast, P. surlyki
mud-rich active fill, reducing reservoir quality can be assumed Dam, 1990 is larger (4–20 mm in tunnel diameter, up to
in places where Paradictyodora occurs more densely. 50 mm deep and up to 120 mm in surface diameter) and
consists of vertically bundled burrows with sand fill and mud
lining, radiating vertically or obliquely upward to the sedi-
5.19 Parahaentzschelinia Chamberlain, 1971 ment surface from a central and vertical main shaft. The
tubes may show a distinct meniscate fill (see Głuszek 1998).
Morphology, Fill and Size: Parahaentzschelinia consists of Ichnotaxonomy: The diagnosis of P. egesheimense Schwei-
numerous small, irregular, mud- and sand-filled tubes radi- gert (1998) is only based on size differences and variations
ating vertically and obliquely upward to the sediment surface in tube density and therefore has to be ascribed to one of the
5.19 Parahaentzschelinia Chamberlain, 1971 107
Fig. 5.99 Parahaentzschelinia ardelia in plan view on the bedding © Paleontological Society, published by Cambridge University Press,
surface (left), cross section of initial development of burrowing (middle), reproduced with permission
and complete perforation of sediment (right). From Chamberlain (1971),
two existing ichnospecies of Parahaentzschelinia. Likewise, producing bivalves are preserved below the funnel-shaped
Rosselia rotatus McCarthy, 1979 with its backfilled main siphonal trace (Fig. 5.103). Furthermore, the burrowing
tube is similar to P. ardelia and may be regarded as junior behavior of particular holothurians may produce similar
synonym of it. traces.
Substrate: Parahaentzschelinia is most common in sili- Ethology: By analogy with modern counterparts, Para-
ciclastic sandy substrates but also occurs in limestone. haentzschelinia can be interpreted as the feeding trace (fo-
Appearance in Core: In vertical core sections, Para- dinichnion) of a tellinid bivalve. The similar trace fossil
haentzschelinia appears as funnel-shaped burrows (if sec- Paradictyodora probably also results from the feeding
tioned in an axial position) with the main tube preserved in activity of a tellinid bivalve (Serpagli et al. 2008).
an apical position (Fig. 5.100). Marginal sections typically Depositional Environment: Parahaentzschelinia was
miss the basal tube and appear in form of an inverted conical originally described from deep-marine deposits (Chamber-
structure. Internally, the burrow is intensely laminated or lain 1971), where it was found subsequently by other
shows irregular convolution. One (the terminal) or more of workers (Uchman 1995, 1998; Tunis and Uchman 1996;
the mud-lined tubes are typically preserved and can be either Monaco 2008; Heard and Pickering 2008; Wetzel 2008;
passively or actively filled with sand. Fig. 5.100c). It is often reported from shallow-marine
Similar Trace Fossils: The overall shape of Para- environments with higher energy conditions (Fig. 5.104),
haentzschelinia commonly gives reason for confusion with including foreshore (Fürsich et al. 2006), tidal flats (Mán-
similar ichnospecies of Rosselia, from which it differs by its gano and Buatois 2004), high-energetic shoal (Knaust
overall funnel-like shape (instead of bulbous or spindle-like) 2009a), shoreface (Bann and Fielding 2004),
and a more irregular and laminated internal structure (instead storm-dominated shelf (Dam 1990) and tide-dominated delta
of a concentric geometry). In addition, more than one ter- (McIlroy 2007). Parahaentzschelinia has also been
minal tube can be preserved and is randomly positioned encountered in lithographic limestone deposited in a coastal
within the funnel-shaped burrow. Thus, R. rotatus McCar- lagoon with influence of turbidity currents (Schweigert
thy, 1979 is better accomodated within Parahaentzschelinia 1998), and from other marginal-marine (estuarine?) settings.
(Fig. 5.101). Paradictyodora is a vertical spreite structure Ichnofacies: Shallow-marine occurrences of Para-
with subvertical folded laminae (Olivero et al. 2004). In haentzschelinia are typical constituents of the Skolithos
vertical sections, compact forms of Paradictyodora may be Ichnofacies, while deep-marine occurrences are associated
confused with Parahaentzschelinia but differ from it by with the Ophiomorpha rudis ichnosubfacies.
curved and meandering spreite segments. Age: Parahaentzschelinia occurs from the Carboniferous
Producers: Originally attributed to the feeding activity of (Chamberlain 1971; Głuszek 1998) to the Holocene (Wetzel
a worm-like animal, Parahaentzschelinia is now best 2008). A wide distribution is known from the Jurassic.
explained as the trace of a tellinid bivalve (e.g. Bromley Reservoir Quality: Due to the incorporation of various
1996; Fig. 5.102). Tellinid bivalves apply various methods amounts of mud into the burrows, Parahaentzschelinia may
of feeding with the means of their siphons, either in a sub- have a slightly negative impact on reservoir quality, which is
surface position or, most commonly, on surface detritus. In partly compensated for by the preferred vertical to slightly
some instances, the casts of the Parahaentzschelinia- oblique orientation of the tubes.
(a) (b) (c)
(d) (e) (f)
(h)
(g)
5.20 Phoebichnus Bromley and Asgaard, 1972 109
b Fig. 5.100 Parahaentzschelinia in sectioned core. Scale bars = 1 cm. North Sea (well 16/2-7, ca. 1967.6 m). e Small specimen in ripple-
(a), (b) and (f) from Knaust (2015a), republished with permission of laminated sandstone. Lower Jurassic (Sinemurian-Pliensbachian)
Elsevier; permission conveyed through Copyright Clearance Center, Tilje Formation (marginal marine), Åsgard Field, Norwegian Sea (well
Inc. a P. surlyki, nicely displaying the central mud-laminated tube and 6506/12-K-3H, ca. 4524.5 m). f Three funnel-shaped specimens of
irregular disturbance with mud incorporated into the funnel-like P. surlyki in ripple-laminated sandstone. The burrows are accompanied
burrow. Middle Jurassic (Bajocian) Ile Formation (marginal marine), with and partly cross-cut by Siphonichnus. Middle Jurassic (Bajocian)
Trestakk Discovery, Norwegian Sea (well 6406/3-2, ca. 4075.35 m). Ness Formation (deltaplain), Valemon Field, Norwegian North Sea
b A funnel-shaped and sideritic specimen of P. surlyki. Middle Jurassic (well 34/10-23, ca. 3236.5 m). g Small specimen in ripple-laminated
(Bajocian) Ile Formation (marginal marine), Norwegian Sea (well sandstone. Lower Jurassic (Pliensbachian) Åre Formation (marginal
6406/8-1, ca. 4382.95 m). c Small specimen emplaced in a thin marine), Skuld Field, Norwegian Sea (well 6608/10-12, ca. 2796.9 m).
turbiditic sand bed, with the bivalve cast preserved below the funnel. h Dense ichnofabric including Siphonichnus, Teichichnus and Schaub-
Lower Cretaceous (Albian, continental slope), off Tanzania. d Large, cylindrichnus, overprinted by a relatively small P. surlyki (middle left).
undulating specimen of P. surlyki (outlined with arrow heads) with Lower Jurassic (Toarcian) Cook Formation (lower shoreface), Norwe-
partly pyritized (brown) tubes. Middle Jurassic (Callovian-Oxfordian) gian North Sea (well 34/2-4, ca. 3832 m)
Hugin Formation (marginal marine), Johan Sverdrup Field, Norwegian
Fig. 5.101 Parahaentzschelinia ardelia (=Rosselia rotatus) from (middle and right), showing the overall funnel-like shape, mud-lined
Permian shoreface to foreshore deposits from the Sydney Basin, tubes, and inclination to bedding. Scale bar = 3 cm. Modified after
Australia. Holotype of R. rotatus in vertical (upper left) and horizontal McCarthy (1979), © Paleontological Society, published by Cambridge
(lower left) section, as well as different burrows in vertical section University Press, reproduced with permission
5.20 Phoebichnus Bromley and Asgaard, modification (e.g. cementation). The entire burrow system
1972 may reach several decimeters in diameter, while the diameter
of individual burrows typically ranges between 1 and 2 cm
Morphology, Fill and Size: Phoebichnus is a horizontal, (Fig. 5.106).
star-like burrow system with a relatively thick, vertical Ichnotaxonomy: Beside the type ichnospecies P. tro-
central shaft (or boss), from which numerous straight, long choides from the Jurassic, three other ichnospecies have
burrows radiate (Fig. 5.105). The burrows have a wall with been added to this ichnogenus. P. minor Li et al., 1999 (from
discrete annuli and are actively filled with a meniscate the Lower Cambrian of China) and P. dushanensis Yang
backfill (Bromley and Asgaard 1972; Evans and McIlroy et al., 2004 (from the Carboniferous of China) remain poorly
2015). The central shaft is often subject to diagenetic defined and probably belong to other ichnogenera, while the
Fig. 5.102 Feeding activity of deep-sea tellinid bivalves Abra nitida,

which may result in Parahaentzschelinia. Zones of siphonal activity
shown as networks of abandoned canals. A. nitida feeds on surface
detritus and places both pseudofeces and feces on the sea floor. After
Wikander (1980), Bromley (1996), reprinted by permission of the
publisher (Taylor & Francis Ltd., http://www.tandfonline.com)
radiating burrows in P. bosoensis Kotake, 2003 (from the

Pleistocene of Japan) lack the diagnostic thick wall and are
filled with pellets.
Substrate: Phoebichnus is predominantly reported from
micaceous sandstone but also from mixed siliciclastic-
carbonate sediments (Joseph et al. 2012).
Appearance in Core: In core, Phoebichnus appears in the Fig. 5.103 Core section displaying the cast of a small bivalve (arrow)
and a series of emerging, funnel-shaped siphon traces above it. Lower
form of bundled burrows sectioned in various directions. Jurassic (Pliensbachian-Toarcian) Cook Formation (shallow marine,
Most common are cross sections more or less vertical to the well 34/5-1S, ca. 3648.65 m). Scale bar = 1 cm
burrow axis, which are accompanied by oblique sections
Fig. 5.104 Parahaentzschelinia

surlyki associated with
hummocky cross-stratified
sandstone in the Lower Jurassic
Neill Klinter Formation (shallow
marine), East Greenland. From
Dam (1990), republished with
permission of the Geological
Society of Denmark
5.20 Phoebichnus Bromley and Asgaard, 1972 111
Fig. 5.105 Bauplan and three-dimensional reconstructions of Phoeb- conveyed through Copyright Clearance Center, Inc. a Structure
ichnus trochoides. (a) and (b) from Bromley and Mørk (2000), showing vertical axis and the proximal parts of the radial branches.
republished with permission of Schweizerbart (www.schweizerbart.de/ b Part of a radial branch, showing the two-zoned backfill. c Central
series/zgp1). (c) and (d) based on serial grinding, from Evans and boss with radial burrows. d Radial burrows with outer wall and
McIlroy (2015), republished with permission of Wiley; permission laminated fill. Radial burrows are approximately 1 cm in diameter.
with an elliptical outline (Fig. 5.107). The radiating burrows particular crustaceans) may also be good candidates (Evans
are relatively large (commonly more than 1 cm in diameter) and McIlroy 2015).
and consist of a thick wall surrounding the actively filled Ethology: P. trochoides is interpreted as the feeding trace
core of the burrow. (fodinichnion) of a sessile deposit-feeding animal, which
Similar Trace Fossils: In outcrop, Phoebichnus could be is thought to live in the central shaft (domichnion) and
confused with other stellate burrows, e.g. Stelloglyphus. In producing the radiating feeding burrows (Bromley and
core, other large burrows with a thick wall or mantle and an Asgaard 1972).
active fill could resemble the radiating burrow segments of Depositional Environment: P. trochoides is a shallow-
Phoebichnus, for instance actively filled and thick-walled marine trace fossil with occurrences in shoreface and delta
Ophiomorpha as well as Macaronichnus. In addition, hori- deposits (e.g. Martin and Pollard 1996; McIlroy 2004;
zontal sections of large Siphonichnus may be confused with Morris et al. 2006; MacEachern and Bann 2008; Pemberton
individual Phoebichnus. Finally, root traces (fossilised root et al. 2012; Joseph et al. 2012) and on the shelf (Heinberg
systems) may be in the size range of Phoebichnus systems and Birkelund 1984; Pemberton and Frey 1984; Dam 1990;
and can produce similar patterns (Gregory and Campbell Bromley and Mørk 2000). The dependence of the producer
2003). on sediment rich in food particles suggests a low-energy
Producers: Vermiform organisms (such as Echiura) have environment (Heinberg and Birkelund 1984), an interpreta-
been assumed as the producers of Phoebichnus (Bromley tion supported by the frequent preservation of entire burrow
and Mørk 2000; Kotake 2003), although arthropods (in systems.
(a) (b)
(c) (d)
Fig. 5.106 Phoebichnus trochoides in outcrop. a Large burrow Same locality as in (a). Scale bar = 1 cm. c Dense ichnofabric mainly
system with a cemented central area (vertical shaft), from which resulting from overlapping burrow systems. Same locality as in (a).
numerous burrows radiate. Middle Jurassic (Callovian) sandstone, d Large stellate burrow system with cemented vertical shaft. Upper
Scarborough, Yorkshire, UK. b Part of a burrow system with the central Cretaceous (Turonian) sandstone, Cardium Formation, Seebe Dam,
area and emerging radials displaying the mantle and meniscate fill. Alberta, Canada. Scale bar = 15 cm. See Pemberton and Frey (1984)
Ichnofacies: Phoebichnus can be a common part of the 5.21 Phycosiphon Fischer-Ooster, 1858
Cruziana Ichnofacies. The “Phoebichnus Ichnofacies” pro-
posed by Heinberg and Birkelund (1984) has not found Morphology, Fill and Size: Phycosiphon is a small spreite
acceptance among subsequent workers. burrow consisting of repeated narrow, U-shaped lobes, each
Age: P. trochoides is a common trace fossil in Mesozoic enclosing a spreite at a millimetric to centimetric scale, and
(Triassic to Cretaceous) deposits (Evans and McIlroy branching regularly or irregularly from an axial spreite of
2015). similar width (Wetzel and Bromley 1994; Fig. 5.108). The
Reservoir Quality: The complex nature of Phoebichnus lobes typically consist of a mud-dominated marginal string
burrows with their actively created mantle and core indicates and a silty or sandy spreite (Fig. 5.109). The burrow systems
increased heterogeneity of the bioturbated substrate, con- are mainly parallel to bedding, although oblique and even
tributing to an overall decrease of potential reservoir vertical sections may occur too. Three-dimensional recon-
properties. structions of “phycosiphoniform” burrows suggest an
5.21 Phycosiphon Fischer-Ooster, 1858 113
(a) (b)
t t
(c) (d)
Fig. 5.107 Phoebichnus trochoides in sectioned core. Scale bars = 1 with a P. trochoides ichnofabric, consisting of a cluster of radiating
cm. a Heterolithic (silty) sandstone with moderate bioturbation and few burrows (center-right). Middle Jurassic (Bajocian) Ile Formation
burrows in cross section in the upper part (t). Middle Jurassic (shallow marine, prodelta), Njord Field area, Norwegian Sea (well
(Bajocian) Ile Formation (shallow marine, prodelta), Norwegian Sea 6407/10-2, ca. 3455.5 m). See also McIlroy (2004). d A cluster of
(well 6406/8-1, ca. 4388 m). b Highly bioturbated silty sandstone with P. trochoides burrows in cross and oblique sections. Lower to Middle
oblique burrow sections in the middle. Middle Jurassic (Bajocian) Ile Jurassic (Toarcian-Aalenian) Stø Formation (offshore), Snøhvit Field,
Formation (shallow marine, lower shoreface), Norwegian Sea (well Norwegian Barents Sea (well 7120/8-3, ca. 2205.5 m)
6406/8-1, ca. 4479 m). c Heterolithic and ripple-laminated sandstone
Fig. 5.108 Conceptual model showing the non-planar orientation of a twisted Phycosiphon burrow lobes (middle and right). From Bednarz
single Phycosiphon burrow lobe with the mantle and spreite shown as and McIlroy (2009), republished with permission of the Palaeontolog-
being transparent to facilitate viewing of the central mudstone strand. ical Association
Lobe parallel to the bedding plane (left) and possible variations of
(a) (b)
Fig. 5.109 Phycosiphon in outcrop. Scale bar = 1 cm. a Bedding- permission conveyed through Copyright Clearance Center, Inc. b Ver-
plane preservation of a burrow system with empty marginal tunnel tical face with various sections of Phycosiphon. Miocene Mount
(mud fill is weathered out) and enclosed spreite area. Eocene Grès Messenger Formation (deep marine, channel-levee system), sea cliff of
d’Annot Formation (deep marine, turbiditic), southeastern France. the Taranaki Peninsula, North Island, New Zealand
From Knaust et al. (2014), republished with permission of Wiley;
Fig. 5.110 Three-dimensional

reconstruction of a
phycosiphoniform burrow (core
with halo). Scale bar = 1 cm.
From Bednarz (2014)
5.21 Phycosiphon Fischer-Ooster, 1858 115
(a) (b)
(c) (d)
b Fig. 5.111 Phycosiphon in sectioned core. Scale bar = 1 cm. layer and is accompanied by Palaeophycus and Teichichnus. Lower
a Heterolithic sandstone with high degree of bioturbation, resulting in Jurassic (Pliensbachian-Toarcian) Cook Formation (offshore transition),
an ichnofabric dominated by Phycosiphon (dark muddy spots sur- Norwegian North Sea (well 34/5-1S, ca. 3654.5 m). c, d Totally
rounded with light sandy haloes), accompanied by Teichichnus and bioturbated silty sandstone with Phycosiphon concentrated in larger,
Schaubcylindrichnus. Upper Jurassic (Oxfordian) Heather Formation sand-filled burrows (Thalassinoides) and accompanied by Teichichnus
(shelf), Fram Field, Norwegian North Sea (well 35/11-9, ca. and Schaubcylindrichnus. Lower Jurassic (Pliensbachian to Bajocian)
2732.85 m). b Heterolithic sandstone with moderate bioturbation. Stø Formation (offshore transition), Iskrystall Discovery, Norwegian
Phycosiphon is concentrated within the fine-grained (muddy to silty) Barents Sea (well 7219/8-2, ca. 2986.5 m)
irregularly winding mud tube surrounded by a sandy halo Depositional Environment: Phycosiphon is a character-
(Naruse and Nifuku 2008; Bednarz and McIlroy 2009; istic component of offshore (shelf) to lower shoreface
Fig. 5.110). deposits, where it commonly occurs in isolation or in higher
Ichnotaxonomy: P. incertum is the only ichnospecies of ichnodiversity within siliciclastic successions (Goldring
Phycosiphon. The three-dimensional Anconichnus horizon- et al. 1991; Pemberton et al. 2012). It occurs in slope
talis is now regarded as a junior synonym of P. incertum deposits (e.g. Savrda et al. 2001), where it can be related to
(Wetzel and Bromley 1994). Bednarz and McIlroy (2009) slump deposits and act as a paleoslope indicator (Naruse and
prepared three-dimensional reconstructions of phycosi- Nifuku 2008). Phycosiphon is also common in deep-marine
phoniform burrows and realized morphological differences settings such as channel-levee complexes, where it occurs in
from the type material as redescribed by Wetzel and marginal channel-levee facies (Callow et al. 2013;
Bromley (1994). Fig. 5.136). Phycosiphon-producing organisms are among
Substrate: Phycosiphon typically occurs in silty or the first colonizers of event deposits which result from
fine-grained sandy, mainly siliciclastic deposits with a soft- storm, turbidity current and bottom current deposition
ground origin. It is more rarely reported from carbonates and
chalk.
Appearance in Core: The occurrence of mud-dominated
“strings”, often in high abundance, in combination with
reworked patches (spreite) of sand or silt, is a characteristic
feature of Phycosiphon in core (Fig. 5.111, see also
Figs. 5.19c, 5.129a and 5.179a). Because the spreite was
often formed slightly deeper in the sediment than the mar-
ginal burrow, these reworked patches often occur slightly
lowered in comparison with the associated mud-dominated
strings. Typical expressions of Phycosiphon in vertical sec-
tions are “… clusters of closely spaced spots or
comma-shaped dots and hooks filled with darker and finer
sediments surrounded by narrow (ca. 1 mm) pale mantles.
Generally, various orientations of the lobes result in a
chaotic arrangement of dark cores in the sections.” (Naruse
and Nifuku 2008).
Similar Trace Fossils: Nereites appears to be similar to
Phycosiphon (particularly in core sections), but differs from
it by the lack of the enclosed spreite and a commonly hor-
izontal course. In general, Phycosiphon is smaller than
Nereites. Phycosiphon could also be confused with Chon- Fig. 5.112 Reconstruction showing how multiple phases of foraging
drites, which shows dichotomous branching and lacks the by an unknown vermiform organism creates phycosiphoniform looped
sandy spreite. burrows composed of marginal tube and spreite. From Bednarz and
Producers: It is likely that the tracemakers of Phyco- McIlroy (2009), republished with permission of the Palaeontological
Association. Different shades of gray represent distribution of silt-sized
siphon are small, vermiform organisms of unknown affinity. (light gray) and mud-sized (dark gray) material. 1 Foraging organism
Ethology: Phycosiphon burrows are best explained as the creates feeding probes lateral to the marginal tube. 2 Successive probes
deposit-feeding activity of small vermiform organisms that are made until the organism has produced a marginal tube the length of
exploit the sediment for organic-rich matter (Wetzel 2010; its body. 3 Outer margin of the loop is produced by the organism
moving along previously produced probes. 4 Second loop is stared after
Izumi 2014; Fig. 5.112). the organism body is straight once again. 5 Completion of second loop
5.22 Planolites Nicholson, 1873 117
(Goldring et al. 1991; Wetzel and Uchman 2001; Wetzel Babcock 2012). The dimensions of Planolites can be quite
et al. 2008). variable and range from millimetric (e.g. Marenco and
Ichnofacies: Phycosiphon is characteristic of the Cruzi- Bottjer 2008) to centimetric.
ana, Zoophycos and Nereites ichnofacies. Ichnotaxonomy: In their review of the ichnogenus
Age: Phycosiphon is a common constituent of Mesozoic Planolites, Pemberton and Frey (1982) recognized only the
and Cenozoic deposits (Goldring et al. 1991) and is formed ichnospecies P. montanus, P. beverleyensis and P. annularis
today (Wetzel 1991, 2008). In their analysis, Callow and as valid. Since then, about ten more ichnospecies have been
McIlroy (2011) observed a dominance of phycosiphoni- introduced, some of which are poorly known and may turn
form ichnotaxa in the Mesozoic and Cenozoic, while out to be junior synonyms.
Nereites prevails in Paleozoic rocks. These authors specu- Substrate: Planolites is a characteristic component of
lated about a change in an ecological niche from Nereites- softgrounds and occurs in siliciclastic and carbonate (in-
dominated Paleozoic to Phycosiphon-dominated Mesozoic cluding chalk), fine- to medium-grained sediment.
and Cenozoic. Appearance in Core: In core, the elongate horizontal
Reservoir Quality: The effect of phycosiphoniform bur- tubes of Planolites have circular to elliptical sections that are
rows on shale hydrocarbon reservoir quality was evaluated actively filled but have no lining. Sand-filled Planolites often
by Bednarz and McIlroy (2012), who documented an occurs in mud-dominated lithologies and thus is easy to
improved reservoir capacity, permeability and fracturability. recognize (Fig. 5.115, see also Fig. 4.3). In contrast,
In conventional reservoirs, the reworked sandy zone (spreite) recognition of Planolites without significant differences in
in phycosiphoniform burrows may also lead to a slight substrate (e.g. sand-filled burrows within sandy host sedi-
increase of porosity and permeability. ment) can be difficult or impossible, and increasing biotur-
bation may just lead to a diffuse bioturbate texture
(Fig. 5.179b, d).
5.22 Planolites Nicholson, 1873 Similar Trace Fossils: Their simplicity leads to confusion
of Planolites burrows with numerous other trace fossils
Morphology, Fill and Size: Planolites is a simple, horizontal when studied in core. Many other cylindrical burrows are
to slightly inclined, cylindrical burrow without branching actively filled and have no lining, but are commonly bran-
and lining, and with an active (homogeneous) fill (Pember- ched, a fact that is often hard to test in core. Confusion with
ton and Frey 1982; Keighley and Pickerill 1995; Figs. 5.113 thinly lined Palaeophycus may occur, but this ichnogenus
and 5.114). In plan view, Planolites appears as straight to differs from Planolites by having a passive fill. Similarities
tortuous burrows (Fig. 5.178b). The cross section of the do also exist between Planolites and Macaronichnus, both
burrows is circular to elliptical, which is partly a function of ichnogenera sharing the same overall geometry. Although
the degree of compaction. An ichnospecies of Planolites Macaronichnus also has an active fill, its surrounding mantle
with both striation and annulation was described by Stanley differs from the smooth margin in Planolites.
and Pickerill (1994). Microbially mediated biomineralization Producers: Planolites is a relatively simple trace fossil
along the burrow margin is interpreted as agent in the pro- that can be produced by a wide range of organisms
moted preservation of Cambrian Planolites (Ahn and belonging to different phyla. Wormlike animals (such as
annelids, hemichordates and priapulids) are often interpreted
as producers of Planolites, but arthropods (e.g. crustaceans)
and molluscs (e.g. bivalves) are likewise able to produce
such traces. Regarding the long stratigraphic range of
Planolites it is likely that this trace fossil was constructed by
various types of animals over time.
Ethology: Planolites is interpreted as the product of
deposit-feeders (fodinichnia), which actively process the
sediment.
Depositional Environment: Planolites has been reported
from all aquatic environments in marine as well as non-
marine settings. It is a common element of shallow-tier
ichnofabrics.
Fig. 5.113 Planolites montanus in a reconstruction by Howard and Ichnofacies: Planolites is a facies-crossing element and
Frey (1984), republished with permission of Canadian Science
Publishing; permission conveyed through Copyright Clearance Center,
can be found in various ichnofacies. It is, however, a com-
Inc. Note the false branching due to overlap of two burrows. Scale mon constituent of the Cruziana Ichnofacies and also occurs
bar = 1 cm in the Nereites Ichnofacies (e.g. Buatois and Mángano
(a) (b)
(c) (d)
(e) (f)
(g) (h)
Fig. 5.114 Planolites from outcrop. Scale bars = 1 cm. a, b Specimen (2004a). e, f Planolites isp. in bioclastic limestone in vertical section
with numerous sand-filled P. montanus covered with mud. Upper (e) and on bedding plane (f). Lower Cretaceous (Barremian) of Cabo
Carboniferous paralic coal measure, western Germany. Senckenberg Espichel, Portugal. g, h Planolites isp. on bedding plane of calcareous
coll., Frankfurt Main (holotype; Richter 1937) in bedding-plane (a) and sandstone. Close association with Lockeia isp. and transitional forms
cross-section view (b). c P. montanus on bedding plane. Lower with Protovirgularia isp. suggest bivalves as producers. Middle
Ordovician sandstone-shale heterolithics (deltaic) near Ritland, Roga- Triassic (Anisian) Udelfangen Formation near Trier, western Germany.
land, southwestern Norway. d Same as (c), vertical section. See Knaust See Knaust et al. (2016) for geological setting
5.22 Planolites Nicholson, 1873 119
(a) (b) (c)
(d) (e)
Fig. 5.115 Planolites in sectioned core. Scale bars = 1 cm. a Dense (Pliensbachian) Åre Formation (marginal marine), Skuld Field, Nor-
occurrence of Planolites in sandstone. Middle Jurassic (Bajocian- wegian Sea (well 6608/10-14S, ca. 2702.5 m). d Planolites in silty
Bathonian) Hugin Formation (shallow marine, shoreface), Sleipner sandstone with coal fragments (debris). Middle Jurassic (Bathonian-
Vest Field, Norwegian North Sea (well 15/9-7, ca. 3565.4 m). Callovian) Hugin Formation (marginal marine), Norwegian Sea (well
b Planolites ichnofabric with large burrows in heterolithic sandstone. 15/6-4, ca. 10626.5′). e Small Planolites in silty mudstone with
Upper Jurassic (Oxfordian) Heather Formation (offshore, shelf), Fram lamination. Upper Cretaceous (Turonian-Santonian) Lange Formation
Field, Norwegian North Sea (well 35/11-11, ca. 2711.5 m). c Planolites (deep marine, basin floor), Norwegian Sea (well 6607/5-1, ca.
in heterolithic (silty) sandstone with coal seams. Lower Jurassic 3408.5 m)
2011). In continental deposits, Planolites occurs in the Reservoir Quality: Little is known so far about the
Scoyenia and Mermia ichnofacies. impact of Planolites on reservoir quality. Dawson (1981)
Age: Planolites is a cosmopolitan trace fossil observed recognized a reduction of reservoir quality when Plano-
from the Ediacarian (Alpert 1975; McCall 2006) throughout lites is present. This fact can be explained by the active
the Phanerozoic. fill of the burrows, which leads to increased
heterogeneity of the sediment. However, sand-filled bur- rarely been recognized in core, and even some of these
rows in muddy matrix may increase connectivity, reported occurrences remain disputable. However, a clear
although preferably in a horizontal direction due to the distinction between the two ichnospecies can be made in
orientation of Planolites. core. R. commune is best recognized in cross section, where
the two marginal tubes with a circular to elliptical cross
section and passive fill enclose the flattened spreite with an
5.23 Rhizocorallium Zenker, 1836 active fill in between (Figs. 5.116d and 5.118). Additional
confidence comes from the association of the burrow with
Morphology, Fill and Size: Rhizocorallium is a U-shaped fecal pellets. This is particular helpful in longitudinal sec-
spreite burrow in which a marginal tube encloses an actively tions, where only a thin stripe of reworked sediment can be
reworked area (spreite; Fürsich 1974c; Basan and Scott seen, sometimes with a pronounced bow-shaped spreite
1979). The burrow plane is horizontal to oblique to the (Fig. 5.116c). In contrast, R. jenense is inclined to various
bedding (Schlirf 2011). Burrow fill can be both active (e.g. degrees and has a dumbbell-like or pouch-shaped cross
spreite) and passive (e.g. marginal tube), or entirely passive section with a contrasting passive fill. Another aspect is the
(Knaust 2013). Delicate scratches can be preserved at the size of some well-developed R. commune, which can reach
burrows surface. Burrow width varies from a few millime- several centimeters in width and thus make it difficult to be
ters to several centimeters, while burrow length is typically recognized in size-limited core sections.
in the range of a few centimeters but occasionally can reach Similar Trace Fossils: The sparse and partly erroneous
several decimeters (Fig. 5.116). The ratio of tube diameter to reports of Rhizocorallium from core may be related to dif-
spreite width is comparatively low and approximately 1:2 to ficulties recognizing this trace fossil and potential confusion
1:5 (Häntzschel 1960; Fürsich 1974c; Worsley and Mørk with similar burrows. Other spreite burrows such as
2001). Diplocraterion and Zoophycos may be mistaken for Rhizo-
Ichnotaxonomy: A high lability in burrow morphology, corallium, but differ from it by their vertical orientation
size and orientation gave reason to the establishment of (Diplocraterion) or whorled morphology with lower
many ichnospecies of Rhizocorallium, which now are tube/spreite ratio (Zoophycos). Tisoa is another U-shaped
regarded as synonyms of only two valid ichnospecies burrow with passive fill, but differs from R. jenense by
(Knaust 2013; Fig. 2.5). R. commune has an active spreite having two narrow marginal limbs without an intervening
fill and often contains elliptical fecal pellets (Coprulus isp.), spreite; in addition it is relatively deep for its width, and is
while R. jenense is passively filled and intensively scratched commonly surrounded by a large concretion. Parts of
(Fig. 5.117). Uchman and Rattazzi (2016) propose the new backfilled horizontal burrows such as Taenidium and Scoli-
combination R. hamatum which, however, differs from cia may also resemple R. commune spreiten but are not
Rhizocorallium by its relatively small marginal tube and U-shaped and do not have a marginal tube.
widespread branching. Despite the abundant fill with fecal Producers: Three groups of organisms commonly qualify
pellets Coprulus oblongus, this form is better accommodated as potential producers of Rhizocorallium: decapod crus-
in the Zoophycos group, with its vertically extended whorls taceans, annelids and larval mayflies (Knaust 2013).
being strongly compacted. The attempt to transfer passively Although Rhizocorallium in some earlier studies was inter-
filled and scratched spreite burrows to the ichnogenus preted as the product of annelids, most workers have since
Glossifungites as done by Belaústegui et al. (2016a) is not favored crustaceans as tracemakers because of the presence of
supported by the evidence of R. jenense as its senior scratches (e.g. Seilacher 2007; Rodríguez-Tovar and Pér-
synonym. ez-Valera 2008; Neto de Carvalho et al. 2010). However,
Substrate: Rhizocorallium is known from siliciclastic and numerous groups of animals are known to produce scratches.
carbonate substrate alike. It occurs in soft and firm sub- Several lines of evidence suggest annelids (e.g. spionids,
strates, which results in the development of contrasting eunicids, terebellids) as producers of Rhizocorallium, an
behaviors that in turn are the ichnotaxobases for differenti- interpretation that is supported by constructional morphol-
ating the two ichnospecies (Fig. 2.5). R. commune is pre- ogy, accompanying features (e.g. fecal pellets), neoichno-
dominantly created in soft to stiff substrates (Fig. 5.116a), logical comparison and remains of the fossilized producers
while R. jenense is characteristic of firmgrounds preserved in situ (Knaust 2013). An interpretation of marine
(Fig. 5.116b). Rhizocorallium as a polychaete burrow is straightforward,
Appearance in Core: Although Rhizocorallium is a while the larvae of emerging mayflies probably account for R.
common trace fossil of worldwide distribution, it has only jenense produced in fluvial settings (Fürsich and Mayr 1981).
5.23 Rhizocorallium Zenker, 1836 121
(a) (b)
(c) (d)
(e) (f)
(g) (h)
b Fig. 5.116 Rhizocorallium in outcrop and in thin sections. Scale tube (left) completely filled with pellets. After Knaust (2013). d Vertical
bars = 1 cm except in (g) = 5 cm. (b), (c) and (d) republished with thin section of a horizontal R. commune displaying an actively created
permission of Elsevier; permission conveyed through Copyright Clear- spreite between the limbs of the passively filled U-shaped marginal
ance Center, Inc. a R. commune in Middle Triassic (Anisian) limestone tube. Lower Jurassic, Grimmen, Germany. After Knaust (2012b).
(type horizon) near Weimar, Thuringia, Germany. b R. jenense with e R. commune in Upper Triassic (Norian) sandstone (storm deposit) of
net-like, crossing and closely spaced scratches. Lower Triassic (Upper Deltaneset, Templefjorden, Svalbard. f Cross section of R. commune as
Buntsandstein, Pelsonian, Rhizocorallium Dolomite, type bed), Jena– in (e). g Sandstone bedding plane with gregarious R. commune. Upper
Ziegenhain, coll. Mägdefrau, Thüringer Landesanstalt für Umwelt und Jurassic (Kimmeridgian, shallow marine), coastal cliffs at Praia do
Geologie, Jena (TLGU 5035-701-202). After Knaust (2013). c Thin Salgado, western Portugal. h R. commune in bedding-plane view. Same
section of parts of R. commune as in (a), displaying the actively filled locality as in (g)
spreite with micritic fecal pellets (Coprulus oblongus) and the marginal
R. commune var. irregulare

Rhizocorallium jenense
- Long and winding, ± horizontal
- Subsurface trace, variably inclined
- Passive fill
- No fecal pellets
- Scratches net-like and closely spaced
- Suspension-feeding
- Firmground
- Polychaetes, crustaceans, insects
- Glossifungites Ichnofacies
R. commune var. auriforme
- Short, slightly inclined

R. commune problematica
- Vertically retrusive spreite
Rhizocorallium commune
- Surface or shallow sub-surface trace

R. commune uliarense - Active spreite fill
- Fecal pellets Coprulus isp.
- Trochospiral - Scratches ± parallel and widely spaced
- Deposit- and suspension-feeding
- Softground, partly firmground
- Polychaetes
- Cruziana Ichnofacies
Fig. 5.117 Classification of Rhizocorallium into two ichnospecies (R. varieties. From Knaust (2013), republished with permission of Elsevier;
commune and R. jenense). Morphological variability in R. commune is permission conveyed through Copyright Clearance Center, Inc.
captured by two ichnosubspecies, while size differences result in two
Ethology: Combined suspension- and deposit-feeding occurrence of R. commune var. irregulare in intertidal and
behavior is favored by most workers, although tests for shallow subtidal environments, whereas R. commune var.
gardening and storage behavior (cache) are inferred from auriforme is present in lagoonal environments and in deeper
oxygenated pyrite framboids found within the Rhizocoral- parts of the basin (Fig. 5.119). Furthermore, R. commune-
lium burrow (Zhang et al. 2016). associated fecal pellets, Coprulus bacilliformis, occur in
Depositional Environment: The two ichnospecies of inter- to supratidal environments and C. oblongus is evident
Rhizocorallium and their varieties may serve as good facies in intertidal and deeper environments (Knaust 2013).
indicators with the potential of supporting sedimentological R. jenense typically occurs along omission surfaces in high-
and paleontological interpretations (Knaust 2013). R. com- energy areas and is often documented along ravinement
mune typically occurs in shelf and nearshore environments surfaces exposed during marine transgressions. It is a com-
(e.g. Farrow 1966; Ager and Wallace 1970; Worsley and mon constituent of transgressive systems tracts (e.g. Knaust
Mørk 2001; Rodríguez-Tovar and Pérez-Valera 2008), and 1998; Knaust et al. 2012; MacEachern et al. 2012).
since the Triassic also in deep-marine deposits. R. jenense is Several studies have utilized Rhizocorallium as a cur-
known from shallow- and marginal-marine environments but rent indicator (Farrow 1966; Schlirf 2000; Worsley and
occurs in fluvial deposits as well (Fürsich and Mayr 1981). Mørk 2001; Rodríguez-Tovar and Pérez-Valera 2008;
Data from the Germanic Basin confirm the preferential Cotillon 2010), with its long axis running parallel or
5.23 Rhizocorallium Zenker, 1836 123
(a) (b)
s
T
s
s
s
B
s
s
(c) (d)
Fig. 5.118 Rhizocorallium commune in sectioned core. Scale bars = Formation (shallow marine), Sleipner Vest Field, Norwegian North Sea
1 cm. a Cross sections with passively filled (sandy) marginal tubes (well 15/9-8, ca. 3474.0 m). c Cross section with passively filled
connected by an actively filled (muddy) spreite (s) co-occurring with (sandy) marginal tubes connected by an actively filled (muddy) spreite.
Bornichnus tortuosus (B). Lower to Middle Jurassic (Aalenian- Lower Jurassic (Aalenian) Stø Formation (upper shoreface), Snøhvit
Bajocian) Stø Formation (shoreface), Snøhvit Field, Barents Sea (well Field, Barents Sea (well 7120/8-1, ca. 2120.4 m). d Weakly defined
7120/6-2 S, ca. 2577.5 m). b Several R. commune cross sections (arrow cross sections (arrow heads). Middle Jurassic (Bathonian-Oxfordian)
heads) are accompanied with a vertical spreite burrow corresponding to Hugin Formation (shallow marine), Gudrun Field, Norwegian North
Teichichnus zigzag (T). Middle Jurassic (Bathonian-Oxfordian) Hugin Sea (well 15/3-9T2, ca. 4497.2 m)
Fig. 5.119 Distribution of Rhizocorallium jenense and R. commune area). From Knaust (2013), republished with permission of Elsevier;
(and varieties) and their paleoenvironmental relationships in the permission conveyed through Copyright Clearance Center, Inc.
Middle Triassic (Lower Muschelkalk) of Thuringia, Germany (type
oblique to the inferred paleocurrent (or the line of the two expected that intense bioturbation with R. commune and
burrow openings perpendicular to it), and the distal their active spreite fill and frequent occurrence of muddy
(curved) end directed onshore. This pattern can be fecal pellets reduces the reservoir quality, whereas passively
obscured by a more random alignment in response to the filled R. jenense certainly would increase it.
exploitation of nutrient-rich ripple troughs by the trace-
maker. Many marine Rhizocorallium are documented as
tolerating a wide range of salinity from hypersaline to 5.24 Rosselia Dahmer, 1937
mesohaline water. R. commune occurs in various oxygen-
depleted deposits with dysoxic conditions and increasing Morphology, Fill and Size: Rosselia includes subvertical,
oxygenation of the sediment (Wignall 1991; Kotlarczyk bulbous- or spindle-shaped burrows with length varying from
and Uchman 2012). few centimeters to over 100 cm (Nara 2002). In most cases,
Ichnofacies: Rhizocorallium commune is a component of Rosselia is unbranched, although occasional side-branches
the Cruziana Ichnofacies, while R. jenense is part of the occur (Fig. 5.120a, c). The interior of the muddy burrow
widespread Glossifungites Ichnofacies. consists of numerous concentric laminae that are arranged in
Age: The ichnospecies R. commune is one of the an onion-like manner (Fig. 5.120b, e–g). The terminal bur-
longest-ranging fossils and is known from the Early Cam- row with passive fill and central or marginal position may be
brian (e.g. Fedonkin 1981; Clausen and Vilhjálmsson 1986; preserved in form of a narrow cylindrical shaft (Fig. 5.120c,
Orłowski 1989) to the Holocene (e.g. Winn 2006), while R. d). Contrast enhancement by early diagenetic iron mineral-
jenense first appeared after the end-Permian mass extinction ization (e.g. goethite, siderite) is common.
(Knaust 2013). Ichnotaxonomy: R. socialis is the most common ich-
Reservoir Quality: The effect of Rhizocorallium on nospecies of the ichnogenus Rosselia and probably the only
reservoir quality is little studied. However, it can be valid one. Other ichnospecies such as R. chonoides Howard
5.24 Rosselia Dahmer, 1937 125
(a) (b) (c)
(d) (e)
(f) (g)
Fig. 5.120 Rosselia in outcrop. Scale bars = 1 cm except in (d) and horizontal (e) section. Lower Ordovician Beach Formation
(b) = 5 cm. a, b Large Rosselia in limonitic preservation and (shoreface), Bell Island, Newfoundland, Canada. See Fillion and
embedded in silty deposits. Flower-like morphology with several Pickerill (1990). f Ichnofabric with vertically stacked R. socialis in
off-branches (a), and individual bulbous specimen displaying the vertical to slightly oblique section. Building stone of unknown age and
onion-like laminated interior (b). Upper Cretaceous (Campanian) origin, Ronda, southern Spain. g Part of R. socialis on a sandstone
Bearpaw-Horseshoe Canyon Formation (shoreface), near Drumheller, bedding plane showing the onion-like lamination with the terminal
Alberta, Canada. See Zorn et al. (2007). c Bulbous Rosselia in limonitic tube. Lower Devonian Taunusquarzit, near Rüdesheim, southern
preservation within coarse-grained sandstone. Middle Jurassic Scar- Germany, the type area of R. socialis. Senckenberg coll., Frankfurt
borough Formation (shoreface), coastal cliff near Scarborough, York- Main. See Dahmer (1937)
shire, UK. d, e R. socialis, crowded in a sandstone bed in vertical
(a) (b)
(c) (d)
5.24 Rosselia Dahmer, 1937 127
b Fig. 5.121 Rosselia in sectioned core. Scale bars = 1 cm. a Slightly Vestland Group (offshore transition in a restricted basin), Johan
off-axial sections of fusiform, mud-lined Rosselia cross-cutting a dense Sverdrup Field, Norwegian North Sea (well 16/2-13S, ca. 1942.0 m).
ichnofabric mainly consisting of Teichichnus zigzag and Chondrites. c Ripple-laminated sandstone with Rosselia overprinting a bioturbate
Middle Jurassic (Callovian) Hugin Formation (offshore transition), texture. Lower Jurassic (Pliensbachian) Cook Formation (shallow
Gina Krog Field, Norwegian North Sea (well 15/6-9S, ca. 3790.0 m). marine), Norwegian North Sea (well 34/5-1S, ca. 3659.5 m).
b Slightly bulbous Rosselia preserving an elongate basal tube. The d Spindle-shaped Rosselia in carbonates of the Upper Permian Khuff
truncation of the top of the burrow indicates an erosive bedding Formation, South Pars Field, Persian Gulf, Iran (well SP9, ca.
boundary that would otherwise not be recognized because of complete 3082.5 m). After Knaust (2014a), republished with permission of
bioturbation (Nereites)) of the host rock. Upper Jurassic (Oxfordian) EAGE
Fig. 5.122 Reconstruction of Rosselia tracemaker as a terebellid through Copyright Clearance Center, Inc. b Illustration after Pemberton
polychaete within its burrow. a Original interpretation from Nara et al. (2001), republished with permission of the Geological Association
(1995), republished with permission of Wiley; permission conveyed of Canada
and Frey, 1984 and R. rotatus McCarthy, 1979 are mor- internal structure makes Rosselia easy to recognize in core
phologically different and may not belong to Rosselia (Fig. 5.121), although confusion may arise from burrows
(Uchman and Krenmayr 1995; Knaust 2015a). with similar morphology. R. socialis with its bulbous swel-
Substrate: Rosselia is a common constituent of silici- ling in the upper part of the burrow is the most common
clastic softground deposits (Fig. 5.121a–c) but occasionally ichnospecies of Rosselia. Funnel-shaped burrows lacking the
also occurs in carbonates (Fig. 5.121d). internal lamination but exhibiting individual vermiform
Appearance in Core: The funnel-, bulb- or spindle-like burrows were attributed to R. chonoides, Howard and Frey
shape of the upper burrow part together with the laminated 1984. It remains unclear if this results from secondarily
Fig. 5.123 Burrow construction

of Rosselia socialis by successive
movements of a vermiform
animal. After Chamberlain
(1971), © Paleontological
Society, published by Cambridge
University Press, reproduced with
permission
reworked burrow fill or if this form belongs to a different Another muddy, concentrically laminated ichnogenus is
ichnogenus (Uchman and Krenmayr 1995). Cylindrichnus, which differs from Rosselia by its
Similar Trace Fossils: The ichnospecies R. chonoides bow-shaped morphology (Goldring 1996; Goldring et al.
Howard and Frey, 1984 and R. rotatus McCarthy, 1979 were 2002). Lingulichnus resembles Rosselia in having a lami-
originally included in Rosselia because of their overall nated, funnel-shaped dwelling chamber, which continuous
similarity with R. socialis, but differ from it by having a downwards into a pedicle trace (Zonneveld et al. 2007).
helicoidal and crescentric fill, respectively. This, in addition Lingulichnus typically shows an elliptical cross section and
to a funnel-shaped instead of bulbous morphology (in R. perhaps could account for less organized forms. The exten-
rotatus) makes those forms rather consistent with the sive shaft of Rosselia may lead to confusion with Skolithos
ichnogenus Parahaentzschelinia than with Rosselia where only partially exposed in core.
(Fig. 5.101). Asterosoma is similar to Rosselia by its active Producers: Terebellid polychaetes are the most likely
muddy, concentrically laminated fill. It differs from it by a producers of Rosselia (Nara 1995, 2002), although other
dominantly subhorizontal orientation and commonly polychaete worms (such as Sabellidae) and sea anemones
star-shaped morphology with individual spindles branching (Actiniaria) have also been considered (Figs. 5.122 and
off from a central area (Bromley and Uchman 2003). 5.123).
Fig. 5.124 Idealized model showing the development of stacked Rosselia socialis in response to changing deposition and erosion. After Frieling
(2007), republished with permission of Springer
5.25 Schaubcylindrichnus Frey and Howard, 1981 129
Ethology: Rosselia is interpreted as the dwelling structure Rosselia contributes to a reduction of the reservoir volume
(domichnion) of detritus-feeding polychaetes. Under rapidly and quality.
changing sedimentation and erosion, the tracemaker tries to
adjust such conditions and produces equilibrium traces
(equilibrichnia). 5.25 Schaubcylindrichnus Frey and Howard,
Depositional Environment: R. socialis is a common trace 1981
in inner shelf areas and typically occurs in lower to middle
shoreface deposits. It is also reported from marginal-marine Morphology, Fill and Size: Schaubcylindrichnus is a
environments, including tidal flats, tidal channels, deltas, U-shaped burrow system typically consisting of three parts:
embayments and lagoons, and flood-tidal deltas (Uchman an isolated single burrow or a bundle of thickly lined, often
and Krenmayr 1995; Carmona et al. 2008). Various studies interpenetrating tubes that were constructed one after
have associated R. socialis with high sedimentation rates and another, a feeding funnel connected to one end of the burrow
repeated erosion (e.g. Nara 1995, 2002; Campbell et al. system, and a fecal mound connected to the other end
2006; Frieling 2007; Netto et al. 2014; Campbell et al. (Löwemark and Nara 2010; Fig. 5.125). Complete tunnel
2016). In such settings, rapid sedimentation leads to verti- systems can reach several decimeters in length, while indi-
cally exaggerated growth of R. socialis, whereas subsequent vidual burrow diameters are typically in the range of a few
erosion events cut down into the spindle-like burrow parts millimeters to about 1 cm (Figs. 5.128 and 5.178a). Burrow
and leave behind funnel-shaped structures (Fig. 5.124). This lining can be around 1–3 mm thick and is typically of white
is common during marine transgressions, where R. socialis color. Burrow fill is passive.
has its highest abundance (Nara 2002), during riverine flood Ichnotaxonomy: Schaubcylindrichnus is now regarded as
events, when plenty of fine-grained particles are in suspen- monoichnospecific; the previously established ichnospecies
sion (Campbell et al. 2006), or during storm events (Netto S. freyi and S. formosus are morphological variants of the
et al. 2014; Campbell et al. 2016). In marginal-marine type ichnospecies S. coronus, and therefore junior synonyms
environments, size reduction may be prevalent in R. socialis of it (Löwemark and Nara 2010, 2013; Figs. 5.126, 5.127
communities as a response to lowered salinity (Frieling and 5.128). The inclusion of Palaeophycus heberti in
2007). Schaubcylindrichnus, as recently proposed by Evans and
Ichnofacies: Rosselia is a common constituent of the McIlroy (2016), is not based on the type material of the
Cruziana Ichnofacies. Given the preferred ethology of its former and therefore becomes void. In the past, Schaub-
producer as a suspension-feeder, it may also be associated cylindrichnus has sometimes been confused with Terebel-
with elements of the Skolithos Ichnofacies to which it can be lina, which however, refers to a large agglutinated
transitional. foraminifer (Miller 1995).
Age: Rosselia is reported from the Lower Cambrian Substrate: Schaubcylindrichnus commonly occurs in
(Silva et al. 2014) to the Holocene (Nara and Haga 2007). sandy and argillaceous (silty to muddy) substrates, more
Reservoir Quality: Because of its suspension-feeding rarely also in carbonates.
behavior, the Rosselia producer incorporates mud-rich sedi- Appearance in Core: Because of its thick lining as diag-
ment into its dwelling and builds a thick lining. This leads nostic feature, Schaubcylindrichnus is relatively easy to
to a local accumulation of external mud and therefore identify in core samples (Fig. 5.129). Typically, clusters of
Fig. 5.125 Schaubcylindrichnus coronus in Miocene offshore- bar = 10 cm. After Nara (2006), republished with permission of
transition deposits outcropping in Japan. Funnel-like and mound-like Elsevier; permission conveyed through Copyright Clearance Center,
structures are associated with the bow-shaped specimen. Scale Inc.
Fig. 5.127 Morphological variability of Schaubcylindrichnus, which

in the past gave reason for the erection of different ichnospecies that
now are regarded as junior synonyms of S. coronus. Individual tunnels
are referred to Palaeophycus heberti. From Löwemark and Hong
Fig. 5.126 Schaubcylindrichnus coronus in a reconstruction by
(2006), reprinted by permission of the publisher (Taylor & Francis Ltd.,
Howard and Frey (1984), republished with permission of Canadian
http://www.tandfonline.com)
Science Publishing; permission conveyed through Copyright Clearance
Center, Inc. Scale bar = 1 cm
Howard 1985, 1990). The intraspecific morphological vari-
burrows with a thick lining are cut in various directions and ability within S. coronus, such as the number of tubes within
thus burrow sections vary from circular to vertically flattened, individual specimens, may be indicators for distinguishing
elliptical and elongate. The characteristically light-colored distal from proximal position on an offshore-shoreface tran-
lining is an eye-catching feature, which hardly can be missed. sect (Löwemark and Nara 2013). An analysis performed by
Schaubcylindrichnus also occurs as a late component in Löwemark and Nara (2013) “… shows a distinct tendency for
complex ichnofabrics and overprints preexisting bioturbation higher tube numbers in the offshore facies where sediments are
(Figs. 4.3, 5.79d, 5.100h, 5.111a, d and 5.156d). characterized by higher silt/mud content, suggesting that S.
Similar Trace Fossils: The appearance of Schaubcylindr- coronus with higher tube numbers were constructed in a calm
ichnus is conspicuous even in core, owing to its thick and environment allowing longer dwelling periods. The increased
pronounced burrow lining. Single occurrences may be con- abundance of nested tubes in settings characterized by thin
fused with Palaeophycus. sand layers indicates that the nested tubes are a reparation
Producers: Nara (2006) and Löwemark and Nara (2010) response to erosional events destroying the feeding funnels at
posit an enteropneust worm as the probable producer of the sediment-water interface.”
Schaubcylindrichnus. Ichnofacies: Schaubcylindrichnus is a common con-
Ethology: A funnel-feeding behavior of a vermiform stituent of the Cruziana and Skolithos ichnofacies.
animal (e.g. enteropneust, polychaete) can be deduced Age: Schaubcylindrichnus occurs in strata at least from the
(Löwemark and Nara 2010; Kikuchi et al. 2016). Carboniferous to the Pleistocene (Löwemark and Nara 2010).
Depositional Environment: Schaubcylindrichnus occurs in Reservoir Quality: No detailed observations about the
a wide range of environments from nearshore to continental influence of Schaubcylindrichnus on reservoir quality are
slope (Frey and Pemberton 1991a; Löwemark and Nara 2010). available. The composition and architecture of the burrows
It is common in shallow-marine settings, where it occurs generally suggest little impact.
especially in lower shoreface to offshore deposits (Frey and
5.26 Scolicia de Quatrefages, 1849 131
(a) (b)
(c) (d)
Fig. 5.128 Schaubcylindrichnus in outcrop. Scale bars = 1 cm. a, d Sectioned surface of a glauconitic, cross-bedded sand within a fault
b Completely bioturbated sandstone with scattered Schaubcylindrich- zone (overturned section) displaying clusters of large Schaubcylindr-
nus as elite trace fossil. The relatively small size of these burrows seems ichnus. Lower Cretaceous Arnager Greensand Formation (storm-
to be related to oxygen deficiency within the sediment. Paleocene dominated shoreface) near Rønne, Bornholm, Denmark
Grumantbyen Formation (shelf) near Longyearbyen, Svalbard. c,
5.26 Scolicia de Quatrefages, 1849 of Scolicia can be outlined (Fig. 5.131): S. prisca, S. plana
and S. strozzii.
Morphology, Fill and Size: Scolicia includes simple (un- Substrate: Scolicia is common in well-sorted, fine-
branched), winding, meandering to coiling, bilobate or trilo- grained sandstone and siltstone.
bate backfilled burrows with two parallel sediment strings Appearance in Core: The appearance of Scolicia in core
along their lower surface and flattened oval cross section is quite diagnostic (Fig. 5.132). Longitudinal and oblique
(Uchman 1995, 1998). Burrow diameter is typically in the sections of the burrows show a densely meniscate or lamellar
range of less than 1 cm up to few centimeters. The traces are backfill, while burrows in oval to reniform cross sections
indefinitely long, capable of reaching at least several centi- differ from the host rock by their active fill and, in an optimal
meters to decimeters in length. situation, reveal two sediment strings at their base.
Ichnotaxonomy: Several trace fossils are now included as Similar Trace Fossils: Bichordites is an echinoid trace
preservational variants in the Scolicia group, namely Taphr- fossil similar to Scolicia but differs by bearing a central
helminthopsis, Laminites, Subphyllochorda and Taphr- bilobate inner zone (Demírcan and Uchman 2012;
helminthoida (Uchman 1995; Fig. 5.130). Three ichnospecies Fig. 5.130). In core, Scolicia may be confused with
(a) (c)
(b)
(d) (e)
b Fig. 5.129 Schaubcylindrichnus in sectioned core. Scale bars = 1 cm. Schaubcylindrichnus in cross section. Upper Jurassic (Oxfordian)
a Totally bioturbated sandstone with Phycosiphon and Cylindrichnus Heather Formation (offshore), Fram Field, Norwegian North Sea (well
accompanied by thickly lined Schaubcylindrichnus, partly compacted. 35/11-11, ca. 2623.9 m). d Highly bioturbated sandstone cross-cut by
Lower Jurassic (Pliensbachian-Toarcian) Ror Formation (offshore), several clusters of Schaubcylindrichnus. Upper Jurassic (Oxfordian)
Åsgard Field, Norwegian Sea (well 6506/12-I-2 H, ca. 4836.7 m). Heather Formation (offshore), Fram Field, Norwegian North Sea (well
b Completely bioturbated siltstone-sandstone with Teichichnus and 35/11-9, ca. 2732.9 m). e Laminated and weakly bioturbated siltstone
Nereites, overprinted with sand-lined Schaubcylindrichnus in the lower (above a sandy turbidite with Cylindrichnus) with abundant sand-lined
part and well-defined Schaubcylindrichnus in the upper part. Upper tubes of Schaubcylindrichnus. Upper Jurassic (Oxfordian) Heather
Jurassic (Oxfordian) Heather Formation (offshore), Fram Field, Nor- Formation (offshore), Fram H-Nord Field, Norwegian North Sea (well
wegian North Sea (well 35/11-9, ca. 2735.5 m). c Completely biotur- 35/11-15ST2, ca. 2970.5 m)
bated sandstone (partly siderite-cemented at top) with bundles of
Fig. 5.130 Characteristic

morphology of Scolicia and the
similar trace fossil Bichordites,
and preservational variants of
Scolicia. From Uchman (1995)
(a) (b)
(c) (d)
(e) (f)
(g) (h)
Fig. 5.131 Scolicia from the Eocene Grès d’Annot Formation (deep d S. plana. e S. strozzii. f Lower bedding-plane with winding S. plana
marine, turbiditic), southeastern France (a–e) and in the Miocene along the sand-mud interface. g Vertical section with Scolicia isp. in
Mount Messenger Formation (deep marine, channel-levee complex) in longitudinal (left) and cross section (right) atop a laminated sandstone
sea cliffs of the Taranaki Peninsula, North Island, New Zealand (f–h). bed. Note the two characteristic sediment strings at the base of the cross
Scale bars = 1 cm except in (f) = 10 cm. (b), (d) and (e) from Knaust section. h Thin-bedded, fine-grained sandstone in vertical section with a
et al. (2014), republished with permission of Wiley; permission dense Scolicia ichnofabric
conveyed through Copyright Clearance Center, Inc. a, b S. prisca. c,
(a) (b)
(c) (d)
S N
Z
S
N
b Fig. 5.132 Scolicia in sectioned core. Scale bars = 1 cm. a Mud- Scolicia in longitudinal section. Upper Cretaceous (Campanian) Nise
stone-sandstone alternation (inclined) with cross sections of Scolicia Formation (deep marine, basin floor), Norwegian Sea (well 6607/5-2,
occurring in the upper part of the thin sandstone layers, where organic ca. 4186.5 m). d Mudstone-sandstone alternation with cross-bedding
sediment is concentrated within ripple troughs. Note the characteristic and intense bioturbation. Nereites (N) in the background is overprinted
sediment strings at the base of the burrows (arrows). Lower Cretaceous by large Scolicia (S), resulting from the exploitation of the sand layers
(Albian, deep marine, channel-overbank), off Tanzania. b Longitudinal by its producer. All other traces are cross-cut by mud-filled Zoophycos
sections of Scolicia in heterolithic turbiditic sandstone. Upper Creta- (Z). Upper Cretaceous (Campanian) Nise Formation (deep marine, fan
ceous (Turonian-Santonian) Kvitnos Formation (deep marine), Aasta fringe), Aasta Hansteen Field, Norwegian Sea (well 6707/10-1,
Hansteen Field area, Norwegian Sea (inclined well 6707/10-2A, ca. 2974.3 m). After Knaust (2009b), republished with permission of
4462.8 m). c Heterolithic intercalation of turbiditic sandstone and Elsevier; permission conveyed through Copyright Clearance Center,
hemipelagic mudstone showing several partly deformed specimens of Inc.
Taenidium, based on the meniscate backfill. However, Depositional Environment: Scolicia is common in
Taenidium is commonly smaller than Scolicia, has less deep-marine deposits belonging to the Nereites Ichnofacies
regularly packed menisci and lacks the diagnostic sediment and in shallow-marine environments assignable to the
strings at the base. Subhorizontal parts of the spreite burrow Cruziana Ichnofacies. Given its preference for sandy to silty,
Zoophycos and Lophoctenium may also resemble Scolicia, organic-rich sediment, Scolicia is a common element of
but belong to a complex three-dimensional burrow system. inner levee deposits within deep-marine channel-levee
Individual Zoophycos spreiten are less regularly packed than complexes (Callow et al. 2013). Similarly, Scolicia is very
Scolicia, are smaller and lack sediment strings. common in proximal fan and related environments but rare
Producers: On the basis of modern analogs and experi- in more distal parts (Heard and Pickering 2008). Scolicia
mental work, irregular echinoids (e.g. spatangoids, heart occurs in a shallow-tier position and therefore is subject to
urchins) are known to produce Scolicia-like traces be destroyed by subsequent burrowers (Figs. 5.9b, 5.135
(Figs. 5.133 and 5.134). and 5.136).
Ethology: Scolicia results from the deposit-feeding (fo- Ichnofacies: Scolicia is a typical element of the deep-
dinichnial) activity of irregular echinoids. marine Nereites and the shelfal Cruziana ichnofacies.
Fig. 5.133 Scolicia in the

Miocene Mount Messenger
Formation (deep marine,
channel-levee complex), one
burrow preserving its producer,
an irregular echinoid. Sea cliffs of
the Taranaki Peninsula, North
Island, New Zealand. Scale
bar = 1 cm
Fig. 5.134 Ichnological

reconstruction of the burrow of
Echinocardium cordatum in
transverse section (center) and
longitudinal section. The latter
shows two funnels to the surface
for deposit-feeding, a functional
one and an abandoned, collapsed
one. Reorientation of
non-spherical sediment grains by
the burrowing activity of the
animal produces a meniscate
backfill. From Bromley and
Asgaard (1975), republished with
permission of the Geological
Society of Denmark
Fig. 5.135 Tiering diagram

with distribution of characteristic
biogenic sedimentary structures
and traces, based on observations
from the modern deep sea.
Note the occurrence of Scolicia
in the upper tier. Scale
bar = 10 cm. Modified after
Wetzel (1981), republished
with permission of Schweizerbart
(www.schweizerbart.de/
9783443190347)
Fig. 5.136 Laminated

heterolithic sandstone with
Phycosiphon. The upper tier of
the ichnofabric is totally
bioturbated with Scolicia, which
leads to a homogenization of the
sediment and thus contributes to
increased reservoir properties.
Parts of the upper tier with
Scolicia are in turn reburrowed by
Phycosiphon. Miocene Mount
Messenger Formation (deep
marine, channel-levee complex),
sea cliffs of the Taranaki
Peninsula, North Island, New
Zealand. Scale bar = 1 cm
Age: Echinoid-produced Scolicia are known since the 1984; Retallack 2001; Fig. 5.137). Faint peristaltic annu-
Jurassic (Seilacher 1986; Fu and Werner 2000), while Paleo- lation of the burrow wall can occur. Burrows are unbran-
zoic Scolicia (e.g. Benton and Gray 1981; Bjerstedt 1988; ched with a slightly varying diameter, typically between 5
Buckman 1992) must have been produced by other organisms and 20 mm.
(such as slug-like animals). Many such Paleozoic Scolicia
have relatively simple morphology and composition com-
pared to the complex endichnial Scolicia from the Mesozoic
onwards (Buatois and Mángano 2004).
Reservoir Quality: Sandy to silty, laminated sediment of
heterolithic composition is the target of irregular echinoids
and becomes homogenized into sand-rich sediment due to
the deposit-feeding activity (Fig. 5.136). This happens at a
broad scale and this effect is amplified by the presence of
relatively large burrows and a dense bioturbate texture
(ichnofabric). Therefore, Scolicia bioturbation may increase
reservoir properties.
5.27 Scoyenia White, 1929
Morphology, Fill and Size: Scoyenia refers to horizontal

to inclined burrow elements with faint and irregular lining, Fig. 5.137 Structure and morphology of Scoyenia gracilis. Scale
longitudinal striation, and a conspicuous meniscate fill, bar = 1 cm. After Bromley and Asgaard (1979), republished with
permission of Elsevier; permission conveyed through Copyright
although some portions of a burrow may show transition Clearance Center, Inc.
to passive fill (Bromley and Asgaard 1979; Frey et al.
5.27 Scoyenia White, 1929 139
(a) (b)
(c) (d)
Fig. 5.138 Scoyenia gracilis in outcrop. Scale bars = 1 cm. Sand- (fluvial), near Cracow, southern Poland. c, d Permian Cutler Formation
stone bedding planes with tunnels and shafts displaying internal backfill (alluvial fan), 191 Canyon north of Moab, Utah, USA. Note association
and external scratches. a Eocene Aspelintoppen Formation (fluvial), with paired burrow apertures likely to be openings of Arenicolites
Brongniartfjellet, Svalbard. b Lower Triassic Buntsandstein Group
Ichnotaxonomy: Only two ichnospecies have been lining typically consists of mud but might be poorly devel-
described so far, of which the type ichnospecies S. gracilis is oped or preserved. The structured meniscate fill is diagnostic
the most common (Fig. 5.138), whereas S. beerboweri and commonly includes fragments of sediment with con-
seems to be restricted to Ordovician paleosols (Retallack trasting color.
2001; Fig. 5.151). Similar Trace Fossils: Taenidium (particularly T. bar-
Substrate: Scoyenia is often encountered in fine-grained retti) is the most similar trace fossil to Scoyenia
sandstone and siltstone with muddy intervals but may also (e.g. S. gracilis) and may be confused with it in core.
occur within calcareous successions, particularly those However, Scoyenia has a striated wall (which is difficult to
related to paleosols, such as horizons with caliche. recognize in core) and typically shows a thin mud lining
Appearance in Core: The relatively large diameter and that is rarely present in Taenidium. The average diameter
the appearance of the burrows make Scoyenia comparatively of Scoyenia is larger than that of Taenidium. Most sig-
easy to recognize in core, although the striated wall is nificantly, the meniscate fill of Scoyenia appears to be
impossible to be proven in sections (Fig. 5.139). Horizontal more disorganized into small pieces, while of Taenidium is
and shallowly inclined burrows are dominant. The faint more homogeneous.
(a) (b)
(c)
Fig. 5.139 Scoyenia gracilis in sectioned core. Scale bars = 1 cm. Formation (fluvial overbank), Snorre Field, Norwegian North Sea
a Heterolithic sandstone with oblique and horizontal, actively filled (well 34/7A-9H, ca. 2784.5 m). c Iron-stained argillaceous deposit with
parts of burrows. Eocene Aspelintoppen Formation (fluvial overbank), burrows displaying packets with discontinuous menisci. Upper Triassic
Brongniartfjellet, Svalbard (Sysselmannbreen well BH 10-2008, ca. (Norian-Rhaetian) Lunde Formation (paleosol), Snorre Field, Norwe-
85.5 m). b Ripple-laminated sandstone with thinly lined and actively gian North Sea (well 34/7-1, ca. 2481.7 m)
filled parts of burrows. Upper Triassic (Norian-Rhaetian) Lunde
5.28 Siphonichnus Stanistreet et al., 1980 141
Producers: Arthropods such as insects (e.g. beetles) and S. lepusaures and S. sursumdeorsum as new ichnospecies. S.
millipedes have been considered as producers of Scoyenia lepusaures and S. sursumdeorsum are probably cross sec-
(e.g. Frey et al. 1984; Retallack 2001; Hasiotis 2010). tions of Teichichnus zigzag and are better accommodated
Ethology: Scoyenia are probably multipurpose feeding, within that ichnogenus. Siphonichnus is the eponym of the
dwelling and breeding burrows (Retallack 2001). More ichnofamily Siphonichnidae, which encompasses burrows of
inclined burrows have been inferred to be escape structures varying morphology consisting of one or more subvertical
(Hubert and Dutcher 2010). tube(s) with passive fill and active lining or mantle (Knaust
Depositional Environment: Scoyenia is a continental 2015a). In addition to Siphonichnus, it includes the ichno-
trace fossil typical of alluvial, lacustrine and fluvial deposits, genera Laevicyclus, Parahaentzschelinia, Scalichnus and
such as paleosols, lakes and overbank deposits. It is often Hillichnus (Fig. 5.141).
related to high soil moisture and inferred to occur in wet Substrate: Siphonichnus is reported from diverse types
seasonal to wet climates (Hasiotis 2010). of substrate including mudstone, siltstone, sandstone and lime-
Ichnofacies: The Scoyenia Ichnofacies was originally stone. It often occurs in variously heterogeneous sandy
designed as almost an afterthought of the marine ichnofacies substrates.
(Seilacher 1967) but now is regarded to be more differenti- Appearance in Core: Although originally described as
ated (see Bromley 1996; Buatois and Mángano 2011; Mel- vertical, Siphonichnus is an unbranched trace fossil that can
chor et al. 2012). also have oblique and horizontal components. It consists of a
Age: Scoyenia is known from the Ordovician (Retallack wall with tight lamination (meniscate backfill) and a central
2001) to the Holocene (Hasiotis 2010). part, which is occupied by a well-defined tube with passive fill.
Reservoir Quality: The sandy fill of Scoyenia burrows In core, these cylindrical burrows appear as elongate, ellipti-
makes them a good conductor for fluids and gas, especially cal or circular sections (Fig. 5.142; see also Fig. 5.100f, h).
if occurring in muddy host sediment. The heterogeneous Similar Trace Fossils: Varying morphology, orientation
composition of the meniscate fill may lead to a slight and size of S. ophthalmoides may give rise to confusion with a
reduction of this property. number of trace fossils being partly similar. First, the vertical
expression of S. ophthalmoides may resemble Skolithos
(Fig. 5.146g) or even Trypanites, particularly in cases where
5.28 Siphonichnus Stanistreet et al., 1980 the lamination of the mantle is poorly visible. For the same
reason, horizontal parts of S. ophthalmoides—as observed in
Morphology, Fill and Size: Siphonichnus includes vertical, core—may appear as Palaeophycus and Macaronichnus. This
oblique or horizontal cylindrical burrows with a linear, situation is further complicated because S. ophthalmoides
winding or bow-shaped morphology and a circular to oval typically co-occurs with Skolithos and Palaeophycus, all of
cross section (Knaust 2015a; Fig. 5.140). It is characterized which may have been produced by a similar tracemaker. The
by a laminated meniscate mantle (active fill), which is lingulide-produced Lingulichnus is probably the trace fossil
penetrated by a homogeneous core (passive fill). The most similar to Siphonichnus. It differs from Siphonichnus by
diameter of the complete burrow is typically in the range of a its circular instead of elliptical cross section, and the internal
centimeter, although much smaller burrows occur as well. lamination is partly penetrated by the passively filled pedicle
Their length can be several decimeters. trace in a less consistent manner. The innermost burrow (core)
Ichnotaxonomy: S. ophthalmoides (Jessen 1950) from the of S. ophthalmoides seems to be wider than the pedicle trace of
Carboniferous of Germany is the senior synonym of S. Lingulichnus, and only preserves a relatively small portion of
eccaensis (Knaust 2014b, 2015a). It is currently regarded as the outer laminated burrow (mantle). S. ophthalmoides has
the only ichnospecies of Siphonichnus, although Zonneveld also been confused with Diplocraterion and, more rarely, the
and Gingras (2013) suggested the inclusion of Scalichnus mantle in S. ophthalmoides could also be mistaken as the mud
phiale within the ichnogenus Siphonichnus and established lining in Ophiomorpha.
(a) (b)
(c) (d)
Fig. 5.140 Siphonichnus ophthalmoides in outcrop. Scale bars = 1 cm. burrows revealing core and mantle on a limestone bedding plane.
From Knaust (2015a), republished with permission of Elsevier; Middle Triassic (Anisian) Meissner Formation (Upper Muschelkalk,
permission conveyed through Copyright Clearance Center, Inc. a Vari- carbonate ramp), Troistedt near Weimar, Thuringia, Germany. d Single
ably oriented burrows with respect to bedding within micaceous, specimen of S. ophthalmoides with a pyritized fill within the core. Late
cross-bedded sandstone of nearshore origin. Lower Jurassic (Hettan- Miocene Mount Messenger Formation (deep marine, slope fan), coastal
gian) Höganäs Formation (nearshore), Helsingborg, southern Sweden. section north of New Plymouth, North Island, New Zealand
b Bedding-parallel burrow part. Same locality as in (a). c Winding
Fig. 5.141 Line drawings of ichnogenera included in the ichnofamily lining or mantle. From Knaust (2015a), republished with permission of
Siphonichnidae and their morphological characteristics, all of them Elsevier; permission conveyed through Copyright Clearance Center,
having one or more subvertical tube(s) with passive fill and active Inc.
5.28 Siphonichnus Stanistreet et al., 1980 143
(a) (b)
(c) (e)
(d)
b Fig. 5.142 Siphonichnus ophthalmoides in sectioned core. a, b and (marginal marine), Åsgaard Field, Norwegian Sea (well 6506/12-K-3H,
e from Knaust (2015a), republished with permission of Elsevier; ca. 4459.0 m). c Bowed burrow in longitudinal section. Lower Jurassic
permission conveyed through Copyright Clearance Center, Inc. Scale (Aalenian) Stø Formation, Snøhvit Field, Norwegian Barents Sea (well
bars = 1 cm. a Heterolithic sandstone with a crowded S. ophthalmoides 7120/8-2, ca. 2097.75 m). d Variably oriented burrows displaying a
ichnofabric consisting of subvertical, inclined and horizontal, partly thick mantle. Lower Jurassic (Aalenian) Stø Formation, Snøhvit Field,
bowed burrows with concave-downwards laminae in the mantle. Lower Norwegian Barents Sea (well 7120/6-2 S, ca. 2583.3 m). e Dense S.
Jurassic (Sinemurian-Pliensbachian) Tilje Formation (marginal marine), ophthalmoides ichnofabric with total bioturbation and discrete burrow
Norwegian Sea (well 6607/12-3, ca. 4217.35 m). b Small and segments trending in all directions. Lower Jurassic (Hettangian-
elongated S. ophthalmoides together with probable casts of the Sinemurian) Nansen Formation, Norwegian North Sea (well 25/10-
producing bivalves (c). Lower Jurassic (Pliensbachian) Tilje Formation 11T2, ca. 4283 m)
Fig. 5.143 Modern bivalve burrows in response to deposition. From Reineck (1958), republished with permission of Schweizerbart (www.
schweizerbart.de/home/senckenberg). a No deposition. b Rapid sedimentation. c Erosion. d Very slow sedimentation
Producers: Contemporary Siphonichnus-like traces are Gingras et al. 2012a; Fig. 5.142b). The relatively long
produced by bivalves adjusting their position within the burrow parts, their wide range of orientations, and the
sediment (Reineck 1958; Fig. 5.143). In some cases, almost perfectly centered core within the mantle are fea-
the cast of the bivalve shell or its cavity is preserved at tures which make other producers than bivalves possible
the base of Siphonichnus (Dashtgard and Gingras 2012; (e.g. arthropods).
5.29 Skolithos Haldeman, 1840 145
Ethology: Siphonichnus is the dwelling trace (domich- Reservoir Quality: Siphonichnus burrows are mainly
nion) of a suspension- and deposit-feeder, such as a bivalve. sand-filled and vertically oriented and therefore can lead to a
It is able to adjust its position with respect to erosion and slight improvement of the vertical connectivity.
deposition (equilibrichnion). The highly variable orientation
of Siphonichnus burrows apparently results from the activity
of a (temporarily?) mobile infaunal tracemaker for the pur- 5.29 Skolithos Haldeman, 1840
pose of locomotion and, at least partly, deposit-feeding.
Therefore, a combined sedentary lifestyle resulting in more Morphology, Fill and Size: Skolithos is a simple, subvertical
or less vertically oriented burrows, and mobile lifestyle cylindrical tube with or without lining and passive fill
resulting in burrows with variable directions of the producer (Alpert 1974; Figs. 5.144 and 5.145). A funnel-shaped
can be inferred. aperture at the top may be developed or preserved (Schlirf
Depositional Environment: Siphonichnus can be regarded 2000). The burrow diameter ranges from millimetric to
as an indicator of shallow-marine and marginal-marine
environments, often related to fluctuating salinity and fresh-
Fig. 5.144 Sketch of Skolithos
water influx (Knaust 2015a). Its distribution ranges from linearis as produced by a modern
proximal offshore and shoreface to deltaic, estuarine and Phoronida (in position) in soft
lagoonal environments. S. ophthalmoides is a good example sediment. Note the sediment
particles adhering to the tube. Not
of an early application of trace fossils in ichnostratigraphy
to scale. After Emig (1982),
and the reconstruction of paleoenvironments in connection republished with permission of
with industrial coal mining. The assumed producer, an Elsevier; permission conveyed
endobenthic marine bivalve with tolerance for freshwater through Copyright Clearance
Center, Inc.
influence and salinity fluctuations, has rapidly colonized the
ecological niche along the paleoshoreline with brackish
influence. This opportunistic colonization resulted in the
abundant occurrence of the “eye”-like burrows (as seen in
cross section), while other macrofossils were too sparse for
quick characterization of the interval. The importance of S.
ophthalmoides for the correlation of coal seams in the mines
of western Germany on the basis of marine intervals, which
are accompanied at their base and top by the “eye”-shale
(Augenschiefer), was realized by early coal investigators
(e.g. Jessen 1950). Later workers have applied S. ophthal-
moides as a facies indicator of marginal-marine deposits as
well as in association with deposits on the lower delta plain
(e.g. interdistributary bay/lagoon) and delta-fed submarine
fans. Siphonichnus nicely reflects the response of its trace-
making bivalve to changing sediment supply. McIlroy (2004)
recognized S. ophthalmoides in situations with tidal-channel
abandonment and delta-lobe abandonment, when a sudden
reduction in sediment supply occurs. S. ophthalmoides is a
common constituent of intertidal and saltmarsh deposits.
Ichnofacies: Siphonichnus belongs to the Skolithos Ich-
nofacies and occurs as accompanying constituent in the
Cruziana Ichnofacies.
Age: Siphonichnus is recorded from the Late Devonian
(e.g. Angulo and Buatois 2012) to Holocene (e.g. Gingras
et al. 2008).
(a) (b)
(c) (d) (e)
(f) (g)
Fig. 5.145 Skolithos in outcrop. Scale bars = 22 cm (a) and 1 cm section (d). Cliff near Arnager, Bornholm, Denmark. e Thick, passively
(b)–(g). a, b S. linearis from its type locality, Chickies Rock, near sand-filled S. verticalis in modern beach deposits. North Island, New
Lancaster, Pennsylvania. A thick sandstone bed (tidal-dune set) is Zealand. f, g Sandstone with trough cross-bedding and abundant
penetrated by numerous long burrows (a), which show the diagnostic burrows. Lower Ordovician Tosna Formation (high-energy subtidal
lining (b). c, d Erratic boulder of Lower Cambrian sandstone completely nearshore environment). Tosna River bank near Nikolskoe village, St.
pierced with burrows (piperock) in longitudinal section (c) and cross Petersburg region, NW Russia (see Dronov and Mikuláš, 2010)
5.29 Skolithos Haldeman, 1840 147
(a) (b) (c)
(f)
(e)
(d)
(h)
(g) (i)
b Fig. 5.146 Skolithos in sectioned core. Scale bars = 1 cm. a Lami- erosion surface. Lower Jurassic (Hettangian) Statfjord Formation
nated micritic limestone with relatively large, deep, sand-filled burrows (fluvial to marginal marine, mixed tidal flat). Johan Sverdrup Field,
originating from an erosion surface. Upper Permian carbonates Norwegian North Sea (well 16/2-17S, ca. 1925.85 m). f Sandstone with
(carbonate platform with tidal flat). South Pars Field, Persian Gulf, a dense cluster (pipe rock) of Skolithos/Siphonichnus cut slightly
Iran (well SP-9, ca. 3082.9 m). From Knaust (2009a), republished with obliquely to their vertical extent. Lower Jurassic (Pliensbachian to
permission of GulfPetroLink. b Laminated micritic limestone with Bajocian) Stø Formation (offshore transition), Iskrystall Discovery,
sand-filled burrows. Upper Permian carbonates (shoal). South Pars Norwegian Barents Sea (well 7219/8-2, ca. 2973.5 m). g Intensely
Field, Persian Gulf, Iran (well SP-9, ca. 2909.75 m). After Knaust bioturbated sandstone with relict bedding preserved (inclined well). The
(2009a, 2014a), republished with permission of GulfPetroLink and dense Ophiomorpha ichnofabric is cross-cut by few Skolithos in the
EAGE. c Bioturbated sandstone with discrete S. linearis (lower left). center and upper right. Middle Jurassic (Callovian) Hugin Formation
Middle Jurassic (Bathonian) Hugin Formation (mixed tidal flat within (upper shoreface). Gina Krog Field, Norwegian North Sea (well 15/5-7,
an estuarine bayhead delta). Gudrun Field, Norwegian North Sea (well 3874.2 m). h Silty sandstone with numerous long burrows. Middle
15/3-9T2, ca. 4493.0 m). d Sandstone-siltstone alternation with thin, Jurassic (Bajocian-Bathonian) Tarbert Formation (marginal marine,
mud-filled Skolithos associated with sparse Polykladichnus and Areni- tidal flat), Valemon Field, Norwegian North Sea (well 34/11-B-13, ca.
colites in the upper part of the siltstone. Lower Jurassic (Pliensbachian) 4594.5 m). i Laminated sandstone with cryptic bioturbation and two
Tilje Formation (tidal flat). Åsgard Field, Norwegian Sea (well mud-filled Skolithos (left), accompanied by Ophiomorpha (right).
6506/12-K-3H, ca. 4537.85 m). e Cross-bedded sandstone with plant Upper Cretaceous (Maastrichtian) Springar Formation (deep marine,
debris and large sand-filled burrows penetrating the substrate from an lobe). Norwegian Sea (well 6604/10-1, ca. 3647.95 m)
centimetric and its length from a few millimeters to several 2012a). In contrast, Trichichnus is characterized by a high
decimeters. The length/diameter ratio can be used to dis- length/width ratio and occasional branching; moreover, it is
criminate Skolithos from morphologically similar trace commonly pyritized. Very thin Siphonichnus ophthal-
fossils. moides may also be mistaken as Skolithos (e.g. Bromley
Ichnotaxonomy: Skolithos is sparse in morphological 1996; Gerard and Bromley 2008). Furthermore, burrow
features. About 20 ichnospecies have been attributed to this parts of other trace fossils such as shafts of Thalassinoides
ichnogenus based on criteria such as morphology, burrow and Ophiomorpha may easily be confused with Skolithos if
wall and size, although some of them do not fit the ichno- the three-dimensional context remains unclear (Fig. 5.146).
generic diagnosis (such as helicoidal and branched forms) Producers: A wide range of organisms is known to
and therefore must be excluded. In addition, ichnospecies produce Skolithos-like traces (Fig. 2.3). Most appropriate are
originally attributed to other ichnogenera such as Tigillites various groups of worms such as priapulids and polychaetes,
and Sabellarifex are now regarded as junior synonyms of as well as phoronids, which may account for many Paleozoic
Skolithos (Alpert 1974; Fillion and Pickerill 1990; Schlirf and also younger Skolithos (e.g. Fenton and Fenton 1934;
2000; Schlirf and Uchman 2005). The ichnospecies S. line- Emig 1982; Sundberg 1983; Dashtgard and Gingras 2012;
aris, S. verticalis and S. annulatus are among the most Figs. 5.144 and 5.147). Another group comprises crus-
important forms of Skolithos. Despite repeated attempts, the taceans (e.g. amphipods), which are known to produce
ichnogenus Skolithos is in need of ichnotaxonomic revision. Skolithos-like traces today (e.g. Dashtgard and Gingras
Substrate: Skolithos occurs in soft (sandy and muddy) 2012). Anthozoa (sea anemones) may be also capable of
substrates, and subordinately in firm siliciclastic and car- producing Skolithos-like traces (e.g. Hertweck 1972). In
bonate sediments. addition, continental Skolithos can be produced by plant
Appearance in Core: Skolithos is relatively easy to recog- roots (e.g. Gregory et al. 2006; Fig. 5.183f) and even insects
nize in core, where it appears as more or less straight burrows and trap-door spiders (arachnids). In deep-marine environ-
arranged normal to the bedding planes (Fig. 5.144). The fill ments, holothurians are known to produce Skolithos
commonly contrasts with the surrounding rock and a lining (Dashtgard and Gingras 2012); aplacophorans (molluscs)
of the burrow wall is characteristic. may also be makers of Skolithos in this setting.
Similar Trace Fossils: Owing to the simplicity of Sko- Ethology: Aquatic Skolithos were predominantly built by
lithos, it may be confused with similar burrows, parts of suspension-feeding organisms for dwelling (domichnia),
other burrows, or even sedimentary and tectonic structures, whereas terrestrial Skolithos also include a scavenging
particularly in core (Fig. 5.189a, f). Skolithos resembles behavior.
Cylindricum Linck, 1949 (and the similar Capayanichnus Depositional Environment: Skolithos is a common indi-
vinchinensis Melchor et al., 2010) from fluvial deposits and cator of relatively high energy, shallow-water, nearshore to
likely produced by freshwater crustaceans. This ichnotaxon marginal-marine environments. In Paleozoic time, the bur-
differs from Skolithos by a lower length/width ratio rows appeared in such high abundance as to build Skolithos
(Hasiotis 2008) and thus justifies its own status (Knaust piperock, with a general decrease in occurrence from
5.30 Taenidium Heer, 1877 149
(a) (b) (c)
Sediment
New sediment mound
layer
Fig. 5.147 Reconstructions of the hypothetical organism inhabiting fully extended to gather food. b Position of organism during time of
Skolithos linearis. Reconstructions based on sedimentary features high-turbidity of sedimentation with tentacles folded inwards. c The
found in and near S. linearis and on life habits of phoronids (Hyman organism emerging and unfolding its tentacles at the end of a high
1959). Scale bar = 1 cm. After Sundberg (1983), © Paleontological sedimentation period. The arrow indicates the direction of sediment
Society, published by Cambridge University Press, reproduced with movement of the tentacles to form the sediment mound
permission. a Organism during low-turbidity conditions with tentacles
Cambrian towards Permian (Droser 1991; Desjardins et al. ranges between 5 and 450 mm (Smith et al. 2008), usually
2010). Skolithos also accompanies other trace fossils in shelfal between 5 and 10 mm. Burrows are unlined or very thinly
and deep-marine environments and is a common constituent lined and unbranched (DʼAlessandro and Bromley 1987).
of fluvial and other continental deposits (e.g. Hasiotis 2010; The backfill menisci are often tightly spaced with little
Melchor et al. 2012). contrast in lithology, grain size or color.
Ichnofacies: Skolithos is the namesake of the Skolithos Ichnotaxonomy: About seven ichnospecies of Taenidium
Ichnofacies, but occurs in other marine and continental are currently distinguished, although this number may
ichnofacies too. change after a thorough review of Taenidium and related
Age: Skolithos is a wide-ranging trace fossil known from meniscate trace fossils (Keighley and Pickerill 1994; Rodrí-
late Precambrian time (Alpert 1975; McCall 2006) through guez-Tovar et al. 2016). The distinction of Taenidium from
the Holocene (Dashtgard and Gingras 2012). the similar trace fossil Beaconites is still debated and finally
Reservoir Quality: Passive burrow fill of Skolithos and may show that Beaconites must be regarded as junior syn-
their tendency of crowded occurrence (so-called piperock) onym of Taenidium (e.g. Goldring and Pollard 1995; Savrda
make such horizons as good candidates to influence reser- et al. 2000). Likewise, Naktodemasis Smith et al., 2008 can
voir quality and vertical connectivity in a positive manner probably also be accomodated in this ichnogenus
(Knaust 2014a). They have the potential to overcome minor (Krapovickas et al. 2009). Ichnospecies of Taenidium
baffles and barriers (at the centimetric-scale) and to connect are differentiated by the style of meniscate backfill, and
reservoir layers. T. serpentinum, T. diesingi (=T. satanassi), T. cameronensis,
T. barretti, T. irregulare (=T. crassum), T. planicostatum and
T. bowni can currently be regarded as valid ichnospecies.
5.30 Taenidium Heer, 1877 Substrate: Taenidium typically occurs in (preferably
fine-grained) sandstone and is associated with heterogeneous
Morphology, Fill and Size: Taenidium is a cylindrical paleosols (Fig. 5.149). Interactions with incipient limestone
meniscate burrow, winding in a predominantly subhorizontal nodules (caliche) and many examples with well-preserved and
but also subvertical direction (Fig. 5.148). Burrow diameter sharp burrow margins indicate cohesive substrate conditions.
Fig. 5.148 Morphological features of Taenidium. a T. serpentinum through Copyright Clearance Center, Inc. b Typical sections of T.
Heer, 1877. Scale bar = 1 cm. From DʼAlessandro and Bromley bowni (Smith, 2007). Scale bar = 2 mm
(1987), republished with permission of Wiley; permission conveyed
Appearance in Core: In core, Taenidium appears as cir- dwelling behaviors (fodinichnia; Hembree and Hasiotis
cular, elliptical to elongate burrow segments with a winding 2008). It is likely that the tracemakers fed on organic matter
course and varying orientation (Fig. 5.150). The meniscate and roots within the soil profile (fodinichnion; Fig. 5.152;
backfill of the burrows is conspicuous. Well-developed Smith et al. 2008).
Taenidium ichnofabrics consist of a mixed layer at the top Depositional Environment: Insect traces of the kind of
and discrete burrows in the lower tier. Taenidium are commonly found in alluvial, fluvial and
Similar Trace Fossils: Taenidium is similar to other marginal-lacustrine environments (Savrda et al. 2000;
meniscate burrows such as Scoyenia and Ancorichnus Bedatou et al. 2009; Hasiotis 2010). They are part of many
(Fig. 5.151; Retallack 2001; Smith et al. 2008). The burrow soil deposits, particularly those with a higher amount of
diameter of Scoyenia is commonly larger (10–30 mm) than moisture but above the water table (A and upper B horizons
in Taenidium (5–10 mm), and Scoyenia is a lined and striated of soil; Hembree and Hasiotis 2008). Taenidium occurs also
burrow with more irregular menisci. Ancorichnus consists of in shallow- and deep-marine deposits (DʼAlessandro and
an unlined, backfilled mantle and thick meniscate backfill. In Bromley 1987).
core, Taenidium may be confused with Zoophycos spreiten, Ichnofacies: Taenidium occurs in marine deposits of the
the latter of which typically alternate vertically (where parts Cruziana Ichnofacies (e.g. Bromley et al. 1999) and is a
of a whorl were cut), or with Rhizocorallium, which contains characteristic element of the Scoyenia Ichnofacies and other
a passively filled marginal tube. Small-scale synsedimentary continental ichnofacies (Buatois and Mángano 2011;
deformation may be responsible for producing Taenidium- Melchor et al. 2012).
like structures (Fig. 5.189c). Age: Ordovician to Holocene (Keighley and Pickerill
Producers: Arthropods are probably the main tracemak- 1994).
ers of Taenidium (Rodríguez-Tovar et al. 2016). By com- Reservoir Quality: No analysis of the influence of
parison with modern analogs and experimental studies, there Taenidium on reservoir quality is available. The uniformly
is good confidence that continental Taenidium is produced meniscate fill of the burrows typically is in the same
by beetle larvae, cicada nymphs, or other insects (Smith grain-size range as the host rock and therefore little differ-
et al. 2008; Hembree and Hasiotis 2008; Fig. 5.152), but ence may be expected. The homogeneous fill of burrows,
also earthworms. Marine Taenidium may result from the which occur in moderate to high density in ripple-laminated
activity of arthropods or worm-like organisms. sandstone with varying grain-size distribution (e.g. overbank
Ethology: Taenidium is interpreted as burrows produced deposits), may favor a better flow behavior than the lami-
by a combination of detritus-feeding, locomotion and nated rock surrounding it (Fig. 5.150c).
(a) (b)
(c) (d)
(e) (f)
Fig. 5.149 Taenidium in outcrop. Scale bars = 1 cm. a, b Completely d Loose slab of sandstone with a reworked caliche nodule and winding
bioturbated sandstone with T. barretti in horizontal (a) and vertical T. barretti with pronounced backfill. Same locality as in (c). From
section (b). Upper Jurassic (Kimmeridgian, fluvial, crevasse splay), La Knaust (2015b). e, f T. irregulare (=T. crassum), freely winding and
Griega Beach in Colunga, Asturias, northern Spain. c Cross-stratified, cross-cutting one another along a sandstone bedding plane. Upper
fine-grained sandstone bed containing brownish T. barretti in its deeper Jurassic (Kimmeridgian, shallow marine), coastal cliffs at Praia do
tier (ca. 10–30 cm below surface). Upper Triassic Kågeröd Formation Salgado, western Portugal
(fluvial overbank), Bornholm, Denmark. From Knaust (2015b).
(a) (b) (c)
(d) (e)
(f) (g)
b Fig. 5.150 Taenidium in sectioned core. Scale bars = 1 cm. a T. Formation (fluvial, overbank), Snorre Field, Norwegian North Sea
barretti in moderately bioturbated sandstone with a reworked caliche (well 34/7-1, ca. 2504.65 m). e Pedogenetically modified carbonate
nodule. Upper Triassic (Norian-Rhaetian) Lunde Formation (fluvial, rock (calichified dolomitic limestone) preserving T. barretti ichnofabric
overbank), Snorre Field, Norwegian North Sea (well 34/7-A-9H, ca. with burrows penetrating the matrix and the cracks. Upper Triassic
2635.55 m). b Taenidium ichnofabric in highly bioturbated micritic (Norian) Skagerrak Formation (alluvial), Johan Sverdrup Field, Nor-
limestone. Upper Permian Khuff Formation (carbonate platform, wegian North Sea (well 16/2-15, ca. 1975.6 m). f Marly limestone with
restricted lagoon), South Pars Field, Persian Gulf, Iran (well SP-9, Taenidium showing false branching. Lower Cretaceous (Berriasian)
3086.5 m). From Knaust (2009a), republished with permission of Asgaard Formation (shelf), Johan Sverdrup Field, Norwegian North
GulfPetroLink. c Selective carbonate cementation mainly following Sea (well 16/2-11A, ca. 2175.5 m). g Chalk with a dense Taenidium
Taenidium burrows as fluid pathways. Triassic Skagerrak Formation ichnofabric, some of the burrows occupying sediment-filled fractures
(alluvial), Johan Sverdrup Field, Norwegian North Sea (well 16/2-9S, and/or large burrows. Paleocene (Danian) Shetland Group (shelf),
ca. 1954.5 m). d Discrete T. barretti burrows with meniscate backfill in Oseberg Field, Norwegian North Sea (well 30/9-B-44B, ca. 4258.5 m)
ripple-laminated sandstone. Upper Triassic (Norian-Rhaetian) Lunde
Fig. 5.151 Differentiation among common backfilled meniscate burrows. After Retallack (2001), republished with permission of Wiley; per-
mission conveyed through Copyright Clearance Center, Inc.
Fig. 5.152 Insect tracemaker

producing meniscate backfill as it
moves through the soil. After
Smith (2007)
5.31 Teichichnus Seilacher, 1955 deposits (e.g. Ekdale et al. 1984; Frey and Bromley 1985;
Savrda 2012), limestone successions (e.g. Farrow 1966; Frey
Morphology, Fill and Size: Teichichnus is a vertical wall- 1970b), and is rarely seen in shale (e.g. Jordan 1985).
like spreite burrow with a straight or curved plan view. It Appearance in Core: Teichichnus is a deep-tier trace
commonly includes unbranched (and rarely branched) ich- fossil and thus has high preservation potential. Because of its
nospecies. Burrow size is quite variable and ichnospecies- as diagnostic features it is relatively easy to recognize in core
well as age-dependent. It ranges from burrow diameters of a (Fig. 5.156; see also Figs. 4.3, 5.100h, 5.111a, b, d, 5.118b,
few millimeters to several centimeters, while burrow length 5.121a, 5.129b and 5.179a). In vertical sections, Teichichnus
can reach more than a meter (e.g. Martinsson 1965). commonly appears as patches of vertical to steeply inclined
Ichnotaxonomy: More than a dozen ichnospecies were spreiten, which may be stacked on top of each other due to
introduced since the erection of T. rectus Seilacher, 1955, different sedimentation and colonization events (Fig. 2.6c).
but many of them are probably morphological variants of a The spreiten are densely laminated and contrast in color with
few ichnospecies (Figs. 5.153, 5.154 and 5.155). T. rectus the host rock. Their width is highly variable and depends on
(Figs. 5.154a and 5.178d) and T. zigzag are the commonest the position of the section with respect to the elongate bur-
ichnospecies. In addition to burrows with only one opening, row. Most spreiten are concave-up, although concave-down
U-shaped burrows with two openings occur (Fig. 5.154b). spreiten may also occur, even within the same burrow (e.g.
Substrate: Teichichnus preferentially occurs in silty and in Teichichnus with a sigmoidal shape, Fig. 5.154a). In
muddy sand (softground). It is also been described from chalk elongate sections, only horizontal lamination may be
Fig. 5.153 Idealized reconstruction of most common ichnospecies of Teichichnus. From Seilacher (2007), republished with permission of
Springer
5.31 Teichichnus Seilacher, 1955 155
Fig. 5.154 Examples of two

different reconstructions of
Teichichnus. Reprinted by
permission of the publisher
(Taylor & Francis Ltd., http://
www.tandfonline.com). a T.
rectus from the Carboniferous of
Ireland. From Buckman (1996).
b U-shaped Teichichnus from the
Holocene of Norway. From
Corner and Fjalstad (1993)
evident. Retrusive spreite burrows are common, although may be confused with the vertical to oblique spreite burrow
protrusive ones may occur too. In some cases, the terminal Paradictyodora, which consists of an irregularly folded,
burrow is preserved as a passively filled lumen and thus fanlike to subconical sheet (Olivero et al. 2004; DʼAlessandro
contrasts with the actively filled spreite. and Fürsich 2005). The entrance burrow of some T. rectus is a
Similar Trace Fossils: Particularly in core, Teichichnus simple, passively filled shaft similar to Skolithos (Buckman
can resemble other spreite traces such as Diplocraterion and 1996). Bann et al. (2004) have documented Rosselia with
Rhizocorallium but differs from them by lacking a wide Teichichnus-like basal extensions, while Teichichnus can be
marginal tube. U- to L-shaped Trichophycus are also similar to part of various trace fossils (e.g. Ophiomorpha and Tha-
Teichichnus but commonly have loosely packed segments lassinoides; Bertling et al. 2006). Many two-dimensional core
with poorly developed spreite (Jensen 1997). Phycodes is sections appear with figures described as Siphonichnus sur-
another trace fossil with off-branching, U-shaped spreite sumdeorsum and S. lepusaures (Zonneveld and Gingras 2013;
burrows, and sections of P. palmatum and P. parallelum may Knaust 2015a; Fig. 5.156e, f, i), both which may turn out
resemble Teichichnus. Some sections of the bulbous, down- simply to be cross sections of broad, bow-shaped burrows
ward tapering to bow-shaped, laminated Cylindrichnus may (i.e. T. zigzag) instead of columnar structures.
be also confused with Teichichnus. The J-shaped cylindrical Producers: Worm-like animals (e.g. annelids) and arthro-
burrow Artichnus displays a vertically stacked spreite struc- pods (e.g. trilobites, crustaceans) are commonly regarded as
ture in its basal part, which differs from Teichichnus in potential producers of Teichichnus. The modern polychaete
wrapping around a central lumen and continuing above it Nereis sp. is known to produce spreite burrows similar to
(Zhang et al. 2008; Ayranci and Dashtgard 2013). Neverthe- Teichichnus (Seilacher 1957; Dashtgard and Gingras 2012)
less, distinction of these ichnogenera in core might be difficult and many Paleozoic forms probably result from the activity of
and Artichnus may be overlooked and misidentified as trilobites. However, X-ray radiography of cores from Holo-
Teichichnus. Sections of many observed specimens suggest cene sediments of the Baltic Sea has demonstrated that some
an overall broad, U-shaped morphology as described in bivalves (e.g. Arctica islandica) may leave Teichichnus-like
Catenichnus McCarthy, 1979, which closely resembles traces by their crawling-plowing mode of locomotion (Werner
Teichichnus and might be synonymous with it (cf. Corner and 2002; Fig. 5.157), while others (e.g. Solecurtus strigilatus)
Fjalstad 1993; Fig. 5.58). Rather oblique Teichichnus spreiten can produce spreite-like structures by the periodic shifting of
(a) (b)
(c) (d)
(e) (f)
Fig. 5.155 Teichichnus in outcrop (a–e) vertical sections, (f) lower Julegård cliff section, Bornholm, Denmark. See Clemmensen et al.
bedding plane. Scale bars = 1 cm. a Individual T. rectus. Lower (2011). d Teichichnus ichnofabric in vertical section in sandy substrate.
Ordovician Beach Formation (shoreface), Bell Island, Newfoundland, Paleocene (shallow marine), south Shikoku, southern Japan. e Retrusive
Canada. See Fillion and Pickerill (1990). b T. palmatus cross sections spreite burrows together with the cast of a large bivalve (b). Same
in silicified limestone. Middle Permian Kapp Starostin Formation locality as in (d). f Straigth specimens of T. duplex. Upper Triassic
(mixed siliciclastic carbonate ramp), Akseløya, Svalbard. c Glauconitic (Middle Keuper, fluvial to lagoonal) of Eppingen near Heilbronn,
sandstone and siltstone with a dense ichnofabric of T. cf. zigzag. Lower southern Germany. Image courtesy of Thomas Schulz (Heilbronn)
Cambrian Norretorp Member (Læså Formation, shallow marine),
5.31 Teichichnus Seilacher, 1955 157
(a) (b) (c)
(d) (e)
(f)
(g) (h)
(i)
b Fig. 5.156 Teichichnus in full (a) and sectioned core (b–i). Scale with longitudinal, oblique and cross sections of several burrows.
bars = 1 cm. a, b Various sections of T. zigzag on the outside of a core Middle Jurassic (Bajocian) Tarbert Formation (sandy tidal flat),
(a) and on two vertical core faces trending perpendicular to each other Oseberg Sør Field, Norwegian North Sea (well 30/9-F-26, ca.
(b). Middle Jurassic (Bathonian-Oxfordian) Hugin Formation (marginal 4462.0 m). g T. zigzag in longitudinal section. Note the retrusive
marine), Norwegian North Sea (well 25/7-2, ca. 4475.15 m). spreite in the lower burrow part and the sand-filled cast of the
c U-shaped Teichichnus isp. Lower Jurassic (Pliensbachian-Toarcian) (holothurian?) producer in the upper part. Middle Jurassic (Bajocian)
Cook Formation (marginal marine), Visund Field, Norwegian North Tarbert Formation (sandy tidal flat), Oseberg Sør Field, Norwegian
Sea (well 34/8-1, ca. 3653.4 m). d Completely bioturbated ichnofabric North Sea (well 30/9-4S, ca. 3346.6 m). h T. zigzag in longitudinal
with T. zigzag, partly reburrowed with Nereites and accompanied by section. Middle Jurassic (Bajocian) Tarbert Formation (sandy tidal flat),
Schaubcylindrichnus. Lower Jurassic (Toarcian) Stø Formation (off- Askja Øst Discovery, Norwegian North Sea (well 30/11-9A, ca.
shore transition), Norwegian Barents Sea (well 7120/8-2, ca. 3846.85 m). i T. zigzag in cross section (lower part) and along bedding
2153.6 m). e Two T. zigzag in cross section. Middle Jurassic (Bajocian) plane (upper part). Middle Jurassic (Bajocian) Tarbert Formation
Tarbert Formation (sandy tidal flat), Oseberg Sør Field, Norwegian (sandy tidal flat), Oseberg Sør Field, Norwegian North Sea (well
North Sea (well 30/9-10, ca. 2813.15 m). f Teichichnus ichnofabric 30/9-13S, ca. 3135.5 m)
(a) (b)
Fig. 5.157 Comparable modern analogs of Teichichnus that were core depth. Courtesy of the Radiography Database, University of
probably produced by bivalves. a X-ray radiograph (negative) of Kiel, http://www.ifg.uni-kiel.de/Radiographien/radiographien.phtml.
Holocene core from the Kiel Bay in the western Baltic Sea, showing b Reconstruction of the crawling-plowing mode of locomotion of A.
spreite-like traces (plow-sole traces, P) similar to Teichichnus, produced islandica, which leads to the origin of a Teichichnus-like trace (“fossil”
by the bivalve Arctica islandica. Ca. 27 m water depth and 0.75–1.0 m plow-sole trace F). After Werner (2002)
the burrow (Bromley 1996; Fig. 5.158). Catenichnus- and Depositional Environment: The ichnogenus Teichichnus
Artichnus-like Teichichnus are consistent with features pro- seems to be a typical element of siliciclastic systems, where
duced by holothurians and might be a result of such. it occurs frequently in association with delta deposits (e.g.
Ethology: Teichichnus mainly results from the deposit- Tonkin 2012). Its producer(s) can be regarded as euryhaline
feeding (fodinichnial) activity of its producer and includes organisms that are able to adapt to a wide range of salinity.
dwelling of a non-vagile tracemaker (MacEachern et al. For this reason, Teichichnus has often been recognized
2012). By adjusting its vertical position, Teichichnus partly in environments associated with lowered salinity. Such
served as an equilibrichnion. conditions are common in marginal-marine settings
5.32 Thalassinoides Ehrenberg, 1944 159
Fig. 5.158 Solecurtus strigilatus

in feeding position. Periodic
shifting of the burrow leaves a
spreite-like structure that may
resemble Teichichnus (although
more oblique). The upper parts of
this structure are rapidly
obliterated by shallower
bioturbation. Depth of burrow
30 cm. Broken lines at lower
right indicate the path taken
during escape reactions. From
Bromley (1996), reprinted by
permission of the publisher
(Taylor & Francis Ltd., http://
www.tandfonline.com)
(e.g. embayments, estuaries, lagoons, ponds etc.), where Age: Teichichnus is known from the Early Cambrian (e.g.
Teichichnus is widespread (Fig. 5.156a–c). Although ichno- Loughlin and Hillier 2010) to the Holocene (e.g. Wetzel
diversity in such assemblages is low, simple deposit-feeding 1981; Corner and Fjalstad 1993).
traces like Planolites may occur along with it (Pemberton and Reservoir Quality: The producer of Teichichnus creates
Wightman 1992; Buatois et al. 2005; MacEachern and Gin- an actively filled spreite by incorporating and packing
gras 2007; Gingras et al. 2012c). Furthermore, Teichichnus is fine-grained material (e.g. mud, silt and organic-rich matter)
often present in tidal deposits (including dunes and bars in from the host sediment into its burrow. This process results
brackish settings; Desjardins et al. 2012; Fig. 5.156e–i) and in locally reduced permeability and thus leads to deteriorated
hyperpycnal flow deposits (Buatois et al. 2011). reservoir quality (Tonkin et al. 2010).
In contrast to such marginal-marine occurrences with low
ichnodiversity, Teichichnus is a characteristic element of
lower shoreface to offshore (shelf) deposits (Pemberton et al. 5.32 Thalassinoides Ehrenberg, 1944
2012). There, however, it occurs in highly diverse
trace-fossil associations (Fig. 5.156d). In this setting, low- to Morphology, Fill and Size: Similar to the genetically related
moderate-energy conditions are prevailing, typical for Ophiomorpha, Thalassinoides consist of boxworks consti-
fair-weather rather than storm deposits (Pemberton et al. tuting horizontal maze with vertical shafts (Fig. 5.159),
1992). Likewise, Teichichnus has been reported from chalky although many related burrow architectures can be produced
shelf deposits (Frey and Bromley 1985). It also occurs as (e.g. Kennedy 1967; Bromley 1967, 1996; Fürsich 1974a).
accompanying trace fossil in assemblages of slope deposits The burrows are circular to elliptical in cross section.
(Hubbard et al. 2012) and deep-sea deposits (Wetzel and Branching is Y- and T-shaped, typically with bulbous
Uchman 2012). Given this latter (lower shoreface, shelf and enlargement of the junctions. Passive fill is common, although
deep marine) distribution, Teichichnus has been used in transitional burrow elements can be actively filled
sequence-stratigraphical analysis as indicator of flooding (Fig. 5.160). Burrows are unlined and contain a sandy, more
events (Pemberton et al. 1992; Taylor et al. 2003) and is rarely also a muddy fill. Thalassinoides is a relatively large
characteristic of transgresssive systems tracts. trace fossil as the boxwork can cover a volume of over 1 m3,
Ichnofacies: Teichichnus is a typical constituent of the although its burrow diameter can range from few millimeters
Cruziana Ichnofacies but also occurs in the Zoophycos to several centimeters (mean about 1 cm). Complete burrow
Ichnofacies. systems can penetrate substrate to several meters.
Ichnotaxonomy: From the about 15 erected ichnospecies, (Fig. 5.161c, d, see also Fig. 2.6b). Because the producer is
the following are most representative (cf. Myrow 1995): capable of bioeroding, Thalassinoides may also occur in
hard substrates such as cemented layers (Fig. 5.161e).
• T. suevicus—predominantly horizontal form that may Similar Trace Fossils: Thalassinoides in core may be
contain enlargements at Y-shaped bifurcations confused with other large and passively filled burrows such
(Figs. 5.159a and 5.160a, b) as Spongeliomorpha (scratched), Psilonichnus and Parma-
• T. paradoxicus—boxwork burrows, highly irregular in ichnus (vertically Y-shaped), Camborygma (chambered,
size and geometry (Fig. 5.159b and 5.160c–f) dominantly vertical), and others. Sections of actively filled
• T. saxonicus—large form with tunnels 5–20 cm in diameter. burrow elements, however, could resemble Planolites,
Asterosoma or even Artichnus. Processes of methane and
Substrate: Thalassinoides occurs in a wide range of soft hydrocarbon seepage may result in branched pipes similar to
to firm substrates and is reported from mudstone, siltstone, Thalassinoides (Fig. 5.189g).
sandstone, conglomerate, limestone and dolomite. It can also Producers: Similar to Ophiomorpha, comparison with
be associated with lithified (hard) substrates. modern analogs shows that the producers of Thalassinoides
Appearance in Core: The three-dimensional geometry of belong to thalassinidean shrimp, particularly callianassids
Thalassinoides is hard to resolve in cored material and (Figs. 5.25, 5.85–5.87). This is principally true for Permian
therefore an unequivocal attribution is seldom possible with to modern Thalassinoides, whereas other arthropods (e.g.
certainty. The relatively large burrow size differs from that of trilobites), sea anemones and worms (e.g. enteropneusts)
associated ichnotaxa and the burrows appear in circular and have been interpreted as tracemakers of Paleozoic Tha-
elliptical cross sections together with (rarely) vertical shafts lassinoides (e.g. Ekdale and Bromley 2003; Cherns et al.
(Figs. 5.161a, b and 5.168). The passive fill usually creates a 2006). Some of these early Thalassinoides probably belong
contrast with the host sediment, although in other cases the to other ichnogenera such as Balanoglossites (Knaust and
fill is the same as the host sediment. It can be subject of Dronov 2013).
reburrowing by other tracemakers, for instance resulting in Ethology: Thalassinoides-producing shrimp are primarily
the occurrence of Chondrites (Fig. 5.35c and 5.161f). suspension-feeders, and their extensive burrow systems were
Depending on the substrate, transitions to related ichno- excavated as dwellings (domichnia). Combined suspension-
genera (e.g. Ophiomorpha and Gyrolithes) are common and deposit-feeding behavior may apply for many Tha-
lassinoides. Other representatives actively collect seagrass
and other organic matter and store it in burrow chambers as
cache (Griffis and Suchanek 1991).
Depositional Environment: Shrimp-produced Thalas-
sinoides is most common in shallow-marine environments
such as shoreface, deltas and others (e.g. Nickell and
Atkinson 1995). Given the ability of the producer to tolerate
fluctuations in salinity, Thalassinoides can be found in
brackish environments, e.g. in estuarine settings and fan
deltas (e.g. Swinbanks and Luternauer 1987). Thalassinoides
is often associated with firm substrates (Glossifungites Ich-
nofacies), where it occurs in a wide range of environments
from marginal marine to the deep marine (Monaco et al.
2009). It is a common constituent of carbonate and chalk
environments (Bromley 1967; Ekdale and Bromley 1991).
Ichnofacies: Thalassinoides is a common constituent of
the Cruziana Ichnofacies, where it occurs in relatively
cohesive substrates. Thalassinoides is also a component of
the substrate-controlled, firmground trace-fossil suite (the
so-called Glossifungites Ichnofacies).
Age: Thalassinoides is frequently reported from the
Ordovician (e.g. Myrow 1995; Ekdale and Bromley 2003;
Fig. 5.159 Morphological variation of Thalassinoides ichnospecies. Cherns et al. 2006) to the Holocene (e.g. Swinbanks and
a T. suevicus (maze). b T. paradoxicus (boxwork). Scale bars = 5 cm.
After Howard and Frey (1984), republished with permission of
Luternauer 1987; Nickell and Atkinson 1995), although
Canadian Science Publishing; permission conveyed through Copyright tracemakers of different phyla must be invoked.
Clearance Center, Inc.
5.32 Thalassinoides Ehrenberg, 1944 161
(a) (b)
(c) (d)
(e) (f)
b Fig. 5.160 Thalassinoides in outcrop and building stone. Scale T. paradoxicus. Cretaceous (platform carbonate), coastal cliff near
bars = 5 cm. a T. suevicus preserved on lower bedding plane. Upper Taghazout, western Morocco. d T. paradoxicus in vertical section.
Jurassic sandstone (Lower Coralline Oolite, shallow marine, shoreface) Same locality as in (c). e Bedded and bioclastic limestone with
coastal cliff near Scarborough, Yorkshire, UK. b T. suevicus preserved stylolites and T. paradoxicus ichnofabric in vertical section. Cretaceous
on lower bedding plane. Upper Jurassic (Upper Oxfordian, lagoonal (shallow marine), Lisbon, Portugal, building stone. f T. paradoxicus
platform carbonate), Krzemionki Opatowskie Neolithic flint mine, burrow system in limestone. Cretaceous (shallow marine), Lisbon,
southern Poland. c Bedding plane with extensive system of Portugal, building stone
(a) (b) (c)
(d) (e)
(f)
Fig. 5.161 Thalassinoides in sectioned core. Scale bars = 1 cm. d Mudstone interlayer between turbidite sandstone, containing large
a Argillaceous sandstone exposing a large-diameter burrow in cross passively filled burrows. Note the occurrence of spreite due to retrusive
section, which is passively filled with coarse-grained sand. Upper burrow adjustment and the transition to actively filled and lined
Jurassic (Tithonian) Draupne Formation (fan delta, slope), Johan Ophiomorpha burrows. Upper Cretaceous (Maastrichtian) Springar
Sverdrup Field, Norwegian North Sea (well 16/2-15, ca. 1943.5 m). Formation (deep marine, fan system), Norwegian Sea (well 6604/10-1,
b Completely bioturbated coarse-grained sandstone with Thalassi- ca. 3647.5 m). e Sandstone with early diagenetic carbonate cementa-
noides boxwork preserved due to partial cementation of the host rock. tion. The originally hard palisade calcite band in the middle is partly
This stratigraphical interval represents an omission surface within the eroded with a sand-filled tunnel coming from the mudstone layer in
region. Upper Jurassic (Tithonian) Draupne Formation (fan delta, between. Upper Cretaceous (Campanian) Nise Formation (deep marine,
slope), Johan Sverdrup Field, Norwegian North Sea (well 16/5-2S, ca. fan system), Aasta Hansteen Field, Norwegian Sea (well 6707/10-1, ca.
1958.5 m). c Chalky limestone with mainly vertical burrow segments 3133.5 m). f Chalky limestone with horizontal burrow parts (tunnels),
(shafts) with a slightly helical appearance as known from the related which are mainly mud-filled and intensively reburrowed with Chon-
ichnogenus Gyrolithes. Lower Cretaceous (Berriasian) Åsgard Forma- drites. Lower Cretaceous (Berriasian) Åsgard Formation (shelf), Johan
tion (shelf), Norwegian North Sea (well 16/4-6S, ca. 1945.2 m). Sverdrup Field, Norwegian North Sea (well 16/2-15, ca. 1899.75 m)
5.33 Tisoa de Serres, 1840 163
(a) (b)
Fig. 5.162 Burrow-enhanced permeability created by Thalassinoides burrow fill consists of sucrosic dolomite, and the matrix consists of
boxworks. From Pemberton and Gingras (2005), republished with mosaic dolomite, resulting in a large difference in permeability. This
permission of AAPG; permission conveyed through Copyright Clear- leads to a charging effect that may become a problem wherein oil in the
ance Center, Inc. a Development of super-K in the Jurassic Arab-D, matrix is bypassed and water is drawn from the aquifer.
Ghawar Field, Saudi Arabia, is a function of the interaction of a b Super-permeability forms where the assemblage attributed to the
ravinement bed developed on a regional transgressive surface of Glossifungites Ichnofacies is emplaced at the surface, with a ravinement
erosion. The Glossifungites Ichnofacies is represented by firmground bed above capped by an offshore bioturbated facies. When present, the
Thalassinoides systems 1–2 cm in diameter, penetrating up to 7 ft flowmeters indicate that 70% of the production comes from this single
(2.1 m) below the transgressive surface of erosion. In some cases, the unit
Reservoir Quality: Its wide distribution, large diameter, 5.33 Tisoa de Serres, 1840
ramification pattern and passive fill make Thalassinoides
susceptible for enhanced fluid flow, which may result in Morphology, Fill and Size: Tisoa refers to very long, elon-
concretional cementation (e.g. Bromley 1967; Fürsich 1973; gate U-shaped burrows with subvertical, oblique and sub-
Fig. 5.89), but also in a good friend of the reservoir geolo- horizontal orientation. The two limbs of the burrow are
gist. The burrows can considerably increase the reservoir positioned very close to each other (Fig. 5.163), which
quality and commonly interconnect otherwise isolated parts results in cross sections with a dumbbell or figure-of-eight
of the reservoir. The reservoir-improving character of Thalas- shape (e.g. Gottis 1954). Tisoa burrows are passively filled,
sinoides has been outlined by various studies. For instance, commonly with sulfide mineralization, and are often encased
Pemberton and Gingras (2005) reported on Thalassinoides- in carbonate concretions. Branching has been recorded but is
perforated firmgrounds that act as “super-permeable” not common. Burrow diameter commonly ranges between 3
horizons in the Arab-D of the world’s largest oil field, the and 20 mm: the length of the burrow is often hard to deter-
Ghawar Field in Saudi Arabia. Sandstone-filled Thalassi- mine because of incompleteness, but can reach several
noides in mudstone of the Ben Nevis Formation off New- decimeters.
foundland (Canada) creates vertical and horizontal Ichnotaxonomy: T. siphonalis is the most common
macropore networks with the potential to act as flow con- ichnospecies (Fig. 5.164), although a few other ichnospecies
duits (Tonkin et al. 2010; Fig. 5.162). Other examples with a were erected subsequently based on morphological variation.
similar size range documenting Thalassinoides-dominated Substrate: T. siphonalis is often reported from argillaceous
ichnofabrics having a critical impact on the fluid-flow substrates, such as black shale. It also occurs in redeposited
properties of hydrocarbon reservoirs and groundwater sediments, for instance debrites and mass-transport deposits,
aquifers (Cunningham and Sukop 2012; Cunningham et al. and coal. In outcrop, Tisoa is often recognized based on its
2012). encasement within a tubular carbonate concretion.
Fig. 5.163 Historical figure of

the type ichnospecies of Tisoa, T.
siphonalis encased in
concretionary limestone, in its
original description by de Serres
(1840)
Appearance in Core: A clear indication of Tisoa in core Confusion may also arise with Rhizocorallium jenense,
is the occurrence of coupled and passively sand-filled bur- which like Diplocraterion bears a spreite and has a much
rows (Fig. 5.165). Beside that, elongate oblique and circular lower length/width ratio compared to Tisoa (Knaust 2013).
cross sections of burrows occur in association. Tisoa ichno- The U-shaped turn of Tisoa resembles that of some Areni-
fabric may reach a relatively high density and may be traced colites, but again their limbs are much closer to each other
to considerable depth (several decimeters). and the length/width ratio is greater. Similar to Tisoa, the
Similar Trace Fossils: Given the extreme length of the vertically oriented trace fossils Paratisoa Gaillard, 1972
burrows, their U-turn is rarely preserved, which may give the (Macsotay et al. 2003) and Bathichnus Bromley et al., 1975
impression that vertically oriented specimens are not trace (Nygaard 1983) are diagenetically enhanced (e.g. by car-
fossils but conduits for escaping seep gases (van de bonate and flint concretions) and can reach extraordinary
Schootbrugge et al. 2010). So far, Tisoa has not been recog- length, but differ from Tisoa by having only one tube.
nized from core, but has been attributed to other trace fossils Producers: Based on modern analogs, tube-dwelling
due to similarities. For instance, the material shown in polychaetes, such as the giant pogonophoran worms, could
Fig. 5.165 was originally described as large Chondrites have produced Tisoa-like traces. In the Pacific Ocean, 1.5 m
(Knaust 2009b). Diplocraterion is another burrow with long pogonophorans grow in heated, sulfur-rich water
similar features as Tisoa, particularly when the limbs of long around warm-water vents (Ruppert et al. 2004).
burrows are very close together and no spreite can be recog- Ethology: A preferred dwelling (domichnial) behavior
nized between them. Such forms were described as can be inferred for the Tisoa producers.
Diplocraterion and D. habichi but some of them may belong Depositional Environment: A striking feature of Tisoa is
to T. siphonalis for the reasons given above (e.g. Heinberg its frequent occurrence in the central part of carbonate
and Birkelund 1984; Bradley and Pemberton 1992; de Gibert concretions related to hydrocarbon seep deposits (Breton
and Martinell 1998; Bann and Fielding 2004; Bann et al. 2006), preferably occurring in deep-marine basins and on
2004; Hubbard and Schultz 2008; Riahi et al. 2014). continental slopes. The Tisoa producer seems to have an
5.34 Trichichnus Frey, 1970b 165
(a) (b)
(c) (d)
Fig. 5.164 Tisoa siphonalis in outcrop. Scale bars = 1 cm. a Large a calcite-cemented sandstone. Same locality as in (a). c Bottom
sandstone clast with cross sections of passively filled T. siphonalis with fragment of a pyritic specimen, with the U-turn to the left. Lower
a dumbbell to figure-of-eight shape. Late Miocene Urenui Formation Cretaceous (Hauterivian), Ulyanovsk District, Volga River (coll.
(deep marine, continental slope), Waiau coastal section north of New Hecker, Paleontological Museum Moscow). d Cross section. Same
Plymouth, North Island, New Zealand. b Elongate T. siphonalis within specimen as in (c)
affinity for sediment with a high organic content and thus Reservoir Quality: The passive fill (commonly consisting
can also be found in mass-transport deposits and even in coal of sand), together with relatively large tubes and extraordi-
seams. Callow et al. (2013) describe abundant, >1 m deep nary penetration depth into mudstone (non-reservoir, source
U-shaped trace fossils from cohesive mudstone below rock), make Tisoa very suitable for overcoming thin barriers
bypass surfaces within the channel axis of a deep-marine and baffles within a given reservoir.
channel, which are similar to those forms reported by Knaust
(2009b, Fig. 5.165).
Ichnofacies: Tisoa does not seem to have a preference to a 5.34 Trichichnus Frey, 1970b
particular ichnofacies because of its close association with
deposits with a high content of organic matter. Such condi- Morphology, Fill and Size: Trichichnus refers to sparsely
tions, however, are typically met on deep-marine basin floors branched or unbranched, hairlike and exceptionally long
(Nereites Ichnofacies) and continental slopes (Zoophycos burrows with a more or less vertical orientation (Fig. 5.166).
Ichnofacies), both preferred sites of seepage. In many cases, The burrows can be lined and are typically filled with sulfide
the sediment already became consolidated before its deep minerals (e.g. pyrite). The morphology of the cylindrical
penetration by the Tisoa producer, and the resulting traces burrows is straight or gently winding to slightly sinuous.
can be assigned the Glossifungites Ichnofacies. Burrow diameter is in the range of 0.1–1.0 mm, whereas
Age: Tisoa is known from the Early Jurassic (van de burrow length of complete burrows commonly exceeds
Schootbrugge et al. 2010) to the Miocene (Frey and Cowles 20 cm but can reach several meters. Scholle (1971) reported
1972). specimens with a length of more than 6 m.
Fig. 5.165 Tisoa in sectioned (a) (b)

core. Scale bars = 1 cm.
a–d Tisoa ichnofabric. Upper
Cretaceous (Campanian)
debris-flow deposit (deep marine,
fan complex), Aasta Hansteen
Field, Norwegian Sea (well
6707/10-1, 3058.7-3059.9 m).
From Knaust (2009b),
Elsevier; permission conveyed
Center, Inc. a Diagram of the
Tisoa ichnofabric in photograph
(left) and line drawing (right),
indicating a maximum
penetration depth on the order of
1 m. Note the occurrence of
pyritized burrows (red) in
addition to sand-filled burrows
(yellow). The top of the
ichnofabric is at a sharp erosion
surface, below which the burrows
appear either as vertical, oblique
or horizontal, occasionally paired (c)
tunnels. Some burrows penetrate
into the underlying sandy
debris-flow deposit, where they
are preferentially concentrated in
large, reworked, sideritic mud
clasts. b Close-up section from
(a), showing subvertical, oblique
and paired subhorizontal burrows.
c Close-up section from (a),
showing predominantly paired
subhorizontal burrows.
d Close-up section from (a),
showing predominantly paired
subhorizontal burrows within the
shale. e Two pairs of
subhorizontal and one oblique
section of Tisoa on top of a
compacted coal seam, all
passively filled with sand from
the layer above. Lower Jurassic
(Pliensbachian) Åre Formation
(top), Heidrun Field, Norwegian
Sea (well 6507/7-A-27, ca.
3235.5 m)
(d) (e)
5.34 Trichichnus Frey, 1970b 167
Ichnotaxonomy: T. linearis seems to be the only valid

ichnospecies of Trichichnus (Fig. 5.167), with T. simplex
being a preservational variant (Uchman 1999). T. appendi-
cus Uchman, 1999 refers to horizontal and oblique burrows
with short lateral appendages, and therefore is rather
excluded from the ichnogenus Trichichnus.
Substrate: Trichichnus is common in fine-grained rocks
such as mudstone, marlstone and limestone with an original
soft to firm consistency, but may also occur in sandy
substrate.
Appearance in Core: Based on its size, morphology and
mineral staining, Trichichnus is easy to recognize in core
(Fig. 5.168). Only fractions of the straight to slightly
winding thin burrows are commonly preserved and appear as
mud-filled or pyritic tubes.
Similar Trace Fossils: Trichichnus has close similarity to
Polykladichnus and Skolithos, but however, differ from them
by having a much smaller length/diameter ratio. In addition,
Polykladichnus has Y- or U-shaped branching and Skolithos
remains unbranched.
Producers: Trichichnus is a meiobenthic trace fossil
(<1 mm in diameter) which has a modern analog produced
by sipunculan worms (Golfingia, Nephasoma) on the Nor-
wegian continental slope (Romero-Wetzel 1987; Shields and
Kedra 2009; Fig. 5.166).
Ethology: Trichichnus has usually been interpreted as the
dwelling (domichnion) of a sediment-feeding chemosymbio-
tic organism. Because of its exceptional length and common
sulfide mineralization, Trichichnus can also be considered as
Fig. 5.166 Sketch of modern Trichichnus from the Vøring Plateau, chemichnion (Uchman 1995; Uchman and Wetzel 2012).
Norwegian Sea, formed by sipunculan worms. From Romero-Wetzel These structures were produced by organisms feeding
(1987), republished with permission of Springer on chemosymbiotic microbes; the organisms maintain a
(a) (b)
Fig. 5.167 Trichichnus linearis in the Miocene Marnoso-arenacea numerous fragmentary specimens (arrow heads). Note the brownish
Formation, Albignano roadcut (Santerno Valley), Italy. Scale bars = 1 cm. staining due to altering of sulfide minerals to limonite. b Close-up view
The hemipelagic background sediment (marlstone) is penetrated by of a burrow
Trichichnus, which is only partially preserved. a Overview picture with
(a) (b)
Fig. 5.168 Trichichnus in sectioned core. Scale bars = 1 cm. Sverdrup Field, Norwegian North Sea. a Well 16/3-4 (ca. 1913.65 m).
Trichichnus together with Thalassinoides and Zoophycos in the Lower b Well 16/5-2S (ca. 1956.95 m)
Cretaceous (Berriasian) chalk of the Åsgard Formation (shelf), Johan
connection to oxygenated water but penetrate into anoxic (Knaust 2010a; Fig. 5.170). The burrows are smooth-walled
sediments rich in methan, the sulfides or ammonium that are and passively filled. The average burrow diameter is about
required for microbial growth. 0.5 mm, whereas burrow length and penetration depth can
Depositional Environment: Trichichnus is common in reach several centimeters.
deep-marine deposits (turbidites and hemipelagites; Wetzel Ichnotaxonomy: V. undulatus so far is the only described
1981, 1991; McBride and Picard 1991) and often associated ichnospecies of Virgaichnus.
with chalk deposits (shelf and deep-water; Frey 1970b; Substrate: V. undulatus has been described to occur in
Savrda 2012). It preferably occurs in oxygen-deficient sedi- micritic or chalky limestone with an originally firm sub-
ment (e.g. Monaco et al. 2012). strate, although diffuse burrow segments may also indicate
Ichnofacies: Trichichnus is associated with the Cruziana, local softground conditions. It is common in sandy sub-
Zoophycos and Nereites ichnofacies. strates too.
Age: Trichichnus is known from Early Ordovician (Fil- Appearance in Core: In core, V. undulatus commonly
lion and Pickerill 1990) to Holocene (Romero-Wetzel 1987). occurs in form of dense boxworks with interconnecting
Reservoir Quality: In their letter to the journal Nature, burrow elements of submillimetric diameter (Fig. 5.171).
Weaver and Schultheiss (1983) demonstrated the profound Individual burrow segments are characterized by a bulbous
effect of open burrows (such as Trichichnus) in deep-sea and undulating appearance, whereas others contrast with a
sediments “… on the overall permeability, and conse- strong contraction that leads to blade-like burrows with a
quently, on the possible flow rates through them in response size less than 0.1 mm. Over larger distances, this feature
to any excess pore pressures”. “The effect of the burrows resembles a kind of boudinage. Consequently, the cross
changes the calculated permeability from the equivalent of a section of the burrows is highly variable and ranges from
clay to that of a coarse sand”. circular to flat elliptical and vertically extended.
Similar Trace Fossils: Morphologically, Virgaichnus
resembles some ichnospecies of Thalassinoides but differs
5.35 Virgaichnus Knaust, 2010a from them by having a much smaller and inconstant burrow
diameter (pinch-and-swell-like features). Virgaichnus shows
Morphology, Fill and Size: Virgaichnus is a meiobenthic some similarities with Chondrites and Pilichnus, from which
trace fossil (burrow diameter less than 1 mm; Knaust, it differs by the lack of dichotomous branching but the
2007a) with a complex, three-dimensional architecture inclusion of T-shaped bifurcations and crossing tunnels.
(Fig. 5.169). The irregular burrow system consists of hori- Bornichnus, another meiobenthic trace fossil, includes small
zontal and inclined elements with Y- and/or T-shaped burrows composed of a crowded tangle of lined tubes that
branching. Burrow sections pinch and swell, which leads to are closely and tortuously branched and have a constant
bulbous enlargements and alternating blade-like contractions diameter. Finally, the twig-like morphology of Virgaichnus
5.35 Virgaichnus Knaust, 2010a 169
(a)
(b) (c)
b Fig. 5.169 Virgaichnus undulatus in outcrop. a Slab with the holotype Clearance Center, Inc. b V. undulatus isp. in bedding-plane view.
specimen showing multiple branching indicated by an arrow head. Glauconitic, completely bioturbated sandstone. Paleocene Grumantbyen
Upper Permian Saiq Formation (carbonate platform, open lagoon), Wadi Formation (shoreface), near Longyearbyen, Svalbard. c Intensely
Bani Awf near Rustaq, Oman. From Knaust (2010a), republished with branched burrow system in limestone. Carboniferous Honaker Trail
permission of Elsevier; permission conveyed through Copyright Formation (deep- to shallow-marine), north of Moab, Utah, USA
(a) (b) (c)
Fig. 5.170 CT-scan of mud-filled Virgaichnus undulatus in glau- Øygard (Bergen). a Clusters of burrows trending in various directions.
conitic, completely bioturbated sandstone (full core). Paleocene Gru- b Individual subvertical burrow element showing an undulating burrow
mantbyen Formation (shoreface), near Longyearbyen, Svalbard (well section with bulbous enlargements. A short and flattened subhorizontal
BH 9-2006, ca. 389.7 m). Main burrow diameter varies between 0.3 and burrow element shows blade-like contractions. c Portion of an ichno-
0.6 mm. Images courtesy of Lars Rennan (Trondheim) and Ørjan Berge fabric consisting of numerous burrows trending in various directions
may be confused with root traces (rootlets) which, however, relative quiet sedimentation regime, such as in a shallow-
preferably bifurcate downwards and may have carbonaceous marine environment on a stable inner shelf (specimens from
material involved. Oman and Norway), open lagoon (specimens from Iran), and
Producers: The pinching and swelling of the burrow deep- to shallow-marine (specimens from Utah).
segments is in agreement with a peristaltic movement of an Ichnofacies: Virgaichnus can be regarded as component
unsegmented, highly deformable vermiform body as known of the Cruziana Ichnofacies.
in the nemerteans (Fig. 5.172). Age: The few reports of marine Virgaichnus range from
Ethology: The passive fill of the burrow system indicates Carboniferous to Paleocene.
an open tunnel boxwork, suitable for a combined dwelling Reservoir Quality: Virgaichnus boxworks may act as
and feeding trace (domichnion and fodinichnion) of a connected pore systems if remaining open (e.g. not pas-
deposit-feeding and subordinate suspension-feeding animal. sively filled or cemented), and thus lead to a considerable
Assuming nemerteans as tracemakers, a predatory lifestyle increase of porosity and permeability in otherwise tight
could also be a possibility for the producer of V. undulatus. rocks (such as micritic carbonates and chalk; Knaust
Depositional Environment: The sparse reports of 2014a). Mud-filled burrows in sandstone, however, reduce
V. undulatus document its occurrence in shelf deposits with the reservoir quality.
5.35 Virgaichnus Knaust, 2010a 171
(a) (b) (c)
A A
C V
V
A
(d) (e)
(f) (g)
Fig. 5.171 Virgaichnus undulatus in sectioned core (a–f) and full core T-like branching and dichotomy. Upper Jurassic (Oxfordian) Heather
(g). Scale bars = 1 cm. a Dolomitic limestone with a Virgaichnus Formation (shelf turbidites), Fram Field, Norwegian North Sea (well
(V) ichnofabric, accompanied by Asterosoma (A) and Cylindrichnus 35/11-11, ca. 2583.5 m). f Dense accumulation of Virgaichnus tubes.
(C). Upper Permian Khuff Formation (open lagoon), South Pars Field, Upper Jurassic (Oxfordian) Heather Formation (shelf turbidites), Fram
Persian Gulf, Iran (well SP9, ca. 3084.4 m). After Knaust (2014a), H-Nord Field, Norwegian North Sea (well 35/11-15ST2, ca. 2979 m).
republished with permission from EAGE. b–d Virgaichnus ichnofabric g Virgaichnus isp. in glauconitic sandstone. Paleocene Grumantbyen
in chalky limestone. Lower Cretaceous (Berriasian) Åsgard Formation Formation (shoreface), near Longyearbyen, Svalbard (well BH
(shelf), Norwegian North Sea (well 16/4-6S, 1943.9–1949.6 m). 10-2008, ca. 820.5 m). Compare with Fig. 5.169b and 5.170
e Loosely arranged tubes and clustered burrows, partly displaying
Fig. 5.172 Selected branch of

Virgaichnus undulatus from the
holotype specimen (see
Fig. 5.169a) with the diagnostic
pinch-and-swell-like features
(photograph), interpreted as the
result of peristaltic movement of
an unsegmented vermiform
organism such as a nemertean
(line drawing). The arrow
indicates the moving direction
and the numbers the peristaltic
movements, made by narrowing
and pushing out the proboscis and
sending back a wave of alternate
swellings and constrictions along
the body. From Knaust (2010a),
Elsevier; permission convoyed
Center, Inc. The sketch in the
lower part illustrates peristaltic
burrowing in Carinoma
tremaphoros, where peristaltic
1
waves (numbered) originate
anteriorly and progress rearward
as the animal burrows. Modified
and redrawn from Turbeville and
1 2
Ruppert (1983), with permission
of Springer
1 2
2 3
5.36 Zoophycos Massalongo, 1855 • Primary lamellae—Arched grooves and ridges charac-
terizing the lamina and interpreted as the subsequent
Morphology, Fill and Size: Zoophycos is a complex trace fossil positions of the marginal tube during its lateral displace-
with wide distribution and a long history in research. It is a ment within the sediment.
spreite burrow with a tremendous morphological variability
(Bromley 1996), which in the past inspired the establishment of Two main forms of Zoophycos can be classified: simple
numerous ichnotaxa. The general characteristics of Zoophycos planar forms and complex spirally coiled forms (Olivero and
are as following (after Olivero and Gaillard 2007; Fig. 5.173): Gaillard 2007; Fig. 5.174 and 5.175). Many Zoophycos appear
with a lobate outline of their lamina. Another feature of some
• Marginal tube—A tubular structure bordering an area of Zoophycos is the abundant occurrence of small ellipsoidal fecal
bioturbated sediment and considered as a tunnel pellets, ca. 1.5 by 0.5 mm in size. The complete Zoophycos
• Lamina—Bioturbated sediment bordered by the marginal burrow systems typically reach a size offew decimeters to more
tube (also referred to as spreite) than 1 m in diameter and several decimeters in depth.
5.36 Zoophycos Massalongo, 1855 173
Fig. 5.173 Schematic drawing

of the Zoophycos spreite. The
“laminaeˮ correspond to the
“primary lamellaeˮ in the
terminology of Olivero and
Gaillard (2007), while the
“impressionsˮ refer to fecal
pellets. From Bischoff (1968), ©
Paleontological Society,
published by Cambridge
University Press, reproduced with
permission
Ichnotaxonomy: The ichnotaxonomic status of Zoophycos with a spirally coiled lamina as diagnostic for the type
is far from being robust and much confusion exists nowa- ichnospecies Z. brianteus, vertical core sections through
days when it comes to the delineation of individual ichno- such a burrow system can cut it either axially (relatively
species. Work done by Olivero (2007) has confirmed that rare) or marginally (common). In axial expression, spreite
Z. brianteus can be regarded as the type ichnospecies. Z. burrows with a conical appearance alternate vertically
brianteus comprises Zoophycos with a spirally coiled lamina (Fig. 5.176a, b). In the marginal expression, individual and
and a slightly lobed outline. Z. villae is also regarded as valid more or less horizontally oriented spreiten become visible
and consists of a lamina furrowed by numerous sinuous and and may be stacked vertically (Fig. 5.176c–h; see also
long lamellae radiating from a raised apex. Many other Figs. 5.39b, 5.132d and 5.168). The spreite burrows are
forms are described (Zhang and Gong 2012), some of which typically a few millimeters to ca. 1 cm thick and display the
diverge considerably from the type ichnospecies and there- characteristic internal lamellae composed of alternating
fore would be best accomodated in other previously estab- sediment packages of different composition. The distance
lished ichnogenera such as Taonurus von Fischer-Ooster, between individual lamellae can be short, leading to a
1858, Cancellophycus de Saporta, 1873 and Echinospira height/width ratio of the lamellae of more than 1
Girotti, 1970 (Bromley and Hanken 2003). Some workers (Fig. 5.176d, g), or may be more stretched horizontally,
refer to the ‘Zoophycos group’ when dealing with those which results in a height/width ration of less than 1
forms with a wide range of morphology (e.g. Uchman and (Fig. 5.176e, f). Some ichnospecies of Zoophycos contain
Demircan 1999; Fig. 5.175). One might also call them the ellipsoidal fecal pellets (about 1.5 mm in length and 0.5 mm
Alectoruridae. in diameter), which belong to Coprulus oblongus and can be
The various forms of Zoophycos show a general evolu- identified in core (Fig. 5.176c). Because of subsequent
tionary trend from relatively simple forms with semielliptical alteration (e.g. reburrowing, diagenetic processes), parts of
lobes and subcircular spreite fields in the Paleozoic to more the spreiten or entire burrows can lack their lamellae
spiral forms with continuous spreiten in the Mesozoic, and (Fig. 5.35b, d–f) and thus appear to be homogeneously filled
finally complex forms with discontinuous spreite fields and with sediment (Fig. 5.176h).
lobes in the Cenozoic (Seilacher 1977, 2007; Bottjer et al. Similar Trace Fossils: There are few other burrows with
1988; Olivero 1996; Chamberlain 2000; Knaust 2004b; which Zoophycos could be confused. Rhizocorallium is
Zhang et al. 2015). another spreite burrow with a U-shaped morphology, and the
Substrate: Zoophycos is known from siliciclastic and horizontal to inclined R. commune constitutes a spreite
carbonate rocks of originally soft to firm consistency. It commonly filled with fecal pellets C. oblongus (Knaust
preferably occurs in fine-grained sediments. 2013). However, it does not develop a spirally coiled lamina
Appearance in Core: Zoophycos spreite burrows are rel- and therefore is laterally restricted. Furthermore, the spreite
atively easy to recognize in core, although identification to of R. commune is generally thicker than in Zoophycos and is
the ichnospecies level requires knowledge about the bounded by a proportionately large marginal tube. Other
three-dimensional burrow architecture. Assuming Zoophycos constituents of the broader ‘Zoophycos group’ may resemble
(a) (b)
(c) (d)
(e) (f)
(g) (h)
5.36 Zoophycos Massalongo, 1855 175
b Fig. 5.174 Zoophycos in outcrop. Scale bars = 1 cm. a, b Relatively From Knaust (2013) and Knaust et al. (2014), republished with
simple, planar form of Zoophycos in bioclastic limestone. Middle permission of Elsevier and Wiley; permission conveyed through
Permian Khuff Formation (shallow marine, carbonate platform), Copyright Clearance Center, Inc. e Vertical section of Zoophycos
Huqf-Haushi Uplift, Oman. From Knaust (2009c), republished with close to the whorl. Middle Permian Kapp Starostin Formation (mixed
permission of Wiley; permission conveyed through Copyright Clear- siliciclastic carbonate ramp), Akseløya, Svalbard. f Lobate spreite part
ance Center, Inc. c Spiral form of Zoophycos in fine-grained carbonates. in oblique section. Same as in (e). g Lobate spreite on carbonate
Middle Jurassic (slope deposits), southeastern France. See Olivero and bedding plane. Upper Jurassic (Kimmeridgian) Alcobaça Formation
Gaillard (1996), Olivero (2003). d Lobate part of a larger Zoophycos (restricted lagoon), coastal cliffs at Praia do Salgado, western Portugal.
system, preserved on a sandstone bedding plane of the Eocene Grès h Same as in (g), cross section of numerous stacked spreiten
d’Annot Formation (deep marine, turbiditic), southeastern France.
Zoophycos sensu stricto, including Spirophyton and Echi- Whereas Paleozoic Zoophycos occur in nearshore deposits,
nospira. The corkscrew-like Spirophyton consists of tightly Mesozoic forms are common on the shelf and later also
arranged whorls winding around an axial shaft, and has an move towards deeper marine environments, and Cenozoic
unlobed edge without marginal tube with an upwards bent Zoophycos seem to be restricted to the deep sea. Aside from
margin (Miller 1991; Gaillard et al. 1999; Seilacher 2007). its typical occurrence within the Zoophycos Ichnofacies on
Echinospira is characterized by a semicircular outline with the slope, Zoophycos is frequently encountered in lagoonal
numerous long, narrow U-shaped spreite burrows that are environments (Fig. 5.176h).
related to a central cylindrical structure. Echinospira is Ichnofacies: Zoophycos is the namesake of the Zoophy-
common in Upper Cretaceous to Miocene deep-marine cos Ichnofacies, which as introduced by Seilacher (1967)
deposits. Individual spreiten laminae of Zoophycos might be was positioned on the continental slope between the
critical to distinguish from elongate sections of the cylin- shallow-marine Cruziana Ichnofacies and the deep-marine
drical and meniscate backfilled burrow Taenidium, in which Nereites Ichnofacies. In many basins, Zoophycos is a con-
case several angles of observation are necessary to differ- stituent of both these adjacent ichnofacies as well.
entiate the two. Age: Zoophycos in the broad sense is a long-ranging trace
Producers: Originally interpreted as marine algae (Mas- fossil with worldwide distribution perhaps from the Cam-
salongo 1855; Olivero 2007) and later as remains of brian (Alpert 1977; Jensen 1997) and certainly from the
sedentary marine worms (Plička 1968), most workers now Ordovician to the Holocene (Wetzels 2008). The report of
agree that the spreite burrow Zoophycos results from the the oldest Zoophycos from the Early Cambrian (Sappenfield
feeding activity of a vermiform animal (Wetzel and Werner et al. 2012) must probably be rejected because of its likely
1981). However, what kind of worm has produced inorganic origin.
Zoophycos remains debatable, and probably organisms of Reservoir Quality: Zoophycos is a common constituent
different phyla must be considered with regard to particular of mud-dominated facies such as lagoonal deposits (e.g. in
forms of this complex trace fossil. Among them, polychaetes the Khuff Formation, see Fig. 5.176h), which can be
(Bischoff 1968, Knaust 2009c), echiurans (Kotake 1992) and intensively perforated by complex spreite burrows. These
sipunculans (Wetzel and Werner 1981; Olivero and Gaillard burrows are actively filled with grainy (peloidal and oolitic)
2007) are good candidates. material introduced from outside the burrows. This phe-
Ethology: The deposit-feeding behavior of a vermiform nomenon is related to the behavior of the Zoophycos
animal was the preferred interpretation of Zoophycos for a maker, supposedly a worm-like animal, which excavates
long time, until particular features of some forms suggested mud in the subsurface, transports it to the seafloor, and fills
alternative interpretations, including the incorporation of its burrow with grainy material from the surface. This
sediment from the seafloor into the burrow by different model was discussed by Kotake (1989), Bromley (1991)
processes (Löwemark 2012). It is likely that the broad and Löwemark et al. (2004), and the resulting introduction
morphological variability of Zoophycos is a reflection of of grainy material in the host rock was recognized by
contrasting ethologies. Pemberton and Gingras (2005) and Knaust (2009a). In the
Depositional Environment: Zoophycos can be regarded as given example, permeability is created by the activity of
a marine trace fossil. Zoophycos (sensu lato) has been the Zoophycos producer, which turns the otherwise tight
proved to be a trace fossil with an evolutionary history mudstone into a reservoir (Knaust 2009a, 2014a;
through the Phanerozoic (Seilacher 1977, 2007; Bottjer et al. Fig. 5.177). A similar situation does also occur in some
1988; Olivero 1996; Neto de Carvalho and Rodrigues 2003). chalk reservoirs of the North Sea.
5.37 Diffuse Bioturbate Texture 177
b Fig. 5.175 Sketches of representative examples of Zoophycos as by successively overlapping U-shaped burrows. Similar forms were
described in the literature. Not to scale. a Helicoidally coiled form with described under the ichnogenus name Echinospira Girotti, 1970. Upper
semicircular to highly lobate spreiten, suggesting upwards construction. Cretaceous to Miocene limestone (deep marine), New Zealand. After
Late Quaternary (deep marine), Celebes Sea. After Löwemark et al. Ekdale and Lewis (1991), reprinted by permission of the publisher
(2004). b Z. rhodensis, a large spiral form comprising a skirt-like zone (Taylor & Francis Ltd., http://www.tandfonline.com). e Simple, lobate
of spreite surrounded by a zone of numerous marginal lobes. Plan view form with planar spreiten extending from a vertical shaft. Middle
(top) and side view (bottom). Pliocene carbonates (deeper marine), Triassic limestone (carbonate ramp), Thuringia, Germany. After Knaust
Rhodes, Greece. After Bromley and Hanken (2003), republished with (2004b), republished with permission of Wiley; permission conveyed
permission of Elsevier; permission conveyed through Copyright through Copyright Clearance Center, Inc. f Coiled protrusive spreite
Clearance Center, Inc. c Four morphotypes of Zoophycos with burrow with convolute margin and long lobes. The spreite is cut by a
increasing complexity. Lower Jurassic to Upper Cretaceous (shelf-to- tubular structure. Side view (top) and plan view (bottom). Upper
basin limestones), southeastern France. After Olivero (2003), repub- Cretaceous White Chalk (shelf), Denmark and southernmost Sweden.
lished with permission of Elsevier; permission conveyed through After Bromley et al. (1999).
Copyright Clearance Center, Inc. d Coiled retrusive spreite burrow built
5.37 Diffuse Bioturbate Texture various lithologies, where sediment consistency is highly
flexible and thus allows for multiple textural reorganizations.
Morphology, Fill and Size: The process of bioturbation not Appearance in Core: The term bioturbate texture is typ-
only results in more or less well defined discrete trace fossils, ically applied to moderately to completely bioturbated rocks
but often leads to an intensely or completely reworked sedi- that do not allow for systematic description and interpreta-
ment with a diffuse texture (e.g. mottled fabric), or even tion of individual trace-fossil components. While in some
complete homogenization of the substrate after repeated cases discrete burrow features are still intact and visible
bioturbation. In that case, the resultant rock contains a diffuse (Figs. 4.5 and 5.121c), in other examples the sediment was
bioturbate texture (Richter 1952; Frey 1973; Frey and subject of repeated bioturbation which resulted in a
Pemberton 1990, 1991b). Although some burrows are still homogenized texture (Fig. 5.179). Yet another example
discernible, they hardly can be identified on an ichnotaxo- contains cryptic bioturbate texture, in which the original
nomical basis (Fig. 5.178). If such remains of burrows are bedding features are still partly intact but were disturbed by
preserved, their size can correspond to the size of all major the burrowing animal (Figs. 5.179d, e and 5.180).
tracemakers and would fall into the two process-related cat- Depositional Environment: Bioturbate texture is com-
egories of macrobioturbation (burrow diameter or width monly related to environments with favorable conditions for
>1 mm) and meiobioturbation (burrow diameter or width colonization, such as good oxygenation, food availability and
between 0.06 and 1 mm). Diffuse bioturbate textures with less sediment starvation (e.g. offshore environments, flooding
discernible burrows (typically of smaller size and as a result of events), but may also occur in conjunction with stressed
the burrowing method) and well expressed physical sedi- environments where rapid and widespread colonization of a
mentary structures may also be referred to cryptic bioturbate limited number of species takes place (e.g. lagoonal deposits).
textures, which results from cryptic bioturbation (Howard and Ichnofacies: Bioturbate texture may occur in a wide range
Frey 1975, 1985; Pemberton et al. 2001, 2008). of ichnofacies, especially the Nereites, Cruziana and Sko-
Ichnotaxonomy: Even highly bioturbated sedimentary lithos ichnofacies (e.g. piperock) within the marine realm.
rocks sometimes reveal burrow parts that can be attributed to Paleosols within the continental Scoyenia Ichnofacies often
particular ichnotaxa. Completely homogenized substrate can also show bioturbate texture.
experience repeated colonization by benthic organisms, Age: Bioturbate texture occurs throughout the Phanerozoic.
which in turn leads to so-called elite trace fossils (Bromley Reservoir Quality Bioturbate texture may have a strong
1996), which may overprint the pre-existing bioturbate tex- impact on reservoir quality (e.g. Knaust 2014a). In most
ture. Bioturbate textures with a more organized style (e.g. cases, the reservoir quality is diminished because intense
tiering, discrete trace fossils) revealing insights into the bioturbation leads to the incorporation of fine-grained mate-
colonization process of the substrate are commonly dealt rial in form of mud particles and fecel material. In some case,
with as ichnofabrics (Ekdale et al. 2012). however, the contrasting effect takes place, if coarser-grained
Substrate: Bioturbate texture is common in softground, particles are churn with a dense matrix and increase vertical
looseground and occasionally firmground substrates of connectivity (e.g. cryptic bioturbate texture, Fig. 5.180).
(a) (b) (h)
(c)
(d)
(e)
(f)
(g)
5.38 Plant Roots and Their Traces 179
b Fig. 5.176 Zoophycos in sectioned core. Scale bars = 1 cm. a Homo- lamellae with different composition, mainly alternating between mud
geneous sandstone with dish-and-pillar structures due to rapid dewa- and glauconitic sand. Upper Cretaceous (Maastrichtian) Springar
tering. Spreiten are readily visible due to their partial fill of black mud Formation (deep marine, fan system), Norwegian Sea (well
and the resulting color contrast with the white to gray sandstone. Some 6604/10-1, ca. 3628.5 m). f Marly limestone with three successive,
conical spreiten indicate close proximity to the axis of a Zoophycos horizontal spreite burrows, overprinting extensively bioturbated back-
burrow system. Lower Cretaceous (Albian, deep marine, channel ground sediment. Lower Cretaceous (Berriasian) Åsgard Formation
system), off Tanzania. b Homogeneous sandstone with conical and (carbonate shelf), Johan Sverdrup Field, Norwegian North Sea (well
planar spreiten, the latter displaying the marginal tube to the right. 16/5-2S, ca. 1951.5 m). g Arenitic limestone with a thick spreite
Lower Cretaceous (Albian, deep marine, channel system), off Tanzania. section, consisting of alternating dark-gray and light-gray lamellae.
c Sandy debrite with two planar spreiten consisting of alternating mud Lower Cretaceous (Berriasian) Åsgard Formation (carbonate shelf),
and sand lamellae, incorporated glauconite grains and sandy fecal Johan Sverdrup Field, Norwegian North Sea (well 16/5-2S, ca.
pellets. Upper Cretaceous (Maastrichtian) Springar Formation (deep 1945.5 m). h Micritic limestone constituting a Zoophycos ichnofabric
marine, fan system), Gro Discovery, Norwegian Sea (well 6603/12-1, with spreite burrows partly diagenetically replaced by anhydrite (white
ca. 3724.5 m). d Ripple-laminated glauconitic sandstone with undu- color). The top is erosional truncated and overlain with grainstone, also
lating spreiten, which are enhanced in visibility by diagenetic iron containing Zoophycos. Upper Permian Khuff Formation (carbonate
staining. Upper Cretaceous (Campanian) Nise Formation (deep marine, platform with open lagoon), South Pars Field, Persian Gulf, Iran (well
channel-levee system), Aasta Hansteen Field, Norwegian Sea (well SP-9, 3097.3-3097.6 m). After Knaust (2009a, 2014a), republished
6707/10-1, ca. 3050.1 m). e Detail of a spreite showing discrete with permission of GulfPetroLink and EAGE
5.38 Plant Roots and Their Traces Substrate: Plants can grow on different substrates
including soft and hard rock of different nature, and not only
Morphology, Fill and Size: Fossil plant roots and their siliciclastics and carbonates. By their nature, plant roots are
traces can appear in a broad range of morphology and size, commonly associated with various kinds of paleosols,
which reflects the variability in shape, size and behavior of including caliche in carbonate rocks.
their tracemakers (Retallack 1988; Fig. 5.181). The fila- Appearance in Core: The diverse appearance of roots and
mentous or tubular roots and root traces range in diameter root traces in rocks depends on the age of the rock (evolu-
from millimeter- to meter-size and taper downwards a few tionary aspect), paleoenvironment (kind of plants, substrate),
centimeters up to several meters (Fig. 5.182). The diage- and diagenesis (preservational aspect) (Fig. 5.181). In core,
netic history and maturity of rooted surfaces and paleosols many roots appear in form of unbranched or downward-
results in different styles of preservation, ranging from branched, irregularly distributed features (Fig. 5.183, see
body preservation or permineralization to deflection of also Fig. 5.28b, d). Their fill can be coaly, carbonaceous,
bedding planes (Pfefferkorn and Fuchs 1991). Rhizocon- sandy or a combination of these. A typical preservation is
cretions (or rhizoliths) are diagenetically encrusted root sand-filled root traces with a thin carbonaceous lining. Other
systems and have a high preservation potential (Owen et al. roots can be enhanced by diagenesis to build rhizoconcre-
2008). tions, some of which hardly reveal the remaining root
Ichnotaxonomy: Fossil plant roots are covered by the structure at their center.
International Code of Nomenclature for Algae, Fungi, and Similar Trace Fossils: Plant-root traces are often the sub-
Plants and can be named independent from other parts of the ject of confusion with morphologically similar animal bur-
plant (e.g. Uchman et al. 2012). There is a transition between rows, particularly in cases where they penetrate marine
fossil plant roots (body fossils) and root traces (trace fossils). substrates (Curran 2015). The simple vertical burrow Sko-
Fossil root traces are fossilized work of organisms and thus lithos is a good candidate for confusion with root traces, and
are covered by the International Code of Zoological some Skolithos actually originate from root penetrations
Nomenclature, but so far only have received few names, (Gregory et al. 2006; Knaust 2014a), while tiny rootlets can be
such as Rhizoichnus D’Alessandro and Iannone, 1982. mistaken for small burrows (e.g. Chondrites, Bornichnus and
Attempts to classify fossil root plants and their traces have Virgaichnus). More complex root traces, for instance those
been proposed by Klappa (1980), Pfefferkorn and Fuchs radiating from a tree stem, may resemble complex burrow
(1991), Bockelie (1994), Wright et al. (1995) and White and systems such as Phoebichnus (e.g. Gregory et al. 2004).
Curran (1997). The classification key proposed by Bockelie Producers: Various groups of plants are potential pro-
(1994) is mainly based on Mesozoic core material from the ducers of root traces (Bockelie 1994), although some com-
Norwegian North Sea. The hierarchical structure of that key pound trace fossils and paleosols can be formed by plant and
involves sediment fill, presence and complexity of branch- animal (e.g. insect) interactions (e.g. Gregory et al. 2004;
ing, size, orientation and morphology. Strullu-Derrien et al. 2012).
Fig. 5.177 Impact of Zoophycos bioturbation on reservoir quality. and feeds the subsurface mud, defecates on the sea floor and fills its
A lagoonal mudstone unit is commonly tight if not bioturbated (upper burrow with grainy (oolitic) material (refuse dump model in the lower
images, core image width is 9.8 cm). However, complete bioturbation image; after Löwemark et al. 2004). In this particular case, the existing
and discrete Zoophycos burrows in the deep tier transfer the mudstone flow unit within the grainstone facies extends upwards with ca. 30 m.
barrier into a mudstone-wackestone flow unit (images in the middle, Upper Permian Khuff Formation (carbonate platform with open
core image width is 9.8 cm). Reason for this is the behavior of the lagoon), South Pars Field, Persian Gulf, Iran. From Knaust (2009a),
Zoophycos producer, a supposed worm-like animal, which excavates republished with permission of GulfPetroLink
5.38 Plant Roots and Their Traces 181
(a) (b)
(c) (d)
Fig. 5.178 Bioturbate texture in outcrop. Scale bars = 1 cm. burrows visible, probably belonging to Macaronichnus segregatis.
a Oblique section with a few discernible burrows in heterolithic Upper Cretaceous (Cenomanian) Arnager Greensand Formation,
sandstone-siltstone (shallow marine), in which discrete Scalichnus and Arnager near Rønne, Bornholm, Denmark. d Vertical section of sand
Schaubcylindrichnus are visible. Upper Jurassic (Early Kimmeridgian), with high amount of bioturbation overprinting partly preserved cross
Brora (Lothberg Point area), Scotland, UK. b Bedding-plane view of a bedding (lower part). Some discrete Teichichnus rectus and Ophiomor-
micritic limestone (carbonate platform) with abundant pencil-like pha nodosa are visible. Lower Cretaceous (Berriasian) Robbedale
burrows resembling Planolites. Middle Triassic (Anisian) Muschelkalk Formation (shallow marine), Madsegrav near Rønne, Bornholm,
Group, south of Roda, southern Spain. c Vertical section of totally Denmark. See Nielsen et al. (1996)
bioturbated glauconitic sand (shelf) with some horizontal tubular
Ethology: Substrate penetration by roots happens for the (paralic) environments including swamps, lagoons and tidal
purpose of plant stabilization, living and obtaining water and flats (e.g. Whybrow and McClure 1980; Knaust 2009a). Rooted
nutrition. horizons are important indicators of subaerial exposure and thus
Depositional Environment: Plants commonly colonize ter- can be used to delineate sequence boundaries (Husinec and
restrial and aquatic environments in continental setting, such as Read 2011) as well as unconformities (Fig. 2.6a).
alluvial, fluvial, lacustrine and eolian deposits (e.g. Glennie Ichnofacies: A wide range of continental and marginal-
and Evamy 1968; Kraus and Hasiotis 2006; Knaust 2015b). marine ichnofacies typically contains plant roots, for
Mangrove and other roots also occur in marginal-marine instance Scoyenia and Psilonichnus ichnofacies.
(a) (b) (c)
(d) (e)
(f)
5.39 Borings 183
b Fig. 5.179 Bioturbate texture in sectioned core. Scale bars = 1 cm. disturbed by numerous sand-filled burrows (Planolites). The sandstone
a Completely bioturbated argillaceous sandstone containing Teichich- includes vertical sand-filled Skolithos burrows (weakly visible). Upper
nus and Phycosiphon. Upper Jurassic (Oxfordian-Kimmeridgian) Jurassic (Oxfordian) Heather Formation (shelf turbidites), Fram Field,
Spekk Formation (shelf), Norwegian Sea (well 6406/12-1S, ca. Norwegian North Sea (well 35/11-11, ca. 2586.5 m). e Argillaceous
3629.9 m). b Intensely bioturbated sandstone (with Planolites?), in sandstone with cryptic bioturbate texture as revealed by many tiny
which remains of the original bedding are poorly preserved. Upper (meiobenthic) burrows. Upper Jurassic (Oxfordian-Kimmeridgian)
Jurassic (Oxfordian-Kimmeridgian) Rogn Formation (offshore sand Sognefjord Formation (marginal marine, deltaic), Vega Field, Norwe-
bar), Norwegian Sea (well 6406/12-1S, ca. 3622.9 m). c Moderately gian North Sea (well 35/11-6, ca. 3184.2 m). f Wholly bioturbated,
bioturbated cross-bedded sandstone. Middle Jurassic (Callovian) Fens- homogenized sandstone with diffuse burrow structures. Upper Jurassic
fjord Formation (tidally influenced delta), Gjøa Field, Norwegian North (Oxfordian-Kimmeridgian) Heather Formation (submarine fan delta),
Sea (well 36/7-1, ca. 2393.1 m). d Argillaceous sandstone with cryptic Fram H-Nord Field, Norwegian North Sea (well 35/11-15ST2, ca.
bioturbate texture, in which the mud laminae appear to be intact but are 2975.0 m)
Age: The oldest records of primitive plants are from the phoronid worms) (Taylor and Wilson 2003; Bromley 2004;
Ordovician, but diversification and complexity gradually Figs. 5.184 and 5.185).
increased through Silurian and Devonian time. From the Substrate: Borings occur in hard substrates that can be of
Devonian until today, root traces have been common con- lithic (e.g. rocks; skeletal material such as shells, corals,
stituents of the rock record. stromatolites, etc.), xylic (i.e. wood), or osteogen (i.e. bone)
Reservoir Quality: Different preservation of plant roots nature (Taylor and Wilson 2003). Lithified substrates can be
may result in contrasting behavior with respect to reservoir formed by the exhumation of deeply buried sediment or by
quality and performance. For instance, large and sand-filled synsedimentary lithification of sediment (Savrda 2012).
roots in a tight matrix considerably enhance vertical com- Appearance in Core: Borings are relatively easy to recog-
munication within the reservoir (Knaust 2014a), while car- nize in core, although their assignment to particular ichno-
bonaceous roots of similar shape and size influence the genera often remains ambiguous as the whole morphology is
reservoir in the opposite way. not known. However, serrated surfaces, sharp boundaries,
cross-cut particles and passive fills are unique features that
prove a bioerosion nature (Fig. 5.186). Bioeroded surfaces
5.39 Borings of hardgrounds are often discontinuity surfaces (such as
omission surfaces and unconformities) and appear as rather
Morphology, Fill and Size: Borings belong to a group of trace sharply defined boundaries with contrasting lithologies
fossils that comprises excavations made by organisms in hard below and above. Because such surfaces often became
substrate in contrast to burrows, which originate in soft or firm subject of prolonged subsequent reworking by currents, they
substrate. They appear with different shapes and degrees of are commonly altered, stained, and disrupted or eroded.
destruction, and include irregular morphologies, networks, Consequently, reworked litho- and bioclasts may also show
pouches, grooves, as well as clavate, bulbous, tubular and evidences of repeated bioerosion phases in form of one or
clubbed forms (Figs. 5.184 and 5.185). The margin of a bor- more generations of borings.
ing, which may be lined, is typically sharp and well-defined, Similar Trace Fossils: A continuous transition exists
creating a high contrast between the host rock and its fill. between cohesive (firm) and hard substrates, and firm-
Borings are passively filled with sediment differing from the grounds may also contain incipient borings characterized by
host rock, occur as open cavities, or are cemented. A wide size sharp boundaries and passive fill (Knaust 2008; Knaust and
range of borings is known, commonly from micrometric to Dronov 2013). Therefore, a clear distinction between both
centimetric in size, with 1 mm as the arbitrary threshold size trace-fossil categories (e.g. burrows in firm and borings in
for the distinction of microborings and macroborings. hard substrates) is not always possible. This is particularly
Ichnotaxonomy: Bioerosion trace fossils (including true in cases where the trace fossils have similar shapes
borings) comprise more than hundred ichnogenera, which and/or were produced by similar kinds of tracemakers that
correspond to about 17% of all invertebrate trace fossils are capable of both, burrowing and bioeroding (e.g. some
(Knaust 2012a). Most common and widespread macrobor- polychaete and bivalve species).
ings are Entobia (borings with networks of chambers and Producers: Many groups of organisms comprise bio-
canals made by sponges), Gastrochaenolites (clavate bi- eroding species, of which cyanobacteria and algae, sponges,
valve borings in lithic substrates), Teredolites (clavate polychaetes, sipunculides, bivalves, echinoids, cirripedes
bivalve borings in lignic substrates), Trypanites and and bryozoans are most relevant for the origin of borings
Palaeosabella (tubular borings made by worms), Rogerella (Warme 1970; Taylor and Wilson 2003; de Gibert et al.
(pouches made by cirripeds), and Talpina (networks of 2012; Tapanila and Hutchings 2012).
5 cm
1 mm
Fig. 5.180 Cross-bedded sandstone, as seen in thin section, reveals Cretaceous (Maastrichtian) Springar Formation (deep marine, fan
disrupted mud-rich laminae due to the process of cryptobioturbation. system), Norwegian Sea (well 6604/10-1)
This phenomenon results in increased reservoir quality. Upper
5.39 Borings 185
Fig. 5.181 Types of rooting structures based on appearance. After parallel or irregular roots. e Fibrous root system with radial root
Pfefferkorn and Fuchs (1991), republished with permission of arrangement. f Vertical or oblique lateral roots coming from surface
Schweizerbart (www.schweizerbart.de/journals/njgpa). a Millime- root, for instance buttress. g Shallow horizontal roots or rhizomes with
ter-thin crust on rocks. b Millimeter-thin crust on sediment. c Thin lateral roots. h Tap root system
layer affected by rhizoids. d Fibrous roots system with more or less
Ethology: Most bioeroders produce their borings for Trypanites Ichnofacies (Frey and Seilacher 1980), although
dwelling (domichnia) by mechanical or chemical bioerosion, transitions to firmgrounds in the Glossifungites Ichnofacies
or a combination of both. (Seilacher 1967) may occur. Bromley and Asgaard (1993)
Depositional Environment: The occurrence of borings proposed to replace or subdivide the Trypanites Ichnofacies
and other bioerosion traces depends on the availability of with the Entobia Ichnofacies for deep-tier borings and the
suitable hardground to be colonized. Thus, preferred loci for Gnathichnus Ichnofacies including superficial sculptures, a
enhanced bioerosion include reefs, bioherms and biostromes concept which still has been debated (MacEachern et al.
(Tapanila and Hutchings 2012), rocky shorelines (de Gibert 2012; de Gibert et al. 2012; Knaust et al. 2012). Borings in
et al. 2012; Fig. 5.187), and shallow-marine carbonate marine woodgrounds are characterized by the Teredolites
platforms (Knaust et al. 2012; Fig. 5.188), although local Ichnofacies (Bromley et al. 1984). In continental settings,
hardgrounds due to synsedimentary cementation (e.g. in borings occur in a variety of unspecified ichnofacies.
delta-front or deep-marine environments) must be also Age: Borings are known from the Early Cambrian (James
considered. Bioeroded surfaces can be indicators of et al. 1977) to the Holocene (Tapanila and Hutchings 2012).
exhumation or synsedimentary lithification with low or no Reservoir Quality: Due to their tightness, hardgrounds
sedimentation over a longer period of time. Thus, borings commonly act as barriers and baffles to fluid flow and can
are valuable indicators in the recognition of omission sur- contribute to layered reservoirs. Similar to structural ele-
faces and unconformities (Fig. 2.6a). Borings in continental ments such as fractures, borings have also the ability pene-
environments are known from living and dead wood, seeds trating such tight horizons and to enhance the vertical
and other plant material (e.g. Sutherland 2003; Feng et al. connection for fluid flow. Passive fill with sediment grains or
2010; Genise et al. 2012). even open borings just contribute to the enhancement of this
Ichnofacies: Marine, bioeroded, rocky or skeletal hard- effect, which is comparable to the firmground burrows of the
grounds are typically associated with the substrate-controlled Glossifungites Ichnofacies (Gingras et al. 2012b).
(a) (b)
(c) (d)
(e) (f)
Fig. 5.182 Plant-root traces in outcrop. a Sandstone bedding plane regressive eolianite. Near Whale Point, North Eleuthera, Bahamas. From
with a large tabular root system with bifurcations. Eocene Aspelintoppen Knaust et al. (2012), republished with permission of Elsevier; permis-
Formation (fluvial), Brongniartfjellet, Van Keulenfjorden, Svalbard. sion conveyed through Copyright Clearance Center, Inc. e Large,
b Vertical section of coaly mudstone (marsh) overlying clean sandstone cylindrical, vertical rhizolith in calichified (pedogenized) dolomitic
(shoreface) with deeply penetrating root traces. Middle Jurassic preservation and subsequent cementation around the root structure with
Scarborough Formation (marginal marine), coastal cliff near Scarbor- calcite (rhizoconcretion). Upper Triassic Kågeröd Formation (fluvial),
ough, Yorkshire, UK. c Sandstone bedding plane (paleosol) with vertical Bornholm, Denmark. Scale bar = 1 cm. From Knaust (2015b).
root traces modified by a diagenetic halo. Same locality as in (a). Scale f Calichified root system within eolian sand. Pleistocene, coastal cliff
bar = 1 cm. d Dense occurrence of rhizomorphs in a Late Pleistocene at Cap Ghir, western Morocco. Scale bar = 5 cm
5.39 Borings 187
(a) (b) (c)
(e)
(f)
(d)
b Fig. 5.183 Plant-root traces in sectioned core. Scale bars = 1 cm. 7220/7-2S, ca. 1152 m). d Paleosol with large system of branched and
a Branched root system with coaly substrate originating from a thin sand-filled root system. Middle Jurassic (Bajocian-Bathonian) Hugin
coal seam and penetrating heterolithic sandstone with marine burrows. Formation (marginal marine), Norwegian North Sea (well 15/6-4, ca.
Lower Jurassic (Rhaetian to Pliensbachian) Åre Formation (tidal flat), 10631 ft). e Large vertical root trace penetrating inclined,
Skuld Field, Norwegian Sea (well 6608/10-14S, ca. 2689.5 m). b Large ripple-laminated sandstone with marine bioturbation. Upper Triassic
fragments of root traces (partly sand-filled and partly coaly) penetrating (Rhaetian) Tubåen Formation (marginal marine), Skavl Discovery,
ripple-laminated sandstone. Lower Jurassic (Hettangian to Sinemurian) Norwegian Barents Sea (well 7220/7-2S, ca. 1152 m). f Individual
Nansen Formation (marginal marine), Norwegian North Sea (well vertical roots partly sand-filled (resembling Skolithos) and partly
25/10-11T2, ca. 4317.5 m). c Dense system of root traces of variable containing carbonaceous material. Upper Triassic (Rhaetian) Tubåen
shape, size and fill. Upper Triassic (Rhaetian) Tubåen Formation Formation (marginal marine), Skavl Discovery, Norwegian Barents Sea
(marginal marine), Skavl Discovery, Norwegian Barents Sea (well (well 7220/7-2S, ca. 1152 m)
(a) (b) (c)
(d) (e)
Fig. 5.184 Line drawings of representative examples of borings, with large rounded chambers and interconnections of the sponge
illustrating their morphology, size, substrate, producer, and ichno- Cliona vermifera in coral (ichnogenus Entobia). b Elongate, narrow
genus. Scale bars = 1 cm except in (d) = 10 cm. Modified after boring of the polychaete Hypsicomus elegans in coral (ichnogenus
Bromley (1978; a, b and d), republished with permission of Elsevier; Trypanites). c Clavate boring of the bivalve Gastrochaena dubia (with
permission conveyed through Copyright Clearance Center, Inc.; Kelly producer shown) in limestone (ichnogenus Gastrochaenolites).
and Bromley (1984, c), republished with permission of Wiley; d Grooves bored by the echinoid Echinometra lucunter in limestone
permission conveyed through Copyright Clearance Center, Inc.; and (ichnogenus Ericichnus). e Clavate borings of the bivalve Martesia
Bromley et al. (1984, e), © Paleontological Society, published by sp. in woodground (ichnogenus Teredolites)
Cambridge University Press, reproduced with permission. a Boring
5.39 Borings 189
(a) (b)
(c) (d)
(e) (f)
b Fig. 5.185 Borings and other bioerosion trace fossils in outcrop. Scale Hajar Mountains, Oman. c Micritic hardground (lower part) in vertical
bars = 1 cm. a Thin hardground layer (Zhelty Bed, ochre) within an section, with large B. triadicus and needle-like T. weisei. The top
arenitic limestone (Dikari Limestone, pinkish-gray) in vertical section, surface is eroded and overlain by arenitic limestone (upper part).
with a complex Balanoglossites triadicus ichnofabric, including Middle Triassic (Anisian) Jena Formation (carbonate ramp), Thuringia,
burrows and borings from different phases of colonization. Middle Germany. d Same hardground as in (c) with deeply penetrating T.
Ordovician (Dapingian) cool-water carbonates, St. Petersburg Region, weisei and one specimen of T. fosteryeomani (next to the left margin).
NW Russia. From Knaust et al. (2012), republished with permission of e Surface of a micritic limestone clast with openings of Caulostrepsis
Elsevier; permission conveyed through Copyright Clearance Center, isp. (small) and Gastrochaenolites isp. (large). Eocene conglomerate
Inc.; and Knaust and Dronov (2013), republished with permission of with reworked Cretaceous limestone clasts, southeastern France.
Springer. b Bedding plane of micritic limestone (dark blue) containing f Another clast with large G. torpedo of different orientation
Entobia isp. (large rounded chambers) and Trypanites weisei (minute (longitudinal and cross sections) due to clast movement and repeated
circular cross sections), both filled with dolomite (ochre). Cretaceous colonization. Same locality as in (e)
(Albian to Cenomanian) Natih Formation (carbonate platform), Misfah,
(a) (b) (c)
(d) (e)
C D
5.39 Borings 191
b Fig. 5.186 Borings in sectioned core. Scale bars = 1 cm. a and layers, which underwent synsedimentary lithification and bioerosion.
b republished with permission of Wiley and EAGE; permission Reworked clasts are concentrated in the turbidite bed above the
conveyed through Copyright Clearance Center, Inc. a Reworked and mudstone layers, one clast (arrow) shows minute borings (Entobia) all
bioeroded limestone clast within grainy substrate. Note the bioerosion around and a larger shaft. Lower Cretaceous (Albian, deep marine,
on upper and lower surface of clast. Lower Triassic Khuff Formation channel-overbank), off Tanzania. d Omission surface (arrow heads)
(storm-reworked sand shoals and sand waves, inner to outer carbonate with reworked phosphatic (brown) pebbles and bioerosion (e.g.
ramp), South Pars Field, Persian Gulf, Iran (well SP9, ca. 2918.3 m). Gastrochaenolites, arrow). Upper Jurassic (Tithonian) Draupne For-
From Knaust (2010b, 2014a). b Biolaminated dolomitic limestone mation (shallow marine), Norwegian North Sea (well 25/10-12ST2, ca.
(stromatolite) with bioeroded surface and Gastrochaenolites (arrows). 2123 m). e Reworked sideritic mudclast having an intensely bioeroded
Lower Triassic Khuff Formation (tidal flat, inner carbonate ramp), margin within sandstone matrix. Middle Jurassic (Callovian) Fensfjord
South Pars Field, Persian Gulf, Iran (well SP9, ca. 2909.8 m). From Formation (delta front), Gjøa Field, Norwegian North Sea (well 36/7-1,
Knaust (2010b, 2014a). c Turbiditic sandstone with thin mudstone ca. 2373.5 m)
(a) (b)
Fig. 5.187 Rocky shoreline deposits. Eocene, southeastern France. of clasts with shallow borings (e.g. Caulostrepsis) and deeper
Scale bar = 10 cm (a) and 1 cm (b). a The well rounded limestone penetrations (e.g. Gastrochaenolites), both distinct with their contrast-
clasts of Cretaceous age are fairly bioeroded and embedded in medium- ing passive sand fill
to coarse-grained sandstone of shallow marine origin. b Close-up view
(a) (b)
Ga Tr
Ga
Th Tw
Tf
Fig. 5.188 Omission suite assemblages from two hardgrounds. From enlarged Gastrochaenolites). Upper Cretaceous (Campanian-
Bromley (1975), republished with permission of Springer. a Cretaceous Maastrichtian transition), Belgium. Height of block ca. 8 cm. b Two
chalk in a relatively shallow-marine setting. Pre-lithification Thalassi- hardground surfaces in limestone are plastered with oysters and contain
noides (Th) is prevalent, accompanied with post-lithification Gas- three types of borings, Gastrochaenolites isp. (Ga), Trypanites weisei
trochaenolites (Ga) and Trypanites (Tr). Borings are partly trapped (Tw) and Trypanites fosteryeomani (Tf). Carboniferous/Middle Jurassic
with intraclasts and debris, which are otherwise unpreserved (see unconformity, England, UK. Height of block ca. 4 cm
(a) (b) (c)
(d)
(e) (f) (g)
Fig. 5.189 Examples of common pseudo-trace fossils as encountered complex), Gudrun Field, Norwegian North Sea (well 15/3-9T2, ca.
in sectioned core. Scale bars = 1 cm. a Sandstone with dewatering 4131.55 m). e Muddy debris-flow deposit (debrite) with floating and
structures (pipes) superficially resembles the ichnogenus Skolithos. poorly consolidated sand clasts mimicking a partly bioturbate texture.
Upper Jurassic (Kimmeridgian-Tithonian) Kimmeridgian Clay Forma- Upper Cretaceous (Maastrichtian) Springar Formation (deep marine,
tion (deep marine, fan system), Kingfisher Field, UK North Sea (well fan system), Gro Discovery, Norwegian Sea (well 6603/12-1, ca.
16/8a-4, ca. 13515.6 ft). b Sandstone-mudstone alternation with 3724.5 m). f Sandstone with microfaults resembling vertical burrows
synaeresis cracks, which could be mistaken as burrows. Middle such as Skolithos. Upper Triassic (Rhaetian) Statfjord Group (fluvial),
Jurassic (Bajocian-Bathonian) Tarbert Formation (marginal marine, Johan Sverdrup Field, Norwegian North Sea (well 16/2-11, ca.
tidal flat), Valemon Field, Norwegian North Sea (well 34/11-B-13, ca. 1940.5 m). g Branched tubular structure filled with coarse sand and
4619.4 m). c Small-scale slump fold (recumbent fold) within a tentatively interpreted to be originated from methane or hydrocarbon
mudstone-sandstone alternation, accompanied by other deformation seepage (cold seep) in shallow subsurface position. Such structures, if
structures, somehow resembling a meniscate burrow such as Taeni- regarded in isolation, are easily mistaken as large burrows such as
dium. Eocene (deep marine, channel-levee system), off Tanzania. shafts of Thalassinoides systems. Upper Jurassic (Tithonian) Rogn
d Small sandstone dyke (injectite) within a thin mudstone layer. Upper Formation (lower shoreface), Norwegian Sea, well 6406/12-1S, ca.
Jurassic (Tithonian) Draupne Formation (deep marine, channel-levee 3631.9 m
5.40 Pseudo-trace Fossils 193
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Index
A Carbonate, 14, 30, 37, 48, 52, 54, 57, 60, 63, 70, 71, 101, 110, 116,
Aerobic, 59 117, 120, 148, 153, 160, 162–164, 170, 173, 179, 185, 190, 191
Alcobaça Formation, 175 Cardium Formation, 112
Almargem Formation, 80 Catenarichnus, 30, 70
Amundsen Formation, 43, 92, 106 Catenichnus, 72, 155
Anaerobic, 59 Caulostrepsis, 190, 191
Ancorichnus, 150 Chenque Formation, 62
Annelid, 58, 102, 117, 120, 155 Chondrites, 22, 44, 53, 54, 55, 57−59, 70, 86, 90, 94, 116, 127, 160,
Aquifer, 2, 100, 163 162, 164, 168, 179
Arachnid, 148 Clay Formation, 192
Åre Formation, 84, 109, 119, 166, 188 Colonization, 6, 9, 11, 22, 30, 34, 57, 70, 84, 95, 99, 100, 145, 154,
Arenicolites, 29, 30, 32, 72, 148, 164 177, 190
Arthropod, 8, 90, 102, 111, 117, 144, 150, 155, 160 Complex trace fossil, 9, 27, 79, 80, 102, 173, 175
Artichnus, 33, 34, 36, 37, 66, 70, 155, 158, 160 Composite trace fossil, 9
Åsgard Formation, 57, 63, 162, 168, 171, 179 Compound trace fossil, 9, 179
Aspelintoppen Formation, 139, 140, 186 Conglomerate, 50, 160, 190
Asterosoma, 34, 36, 37, 39, 41, 66, 80, 81, 128, 160, 171 Conichnus, 41, 59−64, 66, 82, 94, 116
Conostichus, 61, 65
Cook Formation, 15, 35, 84, 104, 109, 110, 116, 127, 158
B Coprolite, 5, 6
Balanoglossites, 160, 190 Coprulus, 72, 74, 76, 93, 120
Bathichnus, 164 Core logging, 22, 24
Battfjellet Formation, 68, 70, 87 Crayfish, 48, 50, 52
Beach Formation, 125, 156 Crustacean, 46, 48, 94
Bearpaw-Horseshoe Canyon Formation, 96, 125 Cruziana Ichnofacies, 9, 30, 34, 39, 47, 70, 80, 85, 91, 100, 102, 104,
Ben Nevis Formation, 59, 100, 163 112, 117, 124, 129, 136, 145, 150, 159, 160, 170, 175
Bergaueria, 41–43 Cryptic bioturbate texture, 92, 177, 183
Bichordites, 131, 133 Cryptobioturbation, 184
Bioerosion, 5, 7, 183, 185, 190, 191 Cutler Formation, 139
Bioerosion trace fossil, 5, 183, 190 Cylindrichnus, 36, 65, 67, 70, 71, 92, 109, 116, 128–130, 133, 155,
Biogenic sedimentary structure, 5, 137 171, 181
Bioturbate texture, 2, 5, 13, 14, 22, 25–27, 32, 92, 117, 127, 138, 177, Cylindricum, 148
181, 183, 192
Bioturbation, 2, 5, 6, 11, 14, 18, 21, 22, 24, 26, 59, 70, 85, 92, 97, 100,
104, 113, 116, 117, 124, 127, 130, 136, 138, 144, 148, 159, 177, D
181, 184, 188 Dictyodora, 102
Bivalve, 11, 14, 37, 54, 58, 61, 80, 82, 104, 106, 107, 109, 110, 117, Diplocraterion, 29, 30, 61, 71, 72, 74, 77, 79, 94, 120, 141, 155, 164
118, 144, 145, 155, 158, 183, 188 Draupne Formation, 162, 191, 192
Boring, 183, 185, 188, 190, 191
Bornichnus, 44, 47, 48, 83, 85, 123, 168, 179
Brachiopod, 82, 83, 85 E
Buntsandstein, 31, 122, 139 Echinoid, 131, 138, 188
Echinospira, 173, 175, 177
Echiuran, 74, 175
C Elite trace fossil, 131, 177
Camborygma, 48, 49, 160 Enteropneust, 90, 130, 160
Capayanichnus, 148 Entobia, 183, 188, 190, 191

DOI 10.1007/978-3-319-49837-9
208 Index
Ericichnus, 188 L
Euflabella, 36, 102 Læså Formation, 156
Laevicyclus, 70, 141
Lange Formation, 119
F Letná Formation, 42
Fecal pellet, 6, 9, 58, 61, 72, 76, 120, 122, 124, 172, 173, 179 Limestone, 57, 60, 70, 73, 82, 96, 101, 107, 118, 122, 141, 142, 148,
Fensfjord Formation, 76, 88, 183, 191 149, 153, 154, 156, 160, 162, 167, 168, 170, 171, 177, 179, 188,
Firkanten Formation, 87 191
Footprint, 61 Lingulichnus, 82–85, 128, 141
Fruholmen Formation, 32 Loloichnus, 48
Lophoctenium, 80, 81, 136
Lunde Formation, 51, 140, 153
G
Gastrochaenolites, 11, 183, 188, 190, 191
Glossifungites, 120 M
Glossifungites Ichnofacies, 9, 78, 124, 160, 163, 165, 185 Macrobioturbation, 177
Glyphichnus, 66, 68, 70 Macroboring, 183
Gordia, 86 Macaronichnus, 11, 62, 85, 86, 88, 90, 102, 111, 117, 141, 181
Greensand Formation, 87, 131, 181 Marnoso-arenacea Formation, 167
Grès dʼAnnot Formation, 25, 31, 33, 34, 34, 38, 96, 114, 134, 175 Meiobenthic, 167, 168, 183
Grumantbyen Formation, 25, 70, 87, 92, 101, 131, 170, 171 Meiobioturbation, 177
Gyrolithes, 46, 160, 162 Meissner Formation, 142
Microboring, 183
Mineralization, 59, 117, 124, 163, 167, 179
H Mount Messenger Formation, 68, 96, 114, 134, 136, 138, 142
Hardeberga Formation, 73, 87 Mudstone, 11, 14, 19, 31, 32, 43, 48, 51, 54, 57, 59, 79, 93, 95, 114,
Hartsellea, 52 119, 136, 141, 160, 162, 163, 165, 167, 175, 180, 186, 191, 192
Heather Formation, 32, 70, 92, 97, 116, 119, 133, 171, 183 Muschelkalk, 8, 42, 83, 124, 142, 181
Heimdallia, 102
Hillichnus, 9, 36, 79, 80, 102, 141
Höganäs Formation, 38, 142 N
Holothurian, 6, 30, 34, 36, 37, 70, 74, 107, 148, 158 Nansen Formation, 35, 144, 188
Honaker Trail Formation, 170 Natih Formation, 190
Hugin Formation, 15, 41, 47, 64, 70, 76, 84, 88, 97, 104, 106, 109, 119, Neill Klinter Formation, 110
123, 127, 148, 158, 188 Nemertean, 8, 172
Hydrocarbon, 1, 19, 59, 117, 160, 163, 164, 192 Nereites, 39, 59, 80, 86, 90, 91, 100, 102, 116, 117, 127, 133, 136, 158,
165, 168, 175, 177
Nereites Ichnofacies, 29, 39, 59, 80, 91, 100, 102, 117, 136, 165, 168,
I 175
Ichnoabundance, 21, 24–26 Neslen Formation, 62, 63, 68, 92
Ichnodiversity, 14, 21, 24–26, 59, 116, 159 Ness Formation, 43, 109
Ichnocoenosis [Ichnocoenoses], 5, 6, 32 Nise Formation, 104, 136, 162, 179
Ichnofacies, 1, 5, 6, 9, 13, 21, 27, 29, 30, 32, 34, 39, 44, 47, 52, 59, 61, Nordmela Formation, 41
70, 78, 80, 85, 90, 91, 100, 102, 104, 107, 112, 117, 119, 122,
124, 129, 130, 136, 139, 145, 149, 150, 159, 160, 163, 165, 168,
170, 175, 177, 181, 185 O
Ichnofabric, 1, 2, 5, 6, 9, 11, 13, 14, 19, 21, 22, 24, 35, 41, 47, 57, 58, Ophiomorpha, 9, 18, 44, 46, 48, 49, 64, 72, 77, 93–95, 100, 107, 111,
66, 67, 70, 71, 76, 77, 84, 86–88, 90, 92, 97, 100, 104, 109, 112, 141, 148, 155, 159, 160, 162, 181
113, 116, 117, 119, 125, 127, 130, 134, 138, 144, 148, 150, 153, Oxygen, 13, 59, 74, 91, 124, 131
156, 158, 162–164, 166, 170, 171, 177, 179, 190
Ichnology, 1, 2, 5, 13, 26
Ile Formation, 41, 63, 84, 97, 109, 113 P
Insect, 6, 30, 141, 148, 150, 153, 179 Palaeophycus, 30, 80, 94, 101, 102, 116, 117, 129, 141
Paleosol, 48, 51, 139−141, 149, 177, 179, 186, 188
Palaeosabella, 183
J Paleontology, 2
Jena Formation, 83, 190 Paradictyodora, 102, 104, 107, 155
Parahaentzschelinia, 80, 106, 107, 110, 141
Paratisoa, 164
K Parmaichnus, 160
Kågeröd Formation, 31, 50, 151, 186 Phoebichnus, 109, 111, 179
Kapp Starostin Formation, 101, 156, 175 Pholeus, 48, 94
Keuper, 156 Phycodes, 155
Khuff Formation, 15, 32, 41, 70, 127, 153, 171, 175, 179, 189, 191 Phycosiphon, 15, 43, 58, 90, 112, 114, 116, 117, 133, 138, 183
Kvitnos Formation, 18, 97, 136 Phymatoderma, 53, 54, 58
Index 209
Pilichnus, 44, 58, 168 Spreite, 9, 11, 15, 30, 33–36, 57, 61, 64, 71–74, 76, 80, 90, 102, 104,
Piscichnus, 41 106, 107, 112, 114, 116, 117, 120, 122–124, 136, 154–156, 158,
Planolites, 58, 86, 102, 117–119, 159, 160, 181 159, 162, 164, 172, 173, 175, 177, 179
Plant root, 5, 6, 27, 44, 148, 179, 181 Springar Formation, 11, 18, 77, 97, 148, 162, 179, 184, 192
Polychaete, 6, 9, 14, 30, 33, 36, 47, 48, 58, 70, 72, 74, 78, 86, 102, 104, Statfjord Formation, 148
120, 127, 128, 130, 155, 183, 188 Stellavelum, 102
Polykladichnus, 30, 148, 167 Stelloglyphus, 111
Pragichnus, 57, 58 Stø Formation, 35, 81, 113, 116, 123, 144, 148, 158
Pseudo-trace fossil, 27, 192, 193 Stratigraphy, 13, 14, 145
Psilonichnus, 48, 94, 160, 181
Psilonichnus Ichnofacies, 29, 181
T
Taenidium, 48, 120, 136, 139, 149, 150, 153, 175
R Talpina, 183
Reservoir characterization, 5 Tarbert Formation, 11, 88, 148, 158, 192
Rhizoconcretion, 179, 186 Tatsukushi Formation, 100
Rhizocorallium, 9, 72, 120, 122, 124, 150, 155, 164, 173 Teichichnus, 34, 72, 102, 154, 155, 158, 159
Rhizolith, 179, 186 Teredolites, 183, 188
Rhizomorph, 186 Teredolites Ichnofacies, 29, 185
Robbedale Formation, 46, 62, 96, 181 Thalassinoides, 9, 57, 100, 159, 160, 163
Rogerella, 183 Tier, 58, 88, 137, 138, 150, 151, 180
Rogn Formation, 183, 192 Tiering, 6, 21, 22, 137, 177
Ror Formation, 70, 133 Tilje Formation, 43, 76, 81, 97, 109, 144, 148
Rosselia, 36, 66, 82, 107, 109, 124, 125, 127–129 Tisoa, 72, 120, 163–166
Rutichnus, 58 Tofte Formation, 41, 84
Tosna Formation, 146
Trace-fossil association, 14, 21, 24, 27, 29
S Trichichnus, 148, 167
Saiq Formation, 170 Trichophycus, 155
San Antonio/San Juan Formation, 35 Trypanites, 183, 190
Sandstone, 41, 47, 54, 79, 93, 110, 140, 150, 170 Trypanites Ichnofacies, 21, 29, 188
Scalichnus, 141, 181 Tubåen Formation, 32, 188
Scarborough Formation, 125, 186
Schaubcylindrichnus, 116, 129–131, 133
Scolicia, 120, 131, 136, 138 U
Scoyenia, 52, 119, 138, 139, 150 Udelfangen Formation, 55, 118
Scoyenia Ichnofacies, 102, 141, 150, 177 Unconformity [unconformities], 11, 181, 183, 185
Sea anemone, 44, 45, 128, 148 Urenui Formation, 165
Sedimentology, 13, 26
Sego Formation, 68
Siltstone, 87, 90, 131, 148 V
Siphonichnus, 82, 111, 139, 141, 142, 144, 145, 148 Virgaichnus, 44, 168, 170–172
Sipunculan, 30, 58
Skagerrak Formation, 153
Skolichnus, 57 W
Skolithos, 6, 30, 32, 44, 70, 80, 85, 107, 130, 145, 148, 149, 167, 192 Worm, 14, 30, 33, 36, 78, 86, 89, 117, 160, 175, 180
Skolithos Ichnofacies, 29, 30, 32, 44, 61, 70, 76, 85, 90, 100, 107, 129,
130, 145, 149, 177
Sognefjord Formation, 84, 183 Z
Sorthat Formation, 46 Zavitokichnus, 102
Spekk Formation, 183 Zoophycos, 9, 47, 52, 58, 59, 91, 102, 120, 136, 159, 168, 172, 173,
Spirophyton, 175 175, 179, 180
Spongeliomorpha, 9, 48 Zoophycos Ichnofacies, 29, 47, 91, 159, 165, 175

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Dirk Knaust

Atlas of Trace Fossils in

ISBN 978-3-319-49836-2 ISBN 978-3-319-49837-9 (eBook)

Library of Congress Control Number: 2016958461

© Springer International Publishing AG 2017

Printed on acid-free paper

This Springer imprint is published by Springer Nature

July 2016 Andrew K. Rindsberg

Stavanger, Norway Dirk Knaust

2 Ichnological Basics, Principles and Concepts . . . . . . . . . . . . . . . . . . . . . . . . 5

3 Applications of Trace-Fossil Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

4 Methodology in Ichnological Core Logging . . . . . . . . . . . . . . . . . . . . . . . . . 21

5 Selected Trace Fossils in Core and Outcrop. . . . . . . . . . . . . . . . . . . . . . . . . 27

5.20 Phoebichnus Bromley and Asgaard, 1972 . . . . . . . . . . . . . . . . . . . . . . . 109

Dirk Knaust is Specialist in Sedimentology in Statoil’s Research and Technology unit in

© Springer International Publishing AG 2017 1

© Springer International Publishing AG 2017 5

Burrow Trail Trackway

Imprint Boring Fecal pellet (coprolite)

Bioerosion Fecal pellets

Arachnids Plant roots

Rhizocorallium commune Rhizocorallium jenense

References Knaust D (2007) Meiobenthic trace fossils as keys to the taphonomic

© Springer International Publishing AG 2017 13

3.2 Stratigraphy role in the development of reservoir quality, including rock

Subaerial Maximum regressive Ravinement Regressive surface of Maximum flooding

Fluvial to Carbonate shoal Shoreface Marginal-marine

Sandy tidal flat

Fig. 3.3 Important processes 1. Sedimentary 2. Ichnological

- Lithological composition - Trace fossils

- Cementation and dissolution - Fractures

Average: 23,9; 4,532

Sandstone (no bioturbation) Sandstone (bioturbated)

Fig. 3.5 Architectural elements

References East Greenland. Grønlands Geologiske Undersøgelse, Rapport 49:

© Springer International Publishing AG 2017 21

80 % 4 4.3 Analysis of Burrow Size and

3 After logging key trace fossils, additional information can be

(a) (b) (c)

Grain size range Cl Si VFS FS MS CS VCS Gr Pe

% area covered 0.1 1 10 100

Order of deposition and

Cylindrical, unbranched and

2 Subhorizontal, gently curved burrows with a thick

(a) Ichnodiversity (b) Ichnoabundance

(a) (b) (c)

© Springer International Publishing AG 2017 27

More common Dominant Dominant

Orientation Subhorizontal Subvertical

Branching Unbranched Branched Unbranched Branched

Ichnotaxa in red font may be branched or unbranched

Teredolites Trypanites Glossifungites

Fig. 5.4 Morphological

Ichnotaxonomy: About 15 ichnospecies of Arenicolites Skolithos (vertical) or Palaeophycus (horizontal). Com-

5.3 Artichnus Zhang et al., 2008

Morphology, Fill and Size: Artichnus is a wide, J-shaped

Fig. 5.10 Artichnus in sectioned (a)

variable in dimension, number, and orientation and may