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1
ICHNOLOGY: PRESENT TRENDS AND SOME FUTURE DIRECTIONS
Introduction
RICHARD G. BROMLEY, LUIS A. BUATOIS, GABRIELA MÁNGANO, JORGE F. GENISE, AND RICARDO N. MELCHOR
2
3
ICHNOLOGY: PRESENT TRENDS AND SOME FUTURE DIRECTIONS
ICHNOLOGY: PRESENT TRENDS AND SOME FUTURE DIRECTIONS
RICHARD G. BROMLEY
Department of Geography and Geology, University of Copenhagen, Øster Voldgade 10, 1350 Copenhagen, Denmark
e-mail: rullard@geol.ku.dk
LUIS A. BUATOIS AND GABRIELA MÁNGANO
Department of Geologic Sciences, University of Saskatchewan, 114 Science Place,
Saskatoon, S7N 5E2, Canada
JORGE F. GENISE
Museo Paleontológico Egidio Feruglio, Av. Fontana 140, 9100 Trelew, Chubut, Argentina
AND
RICARDO N. MELCHOR
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.
INTRODUCTION AND VOLUME LAYOUT
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.
CONCEPTS AND REVIEWS
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
boundary.
ETHOLOGY AND ECOLOGY:
LINKING TRACE FOSSILS AND BEHAVIOR
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-
RICHARD G. BROMLEY, LUIS A. BUATOIS, GABRIELA MÁNGANO, JORGE F. GENISE, AND RICARDO N. MELCHOR
4
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
13
C
org
suggests
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.
ICHNOLOGY AND PALEOENVIRONMENTAL
RECONSTRUCTIONS: FROM THE
CONTINENT TO THE DEEP SEA
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
5
ICHNOLOGY: PRESENT TRENDS AND SOME FUTURE DIRECTIONS
in substrate consistency and the role of microbial degradation are
emphasized.
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.
THE SIGNIFICANCE OF BIOEROSION: EVALUATING
PREDATION AND ENVIRONMENTAL CONTROLS
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.
ICHNOLOGY MOVES OUT OF THE WATER:
TRACES OF INSECTS AND VERTEBRATES
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.
RICHARD G. BROMLEY, LUIS A. BUATOIS, GABRIELA MÁNGANO, JORGE F. GENISE, AND RICARDO N. MELCHOR
6
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