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Sediment-organism interactions: a multifaceted ichnology



The field of Ichnology bridges the gap between the areas of paleontology and sedimentology, but has connections to many subdisdplines within these areas. Biogenic structures record the behavior of their tracemakers and provide valuable information in paleoecologic and paleoenvhnmental analysis. As in situ ethologic structures, trace fossils or ichnofossils yield valuable insights into the paleoecology of ancient benthic communities and the environmental dynamics of depositional systems. Ichnology is truly a multifaceted field, and a broad selection of its facets is represented in the 28 papers of this volume. The papers are the product of lchnia 2004 the First International Congress on Ichnology, convened by Jorge F. Genise and held from 19 to 23 April 2004 at the Museo Paleontológico Egidio Feruglio in Trelew, Patagonia, Argentina. Seven papers connected with the congress, pertaining ichnotaxonomy, were published separately, in Ichos, volume 13, issue 4, edited by J.F. Genise, R.N. Melchor, RG. Netto, and A.K. Rindsberg. Several symposium volumes, books, and short-course notes haw been published in recent years (Pemberton et al, 2001; Buatois et al., 2002; Kowalewski and Kelley, 2002; Hasiotis, 2002; Kelley et al., 2003; Buatois and Mángano, 2003; McIlroy, 2004; Webby et al, 2004; Miller, 2007; SeiIacher, 20071, and ichnology can be considered a particularly active research area in steady growth. The 28 papers herein are arranged in five groups reveal the broad scope of ichnology. One of the aims of Ichnia 2004 was to gather together ichnologists covering different backgrounds and having different interests. The underlying philosophy of the meeting was to explore the multiple aspects of ichnology, trying to establish links between the different subfields. Accordingly, there was a conscious effort to look for common themes while enjoying diversity at the same time. This book attempts to reflect the spirit of that meem. It is devoted to exploring the potential of biogenic structures in a wide variety of fields, such as paleoecology, sedimentology, sequence stratigraphy, biostratigraphy, and evolutionary paleoecology.
Department of Geography and Geology, University of Copenhagen, Øster Voldgade 10, 1350 Copenhagen, Denmark
Department of Geologic Sciences, University of Saskatchewan, 114 Science Place,
Saskatoon, S7N 5E2, Canada
Museo Paleontológico Egidio Feruglio, Av. Fontana 140, 9100 Trelew, Chubut, Argentina
Universidad Nacional de la Pampa, Av. Uruguay 151, L6300CLB Santa Rosa, La Pampa, Argentina
Sediment–Organism Interactions: A Multifaceted Ichnology
SEPM Special Publication No. 88, Copyright © 2007
SEPM (Society for Sedimentary Geology), ISBN 978-1-56576-129-2, p. 3–6.
The field of Ichnology bridges the gap between the areas of
paleontology and sedimentology, but has connections to many
subdisciplines within these areas. Biogenic structures record
the behavior of their tracemakers and provide valuable infor-
mation in paleoecologic and paleoenvironmental analysis. As in
situ ethologic structures, trace fossils or ichnofossils yield valu-
able insights into the paleoecology of ancient benthic communi-
ties and the environmental dynamics of depositional systems.
Ichnology is truly a multifaceted field, and a broad selection of
its facets is represented in the 28 papers of this volume. The
papers are the product of Ichnia 2004, the First International
Congress on Ichnology, convened by Jorge F. Genise and held
from 19 to 23 April 2004 at the Museo Paleontológico Egidio
Feruglio in Trelew, Patagonia, Argentina. Seven papers con-
nected with the congress, containing ichnotaxonomy, were
published separately, in Ichnos, volume 13, issue 4, edited by J.F.
Genise, R.N. Melchor, R.G. Netto, and A.K. Rindsberg.
Several symposium volumes, books, and short-course notes
have been published in recent years (Pemberton et al., 2001;
Buatois et al., 2002; Kowalewski and Kelley, 2002; Hasiotis,
2002; Kelley et al., 2003; Buatois and Mángano, 2003; McIlroy,
2004; Webby et al., 2004; Miller, 2007; Seilacher, 2007), and
ichnology can be considered a particularly active research area
in steady growth. The 28 papers herein are arranged in five
groups that reveal the broad scope of ichnology. One of the aims
of Ichnia 2004 was to gather together ichnologists covering
different backgrounds and having different interests. The un-
derlying philosophy of the meeting was to explore the multiple
aspects of ichnology, trying to establish links between the
different subfields. Accordingly, there was a conscious effort to
look for common themes while enjoying diversity at the same
time. This book attempts to reflect the spirit of that meeting. It
is devoted to exploring the potential of biogenic structures in a
wide variety of fields, such as paleoecology, sedimentology,
sequence stratigraphy, biostratigraphy, and evolutionary pa-
leoecology. In doing so we hope to reflect this more integrated
view of ichnology that intends to construct, extend, or fortify
existing bridges between different subfields, such as
paleoichnology and neoichnology, vertebrate and invertebrate
ichnology, benthic ecology, coprology, and soft- and hard-
substrate ichnology. A long time ago, a journalist asked a jazz
musician about where jazz was going. The answer was “If I
knew that, I would already be there”. There is an implicit risk in
writing introductions that attempt to detect current trends and
decipher future directions. We tackle that challenge here.
We begin with three papers devoted to concepts and reviews
that derive from some of the invited keynote talks. Although a
comprehensive summary of ichnologic concepts is far beyond the
scope of the book, these three papers provide a state of the art of
some of the different subfields and present innovative ideas that
may contribute to a healthy debate within the field. Labandeira
assesses the fossil record of plant–insect associations and com-
pares ichnodata with body-fossil data. His paper provides a
summary of a field that has experienced an explosive develop-
ment during the last decade. Studies of plant–insect interaction
are essential to understand modern land and freshwater ecology,
and their path through the fossil record provides a wealth of
information that helps to decipher the evolution of terrestrial
ecosystems. Lockley describes the morphodynamics of archosaurs
and the trackways they produce. He emphasizes holistic insights
into the relationships between the trace fossils, the feet, the limbs,
and the whole body. He indicates that his morphodynamic
approach provides a less static way of understanding morphol-
ogy. Seilacher reviews new insights into the principles of
ichnostratigraphy. He notes that although the notion that trace
fossils are useless in biostratigraphy is widespread, there are
many exceptions. In particular, his paper underscores the impor-
tance of some Paleozoic ichnotaxa, such as trilobite trace fossils,
arthrophycids, Oldhamia ichnospecies, and Treptichnus pedum,
the latter being particularly important because it marks the main
divide in the stratigraphic record: the Precambrian–Cambrian
The second group comprises four papers devoted to ethology
and ecology, and thereby linking trace fossils and behavior. This
section emphasizes paleobiologic aspects involved in the produc-
tion of biogenic structures and the paleoecological significance of
ichnofaunas. A prime role is given to careful analysis of indi-
vidual ichnotaxa and the associated taphonomic filters, thereby
stressing the importance of the fossilization barrier. Most of these
papers illustrate the necessity of detailed autecologic studies in
ichnology and demonstrate that trace fossils should be under-
stood in terms of their behavioral significance and that ethology
is the stepping-stone from which to infer paleoecologic param-
eters. It is rather disturbing to realize how limited our under-
standing of many biogenic structures still is. Undoubtedly, this is
an area that needs to be further developed in order to produce
more robust trace-fossil models that can be used in paleoecologic
and paleoenvironmental reconstructions. For example, evalua-
tion of the implications of the Zoophycos ichnofacies has been
complicated by uncertainties persisting with respect to the spe-
cific behavior (or behaviors) involved in Zoophycos itself (e.g.,
Ekdale and Lewis, 1991; Bromley, 1991; Kotake, 1994; Olivero and
Gaillard, 1996, 2007; Bromley and Hanken, 2003).
Lanés, Manceñido, and Damborenea have made a detailed
study of the uncommon, doubly spiraled trace fossil Lapispira
based on specimens from Jurassic rocks of Argentina. They
provide an in-depth description of this ichnogenus and analyze
its ethology, trophic type, paleoenvironmental distribution, and
potential producer. They proposed that the complex double
helicoidal tube suggests bacterial farming, although other modes
of behavior may have been involved also. In addition, the tiering
position of Lapispira is evaluated, concluding that it records the
work of a deep-tier crustacean, representing an elite trace fossil.
Löwemark, Lin, Wang, and Schönfeld test the gardening hy-
pothesis that has been used as an explanation of some types of
Zoophycos spreiten. The studied specimens occur in Quaternary
sediments of offshore Portugal. Their analysis of δ
that gardening plays an insignificant role in Zoophycos. Asgaard
and Bromley, on the basis of Pleistocene schizasterid echinoids
that are preserved together with their trace fossil Scolicia, can
confirm the construction of drain tubes behind echinoids of this
family. The study illustrates the importance of combining
paleoichnologic analysis with observations of modern struc-
tures, including experimental neoichnology. Rodríguez, Pazos,
and Aguirre-Urreta document an occurrence of the slender-
rayed Asteriacites lumbricalis, which is ascribed to the activity of
brittle-stars. Sedimentary facies and co-occurring trace fossils
support the notion that ophiuroids may have lived in waters of
less than fully marine salinity.
The third group of papers comprises paleoenvironmental
reconstruction using ichnologic evidence, in a whole range of
settings from the continental to the deep sea. This is the largest
group, including 11 papers, and represents one of the busiest
areas of ichnologic research. Historically, the use of trace fossils
in paleoenvironmental analysis has been the focus of ichnologic
research, particularly since the proposal of the ichnofacies model
by Dolf Seilacher in the fifties and sixties (see reviews and
examples in Frey and Pemberton, 1984; Pemberton et al., 1992;
Bromley, 1996; Buatois et al., 2002, among others). In terms of
conceptual and methodologic tools, most of the authors in this
section attempt to combine both ichnofacies and ichnofabrics in
their approach. The studies included here reveal how an inte-
grated approach combining ichnologic data with sedimento-
logic and stratigraphic data can provide an increased under-
standing of facies, stratigraphic framework, and depositional
setting. Several of these papers provide clues about how this
integrated approach aids in petroleum exploration and devel-
opment. One only needs to look at the amount of high-quality
work that is being produced in this subfield to realize that
sedimentologic and sequence stratigraphic applications of ich-
nology will continue to be the focus of substantial research.
Undoubtedly, the fact that many oil companies have adopted
ichnologic studies (mostly in core logging), as a routine tool will
encourage this applied side of the field. Additionally, the im-
portance of bioturbation in changing the physical and chemical
aspects of the substrate is of paramount importance in reservoir
characterization and represents a growing subfield in applied
ichnology (Pemberton and Gingras, 2005).
The first three papers of this section touch on nonmarine
environments. Netto describes very unusual, nonmarine, Skolithos-
dominated piperock from the Triassic of Brazil. Her study em-
phasizes the importance of substrate consolidation in continental
ichnofaunas. Dense concentrations of vertical burrows are attrib-
uted to opportunistic colonization by insects. This study high-
lights the stratigraphic significance of the colonized surfaces,
which represent breaks in sedimentation. Buatois, Uba, Mángano,
Hulka, and Heubeck have studied ichnofabrics dominated by
Taenidium in fluvial settings from the Miocene of Bolivia. Al-
though the ichnofauna does not display significant composi-
tional variations throughout the succession, ichnofabric analysis
reveals different taphonomic pathways that help to understand
depositional dynamics and environmental conditions. Intense
and deep bioturbation recorded by backfilled trace fossils occurs
in crevasse sandstone and overbank mudstone. Pazos, di Pasquo,
and Amenabar describe near-marine to nonmarine Carbonifer-
ous trace-fossil assemblages of Argentina that record the change
from glacial to nonglacial conditions. Ichnofossils are restricted
to glacial retreat and early postglacial times. Ichnologic data are
integrated with palynologic information to track paleoenviron-
mental changes.
The next paper explores the peculiarities of marginal-marine
depositional systems, specifically estuaries, bays, and deltas.
MacEachern and Gingras document brackish-water trace-fos-
sil suites in the Cretaceous Western Interior Seaway of Alberta.
The so-called “brackish-water model” resulted from the inte-
gration of information from modern environments and Meso-
zoic estuarine deposits. This model has been widely used in
characterization of valley-fill reservoirs. In their contribution,
MacEachern and Gingras refine the model and characterize
assemblages from four main settings: restricted or barrier-barred
bays, open unbarred bays, riverine estuaries, and barred wave-
dominated estuaries.
The next four papers focus on the ichnology of shallow-
marine environments. McIlroy records the ichnology of a Juras-
sic macrotidal, tide-dominated deltaic deposition system from
Argentina. His study documents a diverse ichnofauna that indi-
cates that tidal influence was not accompanied by lowered salini-
ties. Different subenvironments having specific trace-fossil asso-
ciations are characterized. D’Alessandro and Uchman have stud-
ied shallow marine Pleistocene deposits of southern Italy, carry-
ing ichnoassemblages containing Bichordites and Rosselia. They
correlate changes in trace-fossil composition with fluctuations in
environmental stability and energy. Knaust describes unexpect-
edly diverse trace fossils in shallow marine carbonate sediments
near the dawn of the Mesozoic, in the German Triassic
Muschelkalk. His study documents a wide variety of ichnofossils
in softgrounds, firmgrounds, and hardgrounds. Furthermore, he
provides a detailed description of the poorly known ichnogenus
Balanoglossites. Wetzel and Reisdorf make a taphonomic study of
an ichthyosaur skull that has been deposited in the unexpected
beak-downward position (Jurassic, Switzerland). Ichnofabrics
reveal the sedimentary sequence of events that led to this. Changes
in substrate consistency and the role of microbial degradation are
Ponce, Olivero, Martinioni, and López Cabrera report on
Paleogene turbidites from Tierra del Fuego, Argentina, which
show sustained and episodic gravity-flow deposition and related
bioturbation patterns. Historically, deep-marine ichnology has
focused on taxonomic and evolutionary aspects, but compara-
tively little has been written with respect to the application of
trace fossils in detailed facies reconstruction. This study is par-
ticularly relevant because it shows that ichnology may provide
useful information to distinguish between sustained and epi-
sodic gravity flows, a hot topic in deep-marine sedimentology.
Finally, Kakuwa and Webb discuss trace fossils of an Ordovician
pelagic deep-ocean bedded chert in southeastern Australia. The
ichnology of cherts is poorly known, and, therefore, this study
provides original information. Remarkably, the composition of
the ichnofaunas and the styles of bioturbation documented are
very similar to those of much younger deposits, including mod-
ern deep-sea sediments.
This section comprises papers on bioerosion, trace fossils in
hard substrates. Bioerosion is another active subfield within
ichnology, one that is particularly prone to gather researchers
from different backgrounds, including not only paleontologists
but also marine benthic ecologists and reef biologists. The first
International Bioerosion Workshop on the island of Bornholm,
Denmark, produced a group of six papers published as a group
in Historical Biology volume 13, as well as a group of 13 papers
following Bromley (1999). The third workshop in the series, held
at Barcelona, Spain, produced 9 papers following Martinell et al.
(2002), and the fourth workshop published a further 11 papers
following Mikulᢠ(2006). The fifth bioerosion workshop was
held at Erlangen, Germany, in October 2006, and Max Wisshak
and Leif Tapanila are preparing the publication of a proceedings
volume, “Current Developments in Bioerosion”, to be published
by Springer-Verlag, that will contain about 25 papers. Bioerosion,
therefore, very clearly illustrates the multifaceted nature of ich-
nology. In particular, bioerosion yields valuable insights into
predator–prey interactions, and it comes as no surprise that the
majority of recent bioerosion studies have focused on this topic
(see papers in Kowalewski and Kelley, 2002; Kelley et al., 2003).
Certainly, the complexities of predator–prey interactions are
expressed in this section. Additionally, careful analysis of bioero-
sion provides information on environmental controls (e.g.,
Bromley and Asgaard, 1993; de Gibert et al., 1998).
Malumián, López Cabrera, Náñez, and Olivero document
abundant bioerosional penetrations in Cretaceous to Cenozoic
benthic foraminiferal tests from offshore Argentina. This study is
one of the few performed in high latitudes. The authors detect
several trends through the time span analyzed and establish
correlations with different climatic episodes. Farinati describes
fossil burrows from firm sediment and bioerosion trace fossils in
skeletal substrates of Miocene to Pliocene age, Argentina.
Ichnologic information is placed within a sequence stratigraphic
context and used to evaluate the taphonomic history of the shells.
Kelley and Hansen evaluate latitudinal patterns in predatory
borings by naticid gastropods along the east coast of USA. The
patterns resulting from this study are more complex than pre-
dicted, which leads the authors to evaluate possible causes. This
paper provides a modern baseline for interpreting temporal
patterns in the fossil record and underscores the need to examine
multiple samples.
In the last section, ichnology moves out of the water with
seven papers that describe the work of insects and tetrapods.
Continental ichnology, once a neglected subfield, has experi-
enced an explosive development during the last decade. Inverte-
brate ichnology has covered a wide variety of topics, including
the definition of additional ichnofacies (e.g., Buatois and Mángano,
1995; Bromley, 1996; Genise et al., 2000), evaluation of evolution-
ary trends (Buatois et al., 1998; Miller and Labandeira, 2003),
application of trace fossils in continental sequence stratigraphy
(Buatois and Mángano, 2004), ichnofabric analysis of paleosols
(Genise et al., 2004), and documentation and analysis of insect
trace fossils (Genise, 2004), among others. Vertebrate ichnology
was also the focus of significant research leading to the documen-
tation of a widespread database. See, for example, the papers in
Gillette and Lockley (1989), the three-part memorial volume in
honor of W.A.S. Sarjeant (Pemberton et al., 2003–2004) and the
Ichnos Special Issue on Terrestrial Tetrapod Ichnofacies and
Ichnotaxonomy (Conti et al., 2007). In recent years, attempts have
been made to define vertebrate ichnofacies (Lockley et al., 1994;
Hunt and Lucas, 2007). One of the present-day challenges in
continental ichnology is to integrate vertebrate and invertebrate
datasets that have evolved independently and remain essentially
separated (Lockley, 2007).
Voigt discusses tunnel-and-chamber burrows from Carbon-
iferous–Permian alluvial-plain deposits at several localities from
western USA to eastern Europe, as evidence for fossorial behav-
ior of insects in the late Paleozoic. The tracemaker may have
occupied terrestrial muddy to sandy sediments of levee and
proximal overbank subenvironments. Yelinek and Chin show a
relationship between burrows probably made by dung beetles
and large beaver burrows, Daemonelix, in the Miocene of Ne-
braska, USA. They conclude that it was undoubtedly the abun-
dant dung in Paleocastor colonies that attracted the beetles. Milàn,
Bromley, Titschack, and Theodorou describe a diverse mamma-
lian ichnofauna from a Quaternary eolian oolite on the island of
Rhodes, Greece. Tracks attributable to elephants, camels, and
smaller artiodactyls are described, chiefly in vertical section, in
association with rhizoliths and body fossils of land snails.
Rodríguez-Tovar documents the nesting behavior of an extant
burrowing wasp, Bembix oculata. The influence of substrate firm-
ness is underscored, and experiments were performed to reveal
changes in shear strength related to variations in water content in
the substrate. Kulkarni and Borkar detail the architecture of the
arborial nests of extant Crematogaster ants in India. They compare
nests formed in mangrove thickets and deciduous forests, stress-
ing the importance of rainfall and high-speed winds as control-
ling factors. In the Jurassic Morrison Formation of Utah, USA,
Chin and Bishop find bone fragments, occurring within theropod
coprolites, to contain the borings of beetles: these are thus com-
pound trace fossils, the bone substrate having been utilized twice.
Their study demonstrates that by the Jurassic some invertebrates
have developed the ability to exploit dinosaur bone. Last but not
least, Scott, Renaut, Owen, and Sarjeant have made a detailed
study of trace formation and taphonomy of invertebrate burrows
and tetrapod tracks in marginal sediments of saline and alkaline
Lake Bogoria in the Rift Valley of Kenya. Lake-margin environ-
ments, including hot springs and ephemeral streams, provide
favorable areas for the activities of insects, mammals, birds, and
reptiles. The authors address preservational aspects in detail,
discussing the role of efflorescent salt crystallization, substrate
wetting and drying, and the presence of benthic microbial mats
and biofilms.
BROMLEY, R.G., 1991, Zoophycos: strip mine, refuse damp, cache or sewage
farm?: Lethaia, v. 24, p. 460–462.
ROMLEY, R.G., 1996, Trace Fossils: Biology, Taphonomy and Applica-
tions: London, Chapman & Hall, 361 p.
ROMLEY, R.G., 1999, Bioerosion: a collection of workshop papers: Geo-
logical Society of Denmark, Bulletin, v. 45, p. 129.
BROMLEY, R.G., AND A SGAARD, U., 1993, Endolithic community replacement
on a Pliocene rocky coast: Ichnos, v. 2, p. 93–116.
BROMLEY, R.G., AND HANKEN, N.-M., 2003, Structure and function of large,
lobed Zoophycos, Pliocene of Rhodes, Greece: Palaeogeography,
Palaeoclimatology, Palaeoecology, v. 192, p. 79–100.
BUATOIS, L.A., AND MÁNGANO, M.G., 1995, The paleoenvironmental and
paleoecological significance of the lacustrine Mermia ichnofacies: An
archetypical subaqueous nonmarine trace fossil assemblage: Ichnos,
v. 4, p. 151–161.
BUATOIS, L.A., AND MÁNGANO, M.G., EDS., 2003, Icnología: hacia una
convergencia entre geología y biología: Asociación Argentina de
Paleontología, Publicación 9, 184 p.
BUATOIS, L.A., AND MÁNGANO, M.G., 2004, Ichnology of fluvio-lacustrine
environments: Animal–substrate interactions in freshwater ecosys-
tems, in McIlroy, D., ed., The Application of Ichnology to
Palaeoenvironmental and Stratigraphic Analysis: Geological Society
of London, Special Publication 228, p. 311–333.
ichnologic record of the continental invertebrate invasion: Evolution-
ary trends in environmental expansion, ecospace utilization, and
behavioral complexity: Palaios, v. 13, p. 217–240.
BUATOIS, L.A., MÁNGANO, M.G., AND ACEÑOLAZA, F.G., 2002, Trazas Fósiles:
Señales de Comportamiento en el Registro estratigráfico: Museo
Paleontológico Egidio Feruglio, Trelew, 382 p.
CONTI, S., LOCKLEY, M.G., LUCAS, S.G., AND MEYER, C., EDS., 2007, Towards a
renaissance in tetrapod ichnology: Ichnos, v. 14, p. 1–153.
DE GIBERT, J.M., MARTINELL, J., AND DOMÈNECH, R., 1998, Entobia ichnofacies
in fossil rocky shores, Lower Pliocene, northwestern Mediterranean:
Palaios, v. 13, p. 476–487.
EKDALE, A.A., AND LEWIS, D.W., 1991, The New Zealand Zoophycos revis-
ited: Morphology, ethology, and paleoecology: Ichnos, v. 1, p. 183–
FREY, R.W., AND PEMBERTON, S.G., 1984, Trace fossil facies models, in
Walker, R.G., ed., Facies Models, Second Edition: Geoscience Canada,
Reprint Series, v. 1, p. 189–207.
GENISE, J.F., 2004, Ichnotaxonomy and ichnostratigraphy of chambered
trace fossils in palaeosols attributed to coleopterans, ants and ter-
mites, in McIlroy, D., ed., The Application of Ichnology to
Palaeoenvironmental and Stratigraphic Analysis: Geological Society
of London, Special Publication 228, p. 419–453.
Insect trace fossil associations in paleosols: The Coprinisphaera ichno-
facies: Palaios, v. 15, p. 33–48.
GENISE, J.F., BELLOSI, E.S., AND GONZALEZ, M.G., 2004, An approach to the
description and interpretation of ichnofabrics in palaeosols, in McIlroy,
D., ed., The Application of Ichnology to Palaeoenvironmental and
Stratigraphic Analysis: Geological Society of London, Special Publi-
cation 228, p. 355–382.
GILLETTE, D.D., AND L OCKLEY, M.G., 1989, Dinosaur Tracks and Traces: New
York, Cambridge University Press, 454 p.
HASIOTIS, S.T., 2002, Continental Trace Fossils: SEPM, Short Course Notes
51, 132 p.
HUNT, A.P., AND LUCAS, S., 2007, Tetrapod ichnofacies: a new paradigm:
Ichnos, v. 14, p. 59–68.
KELLEY, P.H., KOWALEWSKI, M., AND HANSEN, T.A., EDS., 2003, Predator–Prey
Interactions in the Fossil Record: Topics in Geology series, New York,
Plenum Press/Kluwer, 464 p.
KOTAKE, N., 1994, Population paleoecology of the Zoophycos-producing
animal: Palaios, v. 9, p. 84–91.
KOWALEWSKI, M., AND KELLEY, P.H., 2002, The fossil world of predation: The
Paleontological Society, Papers, v. 8, 398 p.
OCKLEY, M., 2007, A tale of two ichnologies: The different goals and
potentials of invertebrate and vertebrate (tetrapod) ichnotaxonomy
and how they relate to ichnofacies analysis: Ichnos, v. 14, p. 39–57.
OCKLEY, M.G., HUNT, A.P., AND MEYER, C.A., 1994, Vertebrate tracks and
the ichnofacies concept: Implications for palaeoecology and
palichnostratigraphy, in Donovan, S.K., ed., The Palaeobiology of
Trace Fossils: Chichester, John Wiley & Sons, p. 241–268.
MARTINELL, J., DOMÈNECH, R., AND DE GIBERT, J.M., 2002, Bioerosion, recent
and ancient: 3
International Bioerosion Workshop, Barcelona 2000.
Summary of Activities: Acta Geologica Hispanica, v. 37, p. 3–7.
MCILROY, D., ED., 2004, The Application of Ichnology to Palaeoenviron-
mental and Stratigraphical analysis: Geological Society of London,
Special Publication 228, 490 p.
MIKULÁ¢, R., 2006, Bioerosion issue of Ichnos; collection of papers from the
International Bioerosion Workshop (Prague, August 30–Septem-
ber 3, 2004): Ichnos, v. 13, p. 97.
MILLER, M.F., AND LABANDEIRA, C.C., 2003, Slow crawl across the salinity
divide: delayed colonization of freshwater ecosystems by inverte-
brates: GSA Today, v. 12, no. 12, p. 4–10.
MILLER, W., ED., 2007, Trace Fossils; Concepts, Problems, Prospects: Am-
sterdam, Elsevier, 611 p.
OLIVERO, D., AND GAILLARD, C., 1996, Paleoecology of Jurassic Zoophycos
from south-eastern France: Ichnos, v. 4, p. 249–260.
OLIVERO, D., AND GAILLARD, C., 2007, A constructional model for Zoophycos,
in Miller, W., ed., Trace Fossils: Concepts, Problems, Prospects: Am-
sterdam, Elsevier, p. 466–477.
PEMBERTON, S.G., AND GINGRAS, M.K., 2005. Classification and characteriza-
tions of biogenically enhanced permeability: American Association
of Petroleum Geologists, Bulletin, v. 89, p. 1493–1517.
PEMBERTON, S.G., MACEACHERN, J.A., AND FREY, R.W., 1992, Trace fossils
facies models: environmental and allostratigraphic significance, in
Walker, R.G., and James, N.P., eds., Facies Models and Sea Level
Changes: Geological Society of Canada, p. 47–72.
ROBBINS, D., AND SINCLAIR, I.K., 2001, Ichnology and sedimentology of
shallow to marginal marine systems, Ben Nevis and Avalon Reser-
voirs, Jeanne d’Arc Basin: Geological Association of Canada, Short
Course Notes, no. 15, 343 p.
PEMBERTON, S.G., MCCREA, R.T., AND LOCKLEY, M.G., 2003–2004, William
Anthony Swithin Sarjeant (1935–2002): A celebration of his life and
ichnological contributions: Ichnos, v. 10 (2–4)–11 (1–4).
SEILACHER, A., 2007, Trace Fossil Analysis: Berlin, Springer, 226 p.
WEBBY, B.D., MÁNGANO, M.G., AND BUATOIS, L.A., EDS., 2004, Trace Fossils in
Evolutionary Palaeoecology: Fossils and Strata, v. 51, 153 p.
... Ichnofacies and ichofabrics defined in terms of ichnology, diversity, bioturbation level, and colonization order, help in sedimentological interpretation and hydrocarbon explorations (Pemberton et al., 2001;Mcllroy, 2004;Mcllroy et al., 2005;Taylor et al., 2003;Seilacher, 2007;Miller, 2007;Bromley et al., 2007;MacEachern et al., 2007a,b;Avanzini and Petti, 2008;Mcllroy, 2008;Hasiotis, 2010;Buatois and Mángano, 2011;Knaust and Bromley, 2012;Ekdale et al., 2012). The study of bioturbated sediments is complex due to the difference in shapes, sizes and three-dimensional geometries of the burrows (Knaust and Bromley, 2012). ...
... Their usefulness has extended to applied fields, for instance in the geological interpretation of subsurface data by the oil and gas industry (Knaust, 2017 and references therein) and in aquifer characterization (e.g., Cunningham et al., 2012). All these advances have led to the publication of numerous monographs dedicated to ichnology applied to different disciplines (e.g., Pemberton et al., 2001;Hasiotis, 2002;McIlroy, 2004;Bromley et al., 2007;MacEachern et al., 2007;Miller, 2007;Seilacher, 2007;Gerard and Bromley, 2008;Wisshak and Tapanilla, 2008;Buatois and Mángano, 2011;Knaust and Bromley, 2012;Buatois, 2016a, 2016b;Genise, 2017;Knaust, 2017 for some examples from this millennium) and it is becoming more widely recognized by the scientific community. ...
Ichnological studies have become popular during the last decades, particularly those associated with the development of two major concepts —the ichnofacies model and ichnofabric approach. They have driven ichnology into diverse fields of Earth Sciences, including paleoecology, sedimentology, paleoceanography and basin analysis, as well as applied fields for the oil and gas industry and aquifer characterization. Whereas early ichnological analyses focused on outcrops, later the number of ichnological studies on well cores increased noticeably. Still, ichnological research on cores is hampered by certain limitations (i.e., mainly narrow exposed surface), and the characterization of ichnological properties is complicated when cores are involved. To facilitate ichnological analysis in cores from modern deposits, several techniques (among them, X-rays, magnetic resonance and computed tomography) have been used. With the development of computer software, a new high resolution image treatment has emerged as a powerful tool in different branches of ichnological studies, especially for cores from modern marine deposits. Because applications are numerous and perhaps not familiar to all the scientific community, this paper provides an overview of the usefulness of image treatment in ichnological analysis, its first steps and subsequent development, the novel techniques most recently used in the study of cores from modern marine deposits, and some challenges for future research.
... The study of trace fossils, ichnology, has been proven a valuable method in the reconstruction of paleoenvironments (e.g. Curran, 1985;Pemberton, 1992;Bromley, 1996;Pemberton et al., 2001;McIlroy, 2004;Miller, 2007;Bromley et al., 2007;. The early concept of ichnofacies zonation has been continuously refined and the constituent ichnocoenoses were widely used in distinguishing sub-environments (McIlroy, 2008). ...
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In the Upper Permian to Lower Triassic Khuff Formation in the Arabian Gulf, a vast shallow-marine carbonate platform developed broad fades belts with little significant changes in the lithofacies. However, trace fossil assemblages and ichnofabrics, in combination with sedimentological observations, serve in subdividing this platform and in distinguishing sub-environments. From proximal to distal, these are sabkha and salina, tidal flat, restricted lagoon, open lagoon, platform margin, shoreface/inner ramp, slope/outer ramp and basin/deeper intra-shelf. In this way, changes in relative sea level can be better reconstructed and guide the sequence stratigraphic interpretation. Meter- scale shallowing-upward cycles dominate the succession and, in addition to conventional methods, bioturbation, trace fossil assemblages and tiering patterns aid in interpreting subtidal, lower and upper intertidal and supratidal portions of these peritidal cycles. Bioturbation (and cryptobioturbation) have an impact on the primary reservoir quality before diagenetic processes overprint the deposits. For instance, deposit-feeders (such as vermiform organisms) introduce a certain amount of mud and decrease porosity and permeability considerably, whereas others like the Zoop/zycos-producers fill their dwellings with ooid grains and turn a mudstone from a barrier to a flow unit. This novel study demonstrates the value of ichnological information in carbonate reservoir characterization and the significance of trace fossil analysis in fades interpretation, reservoir zonation and the impact of bioturbation on the reservoir quality.
The Faïdja Formation displays mixed siliciclastic-carbonate deposits distributed in three members, which are organised from bottom to top as follows: the Clayey Sandstone Faïdja Member; the Clayey Limestone Bel Aoura Member and the Sandy Claystone Douaouda Member. Sedimentological data indicate the evolution of a subsiding marine environment from the shelf edge to the lower shoreface, sporadically dominated by storms. New age based on recently collected ammonite fauna allowed us to update the age of the Faïdja Formation as Early-Late Kimmeridgian. Ichnological analysis reveals the occurrence of abundant and diverse invertebrate trace fossils, grouped in four ichnoassociations. Ichnoassociation -A- coincides with the lowermost part of the Clayey Sandstone Faïdja Member; it is distinguished by a combination of pre-graphoglyptids and post-depositional traces, ascribed to the distal Cruziana ichnofacies transitional to the Nereites ichnofacies, which indicates a lower offshore zone to shelf edge environment characterised by a low-oxygenated period (anoxic event), frequent storm events and nutrient frequency. Ichnoassociation -B- occupies the middle and upper parts of the Clayey Sandstone Faïdja Member, it is characterised by an abundance of post-depositional traces, corresponding to the archetypal Cruziana ichnofacies that indicates a lower to transitional offshore environment with short phases of low hydrodynamic energy. Ichnoassociation -C- corresponds to the Clayey Limestone Bel Aoura Member that coincides with a drastic facial change and depositional conditions; this association documents a proximal Cruziana ichnofacies, and indicates an upper to transitional offshore zone as a depositional environment. Finally, ichnoassociation -D- coincides with the Sandy Claystone Douaouda Member; it is ascribed to the proximal Cruziana ichnofacies, indicating a lower shoreface environment. The combined sedimentological and ichnological data indicates a relatively subsiding platform with shallowing upwards corresponding to distal, archetypal and proximal Cruziana ichnofacies, respectively.
This chapter focuses on the major types of diversity documented in the insect fossil record. Diversity in the insect fossil record typically is provided either at the locality level, with an enumeration of all resident species (tax‐onomic diversity), or more generally has been counts at the genus or, more commonly, the family level, expressed globally through time (taxic diversity). A wide range of techniques, morphological attributes, and modes of presentation document the morphological disparity of fossil (and modern) insects. The trend of insect diversity during the Paleogene and Neogene periods is best described as a sustained increase to the present day, with a capture rate of 63% of the approximately 980 modern insect families a rate that has not changed appreciably with more recent analyses. The concluding discussion incorporates an understanding of why fossil insect diversity is relevant for the current condition of terrestrial and freshwater ecosystems of the planet.
The Turonian-Coniacian Smoky Hollow Member of the Straight Cliffs Formation in the Kaiparowits basin of southern Utah records a stratigraphic transition from isolated fluvial channel bodies to increasingly amalgamated channel belts capped by the Calico bed, a sheet-like sand-gravel unit. Characteristics of the Smoky Hollow Member are consistent with a prograding distributive fluvial system including: up-section increases in average grain size, bed thickness, and channel-body amalgamation, a fan-shaped planform morphology, and a downstream increase in channel sinuosity. The system prograded to the northeast based on thickness and facies patterns, and paleocurrent indicators. This basin-axial sediment-dispersal trend, which was approximately parallel to the fold-thrust belt at this latitude, is supported by provenance data including detrital zircons and modal sandstone compositions indicating sediment derivation mainly from the Mogollon Highlands and Cordilleran magmatic arc to the southwest, with episodic input from the more proximal Sevier fold-thrust belt to the west. Progradation occurred during a eustatic still-stand, relatively stable climatic conditions, and continuous tectonic subsidence, thus suggesting increased extrabasinal sediment supply as a primary control on basin-fill. Progradation of the Smoky Hollow Member fluvial system culminated in a ~2–3 My hiatus at the top of the lower Calico bed. Correlation with the Notom delta of the Ferron Sandstone, 80 km northeast in the Henry basin, is proposed on the basis of facies relationships and geochronology. The Calico bed unconformity is linked to regional tectonically-driven tilting and erosion observed in both basins. This article is protected by copyright. All rights reserved.
Bioturbation of sediments is one of the most important sources of paleoecological information from the stratigraphic and fossil record. Organisms utilize a variety of behaviors when they interact with the environment. The trace fossil record of the different types of behaviors can commonly be found in marine and terrestrial sedimentary rocks and provides a vast source of information on how animals lived and interacted with each other and the physical and chemical environments surrounding them. This chapter shows the various processes which trace fossils and other biogenic sedimentary structures in marine pelagic/hemipelagic muds pass through as they become a permanent part of the lithosphere. Several schemes have been proposed to assess the amount of bioturbation recorded in sedimentary rock as reflected by ichnofabric. Bioturbation has been an active process in terrestrial environments since animals colonized land by the Ordovician and various other nonmarine settings later in the Paleozoic.
Phoebichnus trochoides is a large, radiating trace fossil most commonly found in shallow marine siliciclastic deposits. The structure consists of a central boss from which extend numerous, lined, radiating burrows which have an active fill. Serial grinding and modelling techniques allow the full three-dimensional morphology of Phoebichnus trochoides to be constructed for the first time. Three-dimensional models of the trace fossil demonstrate that the central zone is composed of stacked disc-shaped layers. The structure is inferred to result from collapse of sediment below a surficial cone created by the trace-maker from excavated sediment produced during burrowing. The fill of the radial burrows is herein determined to be composed of angle of repose laminae that are inclined towards the central zone rather than the meniscate backfill documented in the ichnogeneric diagnosis and all subsequent descriptions. The structure of the fill resulted from the trace-making organism filling its burrows from a dwelling position close to the central boss, probably with material excavated from other parts of the burrow system. This study also reports for the first time subtle conical structures above the radial galleries that are inferred to result from collapse cone feeding. The new fully three-dimensional data set created of the burrow and the near-burrow environment allows for a new palaeobiological understanding of the burrow, which suggests that a crustacean trace-maker is most likely.
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Ichnologic and sedimentologic studies of the Lajas Formation (Middle Jurassic) in Sierra de la Vaca Muerta allowed the recognition of two different types of deltaic mouth bars, each of them showing trace fossil suites with different characteristics. Type I deltaic mouth bars consist of fine to coarse sandstones and fine conglomerates completely reworked by fair-weather and storm wave action, revealing a predominance of basinal hydraulic processes (e.g., waves) during bar construction and progradation. Trace fossil assemblages are composed of Ophiomorpha and Haentzschelinia in the foreset beds, and Polykladichnus, Skolithos, and Arenicolites in the topset beds. Type II deltaic mouth bars comprise sandstones that are fine to coarse and massive or present high angle cross-stratification and current ripples migrating in the opposite direction to the inclination of the foresets. These bars are interpreted to have been deposited during intervals of extraordinary fluvial discharge when wave action was restricted to the topset part of the bars. Whereas equilibrium trace fossils occur in the bottomset beds, escape trace fossils and Ophiomorpha are recorded in the distal foreset beds. In the topset beds, Skolithos and Polykladichnus specimens are very abundant. In general, the two types of mouth bars show low diversity, low abundance of trace fossils and a simple tiering structure. Such traits reflect environmental stresses mainly produced by fluctuating hydraulic energy, salinity, sediment input and high mobility of the substrate.
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The combined study of continental trace fossils and associated sedimentary facies provides valuable evidence of colonization trends and events throughout the Phanerozoic. Colonization of continental environments was linked to the exploitation of empty or under-utilized ecospace. Although the nonmarine trace fossil record probably begins during the Late Ordovician, significant invasion of nonmarine biotopes began close to the Silurian-Devonian transition with the establishment of a mobile arthropod epifauna (Diplichnites ichnoguild) in coastal marine to alluvial plain settings. Additionally, the presence of vertical burrows in Devonian high-energy fluvial deposits reflects the establishment of a stationary, deep suspension-feeding infauna of the Skolithos ichnoguild. The earliest evidence of plant-arthropod interaction occurred close to the Silurian-Devonian boundary, but widespread and varied feeding patterns are known from the Carboniferous. During the Carboniferous, permanent subaqueous lacustrine settings were colonized by a diverse, mobile detritus-feeding epifauna of the Mermia ichnoguild, which reflects a significant palaeoenvironmental expansion of trace fossils. Paleozoic ichnologic evidence supports direct routes to the land from marginal marine environments, and migration to lakes from land settings. All nonmarine sedimentary environments were colonized by the Carboniferous, and subsequent patterns indicate an increase in ecospace utilization within already colonized depositional settings. During the Permian, back-filled traces of the Scoyenia ichnoguild record the establishment of a mobile, intermediate-depth, deposit-feeding in-fauna in alluvial and transitional alluvial-lacustrine sediment. Diversification of land plants and the establishment of ecologically diverse plant communities through time provided new niches to be exploited by arthropods. Nevertheless, most ot the evolutionary feeding innovations took place relatively early, during the Late Palaeozoic or early Mesozoic. A stationary deep unfauna, the Camborygma ichnoguild, was developed in Triassic transitional alluvial-lacustrinbe deposits. Terrestrial environments hosted the rise of complex social behavioral patterns, as suggested by the probable presence of hymenopteran and isopteran nests in Triassic paleosols. An increase in diversity of trace fossils is detected in Triassic-Jurassic eolian deposits, where the ichnofauna displays more varied behavioral patterns than their Paleozoic counterparts. Also, a mobile, intermediate-depth, deposit-feeding infauna, the Vagorichnus ichnoguild, was established in deep lake environments during the Jurassic. In contrast to Paleozoic permanent subaqueous assemblages typified by surface trails, Jurassic ichnocoenoses are dominated by infaunal burrows. High density of infaunal deposit-feeding traces of the Planolites ichnoguild caused major disruption of lacustrine sedimentary fabrics during the Cretaceous. Most insect mouthpart classes, functional feeding groups, and dietary guilds were established by the end of the Cretaceous. Diversification of modern insects is recorded by the abundance and complexity of structures produced by wasps, bees, dung-beetles, and termites in Cretaceous-Tertiary paleosols. The increase in bioturbation migrated from fluvial and lake-margin settings to permanent subaqueous lacustrine environments through time.
From the Foreword: "Predator-prey interactions are among the most significant of all organism-organism interactions....It will only be by compiling and evaluating data on predator-prey relations as they are recorded in the fossil record that we can hope to tease apart their role in the tangled web of evolutionary interaction over time. This volume, compiled by a group of expert specialists on the evidence of predator-prey interactions in the fossil record, is a pioneering effort to collate the information now accumulating in this important field. It will be a standard reference on which future study of one of the central dynamics of ecology as seen in the fossil record will be built." (Richard K. Bambach, Professor Emeritus, Virginia Tech, Associate of the Botanical Museum, Harvard University)
The First International Bioerosion Workshop was held, under the auspices of Geological Institute of the University of Copenhagen, Denmark, at Limensgade Windmill, Aakirkeby, Bornholm in September 1996....
Predation and competition are the primary ecological processes that control the structure and function of communities. Predation affects the distribution and abundance of organisms, the flow of energy through systems, and the diversity and structure of communities. At the level of individuals, predator-prey interactions provide a major arena in which natural selection takes place.
Trace fossils record the behavior of animals at the very spot where they lived millions of years ago. Their growing interest derives from the intimate connection between ichnology and sedimentology and their combined relevance for paleoenvironmental reconstructions, basin analysis, and petroleum exploration. This definitive textbook by a renowned field observer and analyst of trace fossils concentrates on the most distinctive examples, mostly made by infaunal invertebrates in originally soft sediments. It covers the whole geologic column and ranges from deep-sea to shallow-marine and continental environments. Seilacher's Trace Fossil Analysis is designed to foster interpretative skills using the author's own iconic drawings. They are thematically grouped in 75 plates that form the core for the descriptive text and annotated references. A glossary of ichnological terms is also provided.
Well-preserved specimens of the trace fossil Zoophycos are found abundantly in upper Pliocene middle bathyal deposits (Shiramazu Formation) of the southernmost Boso Peninsula, Japan. The Shiramazu Zoophycos represents the product of an inverted conveyor activity of the tracemaker throughout its growth. The census fossil assemblage of Zoophycos contianing tuffaceous pellets is analyzed and indicates a high population density, patchy spatial distribution pattern, and a markedly biased age structure consisting only of adult individuals. Factors considered important in controlling the spatiotemporal pattern and age structure of the Zoophycos animals include enriched organic matter for food and unstable chemicophysical bottom-conditions, and episodic recruitment. The mud bottom was colonized only by individuals derived from an episodic recruitment during the interturbidite period and these subsequently formed patchy aggregations at the nutrient rich seafloor. -from Author
The study of fossilized dinosaur remains, vertebrate paleontology is a well established discipline, but the discovery and rediscovery of numerous and varied dinosaur footprints and nest sites has spurred a renaissance in the associated field of ichnological research. Dinosaur Tracks and Traces is the first book ever to be devoted to this subject, and it represents the work of seventy noted dinosaur ichnologists. Contributors address the history of science and the relevance of dinosaur ichnology to the interpretation of dinosaur behaviour, paleoecology, paleoenvironments, and evolution. Several new preservation, conservation, and documentation techniques are also presented. The book is richly illustrated and is intended for students and professionals in the areas of paleontology, vertebrate zoology, geology, and paleoenvironmental analysis. The historical aspects of the book and the many site descriptions also make Dinosaur Tracks and Traces appealing to amateur fossil collectors and dinosaur enthusiasts.