Sie sind auf Seite 1von 497

The Application of Ichnology to Palaeoenvironmental and

Stratigraphic Analysis
Geological Society Special Publications
Society Book Editors
R. J. PANKHURST (CHIEF EDITOR)
P. DOYLE
F. J. GREGORY
J. S. GRIFFITHS
A. J. HARTLEY
R. E. HOLDSWORTH
J. A. HOWE
P. T. LEAT
A. C. MORTON
N. S. ROBINS
J. P. TURNER

Special Publication reviewing procedures

The Society makes every effort to ensure that the scientific and production quality of its books matches that of its
journals. Since 1997, all book proposals have been refereed by specialist reviewers as well as by the Society's Books
Editorial Committee. If the referees identify weaknesses in the proposal, these must be addressed before the
proposal is accepted.
Once the book is accepted, the Society has a team of Book Editors (listed above) who ensure that the volume
editors follow strict guidelines on refereeing and quality control. We insist that individual papers can only be
accepted after satisfactory review by two independent referees. The questions on the review forms are similar to
those for Journal of the Geological Society. The referees' forms and comments must be available to the Society's
Book Editors on request.
Although many of the books result from meetings, the editors are expected to commission papers that were not
presented at the meeting to ensure that the book provides a balanced coverage of the subject. Being accepted for
presentation at the meeting does not guarantee inclusion in the book.
Geological Society Special Publications are included in the ISI Index of Scientific Book Contents, but they do
not have an impact factor, the latter being applicable only to journals.
More information about submitting a proposal and producing a Special Publication can be found on the
Society's web site: www.geolsoc.org.uk.

It is recommended that reference to all or part of this book should be made in one of the
following ways:
MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Strati-
graphic Analysis. Geological Society, London, Special Publications, 228.
MANGANO, M. G. & BUATOIS, L. A. 2004. Ichnology of Carboniferous tide-influenced
environments and tidal flat variability in the North American Midcontinent. In: MC!LROY,
D. (ed.) The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 157-178.
GEOLOGICAL SOCIETY SPECIAL PUBLICATION NO. 228

The Application of Ichnology to Palaeoenvironmental


and Stratigraphic Analysis

EDITED BY

D. McILROY
Sedimentology & Internet Solutions Ltd, UK

2004

Published by
The Geological Society
London
THE GEOLOGICAL SOCIETY
The Geological Society of London (GSL) was founded in 1807. It is the oldest national geological society in the
world and the largest in Europe. It was incorporated under Royal Charter in 1825 and is Registered Charity
210161.
The Society is the UK national learned and professional society for geology with a worldwide Fellowship (FGS)
of 9000. The Society has the power to confer Chartered status on suitably qualified Fellows, and about 2000 of the
Fellowship carry the title (CGeol). Chartered Geologists may also obtain the equivalent European title, European
Geologist (EurGeol). One fifth of the Society's fellowship resides outside the UK. To find out more about the
Society, log on to www.geolsoc.org.uk.
The Geological Society Publishing House (Bath, UK) produces the Society's international journals and books,
and acts as European distributor for selected publications of the American Association of Petroleum Geologists
(AAPG), the American Geological Institute (AGI), the Indonesian Petroleum Association (IPA), the Geological
Society of America (GSA), the Society for Sedimentary Geology (SEPM) and the Geologists' Association (GA).
Joint marketing agreements ensure that GSL Fellows may purchase these societies' publications at a discount. The
Society's online bookshop (accessible from www.geolsoc.org.uk) offers secure book purchasing with your credit or
debit card.
To find out about joining the Society and benefiting from substantial discounts on publications of GSL and
other societies worldwide, consult www.geolsoc.org.uk, or contact the Fellowship Department at: The Geological
Society, Burlington House, Piccadilly, London W1J OBG: Tel. + 44 (0)20 7434 9944; Fax +44 (0)20 7439 8975;
E-mail: enquiries@geolsoc.org.uk.

For information about the Society's meetings, consult Events on www.geolsoc.org.uk. To find out more about the
Society's Corporate Affiliates Scheme, write to enquiries@geolsoc.org.uk.

Published by The Geological Society from: Typeset by J W Arrowsmith Ltd, Bristol, UK


The Geological Society Publishing House Printed by Cromwell Press, Trowbridge, UK
Unit 7, Brassmill Enterprise Centre
Brassmill Lane Distributors
Bath BA1 3JN, UK USA
AAPG Bookstore
(Orders: Tel. +44(0)1225445046 PO Box 979
Fax +44(0)1225442836) Tulsa
Online bookshop: http://bookshop.geolsoc.org.uk OK 74101-0979
USA
Orders: Tel. +1918 584-2555
The publishers make no representation, express or Fax +1 918560-2652
implied, with regard to the accuracy of the information E-mail bookstore@aapg.org
contained in this book and cannot accept any legal
responsibility for any errors or omissions that may be India
made. Affiliated East-West Press PVT Ltd
G-l/16 Ansari Road, Daryaganj,
© The Geological Society of London 2004. All rights New Delhi 110002
reserved. No reproduction, copy or transmission of India
this publication may be made without written permis- Orders: Tel. +91 112 327-9113
sion. No paragraph of this publication may be repro- Fax +91 112326-0538
duced, copied or transmitted save with the provisions E-mail affiliat@nda.vsnl.net.in
of the Copyright Licensing Agency, 90 Tottenham
Court Road, London W1P 9HE. Users registered Japan
with the Copyright Clearance Center, 27 Congress Kanda Book Trading Co.
Street, Salem, MA 01970, USA: the item-fee code for Cityhouse Tama 204
this publication is 0305-8719/04/S15.00. Tsurumaki 1-3-10
Tama-shi
British Library Cataloguing in Publication Data Tokyo 206-0034
A catalogue record for this book is available from the Japan
British Library. Orders: Tel. +81 (0)423 57-7650
Fax +81 (0)42357-7651
ISBN 1-86239-154-8 E-mail geokanda@ma.kcom.ne.jp
Contents

MclLROY, D. The application of ichnology to palaeoenvironmental and stratigraphic 1


analysis: introduction
MclLROY, D. Some ichnological concepts, methodologies, applications and frontiers 3
PEMBERTON, S. G., MACEACHERN, J. A. & SAUNDERS, T. Stratigraphic applications of 29
substrate-specific ichnofacies: delineating discontinuities in the rock record
GLAUB, I. Recent and sub-recent microborings from the up welling area off Mauritania 63
(West Africa) and their implications for palaeoecology
GOLDRING, R., CADEE, G. C, D'ALESSANDRO, A., DE GIBERT, J. M., JENKINS, R. & POLLARD, 77
J. E. Climatic control of trace fossil distribution in the marine realm
MANNING, P. L. A new approach to the analysis and interpretation of tracks: examples from 93
the Dinosauria
UCHMAN, A. Phanerozoic history of deep-sea trace fossils 125
MARTIN, K. D. A re-evaluation of the relationship between trace fossils and dysoxia 141
MANGANO, M. G. & BUATOIS, L. A. Ichnology of Carboniferous tide-influenced environments 157
and tidal flat variability in the North American Midcontinent
BANN, K. L., FIELDING, C. R., MACEACHERN, J. A. & TYE, S. C. Differentiation of 179
estuarine and offshore marine deposits using integrated ichnology and sedimentology:
Permian Pebbley Beach Formation, Sydney Basin, Australia
BALDWIN, C. T., STROTHER, P. K., BECK, J. H. & ROSE, E. Palaeoecology of the Bright Angel 213
Shale in the eastern Grand Canyon, Arizona, USA, incorporating sedimentological,
ichnological and palynological data
MclLROY, D. Ichnofabrics and sedimentary facies of a tide-dominated delta: Jurassic He 237
Formation of Kristin Field, Haltenbanken, offshore Mid-Norway
BANN, K. L. & FIELDING, C. R. An integrated ichnological and sedimentological comparison 273
of non-deltaic shoreface and subaqueous delta deposits in Permian reservoir units of Australia
BUATOIS, L. A. & MANGANO, M. G. Animal-substrate interactions in freshwater 311
environments: applications of ichnology in facies and sequence stratigraphic analysis of
fluvio-lacustrine successions
MELCHOR, R. N. Trace fossil distribution in lacustrine deltas: examples from the Triassic rift 335
lakes of the Ischigualasto-Villa Union Basin, Argentina
GENISE, J. F., BELLOSI, E. S. & GONZALEZ, M. G. An approach to the description and 355
interpretation of ichnofabrics in palaeosols
DROSER, M. L., JENSEN, S. & GEHLING, J. G. Development of early Palaeozoic ichnofabrics: 383
evidence from shallow marine siliciclastics
TWITCHETT, R. J. & BARRAS, C. G. Trace fossils in the aftermath of mass extinction events 397
GENISE, J. F. Ichnotaxonomy and ichnostratigraphy of chambered trace fossils in palaeosols 419
attributed to coleopterans, ants and termites
BROMLEY, R. G. A stratigraphy of marine bioerosion 455
Index 481
This page intentionally left blank
The application of ichnology to palaeoenvironmental
and stratigraphic analysis: introduction

DUNCAN McILROY

Sedimentology & Internet Solutions Ltd, 29 Proctor Road, Hoylake,


Wirral CH47 4BE, UK (e-mail: dmc@duncanmcilroy.com)

Ichnology is the study of trace fossils, which pre- concept (Seilacher 1964, 1967), which was
serve the activity of animals as recorded by their widely used to determine palaeobathymetry.
tracks, trails, burrows and borings. Rather than Recent work detailed in the paper by Glaub
giving information about the taxonomic affinities extends the use of trace fossils to determine
of a given type of organism, trace fossils yield bathymetry to include microborings in shells.
information about an animal's behaviour in The use of ichnology to determine ancient non-
response to its environment. Trace fossils are marine environments has been taken on in
almost always in situ, are commonly specific to recent years largely through the work of the
a particular suite of environmental conditions, Argentine ichnology groups, as reviewed by
can be readily studied in core and may be Buatois & Mangano and furthered by the detailed
common in strata devoid of body fossils. They study of lacustrine deltas by Melchor.
are invaluable in thorough sedimentological ana- Complementary to the ichnofacies approach to
lysis and are thus of great utility to petroleum the study of trace fossils is the study of
geologists, sedimentologists and palaeontologists ichnofabrics as developed in sedimentary rocks.
alike. Among these, the most complex fabrics probably
Over the last 30 years or so, ichnology has been arise from the combination of pedogenic and bio-
a rapidly developing branch of palaeontology genie processes found in soils. A new approach to
that not only has important applications in clas- the description of such complex fabrics is pro-
sical palaeobiology (e.g. Donovan 1994; Bromley posed by Genise et «/., with illustrated examples
1996), but is also of great value in the more from their recent work. In addition, an ichno-
applied disciplines of palaeoenvironmental and fabric approach to detailed facies characterization
stratigraphical analysis. Much progress has of tidal deltaic facies is adopted by Mcllroy (b).
been made in the development of this discipline, Ancient depositional environments are
but there remain many fascinating and challen- reviewed from the 'bottom up' from deep water
ging issues, particularly in combining ichnology environments - both oxygen rich (Uchman) and
and sedimentology. This book aims to provide oxygen poor (Martin) - through shallow
a summary of recent progress, with an up-to- marine (Bann & Fielding) and marginal marine
date summary of most themes in modern ichnol- (Baldwin & Strother; Bann et aL\ Mangano &
ogy. The volume stems from the 2003 Lyell Buatois; Mcllroy (b)) to non-marine settings
Meeting sponsored by The Geological Society, (Buatois & Mangano). Insights into the forma-
The Palaeontological Association, BP, Shell, tion and preservation of fossil tracks by Manning
Exxon Mobil, Statoil, Total and Amerada Hess. stand to revolutionize the way that ichnologists
The introductory paper by Mcllroy (a) pro- interpret vertebrate tracks, and demonstrate
vides a condensed summary of some ichnological how trace fossils may be used to determine the
themes and frontiers, and outlines a practical saturation of ancient non-marine deposits.
approach for the description of trace fossils and The last theme addressed in the book is how
identification of key stratigraphic surfaces. The trace fossil assemblages have changed through
sequence stratigraphic theme is taken up by time. Evolutionary ichnofaunas are reviewed
Pemberton et «/., illustrated by their work on from the Cambrian by Droser et al. with respect
the Mesozoic of Canada, both in outcrop and to ichnofabrics and substrate changes, and in
in core. The recognition of key (sequence) strati- the aftermath of extinction events by Twitchett
graphical surfaces is addressed in part by the & Barras. Comprehensive Phanerozoic-long
detailed studies of Bann et a/., Mcllroy (b) and reviews of borers and traces of the 'denizens of
Bann & Fielding in shallow to marginal marine the deep' are provided by Bromley and Uchman
depositional systems, and by Buatois & Mangano respectively. The stratigraphic record of the radia-
in their review of non-marine systems. tion of bees, termites, ants and other progenitors
One of the earliest applications of ichnology of chambered burrows is reviewed by Genise,
with widespread use was that of the ichnofacies who also organizes them into ichnofamilies.
From: MC!LROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 1-2. 0305-8719/04/S15.00 © The Geological Society of
London.
2 D. McILROY

References SEILACHER, A. 1964. Biogenic sedimentary structures.


In: IMBRIE, J. & NEWELL, N. (eds) Approaches to
BROMLEY, R. G. 1996.Trace Fossils: Biology, Taphon-Paleoecology. Wiley, New York, 296-316.
omy and Applications. Chapman & Hall, London. SEILACHER, A. 1967. Bathymetry of trace fossils.
DONOVAN, S. K. (ed.) 1994. The Palaeobiology of Trace Marine Geology, 5, 413^28.
Fossils. Wiley, Chichester.
Some ichnological concepts, methodologies,
applications and frontiers

DUNCAN McILROY

Sedimentology & Internet Solutions Ltd, 29 Proctor Road, Hoylake,


Wirral CH47 4BB, UK (e-mail: dmc@duncanmcilroy.com)

Abstract: Ichnology straddles the boundary between palaeontology and sedimentology, and
is becoming an increasingly important tool in both fields. For the palaeontologist, trace
fossils allow insight into behaviour and biomechanics of animals that would otherwise be
the subject of conjecture. For the sedimentologist, trace fossils have a marked impact on
the interpretation of sedimentary rocks in that they destroy primary sedimentary structures,
but can also reveal subtle palaeoenvironmental information beyond the resolution attainable
by analysis of primary physical sedimentary structures. This contribution aims to review the
major developments in the field of ichnology, and to highlight some of the tools and
approaches currently used by ichnologists. A personal ethos for the study of trace fossils
in core is outlined as a model ichnological protocol, and some of the frontiers of the science
as a whole are briefly discussed.

Some landmarks in the history of The approach taken by most ichnologists in


ichnological research response to the 1964 decision of the ICZN was
to continue to apply the rules of the ICZN with-
From nomenclatural chaos to stability out the blessing of officialdom. However, in
(ICZN) anticipation of a revised version of the ICZN,
Sarjeant & Kennedy (1973) published a draft
Structures that we now recognize as trace fossils proposal for a separate ichnological code,
have been recorded and named in the literature adapted from the ICZN and its sister publica-
for centuries. In the early history of palaeontol- tion, the ICBN (International Code for Botanical
ogy, a profusion of names was created for Nomenclature). A less radical approach was
shapes preserved in rocks. In many subdisci- taken by Hantzschel & Kraus (1972) and Sar-
plines of palaeontology, resolving these early jeant (1979), who proposed specific amendments
names was a comparatively simple process. For to the pre-existing ICZN, which were eventually
ichnologists, however, sifting through the integrated into the subsequent version (Melville
plethora of names of taxa commonly misidenti- 1979; ICZN 1985) despite some fierce opposition
fied as sponges, seaweeds and plants has been a to the inclusion of non-reproducing forms (e.g.
monumental task. The individual to whom we Lemeche 1973). The ICZN (1985) therefore
owe the greatest debt is Hantzschel (1962, 1965, overturned the original ruling regarding the
1975) for his monographic works on the identification of trace-making organisms, render-
numerous synonymies of many forms that were ing post-1930 ichnotaxa valid under the code
originally poorly documented and illustrated. regardless of whether a trace-maker could be
The task of ichnotaxonomy was made even identified, and vilifying the personal decision of
more difficult by the International Zoological most ichnotaxonomists to persist with applying
Congress (to whom all taxonomists look for the rules of the ICZN despite not being bound
guidance in taxonomic procedure), who initially by them.
insisted that, to be valid, all trace fossil names In the most recent edition of the ICZN (1999)
erected after 1930 were to be accompanied by a ichnology seems to have been largely embraced
statement identifying the trace-making animal by the zoological community. Trace fossil
(ICZN 1964). The net effect of this was to genera (ichnogenera) established after 1999
render most post-1930 trace fossil names invalid. must have a designated type species (ichno-
This is because ichnologists can seldom identify species) (ICZN 1999, Article 66.1); for earlier
trace-making organisms; indeed a single trace established ichnotaxa no type species need be
may be made by many different taxa and - in designated but may be assigned at a later date
the case of compound trace fossils - a single according to the rules (ICZN 1999, Article 69).
trace may be the work of several organisms The status of ichnotaxonomy is thus now firmly
(e.g. Pickerill & Narbonne 1995; Rindsberg & established as a subdiscipline of taxonomy and
Martin 2003). - thanks to the new provisions within the

From: MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 3-27. 0305-8719/04/S15.00 © The Geological Society of
London.
4 D. McILROY

Table 1. The archetypal Seilacherian ichnofacies

Ichnofacies Predominant trace fossil types Inferred control (Seilacher)

Skolithos Vertical traces of suspension feeders Bathymetry (above fair-weather wavebase,


FWWB)
Cruziana Horizontal and vertical deposit feeders Bathymetry (between FWWB and storm
wavebase, SWB)
Zoophycos Pervasive deposit feeders Bathymetry (shelf and slope below SWB)
Nereites Shallow burrows with complex morphologies Bathymetry (basin-floor with turbidites)
showing highly programmed behaviours
Glossifungites Traces characteristically preserving scratches, Firm surfaces associated with incipient
mostly of suspension feeders submarine lithification
Scoyenia Non-marine traces Freshwater conditions (red-bed deposition)

ICZN (1999) and the sensible taxonomic prac- identification of an ichnofacies, with the types
tices of ichnologists over the last 3(MK) years - of trace fossils/feeding strategy being diagnostic.
ichnology can only grow as a rigorous science. These ichnofacies are widely used in palaeo-
Among the problems that do remain is the environmental interpretation (Table 1).
exclusion of modern traces by the International The archetypal Seilacherian ichnofacies (Table
Code of Zoological Nomenclature (ICZN 1999, 1) were based largely on assemblages of traces in
article 1.2.1), which restricts the use of ichnotaxa a particular lithofacies and related to a bathy-
to fossil and not to modern traces. This regulation metric gradient from shallow Skolithos to deep
complicates the description of modern borings Nereites ichnofacies. The Scoyenia ichnofacies
and burrows, but, as such incipient trace fossils was however created differently, being an envir-
may be incomplete, some taxonomic problems onment-led definition in contrast to the other
may thereby be avoided. The simplest means of behaviourally defined ichnofacies.
making comparisons between modern traces The recognition of these basic ichnofacies
and their ancient counterparts is to use the groupings was of great utility to sedimentologists
prefix 'aff.' to denote their affinity to a trace as an aid to palaeoenvironmental interpretation.
fossil while acknowledging that modern traces This work was immediately grasped by both the
have no validity under the current ICZN. A palaeontological and sedimentological com-
grey area also exists in defining when a modern munities, and was seminal in inspiring refined
trace becomes a trace fossil: at abandonment of sedimentary facies models and in stimulating
the burrow, at burial, or at lithification? Other further classification of associations of trace
issues, particularly the exact definition of what fossils into additional ichnofacies. The sub-
constitutes a trace fossil, are currently under dis- sequent proliferation of ichnofacies has been
cussion (see Bertling et al. 2003). Most modern reviewed recently (Bromley 1996; Pemberton
ichnologists consider rootlets and other plant et al. 2001), and the most important are listed
traces as trace fossils, though the ICZN is reluc- in Table 2. The controls on the distribution of
tant to include non-animal taxa for obvious ichnofacies have been conclusively demonstrated
reasons. It may therefore be that the simplest to be more than simply bathymetric (Ftirsich
solution to these issues is for a separate ichno- 1975; Ekdale et al 1984; Frey et al 1990;
logical code to be created and given some form Bromley & Asgaard 1991; Gierlowski-Kordesch
of approval by the ICZN and ICBN. Such issues 1991; Wetzel 1991; Fig. 1), but the ichnofacies
are as yet unresolved and remain a challenge. themselves retain their usefulness albeit in
modified form.
As can be seen from Table 2 there is no consis-
Ichnofacies approach tent ethos behind the creation of ichnofacies. The
most anomalous of these are the vertebrate
The observations of Seilacher (1964, 1967) that footprint ichnofacies and coprofacies, which are
recurrent associations of trace fossils could be more likely to be trace fossil assemblages related
recognized in the rock record represents the to local palaeoecology and palaeobiology of
first widely applicable use of ichnology. Initially, producers than ichnofacies of inter-regional
six recurrent sets of trace fossils (named ichno- applicability. In addition, Bromley's proposed
facies) were recognized by Seilacher (1967) and Fuersichnus ichnofacies has been demonstrated
named for a characteristic trace fossil. The from a variety of non-marine environments
eponymous trace fossil need not be present for (Buatois & Mangano 2004).
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 5

Table 2. History of the development of the ichnofacies concept, highlighting the inferred palaeoenvironments of the
later ichnofacies

Ichnofacies Palaeoenvironment Author(s)

Paleodictyon and The deep marine ichnofacies was subdivided into Seilacher (1974) recognizes the first
Nereites Paleodictyon for sand-rich proximal turbidites subdivision of one of the
ichnosubfacies and Nereites for mud-rich distal turbidites archetypal ichnofacies. Building
on the work of Ksiazkiewicz
(1970); Crimes (1973)
Rusophycus Fluvial/shallow lacustrine Bromley & Asgaard (1979) as an
ichnocoenosis, and as an
ichnofacies by Bromley (1996)
Fuersiehnus Originally inferred to be representative of shallow Bromley & Asgaard (1979) as an
lacustrine settings, below FWWB. Has ichnocoenosis and as an
subsequently been shown to extent to a variety ichnofacies by Bromley (1996)
of freshwater settings (Buatois & Mangano
2004)
Trypanites Lithic/hardground substrates Frey & Seilacher (1980)
Teredolites Woody (xylic) substrates Bromley et al (1984)
Redefined Scoyenia Freshwater shallow lacustrine and fluviatile See emendation of original definition
settings by Frey et al. (1984); Buatois &
Mangano (1995)
Curvolithus A subset of the Cruziana ichnofacies, found in Lockley et al. (1987) based on the
settings with high sedimentation rates. earlier Curvolithus ichnocoenose
Particularly delta and fan delta deposits of Heinberg & Birkelund (1984)
Psilonichnus Coastal dunes Frey & Pemberton (1987)
Arenicolites A subset of Cruziana ichnofacies (opportunistic Bromley & Asgaard (1991)
colonization of event beds)
Mermia Lacustrine turbidites Buatois & Mangano (1993)
Entobia Subdivision of Trypanites (boring traces) Bromley & Asgaard (1993)
Gnathichnus Subdivision of Trypanites (rasping traces of Bromley & Asgaard (1993)
organisms feeding on the surface of lithic
substrates)
Termitichnus Palaeosol ichnofacies including coprolites, Smith et al (1993), replaced by
rhizoliths and traces in xylic matter e.g. leaves Coprinisphaera of Genise et al.
(2000)
Laoporus The variety of footprint assemblages represent a Lockley et al. 1994
Brasilichnium diverse array of palaeoenvironments, though
Brontopodus their facies specificity is in doubt, and the
Caririchnium separation of Laoporus and Brasilichnium on
and stratigraphic grounds is not well founded
'Shorebird
ichnofacies'
Coprofacies Based on the distribution of various coprolite Hunt et al. (1994)
types. The facies-specificity of coprolites is in
doubt and has not been widely used to date
Coprinisphaera Palaeosols with insect nests Genise et al. (2000)
Ophiomorpha rudis Subdivision of Nereites ichnofacies proposed for Uchman (2001)
ichnosubfacies channel and lobe to lobe fringe environments
but is only recognized from Eocene and
younger strata

The similarities between some non-marine and least in part - to the predominance of anoxia
marine ichnofacies, as highlighted by Bromley resulting from thermal stratification in lakes
(1996), demonstrates parallel behavioural evolu- (Buatois & Mangano 2004). This increased
tion in the non-marine and marine realms, reliance upon interpretation of sedimentary
presumably due to comparable environmental environment before ichnological characteriza-
controls. The notable exception is the near- tion (subjective ichnofacies sensu Reading 1978)
absence of a deepwater mudstone ichnofacies in suggests that there is little utility in continuing
non-marine settings, which is probably due - at to create archetypal ichnofacies - a stance
Fig. 1. Summary diagram showing current thinking on the likely distribution of the main soft/loose and firmground ichnofacies (based on Bromley 1996) and based on
condition of still-stand of sea-level. Ar, Arenicolites ichnofacies; Cu, Curvolithos ichnofacies; Co, Coprinisphaera ichnofacies; Cr, Cruziana ichnofacies; Fu,
Fuersichnus ichnofacies; Gl, Glossifungites ichnofacies; Ne, Nereites ichnofacies; Ps, Psilonichnus ichnofacies; Ru, Rusophycus ichnofacies; Sc, Scoyenia ichnofacies;
Sk, Skolithos ichnofacies; Zo, Zoophycos ichnofacies. During marine flooding events and sea-level fall some ichnofacies become more widespread (e.g. Glossifungites
ichnofacies).
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 7

Table 3. Characteristics that make fossils good zone fossils; the ideal zone fossil would fulfil all criteria

Characteristic Rationale

Rapidly evolving, i.e. narrow stratigraphic range Improves resolution of biozone


Widespread distribution Eases interregional correlation
Good preservation potential Improves chances of occurrence in a given rock unit
Abundance Improves chances of occurrence in a given unit
Facies independence Allows correlation independent of palaeoenvironment
Easy identification Allows use by non-experts

supported by Goldring (1993, 1995) and dis- robust of these have been the schemes of Crimes
cussed further below in terms of ichnofabric (1975, 1987, 1992), which included a vast dataset
analysis and ichnocoenoses. of Neoproterozoic to Cambrian occurrences but
rely on unpublished stratigraphic inferences.
The most widely used ichnozones are those
Ichnostratigraphy based on the Neoproterozoic-Cambrian type
section in southeastern Newfoundland erected
Biostratigraphy is the methodology by which by Narbonne et al. (1987). Indeed the boundary
stratigraphers can subdivide the rock record, itself was defined at the junction between the
and correlate from region to region using the Harlaniella podolica and Phycodes pedum ichno-
record of evolution and extinction of fossil zones (Brasier et al. 1994).
taxa. The basic subdivision of stratigraphic Other radiation events. The early terrestrializa-
time is the zone, and the fossils that define tion event has the potential to yield bio-
those zones are known as zone fossils. stratigraphic data, but such sections do not
In accordance with the criteria in Table 3, the yield abundant ichnological data and there is a
best zone fossils are likely to be rapidly evolving potential problem with likely endemicity and
organisms with a pelagic portion to their lifecycle diachroneity of early non-marine faunas/ichno-
(improves distribution), and should comprise faunas. Rapid radiation is also recorded after
distinctive hard body parts. Thus trace fossils major extinction events (see Twitchett & Barras
generally make poor zone fossils - due largely 2004), such as the Permo-Triassic extinction
to the benthic lifestyle of trace-making organisms event, which is estimated to have eradicated
- except during intervals where benthic organ- 96% of all family-level diversity (Jablonski
isms that produce distinctive burrows evolve 1991). Importantly, however, no phylum is
rapidly. Convergent behavioural evolution is known to have become extinct at that particular
the norm in ichnology, which accounts for the stratigraphic level so - although there was rapid
longevity of most ichnotaxa; convergent evolu- radiation - no fundamentally new body plans
tion of burrowing organisms has also been evolved or died out, which limits the potential
demonstrated (cf. Seilacher 1994). Bio-events usefulness of ichnostratigraphy, but the stepwise
that have the potential to include trace fossils reappearance of ichnotaxa can be of strati-
of biostratigraphic significance include radiation graphic utility (Twitchett & Barras 2004).
events, and the evolution of distinct trace-
making groups. Evolution of distinct trace-making groups
Palaeozoic arthropods. One of the first direct
Radiation events applications of ichnology to petroleum geology
Neoproterozoic-Cambrian. The Cambrian Radia- was the development of an ichnostratigraphic
tion is perhaps the most dramatic of all the scheme for the correlation of peri-Gondwanan
radiation events in the stratigraphic record, shallow marine 'unfossiliferous' quartzites
being the period of time in which most anatomi- (Seilacher 1970, 1985, 1992, 1993; Fig. 2),
cal design becomes established. Many of the which are important reservoir intervals through-
taxa represented by this diversification of body out the Middle East and North Africa. The main
form were benthic in nature, and 'experimenta- trace fossils involved in this ichnostratigraphic
tion' in body form and behaviour are to be scheme are ichnospecies of Cruziana and related
expected. It is thus unsurprising that this period arthropod trace fossils. Such traces are abundant
has been identified by a number of authors as both in core and in outcrop, but body fossils are
having excellent potential for ichnostratigraphy notoriously rare. The concept of basing an ichno-
(Crimes 1975, 1987, 1992; Alpert 1977; Fedonkin stratigraphic scheme on Lower Palaeozoic
et al 1983; Narbonne et al. 1987). The most arthropod traces is thus well founded in that
8 D. McILROY

Fig. 2. The Cruziana ichnostratigraphy of Gondwana (redrawn from Seilacher 1992).

they are abundant and widely distributed, and 22c). Vertebrate footprint ichnotaxonomy is a
the trace-makers (trilobites and other arthro- particularly difficult field, and much work needs
pods) were rapidly evolving during this phase to be done to fully appreciate which characters
in their history. The drawback with the use of are useful for ichnotaxonomy.
the scheme is that the trace-making arthropods Tertiary. The radiation of terrestrial insects is
are benthic and thus prone to provincialism (cf. well reflected in the fossil record of their
Magwood & Pemberton 1988). In addition, burrow chambers, which is a taxonomic charac-
many of the characters used to define the ichno- ter widely used to identify modern insect taxa.
species upon which the scheme relies are only The radiation of insects in the Tertiary was
rarely seen in material other than the exquisitely extremely rapid, and their distribution is highly
preserved type material. The majority of material sensitive to regional climatic shifts (Genise
examined in the field can thus be difficult to 2004). The Insecta are widely dispersed owing
identify by the non-expert. to their commonly airborne adult phase, pre-
Trias sic—Jurassic. During the Permian through sence in a range of non-marine environments,
to the Jurassic footprints of the archosaurs are and their easily characterized egg chambers that
comparatively common, particularly in Europe have a high preservation potential. The Insecta
(Haubold 1984), North America (e.g. Olsen with their staggered first occurrence datums
1980) and South Africa (Ellenberger et al. thus fit the optimal characteristics of a zone
1970). The rapid evolution of the archosaur fossil (see Table 3).
faunas is reflected in their changing footprint
morphologies from early (Triassic) Cheirother-
ium-type footprints to later (Jurassic) tridactyl Seafloor and sediment oxygenation
footprints such as the ichnogenus Grallator.
Such schemes are reliant upon an abundance of One of the most fundamental controls on the dis-
well-preserved surface tracks, and have been tribution of benthic animals and their trace fossils
well calibrated by accessory biostratigraphic in aqueous environments is the availability of
data (e.g. Cornet & Traverse 1975). The difficul- dissolved oxygen. This may be present either in
ties of recognizing surface tracks make an bottom waters or in porewaters, but is essential
awareness of possible under-track artefacts an for all metazoan life. The links between
invaluable skill (Manning 2004). In particular, ichnological/benthic macrofossil distributions
features such as detached heel-like structures and bottom water oxygenation are well estab-
and 'spurs' in some ichnospecies of Brachycheiro- lished (Bromley & Ekdale 1984; Savrda & Bottjer
therium may be related to transmitted heel 1987; Ekdale & Mason 1988), though recent work
structures (see Manning 2004, figs 17c, 21b, (Schieber 2003) has demonstrated the need for
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 9

careful assessment of apparently unbioturbated these parasequences is known as high-resolution


sediments through work with image analysis. sequence stratigraphy (Howell & Aitken 1996).
Appreciation of the role of sediment anoxia in The majority of integrated ichnological/
ancient successions is immature, though the sequence stratigraphic approaches have
impact of sediment anoxia on modern shallow employed the use of trace fossils, either in the
marine taxa is well known (e.g. Pike et al. 2001 recognition of key stratigraphic surfaces (e.g.
and references therein). Likewise, Wignall Bromley & Goldring 1992; Taylor & Gawthorpe
(1993) has correctly highlighted the fact that 1993; Ghibaudo et al. 1996; Oloriz & Rodriguez-
changing substrate conditions can favour depau- Tovar 1999; MacEachern et al. 1999; Malpas
perate ichnofaunas similar to those that typify 2000; Pemberton et al. 2000; Uchman et al.
anoxic bottom-water conditions. 2000) or for improved broad-scale facies inter-
Sediments deposited in low-oxygen settings pretations based on a refined ichnofacies-based
results are generally rich in organic carbon and approach (e.g. Vossler & Pemberton 1988; Frey
thus have a high source rock potential to petro- & Howard 1990; Savrda 1991; Brett 1998;
leum systems (e.g. Oschmann 199la, b; Wignall Siggerud & Steel 1999; Pemberton et al. 2001).
1994). Determination of the palaeo-oxygenation Despite the firm establishment of the ichno-
of such sediments can be approached by geo- fabric/ichnocoenosis approach to improved
chemical means as well as through ichnology facies and stratigraphic analysis (Bockelie 1991;
and palaeoecology (Wignall & Myers 1988; Taylor & Gawthorpe 1993; Taylor & Goldring
Wilkins et al. 1996; Wignall & Newton 2001). 1993; Taylor et al. 2003; Schlirf 2003), this
The organically rich nature of these facies also methodology has been underused (but see Bock-
means that they are a potential treasure-trove elie 1991; Martin & Pollard 1996; Schlirf 2003;
of nutrients for deposit-feeding organisms Mcllroy 2004). The advantage of the ichno-
(Diego & Douglas 1999). Colonization during fabric/ichnocoenosis approach is that its focus
amelioration of low-oxygen conditions by an is improved characterization of facies - the
opportunistic fauna is a common phenomenon fundamental building block of sequence strati-
that is documented in both modern and ancient graphy - through improved understanding of
sediments (e.g. Sagemann et al. 1991; Savrda & trace fossil fabrics and seafloor ecology with
Bottjer 1991; Wignall & Pickering 1993; Bromley respect to the host sediments. Once established,
et al. 1995; Smith et al. 2000; Martin 2004; Fig. an ichnofabric scheme can be used to assess
9). Indeed, the ecology of deep-sea sites in stacking patterns, while simultaneously enhan-
general is now becoming much better known cing ichnological characterization of key strati-
(e.g. Kaufmann & Smith 1997). Recent work graphic surfaces (Mcllroy 2004).
has also highlighted the role of chemosymbiosis It is regrettable that most sedimentology text-
as a feeding strategy in such settings (Paull et al. books focus on Seilacherian ichnofacies (e.g.
1984; Hovland & Thomsen 1989) and has led to Frey & Pemberton 1984; Pemberton et al. 1992,
the reinterpretation of some trace fossils as being 2001), rather than also encompassing the more
the result of such behaviour (Seilacher 1990; Fu flexible ichnofabric/ichnocoenosis approach
1991). outlined below (the notable exception being
Goldring 1999).

Sequence stratigraphy
An ichnological ethos
Perhaps the most significant predictive strati-
graphic tool developed in recent years is that of What follows is a personal approach to the study
sequence (seismic) stratigraphy, which was of trace fossils and ichnofabric, which is designed
developed by Exxon Production Research Co. largely for the study of marine clastic deposi-
in the late 1970s (Vail et al. 1977), and has tional systems (the author's own main focus of
since been further refined and debated (see the research). The basic methodology is of wider
excellent review of Nystuen 1998). The basic application, but should be adapted to the needs
model involves the recognition of unconfor- of the particular ichnological/sedimentological
mity-bound packages of sediment (sequences), problem addressed, e.g. non-marine terrestrial
which can be related to cycles of relative sea- systems (see Genise et al. 2004) and carbonate
level change. Within these sequences higher systems (e.g. Curran 1994).
frequency increases in relative sea-level can be The reader should be aware that there are
recognized (flooding surfaces), which (envelope) many ways to approach ichnological studies,
progradational sedimentary packages known as and that no single approach is correct. All have
parasequences. The study and correlation of their strengths.
10 D. McILROY

Fig. 3. Comparison of two cross-bedded sandstones with the bivalve escape burrow aff. Lockeia (arrowed):
(a) represents a non-marine crevasse splay sandstone from the Carboniferous of Northumberland, England
(scale bar in mm); and (b) a tidally deposited sandstone Jurassic, Neuquen Basin, Argentina.
Ichnotaxonomically, the two specimens are ichnologically similar but sedimentological observations allow
recognition of a tidal depositional environment through tidal bundling.

Scale of observation no information is disregarded. In the same way


that many sedimentologists record trace fossils
As with most sedimentological studies, the ulti- as 'bioturbation', so many palaeontologists/
mate aim of an ichnological study commonly ichnologists record the host sediment as sand-
determines the resolution at which data are stone/shale etc. without due regard to the physi-
recorded. For example, when looking for long cal sedimentary structures contained therein
timescale changes in bioturbation, in a thick, (Fig. 3). The sedimentologist should look to
sedimentologically homogeneous Neoprotero- ichnology to help understand sedimentologically
zoic-Cambrian succession, Mcllroy & Logan homogeneous rocks (e.g. Gowland 1996; Martin
(1999) used decimetre-scale observations of ich- & Pollard 1996; Mcllroy 2004). Likewise, the
nofabric index (sensu Droser & Bottjer 1986, ichnologist may learn about the likely spatio-
1989, 1991). In contrast, when studying the temporal distribution of ichnofabrics by full con-
highly heterogeneous tidal deposits of a tide- sideration of likely sandstone body geometries
dominated deltaic system, the same author and stacking based on sedimentological informa-
made ichnological and sedimentological obser- tion. Although ideally we should all be trans-
vations on a centimetre scale (Mcllroy 2004). disciplinary geoscientists, the reality is that for
The key to producing scientifically valid, most geologists collaboration and open discus-
usable, data is thus to choose an appropriate sion is the way forward; this is especially so
scale at which to collect ichnological and between sedimentologists and ichnologists.
sedimentological data. The constraints are com-
monly time available (often a problem when
working to industry deadlines), volume of data Quantification of bioturbation
required/desired, and the inherent variability
of the sedimentary succession. Observations The need to quantify the extent to which animals
should of course always be made in close detail modify sedimentary fabrics has been recognized
if possible, but if nothing is found in a thick since the early days of modern ichnology.
package of homogeneous sand there is little Many attempts to produce an easy-to-use
advantage in making numerous statements scheme for the documentation of this bioturba-
describing the lack of sedimentological/ichno- tion have been proposed (e.g. Moore & Scrutton
logical features. A conspicuous lack of trace 1957; Reineck 1963; Howard & Frey 1975; Frey
fossils is in itself revealing though, and an impor- & Pemberton 1984; Droser & Bottjer 1986,
tant observation in need of explanation. 1989, 1991; Taylor & Goldring 1993). Most of
these rely upon estimating the proportion of
sedimentary fabric/laminations destroyed by
Sedimentological context the burrowing activity of animals (i.e. bioturba-
tion). These need to quantify the fabric as seen
In getting the maximum amount of information in vertical cross-section is a bias introduced by
from a sedimentary rock it is imperative that the predominance of studies of shallow box
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 11

Fig. 4. Ichnofabric indices exemplified by flashcards according to the scheme of Droser & Bottjer (1989);
redrawn with permission. Shows the proportions of sediment reworked by bioturbation as seen in vertical
cross-section.

cores (modern sediments) and sediment/rock


cores (e.g. in oil field studies).
The most usable of these schemes is the semi-
quantitative flashcards of Droser & Bottjer
(1986, 1989, 1991; Fig. 4), which have been
used with success by a number of authors (e.g.
Droser & O'Connell 1992; Mcllroy & Logan
1999). This approach has recently been extended
to include a flashcard methodology for quantifi-
cation of the extent of bioturbation on bedding
planes (Miller & Smail 1997; Fig. 5). A similar,
but more sophisticated, means of graphically
representing quantitative and semi-quantitative
aspects of trace fossil fabrics in vertical section
has been proposed by Taylor & Goldring
(1993), and is discussed in detail below (Fig. 6).

Ichnofabric analysis
The component of a sediments texture created by
the action of animals is known as its ichnofabric.
Ichnofabric may be created either by bioturba- Fig. 5. Flashcards showing the proportion of bedding
tion (in loose sediment) or by bioerosion (in lithi- planes covered by trace fossils from Miller & Smail
(1997), as classified into 'bedding plane bioturbation
fied sediment) by a diverse array of organisms indices'. Column A represents example bedding
from microbes (e.g. Glaub 2004; Fig. 7a) to planes covered by trace fossils of even size and shape
dinosaurs (e.g. Manning 2004; Fig. 7b). One of with even distribution; Column B represents example
the features of trace-making organisms is that bedding planes covered by trace fossils of different
they are commonly highly sensitive to their sizes and shapes and with uneven distributions
environment and can thus provide a record of (redrawn with permission).
12 D. McILROY

Fig. 6. Example of a modified ichnofabric constituent diagram adapted from Taylor & Goldring (1993)
expressing the ichnofabric of an outcrop from the Lajas Formation, Neuquen Basin, Argentina. Ast,
Asterosoma; Pa, cf. Parahaentzchelinia; Th, Thalassinoides. Note that the horizontal scale is used at the base of
the diagram, which is the author's personal preference.

palaeoenvironmental conditions before, during order to fully characterize their facies and under-
and after deposition of a bed (Fig. 8). When con- stand their palaeoenvironment of deposition.
sidering physical sedimentary structures alone, As discussed above, ichnofabrics are best investi-
information can be gleaned only about condi- gated on a bed-by-bed scale, which normally
tions at the time of deposition, which in many requires sedimentary logging at a scale of at
cases (e.g. hurricane-deposited sandstone beds least 1:50. The features of ichnofabric that
on the normally quiescent proximal shelf) can should be recorded during routine investigation
be anomalous. The modern sedimentologist of sedimentary rocks include:
should therefore not only be able to record the
presence of bioturbation but also be able to com- intensity of bioturbation;
bine information from sedimentary structures diversity;
and other macro/micropalaeontological data in relative abundance;

Fig. 7. Different scales of ichnofabric development: (a) artificial casts of microborings similar to Fasciculus in a
shell clast (Recent of Mauritania courtesy of I. Glaub); (b) vertical cross-section through a dinosaur track
showing bioturbation by a large bipedal dinosaur from the Jurassic Scalby Formation of Yorkshire, UK
('dinoturbation' of some authors) (courtesy of P. Manning).
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 13

Fig. 8. Cored section with Diplocraterion parallelum seen in (a) longitudinal and (b) transverse cross-sections.
The greatly different proportions of the cut surface covered by traces is dependent on the section taken. The
percentage by volume of core bioturbated is in both cases c. 40%.

ichnometry; described in the section above. The present


infaunal tiering; author recommends either the ichnofabric index
succession of bioturbation; schemes of Droser & Bottjer (1986, 1989, 1991),
colonization styles. or bed-by-bed estimation of bioturbation as a
percentage. It is emphasized, however, that the
Intensity of bioturbation parameters outlined below should also be inves-
Sedimentologists should approach this using one tigated in order to make the most of the available
of the many 'bioturbation index' schemes as ichnological data. It must also be remembered
14 D. McILROY

Fig. 9. Example ichnofabrics as developed under a range of palaeoenvironmental and sedimentological


conditions.

that ichnofabric is a 3D phenomenon, and that present within a rock unit can be simply quanti-
examination of a 2D surface (e.g. a cut surface fied, though care must be taken (especially in
of a core) can be misleading; if possible, an core) to ensure that a single trace in different
impression of ichnofabric in the opposite plane orientations (e.g. different cross-sections of the
should be sought in order to estimate the percen- same trace) is not counted twice. Although
tage bioturbation as a volume (see Fig. 8). trace fossil diversity cannot be directly related
to biological diversity it is usually considered as
Diversity a reasonable proxy. For example, a diverse
Ichnological diversity is comparatively simple to fauna of shallow infaunal bivalves may only
measure from both core and outcrop sections, produce two trace fossils (Lockeia as the resting
with experience. The number of trace fossils trace and Protovirgularia as the locomotory
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 15

trace). In most cases, however, high ichnological (but tentative) interpretation of the likely trophic
diversity corresponds to amenable palaeoenvir- niche of the trace-maker.
onmental conditions, and low diversity or lack
of bioturbation to harsh palaeoenvironments. Ichnometry
The exact causes of environmental stress are In addition to documenting the diversity and
diverse and must be assessed on a case-by case abundance of traces, the dimensions of the bur-
basis using ichnological, palaeontological, sedi- rows themselves can be of great importance.
mentological and sometimes geochemical/ The parameter that is most easily recorded is
palynological means. It is also observed that burrow diameter. Studies have shown that trace
ichnological diversity is more difficult to assess fossils become narrower with increased salinity
in core materials where identification below the stress (Hakes 1976; Gingras et al 1999) and
level of ichnogenus is seldom possible and the decreasing dissolved oxygen in porewaters/
morphology of complex branching forms is bottom waters (e.g. Bromley & Ekdale 1984).
difficult if not impossible to recognize (except This reflects well-known biological trends in
on the limited number of bedding surfaces). such settings (e.g. Milne 1940; Remane &
Schlieper 1971). In addition, during some periods
Relative abundance of rapid evolution there is a stratigraphic compo-
Merely presenting the number of different trace nent, in which burrow size increases with time,
fossils present within a rock unit can be a mis- e.g. the Cambrian explosion (Mcllroy & Logan
leading piece of information where the assem- 1999).
blage is dominated by a single ichnotaxon with
minor accessory components. Documentation Infaunal tiering
of the relative volumetric proportions of all The distribution of organisms (and their traces)
traces within an ichnofabric and their relative below the sediment is known as tiering. The
chronology is the fundamental procedure preservation of a tiering profile is reliant upon
behind good ichnofabric analysis. This can be rapid killing off of the community (e.g. burial
particularly instructive if combined with simple under an event bed or death of the community

Fig. 10. Ichnofabrics in outcrops of the Pacoota Sandstone, Amadeus Basin, central Australia showing:
(a) environmental deterioration represented by a decrease in burrow size of Skolithos', (b) development of an
early Arenicolites-dommated ichnocoenosis (Arenicolites ichnofacies) cross-cut by a later, Skolithos-domm&ted,
ichnocoenosis (Skolithos ichnofacies). The upper bed probably represents a single spatfall as all burrows are the
same (small) size. The small size of traces may represent burrows of juveniles buried during a phase of
gradually increasing bioturbation.
16 D. McILROY

due to an anoxic event), because - with continu- in a sediment in which the trace-making organ-
ing deposition - deeper burrows tend to over- isms are vertically partitioned into tiers contain-
print shallower ones (Fig. lOb). This tiering of ing more than one trace fossil (see Fig. 9).
the infaunal community is typically considered Multiple occupancy of a given tier should be
to be a response to partitioning of the infaunal demonstrable by mutual cross-cutting of the
realm into different niches occupied by organ- tier's occupant traces (see below).
isms with different feeding strategies (Bottjer &
Ausich 1982; Bromley 1990, 1996; Wetzel & Succession of bioturbation
Uchman 1998). The occupants of these different Understanding the order of emplacement of
niches, which share similar feeding strategies, burrows and tracks is a fundamental skill that
have been grouped into 'ichnoguilds' by Bromley all modern ichnologists should incorporate into
(1990), though these rely to a large degree on studies aimed at palaeoenvironmental analysis.
interpretation of behaviour, which is notoriously The recognition of successive cross-cutting
difficult to determine with accuracy for most palaeocommunities (ichnocoenoses, see below)
trace fossils. Diagrammatic representation of associated with the same sedimentary unit can
tiering may be done using either the tiering dia- lead to an improved understanding of the coloni-
grams of Bromley (1996, p. 295, fig. 12.11) and zation history and/or changing palaeoenviron-
Wetzel & Uchman (2001) or the more integrated mental conditions subsequent to the deposition
ichnofabric constituent diagrams (ICD) of of a given bed (cf. Wetzel & Uchman 2001).
Taylor & Goldring (1993; see the modified ICD One of the features of such cross-cutting relation-
in Fig. 6). More importantly however, several ships is that later burrows or tracks tend to
authors appear to use 'complex tiering' for all obscure earlier burrows, and deeper burrows
visually complex ichnofabrics (Taylor & Gold- tend to overprint shallower ones with continuing
ring 1993; Taylor et al. 2003). Complex ichno- sedimentation (see Figs 9, lOb). In many cases
logical tiering is defined herein as being formed one or a few trace fossils (termed elite trace

Fig. 11. Styles of colonization of sedimentary surfaces: (a) transport of adult and colonization from above;
(b) spatfall; (c) equilibration and colonization from below.
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 17

fossils by Bromley 1996) visually dominate an


ichnofabric. The elite trace fossil is often pro-
duced by a late-stage bioturbator(s), or may be
visually striking due to a prominent fill/burrow
lining.

Colonization styles
The colonization of sedimentary surfaces may be
achieved in a variety of ways, which can be
basically summarized as being from:
(a) a depositional surface;
(b) an eroded surface;
(c) beneath the sediment surface (usually of a
migrating bedform);
(d) a higher stratigraphic level (Fig. 11).
Options (a) and (b) can be difficult to distinguish
if the eroded sediment is unlithified and thus not
a classical 'Glossifungites surface' (cf. Mac-
Eachern et al. 1992; Fig. 6). Option (c) represents
the behaviour of a fauna well adapted to condi-
tions with high sedimentation rates (cf. the
mouth-bar facies of Mcllroy 2004). Colonization
of sediments from a higher stratigraphic level is a
common feature of turbidites (e.g. Kern 1980;
Buatois & Mangano 2004), though colonization
can be from below (e.g. Uchman 1995).
The ecology of colonization of disturbed/
defaunated sediment has been a focus of
marine ecological research for many years, and
has many applications in understanding the
palaeoecology of marine event beds. The
colonization of marine hardgrounds is generally
considered by ecologists to be largely by larval
settling (Roughgarden et al. 1985). In soft
marine sediments, however, colonization by
larval stages is also supplemented by passive
(currents) or active (locomotion) relocation of
adults (Hall 1994; Snelgrove & Butman 1994;
Cummings et al. 1995; Shull 1997). The potential
contribution to the sediments ichnofabric of
relocated adults is much greater than larval set-
tling (spatfall; see Fig. 1 Ib), and is best developed
in regions with high flow strengths where hydrau-
lic transport of adults is common. In addition,
spatfall may be seasonal in nature, in which
case the colonization window (sensu Pollard Fig. 12. Coarse-grained multi-storey tidal channel
et al. 1993) may be controlled by biological sandstones with rip-up clasts (upper arrow) with
anomalously intense bioturbation (from arrowed
rather than physical phenomena.
surface) by Diplocraterion with a well-developed
A sedimentological/ichnological description teardrop shape reflecting ontogenetic growth. From
that encompasses all the points above should the Tilje Formation, Norwegian Shelf. Core is 10cm
correspond to an excellent basis for further wide.
interpretation. The process of interpreting
ichnofabrics and bioturbated sediments is an For example, it is well known that modern
important step, which should involve careful marine communities are highly patchy in distri-
appraisal of the succession studied. Caution bution and not predictable relative to prevailing
should be exercised when assessing such detailed currents etc. This is demonstrated in the few
datasets, so that over-interpretation is avoided. studies of spatial distributions of ichnofaunas
18 D. McILROY

and ichnofabrics that have been performed thus assemblages, not ichnocoenoses, and it is impor-
far (Palmer & Palmer 1977; Goldring et al. tant that the two are not confused. An element of
1998; Uchman 2001) ichnofabric analysis is needed to recognize the
presence of the Arenicolites ichnofacies (Bromley
& Asgaard 1991), which is commonly over-
Ichnocoenoses printed by the Cruziana or Skolithos ichnofacies
(Fig. 11 a). A number of marine ichnofacies rely
The term 'ichnocoenose' was originally intro- upon ichnofabric analysis to distinguish palimp-
duced by Davitashvili (1945) to mean the traces sest fabrics representing different ichnofacies.
of a biological community or biocoenose (fide Some authors, however, do not generally use
Radwanski & Roniewicz 1970); ichnocoenose in these more subtle ichnofacies (e.g. Pemberton
its original sense thus means the ichnological et al. 1992, 2001, who do not discuss the Arenico-
equivalent of standing crop. Terms for the lites and Curvolithus ichnofacies). In addition,
buried and fossilized ichnocoenose - tapho- the use of the prefixes proximal/distal/depaupe-
coenose and orictocoenose respectively - were rate is increasingly common (e.g. Pemberton
also introduced by Davitashvili (1945), but et al. 2001; Bann & Fielding 2004; Buatois &
have fallen into disuse. The term 'ichnocoenosis' Mangano 2004). It would seem therefore that -
was subsequently independently introduced by although ichnofacies are useful as a starting
Lessertisseur (1955) to encompass fossil assem- point for ichnological studies - ichnologists
blages of ichnotaxa equivalent to biocoenose, working on shallow marine systems have out-
and was adopted in the latter sense by several grown ichnofacies, and indeed some ichnofacies
authors (Hantzschel 1962; Radwanski & actually hide important palaeoenvironmental
Roniewicz 1970). information (cf. Frey & Goldring 1992). The
As noted by Pickerill (1992), communities can future direction of ichnological work in the shal-
rarely, if ever, be recognized in the fossil record, low marine and ultimately the non-marine is
owing to the effects of time averaging. Trace probably through the creation of bespoke ichno-
fossils - though study of their cross-cutting logical models on a basin-by-basin scale, incor-
relationships (ichnofabric analysis) - can give a porating description of assemblages (cf. Fursich
detailed impression of the work of several 1976 for discussion of assemblages) and ichno-
successive communities within a bed. It is coenoses (as resolved by ichnofabric analysis)
useful, therefore, to retain ichnocoenosis as an rather than a reliance on the creation and appli-
approximation of its original meaning of the cation of ever more Seilacherian ichnofacies. The
traces of a biological community. Ichnocoenoses study of the non-marine is, however, still in its
should therefore ideally be considered as com- comparative infancy, and there is still much
prising a group of trace fossils that can be merit in using an ichnofacies-type approach (cf.
demonstrated - by ichnofabric analysis - to Buatois & Mangano 2004; Genise 2004).
have been formed by the action of what approx-
imates to a single benthic community or a succes-
sion of similar communities (based on Ekdale Ichnofabric stacking patterns as a correlative
et al. 1984). tool
The normal condition in the stratigraphic
record is that beds are colonized by a succession Having used some variety of the protocol out-
of different communities, and that several super- lined above to describe and understand the
imposed ichnocoenoses can be recognized. Such ichnology of the stratigraphic succession in ques-
time-averaged trace fossil fabrics are probably tion, the next challenge is to use these data in a
best known as assemblages. An ichnological meaningful manner. As described above, ichno-
assemblage is thus made up of all the trace fossils coenoses are the building block of applied ichno-
found within a given rock unit (usually a bed), logical studies, in both outcrop and core. In
regardless of their relative chronology, and may many cases beds may contain more than one ich-
be composed of one or more ichnocoenoses. nocoenosis - i.e. comprise an assemblage or asso-
ciation that makes up the sediment's ichnofabric.
Ichnocoenoses and Seilacherian ichnofacies In core studies, ichnofabric is the most usable
As highlighted by Bromley (1990, 1996), the use stratigraphic unit. In outcrop, however, where
of ichnocoenosis as being synonymous with good vertical sections are not always available,
Seilacherian ichnofacies (Dorjes & Hertweck associations/assemblages of interface traces and
1975; Frey & Pemberton 1987) is erroneous and the more prominent of the pervasive traces
problematic. Seilacherian ichnofacies were should suffice; quantification of the ichnology is
founded on the recognition of ichnological nonetheless still important.
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 19

By integrating ichnological and sedimento- horizons that, by ichnofabric/ichnofacies


logical data, a sedimentologist should be able analysis, can be demonstrated to be host to a
to produce a refined fades model, which may succession of ichnofaunas recording gradual
then form the basis for stacking pattern analysis deepening or shallowing events (Fig. 13);
at a variety of scales. Through application of horizons showing anomalously large burrows
Walther's law (Walther 1894), trends in relative that evince growth to adult size while living
sea-level within a given sedimentary package at a single horizon, e.g. loop-shaped Diplo-
can be established, progradational trends being craterion (Fig. 12);
recorded by deeper water facies/ichnofabrics/ horizons across which there is a dislocation of
associations being overlain by shallower water facies as evinced by ichnological and/or sedi-
facies/ichnofabrics/associations (as resolved in mentological analysis (Fig. 6).
the integrated conceptual facies/ichnofabric
Hypothetical model examples of the ichno-
model). Analysis of ichnofabric stacking patterns
logical expression of key stratigraphic surfaces
in such a way has been used to great effect to
are by no means intended to be exhaustive.
understand complex or difficult bioturbated
Each depositional system has its own unique
successions (Bockelie 1991; Taylor & Gawthorpe
character, and models such as that of Taylor
1993; Martin & Pollard 1996; Mcllroy 2004) at
et al. (2003) should be used for guidance, but
either the sequence or parasequence scale. The
departures from such idealized cases are to be
methodology is identical to that used by sedi-
expected.
mentologists in routine stratigraphic studies,
but incorporates both palaeontological and sedi-
mentological data for improved facies character-
ization. The above approach may also be used at Ichnological frontiers
a coarser scale with ichnofacies as the building
blocks. Progress
Departures from the expected succession of
facies predicted by applying Walther's law As outlined above, the field of ichnology has
(Walther 1894) to the conceptual facies model progressed apace, especially since the early
need to be explained by either autocyclic or 1970s when the seminal compilations of Crimes
allocyclic means. The surfaces thus identified & Harper (1970, 1977) exemplify the state of
are known as key stratigraphic surfaces and are rapid advancement of the subject and the bur-
discussed below. geoning interest in its applicability. Since the
early 1970s ichnology has become increasingly
relevant to a variety of related disciplines, includ-
ing zoology, ecology, archaeology, geochemistry,
Recognition of key stratigraphic surfaces diagenesis, sedimentology, sequence stratigra-
phy, petroleum reservoir characterization and
The recognition of key stratigraphic surfaces lies petroleum exploration.
at the heart of the sequence stratigraphic In recent years the ichnological understanding
approach to understanding and predicting sedi- of non-marine depositional systems has
mentological phenomena. The classification of improved enormously, largely through the
such surfaces has been gradually refined, partly work of South American ichnological research
through the recognition of the Glossifungites groups (e.g. Buatois & Mangano 1993, 2004;
ichnofacies (e.g. MacEachern et al. 1992; Pem- Genise et al. 2000, 2004). There is, however,
berton et al. 2004), but also through other still much to do by way of integrating this
styles of facies dislocation (Taylor & Gawthorpe improved understanding with sedimentological
1993; Taylor & Goldring 1993; Goldring 1999; and sequence stratigraphic work, as outlined by
Schlirf 2003; Taylor et al. 2003). Buatois & Mangano (2004).
Noteworthy phenomena are generally changes
to the normal ichnological patterns of a given
sedimentological succession, such as:
Ichnology and the petroleum industry
horizons with anomalously intense bioturba-
tion (Fig. 13); The most applied aspect of ichnological work
horizons with anomalously low or high ichno- involves studies that are of relevance to the
diversity (Fig. 13); petroleum industry. The utility of trace fossils
horizons with anomalous ichnofauna, e.g. a stems from the comparatively simple study of
horizon with marine trace fossils in an other- ichnofabrics in core, and the excellent palaeo-
wise non-marine succession (Fig. 13); environmental information that they hold (cf.
Fig. 13. Intercalation of marine and non-marine ichnofabrics from the Cloughton Formation, Jurassic, Yorkshire Coast, UK: (a) field photograph of the interval with
marine flooding and bioturbation; (b) ichnofabrics associated with the marine flooding surface; (c) ichnofabric constituent diagram of the ichnofabric in (b) above.
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 21

Chamberlain 1978 and many authors since) is the sediment, which can be involved in diagenetic
thus invaluable to industry. The demands of reactions that may reduce porosity and perme-
the modern petroleum industry on accurate ability after diagenesis (Worden & Morad
characterization of facies are highly exacting. 2003). Trace fossils may also connect sand
With the increased use of reservoir modelling of layers separated by impermeable mudstones,
sedimentary facies and highly detailed petro- thereby improving reservoir characteristics (Gin-
physical studies of reservoir units, the need for gras et al. 1999; Pemberton et al. 2001, 2004) and
high-quality interpretation of sedimentary systematically clean sandstones of clay (Frey &
facies is greater than ever. If facies are misidenti- Wheatcroft 1989; Prosser et al 1993). Recent
fied, the foundation stone of most other elements experimental ichnology has also demonstrated
of the reservoir characterization process falls that clay mineral authigenesis can occur in the
down, and nasty surprises may lie in store guts of organisms, making the clay mineral
during field development. By using all informa- assemblage of sediments metastable, and thereby
tion available to interpret sedimentary facies, influencing the of creation of clay mineral
such risks are minimised. cements upon burial (Mcllroy et al. 2003).

Facies characterization New technologies


The use of ichnology to characterize sedimentary Of the burgeoning technologies being developed
facies is well developed in shallow and marginal by the petroleum industry perhaps the most
marine depositional systems (e.g. Bockelie 1991; exciting for the ichnologist is that downhole
Mcllroy 2004) but is much less so in the non- imaging (FMI) is now just bordering on a resolu-
marine (see the recent inroads made by Genise tion whereby ichnology may become useful (e.g.
2004; Buatois & Mangano 2004), where deposi- Bourke 1992; Salimullah & Stow 1995), and can
tional systems are much more variable and only get better. Advantages include the potential
prone to the variable effects of climate and the for recovery of image data down the full length of
preservation potential of many trace fossils is the well, which means that the sedimentologist/
comparatively low. Future work on non-marine ichnologist need not work exclusively on cored
ichnology should work towards incorporating intervals. The image data are challenging to
animal and plant trace fossils (Bockelie 1994) in interpret, but ichnologists have been working
sedimentological and sequence stratigraphic with difficult sections of trace fossils in core for
models. many years, and should be adaptable enough to
In recent years much exploration effort has exploit this potentially rich data source.
been directed toward deep water turbidite plays,
e.g. Gulf of Mexico, west of Shetland, west of
Africa. This trend has been reflected in the Experimental and neoichnological studies
increased research into characterization of turbi-
dite architectural elements. These data have not, Experimental and neoichnological studies have a
however, been well integrated with ichnological rich history, including the classical studies of
studies despite there being many ichnologists modern sediments of both intertidal and subtidal
specializing in the ichnology of deep marine settings using box-coring and serial sectioning or
facies. The potential for combination studies X-ray analysis (e.g. Reineck 1958; Howard &
involving sedimentology and ichnology along- Reineck 1972). Such techniques need to be
side data from provenance techniques, palyno- improved and more closely related to deposi-
logical techniques (biostratigraphic and tional events in modern settings to facilitate
palynofacies; see MacEachern et al. 1999) and more informed interpretation of ancient environ-
geochemical techniques (especially in carbo- ments. Facilities for visualizing such data,
nates) is massively underused at present, and including tomography of serial sections (e.g. Fu
provides yet another rich source of potential et al. 1994; Sutton et al. 2001), X-ray and
information for petroleum geologists to use. NMR imaging techniques, have improved. In
addition, time-series X-rays of ichnofabrics in
Reservoir quality laboratory experiments has never been fully
Parameters of interest to the petroleum geologist exploited and should be very revealing.
are the porosity and permeability of potential In addition, ecological information concerning
reservoir intervals. These two parameters are to infauna of many modern environments is well
a large extent controlled by sedimentological established and ideal for incorporation into
heterogeneity but also by diagenesis. One feature models of ancient depositional environments
of bioturbated sandstones is that clay-grade (e.g. Reed 2002 on deep marine traces; Mcllroy
material is commonly mixed into the matrix of 2004 on tidal depositional environments).
22 D. McILROY

A. Martinius, P. Manning and M. Carton are thanked BROMLEY, R. G. & ASGAARD, U. 1993. Two bioerosion
for their reading of an early version of this manuscript. ichnofacies produced by early and late burial
The critical reviews of A. Uchman and M. Schlirf are associated with sea-level change. Geologische
acknowledged with thanks. A. Taylor and S. Gowland Rundschau, 82, 276-280.
are thanked for discussion prior to writing of this BROMLEY, R. G. & EKDALE, A. A. 1984. Chondrites: a
manuscript. Statoil asa and its employees past and trace fossil indicator of anoxia in Sediments.
present are also thanked for nurturing my involvement Science, 224, 872-874.
in the challenges of the modern petroleum geologist BROMLEY, R. G. & GOLDRING, R. 1992. The palaeo-
and for presenting me with interesting and pertinent burrows at the Cretaceous to Palaeocene firm-
challenges over the last six years or so. ground unconformity in southern England.
Tertiary Research, 13, 95-102.
BROMLEY, R. G., PEMBERTON, S. G. & RAHMANI, R. A.
References 1984. A Cretaceous woodground: the Teredolites
ichnofacies. Journal of Paleontology, 58, 488-498.
ALPERT, S. P. 1977. Trace fossils and the basal Cam- BROMLEY, R. G., JENSEN, M. & ASGAARD, U. 1995.
brian boundary. In: CRIMES T. P. & HARPER J. C. Spatangoid echinoids: deep tier trace fossils and
(eds) Trace Fossils 2. Geological Journal Special chemosymbiosis. Neues Jarbruchfur Geologic und
Issue, 9, Seel House Press, Liverpool, 1-8. Paldontologie, Abhandlungen, 195, 25-35.
BANN, K. L. & FIELDING, C. R. 2004. An integrated BUATOIS, L. A. & MANGANO, M. G. 1993. Trace fossils
ichnological and sedimentological comparison of from a Carboniferous turbiditic lake: implications
non-deltaic shoreface and subaqueous delta for the recognition of additional nonmarine ichno-
deposits in Permian reservoir units of Australia. facies. Ichnos,2, 97-114.
In: MclLROY, D. (ed.) The Application oflchnology BUATOIS, L. A. & MANGANO, M. G. 1995. The
to Palaeoenvironmental and Stratigraphic Analysis. palaeoenvironmental and palaeoecological signifi-
Geological Society, London, Special Publications, cance of the Mermia ichnofacies: an archetypal
228, 273-307. subaqueous non-marine trace fossil assemblage.
BERTLING, M., BRADDY, S. ET AL. 2003. Draft proposal Ichnos,4, 151-161.
to emend the code with respect to trace fossils: BUATOIS, L. A. & MANGANO, M. G. 2004. Animal-sub-
request for comments. Bulletin of Zoological strate interactions in freshwater environments:
Nomenclature, 60, 141-142. applications of ichnology in facies and sequence
BOCKELIE, J. F. 1991. Ichnofabric mapping and inter- Stratigraphic analysis of fluvio-lacustrine succes-
pretation of Jurassic reservoir rocks of the Nor- sions. In: MclLROY, D. (ed.) The Application of
wegian North Sea. Palaios, 6, 205-215. Ichnology to Palaeoenvironmental and Strati-
BOCKELIE, J. F. 1994. Plant roots in core. In: DONOVAN, graphic Analysis. Geological Society, London,
S. K. (ed.) The Palaeobiology of Trace Fossils. Special Publications, 228, 309-331.
Wiley & Sons, New York, 177-199. CHAMBERLAIN, C. K. 1978. Recognition of trace fossils
BOTTJER, D. J. & AUSICH, W. I. 1982. Tiering and in cores. In: BASAN, P. B. (ed.) Trace Fossil Con-
sampling requirements in paleocommunity recon- cepts. Society of Economic Paleontologists and
struction. Proceedings of the 3rd North American Mineralogists, Short Course, 5, 119-166.
Paleontological Convention, 1, 57-59. CORNET, B. & TRAVERSE, A. 1975. Palynological contri-
BOURKE, L. T. 1992. Sedimentological borehole image butions to the chronology and stratigraphy of the
analysis in clastic rocks: a systematic approach Hartford Basin in Connecticut and Massachusetts.
to interpretation. In: HURST, A., GRIFFITHS, C. M. Geoscience Manitoba, 11, 1-33.
& WORTHINGTON, P. F. (eds) Geological Applica- CRIMES, T. P. 1973. From limestones to distal turbi-
tions of Wireline Logs. Geological Society, dites: a facies and trace fossil analysis of the
London, Special Publications, 65, 31^2. Zumaya flysch (Paleocene-Eocene), North Spain.
BRASIER, M. D., COWIE, J. W. & TAYLOR, M. E. 1994. Sedimentology 20, 105-131.
Decision on the Precambrian-Cambrian bound- CRIMES, T. P. 1975. The production and preservation of
ary. Episodes, 17, 3-8. trilobite resting and furrowing traces. Lethaia, 8,
BRETT, C. E. 1998. Sequence stratigraphy, paleo- 35^8.
ecology, and evolution: biotic clues and responses CRIMES, T. P. 1987. Trace fossils and correlation of late
to sea-level fluctuations. Palaio, 13, 241-262. Precambrian and early Cambrian strata. Geologi-
BROMLEY, R. G. 1990. Trace Fossils: Biology and cal Magazine, 124, 97-119.
Taphonomy. Unwin Hyman, London. CRIMES, T. P. 1992. The record of trace fossils
BROMLEY, R. G. 1996. Trace Fossils: Biology, Taphon- across the Proterozoic-Cambrian boundary. In:
omy and Applications. Chapman & Hall, LIPPS, J. & SIGNOR, P.W. (eds) Origin and Early
London. Evolution of the Metazoa. Plenum, New York,
BROMLEY, R. G. & ASGAARD, U. 1979. Triassic fresh- 177-202.
water ichnocoenoses from Carsberg Fjord, East CRIMES, T. P. & HARPER, J. C. 1970. Trace Fossils.
Greenland. Palaeogeography, Palaeoclimatology, Geological Journal Special Issue, 3. Seel House
Palaeoecology, 28, 38-80. Press, Liverpool.
BROMLEY, R. G. & ASGAARD, U. 1991. Ichnofacies: a CRIMES, T. P. & HARPER, J. C. 1977. Trace Fossils 2.
mixture of taphofacies and biofacies. Lethaia, 24, Geological Journal Special Issue, 9, Seel House
153-163. Press, Liverpool.
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 23

CUMMINGS, V. J., PRIDMORE, R. D., THRUSH, S. F. & FREY, R. W. & PEMBERTON, S. G. 1984. Trace fossil
HEWITT, J. E. 1995. Post settlement movement facies models In: WALKER, R. G. (ed.) Fades
by intertidal benthic macroinvertebrates: do Models. Geological Association of Canada,
common New Zealand species drift in the water Geoscience Canada Reprint Series 1, 189-207.
column? New Zealand Journal of Marine and FREY, R. W. & PEMBERTON, S. G. 1987. The Psilon-
Freshwater Research, 29, 59-67. ichnus ichnocoenose, and its relationship to adja-
CURRAN, H. A. 1994. The palaeobiology of ichno- cent marine and non-marine ichnocoenoses along
coenoses in Quaternary Bahamian-style carbonate the Georgia Coast. Bulletin of Canadian Petroleum
environments: the modern to fossil transition. In: Geology, 35, 333-357.
DONOVAN, S. K. (ed.) The Palaeobiology of Trace FREY, R. W. & SEILACHER, A. 1980. Uniformity in
Fossils. Wiley, New York. 83-104. marine invertebrate ichnology. Lethaia, 13, 183-
DAVITASHVILI, L. SH. 1945 Tsenozy zhivykh organiz- 207.
mov I organicheskikh ostatkov [Assemblages of FREY, R. W. & WHEATCROFT, R. A. 1989. Organism-
live organisms and of organic remains]. Akade- sediment relations and their impact on sedimen-
miya Nauk Gruzin SSR, 6, 527-534. tary petrology. Journal of Geological Education,
DIEGO, T. D. & DOUGLAS, R. G. 1999. Oxygen related 37, 261-279.
sediment microfabrics in modern 'black shales', FREY, R. W., PEMBERTON, S. G. & FAGERSTROM, J. A.
Gulf of California, Mexico. Journal of Foraminif- 1984. Morphological, ethological, and environ-
eral Research, 29, 453-464. mental significance of the ichnogenera Scoyenia
DORIES, J. & HERTWECK, G. 1975. Recent biocoenoses and Ancorichnus. Journal of Paleontology, 58,
and ichnocoenoses in shallow water marine envir- 511-528.
onments. In: FREY, R. W. (ed.) The Study of Trace FREY, R. W., PEMBERTON, S. G. & SAUNDERS, T. D.
Fossils. Springer, New York, 459-491. 1990. Ichnofacies and bathymetry: a passive
DROSER, M. L. & BOTTJER, D. J. 1986. Trends in the relationship. Journal of Paleontology, 64, 155—158.
depth and extent of bioturbation in Cambrian Fu, S. 1991. Funktion, Verhalen un Einteilung fucoider
carbonate marine environments, western United und lophocteniider Lebenspurren. Institut Senck-
States. Geology, 16, 233-236. enberg, Courier Forschungs, 135, 1-79.
DROSER, M. L. & BOTTJER, D. J. 1989. Ordovician Fu, S., WERNER, F. & BROSSMANN, J. 1994. Computed
increase in the extent and depth of bioturbation: tomography: application in studying biogenic
implications for understanding early ecospace structures in sedimentary cores. Palaios, 9, 116-
utilization. Geology, 17, 850-852. 119.
DROSER, M. L. & BOTTJER, D. J. 1991 Trace fossils and FURSICH, F. T. 1975. Trace fossils as environmental
ichnofabric in Leg 119 cores. Proceedings of the indicators in the Corallian of England and
Ocean Drilling Program, Scientific Results, 119, Normandy. Lethaia, 8, 151-172.
635-641. FURSICH, F. T. 1976. The use of macroinvertebrate
DROSER, M. L. & O'CONNELL, S. 1992. Trace fossils and association in interpreting Corallian (Upper
ichnofabric in Triassic sediments from cores Jurassic) environments. Palaeogeography, Palaeo-
recovered on Leg 112. Proceedings of the Ocean climatology, Palaeoecology, 20, 235-256.
Drilling Program, Scientific Results, 122, 475-485. GENISE, J. F. 2004. Ichnotaxonomy and ichnostratigra-
EKDALE, A. & MASON, T. 1988. Characteristic trace phy of chambered trace fossils in palaeosols
fossil assemblages in oxygen-poor sedimentary attributed to coleopterans, ants and termites. In:
environments. Geology, 16, 720-723. MclLROY, D. (ed.) The Application of Ichnology
EKDALE, A. A., BROMLEY, R. G. & PEMBERTON, S. G. to Palaeoenvironmental and Stratigraphic Analysis.
1984. Ichnology: The Use of Trace Fossils in Geological Society, London, Special Publications,
Sedimentology and Stratigraphy. Society of Eco- 228,417^51.
nomic Palaeontologists and Mineralogists Short GENISE, J. F., MANGANO, M. G., BUATOIS, L. A., LAZA,
Course, 15, 1-317. J. H. & VERDE, M. 2000. Insect trace fossil associa-
ELLENBERGER, F., ELLENBERGER, P. & GINSBURG, L. tions in paleosols: the Coprinisphaera ichnofacies.
1970. Les dinosaurs du Trias et du Lias en Palaios 15, 49-64.
France et en Afrique du Sud, d'apres les pistes GENISE, J., BELLOSI, E. S. & GONZALEZ, M. G. 2004. An
qu'ils on laissees. Bulletin de le Societe Geologique approach to the description and interpretation of
de France, 7, ser., 12, 151-159. ichnofabrics in palaeosols. In: MC!LROY, D. (ed.)
FEDONKIN, M. A., LINAN, E. & PEREJON, A. 1983. Icno- The Application of Ichnology to Palaeoenvironmen-
fossiles de las rocas Precambrico-Cambricas de la tal and Stratigraphic Analysis. Geological Society,
Sierra de Cordoba. Espafla. Boletin Real Sociadad London, Special Publications, 228, 353-380.
Espanola (Geologicd), 81, 125-138. GHIBAUDO, G., GRANDESSO, P., MASSARI, F. &
FREY, R. W. & GOLDRING, R. 1992. Marine event UCHMAN, A. 1996. Use of trace fossils in delineat-
beds and recolonization surfaces as revealed by ing sequence Stratigraphic surfaces (Tertiary Vene-
trace fossil analysis. Geological Magazine, 129, tian Basin, northeastern Italy). Palaeogeography,
325-335 Palaeoclimatology, Palaeoecology, 120, 261-279.
FREY, R. W. & HOWARD, J. D. 1990. Trace fossils and GIERLOWSKI-KORDESCH, E. 1991. Ichnology of an
depositional sequences in a clastic shelf setting, ephemeral lacustrine/alluvial plain system:
Upper Cretaceous of Utah. Journal of Paleontol- Jurassic East Berlin Formation, Hartford Basin,
ogy, 64, 803-820. USA. Ichnos, 1, 221-232.
24 D. McILROY

GINGRAS, M. K., PEMBERTON, S. G., MENDOZA, C. A. & Vardel0ft Formation (Middle Jurassic, Central
HENK, F. 1999. Assessing the anisotropic perme- East Greenland). Journal of Paleontology, 58,
ability of Glossifungitessurfaces.Petroleum 362-397.
Geoscience, 5, 349-357. HOVLAND, M. & THOMSEN, E. 1989. Hydrocarbon-
GLAUB, I. 2004. Recent and sub-recent microborings based communities in the North Sea? Sarsia, 74,
from the upwelling area off Mauritania (West 29-42.
Africa) and their implications for palaeoecology. HOWARD, J. D. & FREY, R. W. 1975. Regional animal-
In: MclLROY, D. (ed.) The Application oflchnology sediment characteristics of Georgia estuaries.
to Palaeoenvironmental and Stratigraphic Analysis. Senkenbergiana Maritima, 7, 33-103.
Geological Society, London, Special Publications, HOWARD, J. D. & REINECK, H. E. 1972. Georgia coastal
228, 63-76. region, Sapelo Island, USA: Sedimentology and
COLORING, R. 1993. Ichnofacies and facies interpreta- biology. IV. Physical and biogenic sedimentary
tion. Palaios, 8, 403^05. structures of the nearshore shelf. Senckenbergiana
GOLDRING, R. 1995. Organisms and the substrate: Maritima, 4, 81-123.
response and effect. In: BOSENCE, D. W. J. & HOWELL, J. A. & AITKEN, J. F. (eds) 1996. High
ALLISON, P. A. (eds) Marine Palaeoenvironmental resolution sequence stratigraphy: innovations and
Analysis from Fossils. Geological Society, applications. Geological Society, London, Special
London, Special Publications, 83, 151-180. Publications, 104.
GOLDRING, R. 1999. Field Palaeontology (2nd edn). HUNT, A. P., CHIN, K. & LOCKLEY, M. G. 1994. The
Longman, Harlow. palaeobiology of vertebrate coprolites. In:
COLORING, R., ASTIN, T. R., MARSHALL, J. E. A., DONOVAN, S. K. (ed.) The Palaeobiology of Trace
GABBOT, S. & JENKINS, C. D. 1998. Towards an Fossils. Wiley & Sons, New York, 221-241.
integrated study of the depositional environment ICZN 1964 International Code of Zoological
of the Bencliff Grit (Upper Jurassic) of Dorset. Nomenclature (SecondEdition). The International
In: UNDERBILL, J. R. (ed.) Development and Trust for Zoological Nomenclature, London,
Evolution of the Wessex Basin. Geological Society, UK.
London, Special Publications, 133, 335-372. ICZN 1985. International Code of Zoological
GOWLAND, A. S. 1996. Facies characteristics and Nomenclature (Third Edition). RIDE, W. D. L.,
depositional models of highly bioturbated shallow SABROSKY, C. W., BERNADI, G. & MELVILLE,
marine siliciclastic strata: an example from the R. V. (eds). The International Trust for Zoological
Fulmar Formation (late Jurassic), UK Central Nomenclature, London, UK.
Graben. In: HURST, A. (ed.) Geology of the ICZN 1999. International Code of Zoological Nomen-
Number Group: Central Graben and Moray Firth, clature (Fourth Edition). RIDE, W. D. L.,
UKCS. Special Publication of the Geological COGGER, H. G., DUPUNIS, C., KRAUS, O., MINELLI,
Society, London, 114, 185-214. A., THOMPSON, F. C. & TUBBS, P. K (eds). The
HAKES, W. G. 1976. Trace fossils and depositional International Trust for Zoological Nomenclature,
environment of four clastic units, Upper Pennsyl- London, UK.
vanian megacyclothems, northeast Kansas. JABLONSKI, D. 1991. Extinctions: a paleontological
University of Kansas Paleontological Contribu- perspective. Science, 253, 375—368.
tions, 63. KAUFMANN, R. S. & SMITH, K. L. 1997 Activity pat-
HALL, S. J. 1994. Physical disturbance and marine terns of mobile epibenthic megafauna at an abys-
benthic communities: life in unconsolidated sedi- sal site in the eastern North Pacific: results from
ments. Oceanography & Marine Biology: An a 17 month time-lapse photographic study. Deep
Annual Review, 32, 178-239. Sea Research, Part 1, Oceanographic Research
HANTZSCHEL, W. 1962. Trace fossils and problematica. Papers, 44, 559-579.
In: MOORE, R. C. (ed.) Treatise on Invertebrate KERN, J. P. 1980. Origin of trace fossils in Polish
Paleontology, Part W, University of Kansas Carpathian flysch. Lethaia, 13, 347-362.
Press, Lawrence, ^777-^245. KSIAZKIEWICZ, M. 1970. Observations on the ichno-
HANTZSCHEL, W. 1965. Vestigia invertebratorum et fauna of the Polish Carpathians. In: CRIMES, T. P.
problematica. Fossilum Catalogus I. Animalia, & HARPER, J. C. (eds) Trace Fossils. Geological
108. Journal Special Issue, 3. Seel House Press, Liver-
HANTZSCHEL, W. 1975. Trace fossils and problematica. pool. 283-322.
In: Teichert, C. (ed.) Treatise on Invertebrate LEMECHE, H. 1973. Comments on the application con-
Paleontology, Part W. Miscellanea, Supplement 1. sidering trace fossils. Bulletin of Zoological
Geological Society of America and University of Nomenclature, 30, 70.
Kansas Press, Boulder & Lawrence. LESSERTISSEUR, J. 1955. Traces fossils d'activite animate
HANTZSCHEL, W. & KRAUS, O. 1972. Names based on et leur significance paleobiologique. Societe Geolo-
trace fossils (ichnotaxa): request for a recommen- gique de France, Memoire, 74.
dation. Z.N.(S.) 1973. Bulletin of Zoological LOCKLEY, M. G., HUNT, A. P. & MEYER, C. A. 1994.
Nomenclature, 29, 137-141. Vertebrate tracks and the ichnofacies concept:
HAUBOLD, H. 1984. Saurierfdhrten. Ziemsen, Witten- implications for palaeoecology and palichno-
berg. stratigraphy. In: DONOVAN, S. K. (ed.) The Palaeo-
HEINBERG, C. & BIRKELUND, T. 1984. Trace fossil biology of Trace Fossils. Wiley & Sons, New York,
assemblages and basin evolution of the 221-241.
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 25

LOCKLEY, M. G., RlNDSBERG, A. K. & ZEILER, R. M. Journal of the Geological Society, London, 160,
1987. The palaeoenvironmental significance of 489^93.
the nearshore Curvolithus ichnofacies. Palaios, 2, MELVILLE, R. V. 1979. Further proposed amendments
255-262. to the International Code for Zoological Nomen-
MACEACHERN, J. A., RAYCHAUDHURI, I. & PEMBERTON, clature Z.N.(G.) 182. Bulletin of Zoological
S. G. 1992. Stratigraphic applications of the Nomenclature, 36, 11—14.
Glossifungites ichnofacies: delineating discontinu- MILLER, M. F. & SMAIL, S. E. 1997. A semiquantitative
ities in the rock record. In: PEMBERTON, S. G. (ed.) field method for evaluating bioturbation on bed-
Applications of Ichnology to Petroleum Explora- ding planes. Palaios, 12, 391-396.
tion: a Core Workshop. SEPM Core Workshop MILNE, A. 1940. The ecology of the Tamar Estuary, IV:
17. Society for Sedimentary Geology, Tulsa, Okla- the distribution of fauna and flora on buoys.
homa, 169-198. Journal of the Marine Biologists Association, UK,
MACEACHERN, J. A., ZAITLIN, B. A. & PEMBERTON, 24, 69-87.
S. G. 1999. A sharp-based sandstone of the MOORE, H. B. & SCRUTTON, P. C. 1957. Minor internal
Viking Formation, Joffre Field, Alberta, Canada: structures of some recent unconsolidated sedi-
Criteria for recognition of transgressively incised ments. American Association of Petroleum Geolo-
shoreface complexes. Journal of Sedimentary gists, Bulletin, 41, 2723-2751.
Research 69, 876-892. NARBONNE, G., MYROW, P. M., LANDING, E. & ANDER-
MAGWOOD, J. P. A. & PEMBERTON, S. G. 1988. Trace SON, M. M. 1987. A candidate stratotype for the
fossils of the Gog Group, a lower Cambrian tidal Precambrian-Cambrian boundary, Fortune
sand body, Lake Louise, Alberta. In: LANDING, Head, Burin Peninsula, southeastern Newfound-
E., NARBONNE, G. M. & MYROW, P. (eds) Trace land. Canadian Journal of Earth Sciences, 24,
Fossils, Small Shelly Fossils and the Precambrian 1277-1293.
Cambrian Boundary. New York State Museum Bul- NYSTUEN, J. P. 1998. History and development of
letin, 463. sequence stratigraphy. In: GRADSTEIN, F. M.,
M ALPAS, J. A. 2000. Integrated sedimentology SANDVIK, K. O. & MILTON, N. J. (eds) Sequence
and palaeoenvironmental analysis of marine Stratigraphy: Concepts and Applications. Norwe-
flooding surfaces: a case study of the Miocene, gian Petroleum Society (NPF) Special Publica-
Nukhul Formation, Gulf of Suez. American tions, 8, 31-116.
Association of Petroleum Geologists, Bulletin, 84, OLORIZ, F. & RODRIGUEZ-TOYAR, F. J. 1999. Diplocra-
1867-1868. terion: a useful marker for sequence stratigraphy
MANNING, P. L. 2004. A new approach to the analysis and correlation in the Kimmeridgian, Jurassic
and interpretation of dinosaur tracks. In: (Prebetic Zone, Betic Cordillera, southern
MclLROY, D. (ed.) The Application of Ichnology Spain). Palaios, 15, 546-552.
to Palaeoenvironmental and Stratigraphic Analysis. OLSEN, P. E. 1980. A comparison of the vertebrate
Geological Society, London, Special Publications, assemblages from the Newark and Hartford
228, 93-123. Basins (Early Mesozoic, Newark Supergroup) of
MARTIN, K. D. 2004. A re-evaluation of the relation- eastern North America. In: JACOBS, L. L. (ed.)
ship between trace fossils and dysoxia. In: Aspects of Vertebrate History. Museum of
MclLROY, D. (ed.) The Application of Ichnology Northern Arizona, Flagstaff, Arizona, 35-53.
to Palaeoenvironmental and Stratigraphic Analysis. OSCHMANN, W. 199 la. Anaerobic-poikiloaerobic-
Geological Society, London, Special Publications, aerobic: a new facies zonation for modern and
228, 141-156. ancient nereitic redox facies. In: EINSELE, G.,
MARTIN, M. A. & POLLARD, J. E. 1996. The role of RICKEN, W. & SEILACHER, A. (eds) Cycles and
trace fossil (ichnofabric) analysis in the develop- Events in Stratigraphy. Springer, Berlin, 565-571.
ment of depositional models for the Upper Juras- OSCHMANN, W. 1991b. Distribution, dynamics and
sic Fulmar Formation of the Kittiwake Field palaeoecology of Kimmeridgian (Upper Jurassic)
(Quadrant 21 UKCS). In: HURST, A. (ed.) Geology shelf anoxia in western Europe. In: TYSON, R. V.
of the Number Group: Central Graben and Moray & PEARSON, T. H. (eds) Modern and Ancient
Firth, UKCS. Geological Society, London, Special Shelf Anoxia. Special Publication of the Geo-
Publications, 114, 163-183. logical Society, London, 58, 381-395.
MclLROY, D. 2004. Ichnology and facies model of a PALMER, T. J. & PALMER, C. D. 1977. Faunal distribu-
tide-dominated delta: Jurassic upper Ror and He tion and colonization strategy in a Middle
Formations of Kristin Field, Halten Terrace, Off- Ordovician hardground community. Lethaia, 10,
shore Mid-Norway In: MclLROY, D. (ed.) The 179-199.
Application of Ichnology to Palaeoenvironmental PAULL, C. K., HECKER, B. et al. 1984. Biological com-
and Stratigraphic Analysis. Geological Society, munities at the Florida Escarpment resemble
London, Special Publications, 228, 237-272. hydrothermal vent taxa. Science, 226, 965-967.
MclLROY, D. & LOGAN, G. A. 1999. The impact of bio- PEMBERTON, S. G., MACEACHERN, J. A. & FREY, R. W.
turbation on infaunal ecology and evolution 1992. Trace fossil facies models: environmental
during the Proterozoic-Cambrian transition. and allostratigraphic significance. In: WALKER,
Palaios, 14, 58-72. R. G. & JAMES, N. P. (eds) Facies Models Response
MclLROY, D., WORDEN, R. H. & NEEDHAM, S. J. 2003. to Sea Level Change. Geological Association of
Faeces, clay minerals and reservoir potential. Canada, 47-72.
26 D. McILROY

PEMBERTON, S. G., MACEACHERN, J. A., GINGRAS, M. SAGEMANN, B. B., WIGNALL, P. B. & KAUFFMANN, E. G.
K. & ZANG, J. 2000. Significance of ichnofossils 1991. Biofacies models for oxygen deficient facies
to genetic stratigraphy: examples from the Cretac- in epicontinental seas: a tool for palaeoenviron-
eous of Alberta, Canada. Science in China, Series mental analysis. In: EINSELE, G., RICKEN, W. &
D, Earth Sciences, 43, 541-560. SEILACHER, A. (eds) Cycles and Events in Stratigra-
PEMBERTON, G. S., SPILA, M., PULHAM, A. J., SAUN- phy. Springer, Berlin, 542-564.
DERS, T., ROBBINS, D. & SINCLAIR, I. K. 2001. Ich- SALIMULLAH, A. R. M. & STOW, D. A. V. 1995. Ichno-
nology and Sedimentology of Shallow to Marginal facies recognition in turbidites/hemipelagites using
Marine Systems. Geological Association of enhanced FMS images: examples from ODP Leg
Canada Short Course Volume 15. 129. The Log Analyst, 36, 38-49.
PEMBERTON, G. S., MACEACHERN, J. A. & SAUNDERS, T. SARJEANT, W. A. S. 1979. Code for trace fossil nomen-
2004. Stratigraphic applications of substrate- clature Palaeogeography, Palaeoclimatology,
specific ichnofacies: delineating discontinuities in Palaeoecology, 28, 147-166.
the rock record. In: MC!LROY, D. (ed.) The Appli- SARJEANT, W. A. S. & KENNEDY, W. J. 1973. Proposal
cation of Ichnology to Palaeoenvironmental and for a code for the nomenclature of trace fossils.
Stratigraphic Analysis. Geological Society, Canadian Journal of Earth Science, 10, 460-475.
London, Special Publications, 228, 29-62. SAVRDA, C. E. 1991. Ichnology in sequence strati-
PICKERILL, R. K. 1992. Carboniferous nonmarine graphic studies: an example from the Lower Paleo-
invertebrate ichnocoenoses from southern New cene of Alabama. Palaios, 6, 39-53.
Brunswick, eastern Canada. Ichnos, 2, 21-35. SAVRDA, C. E. & BOTTJER, D. J. 1987. The exaerobic
PICKERILL, R. K. & NARBONNE, G. M. 1995. Composite zone, a new oxygen deficient marine biofacies.
and compound ichnotaxa: a case example from the Nature, 327, 54-56.
Ordovician of Quebec, eastern Canada. Ichnos, 4, SAVRDA, C. E. & BOTTJER, D. J. 1991.Oxygen-related
53-71. biofacies in maritime strata: an overview and
PIKE, J., BERNARD, J. M., MORETON, S. G. & BUTLER, update. In: TYSON, R. & PEARSON, T. H. (eds)
I. B. 2001. Microbioirrigation of marine sediments Modern and Ancient Continental Shelf Anoxia.
in dysoxic environments: implications for early Geological Society, London, Special Publications,
sediment fabric formation and diagenetic pro- 58, 201-219.
cesses. Geology, 29, 923-926. SCHIEBER, J. 2003. Simple gifts and buried treasures:
POLLARD, J. E., GOLDRING, R. & BUCK, S. G. 1993. Ich- implications of finding bioturbation and erosion
nofabrics containing Ophiomorpha: significance in surfaces in black shales. The Sedimentary Record,
shallow-water facies interpretations. Journal of the 1,4^8.
Geological Society, London, 150, 149-164. SCHLIRF, M. 2003. Palaeoecologic significance of Late
PROSSER, D. J., DAWS, J. A., FALLICK, A. E. & Jurassic trace fossils from the Boulonnais, N.
WILLIAMS, B. P. J. 1993. Geochemistry and diagen- France. Ada Geologica Polonica, 53, 123-142.
esis of stratabound calcite cement layers within the SEILACHER, A. 1964. Biogenic sedimentary structures.
Rannoch Formation of the Brent Group, Murch- In: IMBRIE, J. & NEWELL, N. (eds) Approaches to
inson Field, North Viking Graben (Northern Paleoecology. Wiley, New York, 296-316.
North Sea). Sedimentary Geology, 87, 139-164. SEILACHER, A. 1967. Bathymetry of trace fossils.
RADWANSKI, A. & RONIEWICZ, P. 1970. General Marine Geology, 5, 413-428.
remarks on the ichnocoenose concept. Bulletin de SEILACHER, A. 1970. Cruziana stratigraphy of 'non
I'Academic Polonaise des Sciences, Serie des fossiliferous' Palaeozoic sandstones. In: CRIMES,
Sciences Geologiques et Geographiques, 18, 51-56. T. P. & HARPER, J. C. (eds) Trace Fossils. Geo-
READING, H. G. (ed.) 1978. Sedimentary Environments logical Journal Special Issue, 3, 447-476.
and Facies. Blackwell, Oxford. SEILACHER, A. 1974. Flysch trace fossils: evaluation of
REED, C. 2002. Lighting the mysteries of the abyss. behavioural diversity in the deep-sea. Neues Jar-
Geotimes, 47, 24-25, bruchfur Geologic und Paldontologie Monatshefte,
REINECK, H. E. 1958. Kastengreifer und Lotrohre 4, 233-245
'Schnepfe' Gerate zur Entnahme ungrestorter, SEILACHER, A. 1985. Trilobite palaeobiology and sub-
orientierter Meeresgrundproben. Senckenbergiana strate relationships. Transactions of the Royal
Lethaea, 39, 42^8, 54-56. Society of Edinburgh, 76, 231-237.
REINECK, H. E. 1963. Sedimentgefiige im Bereich der SEILACHER, A. 1990. Aberrations in bivalve evolution
siidlichen Nordsee. Abhandlungen der Senckenber- related to photo- and chemosymbiosis. Historical
gischen Naturforschenden Gesellschaft, 505, 1-107. Biology, 3, 289-311.
REMANE, A. & SCHLIEPER, C. 1971. Biology of Brackish SEILACHER, A. 1992. An updated Cruziana stratigraphy
Water. Wiley, New York. of Gondwanian Palaeozoic sandstones. In: SALEM,
RINDSBERG, A. K. & MARTIN, A. J. 2003. Arthrophycus M. J. (ed.) The Geology of Libya. Elsevier, Amster-
in the Silurian of Alabama (USA) and the problem dam, 1565-1580.
of compound trace fossils. Palaeogeography, SEILACHER, A. 1993. Problems of correlation in the
Palaeoclimatology, Palaeoecology, 192, 187-219. Nubian Sandstone facies. In: THORWEIHE, U. &
ROUGHGARDEN, J., IWASA, Y. & BAXTER, C, 1985. SCHANDELMEIR, H. (eds) Geoscientific Research in
Demographic theory for an open marine popula- Northwest Africa. Balkema, Rotterdam, 329-333.
tion with space-limited recruitment. Ecology, 6, SEILACHER, A. 1994. How valid is Cruziana stratigra-
54-67. phy? Geologische Rundschau, 83, 752-758.
ICHNOLOGY & PALAEOENVIRONMENTAL ANALYSIS 27

SHULL, D. H. 1997. Mechanisms of infaunal polychaete VAIL, P. R., MITCHUM, R. M. et al. 1977. Seismic
dispersal and colonization on an intertidal sand- stratigraphy and global changes of sealevel. In:
flat. Journal of Marine Research, 55, 153—179. Pay ton, C. (ed.) Seismic stratigraphy: Applications
SIGGERUD, E. I. H. & STEEL, R. J. 1999. Architecture and to Hydrocarbon Exploration. American Asso-
trace-fossil characteristics of a 10,000-20,000 year, ciation of Petroleum Geologists, Memoirs, 26,
fluvial-to-marine sequence, SE Ebro Basin, Spain. 49-212.
Journal of Sedimentary Research, 69, 365—383. VOSSLER, S. M. & PEMBERTON, S. G. 1988. Skolithos in
SMITH, C. R., LEVIN, L. A., HOOVER, D. J., MCMURTRY, the Upper Cretaceous Cardium Formation: an
G. & GAGE, J. D. 2000. Variations in bioturbation ichnological example of opportunistic ecology.
across the oxygen minimum zone in the northwest Lethaia, 21, 351-362.
Arabian Sea. In: GAGE, J. D., LEVIN, L. A. & WALTHER, J. 1894. Einleitung in die Geologic als
WOLFF, G. A. (eds) Bent hie Processes in the Deep Historische Wissenschaft, Bd. 3. Lithogenesis der
Arabian Sea: Biogeochemistry, Biodiversity and Gegenwart. G. Fischer, Jena, 535-1055.
Ecology. Deep Sea Research Part II. Topical WETZEL, A. 1991. Ecologic interpretation of deep-sea
Studies in Oceanography, 47, 227-257. trace fossil communities. Palaeogeography,
SMITH, R. M. H., MASON, T. R. & WARD, L. F. 1993. Palaeoclimatology, Palaeoecology, 85, 47-69.
Flash flood sediments and ichnofacies of the WETZEL, A. & UCHMAN, A. 1998. Deep-sea benthic
Late Pleistocene Homeb Silts, Kuiseb River, food content recorded by ichnofabrics: a con-
Namibia. Sedimentary Geology, 85, 579-599. ceptual model based on observations from
SNELGROVE, P. V. R. & BUTMAN, C. A. 1994. Animal- Paleogene flysch, Carpathians, Poland. Palaios,
sediment relationships revisited: cause versus 13, 533-546.
effect. Oceanography & Marine Biology: An WETZEL, A. & UCHMAN, A. 2001. Sequential coloniza-
Annual Review, 32, 111-177. tion of muddy turbidites in the Eocene Beloveza
SUTTON, M. D. BRIGGS, D. E. G., SIVETER, D. J. & Formation, Carpathians, Poland. Palaeogeogra-
SIVETER D. J. 2001. Methodologies for the visuali- phy, Palaeoclimatology, Palaeoecology, 168, 171-
zation and reconstruction of three-dimensional 186.
fossils from the Silurian Herefordshire Lagerstatte. WIGNALL, P. B. 1993. Distinguishing between oxygen
Palaeontologica Electronica, 4, 1-17. and substrate control in fossil benthic assem-
TAYLOR, A. M. & GAWTHORPE, R. L. 1993. Application blages. Journal of the Geological Society, London,
of sequence stratigraphy and trace fossil analysis 150, 193-196.
to reservoir description: examples from the WIGNALL, P. B. 1994. Black Shales. Clarendon Press,
Jurassic of the North Sea. Petroleum Geology of Oxford.
Northwest Europe: Proceedings of the 4th Confer- WIGNALL, P. B. & MYERS, K. 1998. Interpreting benthic
ence, 317-335. oxygenation levels in mudrocks: a new approach.
TAYLOR, A. M. & GOLDRING, R. 1993. Description and Geology, 16, 452-455.
analysis of bioturbation and ichnofabric. Journal WIGNALL, P. B. & NEWTON, R. 2001. Black shales on
of the Geological Society, London, 150, 141-148. the basin margin: a model based on examples
TAYLOR, A. M., COLORING, R. & GOWLAND, S. 2003. from the Upper Jurassic of the Boulonnais,
Analysis and application of ichnofabrics. Earth northern France. Sedimentary Geology, 144, 335-
Science Reviews, 60, 227-259. 356.
TWITCHETT, R. J. & BARRAS, C. G. 2004. Trace fossils WIGNALL, P. B. & PICKERING, K. T. 1993. Palaeoecol-
in the aftermath of mass extinction events. In: ogy and sedimentology across a Jurassic fault
MclLROY, D. (ed.) The Application of Ichnology scarp, NE Scotland. Journal of the Geological
to Palaeoenvironmental and Stratigraphic Analysis. Society, London, 150, 323-340.
Geological Society, London, Special Publications, WILKINS, R., BARNES, H. & BRANTLEY, S. 1996. The
228, 395^15. size distribution of framboidal pyrite in modern
UCHMAN, A. 1995. Tiering patterns of trace fossils in sediments: an indicator of redox conditions.
Paleogene flysch deposits of the Carpathians, Geochimica et Cosmochimica Acta, 60, 3897-
Poland. Geobios, 18, 389-394. 3912.
UCHMAN, A. 2001. Eocene flysch trace fossils from the WORDEN, R. H. & MORAD, S. 2003. Clay minerals in
Hecho Group of the Pyrenees, northern Spain. sandstones: controls on formation distribution
Beringeria, 15, 3-41. and evolution. In: WORDEN, R. H. & MORAD, S.
UCHMAN, A., BUBNIAK, I. & BUBNIAK, A. 2000. The (eds) Clay Mineral Cements in Sandstones. Inter-
Glossifungites ichnofacies in the area of its national Association of Sedimentologists, Special
nomenclatural archetype, Lviv, Ukraine. Ichnos, Publications, 34, 3-41.
7, 183-195.
This page intentionally left blank
Stratigraphic applications of substrate-specific ichnofacies:
delineating discontinuities in the rock record

S. GEORGE PEMBERTON1, JAMES A. MAcEACHERN2 & TOM SAUNDERS1


1
Department of Earth & Atmospheric Sciences, University of Alberta, Edmonton,
Alberta, Canada, T6G 2E3
2
Department of Earth Sciences, Simon Eraser University, Burnaby,
British Columbia, Canada, V5A 1S6

Abstract: Trace fossils represent both sedimentological and palaeontological entities,


providing a unique blending of potential environmental indicators in the rock record.
Trace fossils and trace fossil suites can be employed effectively to aid in the recognition of
various discontinuity types and to assist in their genetic interpretation. Ichnology may be
employed to resolve surfaces of Stratigraphic significance in two main ways: (1) through
the identification of discontinuities using substrate-controlled ichnofacies (the firmground
Glossifungites ichnofacies, the hardground Trypanites ichnofacies and the woodground
Teredolites ichnofacies); and (2) through careful analysis of trace fossils in vertical (soft-
ground) successions (analogous to facies successions). Integrating the data derived from
substrate-controlled ichnofacies (so-called omission suites) with palaeoecological data
from vertically and laterally juxtaposed softground ichnological successions greatly enhances
the recognition and interpretation of a wide variety of stratigraphically significant surfaces.
When this is coupled with conventional sedimentary facies analysis and sequence stratigra-
phy, a powerful approach to the interpretation of the rock record is generated.

Trace fossil assemblages can be employed effec- MacEachern & Burton 2000; Savrda et al
tively to aid in the recognition of various discon- 2001; Taylor et al 2003) and event stratigraphy
tinuity types, as well as to assist in their genetic (Seilacher 1962, 1982; Vossler & Pemberton
interpretations. Ichnology can be utilized to 1988; Frey & Goldring 1992; Pemberton &
resolve surfaces that may have Stratigraphic MacEachern 1997). Genetic stratigraphy lies at
significance in two main ways: (1) through the the core of three main Stratigraphic paradigms:
identification of discontinuities using substrate- genetic Stratigraphic sequences (Galloway
controlled ichnofacies; and (2) through careful 1989a, 1989b), allostratigraphy (Walker &
analysis of trace fossils in vertical (softground) James 1992), and sequence stratigraphy (Van
successions (accomplished by using either ichno- Wagoner et al 1990). The recognition of
facies or ichnofabric analysis). Stratigraphic breaks is essential in any genetic
Though ichnological analysis is a valuable tool, Stratigraphic paradigm, but also is commonly a
it continues to remain highly under-utilized in difficult task, particularly in subsurface analysis.
most sedimentologically driven genetic strati- Discontinuities may reflect processes that are
graphic studies. Integrating the data derived external to the depositional system (allocyclic),
from substrate-controlled, omission-related which may initiate or terminate deposition of
ichnofacies with palaeoecological data from verti- sedimentologically related facies successions
cal ichnological successions greatly enhances the (Walker 1990). Interpreting the origin of the dis-
ability to recognize and interpret a wide variety continuity, essential to sequence stratigraphy
of Stratigraphic surface types. When this is and to genetic Stratigraphic sequences, is vital
coupled with conventional facies analysis and in resolving the depositional environments of
sequence stratigraphy, a powerful approach to associated deposits and in determining the allo-
the interpretation of the rock record is generated. cyclic controls on the depositional systems. To
Trace fossils have proven to be one of the most accomplish this requires the integration of ichno-
important groups of fossils in assisting in the facies relationships (Pemberton et al 2001) or
delineation of stratigraphically important ichnofabrics analysis (Taylor et al 2003; Mcllroy
boundaries related to genetic stratigraphy (e.g. 2004), physical sedimentology and sequence
MacEachern et al 1991, 1992, 1998, 1999b; Stratigraphic techniques. Ichnofacies and recon-
Savrda 199la, 1991b, 1995; Pemberton et al structed ichnocoenoses are part of the total
1992a, 2001; Taylor & Gawthorpe 1993; Pember- aspect of the rock, imparted by the depositional
ton & MacEachern 1995; Ghibaudo et al 1996; environment, and therefore - like lithofacies -

From: MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 29-62. 0305-8719/04/$ 15.00 © The Geological Society
of London.
30 S. G. PEMBERTON ET AL.

are subject to Walther's law. For example, iso- established (Ekdale et al 1984): Glossifungites
lated bored shells or clasts do not, in themselves, (firmground), Trypanites (hardground) and Ter-
constitute the Trypanites ichnofacies. Rather, edolites (woodground). In clastic settings, most
there must be some semblance of stratification, of these trace assemblages are associated with
lateral continuity and/or vertical succession erosionally exhumed (dewatered and compacted
before an ichnofacies can be applied. This or cemented) substrates, and hence correspond
paper expands upon and updates the work to erosional discontinuities. Depositional
already published in Pemberton et al. (2001). breaks, in particular condensed sections, may
also be semi-lithified or lithified, presumably at
their upper contacts (or downlap surfaces), and
may be colonized without associated erosion.
Substrate-controlled ichnofacies and the In general, however, the recognition of sub-
recognition of stratigraphic discontinuities: strate-controlled ichnofacies may be regarded
the use of omission suites as being equivalent to the recognition of discon-
tinuities in the stratigraphic record. Determining
One of the most important factors in the distri- whether these discontinuities are autocyclically
bution of organisms in modern environments is generated or allocyclically generated, and hence
substrate type. In their recent review Taylor stratigraphically important, is considerably
et al. (2003) summarized the different substrate more problematic.
types (Fig. 1) as: Although certain insect and vertebrate bur-
rows in the terrestrial realm may be properly
soupground (water-saturated mudrocks and regarded as firmground in character (e.g.
micrites); Voorhies 1975; Fursich & Mayr 1981; Smith
softgrounds (muddy and micritic sediment 1987; Groenewald et al 2001) or, more rarely,
with some dewatering); hardground suites, they have a low preservation
loosegrounds (sandy sediments where perma-
potential and are relatively minor in the geologic
nent burrows require stabilized margins); record. The overwhelming majority of these
stiffground (stabilized sediment where bur- assemblages originate in marine or marginal
rows are unlined); marine settings. A discontinuity may be gener-
firmground (firm dewatered, often compacted ated in either subaerial or submarine settings,
sediment); but colonization of the discontinuity is most
hardgrounds (lithified substrate surface; varia- likely to occur in association with marine influ-
tions can be shellground, a cemented shell bed; ence, particularly in pre-Tertiary intervals. This
and rockground, with tectonic omission). circumstance has important implications for the
To this list could also be added woodgrounds interpretation of the discontinuity's genesis.
(xylic substrates) and the closely related log- Substrate-controlled ichnocoenoses typically
grounds (substrate composed of distinct logs) cross-cut the pre-existing softground suites, and
(Bromley et al 1984; Savrda et al 1993). Three hence reflect conditions that post-date both the
substrate-controlled ichnofacies have been initial deposition of the unit and the erosion of

Fig. 1. Relationship of substrate type and the distribution of the named ichnofacies.
KEY STRATIGRAPHIC SURFACES 31

that unit. These omission suites actually characteristics of the ichnofauna are thus deter-
correspond to the period of time between the mined by the nature of the substrate and the
erosional event (which exhumes the substrate) palaeoenvironmental conditions prevailing
and final burial of the discontinuity beyond during the hiatus. By observing (a) the under-
reach of the benthic community of the overlying lying and cross-cut softground trace fossil suite
unit. During such a hiatus infaunal organisms (contemporaneous with deposition of the under-
are free to colonize the substrates: the specific lying unit), (b) the omission suite and ichnofacies

Fig. 2. Schematic development of a Glossifungites demarcated erosional discontinuity based on the Jurassic
Arab D interval in Saudi Arabia. 1: The muddy carbonate substrate is deposited and buried. 2: A transgressive
surface of erosion is generated by a sea-level rise. This exposes previously deposited dewatered sediment to the
sediment-water interface, where it is burrowed by firmground organisms. The burrows are filled with
grainstone that is deposited on the surface as a ravinement deposit. 3: After several rises in sea-level a complex
framework results that is characterized by mappable surfaces that are characterized by a Glossifungites
assemblage.
32 S. G. PEMBERTON ET AL.

associated with the exhumed substrate and (c) moderately low diversity, although the borings
the ichnocoenosis of the overlying unit, it is and scrapings of individual ichnogenera may
possible to make some interpretation regarding be abundant;
the origin of the discontinuity and the allocyclic borings oriented perpendicular to the sub-
or autocyclic mechanisms responsible (Fig. 2). strate that may include large numbers of
overhangs.
In contrast to the Glossifungites ichnofacies, the
The Trypanites ichnofacies walls of the borings cut through hard portions of
the substrate rather than skirting around them.
The Trypanites ichnofacies (Fig. 3) develops in
fully lithified substrates such as hardgrounds,
reefs, rocky coasts, 'beach rock' and other
omission surfaces (Pemberton et al. 1980; The Teredolites ichnofacies
Gruszczyhski 1986, 1998). Development of this
The Teredolites ichnofacies (Fig. 4) consists of a
ichnofacies therefore also corresponds to discon-
characteristic assemblage of borings or burrows
tinuities that have major sequence stratigraphic
in woody or highly carbonaceous substrates.
significance. Bromley & Asgaard (1993) sub-
Woodgrounds differ from lithic substrates in
divided the Trypanites ichnofacies into two
three main ways:
ichnofacies: the Entobia ichnofacies for rocky
shorelines (see also De Gibert et al. 1998), and They may be flexible instead of rigid;
the Gnathichnus ichnofacies for bored shells They are composed of carbonaceous material
and boulders found further offshore. Bored instead of mineral matter;
shells and boulders, however, are not substrates They are readily biodegradable (Bromley et al.
that can be correlated, because it is difficult to 1984).
ascertain when the boring activity was initiated.
Such differences dictate that the means by which,
The traces in the Trypanites ichnofacies are
as well as the reasons for which, these two types
characterized by:
of substrate are penetrated are also different. As
cylindrical to vase, tear- or U-shaped to irregu- currents can raft woody substrates, it is impor-
lar domiciles of suspension feeders or passive tant to determine whether the bored substrates
carnivores; are autochthonous or allochthonous. Only the
raspings and gnawings of algal grazers and autochthonous forms are true members of
similar organisms (mainly chitons, limpets the Teredolites ichnofacies. These assemblages
and echinoids); may also be important in defining sequence and

Fig. 3. Trace fossil association characteristic of the Trypanites ichnofacies.


KEY STRATIGRAPHIC SURFACES 33

Fig. 4. Trace fossil association characteristic of the Teredolites ichnofacies.

parasequence boundaries (Savrda 199la). The Exhumation can occur in terrestrial environ-
use of bored logs to define bounding surfaces, ments, as a result of channel meandering or
as in the concept of log-grounds (Savrda et al. valley incision; and in shallow-water environ-
1993) should be avoided, because it is very diffi- ments, as a result of meandering tidal channels,
cult to ascertain when the logs were bored. tidal scour erosion, erosive shoreface retreat
Such logs are clasts and should not be treated associated with wave ravinement, or as a result
in the same manner as the Teredolites ichnofacies of submarine channels cutting through pre-
sensu strieto. viously deposited sediments. Such exhumed
The Teredolites ichnofacies is characterized surfaces commonly correspond to stratigraphic
by: discontinuities, and the specific characteristics
of their colonization are critical to their sequence
sparse to profuse, club-shaped borings;
stratigraphic interpretation (Saunders & Pem-
boring walls that are generally ornamented
berton 1986; MacEachern et al. 1992, 1998,
with the texture of the host substrate (i.e. tree
1999a, 1999b; Pemberton et al. 1992b, 2001;
ring impressions and other xyloglyphs);
Pemberton & MacEachern 1995; MacEachern
stumpy to elongate subcylindrical excavations
& Burton 2000; Gingras and Pemberton 2000;
in marine or marginal marine settings;
Taylor et al. 2003; Mcllroy 2004).
shallower, sparse to profuse non-clavate exca-
The Glossifungites ichnofacies (Fig. 6) is
vations (isopod borings) in freshwater settings.
characterized by:
vertical, cylindrical, U- or tear-shaped, com-
The Glossifungites ichnofacies monly scratch-marked burrows and sparsely
to densely branching dwelling burrows;
The Glossifungites ichnofacies is environ- protrusive spreite in some burrows that
mentally wide ranging, but develops only in develop mostly through animal growth
firm, unlithified substrates such as dewatered (funnel-shaped Rhizocorallium and Diplocra-
muds or highly compacted sands (Fig. 5). De- terion [formerlyGlossifungites})',
watering results from burial, and the substrates animals that leave the burrow to feed (e.g.
are made available to trace-makers if exhumed crabs) as well as suspension feeders;
by later erosion (e.g. Pemberton & Frey 1985). low diversity, but commonly high abundance.
34 S. G. PEMBERTON ET AL.

MacEachern & Burton (2000) and Savrda et al


(2001) have shown omission-related firmground
Zoophycos associated with discontinuities
colonized in very distal settings of the shelf and
slope. Caution should be exercised in distinguish-
ing firmground assemblages from stiffground
assemblages (Wetzel & Uchman 1998) that can
be localized and may be related to deep tiers.
In proximal, high-energy settings, the assem-
blages attributable to the Glossifungites ichno-
facies are dominated by vertical domichnia. The
presence of vertical shafts within shaly intervals
is anomalous, as these structures are not capable
of being maintained in soft muddy substrates.
Glossifungites ichnofacies elements are typically
robust, commonly penetrating 20-100 cm below
the bed junction. Many shafts tend to be large
in diameter (e.g. 0.5-1.Ocm), in particular Diplo-
craterion habichi and Arenicolites. This scale of
burrowing contrasts markedly with the predomi-
nantly horizontal and diminutive ichnogenera
that typify the exhumed shaly intervals. The firm-
ground traces are generally sharp-walled and
unlined, reflecting the stable, cohesive nature of
the substrate at the time of colonization and
burrow excavation. Further evidence of substrate
resilience, atypical of soft muddy beds, is the
passive nature of most burrow fills. This demon-
strates that the structure remained open after the
trace-maker vacated the domicile, thus allowing
material from subsequent depositional events to
infiltrate the open tube. The post-depositional
origin of the Glossifungites ichnofacies is clearly
demonstrated by the ubiquitous cross-cutting
Fig. 5. The Glossifungites ichnofacies is relationships with the previous softground
environmentally wide ranging, but develops only in assemblage. The final characteristic of the suite
firm, unlithified substrates. (A) Develops in dewatered is the tendency to reflect colonization in large
exhumed muds, Albian Viking Formation, Willesden numbers (Fig. 7). In numerous examples, 7-15
Green Field, Alberta, 10-35-40-7W5, 2327m or (B) firmground traces, most commonly Diplocrater-
highly compacted sands. (B) Develops in compacted ion habichi, have been observed on the bedding
sandstone where an inclined Thalassinoides (T) cross-
cuts a lined Ophiomorpha (O) in the Price River A plane of 9 cm (3.5 in) diameter cores, correspond-
core. (C) Gallup Sandstone, San Juan Basin, New ing to a density of between 1100 and 2300 shafts
Mexico with multiple incision surfaces seen in the top per m2. Similar populations have been observed
metre of the unit. from the modern coast of Germany (Schafer
1972), the modern Georgia coast (Basan & Frey
Firmground traces are dominated by vertical 1977; Morris & Rollins 1977; Pemberton &
to subvertical dwelling structures of suspension- Frey 1985), and Willapa Bay (Gingras et al.
feeding organisms (Fig. 7). The most common 1999).
structures correspond to the ichnogenera Diplo-
craterion, Skolithos, Psilonichnus, Arenicolites
and unnamed flask-shaped domichnia compar- Selected case studies of ichnological
able to Gastrochaenolites (Fig. 8). Dwelling applications to sequence stratigraphy
structures of deposit-feeding organisms are also
constituents of the ichnofacies, particularly Regressive surfaces of erosion (RSE) and
where exhumed substrates occupied more sequence boundaries (SB)
sheltered or distal settings during colonization,
and include firmground Thalassinoides, Sponge- Although subaerial exposure and/or erosion
liomorpha and Rhizocorallium. More recently, during relative sea-level lowstand may produce
KEY STRATIGRAPHIC SURFACES 35

Fig. 6. Trace fossil association characteristic of the Glossifungites ichnofacies.

Fig. 7. Characteristics of trace fossils that are associated with the Glossifungites ichnofacies: (A) the burrows
tend to be robust, unlined domiciles; (B) are found in very high densities; (C) commonly display scratch marks;
and (D) cross-cut the original softground trace fossil assemblage.
36 S. G. PEMBERTON ET AL.

Fig. 8. Glossifungites assemblages: (A) Skolithos from the sequence boundary at the base of an interpreted
forced regression, Kaybob Field, Viking Formation, 11-35-61-20W5, depth 1759.1m; (B) Glossifungites suite of
conglomerate-filled Thalassinoides, Cardium Formation, Pembina Field, 12-9-51-10W5, depth 1596.2m;
(C) firmground Arenicolites marking a transgressive surface of erosion (TSE), Cretaceous Viking Formation,
07-19-62-19W5, 1652m; (D) Thalassinoides at surface in the Lower Cretaceous Dun vegan Formation, Jayar
Field, 6-11-62-3W6, 2523.6m; (E) Diplocraterion at transgressive surface in the Upper Cretaceous Horeseshoe
Canyon Formation in outcrops near East Coulee, Alberta; (F) Glossifungites ichnofacies consisting of
Rhizocorallium excavated into offshore shales and cross-cutting a resident softground suite of Helminthopsis,
Planolites, Schaubcylindrichnus, Chondrites and Zoophycos. Albian Viking Formation, Willesden Green Field,
Alberta, 10-35-40-7W5, 2327m.
KEY STRATIGRAPHIC SURFACES 37

widespread development of dewatered, firm or Hayward (1976) interpreted the erosional discon-
cemented substrates (corresponding to regressive tinuity as a submarine canyon wall, excavated
surfaces of erosion and sequence boundaries), into bathyal to neritic inter-arc sediment gravity
most are unlikely to become colonized by sub- flow deposits as a result of basin margin tectonic
strate-controlled trace fossil suites unless the uplift. Colonization of the canyon walls by the
surfaces are subsequently exposed to marine or firmground trace-makers preceded the gradual
marginal marine conditions prior to burial. In burial of the canyon margins by neritic turbidite
most cases, deposition of significant thicknesses deposits of the Tirikohua Formation. The infill
of non-marine strata generally precludes devel- of the submarine canyon probably corresponds
opment of these omission suites on the RSE or to late stage relative sea-level lowstand and
SB themselves. There are a number of scenarios, early transgression.
however, where such discontinuities may be In the subsurface, examples of submarine
preferentially colonized by trace-makers of canyon incision with the development of trace
substrate-controlled assemblages. Such settings fossil assemblages of the Glossifungites ichno-
include RSE developed beneath forced regressive facies have been recognized in the Miocene of
shorefaces, SB underlying lowstand shorefaces, the Nile Delta (Fig. 9). The canyon walls were
SB comprising submarine canyon margins, and excavated during lowstand incision and colo-
SB lying at the estuarine mouths of incised nized by shrimps that constructed robust Thalas-
valleys, prior to transgressive infill. All of these sinoides. The interpretation of the surface is
settings are conducive to colonization of the dis- critical to correct correlation of the canyon fill
continuity because the surfaces were excavated and to the recognition of point source turbidites.
subaqueously in a marine or marginal marine Fine-grained facies outside the canyons are
environment. The marginal marine component totally bioturbated by Phycosiphon, Planolites
of the sequence boundary within an incised and Helminthoida. Similar facies within the
valley is aerially restricted and difficult to discern canyon system reflect episodic mud turbidites
from the transgressively modified sequence and remain virtually unburrowed.
boundary during initial transgression. For that
reason, this latter scenario is discussed in the Forced regressive and lowstand incised shorefaces
context of amalgamated sequence boundaries Forced regressive and lowstand shorefaces con-
and flooding surfaces (FS/SB) of incised valleys. stitute two sequence stratigraphic scenarios by
which sharp-based shoreline sandstones may
Incised submarine canyons form, and both are associated with falling limbs
There are few published ichnological assessments of relative sea-level. Sharp-based shoreface
of an ancient submarine canyon margin. sand bodies, however, have been variably
Outcrops of the lower Miocene Nihotupu and assigned to the progradation of late highstand
Tirikohua formations in Northland, New successions (e.g. Van Wagoner 1995), forced
Zealand, contain a noteworthy firmground regressive (falling stage) systems (e.g. Hunt &
trace fossil assemblage of the Glossifungites Tucker 1992; Walker & Bergman 1993; Bergman
ichnofacies related to submarine canyon incision 1994; Davies & Walker 1993), lowstand systems
(Hayward 1976). The underlying Nihotupu (e.g. Flint et al 1988; Posamentier et al 1992;
Formation consists of volcanogenically derived Posamentier & Chamberlain 1993; Mellere &
siltstones, sandstones and subaqueous mass Steel 1995; Walker & Wiseman 1995), and trans-
flow conglomerates, together with submarine gressively incised complexes (e.g. Downing &
andesite pillow-pile complexes. The underlying Walker 1988; Raychaudhuri et al 1992; Mac-
softground assemblage is sparse and sporadically Eachern et al 1998, 1999b). Despite the wide
distributed, characterized by localized individual range of sequence stratigraphic contexts that
occurrences of Thalassinoides, Planolites and facilitate such deposits, many workers continue
Scalarituba. These deposits are interpreted as to regard sharp-based shoreface sandstone
turbidites that were emplaced at bathyal water bodies to be exclusively of falling stage or low-
depths (based on body fossil content) within an stand origin.
inter-arc basin on the lower eastern flanks of From a facies perspective, however, the sharp-
the west Northland volcanic arc. The contact based shoreface successions generated in all three
with the overlying Tirikohua Formation is systems tracts are virtually identical. Their prin-
sharp and erosional and exhibits visible relief. cipal difference lies in the character of the basal
The exhumed substrate is demarcated by a contact with the underlying facies. One distinc-
firmground omission assemblage consisting of tion, however, is that highstand examples overlie
Skolithos, Rhizocorallium and ?Thalassinoides autocyclic basal surfaces that lack evidence of
attributable to the Glossifungites ichnofacies. incision into and concomitant truncation of
38 S. G. PEMBERTON ET AL

Fig. 9. Glossifungites assemblage associated with submarine canyon incision, West Ahken Field, Nile Delta:
(A) Skolithos filled with anomalous sediment, West Ahken-1 core, 4379.8ft; (B) sequence boundary at base of
submarine canyon fill characterized by a Glossifungites assemblage with Thalassinoides, West Ahken-1 core,
4375.3ft.

regional markers in the underlying succession. of relative sea-level. Likewise, the lowstand
Such sharp-based but non-incised highstand shoreline overlies the marine part of the sequence
shoreface deposits also display a genetic affinity boundary (SB) and is also cut by wave erosion.
of facies across the autocyclic basal surface that The transgressively incised shoreface, however,
commonly corresponds to the erosional bases overlies a wave ravinement surface, cut during
of individual storm beds. In contrast, forced relative sea-level rise. The wave ravinement sur-
regressive, lowstand and transgressive shorefaces face commonly amalgamates with or truncates
overlie allocyclic discontinuities that truncate the earlier sequence boundary (FS/SB). This
regional markers, reflecting incision into under- succession reflects a period of shoreline progra-
lying units. dation during overall transgression, when sedi-
Incised-shoreface deposits may correspond to ment supply outpaced relative rise of sea-level.
forced regressive, lowstand or transgressively All three scenarios favour the development of
incised systems (Fig. 10). The forced-regressive substrate-controlled omission suites on the
shoreface overlies a regressive surface of erosion basal discontinuities, as each is excavated
(RSE) cut by wave action during the falling stage within a marine setting, and permit early

Fig. 10. Differentiation of forced-regressive, lowstand, and transgressively incised shoreface complexes. Sharp-
based, discontinuity-bound (incised) shoreface successions can be ascribed to one of three main sequence
stratigraphic settings. Model 1 reflects forced regression (falling stage), showing the initial fall of relative
sea-level and the development of successive shorefaces sitting on regressive surfaces of erosion (RSE). Note that
although a correlative conformity (CC) may be produced seaward of each RSE, successive sea-level fall makes
these susceptible to erosional removal, and therefore they have a low preservation potential. Model 2 shows the
development of the lowstand shoreface, which reflects the most seaward position of the shoreline associated
with the lowest position of sea-level. Note that the erosional component of the sequence boundary extends only
as far seaward as fair-weather wavebase (FWWB), where it passes into a correlative conformity (CC). In model
3, rise of relative sea-level generates a low-energy flooding surface in basinal positions that passes landward
into a transgressive ravinement surface, as it floods across the lowstand and forced-regressive shorefaces. Where
the surface cuts across or incises through the old sequence boundary, it produces an FS/SB. Note that the rise
of sea-level drowns and preserves the CC of the lowstand shoreface. During a pause in transgression, shoreface
progradation occurs, producing a transgressively incised shoreface. In basinward positions, offshore mudstones
deposited below fair-weather wavebase may directly overlie the erosional component of the FS/SB because the
surface was cut while sea-level occupied a lower position but deposition did not occur until after significant
deepening. (Modified from MacEachern et al. 1998)
40 S. G. PEMBERTON ET AL.

colonization of the exhumed substrate. Differen- sediment supply, rate of change of accommoda-
tiation between these incised complexes is diffi- tion space, basin gradient and duration of
cult, but can be achieved through careful shoreline progradation, caution must clearly be
documentation of the erosional extent of the exercised.
basal discontinuity (MacEachern et al. 1999b). The perceived regional stratigraphic context of
Considerable discussion surrounds the validity the sharp-based shoreface and/or delta deposits
of differentiating lowstand from forced regressive has principally been used as a basis for their
deposits (e.g. Hunt & Tucker 1992, 1995; Kolla interpretation. Ainsworth & Pattison (1994)
et al. 1995). The work of Helland-Hansen & have discussed the problem of attached versus
Gjelberg (1994) and Mellere & Steel (1995), how- detached lowstand complexes, highlighting
ever, illustrates the utility of discriminating fall- some of the difficulties in differentiating between
ing-stage systems tracts associated with forced the two scenarios. Falling-stage (forced regres-
regression from the final lowstand shoreline sive) interpretations are most commonly based
corresponding to maximum fall of sea-level, but on the presence of additional incised shoreface
prior to transgression. A forced-regressive or and/or delta deposits lying basinward of them
falling-stage origin has been proposed for within the same sequence. Lowstand deposits,
sharp-based sandstone bodies of the Viking on the other hand, tend to be identified mainly
Formation in the Garrington Field (Davies & on the basis of the absence of additional basin-
Walker 1993) and Kaybob Field (Pemberton & ward shorefaces. Given that such deposits may
MacEachern 1995). Posamentier et al. (1992) be detached and lying considerably basinward,
and Posamentier & Chamberlain (1993) inter- the sequence stratigraphic interpretation of
preted sharp-based sandstone deposits at these intervals may, in some cases, be highly
Joarcam to reflect a lowstand shoreface deposit. suspect. Walker & Wiseman (1995) concede this
Lowstand shoreface deposits have also been uncertainty with respect to the Viking Formation
interpreted in the Lindbrook and Beaverhill Lindbrook shorefaces of Alberta. MacEachern
Lake fields (Walker & Wiseman 1995), although et al. (1999b) have argued, however, that there
their figure 6 suggests that the 'Lindbrook a' are distinctive facies relationships that can also
deposit probably reflects a falling-stage shore- be employed in order to differentiate the various
face, given that the 'Lindbrook b' shoreface lies sequence stratigraphic scenarios.
farther basinward and likewise overlies a Forced-regressive shoreface and/or deltaic
regressive surface of erosion. The Judy Creek deposits overlie regressive surfaces of erosion
Field, which lies along strike to these deposits, (RSE). These surfaces are cut in submarine con-
contains a lowstand incised shoreface deposit as ditions and pass basinward into conformable
well. surfaces analogous to correlative conformities
The differentiation between sharp-based, (Fig. 10). The RSE are cut by wave erosion as
incised shorefaces and deltas of forced regressive relative sea-level falls, bringing more basinal
versus lowstand origin, however, is problematic. facies into the zone of wave attack. Continued
Both forced regressive and lowstand shoreface sea-level fall results in the subaerial exposure of
deposits tend to be fairly thin, in response to the falling-stage shorefaces, and their subsequent
the diminished accommodation space associated cannibalization by later regressive surfaces of
with relative lowstand of sea-level (Flint 1988; erosion and, ultimately, the sequence boundary.
Posamentier et al. 1992; Van Wagoner 1995). The preservation potential of these deposits is
Walker & Wiseman (1995) have further sug- considerably less than that of the lowstand shore-
gested that the damping of wave energy across face, and the correlative conformities of the RSE
broad shallow platforms lying outboard of are, in particular, unlikely to be preserved in the
these shorefaces contributes to this, because it rock record. Kolla et al. (1995) regard the RSE
inhibits incision into the underlying firmly com- that bound falling-stage deposits merely as
pacted mud. As a result, facies tracts within higher-order sequence boundaries reflecting
these shoreface types may be attenuated or incremental rather than continuous fall of
even absent. Lowstand shorefaces may be relative sea-level.
slightly thicker because they may be developed Lowstand shorefaces directly overlie sequence
during late lowstand, where a slow rise in relative boundaries and, basinward, their correlative
sea-level may be initiated with associated conformities (Plint et al. 1988; Posamentier
increased accommodation space. Sandbody et al. 1992). Landward of the shoreface, the
widths and width/thickness ratios have also sequence boundaries are cut by subaerial ero-
received some consideration as a means of dis- sion. At the base of the shoreface, however,
criminating between these systems, but because sequence boundaries are cut within a marine
of the effects of such controls as variations in setting and therefore favour colonization by
KEY STRATIGRAPHIC SURFACES 41

substrate-controlled assemblages, particularly shoreface deposits. The forced-regressive depos-


where they incise into dewatered offshore muds. its are subjected to subsequent erosion and sub-
As the lowstand shoreface lies in the most aerial exposure during continued fall of relative
seaward position prior to ensuing sea-level rise, sea-level, as well as the potential of transgressive
the marine expression of the sequence boundary ravinement during ensuing rise of relative sea-
and the correlative conformity have a high pre- level (Fig. 10). The lowstand deposits, on the
servation potential, facilitating their differentia- other hand, are produced at the lowest position
tion from forced regressive and transgressively of relative sea-level fall and are unlikely to be
incised counterparts (MacEachern et al. 1999b). subsequently ravined, because water depths are
In weakly storm-influenced settings, the ero- deepened and fair-weather wavebase shifted
sional component of the RSE and sequence landward during later transgression. The
boundaries is unlikely to persist basinward of presence of a correlative conformity might be
fair-weather wavebase and therefore defines a taken as a significant support for a lowstand
sharp base to the lower shoreface. Basinward of interpretation of a deposit.
this position, finer-grained offshore deposits The subsurface Viking Formation of the
overlie the correlative conformity and during Kaybob and Judy Creek fields highlights the
progradation grade upwards into lower shore- similarities between a forced regressive shoreface
face muddy sandstones. As a result, the Glossi- succession and a lowstand one. The Kaybob
fungites ichnofacies and other omission suites Field of central Alberta contains a sharp-based,
are unlikely to occur in positions below fair- incised shoreface excavated into underlying
weather wavebase. In these basinal positions, open marine distal parasequences (Pemberton
coarse-grained lag deposits are likely to be & MacEachern 1995; MacEachern & Pemberton
absent as well. The correlative conformity may, 1997; Fig. 11). The underlying distal para-
however, represent a sharp but depositional sequences consist of moderately to abundantly
facies contact, marked by an abrupt change in bioturbated (BI4-5) mudstones, silty mudstones
proximality of facies, grain size and trace-fossil and sandy siltstones, displaying diverse trace
assemblage. In storm-dominated shorefaces, fossil assemblages attributable to the Zoophycos
however, the extent of the allocyclically gener- ichnofacies and distal and archetypal expressions
ated marine expression of the RSE or sequence of the Cruziana ichnofacies, respectively. The
boundary may be masked by autocyclic storm parasequences reflect progradation of shelf to
erosion surfaces and appear to extend to storm- upper offshore cycles of an underlying highstand
weather wavebase. This scenario results in the systems tract. The erosional discontinuity is
development of a series of vertically stacked principally demarcated by firmground omission
and offlapping, aerially restricted autocyclic sur- suites of Skolithos, Diplocraterion and Thalassi-
faces, rather than a single allocyclic surface, but noides of the Glossifungites ichnofacies (Fig.
recognition of this condition may be problematic 11C, D). Where the discontinuity is excavated
unless outcrop exposure is exceptional. In the into sandier expressions of the underlying
subsurface, recognition of this situation would facies, the surface is demarcated by a palimpsest
be extremely difficult. However, these autocyclic softground suite of Diplocraterion habichi and
surfaces are rapidly buried by tempestites and are Skolithos (Fig. 11 A, B). In all cored intervals
therefore not readily colonized by trace-makers where the discontinuity is preserved, it is ero-
of substrate-controlled ichnofacies. sional and is overlain by silty to muddy sand-
Both forced regressive and lowstand com- stones, interpreted to reflect lower shoreface
plexes are typically sharp based in proximal and middle shoreface deposits (Fig. 11B, D).
positions and gradationally based in basinal The sandstones are moderately and sporadically
positions. Consequently, only the lower shore- bioturbated (BI 2-3), characterized by abundant,
face, middle shoreface and upper shoreface remnant hummocky cross-stratified beds sepa-
deposits directly overlie the erosional expression rated by burrowed beds. The succession shows
of the sequence boundary or the RSE and may an upward decrease in the number and thickness
be demarcated by the Glossifungites ichnofacies. of burrowed beds and a concomitant increase in
Landward, a ravinement surface may become the number and thickness of hummocky cross-
amalgamated with the sequence boundary and stratified beds, consistent with upward shallow-
RSE during the ensuing transgression (e.g. Flint ing (Fig. 11 A). A fully marine, diverse, proximal
et al. 1986; Flint 1988; Pemberton & Mac- expression of the Cruziana ichnofacies domi-
Eachern 1995). It appears reasonable that the nates the more thoroughly burrowed successions
correlative conformity of the RSE has an exceed- and passes upwards into suites consistent with
ingly low preservation potential in falling-stage the Skolithos ichnofacies. Locally, trough cross-
deposits, in contrast to that of lowstand- bedded sandstones containing sporadically
42 S. G. PEMBERTON ET AL.

Fig. 11. Forced regressive shoreface. (A) Box shot of core from the Kaybob incised shoreface; base of the
interval is to the lower left, and top to the upper right (T). The lower unit consists of bioturbated (BI5—6) silty
and sandy mudstones, and (in column 4) muddy sandstones of the underlying regional Viking Formation,
reflecting progradation of lower offshore, upper offshore and lower shoreface environments respectively.
These are truncated by a regressive surface of erosion (RSE), and overlain by coarser-grained lower and
middle shoreface laminated to bioturbated sandstones of a moderately storm-influenced shoreface.
Well 11-35-61-20W5; 1757.6-1762.1 m. (B) Close-up photo of the RSE in photo A, showing a palimpsest
softground suite consisting of Skolithos (S) demarcating the stratigraphic discontinuity. The incised shoreface in
this locality has cut into bioturbated (BI 5) muddy sandstones with softground Ophiomorpha (O), and
Palaeophycus (Pa), reflecting a proximal expression of the Cruziana ichnofacies. The overlying lower shoreface
sandstones of the forced regressive shoreface contain sideritized mudstone rip-up clasts and Palaeophycus (Pa).
Well 11-35-61-20W5, 1759.2m. (C) Box shot of core from the Kaybob incised shoreface in a position more
distal than that of photo A. Base of the interval is to the lower left, and top to the upper right (T). The lower
unit consists of bioturbated (BI 5-6) silty and sandy mudstones of the underlying regional Viking Formation,
reflecting progradation of lower to upper offshore environments. These mudstones are truncated by an RSE,
and overlain by lower shoreface muddy sandstones, passing into middle shoreface sandstones. Well 10-15-62-
19W5; 1667.3-1673.0m. (D) Close-up photo of the RSE in photo C, showing nrmground Thalassinoides (Th) of
the Glossifungites ichnofacies demarcating the discontinuity. The discontinuity in this position is incised into
bioturbated (BI 6) sandy mudstones containing the archetypal Cruziana ichnofacies with Phycosiphon (Ph) and
Chondrites (Ch). The overlying lower shoreface muddy sandstone contains a proximal expression of the
Cruziana ichnofacies with Diplocraterion (D) and Palaeophycus (Pa). Well 10-15-62-19W5, 1671.8m.
KEY STRATIGRAPHIC SURFACES 43

distributed assemblages of the Skolithos ichno- The middle shoreface silty sandstones, however,
facies may directly overlie the sequence bound- contain trace fossils recording a distal expression
ary or grade upward from the middle shoreface of the Skolithos ichnofacies.
sandstones, and are interpreted as upper The Judy Creek incised shoreface deposits
shoreface deposits. The upper contact of the clean and coarsen upward, and erosionally over-
succession is truncated and, locally, cemented lie the distal counterparts of the same shelf to
with haematite-stained siderite, interpreted to offshore parasequences truncated by the land-
reflect subaerial exposure. ward-lying Kaybob forced regressive shoreface.
In basinward positions, where one might The erosional discontinuity is locally demarcated
expect to find the offshore sandy mudstones by well-developed firmground Thalassinoides
and siltstones equivalent to the Kaybob lower and less commonly by Spongeliomorpha, attribu-
shoreface sandstones, the interval has been table to the Glossifungites ichnofacies. In all
removed by later cycles of incision. The basal cored intervals containing an erosional expres-
discontinuity is not preserved seaward of fair- sion of the discontinuity, the overlying facies
weather wavebase, making sequence strati- comprise bioturbated muddy sandstones or
graphic interpretation of the surface and silty sandstones, interpreted to reflect deposition
therefore of the overlying deposit problematic. above fair-weather wavebase (Fig. 12A, B). In
The succession is consistent with an incised the few instances, however, where the Judy
shoreface, but whether forced regressive, low- Creek deposit is initiated by upper offshore
stand or transgressively incised cannot be sandy mudstones, the bounding surface shows
unequivocally demonstrated on the basis of the no evidence of erosional truncation of the
deposits themselves. The presence of additional underlying parasequences (Fig. 12C, D). This is
shorefaces lying basinward of the Kaybob significant, as it records the character of the
deposit strongly suggests that the deposit bounding surface in positions below fair-weather
cannot reflect the lowstand shoreface of the wavebase. This non-erosional surface shows an
sequence (Fig. 10). Furthermore, preferential increase in grain size associated with prograda-
removal of the discontinuity in a seaward posi- tion of the Judy Creek incised shoreface, but
tion would be difficult to accomplish during lying seaward of erosional modification by
transgression, and appears inconsistent with waves. The surface in this position is interpreted
observed relationships in transgressively incised as the preserved correlative conformity of the
examples (e.g. Downing & Walker 1988; Ray- discontinuity lying landward beneath the incised
chaudhuri et al. 1992; MacEachern et al 1998, shoreface sensu stricto (Fig. 12D). In even more
1999b). Additionally, the iron-stained, siderite distal positions the correlative conformity is dif-
cemented sandstone at the upper truncated ficult to identify, and the succession appears to
margin of the interval is consistent with contin- reflect simple progradation. The correlative
ued relative sea-level fall, exposure and subaerial conformity survived in this position because,
erosion of a forced regressive shoreface. As such, after this cycle of progradation, relative sea-
the basal discontinuity is interpreted to reflect a level rose again, thereby flooding the area and
regressive surface of erosion (RSE). causing erosional back-stepping of the shoreline.
By comparison, further basinward of the This is consistent with a lowstand shoreface
Kaybob forced regressive shoreface lies an interpretation, and indicates that the underlying
incised shoreface sandstone in the Judy Creek erosional discontinuity corresponds to the
Field. In contrast to the more storm-influenced sequence boundary, rather than the RSE of a
successions of the Kaybob deposit (MacEachern forced regressive shoreface (Fig. 10).
& Pemberton 1992), the Judy Creek deposit is
weakly storm affected and is characterized by
thoroughly bioturbated (BI5), pebble- and gran- Transgressive surfaces
ule-bearing sandy mudstones, muddy sandstones
and silty sandstones of the upper offshore, lower Transgressive surfaces are manifest by (1) mainly
shoreface and middle shoreface respectively (Fig. non-erosional marine flooding surfaces (MFS)
12). Bioturbation within these facies is generally and bay margin flooding surfaces, and (2) low-
uniformly distributed. All facies contain highly relief, erosional (ravinement) surfaces. The
diverse trace fossil assemblages. The upper off- ravinement surfaces may be produced by either
shore sandy mudstones display suites consistent wave or tidal scour processes, and are referred
with the archetypal Cruziana ichnofacies. The to as transgressive surfaces of erosion (TSE).
lower shoreface muddy sandstones contain Analysis of transgressive surfaces of erosion has
trace fossil assemblages corresponding to proxi- had a relatively long history since Stamp (1921)
mal expressions of the Cruziana ichnofacies. originally defined the term 'ravinement'. A
44 S. G. PEMBERTON ET AL.

Fig. 12. Lowstand shoreface. (A) Box shot of core from the Viking Formation incised shoreface at Judy Creek.
Base of the interval is to the lower left, and top to the upper right (T). In this proximal position, bioturbated
silty mudstones of the regional Viking parasequences, interpreted as lower offshore deposits, are incised into by
moderately to intensely bioturbated (BI4—5) silty sandstones of the Judy Creek lowstand shoreface. The
discontinuity is interpreted as a sequence boundary (SB). Well 02/10-19-63-11W5; 1423.4-1427.1 m. (B) Box
shot of core from the Viking Formation incised shoreface at Judy Creek, slightly distal of that in photo A.
Base of the interval is to the lower left, and top to the upper right (T). The interval shows storm-influenced
lower offshore silty mudstones with rare tempestites (lower offshore) incised into by bioturbated (BI 5) muddy
sandstones of the Judy Creek lowstand shoreface. The discontinuity (SB) corresponds to the same sequence
boundary in photo A. The Judy Creek shoreface sandstones in this position are muddier than those of photo
A, and reflect lower shoreface deposition. Well 12-26-63-11W5; 1453.3-1457.6m. (C) Box shot of core from the
Viking Formation incised shoreface at Judy Creek, distal of the position indicated in photo B. Base of the
interval is to the lower right (B), and top to the upper left (T). The succession shows shelf through upper
offshore mudstones at the base, sharply overlain by coarser-grained upper offshore sandy mudstones and
muddy sandstones of the Judy Creek lowstand shoreface. The sharp contact corresponds to the correlative
conformity (CC) of the sequence boundary present in photos A and B. Well 10-35-64-13W5; 1482.3-1486.7 m.
(D) Close-up of the correlative conformity (CC) in photo C, separating a finer-grained sandy mudstone facies
below from a pebble bearing (Pe), coarser-grained sandy mudstone above. Both facies contain an open-marine,
archetypal Cruziana assemblage with Helminthopsis (H), Terebellina (T), Planolites (P), Phycosiphon (Ph), and
Asterosoma (As), interpreted to reflect upper-offshore environments. Well 10-35-64-13W5; 1483.0m.

number of landmark papers have discussed the pre-existing topography and shoreface depth on
characteristics and implications of ravinement the preservation potential of coastal-plain depos-
surfaces, particularly with respect to their pro- its, surface diachroneity and associated facies
cesses of formation, their depths of incision, the (e.g. Fischer 1961; Swift 1968; Belknap & Kraft
interplay of rate of relative sea-level rise with 1981; Pilkey et al 1981; Nummendal & Swift
KEY STRATIGRAPHIC SURFACES 45

1987). MacEachern et al. (1992) discussed the sandy mudstones (Fig. 13B). Such contacts may
ichnological suites associated with ravinement appear gradational, owing to the biogenic homo-
surfaces and their associated facies. genization of the surface by the more robust and
penetrative trace-makers common in these
Marine flooding surfaces settings. Elsewhere, the upward transition from
Marine flooding surfaces (MFS) are typically shallow to deeper water deposits may occur
abrupt contacts across which there is evidence over intervals of several decimetres or more,
of an increase in water depth. These surfaces reflecting very gradual relative sea-level rise.
are mantled with dispersed sand, granules or Similar stacking of such coarsening upward,
intraformationally derived rip-up clasts, indicat- but thoroughly bioturbated, parasequences sepa-
ing some erosion. The preservation of underlying rated by pronounced marine flooding surfaces
markers indicates, however, that the degree of occurs in the Early Permian of the Sydney
erosion is minimal. MFS are typically character- Basin (Bann 1998; Bann et al. 2004), the Jurassic
ized by the abrupt juxtaposition of offshore, shelf Heather Formation of the Norwegian North Sea
or prodelta shales onto shallow marine sand- (MacEachern & L0seth 2003), and the Turonian
stones, and are easily identified on geophysical Cardium Formation of Alberta (Vossler & Pem-
well logs. Such surfaces may demarcate para- berton 1988, 1989).
sequence boundaries, parasequence sets or even
systems tracts, depending upon their regional Transgressive surfaces of erosion
extent (Bhattacharya 1993). Transgressive surfaces of erosion (i.e. ravinement
The Lower Cretaceous Viking Formation in surfaces) afford the most elegant manner of gen-
western Canada contains numerous MFS separ- erating widespread substrate-controlled trace
ating coarsening-upward, regionally extensive fossil assemblages and palimpsest softground
parasequences. These parasequences are inter- suites, because the exhumed surfaces are both
preted to reflect shelf through distal lower shore- widespread and produced within a marine or
face progradation under fully marine conditions. marginal marine environment. This favours colo-
Three facies comprise a complete coarsening nization by organisms as the ravinement surface
cycle, although the minor cycles rarely comprise is excavated prior to accumulation of significant
a complete cycle. The basal facies consists of thicknesses of overlying sediment (MacEachern
intensely bioturbated (BI5) silty mudstone. et al. 1992a,b; Pemberton & MacEachern
Trace fossils are uniformly distributed and 1995). The upper portion of the Albian Viking
diverse (eight ichnogenera), constituting the Formation in the subsurface of central Alberta
archetypal Zoophycos ichnofacies to a distal contains numerous transgressive surfaces of
expression of the Cruziana ichnofacies. Biotur- erosion (TSE), recording a complex history of
bated sandy mudstone facies grade upward transgression, which culminated in maximum
from the silty mudstones and are intensely bur- flooding of the Western Interior Seaway. Bann
rowed (BI5) with a uniformly distributed and (1998) and Bann et al. (2004) have assessed
highly diverse suite (18-21 ichnogenera) of the the ichnological characteristics of TSE from the
archetypal Cruziana ichnofacies. Muddy sand- Early Permian Pebbley Beach Formation of the
stone facies grade upward from the sandy shale Sydney Basin and found comparable characteris-
facies and are intensely bioturbated (BI5) with tics to those of the Mesozoic successions of the
a diverse (18 ichnogenera) and uniformly distrib- Western Interior Seaway.
uted, proximal expression of the Cruziana ichno- The recognition of discrete TSE is difficult on
facies. The cycles reflect coarsening upward of the basis of sedimentology alone, particularly
facies associated with shoaling, under fully when dealing with the upper Viking Formation,
marine conditions, developed during a highstand where there exist abundant, sharp-based pebble
systems tract. stringers and thin, trough cross-stratified,
The marine flooding surfaces (MFS) in the coarse-grained sandstones, intercalated with
major cycles are commonly marked by the interbedded sandstones, siltstones and shales. A
return to lower offshore or shelf deposition, few of these coarse stringers could reflect the
and are typically abrupt (Fig. 13A). These flood- veneer on transgressive ravinement surfaces,
ing surfaces are rarely significantly disrupted by but owing to their abundance it is difficult to
the diminutive trace-makers that characterize pick those that have regional stratigraphic signif-
the lower offshore and shelf settings. In other icance. Similar complexities have been encoun-
cases, cycles may show considerable biogenic tered in the Early Permian Pebbly Beach
modification of the MFS or transgressive surface Formation of the Sydney Basin, Australia
of erosion (TSE), particularly where lower shore- (Bann 1998), the Middle Jurassic Oseberg
face deposits are overlain by upper offshore Formation (Soegaard & MacEachern 2003),
46 S. G. PEMBERTON ET AL.

Fig. 13. Marine flooding surface/transgressive surface of erosion. (A) Non-erosional marine flooding surface
(MFS) separating upper offshore sandy mudstones below from lower offshore/shelf silty mudstones above.
The sandy mudstones contain the archetypal Cruziana ichnofacies, with Palaeophycus (Pa), Diplocraterion (D),
Asterosoma (As), Planolites (P), Phycosiphon (Ph), Teichichnus (Te) and Rosselia (Ro). The overlying silty
mudstones contain a distal expression of Cruziana ichnofacies and show Phycosiphon (Ph) and Teichichnus (Te)
Well 12-17-39-27W4; 1604.6m. (B) Bioturbated transgressive contact with palimpsest softground Thalassinoides
(Th), subtending into muddy sandstones of the lower shoreface, abruptly overlain by gritty, pebble (Pe) bearing
sandy mudstones and silty mudstones with thin tempestites. The contact probably reflects a TSE with only
minor erosion. The underlying muddy sandstones contain a proximal suite of the Cruziana ichnofacies with
Skolithos (S), Asterosoma (As), Planolites (P), and Phycosiphon (Ph). The overlying mudstones display the
KEY STRATIGRAPHIC SURFACES 47

and the Tarbert and Heather formations habichi), Skolithos, Arenicolites, Rhizocorallium
(MacEachern & L0seth 2003) of the Norwegian and Thalassinoides (Fig. 13D, E), attributable
North Sea. to the Glossifungites ichnofacies. In some
However, in each of these units, virtually every Viking Formation TSE, firmground Zoophycos
TSE incised into, or ravined across, shaly sedi- with associated Thalassinoides and Rhizocoral-
ments exhibits an omission suite attributable to lium have also been identified (Fig. 13F), where
the Glossifungites ichnofacies (Fig. 13D, E F) initial colonization of the discontinuity occurred
or palimpsest softground suites of the Skolithos in more distal or sheltered settings during contin-
ichnofacies (Fig.ISC). Many firmgrounds also ued sea-level rise (MacEachern & Burton 2000).
appear to have been developed on siderite- Savrda (2001) has also identified Zoophycos as
cemented intervals within the shales (Fig. 13D). part of an omission suite in the shelf and slope
Whether the siderite is formed during ravine- deposits of New Jersey. The proximal omission
ment, as a chemical response related to deep sub- assemblages record predominantly suspension-
strate penetration by the firmground trace- feeding behaviours associated with the period
makers, or whether pre-existing, siderite-cemen- of higher energy prevalent during active ravine-
ted bands formed resistant layers through ment and/or the energy conditions at the sub-
which the TSE could not incise, is uncertain. In strate during substrate colonization (Fig. 13C,
the latter scenario, however, soft-bodied fauna D, E). Colonization of these exhumed surfaces
would have to have been capable of penetrating post-dates erosive shoreface retreat but presum-
a highly compacted or cemented layer. Else- ably occurs prior to significant deepening.
where, the TSE have been developed on sandy These higher-energy (proximal) TSE are
substrates and are marked by palimpsest soft- commonly overlain by conglomeratic lags.
ground omission suites (Fig. 13C), typically Transgressive surfaces of erosion that are not
dominated by Diplocraterion habichi and Sko- colonized until after deepening record higher
lithos (e.g. Middle Jurassic Heather Formation, proportions of domichnia of deposit-feeding
Norwegian North Sea, MacEachern & L0seth organisms and are typically overlain by marine
2003; Early Permian Pebbley Beach Formation, pebbly and sandy shales or muddy sandstones.
Australia, Bann et al 2004). In a few exceptional In the most distal settings, the omission suite
cases, TSE excavated across coal layers (Fig. 14) may consist entirely of firmground domichnia
are demarcated by Teredolites longissimus, and feeding structures of deposit-feeding
Diplocraterion parallelum and, more rarely, organisms (Fig. 13F; MacEachern & Burton
Diplocraterion habichi, attributable to the 2000).
Teredolites ichnofacies (e.g. Campanian Horse-
shoe Canyon-Bearpaw transition, Drumheller Transgressively incised shorefaces
Alberta, Saunders & Pemberton 1986; Lower Several Viking Formation oil and gas fields in
Jurassic Neil Klinter Formation, East Green- central Alberta produce hydrocarbons from
land, Dam 1990; and Lower Palaeocene Clayton NW-SE trending, sharp-based sandstones, inter-
Formation, Alabama, Savrda 1991b). preted to rest upon transgressive surfaces of
The firmground omission suites are predomi- erosion incised into underlying facies. These
nantly manifest by Diplocraterion (typically D. include Chigwell (Raychaudhuri et al. 1992),

archetypal Cruziana ichnofacies with Chondrites (Ch), Helminthopsis (H), Phycosiphon (Ph), and Diplocraterion
(D). Well 11-24-65-18W5; 1358.0m. (C) A palimpsest softground of Diplocraterion (D), subtending from a
regionally extensive TSE excavated landward of the Joffre Embayment Complex. The palimpsest suite
cross-cuts remnant lower shoreface sandstones of the regional Viking Fm parasequences. The underlying
sandstones contain a proximal expression of the Cruziana ichnofacies, with Helminthopsis (H), Siphonichnus
(Si), and Zoophycos (Z). Well 16-34-38-25W4; 1433.9m. (D) A proximal expression of a regionally extensive
TSE in the Viking Formation with a pebble lag passively infilling firmground Diplocraterion (D) and Skolithos
of the Glossifungites ichnofacies. The omission suite penetrates siderite cemented silty mudstones with visible
Palaeophycus (Pa) and Chondrites (Ch). Well 12-31-40-02w5; 1860.8m. (E) A proximal TSE overlain by a
pebble lag that passively infills firmground Diplocraterion (D). The omission suite cross-cuts lower offshore silty
mudstones with stacked tempestites, containing abundant Phycosiphon (Ph), Helminthopsis (H), and Chondrites
(Ch), comprising a distal expression of the Cruziana ichnofacies. Well 09-15-39-27W4; 1549.4m. (F) A distal
TSE from the Viking Formation of the Hamilton Field. The underlying mudstones correspond to shelf deposits
and contain the Zoophycos ichnofacies. The TSE is demarcated by a firmground omission suite, consisting of
Thalassinoides and Zoophycos of the Glossifungites ichnofacies. The overlying sandy mudstones are pebble
bearing (Pe) and show the archetypal Cruziana ichnofacies containing Zoophycos (Z), Thalassinoides (Th),
Planolites (P), Helminthopsis (H), and Palaeophycus (Pa). Well 06-13-35-09W4; 904.6m.
48 S. G. PEMBERTON ET AL.

afkldjfhjsdkfhjksdfhdkajsfhksdjfhakdjlfhlaskd
asdfasdfasdfadsfadfadfadfadfadfadsfadfadsfadsf

Fig. 14. Summary diagram showing multiple sites of compacted mud/peat exhumation in a transgressive
barrier-island setting. These include: back-barrier tidal creeks (1) and channels (2), tidal inlets (3), and the open
shoreface (4). In all of these settings, developments of the Glossifungites and Teredolites ichnofacies can
intermingle in response to localized change's in xylic properties. As shown, the erosional resistance of
compacted peat can exert an overriding control on the depths of both channel erosion and shoreface
ravinement. Trace fossil assemblages may therefore reflect complex histories repeated burial and
re-exhumation.

Joffre (Downing & Walker 1988; MacEachern contrast, Walker & Wiseman (1995) reinter-
et al 1998, 1999b), Gilby (Raddysh 1988) and preted it as a transgressive shoreface, primarily
Giroux Lake (Stelck et al 2000). These succes- on the basis of the observation of underlying
sions can be regarded as high-energy parase- and basinward shoreface deposits at Lindbrook
quences bounded by ravinement surfaces. that they regarded as lowstand in origin.
Although transgressively incised shorefaces Furthermore, despite the Lindbrook deposits
tend to display thicker successions than do being given a lowstand interpretation, Walker
falling-stage systems, reflecting the increased & Wiseman (1995) indicated that should an addi-
accommodation space available, the 'transgres- tional shoreface deposit within their sequence 1
sive' interpretation has rested mainly with the be discovered farther to the northeast, the 'inci-
perceived position of the deposits in the regional sion at Lindbrook a would then represent a
stratigrapnic framework rather than with any transgressive incision formed during movement
intrinsic characteristics of the succession itself. of the shoreline to the southwest' (Walker &
For example, Posamentier et al. (1992) and Wiseman 1995, p. 136).
Posamentier & Chamberlain (1993) interpreted It has, however, been suggested by Mac-
the Joarcam deposit as a lowstand shoreface. In Eachern et al. (1999b) that this uncertainty
KEY STRATIGRAPHIC SURFACES 49

could be alleviated through evaluation of the ero- short-lived progradation of upper offshore and
sional extent of the underlying discontinuity. lower shoreface environments of a transgres-
Transgressive ravinement causes an erosional sively incised, weakly storm-influenced shoreface
discontinuity that ultimately lies seaward of complex. In proximal positions (Fig. 15A, B), the
fair-weather wavebase during subsequent peri- transgressively incised shoreface is virtually
odic progradation. This is because the modified indistinguishable from either the forced regres-
surface was cut prior to shoreface progradation, sive (Fig. 10) or lowstand incised shorefaces
while sea-level lay at a stratigraphically lower (Fig. 10); the basal discontinuity shows firm-
position (Fig. 10). Consequently, in the trans- ground suites directly overlain by lower shore-
gressive scenario, lower offshore and upper face sandstones. The transgressive origin of an
offshore deposits, reflecting deposition below incised shoreface's basal discontinuity is demon-
fair-weather wavebase, can overlie the erosional strated, instead, in distal positions. Distally,
component of the basal discontinuity. This is a firmground omission suites of the .Glossifungites
situation that cannot be accommodated by ichnofacies demarcate the discontinuity, indicat-
either a forced regressive or a lowstand scenario ing that it continues to be erosional, even where it
(MacEachern et al. 1999a), and is diagnostic of is overlain by deposits that accumulated below
transgressively incised systems. In fact, Walker fair-weather wavebase (Fig. 15 C, D). This indi-
& Wiseman (1995) noted that an erosional cates that the surface was cut while sea-level
surface always underlies the offshore transition was lower and the area within the zone of wave
mudstone in such settings, though the implica- attack and was colonized during transgressive
tions of that observation were not explored deepening. Ultimate burial of the discontinuity
further. occurred either during a pause in the rate of
As transgressive surfaces of erosion are com- transgression, or when sediment supply to the
monly colonized by firmground omission suites, shoreline outpaced deepening and a period of
widespread firmground assemblages attributable shoreline progradation ensued. In distal posi-
to the Glossifungites ichnofacies can be gener- tions, the transgressively ravined discontinuity
ated, directly overlain by thin gravel lags and was buried beneath offshore sandy mudstone
basinal facies reflecting offshore and shelf deposi- prior to shoreface deposition (Fig. 10), as is
tion. This facies relationship stands in marked diagnostic of a transgressively incised origin. A
contrast to either forced-regression or lowstand comparable succession was described from the
systems where basinal facies overlie the non- Viking Formation of the Chigwell Field by
erosional correlative conformity and lack Raychaudhuri et al (1992).
demarcation by the Glossifungites ichnofacies. The Turonian Cardium Formation of the
Ultimately, the ravinement surfaces of transgres- Pembina field, central Alberta, also contains a
sively incised shorefaces pass seaward into non- series of conglomeratic bodies associated with
erosional marine flooding surfaces (Fig. 10). underlying transgressive surfaces of erosion,
The Viking Joffre Shoreface Complex interpreted as transgressively incised shorefaces
(Sequence 2) of the Gilby-JofTre trend (Mac- (cf. Vossler & Pemberton 1989; Walker & Eyles
Eachern et al 1998, 1999b) contains a sharp- 1991). Below this erosion surface, lower offshore
based transgressively incised shoreface, exca- silty shales are abundantly bioturbated (BI5)
vated into underlying stacked marine parase- and contain a diverse ichnological assemblage
quences. The incision surface (interpreted as an corresponding to a distal expression of the Cruzi-
FS/SB) slopes steeply seaward along its land- ana ichnofacies. The erosional discontinuity is
ward edge and flattens out basinward, forming incised into these silty shales and is marked by
an asymmetric, one-sided erosional scarp. robust, pebble-filled firmground Thalassinoides
Granules and small pebbles of chert locally (Fig. 16) and rare Skolithos of the Glossifungites
mantle the erosional discontinuity. More com- ichnofacies. The conglomerates are largely
monly, the surface is demarcated by firmground devoid of bioturbation but pass upward into
Thalassinoides, Diplocraterion and Skolithos of overlying marine shelf shales that contain trace
the Glossifungites ichnofacies, in both proximal fossil suites attributable to the Zoophycos ichno-
and distal positions (Fig. 15). The FS/SB is facies. The erosional discontinuity corresponds
overlain by a coarsening-upward (shallowing- to the E5 surface of Plint et al. (1986), interpreted
upward) succession of gritty sandy shales and as a surface of initial transgression (Plint 1988;
muddy sandstones containing a fully marine, Plint et al. 1988), upon which the conglomerates
diverse and uniformly distributed trace fossil rest. Firmground colonization of the E5 surface
assemblage that corresponds to an archetypal corresponds to a hiatus in deposition between
to proximal expression of the Cruziana ichno- the initial transgressive generation of E5 and
facies, respectively. These facies reflect the shoreface progradation of the conglomerates
50 S. G. PEMBERTON ET AL.

Fig. 15. Transgressively incised shoreface. (A) Box shot of core from the Joffre Shoreface Complex. Base of the
interval is to the lower left, and top to the upper right (T). The FS/SB here lies in a proximal position. Silty
and sandy mudstones of the regional Viking Formation, reflecting lower offshore and upper offshore conditions
respectively, are erosionally truncated by the basal discontinuity, Overlying the FS/SB are conglomeratic
sandstones of the transgressive lag, passing into bioturbated muddy sandstones of the lower shoreface.
Well 09-16-39-27W4; 1560.8-1565.1 m. (B) Close-up photo of the FS/SB from photo A, demarcated by
firmground Thalassinoides (Th) of the Glossifungites ichnofacies. The discontinuity is overlain by a pebble (Pe)
lag and muddy sandstones of the lower shoreface containing a proximal expression of the Cruziana ichnofacies.
Skolithos (S), Planolites (P), Asterosoma (As), Palaeophycus (Pa), and Siphonichnus (Si) are visible. Well 09-16-
39-27W4; 1562.5m. (C) Box shot of core from the Joffre Shoreface Complex, in a position distal to that of
photos A and B. Base of the interval is to the lower left, and top to the upper right (T). Here, lower and upper
offshore mudstones of the regional Viking Formation are erosionally truncated by the FS/SB, but overlain by
upper offshore sandy mudstones of the Joffre Shoreface Complex. Well 08-14-38-25W4, 1431.6-1434.5 m. (D)
Close-up photo of the FS/SB of photo C, showing the distal expression of the FS/SB. Here, the omission suite
demarcating the discontinuity also consists of firmground Thalassinoides (Th) of the Glossifungites ichnofacies.
However, it is overlain by bioturbated (BI5) gritty, pebble (Pe) bearing sandy mudstones of the upper offshore,
with Phycosiphon (Ph), Chondrites (Ch), Asterosoma (As), Cylindrichnus (Cy), Palaeophycus (Pa), and Planolites
(P) corresponding to the archetypal Cruziana ichnofacies. Well 08-14-38-25W4, 1434.0m.
KEY STRATIGRAPHIC SURFACES 51

such surfaces may also include the discontinuities


at the bases of some transgressively incised
shorefaces (e.g. E-T surfaces of Flint 1988; Flint
et al 1986, 1988; FS/SB of MacEachern et al.
1992, 1998, 1999a, 1999b), the majority are asso-
ciated with incised valley systems. Incised valley
discontinuities may correspond to subaerially
exposed areas, such as delta plains, fluvial flood-
plains, interfluves, or transgressively modified
sequence boundaries within the estuarine valley
fills themselves. Some reserve the term TS/SB'
for discontinuities developed on valley inter-
fluves, where deposition did not occur until the
valley was filled and the area transgressively
ravined during ensuing relative sea-level rise
(e.g. Van Wagoner et al. 1990).

TSE across subaerially exposed surfaces or


Fig. 16. Transgressively incised Cardium shoreface. interfluves
(A) The FS/SB marking the base of the Cardium
Formation conglomeratic shoreface of the Pembina Numerous units in coastal margin (delta plain
field. Moderately bioturbated (BI4-5) sandy and coastal plain) settings display initially sub-
mudstones of the upper offshore, containing
Chondrites (Ch) and Helminthopsis (H), are aerial surfaces subsequently flooded and eroded
erosionally truncated by overlying pebble during transgressive influx of brackish to
conglomerates. The discontinuity is demarcated by marine waters. Such scenarios are conducive to
gravel-filled firmground Thalassinoides (Th) of the the development of substrate-controlled ichno-
Glossifungites ichnofacies. Well 04-13-51-11W5, facies demarcating the discontinuities. The
1636.1 m. (B) The same discontinuity in a nearby Cenomanian Dunvegan Formation consists of
core, showing lower offshore silty mudstones a stacked succession of prograding delta lobes
truncated by pebble conglomerates of the Cardium that varied during its history of deposition
Formation incised shoreface. The firmground from river-dominated to wave-dominated in
omission suite also consists of gravel-filled
Thalassinoides (Th) of the Glossifungites ichnofacies. character (Bhattacharya & Walker 1991). The
Well 12-09-51-10W5, 1596.8m. stacked delta lobes and individual shingles
within the lobes are separated by marine flooding
surfaces of varying scales and which are locally
during a pause in the rate of transgression. The erosional (Bhattacharya 1993). In the subsurface
conglomeratic shoreface was ultimately drowned of the Jayar Field, central Alberta, a TSE, over-
(MFS), and locally removed (TSE), during lain by a transgressive sandstone, cuts across
resumed transgression. rooted and subaerially exposed delta plain
deposits of the underlying delta lobe (Bhatta-
charya & Walker 1991). The erosional dis-
Amalgamated sequence boundaries and continuity is demarcated by a firmground
marine flooding surfaces Thalassinoides of the Glossifungites ichnofacies,
passively filled with coarse-grained sand derived
Amalgamated sequence boundaries and trans- from the overlying transgressive sand sheet.
gressive surfaces are commonly colonized by Similarly, the Lower Albian Mannville Group-
substrate-controlled trace-makers. The lowstand Joli Fou Formation contact in the Kaybob
erosion event typically produces widespread Field of central Alberta is manifest by a region-
firmground, hardground and woodground sur- ally developed FS/SB. In this case, rooted
faces, corresponding to RSE or SB. The follow- incipient palaeosols developed on floodplain
ing transgressive event, commonly accompanied mudstones of the Mannville Group are cross-
by erosion, generates a TSE that tends to cut by robust, firmground Thalassinoides, reflect-
remove most or all of the lowstand deposits ing the Glossifungites ichnofacies (Fig. 17A),
and exposes the original discontinuity to passively filled with muddy sand and large side-
marine or marginal marine conditions. During rite-cemented clasts. The overlying silty shales
this phase of transgression, organisms are able contain a distal expression of the Cruziana
to colonize the re-exhumed substrate. Although ichnofacies and, more rarely, the Zoophycos
52 S. G. PEMBERTON ET AL.

the subsurface, coastal plain to floodplain mud-


stones and siltstones, alternating with palaeosols,
correspond to the Upper Boulder Creek Forma-
tion (post-Cadotte Member but pre-Paddy
Member). These represent the preserved terres-
trial deposits that accumulated during late
Cadotte Member highstand conditions and
initial sea-level fall, which survived incision
during generation of the Paddy Member discon-
formity. The Upper Boulder Creek palaeosols
locally consist of rooted silty light grey mud-
stones with hematite-stained spherulitic siderite
(Leckie et al. 1989). These palaeosols are trun-
cated by a transgressive surface of erosion that
locally displays an omission suite of thin,
sharp-walled Skolithos and more rarely Rhizo-
corallium of the Glossifungites ichnofacies (Fig.
17B). The overlying granule-bearing, sandy
mudstones of the Paddy Member are of low
diversity, archetypal to distal expressions of the
Cruziana ichnofacies, and reflect restricted bay
conditions during regional flooding of the coastal
Fig. 17. FS/SB Interfluve. (A) Glossifungites margin. Regionally, the Paddy Member occupies
ichnofacies-demarcated FS/SB at the Mannville a major estuarine valley excavated during initial
Group-Joli Fou Formation contact, reflecting an lowstand conditions (Leckie & Singh 1991).
interfluve area. Rooted incipient palaeosols are The flooding of the estuary margins during late
colonized with firmground Thalassinoides (Th), and Paddy time resulted in transgressive modification
capped by a transgressive lag. Overlying facies reflect
offshore to shelf deposition. Well 11-03-60-19W5;
of the interfluve area. Continued transgression
1894.3m. (B) Rooted (r), incipient palaeosols of the resulted in the deposition of open marine
Upper Boulder Creek Formation, corresponding to mudstones of the Shaftesbury Formation and
floodplain conditions, are transgressively eroded and the return to shelf al conditions (Leckie et al.
overlain by a thin transgressive lag and brackish- 1991).
water bay mudstones of the Paddy Formation. The
FS/SB of the interfluve is demarcated by firmground
Thalassinoides (Th) of the Glossifungites ichnofacies. Incised valley complexes: demarcation of
The overlying brackish-water mudstones contain an
impoverished Cruziana ichnofacies, with visible valley surfaces
Planolites (P) and exceedingly rare Helminthopsis (H).
Well 08-13-69-11W6; 1907.2m. For the most part, lowstand deposits rarely dom-
inate incised valley complexes, as the system is
largely a zone of sediment bypass during incision
ichnofacies, recording deposition in lower off- (Van Wagoner et al. 1990). Much of the sediment
shore to outer shelf environments. The amalga- accumulation in these systems occurs during late
mated surface corresponds to an interfluve lowstand and transgression, and is therefore
(Van Wagoner et al. 1990) that was transgres- characterized by estuarine infill. The juxtaposi-
sively overrun during basin-wide flooding of the tion of facies into which the valley is incised,
Joli Fou Seaway. This transgression marks a the presence of remnant fluvial deposits within
major period of marine inundation of the the valley, accumulations of estuarine intervals
Western Interior Seaway of North America. during ensuing transgression, and the excavation
The Upper Albian Paddy Member is separated of numerous internal discontinuities within the
from the underlying Cadotte Member in the valley fill, result in highly complex successions
Peace River area of Alberta by a regionally that ichnology is ideally suited to help resolve.
extensive disconformity that was excavated Ichnological suites are also effective at differen-
during a relative sea-level fall and subsequently tiating salinity changes and, more specifically,
transgressively modified during flooding of the salinity reductions, assisting in the differentiation
basin (Leckie & Singh 1991). This continued of fully marine, brackish and freshwater deposits
flooding eventually led to the return of offshore (e.g. Pemberton et al. 1982; Beynon et al. 1988;
to shelf conditions reflected by the overlying Ranger & Pemberton 1992, 1997; MacEachern
Shaftesbury Formation (Leckie et al. 1990). In & Pemberton 1994; MacEachern et al. 1999a).
KEY STRATIGRAPHIC SURFACES 53

Fig. 18. Schematic model of incised valley surface types commonly demarcated by the Glossifungites
ichnofacies (modified after MacEachern & Pemberton 1994).

This, coupled with the presence of substrate- channels within the valley (Fig. 18). The valley
controlled assemblages associated with erosional margins are excavated into coarsening upward,
discontinuities within the valley fill, allows regional Viking highstand marine para-
detailed mapping of valley components and sequences. These parasequences contain fully
assists in the resolution of the sequence strati- marine, high diversity and abundantly burrowed
graphic history of valley excavation and infill. (BI5) distal to proximal expressions of the Cruzi-
The Viking Formation produces hydro- ana ichnofacies. This contrasts markedly with
carbons from estuarine incised valley fills in at the ichnological suites developed within the
least five fields of central Alberta. The facies suc- valley fills. In the Crystal Field the valley margins
cessions and their distributions indicate that they are demarcated by firmground Diplocraterion,
accumulated in a barrier estuary or wave-domi- Thalassinoides and unnamed clavate burrows
nated embayed estuary setting, in the sense of similar in morphology to Gastrochaenolites,
Roy et al (1980) and Dalrymple et al. (1992). assigned to the Glossifungites ichnofacies (Fig.
In most of the incised valley systems of the 19B, C). At Willesden Green, the valley base
Viking Formation the valley margins are demar- contains firmground Arenicolites, Skolithos,
cated trace fossil suites of the Glossifungites ich- Diplocraterion, Rhizocorallium and Thalassi-
nofacies, indicating that the valley probably did noides (Fig. 19F).
not fill until active transgression (i.e. no lowstand
deposits are preserved). Either the valley served
as a zone of sediment bypass and possessed no Incised valley complexes: ichnology of
fluvial deposits, or any lowstand deposits were estuarine valley fills
subsequently eroded and reworked during the
subsequent transgression, producing an amalga- The wave-dominated barred estuary systems can
mated (co-planar) sequence boundary and initial be separated into the bay-head delta, including
transgressive surface. This initial transgressive the active channels and distributary channels,
surface of erosion most likely reflects tidal- the central basin or lagoon, and the estuary
scour ravinement. The base of the estuarine mouth. The bay-head delta complex (Fig. 19A)
valley fill therefore serves both as the sequence is generally characterized by weakly and sporadi-
boundary and as the base of the transgressive cally burrowed, parallel-laminated sandstones.
systems tract. The valley fill, as well, contains a Trace fossil diversities can be high (up to 16
number of internal stratigraphic discontinuities, ichnogenera), although burrow concentrations
locally reflecting re-incision of the valley, erosive are irregular and the numbers of individual
back-stepping of the barred mouth, lateral and forms are low. Bioturbation intensities are also
landward shift of the tidal inlets at the barrier generally low (BI 1-2). The trace fossils comprise
mouth, and lateral shift of tidal creeks and tidal an impoverished expression of the Skolithos
54 S. G. PEMBERTON ET AL.

Fig. 19. Incised Valley Complex. (A) Parallel-laminated sandstone of the bayhead delta front, Crystal Field.
The sandstone is sporadically burrowed with a low-diversity expression of the Skolithos ichnofacies. Unit
shows Bergaueria (Be) and Diplocraterion (D). Well 16-24-45-04W5; 1801.3m. (B) Box shot of core from the
incised valley of the Crystal Field. Base of the core is to the lower left, and top to the upper right (T).
Underlying lower offshore silty mudstones of the regional Viking Formation parasequences have been
truncated by an amalgamated flooding surface and sequence boundary (FS/SB) along the margins of the
Crystal incised valley. The valley fill at this locality consists of sandstone-dominated central basin deposits that
have onlapped the valley margins. Well 04-01-46-04W5; 1802.1-1806.2 m. (C) Close-up of the contact visible in
photo B. Lower offshore silty mudstones of the regional Viking contain a distal expression of the Cruziana
ichnofacies with Helminthopsis (H), Asterosoma (As), and Palaeophycus (Pa). The FS/SB is demarcated by a
firmground omission suite of Skolithos (S), and unnamed flask-shaped domichnia similar to Gastrochaenolites
(G). The overlying bay deposits show dispersed pebbles (pe) at the base with a low-diversity suite attributable
to the mixed Skolithos-Cruziana ichnofacies. Well 04-01-46-04W5; 1804.5m. (D) Sandy central basin deposits
from the Willesden Green incised valley. Current ripple laminated sandstone is intercalated with strongly
burrowed (BI4-5) muddy sandstone showing a low-diversity suite of the mixed Skolithos-Cruziana ichnofacies.
The facies displays Thalassinoides (Th), Macaronichnus (Ma), Palaeophycus (Pa), Siphonichnus (Si), Bergaueria
(Be), Rosselia (Ro) and Planolites (P). Well 06-36-40-07W5; 2322.7m. (E) Bioturbated (BI4) muddy sandstone
from the estuary mouth complex of the Crystal incised valley. The suite corresponds to the Skolithos
ichnofacies, comprising Thalassinoides (Th), Ophiomorpha (O), Skolithos (S), Diplocraterion (D), Planolites (P),
Helminthopsis (H), Palaeophycus (Pa), Siphonichnus (Si), and Teichichnus (Te). Well 08-16-48-03W5; 1529.1 m.
(F) Tidal inlet channel fill deposit from the Willesden Green incised valley. Lower offshore, silty mudstones of
the regional Viking Formation parasequences with visible Phycosiphon (Ph), and Chondrites (Ch). The
discontinuity corresponds to a transgressive scour ravinement (TSR) surface, demarcated by a firmground
omission suite of Rhizocorallium (Rh), and Thalassinoides (Th). The channel fill sandstone displays dispersed
pebbles (pe), and isolated, robust Ophiomorpha (O). Well 11-31-40-06W5; 2285.6m.
KEY STRATIGRAPHIC SURFACES 55

ichnofacies. The central basin complex (Fig. 19B, interbeds. Suites are characterized by sparse
C, D) consists of delicately interstratified sandy Planolites, Palaeophycus and Skolithos, with
mudstones, dark mudstones and thin sandstones. rare Cylindrichnus, Teichichnus and Arenicolites
Synaeresis cracks are present throughout and are comprising the secondary elements. Very uncom-
locally common. The trace fossil assemblages mon ichnogenera include Conichnus, Gyrolithes
show moderate to low diversities of ichnogenera, and fugichnia. IHS intervals are characterized
moderate though variable bioturbation intensi- by stacked, trough cross-stratified beds and
ties (BI2-5; typically BI4), and reflect the current-rippled beds alternating with thin,
mixed Skolithos-Cruziana ichnofacies. Trace depositionally inclined (3-7°) mudstone beds,
fossil suites in the central basins are dominated interpreted to reflect tidal modification of river
by Teichichnus, Planolites, diminutive Rosselia, flow through the channel system. Bioturbation
Cylindrichnus and Palaeophycus (Fig. 19D). intensities are variable but moderate to low
Salinity fluctuations, episodic deposition and (BI2-4), and ichnogenera are sporadically dis-
variable substrate consistency appear to be the tributed. Trace fossil suites are dominated by
dominant stresses imparted on the infauna. The moderate to common Planolites, with moder-
estuary mouth complex (Fig. 19E) consists of ately common to rare Teichichnus, Cylindrichnus
moderately to abundantly burrowed (BI3-5) and Skolithos. Secondary elements comprise
current and oscillation ripple-laminated sand- rare Gyrolithes, Palaeophycus, fugichnia and
stones, with minor intercalated mud beds. The roots. Accessory elements include very rare
trace fossil suites show high diversities of ichno- Arenicolites, Rosselia, Thalassinoides, Chon-
genera, corresponding to the Skolithos ichnofa- drites, Bergaueria, Rhizocorallium, Lockeia and
cies, but the distribution of individual elements Ophiomorpha. Some intervals consist of mono-
reflects the presence of environmental stresses, generic assemblages of Planolites, Cylindrichnus
in particular, episodic deposition. or Gyrolithes, particularly within the better-
Channel-fill fades associations (Fig. 19F) pre- studied McMurray Formation palaeo-valleys.
dominantly consist of the deposits of relatively More saline elements of the suite comprise
small, migrating subaqueous dunes. The amalga- Chondrites, Rosselia, Rhizocorallium, Bergaueria
mation of the trough cross-beds into thick and Ophiomorpha, all of which have only been
intervals supports a high sediment aggradation described from McMurray Formation intervals
rate. The trace fossil suite is sporadically distrib- (Bechtel et al. 1994; Ranger & Pemberton 1997).
uted, low in diversity (seven ichnogenera) and The trace fossil assemblages within the incised
corresponds to the Skolithos ichnofacies. The valley facies associations correspond to simple
most common elements include Ophiomorpha, structures produced by trophic generalists.
Cylindrichnus, Palaeophycus and Diplocraterion. These are referred to as r-selected (opportunistic)
The burrowing demonstrates that most of the behaviours, and are characteristic of stressed
channel complexes accumulated in marine or environmental settings (Pianka 1970), particu-
marginal marine conditions, although the larly those subject to salinity fluctuations. The
degree of salinity stress is difficult to determine. episodic nature of deposition and the variability
The main stress imposed on the trace fossil in substrate consistency lead to the development
suite appears to be related to migration of bed- of trace fossil assemblages that constitute
forms and avalanching of sand into the dwelling an impoverished expression of the mixed
structures. Skolithos-Cruziana ichnofacies (Pemberton
In the more tidally influenced estuaries of the et al. 1992a).
Alberta Basin [e.g. the McMurray Formation
(Pemberton et al. 1982; Ranger & Pemberton
1992, 1997; Bechtel et al. 1994); the Grand Bay-head delta, channel and embayment
Rapids Formation (Wightman et al. 1987; deposits
Beynon et al. 1988); and the Glauconite For-
mation (Leroux et al. 1999)], the valley fills In the Viking Formation of the JofTre Field, an
consist predominantly of channel sandstones amalgamated sequence boundary and flooding
and lateral accretion deposits manifest by surface with a scarp-like geometry truncates
inclined heterolithic stratification (IHS). The underlying regional Viking marine para-
channel sandstones are dominated by stacked sequences and, locally, the transgressively incised
trough cross-stratified beds, lesser low-angle Joffre Shoreface Complex (MacEachern et al.
planar cross-stratified units, and thin current 1998, 1999a). The Glossifungites ichnofacies,
ripple laminated beds. Bioturbation is generally characterized by firmground Skolithos, Diplocra-
very low (BI1-2) and sporadically distributed, terion and Thalassinoides, locally helps to demar-
and typically is associated with thin mudstone cate this erosional discontinuity (Fig. 20A).
56 S. G. PEMBERTON ET AL.

Fig. 20. Joffre Embayment. (A) Joffre Embayment Complex showing regional Viking Formation lower offshore
silty mudstones with Phycosiphon (Ph) and Planolites (P), truncated by a regionally extensive FS/SB, and
overlain by embayment sandstones. The discontinuity is demarcated by firmground Thalassinoides (Th) of the
Glossifungites ichnofacies. Well 14-11-39-27W4; 1572.7m. (B) Glauconitic pebbly (pe) muddy sandstones of
the Joffre Embayment Complex corresponding to a transgressive sand sheet at the base of the regional FS/SB.
The sandstone is moderately well bioturbated (BI4) containing a proximal expression of the Cruziana
ichnofacies. The sandstone contains Planolites (P), Rosselia (Ro), Helminthopsis (H), Palaeophycus (Pa), and
Chondrites (Ch). Well 14-05-38-24W4; 1372.3m. (C) Trough cross-stratified sandstone of the Joffre Embayment
KEY STRATIGRAPHIC SURFACES 57

The deposits overlying the discontinuity con- more pronounced, and parasequence boundaries
stitute the Viking reservoir facies at Joffre and cannot be delineated easily. Near the southern
reflect three stacked, conglomeratic embayment end of the field, these parasequences partition
parasequences that prograded toward the north- the reservoir along the seaward (and structurally
east. The reservoir facies are dominated by up-dip) edge of the deposit. Amalgamation of the
trough cross-stratified and low-angle planar reservoir facies at the north end limits the degree
stratified sandstones, pebbly sandstones and of partitioning.
conglomerates, concentrated along the southern The final parasequence of the embayment
margin of the amalgamated sequence boundary complex is truncated by a regionally extensive
and flooding surface. The coarse elastics progres- flooding surface, typically manifest as a wave
sively inter-finger with, and ultimately pass into, ravinement surface. The wave ravinement sur-
interbedded mudstones and fine-grained sand- face is commonly demarcated by the Glossi-
stones in a northward and eastward direction. fungites ichnofacies, or where excavated across
Near the base, the coarse elastics contain glauco- sandy underlying facies, a palimpsest softground
nite, siderite-cemented mudstone interbeds, mud suite of Diplocraterion (Fig. 13C). Facies over-
inter-laminae and resistant mudstone rip-up lying the marine flooding surface reflect fully
clasts, and display moderate to low degrees of marine conditions.
burrowing, diminishing in intensity upward
(Fig. 20B). The trace fossil suite corresponds to
the Skolithos ichnofacies. Overlying facies are Conclusions
dominated by well-sorted, unidirectional trough
cross-bedded and low-angle planar stratified The main applications of ichnology to genetic
coarse elastics, locally in fining upward cycles stratigraphy are twofold. The most obvious use
with scoured bases (Fig. 20C, D). The elastics lies in the demarcation of erosional discontinu-
contain mudstone rip-up clasts and thin mud- ities. To date, substrate-controlled ichnofacies
stone interbeds. Burrowing is of low abundance have been underutilized but are gaining recogni-
(BI1-2), and reduced diversity, with Diplocrater- tion as a viable means of recognizing and map-
ion, Skolithos, Palaeophycus and Ophiomorpha of ping these stratigraphically important surfaces,
the Skolithos ichnofacies. The interbedded mud- both in outcrop and subsurface. Locally, many
stone and sandstone deposits contain oscillation surfaces are obvious on the basis of sedimentol-
ripples, wavy lamination, combined flow ripples ogy alone; however, their appearance can
and rare current ripples (Fig. 20E, F). These change markedly across the study area, making
heterolithic intervals are weakly burrowed correlation difficult. Substrate-controlled ichno-
(BI 1-3) with a sporadically distributed, low- facies, such as the Glossifungites ichnofacies,
diversity (salinity stressed?) trace fossil suite of are also important to the genetic interpretation
the mixed Skolithos-Cruziana ichnofacies of erosional discontinuities in marine-influenced
(MacEachern et al 1998, 1999a). Dominant siliciclastic intervals, as the many examples
elements comprise Teichichnus, Cylindrichnus, cited in the paper demonstrate. Hence, even
Planolites and Palaeophycus. where discontinuities can be recognized purely
Detailed ichnological, sedimentological and sedimentologically or stratigraphically, the asso-
stratigraphic analyses demonstrate that the ciated trace fossils enhance their sequence strati-
coarse elastics overlying the discontinuity com- graphic interpretation. In many cases, the genetic
prise at least three parasequences. These parase- interpretation of the discontinuity has come prin-
quences onlap the discontinuity in a southwest cipally from the trace fossil assemblages that are
direction and inter-finger with mudstones to the associated with the discontinuity and the over-
northeast. Toward the north end of the field, lying units. The continued integration of
erosional amalgamation of the coarse elastics is substrate-controlled ichnofacies with detailed

Complex with sporadically distributed lined Diplocraterion (D). Well 02-05-39-26W4; 1573.3m. (D) Trough
cross-stratified sandstone of the Joffre Embayment Complex with sporadically distributed lined Diplocraterion
(D). Well 11-07-39-26W4; 1548.1 m. (E) Heterolithic succession of oscillation rippled, combined flow rippled
and low-angle parallel-laminated sandstone and dark, weakly burrowed (BI 2-3) mudstone corresponding to
open bay deposits. Trace fossils reflect a stressed, low-diversity expression of the mixed Skolithos-Cruziana
ichnofacies, characterized by Teichichnus (Te), Cylindrichnus (Cy), and Planolites (P). Well 02-18-38-24W4;
1459.2m. F) Heterolithic succession of oscillation rippled (osc), current-rippled (cr), and low-angle parallel-
laminated sandstone, with dark, weakly burrowed (BI 2) mudstone corresponding to open bay deposition.
Trace fossils reflect a stressed, low-diversity expression of the mixed Skolithos-Cruziana ichnofacies,
characterized by Teichichnus (Te), and Planolites (P). Well 03-24-38-25W4; 1437.7m.
58 S. G. PEMBERTON ET AL.

stratigraphic and sedimentological analysis will USA) reinterpreted as lowstand shoreface depos-
undoubtedly enhance and refine the developing its. American Association of Petroleum Geologists
genetic stratigraphic paradigms. Bulletin, 64, 184-201.
The second use is more subtle, and is con- BEYNON, B. M., PEMBERTON, S. G., BELL, D. A. &
LOGAN, C. A. 1988. Environmental implications
cerned with trace fossil behaviours and their of ichnofossils from the Lower Cretaceous
palaeoenvironmental implications. Trace fossils, Grand Rapids Formation, Cold Lake Oil Sands
when used in conjunction with primary sedimen- Deposit. In: JAMES, D. R. & LECKIE, D. A. (eds)
tary structures, are useful in the delineation and Sequences, Stratigraphy, Sedimentology: Surface
interpretation of facies and facies associations. and Subsurface. Canadian Society of Petroleum
When these behavioural and substrate-con- Geologists, Memoirs, Calgary, Alberta, 15, 275-
trolled aspects of ichnology are integrated fully 290.
with other sedimentological and stratigraphic BHATTACHARYA, J. P. 1993. The expression and inter-
analyses, the result is a powerful approach to pretation of marine flooding surfaces and ero-
the recognition and genetic interpretation of dis- sional surfaces in core: examples from the Upper
Cretaceous Dunvegan Formation, Alberta fore-
continuities in the rock record. land basin, Canada. In: POSAMENTIER, H. W.,
SUMMERHAYES, C. P., HAQ, B. U. & ALLEN, G. P.
The authors would like to thank the Natural Science (eds) Stratigraphy and Facies Associations in a
and Engineering Research Council of Canada
Sequence Stratigraphic Framework. International
(NSERC) for research funding. The senior author
Association of Sedimentologists, Special Publica-
would like to acknowledge the Canada Research
tions, Oxford, 18, 125-160.
Chairs programme for their support of his research.
BHATTACHARYA, J. & WALKER, R. G. 1991. Allostrati-
D. Robbins assisted with some of the drafting, and
graphic subdivision of the Upper Cretaceous
we are grateful for his contribution.
Dunvegan, Shaftesbury and Kaskapau forma-
tions, northwestern Alberta subsurface. Bulletin
of Canadian Petroleum Geology, 39, 145-164.
References BROMLEY, R. G. & ASGAARD, U. 1993. Two bioerosion
ichnofacies produced by early and late burial
AINSWORTH, R. B. & PATTISON, S. A. J. 1994. Where associated with sea level change. Geologische
have all the lowstands gone? Evidence for attached Rundschau, 82, 276-280.
lowstand systems tracts in the Western Interior of BROMLEY, R. G., PEMBERTON, S. G. & RAHMANI,
North America. Geology, 22, 415-418. R. A. 1984. A Cretaceous woodground: the
BANN, K. L. 1998. Ichnology and sequence stratigraphy Teredolites ichnofacies. Journal of Paleontology,
of the Early Permian Pebbley Beach Formation 58, 488-498.
and Snapper Point Formation in the southern DALRYMPLE, R. W., ZAITLIN, B. A. & BOYD, R. 1992.
Sydney Basin. PhD thesis, University of Wollon- Estuarine facies models: conceptual basis and
gong. stratigraphic implications. Journal of Sedimentary
BANN, K. L., FIELDING, C. R., MACEARCHERN, J. A. Petrology, 62, 1130-1146.
& TYE, S. C. 2004. Differentiation of estuarine DAM, G. 1990. Paleoenvironmental significance of trace
and offshore marine deposits using integrated fossils from the shallow marine Lower Jurassic
ichnology and sesdimentology: Permian Pebbley Neill Klinter Formation, East Greenland. Palaeo-
Beach Formation, Sydney Basin, Australia. In: geography, Palaeoclimatology, Palaeoecology, 79,
MclLROY, D. (ed.) The Application of Ichnology 221-248.
to Palaeoenvironmental and Stratigraphic Analysis. DAVIES, S. D. & WALKER, R. G. 1993. Reservoir geome-
Geological Society, London, Special Publications, try influenced by high-frequency forced regres-
228, 179-212. sions within an overall transgression: Caroline
BASAN, P. B. & FREY, R. W. 1977. Actual-palaeontol- and Garrington fields, Viking Formation (Lower
ogy and neoichnology of salt marshes near Cretaceous), Alberta. Bulletin of Canadian Petro-
Sapelo Island, Georgia. In: CRIMES, T. P. & leum Geology, 41, 407-421.
HARPER, J. D. (eds) Trace Fossils 2, Geological DE GIBERT, J. M., MARTINELL, J. & DOMENECH, R.
Journal, Special Issue 9, 49-90. 1998. Entobia ichnofacies in fossil rocky shores,
BECHTEL, D. J., YUILL, C., RANGER, M. J. & PEMBER- Lower Pliocene, northwestern Mediterranean.
TON, S. G. 1994. Ichnology of inclined heterolithic Palaios, 13, 476-487.
stratification of the McMurray Formation, north- DOWNING, K. P. & WALKER, R. G. 1988. Viking For-
eastern Alberta. In: PEMBERTON, S. G., JAMES, D. mation, Joffre Field, Alberta: shoreface origin of
P. & WIGHTMAN, D. M. (eds) Mannville Core long, narrow sand body encased in marine mud-
Conference. Canadian Society of Petroleum stones. Bulletin American Association Petroleum
Geologists, Calgary, Alberta, 351-368. Geologists, 72, 1212-1228.
BELKNAP, D. G. & KRAFT, J. C. 1981. Preservation EKDALE, A. A., BROMLEY, R. G. & PEMBERTON, S. G.
potential of transgressive coastal lithosomes on 1984. Ichnology: Trace Fossils in Sedimentology
the US Atlantic shelf. Marine Geology, 42,429-442. and Stratigraphy. Society of Economic Paleontolo-
BERGMAN, K. M. 1994. Shannon Sandstone in Hartzog gists and Mineralogists, Short Course Notes,
Draw-Heldt Draw fields (Cretaceous, Wyoming, Tulsa, Oklahoma, 15.
KEY STRATIGRAPHIC SURFACES 59

FISCHER, A. G. 1961. Stratigraphic record of transgres- KOLLA, V., POSAMENTIER, H. W. & ElCHENSEER, H.
sing seas in light of sedimentation on the Atlantic 1995. Stranded parasequences and the forced
coast of New Jersey. American Association of regressive wedge systems tract: deposition during
Petroleum Geologists Bulletin, 45, 1656-1666. base level fall - discussion. Sedimentary Geology,
FREY, R. W. & GOLDRING, R. 1992. Marine event beds 95, 139-145.
and recolonization surfaces as revealed by trace LECKIE, D. A. & SINGH, C. 1991. Estuarine deposits of
fossil analysis. Geological Magazine, 129, 325-335. the Albian Paddy Member (Peace River Forma-
FURSICH, F. T. & MAYR, H. 1981. Non marine Rhizo- tion) and lowermost Shaftesbury Formation,
corallium (trace fossil) from the Upper Freshwater Alberta, Canada. Journal of Sedimentary Petrol-
Molasse (Upper Miocene) of southern Germany. ogy, 61, 825-849.
Neues Jahrbuch Geologie Palaontologie, Monat- LECKIE, D. A., Fox, C. & TARNOCAL, C. 1989. Multiple
shefte, 6, 321-333. palaeosols of the late Albian Boulder Creek
GALLOWAY, W. E. 1989a. Genetic Stratigraphic Formation, British Columbia, Canada. Sedimen-
sequences in basin analysis I: Architecture and tology, 36, 307-323.
genesis of flooding surface bounded depositional LECKIE, D. A., STANILAND, M. R. & HAYES, B. J. 1990.
units. American Association of Petroleum Geolo- Regional maps of the Albian Peace River Arch,
gists Bulletin, 73, 125-142. northwestern Alberta and northeastern British
GALLOWAY, W. E. 1989b. Genetic Stratigraphic Columbia. Bulletin Canadian Society of Petroleum
sequences in basin analysis II: Application to Geologists, 38, 176-189.
Northwest Gulf of Mexico Cenozoic Basin. Amer- LEROUX, M. S., MACEACHERN, J. A. & ZAITLIN, B. A.
ican Association of Petroleum Geologists Bulletin, 1999. Style contrast of estuarine incised valley
73, 143-154. complexes: ichnological and sedimentological ana-
GHIBAUDO, G., GRANDESSO, P., MASSARI, F. & lysis of valley trends in the Glauconite Member
UCHMAN, A. 1996. Use of trace fossils in delineat- and Viking Formation, central Alberta. In:
ing sequence Stratigraphic surfaces (Tertiary Vene- WRATHALL, B., JOHNSTON, G., ARTS, A., Rozsw,
tian Basin, northeastern Italy). Palaeogeography, L., ZONNEVELD, J.-P., ARCURI, D. & McLELLAN,
Palaeoclimatology, Palaeoecology, 120, 261-279. S. (eds) Digging Deeper, Finding a Better Bottom
GINGRAS, M. K. & PEMBERTON, S. G. 2000. Firmness Line. Canadian Society of Petroleum Geologists,
profiles associated with tidal creek deposits: the Core Conference, Calgary, Alberta, paper 99-
temporal significance of Glossifungites assem- 127C.
blages. Journal of Sedimentary Research, 70, MACEACHERN, J. A. & BURTON, J. A. 2000. Firm-
1025-1033. ground Zoophycos in the Lower Cretaceous
GINGRAS, M. K., PEMBERTON, S. G., SAUNDERS, T. & Viking Formation, Alberta: a distal expression of
CLIFTON, H. E. 1999. The ichnology of brackish the Glossifungites ichnofacies. Palaios, 15, 387-
water Pleistocene deposits at Willapa Bay, 398.
Washington: variability in estuarine settings. MACEACHERN, J. A. & LOSETH, T. M. 2003. Sedimen-
Palaios, 14, 352-374. tology and ichnology of transgressively back-
GROENEWALD, G. H., WELMAN, J. & MACEACHERN, stepped wave-dominated deltaic reservoir:
J. A. 2001. Vertebrate burrow complexes from Middle Jurassic Tarbert and Heather Formations,
the Early Triassic Cynognathus Zone (Driekoppen North Sea, Norway. Abstract Volume, AAPG
Formation, Beaufort Group) of the Karoo Basin, Annual Convention, Salt Lake City, Utah, May
South Africa. Palaios, 16, 148-160. 2003, p. Al 10.
GRUSZCZYNSKI, M. 1986. Hardground and ecological MACEACHERN, J. A. & PEMBERTON, S. G. 1992.
succession in the light of early diagenesis (Jurassic, Ichnological aspects of Cretaceous shoreface
Holy Cross Mountains, Poland). Acta Palaeonto- successions and shoreface variability in the
logica Polonica, 31, 163-212. Western Interior Seaway of North America. In:
GRUSZCZYNSKI, M. 1998. Chemistry of Jurassic seas PEMBERTON, S. G. (ed.) Applications of Ichnology
and its bearing on the existing organic life. Acta to Petroleum Exploration: A Core Workshop.
Geologica Polonica, 48, 1-29. Society of Economic Paleontologists and Mineral-
HAYWARD, B. W. 1976. Lower Miocene bathyal and ogists, Core Workshops, Tulsa, Oklahoma, 17,
submarine canyon ichnocoenoses from North- 57-84.
land, New Zealand. Lethaia, 9, 149-162. MACEACHERN, J. A. & PEMBERTON, S. G. 1994.
HELLAND-HANSEN, W. & GJELBERG, J. G. 1994. Con- Ichnological character of incised valley fill systems
ceptual basis and variability in sequence stratigra- from the Viking Formation of the Western
phy: a different perspective. Sedimentary Geology, Canada Sedimentary Basin, Alberta, Canada. In:
92, 31-52. DALRYMPLE, R., BoYD, R. & ZAITLIN, B. (eds)
HUNT, D. & TUCKER, M. E. 1992. Stranded parase- Recognition and Fades of Incised- Valley Fills.
quences and the forced regressive wedge systems Society of Economic Paleontologists and Mineral-
tract: deposition during base level fall. Sedimen- ogists, Special Publications, Tulsa, Oklahoma, 51,
tary Geology, 81, 1-9. 129-157.
HUNT, D. & TUCKER, M. E. 1995. Stranded parase- MACEACHERN, J. A. & PEMBERTON, S. G. 1997.
quences and the forced regressive wedge systems Ichnology: biogenic utility in genetic stratigraphy.
tract: deposition during base level fall - reply. Canadian Society of Petroleum Geologists 1997
Sedimentary Geology, 95, 147-160. Core Conference, 387—412.
60 S. G. PEMBERTON ET AL.

MACEACHERN, J. A., PEMBERTON, S. G. & RAYCHAUD- Paleontologists and Mineralogists, Special Publi-
HURI, I. 1991. The substrate-controlled Glossifun- cations, Tulsa, Oklahoma, 41, 241-260.
gites ichnofacies and its application to the PEMBERTON, S. G. & FREY, R. W. 1985. The Glossi-
recognition of sequence stratigraphic surfaces: fungites ichnofacies: modern examples from the
subsurface examples from the Cretaceous of the Georgia coast, USA In: CURRAN, H. A. (ed.)
Western Canada Sedimentary Basin, Alberta, Biogenic Structures: Their Use in Interpreting
Canada. In: LECKIE, D. A. , POSAMENTIER, H. W. Depositional Environments. Society of Economic
& LOVELL, R. W. W. (eds) 1991 NUN A Conference Paleontologists and Mineralogists, Special Publi-
on High Resolution Sequence Stratigraphy. Geolo- cations, Tulsa, Oklahoma, 35, 237-259.
gical Association of Canada, Program, Proceed- PEMBERTON, S. G. & MACEACHERN, J. A. 1995. The
ings and Guidebook, Calgary, Alberta, 32-36. sequence stratigraphic significance of trace fossils:
MACEACHERN, J. A., RAYCHAUDHURI, I. & PEMBERTON, examples from the Cretaceous foreland basin of
S. G. 1992. Stratigraphic applications of the Alberta, Canada. In: VAN WAGONER, J. C. &
Glossifungites ichnofacies: delineating discontinu- BERTRAM, G. (eds) Sequence Stratigraphy of
ities in the rock record. In: PEMBERTON, S. G. Foreland Basin Deposits: Outcrop and Subsurface
(ed.) Applications of Ichnology to Petroleum Examples from the Cretaceous of North America.
Exploration: A Core Workshop. Society of American Association of Petroleum Geologists,
Economic Paleontologists and Mineralogists, Memoirs, Tulsa, Oklahoma, 64, 429^75.
Core Workshop Notes, Tulsa, Oklahoma, 17, PEMBERTON, S. G. & MACEACHERN, J. A. 1997. The
169-198. ichnological signature of storm deposits: the use
MACEACHERN, J. A., ZAITLIN, B. A. & PEMBERTON, S. of trace fossils in event stratigraphy. In: BRETT,
G. 1998. High resolution sequence stratigraphy C. E. (ed.) Palaeontological Event Horizons Eco-
of early transgressive incised shoreface and early logical and Evolutionary Implications. Columbia
transgressive valley/embayment deposits of the University Press, New York, 73-109.
Viking Formation, Joffre Field, Alberta, Canada. PEMBERTON, S. G., KOBLUK, D. R., YEO, R. K. & RISK,
American Association of Petroleum Geologists M. J. 1980. Trypanites borings at the Silurian-
Bulletin, 82, 729-756. Devonian disconformity in southern Ontario.
MACEACHERN, J. A., PEMBERTON, S. G. & ZAITLIN, Journal of Paleontology, 54, 1258-1266.
B. A. 1999a. Coarse-grained, shoreline-attached, PEMBERTON, S. G., FLACH P. D. & MOSSOP, G. D. 1982.
marginal marine parasequences of the Viking Trace fossils from the Athabasca Oil Sands,
Formation, Joffre Field, Alberta Canada. In: Alberta, Canada. Science, 217, 825-827.
BERGMAN, K. (ed.) Isolated Marine Sand Bodies: PEMBERTON, S. G., MACEACHERN, J. A. & FREY, R. W.
Sequence Stratigraphic Analysis and Sedimento- 1992a. Trace fossil facies models: environmental
logical Interpretation. Society of Economic and allostratigraphic significance. In: WALKER,
Paleontologists and Mineralogists, Special Publi- R. G. & JAMES, N. (eds) Facies Models: Response
cations, Tulsa, Oklahoma, 64, 273-296. to Sea Level Change. Geological Association of
MACEACHERN, J. A., ZAITLIN, B. A. & PEMBERTON, Canada, St John's, Newfoundland, 47-72.
S. G. 1999b. A sharp-based sandstone succession PEMBERTON, S. G., REINSON, G. E. & MACEACHERN,
of the Viking Formation, Joffre Field, Alberta, J. A. 1992b. Comparative ichnological analysis
Canada: criteria for recognition of transgressively of Late Albian estuarine valley fill and shelf shore-
incised shoreface complexes. Journal of Sedimen- face deposits, Crystal Viking Field, Alberta. In:
tary Research, 69, 876-892. PEMBERTON, S. G. (ed.) Applications of Ichnology
MclLROY, D. 2004. Ichnofabrics and sedimentary to Petroleum Exploration: A Core Workshop.
facies of a tide-dominated delta: Jurassic He Society of Economic Paleontologists and Mineral-
Formation of Kristin Field, Haltenbanken, Off- ogists, Core Workshops, Tulsa, Oklahoma, 17,
shore mid-Norway. In: MC!LROY, D. (ed.) The 291-317.
Application of Ichnology to Palaeoenvironmental PEMBERTON, S. G., SPILA, M. V., PULHAM, A. J.,
and Stratigraphic Analysis. Geological Society, SAUNDERS, T., MACEACHERN, J. A., ROBBINS, D.
London, Special Publications, 228, 237-272. & SINCLAIR, I. 2001. Ichnology and Sedimentology
MELLERE, D. & STEEL, R. 1995. Facies architecture and of Shallow and Marginal Marine Systems: Ben
sequentiality of nearshore and 'shelf sandbodies: Nevis and Avalon Reservoirs, Jeanne D'Arc Basin.
Haystack Mountains Formation, Wyoming, Geological Association of Canada, St John's,
USA. Sedimentology, 42, 551-574. Newfoundland, Short Course Notes, 15.
MORRIS, R. W. & ROLLINS, H. B. 1977. Observations PIANKA, E. R. 1970. On r and k selection. American
on intertidal organism associations of St Cathe- Naturalist, 104, 592-597.
rines Island, Georgia. 1. General description and PILKEY, O. H., BLACKWELDER, B. W., KNEBEL, H. J. &
paleoecological implications. Bulletin American AYERS, M. W. 1981. The Georgia Embayment
Museum of Natural History, 159, 89-128. continental shelf: stratigraphy of a submergence.
NUMMEDAL, D. & SWIFT, D. J. P. 1987. Transgressive Geological Society of America, Bulletin, 92, 52-63.
stratigraphy at sequence-bounding unconformi- PLINT, A. G. 1988. Sharp-based shoreface sequences
ties: some principles derived from Holocene and and 'offshore bars' in the Cardium Formation of
Cretaceous examples. In: NUMMEDAL, D., PILKEY Alberta: their relationship to relative changes in
O. H. & HOWARD, J. D. (eds) Sea-level Fluctuation sea level. In: WILGUS, C. K., HASTINGS, B. S.,
and Coastal Erosion. Society of Economic KENDALL, C. G. ST C., POSAMENTIER, H. W.,
KEY STRATIGRAPHIC SURFACES 61

Ross, C. A. & VAN WAGONER, J. C. (eds) Sea-Level SAUNDERS, T. & PEMBERTON, S. G 1986. Trace Fossils
Changes: An Integrated Approach. Society of and Sedimentology of the Appaloosa Sandstone:
Paleontologists and Mineralogists, Special Publi- Bearpaw-Horseshoe Canyon Formation Transition,
cations, Tulsa, Oklahoma, 42, 357-370. Dorothy, Alberta. Canadian Society of Petroleum
PLINT, A. G., WALKER, R. G. & BERGMAN, K. M. 1986. Geologists, Field Trip Guide Book, Calgary,
Cardium Formation 6. Stratigraphic framework Alberta.
of the Cardium in subsurface. Bulletin of Canadian SAVRDA, C. E. 199la. Teredolites, wood substrates, and
Petroleum Geology, 34, 213-225. sea-level dynamics. Geology, 19, 905-908.
FLINT, A. G., WALKER, R. G. & DUKE, W. L. 1988. SAVRDA, C. E. 1991b. Ichnology in sequence strati-
An outcrop of subsurface correlation of the graphic studies: an example from the lower
Cardium Formation in Alberta. In: JAMES, D. P. Paleocene of Alabama. Palaios, 6, 39-53.
& LECKIE, D. A. (eds) Sequences, Stratigraphy, SAVRDA, C. E. 1995. Ichnologic applications in paleo-
Sedimentology: Surface e. Canadian ceanographic, paleoclimatic, and sea-level studies.
Society of Petroleum Geologists, Memoirs, Palaios, 10, 565-577.
Calgary, Alberta, 15, 167-184. SAVRDA, C. E., OZALAS, K., DEMKO, T. H., HUTCHIN-
POSAMENTIER, H. W. & CHAMBERLAIN, C. J. 1993. SON, R. A. & SCHEIWE, T. D 1993. Log grounds
Sequence Stratigraphic analysis of Viking Forma- and the ichnofossil Teredolites in transgressive
tion lowstand beach deposits at Joarcam field, deposits of the Clayton Formation (Lower Paleo-
Alberta, Canada. In: POSAMENTIER, H. W., cene), western Alabama. Palaios, 8, 311-324.
SUMMERHAYES, C. P., HAQ, B. U. & ALLEN, G. P. SAVRDA, C. E., BROWNING, J. V., KRAWINKLE, H. &
(eds) Stratigraphy and Fades Associations in a HESSELBO, S. P. 2001. Firmground ichnofabrics
Sequence Stratigraphic Framework. International in deepwater sequence stratigraphy, Tertiary
Association of Sedimentologists, Special Publica- clinoform-toe deposits, New Jersey slope. Palaios,
tions, Oxford, 18, 469-485. 16, 294-305.
POSAMENTIER, H. W., ALLEN, G. P., JAMES, D. P. & SCHAFER, W. 1972. Ecology and Palaeoecology of
TESSON, M. 1992. Forced regressions in a sequence Marine Environments. Oliver & Boyd, Edin-
Stratigraphic framework: concepts, examples, and burgh/University of Chicago Press, Chicago.
exploration significance. American Association of SEILACHER, A. 1962. Paleontological studies in turbidite
Petroleum Geologists Bulletin, 76, 1687-1709. sedimentation and erosion. Journal of Geology, 70,
RADDYSH, H. K. 1988. Sedimentology and 'geometry' 227-234.
of the Lower Cretaceous Viking Formation, SEILACHER, A. 1982. Distinctive features of sandy tempes-
Gilby A and B Fields, Albert. In: JAMES, D. P. tites. In: EINSELE G. & SEILACHER, A. (eds) Cyclic and
& LECKIE, D. A. (eds) Sequences, Stratigraphy, Event Stratification. Springer, Berlin, 333-349.
Sedimentology: Surface and Subsurface. Canadian SMITH, R. M. H. 1987. Helical burrow casts of therap-
Society of Petroleum Geologists, Memoirs, sid origin from the Beaufort Group (Permian) of
Calgary, Alberta, 15, 417^30. South Africa. Palaeogeography, Palaeoclimatol-
RANGER, M. J. & PEMBERTON, S. G. 1992. The sedimen- .
tology and ichnology of estuarine point bars in the SOEGAARD, K. & MACEACHERN, J. A. 2003. Integrated
McMurray Formation of the Athabasca Oil Sands sedimentological, ichnological and sequence
Deposit, northeastern Alberta, Canada. In: PEM- Stratigraphic model of a coarse clastic fan delta
BERTON, S. G. (ed.) Applications of Ichnology to reservoir: Middle Jurassic Oseberg Formation,
Petroleum Exploration: A Core Workshop. Society North Sea, Norway. Abstract Volume, AAPG
of Economic Paleontologists and Mineralogists, Annual Convention, Salt Lake City, Utah, May
Core Workshops, Tulsa, Oklahoma, 17, 401-421. 2003, p. A160.
RANGER, M. J. & PEMBERTON, S. G. 1997. Elements of a STAMP, L. D. 1921. On cycles of sedimentation in the
Stratigraphic framework for the McMurray Eocene strata of the Anglo-Franco-Belgian basin.
Formation in south Athabasca. In: PEMBERTON, Geological Magazine, 58, 108-114, 146-157, 194-
S. G. & JAMES, D. P. (eds) Petroleum Geology of 200.
the Cretaceous Mannville Group, Western STELCK, C. R., MACEACHERN, J. A. & PEMBERTON, S. G.
Canada. Canadian Society of Petroleum Geolo- 2000. Foraminiferal biostratigraphic analysis of
gists, Memoirs, Calgary, Alberta, 18, 263-291. the Viking Formation, Kaybob North and
RAYCHAUDHURI, I., BREKKE, H. G., PEMBERTON, S. G. Giroux Lake fields, central Alberta: a comparison
& MACEACHERN, J. A. 1992. Depositional facies with the Hasler Formation biostratigraphy of
and trace fossils of a low wave energy shoreface northeastern British Columbia. Canadian Journal
succession, Albian Viking Formation, Chigwell of Earth Science, 37, 1389-1410.
Field, Alberta, Canada. In: PEMBERTON, S. G. SWIFT, D. J. P. 1968. Coastal erosion and transgressive
(ed.) Applications of Ichnology to Petroleum stratigraphy. Journal of Geology, 76, 444-456.
Exploration: A Core Workshop. Society of TAYLOR, A. M. & GAWTHORPE, R. L. 1993. Application
Economic Paleontologists and Mineralogists, of sequence stratigraphy and trace fossil analysis
Core Workshops, Tulsa, Oklahoma, 17, 319-337. to reservoir description: examples from the
ROY, P. S., THOM, B. G. & WRIGHT, L. D. 1980. Holo- Jurassic of the North Sea. In: PARKER, J. R. (ed.)
cene sequences on an embayed high-energy coast: Petroleum Geology of Northwest Europe, Proceed-
an evolutionary model. Sedimentary Geology, 26, ings of the 4th Conference. Geological Society of
1-19. London, 317-335.
62 S. G. PEMBERTON ET AL.

TAYLOR, A. M., GOLDRING, R. & GOWLAND, S. 2003. WALKER, R. G. & BERGMAN, K. M. 1993. Shannon
Analysis and application of ichnofabric. Earth sandstone in Wyoming: a shelf ridge complex
Science Reviews, 60, 227-259. reinterpreted as lowstand shoreface deposits.
VAN WAGONER, J. C. 1995. Overview of sequence Journal of Sedimentary Petrology, 63, 839-851.
stratigraphy of foreland basin deposits: terminol- WALKER, R. G. & EYLES, C. H. 1991. Topography and
ogy, summary of papers and glossary of sequence significance of a basin wide sequence-bounding
stratigraphy. In: VAN WAGONER, J. C. & BERTRAM, erosion surface in the Cretaceous Cardium
G. T. (eds) Sequence Stratigraphy of Foreland Formation, Alberta, Canada. Journal of Sedimen-
Basins. American Association of Petroleum tary Petrology, 61, 473-496.
Geologists, Memoirs, Tulsa, Oklahoma, 64, 9-21. WALKER, R. G. & JAMES, N. P. (eds) 1992. Facies
VAN WAGONER, J. C., MITCHUM, R. M. JR, CAMPION, Models: Response to Sea Level Change. Geological
K. M. & RAHMANIAN, V. D. 1990. Siliciclastic Association of Canada, St John's, Newfoundland.
Sequences, Stratigraphy in Well Logs, Cores and WALKER, R. G. & WISEMAN, T. R. 1995. Lowstand
Outcrops. American Association of Petroleum shorefaces, transgressive incised shorefaces, and
Geologists, Methods in Exploration, Review 7, forced regressions: examples from the Viking
Tulsa, Oklahoma. Formation, Joarcam area, Alberta. Journal of
VOORHIES, M. R. 1975. Vertebrate burrows. In: FREY, Sedimentary Research, 65, 132-141.
R. W. (ed.) The Study of Trace Fossils. Springer, WETZEL, A. & UCHMAN, A. 1998. Biogenic sedimentary
New York, 325-350. structures in mudstones: an overview. In:
VOSSLER, S. M. & PEMBERTON, S. G. 1988. Skolithos in SCHIEBER, J., ZlMMERLE, W. & SETHI(eds)
the Upper Cretaceous Cardium Formation: an Shales and Mudstones 1. E. Schweizerbart'sche
ichnofossil example of opportunistic ecology. Verlagsbuchhandling, Stuttgart, 351-369.
Lethaia, 21, 351-362. WIGHTMAN, D. M., PEMBERTON, S. G. & SINGH, C. 1987.
VOSSLER, S. M. & PEMBERTON, S. G. 1989. Ichnology Depositional modeling of the Upper Mannville
and paleoecology of the offshore siliciclastic (Lower Cretaceous), central Alberta. Implications
deposits (Cardium Formation). Palaeogeography, for the recognition of brackish water deposits. In:
Palaeoclimatology, Palaeoecology, 74, 217-239. TILLMAN, R. W. & WEBER, K. J. (eds) Reservoir
WALKER, R. G. 1990. Facies modeling and sequence Sedimentology. Society of Economic Paleontolo-
stratigraphy. Journal of Sedimentary Petrology, gists and Mineralogists, Special Publications,
60, 777-786. Tulsa, Oklahoma, 40, 189-220.
Recent and sub-recent microborings from the upwelling area off
Mauritania (West Africa) and their implications for palaeoecology

INGRID GLAUB

Geologisch-Palaontologisches Institut, Senckenberganlage 32-34,


D-60325 Frankfurt am Main, Germany (e-mail: LGlaub@em.uni-frankfurt.de)

Abstract: Late Quaternary dead molluscan shells off Mauritania (West Africa) from the
intertidal zone to 220-300 m water depth were studied for microborings. The study gives
preliminary data on microborings in upwelling areas and their implications for the fossil
record. In total 18 ichnotaxa are described. They are considered to be produced by cyano-
bacteria, green algae, red algae, fungi and foraminifera. The ichnotaxonomic composition
shows minor differences relative to tropical/subtropical areas of investigation. No ichnotaxa
are believed to be specific to upwelling areas. Bathymetrical distribution patterns revealed
different depth ranges for individual ichnotaxa. Relative to areas with similar latitude but
not influenced by upwelling, the absolute depth of the photic zone is shallower. The majority
of ichnotaxa observed are already known from the fossil record (tropical and subtropical
study areas) and should also be expected from ancient upwelling areas.

The term 'microborings' is used for boring The sampling activity of the Meteor Cruise 25/
systems in hard substrates with individual tunnel 1971 off Mauritania (West Africa) provided an
diameters of less than 100 urn. They are com- excellent database from which to obtain an initial
monly found in calcareous substrates, such as impression of the microboring inventory in an
shells and ooids (e.g. Golubic et al 1975; Budd upwelling area (initial documentation in Glaub
& Perkins 1980; Glaub 1994; Bundschuh 2000; et al. 2002), A great amount of Quaternary
Vogel et al. 2000), but are also rarely observed shell material (mainly molluscan shells) was
in phosphatic substrates, such as teeth and collected, ranging from the intertidal down to
bones (e.g. Konigshof & Glaub in press). 300 m water depth. The present study addresses
Research on modern and fossil microborings has the following questions:
intensified since the development of the casting
embedding technique (Golubic et al. 1970). This (1) Does the ichnotaxonomic composition differ
preparation method is based on the filling of from non-upwelling localities?
boring systems by polymer resin and subsequent (2) What information do the samples give on
dissolution of the infested substrate. The resulting bathymetric distribution patterns of micro-
artificial casts allow a three-dimensional visualiza- borings in upwelling areas?
tion of the various borings by SEM. The fully (3) What are the palaeoecological implications?
detailed morphology of microborings yields
information on the producing microbial endoliths
(belonging to cyanobacteria, green algae, red Material and methods
algae, fungi etc.) and provides the basis for com-
parison with other fossil and recent microborings. The cruise 25/1971 of R.V. Meteor collected mol-
Studies of tropical to subtropical modern and luscan shells at 24 stations (Fig. 1). The activity of
fossil microborings are numerous (e.g. Radtke the scientific crew members included additional
et al. 1997 and references therein; Gektidis coastal field trips. Sampling depths range from
1997; Vogel et al. 1999). In contrast, studies on the intertidal to 300m water depth. Samples
modern and fossil microborings from high were taken by dredges, grab samplers, box sam-
latitudes are still rare (Bromley & Hanken 1981; plers and vibrocorers (Einsele et al. 1977). The
Akpan & Farrow 1984; Akpan 1986; Young & present investigation is based on approximately
Nelson 1988; Schmidt & Freiwald 1993; Glaub 150 Holocene molluscan shells, examined by
et al. 2002; Vogel & Marincovich in press). In SEM after application of the casting embedding
this context, microborings from upwelling areas technique (Golubic et al. 1970).
in low latitudes are of great interest. They are The first intention was to focus on Recent
considered to display the influence of tempera- material, because of the good quality hydro-
ture on microboring distribution patterns under graphic data (light, temperature, currents)
a similar angle of light incidence as in tropical available. However, study of the upper layers
to subtropical study areas. of profiles (box sampler, vibrocorers) produced
From\ MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 63-76. 0305-8719/04/S 15.00 © The Geological Society
of London.
64 I. GLAUB

Fig. 1. Left: area of investigation. Right: ship track of sampling stations and bathymetry (in metres).
Below: shelf zonation, stations and sampling equipment (GS, grab sampler; BS, box sampler;
Dr, dredge; VC, vibrocorer). Reproduced with permission from Einsele et al (1977).
MICROBORINGS IN UPWELLING AREAS 65

no results, even from fresh-looking samples. As a given (e.g. 'Tripartitum Form'). It was decided
result the studied substrates derive from to apply the names of existing ichnotaxa to
sampling with a dredge and grab sampler and modern traces for the following reasons: (1) if a
thus represent a time interval of some thousands modern trace and a fossil trace are identical, it
of years (Einsele et al. 1977). This revised is consistent to give both the same name; and
sampling method is, however, a more realistic (2) ichnological studies on modern microborings
analogue for fossil assemblages, which tend to have a high potential to aid palaeoecological
be time-averaged to some degree. reconstructions, even if the producer is
The area of investigation is characterized by unknown. As long as the trace-maker is not
upwelling water masses and by a cold, mainly identified, one has to find a name for the trace
south-directed surface current belonging to the observed and cannot use biotaxonomy. In this
Canary current system. Water temperatures of case the application of established ichnotaxa is
about 15°C to 17°C are reported from 50m more precise and less wordy than informal
water depth and salinity values are about 35.7%o names (e.g. Tripartitum Form', 'Fasciculus
(Mittelstaedt 1972). The area currently belongs dactylus-lis
to the cold-temperate region. Near the coast the should not be used in establishing new ichnotaxa,
suspension load is high. Plankton blooms may because there is still the opportunity to find the
also cause local, temporary reduction of light trace-maker in future and to use a biological
transparency conditions in the water column. binomen. This approach serves to avoid
For the naming of the borings observed, estab- the creation of ichnotaxa based on parallel
lished ichnotaxa were used where available. As taxonomy that could otherwise increase in an
for the remaining taxa, informal names were irresponsible way.

Fig. 2. Artificial casts of microborings in molluscan shells off Mauritania; SEM pictures. Simplified drawings
are added where needed, (a) Caverna pediculata. Station 69 (41 m water depth), (b) Saccomorpha clava. Station
66 (88-89 m water depth), (c) Fasciculus acinosus. Intertidal Baie de St Jean, (d) Fasciculus dactylus together
with Tripartitum Form (thinner tunnels). Station 70 (20m water depth), (e) Fasciculus isp. 1. Station 69 (41 m
water depth), (f) Fasciculus isp. 2. Station 58 (74m water depth), (g) Polyactina araneola. Station 69 (41 m
water depth), (h) Orthogonum fusiferum. Station 70 (20m water depth), (i) Orthogonum lineare together with
Saccomorpha clava. Station 77 (151m water depth).
66 I. GLAUB

Description measure 20-30 um in diameter at their greatest


width and are 30-60 um in length. They are con-
In total, 18 ichnotaxa are characterized by a brief nected to the substrate surface by thin, short,
morphological description, complemented by occasionally ramified tunnels. One of these
taxonomic comments and data on their geo- tunnels is usually 6-7 um wide and 6-7 um
graphical distribution and stratigraphic range. long, whereas the others display 1-2 um in
There are several other borings that are not diameter and are of similar length.
described herein because their rarity precludes Taxonomic comment. Similar modern borings
confident characterization of their morphology. are affiliated to Codiolum-stages of the green
In addition, borings similar to those of endo- alga Gomontia polyrhiza (Lagerheim) Bornet &
lithic bryozoa and endolithic sponges are Flahault 1889.
observed. Distribution in modern and ancient environ-
ments. Records of its modern geographic distri-
Cavernula pediculata Radtke 1991 (Fig. 2a) bution concentrate on the northern hemisphere,
Description. Cavernula pediculata is character- where it inhabits tropical to non-tropical envir-
ized by bag-shaped cavities orientated perpendi- onments (e.g. Nielsen 1972; Radtke 1993; Guiry
cular to the substrate surface. The borings & Nic Dhonncha 2002). The oldest fossil record

Fig. 3. Distribution of microborings at different stations, sorted by water depth.


MICROBORINGS IN UPWELLING AREAS 67

dates back to the Triassic (Schmidt 1992). The 1997) and Mediterranean (Le Campion-
occurrence of Cavernula pediculata off Maurita- Alsumard 1978). The fossil record of Fasciculus
nia is rare (Fig. 3). acinosus dates back to the Permian (Glaub et al.
1999). Fasciculus acinosus is mainly restricted to
Saccomorpha clava Radtke 1991 (Fig. 2b) the shallow euphotic zone II, which corresponds
Description. The boring system consists of club- to the intertidal (sensu Glaub 1994). Observa-
shaped borings (10-30 jam in diameter at their tions in Mauritanian sample material confirm
greatest width, interconnected by small tunnels this environmental restriction and extend the
1 um in diameter). Several varieties of club- geographic distribution of the ichnotaxon to
shaped borings are developed, in some cases up welling areas.
occurring within a single branching system:
some clubs show gradually increasing width Fasciculus dactylus Radtke 1991 (Fig. 2d)
from the proximal area near the substrate surface Description. The Fasciculus dactylus borings are
to the distal tips, whereas others look like a ball bunches of tunnels radiating from an area of
or an ellipsoid on a stem, and still other clubs entry into the substrate. Individual tunnels
are heart-shaped, caused by a small distal measure 5-9 jam in diameter. Tunnels usually
notch. A branching system mainly connects the display rounded tips and can only rarely be
club-shaped borings in their proximal portions, demonstrated to branch. Fasciculus dactylus
but some tunnels branch off at the distal part may also be developed spreading parallel to the
of the club. No collar was observed. substrate surface with smaller (5-6 jam), com-
Taxonomic comment. The ichnotaxon Sacco- monly branching tunnels.
morpha clava Radtke 1991 is used for borings Taxonomic comment. The boring pattern is
morphologically similar to those of the modern known to be produced by Hyella caespitosa
fungus Dodgella priscus Zebrowski 1936. Bornet & Flahault 1889 in the Recent, but may
Distribution in modern and ancient environ- also be produced by other Hyella or Solentia
ments. Dodgella priscus is known from tropical species (cyanobacteria). Fasciculus dactylus was
and non-tropical modern environments down introduced by Radtke (1991).
to 2350m water depth (Hohnk 1969; Zeff & Distribution in modern and ancient environ-
Perkins 1979; Budd & Perkins 1980; Golubic ments. Hyella caespitosa is known from many
et al. 1984). Saccomorpha clava is recorded tropical and non-tropical study areas: NE
from the Triassic (Schmidt 1992), Jurassic Atlantic, SE Pacific, Indian Ocean (Guiry &
(Glaub 1994), Cretaceous (Hofmann 1996) and Nic Dhonncha 2002). Fasciculus dactylus is
Tertiary (Radtke 1991). quite abundant off Mauritania (Fig. 3). It is
Saccomorpha clava borings are very abundant index ichnotaxon for two subzones of the eupho-
in the sampling area off Mauritania. These find- tic zone: Fasciculus acinosus j Fasciculus dactylus -
ings confirm earlier data, which indicate that ichnocoenosis of the upper euphotic zone II for
Saccomorpha clava borings usually become the intertidal and Fasciculus dactylus jPalaeo-
more common with increasing water depth. Sac- conchocelis starmachii - ichnocoenosis for the
comorpha clava is a key ichnotaxon of the index well-illuminated subtidal (Glaub 1994). It is
microboring ichnocoenosis for the aphotic zone known since the Permian (Balog 1996; Glaub
(Glaub 1994; see below). et al. 1999).

Fasciculus acinosus Glaub 1994 (Fig. 2c) Fasciculus isp. 1 (Fig. 2e)
Description. The most characteristic feature of Description. Fasciculus isp. 1 is a unique cluster-
Fasciculus acinosus is that small sphaeroid forming boring system characterized by tunnels
cavities (4-8 um in diameter) are arranged close radiating from a central area of penetration.
to each other, so as to resemble a bunch of The tunnels of some clusters display perpendicu-
grapes. At the distal portion a prominent lar to parallel orientation to the substrate
tunnel is developed (6-8 jam in diameter, 20- surface, whereas other clusters are typified by
40 jim in length). only substrate-parallel tunnels. In both cases,
Taxonomic comment. Fasciculus acinosus three tunnels are typically visible at the entrance
Glaub 1994 is an ichnotaxon displaying a similar area. The tunnels measure 7-9 jim in diameter.
boring pattern to the modern cyanobacterial Tunnel constrictions are present in some cases.
species Hyella balani Lehmann 1903. The commonly developed regular dichotomous
Distribution in modern and ancient environ- branching is distinctive.
ments. Hyella balani is widely reported from the Taxonomic comment. Fasciculus isp. 1 is prob-
Indian Ocean (Guiry & Nic Dhonncha 2002), ably produced by Hyella Stella, a modern cyano-
NE Atlantic (Nielsen 1972), Caribbean (Gektidis bacterium classified in the order Pleurocapsales
68 I. GLAUB

and first described by Al-Thukair & Golubic run parallel to the surface for several lOOum,
(1991). to be connected with another Polyactina araneola
Distribution in modern and ancient environ- boring. Together with boring systems clearly
ments. The first description of Hyella Stella was identifiable as being Polyactina araneola are
based on specimens in modern ooids from the those that might represent initial stages. These
Arabian Gulf, Indian Ocean (Al-Thukair & possible initial stages are stemmed globular
Golubic 1991). Gektidis (1997) registered borings that widen distally, where one to four
Hyella Stella rarely while evaluating study thinner tunnels are developed.
experiments off the Bahamas. A further report Taxonomic comment. Conchyliastrum enderi
from modern environments is given for similar Zebrowski 1936, a lower fungus (order Chytri-
borings (Radtke 1993, Fasciculus sp). For diales), has been proposed as the producer of
records from ancient environments, refer to the boring Polyactina araneola by Radtke (1991).
Green et al. (1988). They reported findings of Distribution in modern and ancient environ-
microborings with preserved organic remains ments. Conchyliastrum is known from tropical
similar to Hyella Stella, named Eohyella dichot- and non-tropical environments (Hohnk 1969;
oma Green et al. 1988. Their studies are based Zeff & Perkins 1979; Budd & Perkins 1980).
on Proterozoic (700-800 Ma) silicified pisoliths The corresponding ichnotaxon Polyactina ara-
in Greenland. neola has a long geological record back to the
Silurian (Bundschuh 2000). Boring activity of
Fasciculus isp. 2 (Fig. 2f) marine endolithic Conchyliastrum species seems
Description. Tunnels 1—4 um in diameter are to increase in the deep euphotic zone and
developed parallel to the substrate surface deeper parts of the water column. Off Maurita-
(rarely observed in perpendicular position). nia, its distribution is exclusive to samples from
Ramification abundantly occurs, following station 58, 59, 68, and 69 (Fig. 3)
angles of up to 90°. The boring system forms den-
sely infested patches. Poorly developed tunnel Orthogonum fusiferum Radtke 1991 (Fig. 2h)
constrictions are visible every 2-8 um. Description. Orthogonum fusiferum borings are
Taxonomic comment. Fasciculus isp. 2 displays characterized by thin tunnels (l-2um in dia-
the characteristics described by Radtke (1991) meter) with typical spindle-shaped enlargements
for the ichnogenus Fasciculus. The producer of (5-7um in diameter, lOum long). Besides this
this boring is unknown, but the size and ramifica- type of development there is a smaller one. It
tion pattern suggest a bacterium. displays tunnel diameters of approximately
Distribution in modern and ancient environ- 0.5 jam and enlargements 4um in diameter and
ments. The record in modern and ancient envir- 6um long. The smaller version shows up to
onments is confused. In some respects, borings four tunnels branching off at the spindle-shaped
of the Tertiary Fasciculus parvus (Radtke 1991, broadening, whereas the bigger variety is
taf. 10, fig. 6), orientated parallel to the substrate unbranched at widenings, or a single tunnel
surface, look similar. In addition, Radtke (1993) may originate.
observed borings called Fasciculus parvus show- Taxonomic comment. Orthogonum fusiferum
ing parallel development in modern substrates Radtke 1991 is the ichnotaxon for borings simi-
off Lee Stocking Island (Bahamas). Fasciculus lar to those of the modern lower fungus Ostraco-
isp. 2 is very abundant off Mauritania (Fig. 3) blabe implexa Bornet & Flahault 1889.
Distribution in modern and ancient environ-
Polyactina araneola Radtke 1991 (Fig. 2g) ments. Ostracoblabe implexa is known from
Description. The Polyactina araneola boring is studies in the southern part of the NW Atlantic
composed of three morphological elements: (e.g. Radtke 1993) and the Mediterranean (Le
entry tunnel; distal globular enlargement; and Campion-Alsumard 1978). According to studies
thinner tunnels with tapering ends. The entry of Le Campion-Alsumard, the boring activity
tunnel (8-10 jam in diameter) is orientated per- of Ostracoblabe implexa extends down to 200 m
pendicular to the substrate surface. Deeper in water depth. Orthogonum fusiferum is only
the substrate the tunnel extends its diameter to rarely documented off Mauritania (Fig. 3). As
a nearly globular enlargement 30-40 jam in dia- for ancient environments, the oldest record of
meter. All around this widening, thinner tunnels Orthogonum fusiferum is that of Bundschuh
(8-10 jam in diameter) radiate in different direc- (2000) from the Silurian.
tions. These tunnels are characterized by
gradually decreasing diameters and tapering Orthogonum lineare Glaub 1994 (Fig. 2i)
ends 1-2 um in diameter. Many tunnels turn Description. Tubular borings, 5-15 um in dia-
back towards the substrate surface, where they meter, display a highly organized boring
MICROBORINGS IN UPWELLING AREAS 69

system, orientated parallel to the substrate sur- Hofmann (1996, taf. X, fig. 5-6, taf. XI, fig. 1)
face, dominated by ramification angles of 90°. from the Cretaceous of northern Europe.
Tunnels running parallel to each other are Orthogonum lineare is, together with Sacco-
common. morpha clava, a key ichnotaxon for recognition
Taxonomic comment. The ichnotaxon Ortho- of the aphotic zone ichnocoenosis (of Glaub
gonum lineare was erected by Glaub (1994) for 1994). Off Mauritania, Orthogonum lineare is
fossil borings. Its producer is unknown (see common in samples from station 77 (151m)
below). (Fig. 3).
Distribution in modern and ancient environ-
ments. Boring patterns similar to that of Ortho- Orthogonum spinosum Radtke 1991 (Fig. 4a)
gonum lineare are reported from several places Description. Tubular tunnels, characterized by
under numerous informal names - modern and diameters of 10-3 8 um and mainly rectangular
fossil, topical and non-tropical - mainly from ramifications. The tunnels are in some cases
the aphotic zone. Reports on modern borings developed in some distance to the substrate
are given by: Zeff& Perkins (1979) from the Car- surface and are in that case abundantly con-
ibbean at 435-775 m as Tubular branching bor- nected with it by perpendicular tunnel junctions.
ings'; Budd & Perkins (1980) from the Caribbean They display a conspicuous boring surface,
at 119-530 m as Tubular-Form F; Schmidt & which is surrounded by short hair-like extensions
Freiwald (1993) from Norway as Type C at (1-2 jam in diameter, 10-15 um maximum length)
20-90 m; and Krutschinna (1997) from Norway all around the tunnels, but in some cases
at 230-300 m as Tubulare Spur 1'. Fossil forms restricted to distinct portions of the tunnel
have been described by Glaub (1994) from the (Fig.4a).
Jurassic of Europe (inferred to be from the Taxonomic comment. The first description of
deep euphotic zone to aphotic zone) and by Orthogonum spinosum was given by Radtke

Fig. 4. Artificial casts of microborings in molluscan shells off Mauritania; SEM pictures. Simplified drawings
are added where needed, (a) Orthogonum spinosum. Station 66 (88-89 m water depth), (b) Orthogonum tubulare.
Station 77 (151 m water depth), (c) Reticulina elegans. Station 69 (41 m water depth), (d) Scolecia filosa. Station
69 (41 m water depth), (e) Scolecia serrata. Station 69 (41 m water depth), (f) Globodendrina monile; thinner
tunnels belong to other taxa. Station 70 (20m water depth), (g) Tripartite Form. Station 69 (41 m water depth).
70 I. GLAUB

(1991) based on Tertiary samples. Similar ally organized green alga Ostreobium quekettii
modern borings are known, but yet not attribu- Bornet & Flahault 1889.
ted to a distinct producer. Distribution in modern and ancient environ-
Distribution in modern and ancient environ- ments. A long list of distributional data exists
ments. Spinose borings, usually characterized for Ostreobium quekettii, which indicates nearly
by longer hairs than described above, are very global distribution. This endolithic chlorophyte
abundant in modern waters, mainly in deep is eurybathic and can survive in euphotic inter-
water (e.g. Zeff& Perkins 1979; Budd & Perkins tidal habitats (shaded positions) as well as
1980; Glaub 1994; Krutschinna 1997). In under dysphotic conditions (deepest record in
contrast, records of fossil spinose borings are 220 m water depth near the Bahamas at approxi-
rare: Radtke (1991, Tertiary), Schmidt (1992, mately 0.01 % of surface light; Fredj & Falconetti
Triassic). 1977). The maximum density of occurrence is
recorded from the deep euphotic zone. The corre-
Orthogonum tubulare Radtke 1991 (Fig. 4b) sponding ichnotaxon Reticulina elegans is known
Description. The boring systems usually spread from Silurian times (Bundschuh 2000). Reticu-
parallel to the substrate surface, branching lina elegans is key ichnotaxon of the index ichno-
rectangular. The main characteristic feature is coenosis for the deep euphotic zone (Glaub
the extremely varying tunnel diameter. At the 1994).
point of ramification irregularly-shaped enlarge-
ments are developed. The diameters of the inter- Scolecia filosa Radtke 1991 (Fig. 4d)
connecting tunnels measure 10-30|im, whereas Description. Scolecia filosa is characterized by
the enlargements reach more than 40 jim in dia- curved running tunnels of 1-3 jam in diameter.
meter. The tunnel surface of both elements, The borings may form dense networks com-
which is more clearly visible at the enlargements, monly displaying loops. Ramification rarely
is verrucose. occurs but, if observed, it shows the typical X-
Taxonomic comment. The ichnotaxon Ortho- or Y-ramification patterns.
gonum tubulare Radtke 1991 was established Taxonomic comment. The ichnotaxon Scolecia
from Tertiary study material. The biological filosa was named by Radtke (1991). Its boring
identity of its producer is as yet unknown. pattern is comparable with that of the modern
Distribution in modern and ancient environments. cyanobacterium Plectonema terebrans Bornet &
Borings with similar morphology but smaller Flahault 1889.
dimensions have been documented in foramini- Distribution in modern and ancient environ-
feran tests from the Atlantic Ocean at 2195- ments. In modern environments Plectonema
2323m water depth (Golubic et al. 1984). As for terebrans is abundantly observed. Its bathymetri-
the fossil record, besides the description of Ortho- cal distribution ranges from the intertidal to
gonum tubulare from the Tertiary (Radtke 1991), it 370m (Lukas 1978). It has been suggested that
is known from the Cretaceous (Hofmann 1996) Plectonema terebrans may live as a facultative
and the Jurassic (Glaub 1994). Off Mauritania, heterotroph (Glaub 1994; Glaub et al. 2001).
Orthogonum tubulare borings were observed in Fossil borings classified as Scolecia filosa are
samples from five stations (Fig. 3). known from deposits as old as the Silurian
(Glaub et al. 1999; Bundschuh 2000). The general
Reticulina elegans (Radtke) Bundschuh 2000 observation that the boring pattern described is
(Fig. 4c) widespread in modern as well as in ancient envir-
Description. Reticulina elegans borings occur as onments is confirmed by the findings off
densely ramified tunnel networks. They are Mauritania. Also in good accordance with earlier
characterized by dichotomous branching, which observations (see discussion in Glaub et al. 2001)
gives the boring system a zigzag pattern. The is the fact that these borings represent bathy-
tunnel diameter ranges from 2um to 5|im. In metrically the deepest cyanobacterium off
the Mauritanian samples studied, Reticulina Mauritania.
elegans was generally developed in the substrate
around sponge macroborings. There is no clear Scolecia serrata Radtke 1991 (Fig. 4e)
explanation for this, but it seems as though Description. The boring system is identified by
Reticulina elegans borings were developed after thin tunnels 0.5-2um in diameter, running in a
the sponge tissue decomposition. slightly zigzag pattern. The borings may form
Taxonomic comment. The characteristics of mat-like patches, caused by meandering tunnels
the ichnotaxon Reticulina elegans (Radtke) developed close to each other. In general, the
Bundschuh 2000 correspond to those of the boring systems are arranged parallel to the
boring pattern known from the modern siphon- substrate surface.
MICROBORINGS IN UPWELLING AREAS 71

Taxonomic comment. Boring systems in fossil Tripartitum Form (Fig. 4g)


substrates developed similarly to those described Description. The boring system is composed of
above were called S cole da serrata by Radtke three parts: (a) interwoven tunnels 1-8 um in dia-
(1991). According to her study the producer is meter; with (b) episodically integrated ellipsoidal
unknown, but most probably a bacterium. to globular enlargements 3-12um in diameter;
Distribution in modern and ancient environ- and (c) in some cases connected to perpendicu-
ments. Scolecia serrata is known from different larly orientated clusters of three or more tunnels
modern environments. Radtke (1991) gives (8-20 um in diameter) with slightly pointed tips.
citations for observations in the Atlantic and Taxonomic comment. The informal name
the Pacific Ocean. According to Radtke (1993), Tripartitum Form' was chosen to describe
Scolecia serrata is found as deep as 1600m boring patterns known from the endolithic con-
water depth. Although modern borings like chocelis stage of the modern rhodophyte taxa
Scolecia serrata are recorded from many places Porphyra and Bangia. Variations of tunnel dia-
(tropical and non-tropical), its taxonomic affinity meters suggest that different species may have
remains unknown. Findings in ancient deposits caused the borings observed. Chain-like arrange-
are so far restricted to the Tertiary Period ments of ellipsoidal to globular widenings, which
(Radtke 1991; Glaub et al 2002). are typical for the fossil biotaxon Palaeoconcho-
celis starmachii Campbell et al. 1979 and its
Globodendrina monile Plewes et al. 1993 (Fig. 4f) modern counterpart Porphyra nereocystis Ander-
Description. The boring consists of a spheroid, son 1892 (in Blankinship & Keeler 1892), were
verrucose swelling (60-100 um in diameter) observed only in one sample of the Mauritanian
from which a single tunnel (20-30 um in study material.
diameter) branches off, running parallel to the Distribution in modern and ancient environ-
substrate surface. This tunnel continues ments. As for endolithic red algae, more than
branching at low angles, forming anastomosing 100 modern Porphyra and some Bangia species
tunnel fusions in some cases. The branching are known (Guiry & Nic Dhonncha 2002),
system developed on one side of the globular but their endolithic conchocelis stages are
swelling may be 400 um wide and 500 um long. usually not documented with the help of the
Hair-like extensions are observed all over the casting embedding technique. Therefore the
boring system and are also developed as comparison of modern and ancient red algal
connections to the substrate surface. borings is usually performed only for Porphyra
Taxonomic comment. The taxon Globoden- nereocystis and Palaeoconchocelis starmachii.
drina monile was erected by Plewes et al. (1993) Borings, so far summarized under the term
for borings of a foraminifer. The modern analo- Tripartitum Form, occur abundantly off Mauri-
gue is so far not identified at genus or species tania (Fig. 3).
level (Cherchi & Schroeder 1991).
Distribution in modern and ancient environ- Pygmy Form (Fig. 4h)
ments. There are a few reports on modern Description. Thin tunnels approximately 0.3—
boring foraminifera (e.g. Smyth 1988; Cherchi 0.5um in diameter running parallel to the sub-
& Schroeder 1991; Freiwald & Schonfeld 1996). strate surface. They display ramification at differ-
The X-ray documentation of Cherchi & Schroe- ent angles, mainly between 60° and 90°, and are
der (1991) shows both the globular boring slightly curved.
system and the producing foraminifera within, Taxonomic comment. The name 'Pygmy Form'
although fine details such as hair-like extensions was chosen according to the term for those bor-
are not visible. Their sample material derives ings observed near the Bahamas (Radtke 1993).
from 180m water depth off Scotland. Hitherto, The boring organism is yet not identified, but a
the casting embedding technique has rarely been bacterium is suggested, owing to the extremely
used for documentation of modern foraminifera small tunnel diameter.
and their borings, which makes comparison of Distribution in modern and ancient environ-
modern and fossil boreholes hard to access. The ments. The Pygmy Form is commonly observed
data on the fossil record of Globodendrina in the Mauritanian samples. It holds the deepest
monile date back to the Jurassic (Plewes et al. boring record in the Mauritanian samples
1993; Glaub 1994, see 'Semidendrina Form') or (station 62, 220-300 m). Together with Radtke's
even the Carboniferous (personal communica- observation near the Bahamas this seems to be
tion K. P. Vogel, Frankfurt). Future studies only the second record. No fossil record of the
should focus on determining its geographic distri- Pygmy Form is known, and it is most probable
bution more precisely as it has considerable that the extreme small tunnel diameter reduces
potential to aid palaeoecological reconstructions. its preservation potential.
72 I. GLAUB

Echinoid Form (Fig. 4i) cyanobacteria, two to green algae, one to a red
Description. The boring system consists of an alga).
entry tunnel connected with a fan-like widening.
The entry tunnel measures 15-30 um in diameter
and is up to 60-70 um long. At the point of entry Borings and bathymetric interpretation
it is usually orientated at a high angle to the
substrate surface, and is recurved back to the Microborings have been demonstrated to be
surface. Near the latter contact to the surface important tools for palaeobathymetric recon-
the tunnel broadens to a flat, lobed, fan-like struction (Glaub 1994, 1999; Vogel et al. 1995;
widening (60-120 um wide and 45-80 um long). Gektidis 1997). This scheme is based on depth-
The Echinoid Form is characterized by a slightly related assemblages of traces made by micro-
verrucose surface texture to the boring and has endoliths. Such assemblages have enabled the
hair-like extensions, which may occur all over identification of subzones of the euphotic zone
the boring, including the contact between the and the aphotic zone. Subsequent studies have
fan-like area and the substrate surface. demonstrated the applicability of the scheme to
Taxonomic comment. The informal name sedimentary basins from the Palaeozoic onwards
'Echinoid Form' was used by Radtke (1993) for (e.g. Vogel et al. 1995, 1999; Glaub & Bundschuh
similar modern borings. So far nothing is 1997; Bundschuh 2000).
known about either the taxonomic affinity of The euphotic zone covers the supratidal, the
the borer or the fossil record of the trace. intertidal and the well-illuminated sub tidal. The
Distribution in modern and ancient environ- lower limit of the euphotic zone is defined as
ments. Besides its occurrence off Mauritania, the depth where the surface light (between 350
the Echinoid Form is so far only recorded from and 700 nm wavelength) is reduced to approxi-
modern environments near Mexico (Giinther mately 1% (for definition and further literature
1990) and near the Bahamas (Radtke 1993). see Glaub 1994). This depth is almost identical
with the depth at which the photosynthesis rate
equals the respiration rate of photoautotrophic
Interpretation organisms.
Photoautotrophic endoliths dominate the
Ichnotaxonomic composition euphotic zone (e.g. Golubic et al. 1975), many
of them being obligate photoautotrophs. How-
The ichnotaxonomic composition is in good ever, one living cyanobacterial species (Plecto-
accordance with data from tropical study areas nema terebrans, which produces a boring
(e.g. Puerto Rico, Budd & Perkins 1980; Baha- similar to the ichnotaxon Scolecia filosa) and
mas, Radtke 1993). All ichnotaxa described one living chlorophycean species (Ostreobium
were also found by Radtke in the Bahamas quekettii, with its boring being comparable to
except Orthogonum spinosum. In contrast to her the ichnotaxon Reticulina elegans) can cope
studies, borings similar to the following ichno- with less than 1% of surface light (between 350
taxa are lacking off Mauritania: Scolecia maean- and 700 nm wavelength). Consequently, they
dria, Fasciculus parvus, Fasciculus grandis, are not restricted to the euphotic zone (Glaub
Eurygonum nodosum and Saccomorpha sphaer- 1994, Gektidis 1997, Vogel et al. 1999, Glaub
ula. Comparison with high-latitude assemblages et al. 2001). It demands further investigations
is not possible, because of the small database to understand how Plectonema terebrans and
available. As a result, none of the ichnotaxa Ostreobium quekettii can exist under these low
observed can be considered to be restricted to light conditions (see discussion in Glaub 1994).
the up welling area. On the basis of microboring assemblages, a
The taxonomic inventory of inferred trace- subdivision of the euphotic zone is given. The
makers is dominated by heterotrophs. Three ich- shallow euphotic zone I is equivalent to the
notaxa are considered to be produced by fungi, supratidal zone. In modern environments this is
whereas one is believed to originate from forami- dominated by cyanobacteria capable of protect-
niferal boring and seven are of unknown origin ing themselves from sunburn damage by sheath
(most probably bacterial or fungal hetero- pigmentation. No index ichnocoenosis for micro-
trophs). In addition, the endolith producing borings was defined for this zone, because sam-
Scolecia filosa borings is suspected to live as a pling has not been extended to this euphotic
facultative heterotroph, among other reasons subzone so far. The shallow euphotic zone II
because of its deep occurrence (see discussion in corresponds to the intertidal zone; its microbor-
Glaub 1994, Glaub et al. 2001). Six ichnotaxa ing index ichnocoenosis is called the Fasciculus
are attributed to photoautotrophs (three to acinosus I Fasciculus dactylus-ichnocoenosis. It is
MICROBORINGS IN UPWELLING AREAS 73

characterized mainly by cyanobacterial borings developed perpendicular to the substrate surface,


orientated perpendicular to the substrate surface, and a decrease in cyanobacterial borings is
which are able to deal with the changing hydro- observed compared with the intertidal zone. In
graphic conditions in the intertidal. For shallow contrast to data from tropical study areas (sum-
euphotic zones I and II the endolithic commu- marized in Glaub 1994), off Mauritania hetero-
nities are influenced by both photic and hydro- trophs dominate over photoautotrophs (one
dynamic factors. For this reason, the photic cyanobacterium, two algae, six heterotrophs
and hydrodynamic zonation units are identical. and the probably facultative heterotroph
Shallow euphotic zone III encompasses well- producer of Scolecia filosa).
illuminated sub tidal settings. In contrast to the The typical change in microendolithic associa-
aforementioned subzones, its microboring tion from the shallow euphotic zone to the deep
community is characterized by cyanobacteria euphotic zone is also hard to follow off Maurita-
along with borings of red and green algae. The nia. Only one sample from station 55 (25m)
predominant boring pattern is perpendicular to clearly shows the Palaeoconchocelis starmachii/
the substrate surface. The Fasciculus dactylus/ Reticulina elegans-ichnocoQnosis of the deep
Palaeoconchocelis starmachii-ichnocoQnosis is euphotic zone with parallel development of
the corresponding index assemblage. Fasciculus dactylus.
The deep euphotic zone is defined as the less- Samples from 41 m water depth down to 74m
illuminated part of the euphotic zone down to water depth display a rather confused distribu-
approximately 1% of surface light. The endo- tion pattern of microborings. These stations are
lithic community is dominated by red and green characterized by Einsele et al. (1977) as mixed
algae. Boring parallel to the substrate surface assemblages. Older shell material indicating
typifies this association. The index assemblage lower water depth is mixed with remains of
of this euphotic subzone is called Palaeo- modern shell-bearing organisms. This fact is
conchocelis starmachii/Reticulina elegans- confirmed by the findings of the present study.
ichnocoenosis. Several samples show a mixture of ichnocoenoses
The dysphotic zone extends from the 1 % level from the shallow euphotic zone, the deep eupho-
to about 0.01% or 0.001% of surface light. The tic zone and probably the dysphotic zone, which
most characteristic feature of the dysphotic reflect exposure to different light conditions.
zone is the dominance of chemoheterotrophic Despite this diffuse picture, the occurrence of
endoliths (mainly fungi), which are accompanied some microboring taxa is restricted to this bathy-
by Scolecia filosa and Reticulina elegans. The metric interval (the borings affiliated to hetero-
endolithic assemblage of the aphotic zone con- trophs: Polyactina araneola and the Echinoid
sists of heterotrophs only. The index assemblage Form), whereas others have their shallowest
for this zone is called the Saccomorpha clava/ occurrence in this mixing zone (the borings
Orthogonum lineare-ichnocoenosis. affiliated to heterotrophs: Orthogonum spinosum
and Orthogonum tubulare) or disappear here
(the borings affiliated to photoautotrophs:
Application of the bathymetric model Cavernula pediculata, Fasciculus dactylus. Fasci-
culus isp. 1, Reticulina elegans and Tripartitum
Several samples collected in the intertidal zone by or
the land excursion related to the Meteor cruise lineare, which are considered to have hetero-
were considered to be recent. Their microboring trophic trace-makers, are observed in samples
assemblage shows the typical elements of the from shallower stations as well as in those from
Fasciculus acinosus/Fasciculus dactylus-ichno- deeper stations, but not in the mixing zone.
coenosis (Glaub 1994). In total, 11 microboring The zone between 41 m and 74 m shows the
ichnotaxa are observed (three affiliated to cyano- usually observed displacement of photoauto-
bacteria, three to algae, four to heterotrophs and trophs by heterotrophs, but there is no clear
the probably facultative heterotroph producer of gradient visible within this zone. The lower
Scolecia filosa) (Fig. 3). Thus the identification of boundary of the euphotic zone is expected most
the upper euphotic zone II by means of micro- probably at the top of the mixing zone or
borings is clearly applicable. between the 25m and the 41m station. The
Demonstration of upper euphotic zone III parallel development of Fasciculus dactylus is
is, however, more problematic. Only three considered as an indicator of this boundary.
samples from station 70 (20 m) clearly show the Further support derives from light measurements
typical composition of the Fasciculus dactylus/ (Morel 1982) (see below).
Palaeoconchocelis starmachii-ichnocoQnosis. For Samples deriving from 88-89 m (station 66)
further confirmation, Fasciculus dactylus is down to sampling depth 220-300 m (station 62)
74 I. GLAUB

clearly display the characteristics of the Sacco- ichnotaxonomy with them and by their competent
morpha clava/Orthogonum linear e-ichnocoenosis. suggestions. An anonymous reviewer also deserves
A variety of borings known from heterotrophs my appreciation.
(mainly fungi) were identified, associated by
Scolecia filosa (down to 148-149 m). Two sam-
ples from 74m (station 58) displaying no features References
of reworking, in contrast to the other samples
from this depth, indicate aphotic conditions. AKPAN, E. B. 1986. Depth distribution of endolithic
Thus the boundary between the dysphotic zone algae from the Firth of Clay: implications for
and the aphotic zone is inferred between 74m delineation and subdivision of the photic zone.
and 88-89 m water depth. Journal of the Marine Biological Association of
The absolute depths estimated for the bound- the United Kingdom, 66, 269-275.
aries between the individual zones and subzones AKPAN, E. B. & FARROW, G. E. 1984. Shell-boring
algae on the Scottish continental shelf: identifica-
are in good accordance with measurements of the
tion, distribution, bathymetric zonation. Trans-
optical properties off Mauritania (Morel 1982). actions of the Royal Society of Edinburgh: Earth
Remarkably, they are shallower than those Science, 75, 1-12.
found in other areas of similar latitudinal posi- AL-THUKAIR, A. A. & GOLUBIC, S. 1991. Five new
tion. One explanation for this observation is a Hyella species from the Arabian Gulf. Algological
high load of planktic content and resuspended Studies, 64, 167-214.
sediment. Morel's long-term light measurements BALOG, S.-J. 1996. Boring thallophytes in some
off Mauritania record 10-60 m for the lower limit Permian and Triassic reefs: bathymetry and
of the euphotic zone. According to Jerlov's water bioerosion. In: REITNER, J., NEUWEILER, F. &
type III (Jerlov 1976) the euphotic zone extends GUNKEL, F. (eds) Global and Regional Controls
on Biogenic Sedimentation. I. Reef evolution. Got-
to approximately 30 m water depth, whereas the tinger Arbeiten fur Geologic und Palaontologie,
aphotic zone onset is in approximately 70m Research Reports, Sb2, 305-309.
water depth. BLANKINSHIP, J. W. & KEELER, C. A. 1892. On the
natural history of the Farallon Islands. Zoe, 3,
14^173.
Geological significance BORNET, M. E. & FLAHAULT, C. 1889. Sur quelque
plantes vivant dans le test calcaire des mollusque.
Most of the ichnotaxa observed in this study are Bulletin de la Societe Botaniques de France, 36,
known from the fossil record (14 taxa). Only for 147-176.
the following ichnotaxa are there no clear data or BROMLEY, R. G. & HANKEN, N.-M. 1981. Shallow
marine bioerosion at Vard0, arctic Norway.
no data at all from ancient deposits: Fasciculus Bulletin of the Geological Society of Denmark, 29,
isp. 2, Tripartitum Form, Pygmy Form and 103-109.
Echinoid Form. The high number of Maurita- BUDD, D. A. & PERKINS, R. D. 1980. Bathymetric
nian ichnotaxa with fossil equivalents makes zonation and paleoecological significance of
the results meaningful for palaeoenvironmental microborings in Puerto Rican shelf and slope
studies. As for palaeo-depth reconstructions by sediments. Journal of Sedimentary Petrology, 50,
microborings, the main requirement is the 881-903.
autochthony of the samples studied, because BUNDSCHUH, M. 2000. Silurische Mikrobohrspuren.
only in this case do borings of photoautotrophic Ihre Beschreibung und Verteilung in verschie-
denen Faziesraumen (Schweden, Litauen und
endoliths reflect light conditions at the depth at den USA). PhD thesis, Geologic, Johann Wolf-
which bioerosion occurred. The studies of gang Goethe-Universitat, Frankfurt am Main.
Mauritanian samples demonstrate that investiga- CAMPBELL, S. E., KAZMIERCZAK, L. & GOLUBIC, S.
tions on microboring itself can aid in identifica- 1979. Palaeoconchocelis starmachii gen.n., sp.n.,
tion of allochthonous samples, if mixing of an endolithic Rhodophyte (Bangiaceae) from the
ichnocoenoses is observed. Silurian of Poland. Ada Palaeontologica Polonica,
24, 405^08.
I am very indebted to D. Herm, K. Vogel, M. CHERCHI, A. & SCHROEDER, R. 1991. Perforations
Gektidis and G. Radtke for information, discussion branchues dues a Foraminiferes cryptobiotiques
and for making available important samples. My dans des coquilles actuelles et fossiles. Compte
gratitude is extended to A. Mutter, O. Sagert, R. Rendus de I'Academic des Sciences, Serie II, 312,
Schade and J. Tochtenhagen for their technical 111-115.
assistance. The investigations were kindly funded EINSELE, G., ELOUARD, P., HERM, D., KOGLER, F. C. &
by the German Research Foundation (DFG-Projekt SCHWARZ, H. U. 1977. Source and biofacies of
Vo 90/21). Sincere thanks go to D. Mcllroy and Late Quaternary sediments in relation to sea
R. Bromley. The manuscript was very much level on the shelf off Mauritania, West Africa.
improved by an intensive, friendly discussion on 'Meteor'-Forschungsergebnisse, C 26, 1^3.
MICROBORINGS IN UPWELLING AREAS 75

FREDJ, G. & FALCONETTI, C. 1977. Sur la presence GUIRY, M. D. & NIC DHONNCHA, E. 2002. AlgaeBase.
d'Algues filamenteuses perforantes dans le test www. algaebase. org
des Gryphus vitreus (BORN) (Brachiopodes, GUNTHER, A. 1990. Distribution and bathymetric zona-
Terebratulides) de la limite inferieure du plateau tion of shell-boring endoliths in recent reefs and
continental mediterraneen. Compte Rendus de shelf environments. Cozumel, Yucatan (Mexico).
I'Academic des Sciences, Serie D, 284, 1167—1170. Fades, 22, 233-262.
FREIWALD, A. & SCHONFELD, J. 1996. Substrate pitting HOFMANN, K. 1996. Die mikro-endolithischen Spuren-
and boring pattern of Hyrrokkin sarcophaga fossilien der borealen Oberkreide Nordwest-
Cedberg, 1994 (Foraminifera) in a modern deep- Europas. Geologisches Jahrbuch, A 136, 1-151.
water coral reef mound. Marine Micropaleontol- HOHNK, W. 1969. Uber den pilzlichen Befall kalkiger
ogy, 28, 199-207. Hartteile von Meerestieren. Bericht Deutsche Wis-
GEKTIDIS, M. 1997. Vorkommen, Okologie und Taxo- senschaftliche Kommission fur Meeresforschung,
nomie von Mikrobohrorganismen in ausgewahl- 20, 129-140.
ten Riffbereichen um die Inseln Lee Stocking JERLOV, N. G. 1976. Marine Optics. Elsevier, Amster-
Island (Bahamas) und One Tree Island (Austra- dam, 1-231.
lien). PhD thesis, Biologic, Johann Wolfgang KONIGSHOF, P. & GLAUB, I. (in press). Traces of
Goethe-Universitat, Frankfurt am Main. microboring organisms in Palaeozoic conodont
GLAUB, I. 1994. Mikrobohrspuren in ausgewahlten elements. Geobios.
Ablagerungsraumen des europaischen Jura KRUTSCHINNA, J. 1997. Untersuchungen an der
und der Unterkreide (Klassifikation und Paloko- Tiefwasserkoralle Lophelia pertusa in verschie-
logie). Courier Forschungsinstitut Senckenberg, denen pra- und postmortalen Stadien unter
174, 1-324. besonderer Beriicksichtigung des mikroendo-
GLAUB, I. 1999. Microborings and bathymetrical lithischen Befalls. Diploma thesis, FB Biologic,
reconstructions. Bulletin of the Geological Society Johann Wolfgang Goethe-Universitat, Frankfurt
of Denmark, 45, 143-146. am Main.
GLAUB, I. & BUNDSCHUH, M. 1997. Comparative LE CAMPION-ALSUMARD, T. 1978. Les eyanophycees
studies on Silurian and Jurassic/Lower Cretaceous endolithes marines. Systematique, ultrastructure,
microborings. Courier Forschungsinstitut Sencken- ecologie et biodestruction. PhD thesis, Universite
berg, 201, 123-135. d'Aix-Marseille II, Marseille.
GLAUB, I., BALOG, S.-J. et al 1999. Euendolithic cyano- LEHMANN, E. 1903. Uber Hyella balani nov. spec. Nyt
bacteria/cyanophyta and their traces in earth Magazinfor Naturvidenskap, 41, 77-87.
history. In: CHARPY, L. & LARKUM, A. W. D. LUKAS, K. J. 1978. Depth distribution and form among
(eds) Marine Cyanobacteria, Bulletin de I'lnstitut common microboring algae from the Florida
Oceanographique, Monaco, Numero Special, 19, continental shelf. Geological Society of America,
135-142. Abstracts, 10, 448.
GLAUB, L, VOGEL, K. & GEKTIDIS, M. 2001. The role of MITTELSTAEDT, E. 1972. Der hydrographische Aufbau
modern and fossil cyanobacterial borings in und die zeitliche Variabilitat der Schichtung und
bioerosion and bathymetry. Ichnos, 8, 185-195. Stromung im nordwestafrikanischen Auftriebsge-
GLAUB, L, GEKTIDIS, M. & VOGEL, K. 2002. Micro- biet im Fruhjahr 1968. 'Meteor'-Forschungsergeb-
borings from different North Atlantic shelf areas nisse, A, 11, 1—57.
- Variability of the euphotic zone extension and MOREL, A. 1982. Optical properties and radiant energy
implications for paleodepth reconstructions. in the waters of the Guinea Dome and the
Courier Forschungsinstitut Senckenberg, 237, 25- Mauritanian upwelling area in relation to primary
37. production. Rapports et Proces-Verbaux des
GOLUBIC, S., BRENT, G. & LE CAMPION-ALSUMARD, T. Reunions, 180, 94-107.
1970. Scanning electron microscopy of endolithic NIELSEN, R. 1972. A study of shell-boring marine algae
algae and fungi using a multipurpose casting- around the Danish island Laeso. Botanisk
embedding technique. Lethaia, 3, 203-209. Tidsskrift, 67, 245-269.
GOLUBIC, S., PERKINS, R. D. & LUKAS, K. J. 1975. PLEWES, C. R., PALMER, T. J. & HAYNES, J. R. 1993. A
Boring microorganisms and microborings in boring foraminiferan from the Upper Jurassic
carbonate substrates. In: FREY, R. W. (ed.) of England and Northern France. Journal of
The Study of Trace Fossils. Springer, Berlin, Micropalaeontology, 12, 83-89.
229-259. RADTKE, G. 1991. Die mikroendolithischen Spuren-
GOLUBIC, S., CAMPBELL, S. E., DROBNE, K., CAMERON, fossilien im Alt-Tertiar West-Europas und ihre
B., BALSAM, W. L., CILMERAN, F. & DUBOIS, L. palokologische Bedeutung. Courier Forschungsin-
1984. Microbial endoliths: a benthic overprint in stitut Senckenberg, 138, 1-185.
the sedimentary record, and a palaeobathymetric RADTKE, G. 1993. The distribution of microborings in
cross-reference with foraminifera. Journal of molluscan shells from recent reef environments
Paleontology, 58, 351-361. at Lee Stocking Island, Bahamas. Fades, 29, 81-
GREEN, J. W., KNOLL, A. H. & SWETT, K. 1988. 92.
Microfossils from ooliths and pisoliths of the RADTKE, G., HOFMANN, K. & GOLUBIC, S. 1997. A
Upper Proterozoic Eleonore Bay Group, Central bibliographic overview of micro- and macroscopic
East Greenland. Journal of Paleontology, 62, bioerosion. Courier Forschungsinstitut Sencken-
835-852. berg, 201, 307-340.
76 I. GLAUB

SCHMIDT, H. 1992. Mikrobohrspuren ausgewahlter VOGEL, K., BALOG, S.-J., BUNDSCHUH, M., GEKTIDIS,
Faziesbereiche der tethyalen und germanischen M., GLAUB, I., KRUTSCHINNA, J. & RADTKE, G.
Trias (Beschreibung, Vergleich und bathy- 1999. Bathymetrical studies in fossil reefs with
metrischen Interpretation). Frankfurter geowis- microendoliths as paleoecological indicators.
12, 1-228. Profil, 16, 181-191.
SCHMIDT, H. & FREIWALD, A. 1993. Rezente gesteins- VOGEL, K., GEKTIDIS, M., GOLUBIC, S., KIENE, W. E. &
bohrende Kleinorganismen des norwegischen RADTKE, G. 2000. Experimental studies on
Schelfs. Natur und Museum, 123, 149-155. microbial bioerosion at Lee Stocking Island,
SMYTH, M. J. 1988. The foraminifer Cymbaloporella Bahamas and One Tree Island, Great Barrier
tabellaef Reef, Australia: implications for paleoecological
shells. Journal of Foraminiferal Research, 18, reconstructions. Lethaia, 33, 190-204.
277-285. YOUNG, H. & NELSON, C. 1988. Endolithic biodegrada-
VOGEL, K. & MARINCOVICH, L. in press. Paleobathy- tion of cool-water skeletal carbonates on Scott
metric implications of microborings in Tertiary shelf, northwestern Vancouver Island, Canada.
strata of Alaska, USA. Palaeogeography, Palaeo- Sedimentary Geology, 60, 251-267.
climatology, Palaeoecology. ZEBROWSKI, G. 1936. New genera of Cladochytriaceae.
VOGEL, K., BUNDSCHUH, M., GLAUB, L, HOFMANN, K., Annals of the Missouri Botanical Garden, 23, 553-
RADTKE, G. & SCHMIDT, H. 1995. Hard substrate 564.
ichnocoenoses and their relations to light intensity ZEFF, M. L. & PERKINS, R. D. 1979. Microbial altera-
and marine bathymetry. Neues Jahrbuch fur tion of Bahamian deepsea carbonates. Sedimentol-
Geologie und Palaontologie, Abh., 195, 49-61. ogy,26, 175-201.
Climatic control of trace fossil distribution in the marine realm

ROLAND GOLDRING1, GERHARD C. CADEE2, ASSUNTA D'ALESSANDRO3,


JORDI M. DE GIBERT4, RICHARD JENKINS5 & JOHN E. POLLARD6
l
Geoscience Building, School of Human and Environmental Sciences, University of Reading,
Whiteknights, PO Box 227, Reading RG6 6AB, UK (e-mail: r.goldring@reading.ac.uk)
2
Netherlands Institute for Sea Research, NIOZ, Paleobiology Department, PO Box 59,
1790 AB Den Burg, Texel, The Netherlands (e-mail: cadee@nioz.nl)
3
Dipartimento di Geologia e Geofisica, Universita di Bari, Campus universitario,
Via E. Orabona 4, 70125 Bari, Italy (e-mail: tina@geoMniba.it)
4Departament d'Estratigrafia, Paleontologia i Geociencies Marines, Universitat de Barcelona,
Marti Franques s/n, 08028, Barcelona, Spain (e-mail: gibert@geo.ub.es)
5
The South Australian Museum, North Terrace, Adelaide, South Australia 5000
(e-mail: jenkins.richard@saugov.sa.gov.au)
6Department of Earth Sciences, University of Manchester, Manchester Ml 3 9 PL, UK
(e-mail: John .pollard@ man.ac. uk)

Abstract: Modern coastal and shoreface faunas exhibit strong latitude (climate) controlled
distributions. In contrast, most ichnotaxa are long-ranging, and ichnofacies are widely dis-
tributed geographically. This is readily explained by the dominantly warmer and more
equable climates of much of the past, as well as the diversity of the producers of most ichno-
taxa. Nevertheless, in the Pleistocene, and in the Eocene, cool-water ichnofabrics can be
recognized. The latitudinal distributions of thalassinidean crustaceans and infaunal spatan-
goid echinoids are examined because of their propensity to form distinctive and often
abundant trace fossils. Three climatic zones are tentatively recognized from modern shore
and shoreface sediments, and which are considered to extend back to the Mesozoic: tropical
and subtropical with pellet-lined burrows (Ophiomorphd), echinoid burrows and other traces;
temperate with echinoid burrows and mud-lined or non-lined thalassinidean burrows
(Thalassinoides), but without Ophiomorpha; and arctic (cold waters) with only a molluscan
and annelid trace fossil association. Examples demonstrating this climatic trend are drawn
from the Cenozoic and Pleistocene.

Trace fossils, with the exception of those utilized not its burrow. Diplocraterion is, of course, gener-
in Palaeozoic ichnostratigraphy and footprint ally recognized as the work of a range of animals,
stratigraphy, are well known for their long strati- and at present it is impossible to recognize any
graphic ranges, and sedimentological research has general climatic control in its distribution. Differ-
focused on the ichnofacies concept and the rela- ences in trace fossil suites of climatically distinct
tive constancy of the Seilacherian ichnofacies continental sediments are well established for the
through the Phanerozoic. Ichnotaxa are generally Permian (e.g. desert sandstones, Brady 1947;
regarded as having wide geographical extent as Braddy 1999), lacustrine red-beds (Walter 1980,
well as long time ranges. This is undoubtedly, in 1982, 1983), proglacial lakes (Savage 1971;
part, attributable to the more equable climates Anderson 1981), and Triassic lacustrine red-beds
that prevailed over much of the Phanerozoic, as (Clemmensen 1979). The evolutionary trends in
well as the recognition that individual long- environmental expansion and ecospace utilization
ranging ichnotaxa were formed by a variety of seen in continental ichnofaunas (Buatois & Man-
different animals. However, there are rare to gano 1993; Buatois et al. 1998), especially insect
uncommon ichnotaxa, and others that are more trace fossils in palaeosols (Genise et al. 2000), in
common, that have more restricted distributions, part reflect their response to climate. Pemberton
For example, Diplocraterion of decimetre depth et al. (1992) referred to the distinctions between
and width, are known from the Cambrian to the early Cretaceous, late Cretaceous and Cenozoic
Miocene. Taylor et al. (2003) pointed out that suites of trace fossils in non-marine environments,
specimens with these dimensions are unknown which may be readily attributable to the evolution
from modern environments, and hinted that the of angiosperms, insects and crustaceans rather
originator may be known off tropical deltas, but than climate per se. Savrda (1995) discussed the

From: MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 77-92. 0305-8719/04/S15.00 © The Geological Society
of London.
78 R. COLORING ET AL.

application of ichnology to understanding climati- stratigraphy greatly influence the degree of bio-
cally controlled decimetre-scale bedding rhythms, turbation, as discussed by Taylor et al. (2003)
scour and redox cycles. Stratigraphic facies shifts and Mcllroy (2004). Ausich & Bottjer (1982)
of Ophiomorpha and Zoophycos have been and Bottjer & Ausich (1982) showed that the
documented (Bottjer et al 1987, 1988), but these beginning of the Mesozoic witnessed a massive
do not refer to climatic control; rather Glaub increase in the depth of bioturbation, though
(1994, 1999, 2004), Perry (1998) and Vogel et al this may be somewhat modified if the 1 m depth
(1995) showed how the distribution of micro- of Thalassinoides reported by Droser & Bottjer
borings in skeletal carbonates is determined by (1989) in Ordovician limestones is included.
photic control (depth), and Glaub et al (2002) Droser & Bottjer (1988, 1993) and Bottjer et al
and Vogel & Marincovich (in press) were able to (2000) considered that they could detect a general
extend this to latitudinal trends. increase in the degree of bioturbation seen
Climatic effects on trace fossil distributions will between late Neoproterozoic-Cambrian and
obviously be most apparent in continental, Ordovician sediments of the shoreface, which
marginal marine and neritic facies, because of they attributed to the increase in diversity and
the greater diversity and disparity present in the abundance of the infauna.
equivalent environments than in pelagic facies. Secondly, Cadee (2001) referred to, and
To establish such effects in the fossil record it is expanded on, the increase in diversity of coastal
necessary to show that one is dealing with ichno- life from high to low latitudes. Diversity is not
logical differences between similar depositional readily or reliably applied to ancient bioturbated
environments that are of similar age. It is also sediments, because the effects of salinity and the
important to attempt to demonstrate that absence colonization window have to be eliminated
of an ichnotaxon is due to climatic rather than to (Taylor et al. 2003), and because ichnodiversity is
other palaeoecological factors. Absence of an quite distinct from the diversity of body fossils.
ichnotaxon from a facies in which it would be But changes in diversity through a succession are
expected to occur may be due to real absence most useful, especially when applied to the
(impossible to prove positively), or apparent recognition of marginal marine facies. To apply
absence due to hydraulic factors (i.e. it was once diversity change to ancient sediments requires
there, but was removed by penecontemporaneous extensive examples of more or less contem-
erosion), or apparent absence due to loss of poraneity. Cadee (2001) drew attention to the
identity by reworking. It is in the last situation lack of a diverse fauna of callianassids and crabs
that the role of tiering in aggradational settings and their activities from arctic and temperate
is significant, because the traces of deeper-tier coasts in contrast to their richness in warm
burrows tend to overprint and eliminate the waters. This is of much geological interest because
activities of shallow-tier bioturbators from the trace fossils formed by these crustaceans are so
the rock record, leading to a distinct bias in the conspicuous in the fossil record. But body fossils
ichnofabric and preferential preservation as elite are very uncommon, and there are only a few
traces (Bromley 1996; Goldring et al. 2002). distinctive types of burrow.
Cadee (2001), in a section of his review paper Independent criteria that may be used to assess
on the sediment dynamics of bioturbating organ- palaeoclimate of an ichnological section are:
isms in the coastal zone, drew attention to the
overall palaeogeography and regional climate;
modern latitudinal variation in bioturbation,
facies, especially those that are climatically
and certain changes in the diversity of groups of
sensitive - glaciogenic, desert;
organisms with latitude. The purpose of our con-
information from associated body fossils,
tribution is to demonstrate that such changes may
particularly diversity, and information from
with confidence be related to climatically induced
modern and ancient associations in carbonates
latitudinal change, and can be recognized in the
(foramol, chlorozoan, Lees & Buller 1972;
fossil record. We extend this discussion to include
Lees 1975);
the shoreface, which is of greater geological
morphological factors such as shell thickness,
significance, but lack of information prevents
growth banding in plants, corals and shells
extension to arctic shoreface settings.
(Insalaco 1996), and geochemistry/isotope
Cadee (2001) considered two aspects that
geochemistry of shells;
might be applied to modern bioturbated sedi-
abundance (frequency) - a complex ecological
ments: first, that the overall degree of sediment
factor, but one that may be of value when the
reworking between arctic coasts and warm
overall picture is taken.
coasts increased more than fivefold. But such
measurements cannot be applied to fossil We first discuss aspects of the distributions and
examples, where sedimentation rate and event ichnology of typical crustaceans and spatangoid
CLIMATIC CONTROLS ON MARINE ICHNOLOGY 79

echinoids, before considering some case studies burrow margin, and then smoothed off on the
and drawing conclusions. inside (Frey et al. 1978). Type 4 burrows are distin-
guished by the absence of sediment mounds, but
with restricted apertures, and a deep reticulate
Ichnology of certain crustaceans system of galleries. It is the pelletal lining that is
so readily observed in the fossil record. Such
Dworschak (2000) reviewed the present-day traces are often referred to Ophiomorpha, even
latitudinal distribution of thalassinidean species, though the diagnostic burrow morphology of
which show the highest diversity between 30- Ophiomorpha may not be apparent. The pellets
40°N and 20-30°S (Fig. 1). Thalassinideans are of Ophiomorpha vary considerably in composition
unknown from coastal and shoreface sediments between occurrences, though without apparent
in latitudes higher than about 70°N and 50°S. facies change. Most commonly they are of
Many species construct substantial burrow sys- muddy sand (as described by Frey et al. 1978), in
tems to appreciable depths (partially reviewed by which case they do not compact. In other cases
Bromley 1996). Griffis and Suchanek (1991) they may be more muddy, or rich in plant debris,
recognized four types of thalassinidean burrow, and may become strongly compacted on burial
to which we add additional types (of unproven (Pollard et al. 1993). Frey et al. (1987) also
thalassinidean origin) from the fossil record described a combination of a pelletal lining, and
(below). The burrows are commonly, but not a further mud inner lining. Other crustaceans
invariably, lined (when excavating a relatively make lined burrows, and in the stomatopod
loose substrate). The lining can be of several Squilla, the mud lining may be up to 5 mm thick
types: mucus, which is unlikely to be preserved for a burrow 10-20 mm diameter (Hertweck
fossil; a mud lining (see Bromley 1996), which 1972). Unattributed crustacean burrows con-
may be of different thickness in different taxa; a structed in Cretaceous Chalk, representing a
mud lining pressed into the burrow margin (tamp- lithified coccolith ooze, may also be lined by fish
ing, Stamhuis et al 1996); or a pelleted lining (the bones and scales (Bromley 1996), gymnospermous
hard lining of Griffis & Suchanek 1991), associated leaves or echinoderm elements (Bather 1911), or
with type 4 burrows (Griffis & Suchanek 1991). remain unlined. It is the lining, or its absence, as
The pellets are inserted by the animal into the well as the overall burrow morphology, that is of

Fig. 1. Global map with indications of latitudinal distributions of Callichurus major, thalassinideans and
infaunal echinoids. (1) 70°N to 50°S, northern and southern limits of thalassinidean, and northern limit of
infaunal echinoids. (2) 34°N to 27°S, northern and southern limits of Callichurus major. (3) 42°S probable
southern limit of infaunal echinoids.
80 R. GOLDRING ET AL.

main concern to palaeontology, and it is unfortu- similar burrows as the work of the ghost crab Ocy-
nate that these features have been less considered pode (ocypodid). The upright bow-form burrow
by biologists, for obvious practical considerations. Glyphichnus (Bromley & Goldring 1992), with
Three ichnogenera are widely recognized, and crustacean-type bioglyphs, from lower Cenozoic
attributed to the work of thalassinidean, or sediments, may be linked with certain Cylindrich-
thalassinidean-like crustaceans: Ophiomorpha, nus (Goldring et al. 2002). Associated Meyeria
Thalassinoides and Spongeliomorpha (but see sp. (Glypheoidea) were probably responsible for
Fursich 1973; Schlirf 2000). The overall burrow Lower Cretaceous examples (Goldring 1996).
morphology generally relates to type 4 of Griffis Thalassinoides, Ophiomorpha and Spongelio-
& Suchanek (1991). In Thalassinoides the margin morpha may all be present in one specimen to
of the burrow is smooth, whereas in Spongelio- form a compound trace fossil, reflecting the
morpha the margin displays distinct bioglyphs. need of the constructor to deal with the burrow
Other features used to distinguish crustacean margin in different ways. Clearly, with a firm
ichnotaxa are discussed by Schlirf (2000, and substrate the organism needs only to render the
references therein). Five other crustacean ichno- margin attractive/unattractive to intruders or
taxa may also be considered. The upright spiral microorganisms. The common observation in
Gyrolithes may lead into one or other of the sandy sediments is for a pellet-lined upper section
three ichnotaxa. It generally does not have a of the burrow system to pass down to unlined or
pelletal lining. The sinuous Sinusichnus (Gibert top-lined galleries, reflecting passage into a
1996a, Gibert et al 1999) from the Pliocene of firmer level.
the northwestern Mediterranean, which has an Some species of callianassid thalassinideans
unlined burrow, was probably constructed by a appear to colonize a range of substrates: sand,
decapod crustacean. Pliocene examples of sandy mud, silty mud. Those that pellet-line
Psilonichnus (Fursich 1981), with a Y-, J- or U- their burrows and live in coastal sands, or in
form burrow, were linked to Pholeus (Nesbitt & sandy shoreface settings (Fig. 2 right), appear
Campbell 2002), and to thalassinideans (Gingras to have a restricted latitudinal range. Thus,
et al 2000), though Frey et al. (1984) interpreted on western Atlantic coasts, Callichurus major,

Fig. 2. (a) Diagram to show the formation of meniscate backfill by the modern, globular echinoid
(Echinocardium cordatum) in longitudinal and transverse (centre) section (modified from Bromley & Asgaard
1975), and a wedge-shaped echinoid (Schizaster). Spines are shown diagrammatically. An active respiratory
shaft (funnel) and an abandoned shaft indicate advance of Echinocardium. (b) Composite diagram of an
Ophiomorpha-Planolites-mottlQd ichnofabric, with restricted shaft, constricted aperture, and lined, top-lined
and unlined portions of shafts and galleries.
CLIMATIC CONTROLS ON MARINE ICHNOLOGY 81

forming Ophiomorpha-typeburrows, does not relatively poorly sorted sandy sediment (Kana-
today extend north of the North Carolina- zawa 1992, 1995). Sand dollar activity has not
South Carolina state boundary (about 34°N) yet been recorded from ancient sediments.
(personal communication Curran 2002) or Echinoid burrows are found in lower beach,
south of southern Brazil (about 27° S) upper and middle shoreface settings, and, off
(Dworschak 2000). Thalassinideans in muddier the Georgia coast (Dorjes 1972), in offshore
sediment are present to higher latitudes: e.g. relict sands, as well as deeper-water environ-
Callianassa subterranea in the North Sea ments (below). The latitudinal distribution of
(Reineck 1963; Stamhuis et al 1997) and their shallow marine infaunal spatangoids extends
burrows, which are not pellet-lined, may be from the tropics to 70°N (North Cape) for
referred to Thalassinoides or Spongeliomorpha. African and European species (Hayward &
For fossil distributions the question to be Ryland 1990), and to at least 42°S (Fell 1948)
raised is whether modern distributions can be (Fig. 1), thus considerably greater than that of
applied to ancient occurrences, because different crustaceans forming pellet-lined burrows.
taxa will have been responsible (at least at lower The ichnotaxonomy of spatangoid loco-
orders). The question that must also be posed is motion-feeding trace fossils was reviewed by
whether the recognized ichnospecies of Ophio- Uchman (1995). Bichordites refers to traces
morpha, distinguished on the form and arrange- made by Echinocardium-typQ spatangoids, with
ment of the pellets lining the burrow margin, a single drain, and Scolicia (and associated
had the same or different ecologies, e.g. O. preservational variants) to Spatangus-typQ
nodosa, O. annulata and O. irregulaire. Tenta- echinoids, with a double drain (Fig. 2 left). The
tively, we suggest that occurrences of pellet- resting trace Cardioichnus may be found in
lined burrows in shoreface, lower beach and close association with Scolicia or Bichordites
estuarine sandy sediments represent complex (Smith & Crimes 1983).
constructions in warm tropical to subtropical
environments by decapod crustaceans. Although
there is little likelihood for confusing Ophiomor- Discussion on modern distributions of
pha and Thalassinoides, it is useful to keep in infaunal echinoids and Ophiomorpha-
mind the nature of the substrate associated forming crustaceans
with each occurrence.
The Upogebiidae appear to have a similar The present latitudinal distributions of infaunal
latitudinal range to the Callianassidae echinoids and burrowing crustaceans would
(Dworschak 2000). The burrows are of type 5 seem to offer opportunities for application to
(Griffis & Suchanek 1991), and of the form gen- fossil occurrences, especially because of their dis-
erally referred to the ichnotaxon Psilonichnus. tinctive traces. But there are several problems,
The type ichnospecies, P. tubiformis, is unlined, not only with respect to the well-known difficulty
as are other ichnospecies, though P. upsila of extending modern ecologies to those of fossil
(Frey et al. 1984) and P. lutimuratus (Nesbitt & taxa. It is important to understand the preserva-
Campbell 2002) have a distinctive mud lining. tion potential and regional facies distribution of
the traces. We thus address ecological aspects,
and the sedimentologically related aspects of
Spatangoid echinoid ichnology tiering and tier preservation.
Perhaps the first consideration is in respect of
Burrowing echinoids evolved rapidly in the the salinity tolerance of each group: echinoids,
Cenozoic, though many Mesozoic echinoids being stenohaline, are normally excluded from
were able to plough through the sediment. estuarine environments, whereas crustaceans
Tchoumatchenco & Uchman (2001) record the enjoy a wider salinity tolerance. Where infaunal
oldest deepwater Scolicia that is reasonably echinoids and burrowing crustaceans are present
attributable to an echinoid, from the Tithonian in the same general area, such as the southern
(latest Jurassic), though the age of the oldest shal- North Sea (Reineck 1963) and Georgia coast
low-water three-dimensional backfilled burrow (Dorjes 1972), they tend to occupy different
that can be related to an echinoid is much less specific areas, with different sediment character-
certain, probably because of the low preservation istics, and penetrate to different tier levels. Two
potential (below). Spatangoid echinoids burrow distinct infaunal activities are represented: loco-
to depths no greater than 20cm, and mostly motion/feeding traces by echinoids, and more
shallower (reviewed by Bromley & Asgaard or less permanent to semi-permanent dwelling
1975) (Fig. 2 left). Their burrows can be very structures by crustaceans. The latter require
abundant as well as prominent, especially in wall stabilization and thus the availability,
82 R. GOLDRING ET AL.

either in suspension, within the sediment, or at or amensalism by other bioturbators. Although


at the sediment-water interface, of fine-grained forming relatively deep burrows in the lower
mud or organic matter. beach and shoreface they must maintain a con-
Both groups construct burrows to an appreci- nection with the substrate surface by construct-
able depth, but whereas many fossil and recent ing a ventilation shaft, which must be regularly
crustaceans' burrows extend to a metre or more, replaced as the animal progresses forward (Fig.
and may be regarded as deep-tier structures, echi- 2 left). Any obstruction to this upwardly built
noid burrows are relatively shallow-tier. This is anshaft is disadvantageous to the animal below.
aspect that relates to the relative preservation of Burrowing echinoids would be inhibited from
the tiers, especially, under conditions of more or colonizing areas with pellet (hard)-lined shafts
less continuous sediment aggradation. It is to be of Ophiomorpha, or lined tubes of Skolithos (cf.
expected that spatangoid traces would be readily sand mason Lanice). On the other hand, popula-
overprinted by other deeper traces, and possibly tion disturbance by the activity of the burrowing
eliminated from the fossil record. Indeed, observa- echinoid Brissopsis lyrifera (Widdicombe &
tions by JMdeG and RG on the Miocene shelfal Austen 1998) had a marked negative effect on
sandy biosparites of Alicante (Spain) (below) overall diversity. We also note the effect on
show that this is indeed the case. But only in the deepwater spatangoids on the preservation of
Miocene of the Maltese Islands have echinoid Zoophycos, with truncation of the initial stages
burrows been strongly overprinted to apparent of construction (Kotake 1989).
obscurity by deeper-tier crustacean activity We restrict our discussion to shallow marine
(below). Preservation of shallow tiers may be environments and facies, because arthropod
favoured by several situations (Taylor et al. and echinoid distributions in modern deepwater,
2003; Mcllroy 2004), in which event beds are basinal environments have not yet been fully
particularly involved (we do not refer to tier analysed in respect of ecological parameters.
preservation associated with geochemical changes, Mesozoic and younger turbidites yield many
such as oxygenation, e.g. Wignall 1994): examples of Ophiomorpha and Scolicia, but the
environmental factors that determine their distri-
Below a distal turbiditic event bed, or storm
butions are largely unknown. Furthermore it is
event bed, where little (minimal) penecontem-
not clear whether constructor populations were
poraneous erosion had taken place prior to
necessarily endemic. Occurrences of Ophiomor-
the event sedimentation. This is the classic
pha in basin-floor settings have been frequently
situation for the preservation and casting of
regarded as the work of relocated individuals
pre-event, shallow graphoglyptids on the
(e.g. Follmi & Grimm 1990), though Uchman
soles of distal turbidites (Orr 1994; Uchman
(1995) showed that Ophiomorpha in Miocene
1995).
flysch must be regarded as a normal component
Close to the upper surface of event beds, where
of the deep-sea trace fossil assemblage. The
the event bed was colonized temporarily prior
same problem applies to deep-sea echinoids,
to the renewal of 'normal' sedimentation.
but it may be considered that, following initial
In inclined heterolithic sedimentation, with
relocation from the shelf, populations became
thin amalgamated or closely spaced event
adapted to the deep seafloor (Wetzel &
beds, where little erosion and little sedimenta-
Uchman 2001). Trace fossils are notoriously
tion took place between successive events.
poorly preserved in hemipelagic mudrocks
Gibert (1996b) described this situation from
between turbidite beds. But photographic studies
the Middle Jurassic of Central England, in
and modern deep-sea cores (Fu & Werner 2000),
oolitic and bioclastic grainstones, where each
from coarse silts and fine sands from the north-
'event' is capped by a thin micrite. Deposition
east Atlantic, show that infaunal echinoids, and
took place in a marginal marine environment,
their activity, are of common occurrence. Over-
possibly representing lateral deposition on a
printing of shallower tiers is to be expected,
'point bar'.
though Baldwin & McCave (1999) recorded
In association with large-scale cross-stratifica-
from their sampled area off Nova Scotia the
tion, with impure packstone event units (0.1-
ephemeral nature of the shallower tiers due to
1.0m thick), as described from Rhodes and
penecontemporaneous erosion associated with
Italy (below), where the 'events' represent
frequent benthic storm events. Fu (personal com-
grain-flow avalanching off the edge of a pro-
munication 2000) suggested that the tiering in
grading platform. Similar preservation is to be
deep-sea sediments is often a temporal matter,
expected under avalanching in other situations.
and deeper-tier burrowing organisms are inhib-
A further problem for the preservation of ited by an abundance of burrowing echinoids.
echinoid burrows is that they are subject to Temporal aspects, amensalism and population
CLIMATIC CONTROLS ON MARINE ICHNOLOGY 83

density are not readily assessed for ancient body but with caution to Cretaceous and older settings
or trace fossils, but are relevant factors in respect in respect of the distribution of Ophiomorpha
of shallow as well as deep marine environments. producers.
It follows from the above discussion that turbi-
dites with Ophiomorpha, owing to their being
relocated animals, might have originated in Ophiomorpha and spatangoid trace fossils
tropical or subtropical waters. However, we
reserve judgement on endemic populations, With infaunal echinoid trace fossils not being
except that, following Dworschak (2000), it is recorded prior to the Tithonian (Tchoumatch-
unlikely that water depth would be greater than enco & Uchman 2001) (and mostly later), their
2000m. Further, Tchoumatchenco & Uchman usefulness as climatic indicators is stratigraphi-
(2001) pointed out that the environmental cally limited. Echinoid burrows have been exten-
range of trace fossils is considered mainly at the sively described from the Pleistocene of the
ichnogenus level, but the ichnospecies of Ophio- Mediterranean, though less frequently from Cen-
morpha in deepwater facies differ from the ozoic strata. Ophiomorpha is commonly recorded
common O. nodosa of shallow water, in being from older Mesozoic sediments and onwards.
mostly hypichnia with long, smooth segments.
Uchman (1999) attributed this to substrate
conditions, but barrier prospecting (Jensen & Tropical and subtropical associations
Atkinson 2001) may also be an explanation. (Table 1)
Off the Georgia coast (Dorjes 1972), where
Callichirus major is common in the beach envir- Eocene of southern England
onments, the echinoid Moira bioturbates Pollard et al. (1993) described several ichnofab-
medium- to coarse-grained relict sand, which rics containing Ophiomorpha from units of the
may be too clean for thalassinidean burrowing. Eocene in southern England where the ichno-
In contrast, in the southern North Sea Callia- taxon is prominent (Fig. 2 right), and made com-
nassa subterranea occupies silty mud, whereas parisons with other occurrences, especially from
Echinocardium cordatum occupies fine- to the southeast coast of the USA (Frey et al.
medium-grained sand. In the Gulf of Gaeta 1978). Muddier units contain abundant spatan-
(Mediterranean) (Dorjes 1971) recorded Echino- goid echinoids (Lewis 1989). Echinoid burrows
cardium cordatum but no extensive activity by were tentatively recognized in the highly bio-
thalassinideans. The spatial separation of turbated mudstones (Pollard et al. 1993). The
modern echinoids and thalassinideans seems to climate of the Eocene is generally recognized as
be related to substrate preference, but it might subtropical (Purton & Brasier 1997), with a
be expected that pellet-lined burrows would be flora that is compared to that of southeast Asia
formed in sandy substrates in the North Sea if today (e.g. Collinson 1983).
higher temperatures were present! The thalassini-
dean present in the North Sea forms Thalassi- Oligocene of New Zealand
noides-type burrows, but not Ophiomorpha. Ward & Lewis (1975) recognized well-preserved
We have few data from modern coastal waters, spatangoid burrows in the Arno Limestone, a
and there are probably many more records avail- bioturbated calcarenite with associated large-
able from the Pacific and Indian oceans. We can scale cross-beds, and scour channels. Ophiomor-
tentatively recognize three latitudinal zones in pha nodosa is uncommon, and Ward & Lewis
modern coasts and shoreface settings: (1975) noted the somewhat disparate occurrence
of echinoid burrows (Scolicid) and Ophiomorpha,
in the tropics and sub tropics, where pellet-
though with extensive overlap. Scolicia is also
forming thalassinideans and burrowing echi-
recorded in association with Cardioichnus from
noids are present, though in different facies;
the Upper Miocene to Lower Pliocene (Gregory
in temperate latitudes, where spatangoid echi-
1985).
noids are present with Thalassinoides producers
(in different facies), but not Ophiomorpha
Miocene of Austria, Denmark and Poland
producers;
Radwanski et al. (1975) and Radwanski (1977)
in arctic latitudes, where neither group is
described echinoid burrows, which can be
present, and the principal bioturbators are
assigned to Bichordites, in shallow marine het-
annelids and bivalves (Aitken et al. 1988).
erolithic sands and muds from Denmark, in
We can apply this model to Pleistocene and association with a diverse ichnofauna, including
Cenozoic occurrences with a certain degree of Ophiomorpha nodosa. O. nodosa is not, however,
confidence, depending on the data available, present in the same lithology as Bichordites.
84 R. GOLDRING ET AL.

Table 1. Occurrences of Bichordites/Scolicia and Ophiomorpha in tropical/


subtropical, temperate and arctic climatic zones in the Cenozoic. For most,
the palaeoclimate can be corroborated from associated body fossils

Arctic ? Pliocene, Alaska

Temperate Eocene/Oligocene, South Australia


Pliocene, Washington State, USA
?Pleistocene, Washington State, USA
Pleistocene, Mediterranean
?Pleistocene, Korea

Tropical/subtropical Eocene, southern England


Oligocene, New Zealand
Miocene, Amazonia, Brazil
Miocene, Austria, Denmark and Poland
Miocene, Mediterranean
Miocene, Patagonia, Argentina
Pliocene, northwestern Mediterranean
Pleistocene (Tyrrhenian), Tunisia
Pleistocene, Jamaica

Radwanski et al. (1975) noted that the deposi- Limestone Formation of the Maltese Islands,
tional environment was similar to that described neither echinoid burrows nor O. nodosa are
from the modern southeastern North Sea coast, common. This is in spite of the high frequency
though recognizing the warmer climatic condi- and diversity of infaunal echinoids (especially
tions from the presence of warm water molluscs, Schizaster), which are in or close to life position
and burrows that they attributed to warm water (Rose 1974; Rose et al. 1992). Echinoid burrows
holothurians and amphipods. Uchman & Kren- have been observed (Goldring et al. 2002) at a
mayr (1995) described occurrences of Ophiomor- few specific levels associated with event beds,
pha and Bichordites from shoreface sands and where the shallow tiers have not been over-
muds (molasse facies) of the Lower Miocene of printed by deep-tier, bow-form burrows in Cylin-
Austria. This represents the oldest record of drichnus-mode preservation (Goldring et al.
Bichordites. IScolicia and Ophiomorpha were 2002).
also recorded from transgressive siliciclastics in The Pliocene marginal marine basins of the
SE Poland (Rajchel & Uchman 1999). northwestern Mediterranean were described by
Gibert & Martinell (1998, 1999). Echinoid bur-
Miocene and Pliocene of the Mediterranean area rows are present in the Roussillon Basin in
The molluscs and corals show that the climate of
the Mediterranean area during the Middle to late
Miocene was warm to subtropical. Ophiomorpha
nodosa has been described from a number of sites
(Table 1).
In Alicante (Spain) Bichordites dominates the
sandy biosparites of the Middle to late Miocene
in the Bateig Hill area of Novelda, where the
Miocene sediments occupy one of the small
foreland Eastern Prebetic Basins (Sanz de
Galdeano & Vera 1992). Bateig Hill is extensively
quarried for architectural stone (Bland et al.
2001), and the variety Bateig Fantasia displays
Bichordites ichnofabric (Fig. 3) in 5-20 cm
thick, planar event beds, occasionally associated
with Maretia sp. O. nodosa is occasionally
present, and is more prominent in stratigraphi-
cally adjacent facies, where it may be the
dominant ichnotaxon. Fig. 3. Bateig Fantasia, Miocene, Alicante. Slab cut
In the Maltese Islands, in contrast to Alicante, parallel to stratification with Bichordites isp. cut by
shallow, pelagic wackestones of the Globigerina other traces.
CLIMATIC CONTROLS ON MARINE ICHNOLOGY 85

sandy blue clays in the Planolites—Teichichnus- Ophiomorpha, Monocraterion, Skolithos and


Thalassinoides association. In the contempora- small, curved burrows, with the only mollusc
neous Baix Llobreget Basin, Scolicia is present being Loripes lacteus (a lucinoid bivalve). The
in the more distal deposits whereas Ophiomorpha Tyrrhenian of the Mediterranean represents
is present in more proximal and coarser sandy warm-water conditions (Amore et al. 2000).
channel sediments, associated with Macaronich-
nus isp. and Skolithos linearis. The bioturbation
index does not exceed BI2. The Lower Pliocene Temperate associations
sediments are regarded as having been deposited
under warm water conditions (Martinell et al. The Lower Tertiary of South Australia
1984). Ophiomorpha is not observed in the well-known
and extensive Tertiary deposits of southeastern
Miocene of Patagonia Australia, probably a reflection of the high to
In the Lower Miocene of Patagonia (Argentina), mid-latitudinal placement of this region, lending
Ophiomorpha nodosa is prominent in shoreface support to James's (1997) and James et al.'s
silty sandstones of Chubut Province (Buatois (1997, 2001) recognition of many marine calcar-
et al. 2003; Carmona & Buatois 2003), and a enites of this province as 'cool-water carbonates'.
diverse suite of trace fossils is present in asso- Most of these carbonates are principally made of
ciated lower shoreface sediments. Scolicia is current transported bryozoan remains, and dis-
recorded. The authors suggested that the climatic tinctive trace fossils are rare in the more offshore
signature was cold-temperate due to polar shelf settings. However, distinctive and exqui-
currents, based on independent analysis of sitely preserved crustacean remains, generally of
microfossil elements, and cetacean and penguin characteristic Indo-West-Pacific aspect, are
skeletal remains. This would appear to question common (Jenkins 1977, 1985).
our model. However, the palaeoclimatic model Despite the abundance of echinoids in Austra-
for the Middle Miocene of the area (Valdes lian basinal calcarenites, examples of their traces
et al. 2000) predicts warmer conditions, not are rare. A spectacular bed of Scolicia with
dissimilar from those of the Miocene of the Med- straight and closely meandering backfilled
iterranean area (above). Also, evidence from burrows 8-10 cm wide forms the upper three-
insect trace fossils in palaeosols, fossil plants quarters of a 1 m thick cross-bedded shoreface,
and mammals in the contemporary Punturas sand-rich calcarenite above the major flooding
Formation in inland Patagonia at the same surface, marking the base of the transgressive
latitude indicates a tropical to subtropical cli- Early Oligocene Aldinga Member of the Port
mate (Bown & Laza 1990). Willunga Formation at Port Willunga (Fig. 4).
A likely echinoid producer is occasionally
Miocene of Brazil found in association with the traces, the bigger
Gingras et al. (2002) described inclined hetero- examples of which can be attributed to Meoma
lithic stratification (tidally influenced point bar
sediments) from the Miocene of Amazonia,
deposited in an environment of fluctuating sali-
nity, with the co-occurrence of Ophiomorpha
and Scolicia, the latter indicating colonization
during saline incursions.

Pleistocene of Jamaica and Tunisia


Bichordites is also recorded from the Pleistocene
of Jamaica (Pickerill et al. 1993), in storm-
dominated, medium- to coarse-grained, com-
monly amalgamated sandstones. Most examples
are seen only on upper surfaces, but in associa-
tion with Ophiomorpha, Thalassinoides, Skolithos
and rare Chondrites. The occurrence probably
represents warm conditions and thus a mixed
association.
In the late Pleistocene (Tyrrhenian) of Khunis
(Tunisia), Plaziat & Mahmoudi (1988) and Mah- Fig. 4. Scolicia at the base of the transgressive Early
moudi (1988) drew their examples of Bichordites Oligocene Aldinga Formation at Port Willunga.
from a Skolithos ichnofacies association with Hammer graduated in 5 cm intervals.
86 R. GOLDRING ET AL.

tuberculata (McNamara et al. 1986), but frag- Swinbanks & Murray (1981) noted that the
ments of another unidentified smaller spatangoid former lines its burrow with mud and mucus,
are also present. whereas the latter does not line its burrow.
Thalassinoides appears with the transgression Only the Pleistocene deposits contain Ophio-
at the beginning of the Late Eocene, and at morpha associated with bay sediments. Although
Port Noarlunga, south of Adelaide, metre-deep the authors do not explain whether the Pleisto-
examples penetrate into cracks in the pre-lithified cene deposits represent glacial or interglacial
Tortachilla Limestone before ramifying the soft, sedimentation, we suggest that they may repre-
decalcified underlying sands. Subpolygonal sent interglacial sedimentation, warmer than at
developments of the ichnotaxon are silicified in present.
the overlying lower parts of the Gull Rock
Member of the Blanch Point Formation. How- Pleistocene of Mediterranean
ever, it is in the Late Oligocene to older Miocene The Pleistocene of the Mediterranean offers a
of the Port Phillip basin, Victoria, where number of examples of shallow water, shoreface
common phosphatized examples of Thalassi- facies, several of which show abundant traces
noides include associated remains of producers due to burrowing echinoids. D'Alessandro &
such as Ctenocheles (Callianassidae). Paired Massari (1997), in a comprehensive and inte-
claws of Callianassa that died in their burrows grated study of the Pliocene and Pleistocene
are seen in the shoreface Pliocene at Port Will- deposits in southern Italy, showed that the
unga, south of Adelaide. Middle Pleistocene (Calcarenite della Casarana)
The fully articulated, stalk-eyed crab Ommato- contains a cool-water (foramol) fauna including
carcinus, clearly indicative of a warm-water Arctica islandica (Raffi 1986). Facies 1 includes
influence during the late Early and Middle metre-scale cross-stratification with laminated-
Miocene, apparently died in sloping burrows to-bioturbated calcarenitic event beds. The bio-
which show a stacked septation (Jenkins 1975). turbated intervals are 5-10 cm thick, with Bichor-
The abundant foraminiferal, faunal and floral dites isp., whereas the laminated part can be
indications of warm climatic episodes in south- much thicker. In some intervals the bioturbated
ern Australia during the Cenozoic are well beds are complex and up to a metre thick. The
documented (Wright et al. 2000). laminated-to-bioturbated units represent grain
flow avalanche deposits, subsequently burrowed.
Pliocene of Washington State, USA Ophiomorpha is recorded as very rare. The clino-
Campbell & Nesbitt (2000) and Nesbitt & Camp- form stratification is truncated by a unit (up to
bell (2002) have provided a convincing account 2 m thick) of fine-grained calcarenites, intensely
of an active margin, storm-flood influenced bioturbated by Thalassinoides boxworks. The
Pliocene estuary fill, grading to shelfal (basinal) Calcarenite della Casarana as a whole represents
muddy sediments. Whereas the shelfal sediments coarse sediments deposited at relatively lower
contain echinoid burrows, more proximal sandy, sea-level during forced regression.
estuarine sediments contain Psilonichnus latimur- The succeeding Middle Pleistocene (pre-
atus. Campbell & Nesbitt (2000) recorded only Tyrrhenian) Sabbie della Serrazza is a more
rare Ophiomorpha. Psilonichnus appears to have heterogeneous unit that coarsens upwards from
a wider latitudinal distribution than Ophiomor- micaceous sandy muds to coarse-grained
pha, and, as Campbell & Nesbitt (2000) note, calcarenites. It is associated with a relatively
its producers appear to have a tolerance to cool-water (D'Alessandro & Massari 1997)
lower salinity. (cooler than present-day) shelly fauna with
numerous Pseudamussium septemradiatum and
Pleistocene of Washington State, USA A. islandica. Thalassinoides networks and
Gingras et al. (1999, 2000) compared the modern boxworks are present, which in some instances
trace-forming biota from Willapa Bay with the are of Spongeliomorpha type when piped into
trace fossils in closely similar Pleistocene Early Pleistocene marls.
estuarinent Similar large-scale cross-stratification in grain-
nized as of common occurrence in several stones to packstones was described by Bromley
Pleistocene facies. This appears to refute our & Asgaard (1975), and Hanken et al. (1996)
model, as it is unlikely that subtropical condi- from the Pleistocene Cape Arkhangelos calcar-
tions reached to 47°N, even during warmer inter- enite facies of the Rhodes Formation of
glacials. Thalassinoides and 'Ophiomorpha-\ike' Rhodes. The unit represents a spectacular, and
burrows were recognized from the modern unusual, example of such stratification, which
sediments at Willapa Bay and linked to Upogebia forms large asymptotic clinoforms in beds with
pugettensis and Callianassa californiensis, though dips up to 30°. The facies is composed of event
CLIMATIC CONTROLS ON MARINE ICHNOLOGY 87

beds, from less than 10 cm to over 1 m thick, each rising sea-levels. Most of the sequences are
fining upwards. As in Italy, the beds appear to bounded by subaerial, karstic unconformities.
represent grain-flow avalanches on clinoforms A different association is recorded in another
constructed at the outer margin of a shallow- sequence with O. nodosa associated with Arctica
water, carbonate sand body that was prograding islandica, a bivalve that today is found only in
into deeper water. The beds are intensely biotur- waters where the February temperature are
bated by Echinocardium, which forms winding equal or lower than 5-6 °C, i.e. cool to cool-
traces of Bichordites. Ophiomorpha nodosa is temperate (Raffi 1986). However, O. nodosa is
absent, but is recorded from lower units of the atypical, being less than 1 cm in diameter with a
Rhodes Formation, as are large corals, which lining of numerous small pellets, and a different
are of late Pliocene age, representing warm- overall morphology, with scarcely branched
water sedimentation. long shafts.
Ophiomorpha is well known from ancient
cross-stratified siliciclastic and carbonate sands, Pleistocene of Korea
though often relatively sparse (Pollard et al. Kim & Heo (1997) described marginal marine
1993). There are several possible explanations and shelfal siliciclastics from the Early Pleisto-
for the absence of crustacean activity cross- cene Seogwipo Formation of Jeju Island
cutting echinoid burrows in the clinoform strati- (Cheju-do) (latitude 33°N), Korea. Probable
fication of Rhodes: Bichordites or Scolicia (recorded as Laminites)
are present in association with a Cruziana ichno-
(a) temporal exclusion as in the deep sea
facies assemblage. Thalassinoides is recorded, but
(above);
not Ophiomorpha. The palaeoenvironment and
(b) the short duration of the colonization
molluscan assemblage were described by Kang
window between successive avalanches,
(1995), who recognized the influence of cold-
though this does not seem to have pre-
vented the formation of Ophiomorpha in water currents, and a southward migration of
Cretaceous and Cenozoic cross-stratified boreal species.
strata (Pollard et al. 1993);
(c) climatic control. Arctic associations
The association with foramol facies in Italy sug-
gests that a climatic control is the most likely Eyles et al. (1992) described Pliocene? glacially
explanation. influenced continental shelf and slope trace fos-
In these examples from Italy and Rhodes sils from the Yakataga Formation of Alaska.
Ophiomorpha appears to be virtually absent, We are not convinced by their identification of
but Thalassinoides is present in the same strati- Ophiomorpha (Eyles et al. 1992, fig. 10), or that
graphic units associated with echinoid activity, the backfilled burrow (op. cit. fig. 11) is attri-
thus corresponding to present-day occurrences. butable to echinoid activity. Thus their trace
Hanken et al. (1996, figs 8, 14) interpreted the fossil assemblages may be better interpreted as
Cape Arkhangelos as representing highstand attributable to annelid and molluscan activity
sedimentation, but this is somewhat inimical in an arctic setting.
for a cool-water (glacial) interval, and may be
better attributed to local tectonic activity.
Confirmation of the climatic control is seen in Conclusions
the Novoli Graben (Salento Peninsula, Puglia,
southern Italy), where 9m-scale sequences have Our analysis of a number of Cenozoic and Pleis-
been described by D'Alessandro et al. (in tocene occurrences of echinoid and crustacean
burrows, together with modern examples, sug-
press). The sediments predominantly comprise
gests a general model of climate control for this
bioclastic packstones and grainstones to rud-
geological interval. We tentatively recognize
stones, and represent foramol-type calcarenites.
Biogenic mottling is ubiquitous, and discrete three latitudinal zones in modern coastal and
shoreface settings:
trace fossils include Thalassinoides, Bichordites,
and (restricted to certain horizons) Ophio- Tropical and Subtropical Zone, where
morpha, Cylindrichnus, Gyrolithes and Tasselia. pellet-forming thalassinideans and burrowing
The vertical changes in the body and trace echinoids are present, though in discrete facies;
fossils within the sequences indicate water Temperate Zone, where spatangoid echinoids
depth changes corresponding to known sixth- are present with facially separated Thalas-
order glacio-eustatic climate fluctuations, with sinoides producers, but not Ophiomorpha
Ophiomorpha entering the stratigraphy with producers;
88 R. GOLDRING ET AL.

Arctic Zone, where neither group is present, References


and the principal bioturbators are annelids
and molluscs. AITKEN, A. E., RISK, M. J. & HOWARD, J. D. 1988.
Animal-sediment relationships on a subarctic
Although we have referred to relatively few intertidal flat, Pangnirtung Fiord, Baffin Island,
examples, and with differing degrees of confi- Canada. Journal of Sedimentary Petrology, 58,
dence, our model for climatic distribution of 969-978.
shallow marine trace fossils appears to be AMORE, F. O., CIAMPO, G., Di DONATO, V., ESPOSITO,
robust. There are two anomalous examples P., ERMOLLI, E. R. & STAITI, D. 2000. An inte-
from the Pliocene and Pleistocene. The Pliocene grated micropalaeontological approach applied
to Late Pleistocene-Holocene palaeoclimatic and
example we regard as probable misidentification palaeoenvironmental changes (Gaeta Bay, Tyrrhe-
of both the crustacean and echinoids traces, nian Sea). In: HART, M. B. (ed.) Climates: Past and
though we must reserve judgement on the Present. Geological Society, London, Special
Pleistocene example from Washington State. Publications, 181, 95-111.
The occurrence in Italy of O. nodosa in associa- ANDERSON, A. M. 1981. The Umfolozia arthropod
tion with Arctica islandica is also anomalous, trackways in Permian Dwyka and Ecca Series
but the size and overall morphology of the of South Africa. Journal of Paleontology, 55, 84—
trace fossil are quite atypical. The association 108.
of Ophiomorpha in apparently cold-temperate AUSICH, W. I. & BOTTJER, D. J. 1982. Tiering in suspen-
sion-feeding communities on soft substrata
sediments in the Miocene of Patagonia is anom-
throughout the Phanerozoic. Science, 216, 173—
alous, but the palaeoclimate model suggests 174.
much warmer conditions than those suggested BALDWIN, C. T. & McCAVE, I. N. 1999. Bioturbation in
by the micropalaeontological data. an active deep-sea area: implications for models of
We also tentatively suggest that these assess- trace fossil tiering. Palaios, 14, 375-388.
ments may be applied to deep water (turbiditic) BATHER, F. A. 191L Upper Cretaceous terebelloids
occurrences. We recognize the problem of from England. Geological Magazine, 8, 481^87,
extending this zonation further back in the geo- 549-556.
logical record to Mesozoic occurrences of Ophio- BLAND, B. H., GIBERT, J. M. DE & COLORING, R. 2001.
morpha because of the problems in knowing what A fossil whodunnit. Geology Today, 17, 229-230.
BOTTJER, D. J. & AUSICH, W. I. 1982. Tiering and sam-
the constructor(s) were, and whether they were pling requirements in paleocommunity reconstruc-
capable of forming a pellet lining to the tion. Proceedings of the Third North American
burrow. We also raise some of the questions Paleontological Convention, 1, 57-59.
that need to be addressed by ichnology: the BOTTJER, D. J., DROSER, M. L. & JABLONSKI, D. 1987.
types and methods of construction of burrow Bathymetric trends in the history of trace fossils.
linings, turbiditic distributions of Ophiomorpha In: BOTTJER, D. J. (ed.) New Concepts in the Use
and related crustacean traces, and ecological tol- of Biogenic Sedimentary Structures for Paleoenvir-
erances of spatangoid echinoids. The distribution onment Interpretation. Society of Economic
of other ichnotaxa should be investigated for Paleontologists and Mineralogists, Pacific Section,
Los Angeles, California, 57—65.
possible latitudinal distributions. We also
BOTTJER, D. J., DROSER, M. L. & JABLONSKI, D. 1988.
emphasize the necessity for careful identification Paleoenvironmental trends in the history of trace
of ichnotaxa, and the identification of the fossils. Nature, 333, 252-255.
sedimentary surface actually colonized. BOTTJER, D. J., DROSER, M. L. & DORNBOS, S. Q. 2000.
The Cambrian susbstrate revolution. GSA Today,
We are grateful to the management and quarrymen of 10, 1-7.
Bateig Laboral SA (Alicante) for access to the quarries BOWN, T. M. & LAZA, J. H. 1990. A Miocene termite
of Bateig Hill, Novelda. E. Stamhuis (Groningen) gen- nest from southern Argentina and its paleo-
erously provided a copy of his thesis, and L. Buatois climatological implications. Ichnos, 1, 73-79.
(Tucuman), P. Dworschak (Vienna) and J.-Y. Kim BRADY, L. F. 1947. Invertebrate tracks from Coconino
(Chungbuk, Korea) kindly provided relevant literature. Sandstone of northern Arizona. Journal of Paleon-
B. Sellwood (Reading) discussed and provided a CD- tology, 21, 466^72.
ROM for Miocene climates. A. Curran (Smith College, BRADDY, S. J. 1999. Terrestrial trace fossils from the
Massachusetts), K. Campbell and M. Gregory (Auck- Robledo Mountains ichnofauna (Lower Permian)
land), Shaoping Fu (Bochum) and L. Nesbitt of southern New Mexico. Geoscientist, 9, 5-6.
(Washington) are acknowledged for generous discus- BROMLEY, R. G. 1996. Trace Fossils: Biology, Taphon-
sion. L. Buatois and J. Genise constructively reviewed omy and Applications (2nd edn). Chapman &
the paper. Participation of JMdG forms part of the Hall, London.
activities of the consolidated research group BROMLEY, R. G. & ASGAARD, U. 1975. Sediment
2001SGR-00077 of the University of Barcelona, and structures produced by a spatangoid echinoid: a
research project BTE 2000-0584 of the Ministerio de problem of preservation. Bulletin of the Geological
Ciencia y Technologia of Spain. Society of Denmark, 24, 261-281.
CLIMATIC CONTROLS ON MARINE ICHNOLOGY 89

BROMLEY, R. G. & GOLDRING, R. 1992. The palaeo DROSER, M. L. & BOTTIER, D. J. 1989. Ordovician
burrows at the Cretaceous to Palaeocene firm- increase in extent and depth of bioturbation:
ground unconformity in southern England. implications for understanding early Paleozoic
Tertiary Research, 13, 95-102. ecospace utilization. Geology, 17, 850—852.
BUATOIS, L. A. & MANGANO, M. G. 1993. Ecospace DROSER, M. L. & BOTTIER, D. J. 1993. Trends and
utilization, paleoenvironmental trends and the patterns of Phanerozoic ichnofabrics. Annual
evolution of early nonmarine biotas. Geology, 21, Review of Earth and Planetary Science 1993, 21,
595-598. 205-225.
BUATOIS, L. A., MANGANO, M. G., GENISE, J. F. & DWORSCHAK, P. C. 2000. Global diversity in the
TAYLOR, T. N. 1998. The ichnological record of Thalassinidea (Decapoda). Journal of Crustacean
the continental invertebrate invasion: evolutionary Biology, 20, special number 2, 238-245.
trends in environmental expansion, ecospace EYLES, N., VOSSLER, S. M. & LAGOE, M. B. 1992.
utilization and behavioral complexity. Palaios, Ichnology of a glacially-influenced continental
13, 217-240. shelf and slope; the late Cenozoic Gulf of Alaska
BUATOIS, L. A., BROMLEY, R. G., MANGANO, M. G., (Yakataga Formation). Palaeogeography, Palaeo-
BELLOSI, E. & CARMONA, N. 2003. Ichnology of climatology, Palaeoecology, 94, 193—221.
shallow marine deposits in the Miocene Chenque FELL, H. B. 1948. A key to the sea-urchins of New
Formation of Patagonia: complex ecologic Zealand. Tuatara, 1, 6-13.
structure and niche partitioning in Neogene eco- FOLLMI, K. B. & GRIMM, K. A. 1990. Doomed pioneers:
systems. Asociacion Paleontologica Argentina. gravity-flow pioneers and bioturbation in marine
Publication Especial, 9, 1-11. oxygen-deficient environments. Geology, 18,
CADEE, G. C. 2001. Sediment dynamics by bioturbating 1069-1072.
organisms. In: REISE, K. (ed.) Ecological Compari- FREY, R. W., HOWARD, J. D. & PRYOR, W. A.
sons of Sedimentary Shores. Ecological Studies, 1978. Ophiomorpha: its morphologic, taxonomic,
Springer, Berlin, 151, 127-148. and environmental significance. Palaeogeo-
CAMPBELL, K. A. & NESBITT, E. A. 2000. High- graphy, Palaeoclimatology, Palaeoecology, 23,
resolution architecture and paleoecology of an 199-229.
active margin, storm-flood influenced estuary, FREY, R. W., CURRAN, H. A. & PEMBERTON, S. G. 1984.
Quinault Formation (Pliocene), Washington. Tracemaking activities of crabs and their environ-
Palaios, 15, 553-579. mental significance: the ichnogenus Psilonichnus.
CARMONA, N. B. & BUATOIS, L. A. 2003. Estructuras Journal of Paleontology, 58, 333-350.
biogenicas de crustaceos en el Mioceno de la FREY, R. W., HOWARD, J. D. & HONG, J.-S. 1987.
cuenca del Golfo San Jorje: implicancias paleo- Prevalent lebensspuren on a modern macrotidal
biologicas y evolutivas. Asociacion Paleontologica flat, Inchon, Korea: ethological and environ-
Argentina. Publication Especial, 9, 97-108 mental significance. Palaios, 2, 571-593.
CLEMMENSEN, L. B. 1979. Triassic lacustrine red-beds Fu, S. & WERNER, F. 2000. Distribution, ecology and
and palaeoclimate: The 'Buntsandstein' of taphonomy of the organism trace, Scolicia, in
Helgoland and the Malmos Klint Member of northeast Atlantic deep-sea sediments. Palaeogeo-
East Greenland. Geologische Rundschau, 68, 748- graphy, Palaeoclimatology, Palaeoecology, 156,
774. 289-300.
COLLINSON, M. E. 1983. Fossil Plants of the London FURSICH, F. T. 1973. A revision of the trace fossils
Clay. Palaeontological Association Field Guides Spongeliomorpha, Ophiomorpha and Thalassi-
to Fossils, No. 1, London. noides. Neues Jahrbuchfur Geologic und Palaonto-
D'ALESSANDRO, A. & MASSARI, F. 1997. Pliocene and logie, Monatshefte 1973, 719-735.
Pleistocene depositional environments in the FURSICH, F. T. 1981. Invertebrate trace fossils from the
Pesculuse area (Salento, Italy). Revista Italiana Upper Jurassic of Portugal. Communiques Servicos
de Paleontologia e Stratigrafia, 103, 221-258. Geologicos de Portugal, 67, 53-168.
D'ALESSANDRO, A., MASSARI, F., DAVAUD, E. & GENISE J. F., MANGANO, M. G., BUATOIS, L. A., LAZA,
GHIBAUDO, G. (in press) Pliocene-Pleistocene J. H. & VERDE, M. 2000. Insect trace fossil associa-
sequences bounded by subaerial unconformities tions in paleosols: the Coprinisphaera ichnofacies.
within foramol ramp calcarentites and mixed Palaios, 15, 49-64.
deposits (Salento, SE Italy). Sedimentary Geology. GIBERT, J. M. DE. 1996a. A new decapod burrow system
DORIES, J. 1971. Der Golf von Gaeta (Tyrrhenisches from the NW Mediterranean Pliocene. Revista
Meer). IV. Das Makrobenthos und seine kiisten- Espanola de Paleontologia, 11, 251-254.
parallelen Zonierung. Senckenbergiana Maritima, GIBERT, J. M. DE. 1996b. Diopatrichnus odlingi n. isp.
3, 203-246. (annelid tube) and associated ichnofabrics in the
DORIES, J. 1972. Georgia coastal region, Sapelo Island, White Limestone (M. Jurassic) of Oxfordshire:
USA: sedimentology and biology. VII, Distribu- sedimentological and palaeoecological signifi-
tion and zonation of macrobenthic animals. cance. Proceedings of the Geologists' Association,
Senckenbergiana Maritima, 4, 183—216. 107, 189-198.
DROSER, M. L. & BOTTIER, D. J. 1988. Trends in depth GIBERT, J. M. DE & MARTINELL, J. 1998. Ichnofabrics
and extent of bioturbation in Cambrian carbonate of the Pliocene marginal marine basins of the
marine environments, western United States. northwestern Mediterranean. Revista de la
Geology, 16, 233-236. Societal Geologica de Espana, 11, 43-56.
90 R. GOLDRING ET AL.

GIBERT, J. M. DE & MARTINELL, J. 1999. Proximal- HAYWARD, P. J. & RYLAND, J. S. (eds) 1990. The
distal variations of trace fossil assemblages in a Marine Fauna of the British Isles and North- West
Pliocene ria, Baix Llobregat, northeastern Spain. Europe. Vol. 1: Introduction and Protozoans to
Revista de la Societat Geologica de Espana, 12, Arthropods. Clarendon Press, Oxford
209-214. HERTWECK, G. 1972. Georgia coastal region, Sapelo
GIBERT, J. M. DE, JEONG, K. & MARTINELL, J. 1999. Island, USA: sedimentology and biology. V,
Ethologic and ontogenetic significance of the Distribution and environmental significance of
Pliocene trace fossil Sinusichnus sinuous from lebensspuren and in-situ skeletal remains. Senck-
the northwestern Mediterranean. Lethaia, 32, enbergiana Maritima, 4, 125-166.
31-40. INSALACO, E. 1996. The use of Late Jurassic coral
GINGRAS, M. K., PEMBERTON, S. G., SAUNDERS, T. & growth bands as palaeoenvironmental indicators.
CLIFTON, H. E. 1999. The ichnology of modern Palaeontology, 39, 413^431.
and Pleistocene brackish-water deposits at JAMES, N. P. 1997. The cool-water carbonate deposi-
Willapa Bay, Washington: variability in estuarine tional realm. In: JAMES, N. P. & CLARKE, J. A. D.
settings. Palaios, 14, 352-374. (eds) Cool-Water Carbonates. Society for Sedi-
GINGRAS, M. K., HUBBARD, S. M., PEMBERTON, S. G. & mentary Geology, Special Publications, Tulsa,
SAUNDERS, T. 2000. The significance of Psilonich- Oklahoma, 56, 185-203.
nus at Willapa Bay, Washington. Palaios, 15, JAMES, N. P., BONE, Y., HAGEMAN, S., GOSTIN, V. A. &
142-151. FEARY, D. A. 1997. Cool-water carbonate
GINGRAS, M. K., RADANEN, M. & RANZI, A. 2002. The sedimentation during the terminal Quaternary,
significance of bioturbated inclined heterolithic high-amplitude sea-level cycle: Lincoln Shelf,
stratification in the southern part of the Miocene southern Australia. In: JAMES, N. P. & CLARKE,
Solimoes Formation, Rio Acre, Amazonia J. A. D. (eds) Cool-Water Carbonates. Society
Brazil. Palaios, 17, 591-601. for Sedimentary Geology, Special Publications,
GLAUB, I. 1994. Mikrobohrungen in ausgewahlten Tulsa, Oklahoma, 56, 53-76.
Ablagerungensraumen des europaischen Jura JAMES, N. P., BONE, Y., COLLINS, L. B. & KYSER, K.
und der Unterkreide (Klassifikation und Paloko- 2001. Surficial sediments of the Great Australian
logie). Courier Forschungsinstitut Senckenberg, Bight: facies dynamics and oceanography on a
174, 1-324. vast cool-water carbonate shelf. Journal of Sedi-
GLAUB, I. 1999. Microborings and bathymetrical mentary Research, 71, 549-567.
reconstructions. Bulletin of the Geological Society JENKINS, R. J. F. 1975. The fossil crab Ommatocarcinus
of Denmark, 45, 143-146. corioensis (Cresswell) and a review of the related
GLAUB, I. 2004 Recent and sub-recent microborings Australasian species. Memoirs of the National
from the upwelling area off Mauritania (West Museum of Victoria, 36, 33—62.
Africa) and their implications for palaeoecology. JENKINS, R. J. F. 1977 A new fossil homolid crab (Dec-
In: MclLROY, D. (ed.) The Application of Ichnol- apoda, Brachura), middle Tertiary, southeastern
ogy to Palaeoenvironmental and Stratigraphic Australia. Transactions of the Royal Society of
Analysis. Geological Society, London, Special South Australia, 101, 1-10.
Publications, 228, 63-76. JENKINS, R. J. F. 1985. Fossil spider crabs from
GLAUB, I., GEKTIDIS, M. & VOGEL, K. 2002. Micro- Australia. In: MURRAY LINDSAY (ed.) Stratigraphy,
borings from different North Atlantic shelf areas: Palaeontology, Malacology Papers in Honour of
variability of the euphotic zone extension and Dr Nell Ludbrook. Department of Mines and
implications for palaeodepth reconstructions. Energy South Australia, Special Publications, 5,
Courier Forschungsinstitut Senckenberg, 237, 25— 145-165.
37. JENSEN, S. & ATKINSON, R. J. A. 2001. Experimental
GOLDRING, R. 1996. The sedimentological significance production of animal trace fossils, with a discus-
of concentrically laminated burrows from Lower sion of allochthonous trace fossil producers.
Cretaceous Ca-bentonites, Oxfordshire. Journal Neues Jahrbuch fur Geologic und Palaontologie,
of the Geological Society, London, 153, 255—263. Monatshefte, 2001, 594-606.
GOLDRING, R., GRUSZCZYNSKI, M. & GATT, P. A. KANAZAWA, K. 1992. Adaptation of test shape for bur-
2002. Bow-form burrow and its sedimentological rowing and locomotion in spatangoid echinoids.
and paleoecological significance. Palaios, 17, Palaeontology, 35, 733-750.
622-630. KANAZAWA, K. 1995. How spatangoids produce their
GREGORY, M. R. 1985. The 'Bow Tie' trace fossil from traces: relationship between burrowing mechan-
East Cape, North Island, New Zealand. New ism and trace structure. Lethaia, 28, 211—219.
Zealand Geological Survey Record, 9, 56-58. KANG, S. S. 1995. Reconstruction of the paleoenviron-
GRIFFIS, R. B. & SUCHANEK, T. H. 1991. A model of ment and molluscan assemblage of the Lower
burrow architecture and trophic modes in thalassi- Pleistocene Sogwipo Formation, Cheju Island,
nidean shrimp (Decapoda: Thalassinidean). Korea. PhD thesis, Niigata University, Japan.
Marine Ecology Progress Series, 79, 171-183. KIM, J.-Y. & HEO, W.-H. 1997. Shell beds and trace
HANKEN, N.-M., BROMLEY, R. G. & MILLER J. 1996. fossils of the Seogwipo Formation (Early Pleisto-
Plio-Pleistocene sedimentation in coastal grabens, cene), Jeju Island, Korea. Ichnos, 5, 89-99.
north-east Rhodes, Greece. GeologicalJournal, 31, KOTAKE, N. 1989. Paleoecology of the Zoophycos
393-418. producers. Lethaia, 22, 327-341.
CLIMATIC CONTROLS ON MARINE ICHNOLOGY 91

LEES, A. 1975. Possible influence of salinity and tem- RADWANSKI, A., FRIIS, H. & LARSEN, G. 1975. The
perature on modern shelf carbonate sedimenta- Miocene Hagen0r-B0rup sequence at Lillebaelt
tion. Marine Geology, 19, 159-198. (Denmark): its biogenic structures and deposi-
LEES, A. & DULLER, A. T. 1972. Modern temperate- tional environment. Bulletin of the Geological
water and warm-water shelf carbonate sediments Society of Denmark, 24, 229-260.
contrasted. Marine Geology, 13, M67-M73. RAFFI, S. 1986. The significance of marine boreal
LEWIS, D. N. 1989. Fossil Echinoidea from the Barton molluscs in the Early Pleistocene faunas of the
Beds (Eocene, Bartonian) of the type locality at Mediterranean area. Palaeogeography, Palaeo-
Barton-on-Sea in the Hampshire Basin, England. climatology, Palaeoecology, 52, 267-289.
Tertiary Research, 11, 1-47. RAJCHEL, J. & UCHMAN, A. 1999. Trace fossils from the
MclLROY, D. 2004. A review of some ichnological con- Miocene transgresive siliciclastics near Dynow, SE
cepts, methodologies, applications and frontiers. Poland. Ada Palaeontologica Romaniae, 2, 433-
In: MclLROY, D. (ed.) The Application of Ichnol- 440.
ogy to Palaeoenvironmental and Stratigraphic REINECK, H.-E. 1963. Sedimentgefiige im Bereich der
Analysis. Geological Society, London, Special siidlichen Nordsee. Abhandlungen der Senckenber-
Publications, 228, 3-27. gischen Naturforschenden Gesellschaft, 505, 1—138.
MCNAMARA, K. J., PHILLIP, G. M. & KRUSE, P. O. ROSE, E. R. 1974. Stratigraphical and facies distribu-
1986. Tertiary brissid echinoids of southern tion of irregular echinoids in Miocene limestones
Australia. Alcheringa, 10, 55-86. of Gozo, Malta and Cyrenaica, Libya. Memoire
MAHMOUDI, M. 1988. Nouvelle proposition de sub- de Bureau Recherches Geologic et Mineral, 78,
divisions stratigraphiques des depots attribues au 349-355.
Tyrrhenien en Tunisie (region de Monastir). Bulletin ROSE, E. R., PRATT, S. K. & BENNETT, S. M. 1992.
Societe geologique de France, 1988 (8), IV, 431-435. Evidence for sea-level changes in the Globigerina
MARTINELL, J., DOMENECH, R. & MARQUINA, M. J. Limestone Formation (Miocene) of the Maltese
1984. Molluscan assemblages in the north-east Islands. Paleontologia i Evolucio, 24-25, 265-276.
Spanish Pliocene. Annales Geologiques des Pays SANZ DE GALDEANO, C. & VERA, J. A. 1992. Strati-
Helleniques, 32, 35-56. graphic record and palaeogeographical context
NESBITT, E. A. & CAMPBELL, K. A. 2002. A new Psilo- of the Neogene basins in the Betic Cordillera,
nichnus ichnospecies attributed to mud-shrimp Spain. Basin Research, 4, 21-36.
Upogebia in estuarine settings. Journal of Paleon- SAVAGE, N. M. 1971. A varvite ichnocoenosis from the
tology, 76, 892-961. Dwyka Series of Natal. Lethaia, 4, 217-233.
ORR, P. J. 1994. Trace fossil tiering within event beds SAVRDA, C. E. 1995. Ichnologic applications in paleo-
and preservation of frozen profiles: an example oceanographic, paleoclimatic, and sea-level
from the Lower Carboniferous of Menorca. studies. Palaios, 10, 565-577.
Palaios, 9, 202-210. SCHLIRF, M. 2000. Upper Jurassic trace fossils from the
PEMBERTON, S. G., MACEACHERN, J. A. & FREY, R. W. Boulonnais (northern France). Geologica et
1992. Trace fossil facies models: environmental Palaeontologica, 34, 145-213.
and allostratigraphic significance. In: WALKER, SMITH, A. & CRIMES, T. P. 1983. Trace fossils formed by
R. G. & JAMES, N. P. (eds) Facies Models. Geo- heart urchins: a study of Scolicia and related
logical Association of Canada, 47-72. traces. Lethaia, 16, 79-92.
PERRY, C. T. 1998. Grain susceptibility to the effects of STAMHUIS, E. J., REEDE-DEKKER, T., ETTEN, Y. VAN,
microboring: implications for the preservation of WILJES, J. J. DE & VIDELER, J. J. 1996. Behaviour
skeletal carbonates. Sedimentology, 45, 39-51. and time allocation of the burrowing shrimp
PICKERILL, R. K., DONOVAN, S. K., DIXON, H. L. & Callianassa subterranea (Decapoda, Thalassinidea).
DOYLE, E. N. 1993. Bichordites monastirensis Journal of Experimental Marine Biology and
from the Pleistocene of southeast Jamaica. Ecology, 204, 225-239.
Ichnos, 2, 225-230. STAMHUIS, E. J., SCHREURS, C. E. & VOLELER, J. J. 1997.
PLAZIAT, J.-C. & MAHMOUDI, M. 1988. Trace fossils Burrow architecture and turbative activity of the
attributed to burrowing echinoids: a revision thalassinid shrimp Callianassa subterranea from
including new ichnogenus and ichnospecies. the central North Sea. Marine Ecology Progress
Geobios, 21, 209-233. Series, 151, 155-163.
POLLARD, J. E., COLORING, R. & BUCK, S. G. 1993. Ich- SWINBANKS, D. D. & MURRAY, J. W. 1981. Bio-
nofabrics containing Ophiomorpha: significance in sedimentological zonation of Boundary Bay tidal
shallow-water facies interpretation. Journal of the fiats, Fraser River Delta, British Columbia. Sedi-
Geological Society, London, 150, 149-164. mentology, 28, 201-237.
PURTON, L. & BRASIER, M. 1997. Gastropod carbonate TAYLOR, A., GOLDRING, R. & GOWLAND, S. 2003. Ana-
618O and 813C values record strong seasonal pro- lysis and application of ichnofabrics. Earth Science
ductivity and stratification shifts during the late Reviews, 60, 227-259.
Eocene in England. Geology, 25, 871-874. TCHOUMATCHENCO, P. & UCHMAN, A. 2001. The oldest
RADWANSKI, A. 1977. Present-day types of trace in the deep-sea Ophiomorpha and Scolicia and associated
Neogene sequence; their problems of nomen- trace fossils from the Upper Jurassic-Lower
clature and preservation. In: CRIMES, T. P. & Cretaceous deep-water turbidite deposits of SW
HARPER, J. C. (eds) Trace Fossils 2, Geological Bulgaria. Palaeogeography, Palaeoclimatology,
Journal, Special Issue, 9, 227-264. Palaeoecology, 169, 85-99.
92 R. GOLDRING ET AL.

UCHMAN, A. 1995. Taxonomy and palaeoecology of WALTER, H. 1983. Zur Taxonomie, Okologie und Bio-
flysch trace fossils: the Marnoso-arenacea Forma- stratigraphie der Ichnia limnisch-terrestrischer
tion and associated facies (Miocene, northern Arthropoden des mitteleuropaischen Jungpalao-
Apennines, Italy). Beringeria, 15, 3-115. zoikums. Freiberger Forschungsheft, C382, 146-
UCHMAN, A. 1999. Ichnology of the Rhenodanubian 193.
Flysch (Lower Cretaceous-Eocene) in Austria WARD, D. M. & LEWIS, D. W. 1975. Paleoenviron-
and Germany. Beringeria, 25, 67—173. mental implications of storm-scoured, ichno-
UCHMAN, A. & KRENMAYR, H.G. 1995. Trace fossils fossiliferous mid-Tertiary limestones, Waihao
from Lower Miocene (Ottangian) molasse depos- District, South Canterbury, New Zealand. New
its of Upper Austria. Palaontologisches Zeitschrift, Zealand Journal of Geology & Geophysics, 18,
69, 503-524. 881-908.
VALDES, P. J., SPICER, R. A., SELLWOOD, B. W. & WETZEL, A. & UCHMAN, A. 2001. Sequential coloni-
PALMER, D. C. 2000. Understanding Past Climates: zation of muddy turbidites in the Eocene Beloveza
Modelling Ancient Weather. CD-ROM, Gordon & Formation, Carpathians, Poland. Palaeogeogra-
Breach Science Publishers/OPA, Amsterdam. phy, Palaeoclimatology, Palaeoecology,168, 171—
VOGEL, K. & MARINCOVICH, L. (in press) Paleo- 186.
bathymetric implications of microborings in WIDDICOMBE, S. & AUSTEN, M. C. 1998. Experimental
Tertiary strata of Alaska, USA. Palaeogeography, evidence for the role of Brissopsis lyifera (Forbes
Palaeodimatology, Palaeoecology. 1841) as a critical species in the maintenance of
VOGEL, K., BUNDSCHUH, M., GLAUB, I., HOFMANN, K., benthic diversity and the modification of sediment
RADTKE, G. & SCHMIDT, H. 1995. Hard substrate chemistry. Journal of Experimental Marine Biology
ichnocoenoses and their relations to light intensity and Ecology, 228, 241-255.
and marine bathymetry. Neues Jahrbuchfur Geolo- WIGNALL, P. B. 1994. Black Shales. Oxford Mono-
gie und Palaontologie. Abhandlungen, 195, 49—61. graphs on Geology and Geophysics, 30, Oxford
WALTER, H. 1980. Zur Kenntnis der Ichnia limnisch- Science Publications, Oxford.
terrestrischer Arthropoden des Rotliegenden. WRIGHT, A. J., YOUNG, G. C., TALENT, J. A. & LAURIE,
Freiberger Forschungsheft, C357, 61-68. J. R. (eds) 2000. Palaeobiogeography of Australa-
WALTER, H. 1982. Zur Ichnologie der Oberen Hornber- sian Faunas and Floras. Association of Australa-
ger Schichten des ostlichen Harzvorland. Freiber- sian Palaeontologists Memoirs, 23, 470pp.
ger Forschungsheft, C366, 45^48.
A new approach to the analysis and interpretation of
tracks: examples from the dinosauria
PHILLIP L. MANNING

The Manchester Museum, University of Manchester, Oxford Road,


Manchester, M13 9PL, UK

Abstract: Tracks can potentially offer unique sources of information, providing insight into
the environments, gait and posture, locomotion and behaviour. Track preservation can yield
important information on substrate consistency and enable the recognition of transmitted
subsurface tracks. The ability to recognize transmitted tracks has broad implications for
the understanding of palaeoenvironments and interpretation of ichnological assemblages.
In order to gain an understanding of how tracks are formed in three dimensions, and of
their variability of expression in different substrates, controlled laboratory simulations
were undertaken. Experiments were designed to recover subsurface track layers, yielding
for the first time detailed information on subsurface morphology that could be related to
'true' surface track features. It was found that subsurface track relief can be correlated
with the magnitude and distribution (across a foot) of load acting on the surface sediment.
This pressure is transmitted through the sediment, and deforms successive layers at depth,
producing an undertrack. The most significant factor controlling track morphology, whether
surface or subsurface, was found to be the moisture/density relationship within the substrate
at the time of track formation. Variability in the dimensions of simulated tracks, relative to
the 'true' surface track, indicates that caution should be exercised when using fossil tracks to
calculate hip height, speed, age, and population dynamics. In addition, comparison of experi-
mental tracks with dinosaur tracks from the Yorkshire coast suggests that many morpho-
logical differences between vertebrate ichnotaxa reflect sediment rheology and taphonomy
rather than taxonomy of the track-maker.

Fossil tracks have the potential to reveal informa- Field study of Middle Jurassic dinosaur tracks
tion on the size, gait and speed of individuals, as was undertaken on the Yorkshire Coast and
well as clues to their behaviour and the environ- compared with experimental tracks. Recent
ments in which the animals lived. However, the work has highlighted the importance of under-
interpretation of tracks is often difficult, in that standing and interpreting the formation and
what is available for study is, in many cases, not preservation of dinosaur tracks in the field
an original track surface. If fossil tracks are to (Romano & Whyte 2003 and references therein).
be a useful tool for interpreting behaviour and The diverse dinosaur track morphologies found
environments, it is essential that preservational within the Middle Jurassic track assemblages of
types are recognized and can be related to a the Yorkshire coast were studied with a view to
realistic surface trace. Improved understanding improving their interpretation through compari-
of track formation and preservation could also son with laboratory track simulations.
assist in diagnosing the properties and behaviour
of sediment at the time of track formation. This
study provides some insight into, and interpreta- History of experimental vertebrate
tion and understanding of, the complex, three- palaeoichnology
dimensional processes occurring beneath the
surface of a track (subsurface deformation) at The study of vertebrate tracks and traces - verte-
the time of track formation. Guidelines are also brate palaeoichnology - has concentrated on
provided for the interpretation of fossil tracks describing the trace, with little or no interpreta-
and their use. tion of track formation/preservation.
Experimental equipment was designed and The first laboratory investigations into verte-
built to optimize repeatability of experiments, brate track formation and preservation were
to control rheological conditions, and to enable carried out in December 1827 at Oxford Univer-
recovery of subsurface track layers. Foot tem- sity by William Buckland (Sarjeant 1974). Buck-
plates used were designed to be comparable to land persuaded a crocodile and then a tortoise
dinosaur feet, though a similar methodology to walk across a soft pie-crust (presumably of
could be used for any foot morphology or gait. dough) and also over wet sand and soft clay
From'. MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 93-123. 0305-8719/04/S 15.00 © The Geological Society
of London.
94 P. L. MANNING

Preservation of skin/scale impressions (Lock-


ley 1991).
Presence of 'messy tracks' in which clay adher-
ing to the foot distorts the foot outline (Bird
1944).
Detached mud clasts (presumed to have been
once adherent to the foot) within the track
(Whyte & Romano 1993a, 1994a; Allen 1997).
Collapse/flow features in which the track
margin overlies the sole of the track (e.g.
Lockley et al 1989; Allen 1997).
The presence of a raised displacement rim or
bourrelet, which is a direct reflection of the
sediment's consistency at the time of track
formation. Cohesive sediments tend to bulge
Fig. 1. Stacked tracks as recognized by Hitchcock into an unbroken, smooth rim, but more
(redrawn from Hitchcock 1858). friable sands tend to bulge into a radially
cracked displacement rim, and spill onto the
track surface (Thulborn 1990; Allen 1997).
(Buckland 1828). Later work by Hitchcock (1858)
on fossil tracks from the Lower Jurassic of A scenario that clouds this issue of surface track
Connecticut, USA, astutely recognized trans- recognition is created when a limb punctures into
mitted tracks (Fig. 1). It took over 100 years for underlying laminated strata, causing the presence
ichnologists to rediscover this phenomenon of an underprint in which only the deeper portion
through experimental work by Lingen & Andrews of the limb is impressed into the sediment (e.g.
(1969) on horse tracks. Thulborn 1990). These underprints are distinct
Subsequent work has refined our understand- from transmitted tracks (Fig. 1), which are the
ing of tracks and track formation through a primary focus of this paper.
variety of approaches.
Experimental neoichnology
Detailed study of vertebrate trace fossils A second approach to understanding trace fossil
preservation has come through observations of
A number of authors have studied deposits with known organisms in identifiable sedimento-
abundant well-preserved tracks from which a logical/rheological conditions. As mentioned
variety of broad-scale observations about sedi- above, experimentation has been a fundamental
ment conditions and track preservation have tool in vertebrate ichnology since the early
been made. It is generally considered that experiments of Buckland (1828). Recent
firmground conditions produce the best preser- advances in reptilian, avian and mammalian
vational conditions for surface tracks (e.g. neoichnology have come through careful consid-
Tucker & Burchette 1977) and that soft/soup- eration of foot anatomy and the kinematics of
ground conditions resulted in poor surface locomotion. Of particular importance to this
track preservation (e.g. Whyte & Romano study is the work on a variety of dinosaur-like
1981) (terminology for sediment consistency modern taxa, from komodo dragons to ostrich
from Dodd & Stanton 1990). Detailed study of (e.g. Padian & Olsen 1984; Demathieu 1987;
Neogene avian and mammalian ichnofaunas by Farlow 1989; Gatesy et al, 1999).
Scrivner & Bottjer (1986) documented wide Preservation of surface tracks is improved by
morphological diversity in artiodactyl tracks. microbial mat growth on the track surface
They attributed the difference in morphology to (Thulborn 1990) and, in desert environments,
variations in sediment water content at the time by moistening by dew before casting by sand
of formation. It was also noted by Scrivner & (McKee 1947).
Bottjer (1986) that surface track preservation
was improved by rapid casting by sand.
Application ofindenter theory
Determination of surf ace tracks Experimentation using artificial indenters in
order to understand track formation and mor-
The recognition of surface tracks is facilitated by: phology was first undertaken by Allen (1989).
CONTROLS ON PRESERVED TRACK MORPHOLOGY 95

He suggested that mechanical theory offered a was noted as a common phenomenon, with
number of insights into the likely character of underprints often differing greatly from the
animal tracks in the field. In particular, it surface tracks. They noted that digit III was
suggests that: probably the thickest in the track-maker's foot,
and normally produced a correspondingly
the track shaft is surrounded by a substantial broad impression. However, sometimes this prin-
deformed zone;
cipal load-bearing digit generated a thixotropic
the deformed zone is likely to include faulted
reaction in the underlying sediment, where on
as well as folded layers; and
withdrawal of the foot the walls of digit III
the character of the track is likely to vary with
were sucked inwards, generating a narrow
the stratigraphy of the affected sediment.
middle digit (cf. Figs 6c, 19). In addition the
Allen (1989) found that the use of an indented authors found that the foot did not sink into
plastic material in laboratory tests qualitatively the substrate during their T-phase or the W-
reproduced all the essential features of real phase, leading to gaps in the trackways.
tracks. In the laboratory experiments the limb The first attempt at interpreting complex
and foot cut a shaft into the sediment, creating three-dimensional surface failure in association
an extensive zone of deformed sediment around with dinosaur tracks was undertaken by Gatesy
and below the shaft; the degree of deformation et al (1999). Theropod tracks from the Fleming
increased with increased depth of penetration. Fjord Formation (Norian-Rhaetian), East
The deformed zone comprised an axial down- Greenland, were compared with tracks produced
fold, in which downward-decaying undertraces by a modern avian theropod, a helmeted guinea-
were preserved, and a marginal upfold, with fowl (Numida meleagris). The Triassic tracks
associated shear and fracture zones. Allen indicated that the animals had walked over
(1989) found that, where the sole of the foot substrates in quite variable conditions (dry to
was complex in shape, anatomical details - in saturated), producing a diverse assemblage.
the form of cross-folds (sensu Allen 1989) - The serial sectioning of tracks confirmed that
were preserved in the undertrack. Triassic theropod feet sank, moved forward
and were extracted with convergent toes, in a
similar fashion to guinea-fowl (Gatesy et al.
Extrapolation of biomechanics from tracks 1999).
Although great inroads have been made into
An initial contribution to the understanding of understanding vertebrate tracks, a thorough
tracks in relation to biomechanics was the study of variability of tracks in three dimensions
suggestion that the greater the force borne by a in relation to transmitted features is lacking.
limb, the deeper will be its underprint impression Presented herein is a first attempt to link surface
(Demathieu 1987). Hence 'relative foot [track] features described from dinosaur tracks Thul-
depth' (relative to the surface on which the born & Wade (1989), modern tracks (Allen
animal walked) was proposed as a crucial 1997; Gatesy et al. 1999) and the experimental
parameter for understanding the mechanics of indenter approach (Allen 1989) to the sub-
track formation, and for estimating the centre surface.
of gravity of track-makers. This statement pre- A continuous trackway from dry to saturated
dicts that deeper underprint impressions will sediment is easy to follow on a modern beach;
represent the dominant weight-bearing limbs, however, the interpretation of fossil tracks in
such as the manus or even particular parts of various stages of water cover is not always
the limb (e.g. distal portion of digits). obvious, especially with less complete track
Through study of a rich dinosaur track deposit material. The depth of the water may have
in Australia, Thulborn & Wade (1989) distin- affected a dinosaur's method of locomotion and
guished three distinct phases of track creation the resultant track morphology. The tracks of
during footfall, termed: wading dinosaurs would be quite different from
those of 'swimming' dinosaurs. The 'swimming'
touch down (T-phase);
(actually punting of a partially buoyant indivi-
weight bearing (W-phase); and
dual off the bed of a water body) animals may
kick-off (K-phase).
take longer strides, because they tended to be
Some of the morphological variability was buoyed up between one footfall and the next,
attributed to sediment consistency, but Thulborn and their tracks may show only the tips of the
& Wade (1989) also related some differences toes (Coombs 1980; Thulborn 1990; Whyte &
between tracks to variability in the footfall Romano 1994b; Romano & Whyte 1996).
cycle. Variation in the track shape with depth Current strength would also affect stride length.
96 P. L. MANNING

Experimental tracks
The methodology used herein is designed to
recover data from transmitted tracks generated
by experimentation. The transmitted tracks can
then be studied and compared with respect to
the sediment conditions under which they were
formed.
The analysis of the distribution of ground
reaction forces on indenter templates was under-
taken to assist interpretation of load and
pressure distribution during track formation
and its bearing on final track morphology. The
equipment used, an optical pedobaragraph,
also allowed determination of the centre of pres-
sure during footfall. This provided a dynamic
insight into the distribution of pressure across
the indenter foot for comparison with track
morphology.
Laboratory equipment included indenter tem-
plates, indenter assembly, Newton compression
meter, and various sediment-testing boxes in Fig. 2. Templates used in creation of surface tracks.
which sediments could be constructed and Surface areas for the templates were: Tl, 750.6mm2;
moisture content could be varied. Indenting a T2, 764mm2; T3, 624.2mm2; T4, 1294mm2; T5,
substrate to record the formational and preser- 1257mm2; T6, 757mm2; T7, 1005mm2.
vational history of a track appeared a simple
task at the outset. However, the surface features
were analogous to icebergs, with most of the similar to the tridactyl tracks of the Yorkshire
information locked beneath the surface of the Coast.
track as a complex, three-dimensional structure.
The indenter templates varied a great deal in Porous plate box
form and functional ability, allowing a study of Previous studies have only allowed the introduc-
the relationship between form, function and tion of fluids to a sediment from above, often dis-
substrate. The grain size and moisture content rupting or destroying any constructed laminae. A
were varied between individual track simula- better approach is to introduce water in a con-
tions, as Scrivner & Bottjer (1986) recognized trolled fashion from underneath via a porous
these as important variables affecting track plate. A large plastic box (with reinforced struts
preservation. to prevent deformation of the box under load)
had a porous plate inserted 40mm above the
floor of the box. The porous plate was con-
Experimental equipment structed from a piece of 5mm thick plastic,
drilled with a close, regular array of 3 mm dia-
Construction of indenter templates meter holes and covered in a fine (105 um) poly-
ester monofilament mesh before its insertion into
Both simple and complex shape templates were the box. Separate inlet and outlet valves were
cut from 50mm thick, rigid plastic. The shapes inserted into the box, beneath the level of the
were chosen to represent a variety of dinosaur porous plate. The outlet valve allowed the
pes morphologies in plan view. The surface water level to be monitored, and hence the
area of the seven (templates T1-T7) rigid, plastic saturation of the sediment could be controlled.
templates (Fig. 2) was calculated using a When the sediment had reached the desired
planimeter. saturation point, flooding of the sample could
This study primarily concerns a semi-rigid T8 be stopped immediately. When it was required
that consists of three converging steel bars to expel water from the sample, the hydrostatic
threaded into an aluminium heel with a moulded head tube was removed and the water could
silicone rubber outer. Silicone rubber was drain away freely (Fig. 3).
applied to the frame when held within a plaster The porous-plate testing box had to withstand
mould. The two-part mould had been formed temperatures of at least 100°C during oven
around a sculpted foot, of generic tridactyl type drying. The temperature was incrementally
CONTROLS ON PRESERVED TRACK MORPHOLOGY 97

allowing the template to be indented by hand


to a given force. The freedom of the NCM
allowed more natural movements of the tem-
plates to be mimicked, at the price of some
experimental control. The NCM was invaluable
with the porous plate generated tracks, as the
indenter assembly would have proved difficult
to apply within the confines of the testing box.
This method was employed for most of the
experiments discussed below.

Experimental methods

The optical pedobaragraph (OPB)


Fig. 3. The porous-plate testing box as used for
subsurface track simulations. The OPB experiments were either dynamic or
static loaded tests. In some experiments addi-
tional load was exerted on a specific digit or
increased from 30 °C to 100 °C over 8-10 days in digits to record possible effects of an uneven
an oven. gait. The OPB recorded the dynamically loaded
tests at a frame capture rate of either 25 frames
Indenter assembly per second (25 Hz) or 16.7 frames per second
An indenter assembly was constructed, and (16.7 Hz).
initially used in track simulations, so that the The apparatus consists of a glass plate illumi-
templates were driven into the sediments in the nated at two opposing edges by strip lights and
same way and at a controlled rate (Fig. 4). covered by a thin sheet of white deformable
The indenter assembly allowed a template to film, which is the surface on which a force can
be attached and pulled against a counterbalance be applied (Fig. 5). Light rays are totally intern-
by means of a spring balance, so that the ally reflected within the glass plate, except at
template indented the sediment below. The points where the white film touches the glass.
disadvantage of this assembly was that it gave At these points the light rays are refracted out
the template a forward shunt on initial contact of the glass plate and scatter back from the
with a sediment surface, and very unnatural white film. The film surface is undulating in
reverse motion on retraction of the template. It appearance on the microscopic scale, and an
was also difficult to generate large forces through increase in pressure on the film results in these
the template without the apparatus moving or undulations being deformed into intimate con-
lifting with the template as the pivot. tact with the glass plate. The greater the pressure
Owing to the natural rolling motion observed applied to the film, the greater the area of contact
in almost all bipedal vertebrates when walking, with the film and glass, and the greater the
a second indenting device was adapted to quantity of light that escapes from the glass.
mimic this motion. A Newton force compression
meter (NCM) was adapted so that a template
could be threaded onto the end of the device,

Fig. 4. Indenter assembly used for surface track


simulations. Fig. 5. The optical pedobaragraph.
98 P. L. MANNING

The result is a continuous grey-scale image of the the initial strike of an indenter to its toe-off.
indenter, in which intensity is proportional to the Pressure is expressed in kgcm~ 2 , which can
applied pressure (Betts et al. 1991). be converted to the official SI unit of pressure,
A colour image is generated from the digitized the pascal (Pa). The conversion factor is:
data, with the colour of the image being related lkgcnT 2 = 98.1kPa.
to the intensity and distribution of pressure at a It is also possible to calculate the progression
given time. From such images it is possible to of the centre of pressure throughout the sequence
see where high pressures occur, and to assess of frames, enabling assessment of the load-
the general pressure-distribution changes from bearing properties of any indenter, as well as to

Fig. 6. (a) Tridactyl dinosaur track (F00813) sectioned parallel to digit III. Scalby Formation, Scalby Bay,
Yorkshire, (b) Foot template 8 (T8) used in laboratory track simulations (scale 10cm). (c) Track simulation
T8/F13 layer 3, top surface of plaster track layer 3T, from a depth of 1.3cm below the surface track layer
(scale bar increments 1 cm), (d) Track simulation T8/D1.7b using T8, indented into sediment D with a force of
TON when dry (moisture content 0.3%). (e) Track simulation T8/D2.7 using T8 indented into sediment D with
a force of 49 N when moist (moisture content 7.3%).
CONTROLS ON PRESERVED TRACK MORPHOLOGY 99

produce a combined frames image. The com-


bined frames image is a useful indicator of pres-
sure distribution over an indenter throughout a
complete dynamic cycle, which can then be com-
pared with static loading of the same indenter.
Template 8 was tested in both static and
dynamic runs on the OPB. Sediment was also
placed on top of the transducer material to
measure the transmission of forces. Templates
1-7 were found to be too rigid for the OPB trans-
ducer material, and gave no usable data.
The results generated by the OPB provided
useful data on the relationship between the
distribution of load over the sole of a foot in
both static and dynamic load situations. Such
information is applied to the interpretation of
preservation features observed in both fossil
and laboratory-simulated tracks below.

Recovery of surface track features


Fig. 7. Generic tridactyl track showing position of
The initial indenter tests were undertaken to surface features recorded that relate to track
record surface features relating to the formation morphology: digit length and interdigital angle (IDA).
and preservation of tracks, using characterized,
homogeneous sediments. The sediment was
placed in a large tray 1 m long by 500 mm wide constructed for each track simulation. Kaffir
and 150mm deep. The sediment was at first D™ plaster of Paris (British Gypsum™) was
indented dry, and then moisture content was the material used to inter-laminate with the char-
increased incrementally. Moisture content was acterized sand and clay samples. Kaffir D™
calculated at the beginning and end of each plaster has suitable properties, including a setting
indenter test. For the surface tests the static time of 8 min, a high compressive strength (once
indenter assembly was used. Templates 1-8 dry), minimal effect on the behaviour of sur-
were indented into the substrate, with the sedi- rounding sediments, and a very low linear expan-
ment mixed and levelled before each experiment. sion on drying (around 0.25%). Other materials
A 300mm x 300mm wire grid of 10mm squares were tried, including cement, various muds and
was lightly impressed on the surface of the even flour, but none of them fulfilled all the
sediment before indentation, to emphasize the above criteria.
surface displacement features (e.g. Fig. 6d, e Laminated sediments were constructed when
below). The indenter force applied and the dry, as this enabled up to 16 layers of sand and
depth, width and length of each resultant track plaster to be constructed (sometimes including
were recorded, as was the slope angle of the laminae of damp clay sandwiched between dry
track wall and other distinct characteristics. sand), without affecting the dry plaster. The
Any transient features observed at the time of alternating layers of sand and plaster were indivi-
track formation were also noted, such as dually sieved into the porous-plate testing box,
dewatering of sediment around a track, or with the uppermost and lowermost layers
suction of the floor of a track above true track always consisting of sand. Equal volumes of sedi-
surface. The main surface features relating ment were used in all layers (approximately
directly to the foot are as shown in Figure 7. 900 ml per layer) and equal volumes of plaster
(approximately 200ml per layer). Plaster was
not sieved right up to the edges of the porous-
Recovery of subsurface track features plate testing box, so that water would pass
freely through the sand, around and between
A prerequisite for retrieving subsurface the plaster layers.
deformed track layers is that separable laminae Once the laminated sediment had been con-
be present. Homogeneous sediments were not structed, it could be indented either dry or moist
suitable for recording subsurface deformation, (by introducing water via the porous plate).
and so laminated sediments had to be There was a time interval of approximately
100 P. L. MANNING

4min (with Kaffir D™ Plaster of Paris) before defined as those that have not been transmitted
the plaster began to harden. In this time water from overlying sediments. In the laboratory-
had to be introduced via the porous plate, and simulated tracks this category encompasses
the sediment had to be indented. tracks from the top sand layer, infilled by the
As soon as a sample had been indented (using uppermost plaster layer and best seen as a cast.
template 8 for this set of experiments), four 3 mm The nomenclature used here is for laboratory-
thick copper rods were inserted vertically generated tracks, where the unique situation
through the sediment, puncturing each plaster exists of knowing the exact dimensions and mor-
layer immediately around the track to leave phology of the track-maker's pes. Nomenclature
reference points. A further thin layer of sand is used in this study to include the track dimen-
was added to the surface track feature to act as sions and angles recorded from track morphol-
a release agent, enabling an additional layer of ogy (Figs 7, 8a). Many of these terms have
wet plaster to be poured into the surface track been used to describe fossil tracks (Haubold
to record its morphology (relative to the copper 1984; Leonardi 1987; Gillette & Lockley 1989;
reference rods). When the top layer of plaster Thulborn 1990). It is difficult, if not impossible,
was dry, the copper rods were removed from to associate some features of fossil tracks with
the sample. the morphology of a track-maker's pes or
The porous-plate testing box, sediment and manus, owing to the vagaries of preservation.
track were placed in an oven at 30 °C for 24 h.
The temperature was then increased every daily
by 10 °C until the oven reached 100 °C. After Descriptive terminology for subsurface track
8—10 days the porous-plate testing box was morphology
removed from the oven, and the sample was
removed. The sample then had to be returned This terminology relates to transmitted track
to the oven for a period of 5 days to expel any features from the recovered subsurface layers
remaining moisture. The sediment had to be and puncture features associated with under-
completely dry because of the delicate plaster tracks (Fig. 8b). It is important that, if a term
layers, which are easily destroyed when damp. is used, the layer from which the feature is
In addition, cohesive sands hamper recovery of described is made clear, i.e. Aw, Bw, C" etc.,
plaster layers. where n is the layer number and surface type
When removed from the oven, the sides of the (upper or lower).
sample were brushed clear of sand until the layers Subsurface tracks were all produced using
of plaster appeared as relief ledges. The depth of template 8 (T8). As discussed earlier, the tem-
each layer, relative to the surface-track layer, was plate was very basic in form, representing the
measured. The lowermost sand layer was first simplified morphology of a theropod or very
brushed away, revealing a cast of the upper gracile ornithopod dinosaur pes. The proximal
surface of the lowermost horizon. Any feature convergence of the digits produced an unnatural
that was associated with transmitted or under- intersection of the median lines for each digit.
print features was mapped onto an acetate The median lines of dinosaur's digits do not
sheet and photographed. The reference points usually, if ever, converge in any example of a
made by the copper rods were also mapped dinosaur's skeletal pes. However, it was conveni-
onto the acetate sheet, and a track layer reference ent in the current study that the position of the
number was assigned. A spatula was then used to heel and digits of the template made measure-
gently lift the plaster layer, so that the lower sur- ments of track dimensions much easier, as well
face could be brushed clean, mapped onto an as being able to relate track morphologies
acetate sheet and photographed. The process of between layers more easily.
acetate sheet mapping and photographing was
repeated for each successive track layer.
Data recording
Track description Experiments were recorded using the abbrevia-
tion for the template followed by the experiment
Descriptive terminology for surface track number: for example, T8/F2 was the second
features experiment carried out using template 8 (T8).
Six types of data were recorded from the recov-
Surface track features are taken to include the ered plaster track layers.
upper surface of the base of shafts of undertracks The maximum zone of deformation (MZD)
(sensu stricto Thulborn 1990). Surface tracks are (Fig. 8b) was recorded for the top (T) and base
Fig. 8. (a) Track dimensions recorded for surface tracks; (b) features recorded from recovered subsurface layers. Data were used for subsequent ichnometric analysis
and measured relative to known points described by puncture holes made by copper rods.
102 P. L. MANNING

(B) of each plaster track layer (except for T8/F2, However, the volume (V) increases even faster,
where only one of the basal track surfaces could in proportion to the cube of its linear dimensions
be recorded). The coordinates were plotted for (/):
each track simulation on a standard grid
(300 mm x 300 mm), to allow comparison
between layers within an individual track and Each time the dimensions of T8 are doubled, the
their relationship with other track simulation surface area of the foot increases by a factor of 4
experiments. Individual track surface tracings (FL2) FL (Footlength) and the volume increases
were generated and placed in successive order, by a factor of 8 (FL ). This means that, if the
for ease of interpretation. The direction of dimensions of the template 8 are doubled, the
travel was marked on the first plot of each surface area increases from 470mm (125mm
track data series. long foot) to 1880mm2 (250mm long foot) and
The position of the 'heel' was recorded in all the weight of the animal (load) increases from
successive layers, as was the most anterior point 5kg to 40kg. This means that eight times the
attributable to the distal end of each digit (Fig. load is exerted over four times the area, so that
8b). The coordinates were recorded and plotted the pressure under the bigger foot is twice that
in the same way as for the MZD. of the smaller foot. The application of a 40 kg
Track length was recorded for the top and base load was not possible in the existing experimental
surfaces of each plaster track layer. The track testing frame, so the scaled load of 5kg was
length was taken as the length of the MZD applied. The larger foot would be dynamically
(Fig. 8b), as this was the most persistent track similar to the smaller one, but twice the load
length parameter to be recorded at any depth. would be transmitted through the sediment,
Track width was recorded for the top and base affecting the resulting scale but not the morphol-
surfaces of each plaster track layer. The width of ogy of a resultant track.
the track was taken as MZD track width, as It is impossible to achieve a quantitative test
shown on Figure 8b. The width of the MZD for all fossil dinosaur tracks as there are too
was taken as the maximum width of the track many variables to account for. These variables
as measured perpendicular to the track axis include the moisture content at the time of
(axis of digit III) (as shown on Fig. 8b). track formation, the weight of the dinosaur, the
Digit length was also studied, taking the length true morphology of the dinosaur's foot, and the
measured from the most anterior point attributa- exact gait of the dinosaur at the time of track
ble to the distal end of each digit to the posterior formation. Although these variables are con-
margin of the 'heel' from each track layer trolled in the current laboratory track simula-
(Fig. 8b). tions, it is impossible to know them from a
The IDA was measured as the angle of divari- fossil track. This means that the quantitative
cation at the point of intersection of the median data produced in this study can be used only as
axis of the digits about the 'heel' (Fig. 8b). a qualitative guide to the conditions prevailing,
and the foot morphology of the track-maker, at
the time of track formation.
Interpretation of results The magnitude of the maximum zone of defor-
mation (MZD) (e.g. Fig. 6d) and related features
Indenter theory was in proportion to the load applied to the
isotropic sediment. The MZD marked the distri-
The force exerted on T8 (Fig. 6b) during each of bution of vertical pressure at the point of failure
the subsurface track experiments was consider- at the surface and within the sediment. In trans-
ably smaller than those expected for a dinosaur verse cross-section the deformation has an
with a similar-sized foot. A load of 5 kg (49 N) onion-shaped force bulb, as reflected in the
was applied over the surface area of template 8 width of the MZD with depth (Fig. 9). In longi-
(470mm2), producing a force of 1.06kN/m2 tudinal section the force bulb is found to be
(1.06kPa). distorted, owing to the dynamic nature of track
When different-sized structures retain the same formation (Fig. 10).
shape, they are considered to scale with isometry These results are in line with the predictions
or to be geometrically similar (Swartz & of Boussinesq (1883), who solved the problem
Biewener 1992). The surface area (S) increases of stresses reproduced at any point in a homo-
in proportion to the square of its linear dimen- geneous, elastic and isotropic medium as the
sions (/): result of a point load applied on the surface
of an infinitely large half-space. Boussinesq's
elastic analysis is represented by the following
CONTROLS ON PRESERVED TRACK MORPHOLOGY 103

at a horizontal distance r from the line of action.


Application of Boussinesq's equation to a
hypothetical homogeneous sediment with a bear-
ing capacity of approximately 0.4 N m"2 loaded
with a 49 N force produces a failure zone (force
bulb) (Fig. 11) that is remarkably similar to
that seen during experimentation (Fig. 9).
Boussinesq's theory relates the distribution of
a static load at depth, but the current study
applied dynamic loading, and therefore the
application of Boussinesq's theory was qualita-
tive rather than quantitative. Variations from
Fig. 9. MZD length/depth plot for track simulation the expected onion-shaped bulb were produced
T8/F8 showing a characteristic onion-shape force by the experiments, owing to the dynamic
bulb. (forward-directed) loading (Fig. 10). One of the
longitudinally cross-sectioned fossil tracks
studied (F00813) also displayed a distorted force
bulb (Fig. 6a). Experiments T8/F6 and T8/F7
were the only interlaminated clays and sands
(non-homogeneous sediments) tested, and it
was observed that they also generated a distorted
force bulb consistent with possible failure predic-
tions using Boussinesq's theory.

Fig. 10. MZD length/depth plot for simulation T8/F8


displaying an anteriorly displaced force bulb. Kinematics
The standardized method for indenting template
T8 into the sediment was based on kinematic
analysis of modern bipedal vertebrates (Clark
& Alexander 1975; McMahon 1984; Gatesy &
Biewener 1991). As the laboratory simulations
were to be compared with tracks of bipedal dino-
saurs from the Middle Jurassic of the Yorkshire
where P is the point load (P), and <rv is the verti- coast, the force needed to be applied to the
cal stress at a point depth z below the point load experiment's sediment surface in a realistic

Fig. 11. Pressure bulb for a hypothetical 49 N impact on a homogeneous sediment, as predicted by
Boussinesq's equation. Values for av are in Nm~ 2 . Dotted line delineates failure zone (force bulb) for sediment
with a hypothetical bearing capacity of approximately 0.4 Nm~ 2 .
104 P. L. MANNING

manner. This was facilitated by use of the (1) the heel-down phase;
Newton compression meter, as described above. (2) the rotation phase, where the foot rolls for-
All known Middle Jurassic bipedal dinosaurs ward and the centre of mass of the animal
locomoted with a primitive hip-based retraction passes over the foot; and
mechanism for the movement of the hind-limb. (3) the toe-off phase, where the animal places its
The support phase of the hip-based step cycle weight on the distal ends of its digits before
(sensu Gatesy & Biewener 1991) has three distinct pushing off the ground (Fig. 12).
phases:
The angle of action of the force acting on the
foot at the heel-down phase of a limb cycle
appears remarkably similar among living bipedal
animals, and ranges from 56° to 73° (measured
from the horizontal), depending on the type of
animal and the speed at which it is travelling.
The angle to which the limb rotates forward
before the toe-off phase of the step cycle also
shows little variation between living bipedal ani-
mals, and ranges between 106° and 138° from the
horizontal in the direction of travel (Gatesy &
Biewener 1991; Fig. 12).
For the purpose of experimentation, the limb
angle at the heel-down phase of the step cycle
for this study was emplaced at approximately
70° (Fig. 12a). The limb was then loaded and
rotated to the toe-off phase and withdrawn at
an angle of 120° (Fig. 12c). It is noted that sub-
strate type and condition can cause organisms
to alter the angle of limb position at the heel-
down and toe-off phase of a step cycle. For
example, a slippery surface might necessitate a
deliberate flat placement of the foot (at a high
angle, approximately 90°) to prevent instability.

Subsurface failure
Indenting sediment causes a variety of failure
types. There are four distinct types of track-
related failure recognized in the present experi-
mental study:
In general shear failure, continuous failure sur-
faces develop between the edge of the indenter
and the sediment surface, causing distinct sur-
face anterior displacement rim. The distribu-
tion of pressure (load) through the sediment
is spread downwards and outwards, delineated
by the position of subsurface shear surfaces
(Fig. 13a);
In local shear failure there is a significant com-
pression of the sediment under the template/
foot. Associated failure surfaces (ISZ and dis-
placement rims) do not reach the soil surface
(Fig. 13b);
Puncture shear failure comprises vertical
shearing (failure) that occurs immediately
Fig. 12. The phases of track formation, showing the around the template, creating a shaft. There
limb angle at: (a) heel down; (b) forward rotation; is no surface development of failure surfaces
(c) toe-off; (d) withdrawal of foot. such as ISZ or displacement rims (Fig. 13c);
CONTROLS ON PRESERVED TRACK MORPHOLOGY 105

The word 'soil* has different meanings to workers


of various disciplines. The definition for soil used
herein is an engineering one, where a soil is
defined as any loose sedimentary deposit, such
as gravel, sand, clay or a mixture of these
materials (Smith 1981).
The size and distribution of soil particles and
the air or water occupying voids between the
solid particles affect the mechanical properties
of a soil, as do its porosity and permeability.
The sediments tested were silty-fine to medium-
grained sands that had little resistance to shear-
ing when dry, but when the moisture content
was increased so too did shear strength, up to
the critical hydraulic gradient. Increasing moist-
ure content effectively increases the bulk density
of sediment, as water replaces air contained in
the voids between soil particles. The denser a
soil becomes, the greater its shear strength
(Karafiath & Nowatzki 1978). However, if
moisture content increases beyond the soil's
critical saturation point (critical hydraulic gradi-
ent), where the soil particles no longer come into
contact owing to porewater pressure, the soil
fails and becomes a soupground (sensu Dodd &
Stanton 1990).

Track preservation and moisture/density


relationships
Fig. 13. Modes of failure: (a) general shear; (b) local
shear; (c) puncture shear (after Craig 1992); Footprints have the effect of mechanically com-
(d) liquefaction failure (Atkinson & Bransby 1978). pressing (densifying) a soil. Although the
Shear/failure surfaces are shown in dotted lines, and mechanisms that control compaction are not
displaced sediment is shaded dark grey. fully understood, it has been demonstrated that
soil moisture content at the time of compaction
is critical, especially for fine-grained soils such
Liquefaction failure occurs where areas of as silts (Karafiath & Nowatzki 1978). The sim-
intense pressure are created quickly (e.g. end plest and perhaps most widely accepted hypoth-
of push-off phase in a step cycle), causing the esis (Lambe 1961) for the relationship between
sediment to reach its liquid limit, whereupon moisture and density at a given compressional
the sediment flows into the track, destroying event is that, as water is added to a soil, air is
all evidence of foot morphology of the track- expelled. As soil particles adsorb water a surface
maker (Fig. 13d). film is formed that permits particles to slide over
each other more easily. Failure occurs when an
external load is applied if the critical saturation
Soil mechanics point of the soil is exceeded. As the thickness of
this water film is negligible compared with the
Track preservation is strongly related to the diameter of particles in coarse-grained soils, e.g.
mechanical and physical properties of a substrate sand-grade sediments, the effect is not as pro-
and their effect on a resultant track. A track is nounced as for finer-grained silty soils (Karafiath
formed when the yield strength of the substrate & Nowatzki 1978).
is exceeded owing to the locomotion of the In fine-grained soils the lubricating effect of
track-maker. The mechanical properties of a the water on the soil exists up to a certain thresh-
substrate directly influence the resultant features old. When additional water no longer replaces
associated with footfall. To fully understand the air in the soil voids, and the amount of entrapped
formation and preservation of tracks, an under- air remains essentially constant, the water has the
standing of the mechanics of soils is required. opposite effect: that is, it occupies pore space that
106 P. L. MANNING

could be filled with soil particles upon the appli- enabled comparison of the moisture/density rela-
cation of load (Karafiath & Nowatzki 1978). tionship of each simulation, as permeability
When a load is applied to such a sediment the could be treated as a constant.
water escapes between the soil particles. The The internal angle of shearing resistance of the
degree to which this occurs is controlled by the dry sands was dependent on the grading and
permeability of the sediment. The load is then relative density of each sample: therefore the
transferred from the virtually incompressible loose-packed character of each test run resulted
water to the compressible skeleton of soil parti- in a relatively low density. However, the intro-
cles, which takes up the load. duction of moisture to the sand had the effect
There is an optimum moisture content for a of initially reducing the volume of the sand
given soil under a given compaction energy that owing to consolidation, by increasing the relative
will allow the wetted soil particles to come as density of the sample. The position of the water
close together as possible. As the subsequent table relative to the indent of the template had
drying of the soil does not affect this spacing, an important effect on the ultimate bearing
the compaction of the soil at this optimum moist- capacity of the sands.
ure content results in a maximum dry density. The track simulations indicated that the bear-
The variation of dry density over a range of ing capacity of fine-grained sands increased with
mixing moisture contents constitutes the moist- water content, but the medium-grained sand
ure/density relationship (moisture condition decreased in strength with higher moisture con-
value) for a given soil compacted under a given tent. The saturated fine and medium-grained
type of load (related to foot morphology and sediments displayed comparable shear strength,
gait) and the amount of energy exerted on that as opposed to their dry state, where the fine-
load (a function of size and speed of organism). grained sediments were weaker. Non-cohesive
As moisture is removed (or added) from a silt soils, such as sand, have a low bearing capacity
to silty fine to medium-grained sand, as in this under static loading (British Standard 8004:
study, the sediment passes through a series of 1986. Given that the load (force) delivered by a
states: liquid, plastic, semi-solid and solid. The footfall has direction, the resulting failure has
transition from one state to another occurs at few comparisons in the field of soil mechanics,
points known as consistency limits (Smith 1981). owing to the speed and dynamic nature of
The important role that moisture content has the compaction event. However, the types of
in affecting (along with grain size) track mor- failure for static loading situations bear a
phology has been recognized before (Tucker & resemblance to the failure observed in the track
Burchette 1977; Scrivner & Bottjer 1986; Allen simulations in this study, where the template
1997). This study has, however, generated and indented the sediment beyond its ultimate
recovered well-defined subsurface tracks in both bearing capacity.
dry and saturated sediments, previously consid- The ultimate bearing capacity of a sediment is
ered poor preservational media (Tucker & defined as the minimum pressure required to
Burchette 1977; Scrivner & Bottjer 1986; cause shear failure along a surface (shear surface)
Gatesy et al. 1999). The poor surface develop- of the supporting soil immediately below or
ment of a track should, in many cases, be treated adjacent to a foundation (Craig 1992) or, in the
as an indicator that a definable subsurface track current study, under a foot. Prandtl (1920,
feature has been transmitted or punctured at 1921) recognized that the failure surface was
depth. not linear throughout the soil, but consisted of
active and passive Rankine zones separated by
a radial zone (Fig. 14).
Permeability and residual strength Craig (1992) identified three distinct modes of
failure (Fig. 13a-c): general shear, local shear
The sediments chosen for the laboratory track and puncture shear. The track morphologies
simulations were within the range of silty fine and surrounding maximum zone of deformation
to medium-grain sands, but clays and silts were (MZD) in the current study can be explained as
additionally used in some simulations (T8/F6 & the result of one or a combination of the three
T8/F7). failure modes identified by Craig (1992). The
The values of permeability for the sediments MZD of each of the track simulations and the
used in this study were typical for silty fine to MZD length/depth and MZD width/depth
medium-grade sands (from 1.3xlO~ 0 5 to plots help differentiate between modes of failure.
2.4 x lO^^ms" 1 ), displaying moderate to high In the case of general shear failure, continuous
permeability (Atkinson & Bransby 1978). The failure surfaces develop between the edges of the
similarity in permeability of the sediments template and the ground surface (Fig. 13a). As
CONTROLS ON PRESERVED TRACK MORPHOLOGY 107

Fig. 14. Shear surfaces (failure surface) beneath a loading point, showing the relative position of the Rankine
zones: I, active Rankine zone; II, radial Rankine zone, arrows indicate direction of force dissipation; III,
passive Rankine zone, arrows depict direction of sediment displacement; IV, displacement rim (after Karafiath
& Nowatzki 1978).

Fig. 15. Shear surfaces as developed in a soil with an upper strong sand underlain by a weaker sand, showing
the preferential development of local shear zones in the weaker sediment (after Braja 1994).

pressure increases toward the bearing capacity of the liquid limit is reached and sediment is
the sediment, a state of plastic equilibrium is saturated, causing liquefaction and flow (Fig.
reached, initially in the sediment around the 13d), resulting in contorted laminae (when pre-
edges of the template and gradually spreading sent) and squelch features (Tucker & Burchette
downwards and outwards. When the condition 1977).
of plastic equilibrium has been reached in a Sediment strength affects its bearing capacity
sediment, shear failure is imminent through the and the scale of shear failure (Braja 1994).
whole sediment mass (Craig 1992). Ultimately When a strong layer under a point load is
the state of plastic equilibrium is fully developed relatively thin, failure takes place by the force
through the sediment above the failure surfaces. being transmitted through the strong sand
Heaving (bulging) of the ground surface occurs layer, followed by general shear failure in the
on all sides of the template. In the present experi- underlying, weaker sand layer (Fig. 15). When
ments the effect is most prominent anterior to an upper strong sand layer is relatively thick,
and between digits II and III, and III and IV. failure may be fully located in the strong sand
Local shear failure is typified by a significant (Fig. 14).
compression of the soil under the template, Static loading of sediment causes predictable
accompanied by only a partial development of failure at an ultimate bearing capacity, but a
the state of plastic equilibrium. The failure sur- dynamic load causes variation in the distribution
faces do not reach the ground surface, and only of pressure through sediment. Tracks are formed
slight heave occurs. Local shear failure (Fig. by the action of such dynamic loads, distorting
13b) is associated with soils of high compressi- the distribution of forces transmitted through
bility, and is characterized by the occurrence sediment, in the direction of travel of the
of relatively large settlements. Puncture shear animal. This is further complicated by the move-
failure (Fig. 13c) occurs when there is relatively ment of the foot in relation to the body and the
high compression of the soil under the template, kinematics of the step cycle (Fig. 12).
accompanied by shearing in the vertical direction
around the edges of the template. There is no
heaving of the ground surface away from the Experimental results
edges. This study additionally included a fourth
mode of track failure, liquefaction failure The tracks generated in the laboratory-
(Atkinson & Bransby 1978). This occurs when controlled conditions yielded a variety of track
108 P. L. MANNING

morphologies (surface and subsurface) indicative A displacement rim was commonly generated
of the prevailing rheological conditions. along the posterior margin of digit IV (Fig. 6d).
Several of the laboratory-simulated tracks This was a result of an unnatural posterior
(T8/F4, T8/F9, T8/F11, T8/F13) were formed shunt caused by the indenter assembly used for
while the sediment was saturated. The rapid the surface track simulations. It was because of
loading associated with track formation caused this unnatural type of locomotion and the static
excess pore pressure to develop within the sedi- step cycle that the indenter assembly was not
ment, as the fluid in the pore spaces had limited used for subsurface track simulations, and the
travel at its predetermined rate of permeability NCM assembly was used instead.
(&), and was therefore unable to keep up with The surface track simulations also generated
the rate of loading. Thus, in some cases, track radiating surface tension features within a moist-
formation generated excess pore pressure in the ure content range of 3-10%. These radiating
sediments, which caused liquefaction failure features were associated with the posterior
(Fig. 13d), whereupon the sediment could poten- margin of the heel, distal end of digits, and
tially flow under its own weight (Smith 1981), as hypices of the track. A radial tension feature
observed in track simulation T8/F13 at a depth (radial crack) anterior to digit III was observed
of 15mm (Fig. 6c). on several tracks (Fig. 6e), and is interpreted as
Dinosaur tracks cannot be described in terms a surface development of an ISZ. The radial
of simple soil mechanics, although the mode of tension features occurred at a sediment moisture
failure (Craig 1992; Atkinson & Bransby 1978) content of between 4% and 8% and was poorly
for the sediment can be recognized. developed, although a larger load would increase
the development of the feature, as seen in the ISZ
feature in Figure 6d. The failure surface for this
Laboratory-simulated surface tracks feature was considered brittle, as the surface
feature did not plastically deform, more com-
The surface track simulations indicated that the monly observed with higher moisture contents.
shape and rigidity of a template affected the Recognizable surface tracks were formed at a
behaviour of sediment into which it was indented. sediment moisture content range of 2—24%,
The initial eight templates used had flat, rigid beyond which surface track features collapsed.
surfaces, except for T8. The flexible digits of The coarser-grained sands retained track defini-
template 8 meant that, on impact with sediment, tion at the highest moisture contents (up to
penetration was to a greater depth, resulting in 25% moisture content), whereas fine-grained
deeper tracks than for any of the other templates. sands collapsed at a moisture content of 25%.
The shape of template 8 promoted the develop- The relationship between moisture content and
ment of radial shear zones (Fig. 14), indicative grading of the sediment suggested that, as grain
of general and local shear failure. However, size increased, the liquid limit of the sediment
templates 1-7 compacted sediment beneath also increased; the opposite applied to finer-
them (in the active Rankine zone: Fig. 14) result- grained sediments.
ing in puncture shear failure (Fig. 13c). Fine-grained sediments retained an outline of
It was decided at an early stage of the study the track, even if some of the surface track
that the behaviour of the sediment and resultant digits had partially collapsed. However, coarser-
tracks created by T8 was most natural, so the grained sediments quickly collapsed (<10min),
template was used for all subsequent simulations. leaving a low-relief surface track feature. The
A feature commonly recorded in dry and occa- collapse of the tracks was related to the perme-
sionally for some moist sediments (<2% moist- ability of the indented sediment.
ure content) was the development of interdigital Medium-grained sediments had a higher per-
shear zone (ISZ) features (Fig. 6d). The surface meability than the fine-grained sediments. The
ISZ features tended to be more developed in coefficient of permeability of the fine-grained
dry sediments (<1% moisture content), some- sediment (sediment A) used in surface track
times resulting in multiple ISZ features (Fig. simulations T8/A1.7-T8/A4.7b (Fig. 16a-d),
6d): these were usually observed as a subsurface was in the order of 1.36 x KT^ms"1. The
features. The surface development of multiple coefficient of permeability of the medium-
ISZ features was usually due to an increase in grained sediment (sediment D) used in surface
the load applied through the template, allowing track simulations T8/D1.7-T8/D4.7 (Fig. 6d),
the surface development of subsurface shear was in the order of SxKT^ms" 1 . Track
fronts, as a result of a more developed general features in medium-grained sediments flooded
shear failure than that observed with smaller more rapidly than those in fine-grained sedi-
loads. ments. This caused surface track features to
CONTROLS ON PRESERVED TRACK MORPHOLOGY 109

Fig. 16. (a) Track simulation T8/A1.7 using T8 indented into sediment A with a force of 49 N when dry
(moisture content 0.2%). (b) Track simulation T8/A2.7 using T8 indented into sediment A with a force of 49 N
when moist (moisture content 4.1%). (c) Track simulation T8/A3.7 using T8 indented into sediment A with a
force of 49 N when moderately moist (moisture content 26%). (d) Track simulation T8/A4.7b using T8
indented into sediment A with a force of 49N when sediment was saturated (moisture content 29.3%).

collapse and infill with water, resulting in the permits the particles to slide over each other
leaching of a fine intergranular component of more easily when an external load is applied.
the sediment into the track (matrix infiltration).
The fine-grained sediments used in surface
track simulations allowed the template to indent Laboratory-simulated subsurface tracks
the sediment to a depth of 110mm when satu-
rated (Fig. 16d). However, similar moisture Template 8 was applied using the NCM appara-
contents (30%) for coarser-grained sediments tus for all the subsurface simulations, with a load
allowed the same template, under the same of 5 kg applied during each experimental run.
load, to indent to a depth of only 50 mm. The Dry fine-grained sands (sediments A, B and E)
strength of the sediment decreased with increas- had lower shear strength values than dry
ing moisture content, although a larger grain medium-grained sands (sediments C, D and F),
size increased the threshold at which the optimum which allowed template 8 to affect layers up to a
moisture content of the sediment is reached. This maximum of 92mm below the surface track
effectively increases the strength of coarser- layer. The minimum affected sediment depth for
grained sediments, as the water film around the the template in dry fine-grained sediments was
sediment particles is negligible compared with 75mm below the surface track layer. The
the diameter of the sand grains. However, the dry, fine-grained sediments of track simulations
surface film of water on fine-grained sands T8/F2, T8/F10 and T8/F12 all displayed features
110 P. L. MANNING

typical of general shear failure (similar to Fig. was the transmitted position of overlaying
6d). Well-developed interdigital shear (ISZ) posterior margins of digits II and IV, and was
surfaces spread downwards and in front of the named a multiple digital transmission (MDT)
tracks, and an anterior displacement rim was feature (Fig. 8b). The MDT feature was observed
displayed on the surface track layer of track simu- only as a transmitted subsurface feature, and
lations T8/F10 and T8/F12. However, track with dry, fine-grained sands occurred on layers
simulation T8/F2 showed surface puncture of at depths of 30-60 mm. The deeper MDT feature
the top track layer caused by the template, appar- (92mm) observed with track simulation T8/F2
ent by the presence of separate entrance and exit resulted from template 8 puncturing layer 9 by
points for digits II and IV. The depth of puncture up to 40 mm, generating a deeper MDT feature.
was not more than 40 mm, or the template would Key features of track simulations in dry, fine-
have punctured layer 5 (Fig. 17b). grained sediments:
A distinct subsurface feature was observed
on the anterior track margins of all the dry, Lower shear strength than dry medium-
fine-grained sands tested (Fig. 17a). The feature grained sediment.

Fig. 17. (a) Track simulation T8/F2, layer 2, top surface of plaster track layer 3 at a depth of 8 cm below the
surface track layer, showing multiple digital transmission (MDT) features and well-developed interdigital shear
zones, (b) Track simulation T8/F3, layer 5, basal surface of plaster track layer, at a depth of 2.9 cm below the
surface track layer with interdigital shear zones, (c) Track simulation T8/F7, layer 9, basal surface of plaster
track layer at a depth of 8 cm below the surface track layer. Shows collapse of digits and posteriorly directed
projections (cf. Fig. 21b). (d) Track simulation T8/F6, layer C, basal surface of plaster track layer from a depth
of 2.3 cm below the surface track layer.
CONTROLS ON PRESERVED TRACK MORPHOLOGY 111

Features deeper and better developed (MDT, of these sediments. The saturated strength of
ISZ) in fine-grained than medium-grained the fine-grained sands was greater than that of
sediment. the dry sediments. However, the saturated
Simulations typically displayed general shear strength of the medium-grained sand decreased,
failure. compared with its dry state. Track simulation
Well-developed ISZ and displacement rim T8/F7 (Fig. 17c) was an exception to this trend,
features. inasmuch as it exhibited a decrease in strength,
Digital positions transmitted to 69-92 mm with a maximum depth track feature visible at
depth. 99mm below the surface. However, this was
Track relief increased anteriorly from 30— due to puncture failure to a depth of 58mm,
40 mm depth. owing to the influence of interlaminated clays
Distal end of digit III compacted track layers in this track simulation, given that the failure
to a depth of 60-90 mm. terminated at a clay horizon.
MDT feature occurred at 30-60 mm depth. Key features of track simulations in fine-
Clear anterior track displacement with increas- grained saturated sediment:
ing depth.
Increased shear strength compared with dry
Increasing interdigital angle with depth.
state.
Decrease in digit length with depth.
Liquefaction failure displayed at depths up to
For dry, medium-grained sands (sediments C, D 15mm.
and F), track simulations influenced layers up to a Local shear failure occurred from 15mm
maximum depth of only 65 mm below the surface downwards.
track layer. The minimum depth of influence in dry Poorly developed (low-relief) ISZ and dis-
medium-grained sediments was 46 mm below the placement rim features.
surface track layer. Simulations T8/F3 and T8/ Digital positions transmitted to a depth of
F8 displayed features typical of general shear 49-57 mm.
failure. Well-developed interdigital shear (ISZ) Track relief increased anteriorly from a depth
surfaces spreading downwards and forwards of 30 mm.
were also evident in the tracks, combined with an Distal end of digit III was compacted to a
anterior displacement rim displayed at the surface depth of 60 mm.
of track layer 7 of simulation T8/F3. The ISZ MDT features occurred at 33-52 mm depth.
features of simulation T8/F3 (Fig. 17b) were well Anterior track displacement occurred at
developed in layer 5B, giving the misleading depths below liquefaction or puncture failure.
appearance of interdigital webbing. Interdigital angles and digit length varied with
Key features of track simulations in dry, depth, displaying no discernible trend.
medium-grained sediments: Saturated medium-grained sands (sediments C,
Simulations typically displayed general shear D and F) exhibited shear strengths comparable
failure. to those of saturated fine-grained sands (sedi-
Features shallower and less developed than in ments A, B and E). However, the shear strength
fine-grained sediment. of the medium-grained sands decreased when
Well-developed ISZ feature and sometimes an saturated (track simulations T8/F4 and T8/F9),
anterior displacement rim. compared with the dry strength of the sediments.
Digital positions transmitted to a depth of Saturated medium-grained sediments had lower
39-45 mm. shear strength than dry medium-grained sands
Track relief increased anteriorly from 29- (sediments C, D and F), allowing template 8 to
32 mm depth. affect layers up to a maximum of 94 mm below
Distal end of digit III compacted track layers the surface track layer (layer 9 of simulation
to depth of 39-54 mm. T8/F4). The minimum affected sediment depth
MDT feature occurred at 32-56 mm depth. for the template in saturated medium-grained
Tracks displayed clear anterior displacement sediments was 40mm below the surface track
with increasing depth. layer of simulation T8/F6. The saturated sedi-
Interdigital angle increased with depth. ment (F) used for track simulation T8/F6 was
Digit length decreased in length with depth. medium-grained; however, an underlying layer
(10 mm thick) of fine clay prevented the transmis-
Saturated fine-grained sands (sediments A, B sion of track features beyond a depth of 40 mm
and E) had shear strengths comparable to those (Fig. 17d).
of the saturated medium-grained sands (sedi- Key features of track simulations in medium-
ments C, D and F), in contrast to the dry state grained, saturated sediments:
112 P. L. MANNING

Shear strength decreased compared with dry Dinosaur anatomy and gait
state of sediment.
Surface layers (up to 33-34 mm depth) display The form of the vertebrate skeleton has a direct
puncture failure. influence on the function of a limb, as it provides
General shear failure occurs below 34mm the anchor points for the musculature that drives
depth. locomotion, and delineates the degree of possible
Well to poorly developed ISZ features and dis- movements of a limb. The body fossil record
placement rims. preserves information on the skeletal anatomy,
Digital positions transmitted to a depth of size, and hence the inferred gait of some
40-66 mm. dinosaurs. This has enabled workers to generate
Track relief increased anteriorly from a depth functional models and to reconstruct the loco-
of 3 5-46 mm. motion of many dinosaurs (Romer 1923; Alexan-
Distal end of digit III compacted to a depth of der 1985; Norman 1986; Johnson & Ostrom
60mm. 1995).
Development of MDT features was restricted
to 10-15 mm depth.
Anterior track displacement occurred at Comparison of experimental tracks and OPB
depths below puncture failure.
Interdigital angle and digit length displayed data
little variation with depth. The results from the optical pedobaragraph
(OPB) confirmed that the distribution of pressure
over the sole of T8 corresponded with the loca-
tion of many track features. High pressure
Discussion values corresponded with high-relief features,
and conversely low pressures with shallow or
This study recognizes two main groups of influ- missing features were associated with the original
ences on the morphology of dinosaur tracks, template morphology.
whether fossil or laboratory-simulated: (1) kine- The dynamically loaded OPB track simula-
matic and morphological influences of the tions all recorded the highest point of pressure
animal on the track, and (2) preservational exerted over the sole of the template beneath
influences of rheological conditions etc. on the the distal end of digit III. A consistent feature
track. observed in almost all the subsurface track simu-
Track-maker morphology influences the size, lations was the compaction of track layers at
gait, step cycle (kinematics) and distribution depth under the area of the distal end of digit
of weight over a foot, affecting final track III. The OPB traces also indicate that the main
formation. The preservational influences load-bearing digits in both static and dynamic
account for the mechanical and physical prop- loading situations were digits II and III (III
erties of a substrate and their effect on resultant being the dominant load-bearing digit), with
tracks. digit IV often leaving little or no pressure trace.
The laboratory-simulated tracks provided The most common digits to be transmitted to
information not only on the preservation of lower subsurface layers in the laboratory track
tracks, but also on the influence of track mor- simulation were likewise digits II and III.
phology.
How an animal interacts with a substrate is
integral to understanding any resultant preserva-
tional features. How an organism's foot interacts Comparison of laboratory-simulated tracks
is dependent upon the anatomy of its limb, the with fossil tracks
morphology of the foot, the movement of a
limb during the step cycle, and the speed and The current study interpreted the morphological
behaviour of the animal at the time of track features generated in laboratory-simulated
formation. tracks by a tridactyl template, while varying
The substrate in which tracks are made also sediment grain size and moisture content. The
affects their formation, inhibiting or enhancing Yorkshire Coast Middle Jurassic track assem-
the animal's ability to traverse the substrate, blage is dominated by tridactyl tracks (see
and the gait adopted in some cases. The com- Romano & Whyte 2003 for review), and many
position, properties and condition of a substrate of the preservation features observed in the
also have implications for the preservation tridactyl tracks were comparable to those from
potential of a track. the laboratory simulations.
CONTROLS ON PRESERVED TRACK MORPHOLOGY 113

Burniston dinosaur tracks The tracks displayed a variety of features eom-


Tridactyl dinosaur tracks from the Middle parable to some of the laboratory-simulated
Jurassic Scalby Formation, Burniston Bay, tracks, generating seven categories of tracks
North Yorkshire, were observed in the field, studied at this locality:

Fig. 18. (a) Tridactyl dinosaur track (A) and additional digit (B) from an adjoining track. Track preserved as a
cast feature (5 m south of steps, towards Cromer Point), Burniston Footprint Bed, Scalby Formation,
Long Nab Member, Burniston Bay, Yorkshire (scale bar 10cm). (b) Track simulation T8/F4, layer 4,
basal surface of plaster track layer (4B) from a depth of 4.6cm below the surface track layer, showing distinct
anterior displacement (scale bar 10cm). (c) Tridactyl dinosaur track A displays pronounced digits II and III
with digit II leaving almost no mark. Tridactyl dinosaur track B displays collapsed digits, leaving only a
pronounced 'heeF cast. Tracks preserved as cast features on the sole of a sandstone bed. Burniston Bay
(15m south of steps, towards Cromer point), Burniston Footprint Bed, Scalby Formation, Long Nab Member,
Burniston Bay, Yorkshire, (d) Detail of a single track preserved on the underside of a sandstone block as cast
in relief. Burniston Bay, Scalby Formation, Long Nab Member, Burniston Bay, Yorkshire. Radiating from the
track and related to its formation are desiccation track-like features.
114 P. L. MANNING

1. High moisture content. The single tridactyl


track (A) from the Scalby Formation (Fig. 18a)
- with an additional digit (B) cutting into the
complete track - shows a faint interdigital
shear zone feature (ISZ). Similar surface collapse
features were observed with simulation T8/D4.7,
when medium-grained sand was saturated at the
time of indentation. The surface track indicates
that the foot of the dinosaur penetrated at least
as far as the upper surface of the track block,
which subsequently collapsed on withdrawal of
the foot.
The basal tridactyl track (A) increases in relief
anteriorly, indicating that it was transmitted
from a depth within the thickness of the track
block (70-80 mm). Laboratory-simulated tracks
in similar saturated medium-grained sediments Fig. 20. Sketch of the upper surface of a track block
in simulation T8/F4 (Fig. 18b) displayed punc- displaying collapsed digits. The faint interdigital shear
ture failure, with collapsed digit features similar zone (ISZ) is highlighted.
to track A.
As measured from the most posterior point of
the heel to the distal end of digit III on the top A second specimen shows collapsed digits and
surface, the track length of the specimen in development of a faint ISZ feature (Fig. 20) on
Figure 18a is 150mm. This increases to 175mm the upper surface of a track block, indicating
on the underside of the block, and shows an ante- saturated conditions at the time of track forma-
rior displacement of at least 25 mm relative to the tion (cf. simulation T8/F4: Fig. 18b). The trans-
faint tridactyl track on the upper surface. The mitted track on the basal surface of the block
increase in track length on the underside of the displays increasing relief anteriorly and no ISZ
block could have been caused by an underlying or MDT features; however, the median line of
layer of clay. Similar increase in track length digit III indicates that it has collapsed.
was observed in simulation T8/F6, which 2. Moisture content 20% (saturated). Several
diverted the downward-directed transmission tridactyl dinosaur tracks from the Burniston
forces, causing anterior displacement of the Footprint Bed had only partial digits preserved
weaker sediment above the lowermost clay. The (Fig. 18c), displaying collapsed distal ends to
fossil track (Fig. 18a) shows evidence of having digits, often leaving pronounced heel casts. The
been transmitted to the top of a thick clay Burniston Footprint Bed sandstones (fine- to
layer, which has subsequently been weathered, medium-grained) are underlain by silty shale,
leaving traces of the clay adhering to the margins which played an important role in the excellent
of the track digits. Such a track is likely to have preservation displayed by many of the dinosaur
been formed as shown diagrammatically in tracks from this locality. The marked phalangeal
Figure 19 through comparison with laboratory nodes (pads) of track digits and presence of heel
simulation T8/F6. impressions indicate that the tracks were formed

Fig. 19. Hypothetical cross-section of a track along the median line of digit III, showing variation in track
length (Li surface track and L2 basal track) and displacement (D) with depth. The diagram is based on the
fossil specimen in Figure 6a and the experimental track in Figure 18b.
CONTROLS ON PRESERVED TRACK MORPHOLOGY 115

near, but not at, the surface. The partial collapse experimentation, the compacted area of sand
of the most distal end of digits and the complete was pushed into the clay layer, producing well-
collapse of digits indicate that tracks were defined tracks (Fig. 17d); however, the phalan-
formed at different moisture conditions (approxi- geal nodes (pads) are better defined at a lower
mately 20% and 30% respectively). None of the moisture content (Fig. 6e), which suggests that
tracks shows any transmitted features (such as tracks with pronounced heels and collapsed
relief increasing anteriorly with depth or MDT digits formed while the sediment had a high
features), and they display a consistent depth of moisture content (saturated), and that the
relief for the length of track features. better-delineated phalangeal tracks formed at
The dinosaurs that made the Burniston tracks lower moisture contents (c. 20%).
walked over a fine- to medium-grained sand, into 3. Transmitted tracks with decreasing moisture
which their feet sunk by approximately 10- content. Detail of one of the tracks studied
40mm, until encountering the stronger sub- (Fig. 18d) shows that the desiccation cracks
surface silty horizon, which plastically deformed, often originate from the distal end of track
rather than being punctured. The tracks with a digits, indicating that the tracks formed before
distinct heel but collapsed digits were the the desiccation cracks. The tridactyl tracks
shallower features (10-20 mm), where sand col- (Fig. 18d) increase in relief anteriorly and are
lapsed around the digits (Fig. 18c) on extraction poorly defined, indicating that they were trans-
of the foot, leaving only a distinct heel impres- mitted features. The distal ends of the digits
sion (cast). The well-defined transmitted tracks were joined by what might be the transmitted
show distal or no collapse features on digits. remnant of an interdigital shear zone (ISZ)
These were deeper (20-40 mm) track features, feature. The distal end of digit III (Fig. 18d)
where the foot of the dinosaur compacted the undercut the siltstone into which the basal cast
sand under its feet into the silty horizon, along feature was transmitted, from within the 600-
the whole length of its foot. There were no 700 mm depth of the sandstone block. The silt-
shear features (ISZ or MDT) associated with stone probably acted as a termination point for
these particular tracks from Burniston. In terms transmitted forces from the footfall, owing to
of soil mechanics, the pressure of footfall - its relatively high shear strength compared with
equivalent to the active Rankine zone (Fig. 20) the overlying sands.
- compacted an image of the foot onto the A similar transmitted track was observed on a
upper surface of the silt. However, the greater basal clay layer of simulation T8/F6. The track
shear strength of the silts did not permit the was formed while an overlying, medium-grained
development of the radial shear zone, resulting sand layer was saturated (as was the underlying
in a well-defined track image. clay layer). The clay layer had the effect of divert-
The absence of a toe-off phase feature in any ing the force bulb laterally, expanding the track's
of the tracks (e.g. Fig. 18c) suggests that the dimensions, and causing the distal end of the
substrate may have affected the way in which ani- digits to almost join. On drying, desiccation
mals traversed the area. It was noted that, during cracks formed in the clay 40mm below the
experimentation, when T8 was applied in a three- sand. This indicates that desiccation cracks are
phase step cycle the template would slide on not necessarily a good indicator for recognition
moist or saturated clays underlying a 20-40 mm of a true surface track horizon.
thick sand. Only when the template was exerted 4. Dry sands punctured to lower horizons with
at right angles to the sediment (simulation T8/ higher moisture content. An excellent tridactyl
F6) did the template leave a stable track, which pes track (Fig. 2la) was observed on the under-
in turn allowed the recovery of subsurface track side of a fine-grained sandstone block preserved
layers. in hyporelief. Associated with the transmitted
Track simulation T8/F6, in saturated medium- pes imprint, an anterior displacement rim -
grained sands underlain by a clay layer (at depth comprising multiple interdigital shear zones
40 mm), exhibited many track features observed (ISZ) - resulting in an extensive maximum zone
in the Burniston Footprint Bed (Fig. 17d). The of deformation (MZD).
pronounced heel, with collapsed digits formed The distal end of digit III clearly displays a
as a near-surface feature, in which the heel broad hoof impression (Fig. 2la), typical of
compacted the sands beneath it, but did not those expected of an ornithopod dinosaur, such
puncture the underlying clays, the overlying as Camptosaurus from the Middle Jurassic
saturated sand caused the digits to collapse (Weishampel et al. 1990). The sediment between
on withdrawal of the template. The resulting and anterior to digits II and III displays a faint
track was almost identical to some Burniston ISZ feature, but this is swamped by the anteriorly
Footprint Bed tracks (e.g. Fig. 18c). During displaced, concentric failure feature. The
116 P. L. MANNING

Fig. 21. (a) Tridactyl pes (right foot) preserved as a cast in hyporelief, associated with an anterior displacement
rim consisting of several concentric failure surfaces, yielding an extensive maximum zone of deformation
(MZD) around the pes. A well-defined 'hoof is also present on digit III, which has punctured to the base of
this sandstone. From the Scalby Formation, Long Nab Member, Burniston Bay, Yorkshire (scale bar 10cm).
(b) Transmitted tridactyl dinosaur track, preserved on the underside of a sandstone block as a cast in
hyporelief, with digit IV only partially recovered. Arrow indicates a shear surface in the interdigital shear zone
(ISZ). Compare the distinct posterior projection with that in Figure 17c. From the Scalby Formation, Long
Nab Member, Burniston Bay, Yorkshire, (c) Large tridactyl dinosaur track with associated smaller tridactyl
dinosaur track (top left) preserved as a cast in hyporelief. A faint interdigital shear zone (ISZ) feature is
present, and is cut by the isolated digit. From the Scalby Formation, Long Nab Member, Scalby Bay,
Yorkshire (scale bar 10cm).

anterior displacement rim is a development of performed upon a dry, fine-grained sand, sug-
the combined ISZ features of digits IV-III and gesting similar soil conditions for the fossil
III-II, resulting in a combined multiple inter- track. The presence of such well-developed ISZ
digital shear zone feature. features was a typical feature of dry, fine-grained
The type of failure seen in the fossil track was track simulations. The broad hoof (digit III)
observed in track simulation T8/F2, which was marks the maximum penetration of the foot
CONTROLS ON PRESERVED TRACK MORPHOLOGY 117

(punctured to the base of the sand) at the toe-off track, also formed in similarly fine-grained
phase of the step cycle. The rest of the track is a sands. The fossil track (Fig. 21b) was most
transmitted feature. The preservation of the probably formed when the sediment had a high
'hoof suggests perforation to a horizon with moisture content (if not saturated), and is a
higher moisture content than the overlying dry deep (at least 60 mm) transmitted feature.
sands, accounting for the preservation of this 6. Liquefaction failure close to true surface
deep track feature. track. A large (300mm long) tridactyl track
Similar anterior concentric failure rings have (Fig. 21c) collected from the Scalby Formation
been observed in previous studies, attributing was preserved as a low-relief cast on the under-
the sand crescents to upslope and downslope fea- side of a fallen block. Digits II and III were
tures generated by locomotion over sand dunes clearly defined, with digit IV being relatively
(Reynolds 1989). However, several horizontally faint. The track increases in relief anteriorly.
bedded tracks observed in the current study dis- The track infill was paler than the surrounding
played anterior sand crescents, suggesting that sediments. A smaller tridactyl track was located
their formation was influenced by the dynamics anterior of digit II of the large track. The smaller
of locomotion combined with the foot morphol- track post-dated the large track, as it deforms the
ogy of the track-maker and the mechanical prop- interdigital shear zone.
erties of the substrate, rather than by local The serial-sectioned track (F00813), also from
topography. Similar anterior displacement rims the Scalby Formation (Fig. 6a), displayed a simi-
(sand crescents) were generated in laboratory lar track prior to sectioning. The cross-section
simulations (Fig. 17a). The laboratory-simulated along the median line of digit III (Fig. 6a)
tracks also indicate that the displacement rims shows a region of down-warped sediments in
were not affected by the surface topography of the heel area (A) that represents deformation
the sediments in the porous-plate testing box, caused at the heel-down phase of the step cycle
as they were horizontally bedded. (Fig. 12a). The mid-region (B) of the section of
5. Deep transmitted in saturated conditions. A digit III represents the second forward rotation
tridactyl track (Fig. 21b) from Burniston Bay phase of the step cycle (Fig. 12b). The third
displays features typical of a transmitted track. and most distinct point of the step cycle, the
The track increases in relief anteriorly and the toe-off phase, is clearly represented by a severely
heel is poorly transmitted, represented only by down-warped area (C) of sediment (Fig. 12c, d).
a faint, posteriorly projecting feature. Digit III The track displays all three phases of a step cycle
is the dominant digit, indicating that it was the that would be expected for walking using a hip-
main load-bearing digit of the track-maker. The based retractor mechanism.
mixed nature of the sediment within digits II The basal features for the tracks in Figure 6a
and III suggests that the foot punctured or com- and Figure 21c show some evidence that they
pacted underlying layers to penetrate the exposed were transmitted: the increase in relief anteriorly,
horizon. A faint multiple interdigital shear zone the faint heel feature, and the dominant relief of
(ISZ) feature between digits II and III is partially digits II and III (load-bearing digits). However, it
preserved at the hypex of the digits (Fig. 21b, is the cross-section along the median line of digit
arrowed). III of track F00813 (Fig. 6a) that indicates that
The track simulation T8/F7 produced a deep track F00813 and possibly that in Figure 21c
(80mm) transmitted track feature (Fig. 17c), are deep transmitted features, with MZD (track
very similar to the transmitted track described length) reducing incrementally with depth.
above (Fig. 21b). The simulated track displayed Track F00813 was formed in fine-grained sand,
all the features observed in the fossil track, with very fine interlaminated silt and organic-
including the faint posteriorly projecting feature rich horizons. Based on the data generated
at the heel. The simulated track indicates that from the laboratory track simulations, the
the posteriorly projecting feature may relate to incremental reduction in MZD length in a fine-
a multiple digital transmission feature (MDT), grained sand indicates that the sand was satu-
unique to transmitted tracks. Digit III of the rated (or had a high moisture content) at the
simulated track also displays puncture features, time of track formation. The presence of what
where overlying layers were compacted into appears to be liquefaction failure at the anterior
lower layers. The fossil track indicated that the margin of the track F00813 (Fig. 6a (C)) also
exposed basal layer was at least 60mm below supports the notion that the sediment had a
the true surface on which the organism had high water content at the time of track forma-
walked. The saturated, fine-grained sands used tion. The presence of a very faint ISZ feature
in the laboratory simulation reproduced (at on the larger track (Fig. 21c) suggests that it
depth) a track feature very similar to the fossil was also formed while the sediment had a high
118 P. L. MANNING

moisture content, but closer to the surface than suggesting that the sediment underwent lique-
track F00813. faction failure at the compactive event of track
7. Transmitted with liquefaction failure. Three formation. All digits display medial collapse
tridactyl tracks (Fig. 22) preserved as surface structures, indicating that the track collapsed
tracks were defined only by the highly on withdrawal of the dinosaur's foot. Figure
deformed/contorted sediment within the tracks, 22b also shows a natural, end-on cross-section

Fig. 22. (a) Tridactyl dinosaur track (dorsal surface of track) defined only by the highly deformed/contorted
sediment within the track. Scalby Formation, Long Nab Member, Scalby Bay, Yorkshire (scale bar 10cm). The
outline is marked with chalk for clarity, (b) Tridactyl dinosaur track (dorsal surface of track) defined only by
the highly deformed/contorted sediment within the track. Scalby Formation, Long Nab Member, Scalby Bay,
Yorkshire (scale bar 10cm). (c) Tridactyl dinosaur track (lower surface of track) defined by the deformed
sediment within the track. Anterior transmitted track (A) with transmitted posterior displacement feature (B).
Scalby Formation, Long Nab Member, Scalby Bay, Yorkshire (scale bar 10cm). The outline is marked with
chalk for clarity.
CONTROLS ON PRESERVED TRACK MORPHOLOGY 119

of the heel area of the track, defined by the highly tracks and morphology to determine the taxo-
deformed/contorted down-warped sediment, nomic affinities require supporting evidence
more pronounced near the surface of the track. generated by laboratory track simulations, and
Fragments of the paler overlying fine-grained an appreciation of morphological differences
sandstone have been pushed down into the heel between surface and transmitted track features.
area, floating in the darker, grey silty sand infill- The interpretation of features observed in
ing this feature. The grey, silty sand infilling laboratory-simulated tracks from this study can
delineates a possible feature relating to an incre- be usefully applied to fossil tracks.
mental decrease in the maximum zone of defor-
mation (MZD) with increased depth. Owing to Vertebrate ichnofacies
the fine grade of the sediment, this suggests that Lockley et al. (1994) suggested that vertebrate
the sediment had a very high moisture content, tracks could be compared to invertebrate
and was possibly saturated, at the time of track traces, claiming that their distributions were
formation. both controlled by the same sedimentological
The track in Figure 22c displays two distinct and stratigraphic principles. They reasoned that
areas relating to its formation. Area B could be repeatedly occurring track assemblages (ichno-
interpreted as a fleshy pad at the back of a coenoses) in various terrestrial deposits might
foot, or as a transmitted heel feature. Area A help to define distinctive vertebrate ichnofacies.
represents the distorted area of sediment created The term 'ichnofacies' was considered to equate
on extraction of the track-maker's foot. Area A to recurrent ichnocoenoses associated with parti-
is interpreted as the dynamic result of a foot cular ancient environments, preserved as a dis-
puncturing the saturated sediment, with the tinctive lithofacies (Lockley et al. 1994).
resultant track feature migrating anteriorly with The term 'ichnofacies' was originally intro-
depth. Area B, if interpreted as a transmitted duced by Seilacher (1964,1967) to describe recur-
feature, represents the entrance point of the ring associations of trace fossil assemblages
track-maker's heel at the true track surface, associated with specific lithofacies and deposi-
transmitted to depth. tional environments. The ichnofacies reflected
The track simulation T8/F11 and simulation the environmental conditions (salinity, substrate
T8/F13 (Fig. 6c) produced features comparable character and bathymetry) and the organism's
to the three fossil tracks, in saturated, fine- often intimate relationship with a specific sub-
grained sediments. The simulated track showed strate (see review in Mcllroy 2004).
liquefaction failure of the surface track. The nat- The current study suggests that the composi-
ural section of the heel of the fossil track also tion, properties and condition of a substrate
indicates that the intensely deformed sediments directly control the 'type' of track preservation
were restricted to the top 30mm of the track. and resultant morphology. The use of a single
Track simulation T8/F13 clearly shows the over- or recurring vertebrate ichnocoenosis to diag-
turned and contorted elements of the layers that nose specific facies relationships (ichnofacies) is
underwent liquefaction failure (Fig. 6c). rejected, on the grounds that vertebrate tracks
The three fossil tracks all display similar fail- are not substrate-specific and are a function of
ure; however, those in Figure 22a and 22b are the soil conditions (controlled by the prevailing
surface tracks, whereas that in Figure 22c is a moisture/density relationship at a given time).
transmitted track. The incremental decrease in Laboratory simulation generated a diverse
MZD with depth, combined with the fine grade range of track morphologies that could assist in
of the sediment, suggests that the sediment had the interpretation of substrate conditions prevail-
a very high moisture content, and was possibly ing in a specific environment at the time of track
saturated, at the time of track formation. The formation. However, the transient and often
liquefaction failure of the surface sediments and ephemeral relationship between vertebrates and
digit collapse features also support the inferred specific environments makes it difficult, if not
interpretation of high moisture content at the impossible, to define environments based on
time of track formation. ichnocoenoses, given that many terrestrial verte-
brates are not restricted to a particular palaeo-
Utilization of fossil tracks environment - particularly the Dinosauria.
An understanding of laboratory-simulated
tracks can assist in unravelling how fossil A tangled tale of vertebrate track taxonomy
tracks are formed and the processes that subse- Fossil vertebrate track nomenclature and that of
quently altered them. The use to which many other trace fossils is hotly debated (see Sarjeant
workers have applied fossil tracks must be 1990 for review). The diverse track morphologies
reviewed critically. Studies using size of fossil generated using a single template in the course of
120 P. L. MANNING

this study have shown how little, if any, of the track, its formational history, and preservation
track-maker's morphology is preserved in trans- features (Peabody 1948). The results of this
mitted tracks. When naming tracks, are ichnolo- study, however, indicate that an ichnological
gists identifying a specific animal's track or the approach has merit over an osteologically
trace of an action? based approach, and suggests that many track
A trace fossil is a sedimentary structure that is features have little use as ichnotaxobases.
produced by biological activity, and has a form Sarjeant (1990) concluded similarly that:
that is determined in part by that activity, often
we should remember at all times that we are
reflecting the morphology of the track-maker to
naming, not the animal itself, but the trace of its
only a limited extent (Sarjeant 1990). However,
act as reflected in the sediment and preserved in
one of the aims of a vertebrate ichnologist is
the fossil record.
still to identify the track-maker, a goal that is
seldom possible. The morphological variability The need for a standard approach to the
on which ichnotaxa are often erected relates to description of tracks is vital if the trace of an
the often complex formation and preservation action or preservational features of an animal's
of a track, not the foot morphology of the pro- activity are to be the criterion by which it is
genitor. named. Leonardi (1987) and Thulborn (1990)
Invertebrate and vertebrate palaeoichnologists provide the most consistent approach to track
have for many years followed the guidelines set description and terminology, but terms were
down in the International Code of Zoological not available to describe some of the features
Nomenclature (ICZN) when naming new ichno- observed in the current study. The terminology
taxa. Modifications to the ICZN guidelines adopted in this study has utilized some existing
(Basan 1979) have provided a defined status for terms and has adapted and developed terms
trace fossils within zoological taxonomy (see specifically relating to the surface and subsurface
Mcllroy 2004). However, some authors have morphology of a track (see Figs 7, 8). Terms such
suggested that a completely separate code of as the maximum zone of deformation (MZD),
nomenclature is still required for trace fossils, interdigital shear zone (ISZ), and multiple digital
in which ichnotaxa are differentiated from transmission (MDT) describe morphological
other natural animal taxa, and that the aim of features that are the result of sediment behaviour
the ichnologist should be restricted to identifying caused by a limb's interaction with a substrate
either the track-maker or its activity (Sarjeant during track formation. The MZD, ISZ and
1990; Mcllroy 2004). MDT features should not be used as characters
Some ichnologists (Farlow 1992; Farlow & to define an ichnotaxon, only to help interpret
Lockley 1993) disagree with Sarjeant (1990), them, as they relate to sediment behaviour in
and have placed ichnotaxa within osteologically relation to the dynamic formation of a track,
based orders and classes. This approach has and not to animal behaviour. The variations in
potential problems, such as when the tracks of MZD, ISZ and MDT are useful tools for
a tyrannosaur were described (Tyrannosauropus assisting in the interpretation of a track
Haubold 1971) that have since been attributed (vertebrate or invertebrate), as are the digital
to a hadrosaur (Lockley 1991). Under the rules lengths and interdigital angles (IDA). The use
laid down in the ICZN code, the first name of digit length and IDA as descriptive termin-
given to a taxon takes priority over subsequent ology has proved useful in the current study,
reinterpretation, however unsuitable or mislead- but the instability of these values within a
ing the name. Sarjeant (1990) argues clearly that single track also makes them unsuitable ichno-
such an approach has an element of absurdity, taxobases.
because it gives equal systematic status to the In summary it is considered that vertebrate
animal itself and to the effect of its actions. ichnotaxa should reflect the morphological dif-
The current study has produced many over- ferences resulting from track formation and
print, true-surface and subsurface track features, preservation and not the affinity of an alleged
all of which assist with the understanding of the maker.
interaction of a foot with the substrates tested
in this study. The information gained on the pre-
servational circumstances and effects, from the Conclusions
laboratory track simulations, far outweighed
the information gained on the track-maker's A controlling factor in the formation of track
foot (template) morphology. An ichnotaxon, features is the moisture/density relationship
whether an ichnospecies, ichnogenus or ichno- prevailing in the sediment at the time of
family, should reflect the morphology of the indentation.
CONTROLS ON PRESERVED TRACK MORPHOLOGY 121

Subsurface track (simulated) morphology Desiccation cracks post-dating a track do not


demonstrated the dynamic nature of track always indicate aerial exposure of the track
formation by the anterior displacement of surface. Cohesive soils overlain by moist to
track features with depth, producing 'stacked' saturated non-cohesive soils can be indented
tracks. The direction of the displacement of by a foot generating a transmitted track at
the subsurface track features corresponded depth. On drying, the transmitted track in
with the direction and magnitude of the force the cohesive soil cracks, and is subsequently
applied and the physical properties of the infilled by the overlying non-cohesive soils,
sediment through which the force travelled. giving the false impression of a surface track
Multiple digital transmission (MDT) features feature (albeit with transmitted track mor-
were unique to subsurface deformation, indi- phology).
cating a transmitted track layer. The MDT The use of dinosaur tracks in comparative
features were developed over a greater depth multivariate studies should be restricted to
of track within dry sediments, with a vertical surface track features, for comparison with
reduction of MDT features with increasing other surface track features. The inclusion of
moisture content. transmitted tracks in such studies invalidates
Alteration of grain size and moisture content any inferred taxonomic or osteological rela-
in the laboratory-simulated tracks affected tionships, owing to the disparity between
the type and extent of the development of the surface and subsurface track morphology
maximum zone of deformation track length and size.
(MZD track length). Dry medium-grained Any multivariate study based on morpho-
sands displayed a trend towards large incre- logical variability in tracks and trackways
mental reduction in MZD track length with will be viable only if the three-dimensional
increasing depth. Dry fine-gained sands dis- variability of track morphology is understood.
played trends towards little or no reduction Future multivariate studies must approach the
in MZD track length with depth. task of understanding the three-dimensional
Saturated (moisture content >29%) medium- components of a track before valid compari-
grained sands displayed trends towards little or son can be made with other tracks within a
no reduction in MZD track length with depth. three-dimensional framework.
Saturated (moisture content >29%) fine- Vertebrate ichnotaxa should reflect the mor-
grained sands displayed trends towards a phological differences resulting from behaviour,
large incremental reduction in MZD track not the affinity of an alleged track-maker or
length with depth. artefacts of formation and preservation.
Optical pedobaragraph (OPB) studies con- The methods developed for the recovery and
firmed that high pressure values corresponded interpretation of laboratory subsurface track
to high-relief features in tracks, and conversely layers may go some way to assisting with the
low pressures corresponded to shallow or interpretation of fossil tracks. However, many
absent track features. more track simulations are required in varying
The distribution of pressure across a foot sediment types, and with realistic indenters, to
relates to the kinematics of the step cycle, allow a more complete understanding of
and implies the limb retraction mechanism processes that form and subsequently alter
adopted by that animal. The correlated track surface and subsurface track morphology and
relief of simulated and fossil tracks can possi- composition.
bly be used to define the limb retraction
mechanism adopted by dinosaurs from trans- This research would have not been possible were it
mitted track features that display the distribu- not for a studentship awarded by the University of
tion of pressure over the sole of the foot. Sheffield and supervised by M. Whyte and M.
Romano. K. Padian, P. Ensom, M. Benton, J. Pollard
Track length within different 'layers' of a single are thanked for critical comments and J., A. & K. Man-
transmitted track were dependent on sediment ning for their support. Thanks also to D. Mcllroy for
consistency and properties. Only the 'true' sur- editorial help, for his patience, and for redrafting the
face track 'layer' should be used to calculate the figures.
hip height, based only on the foot length, as
transmitted track lengths can vary greatly
from the 'true' surface track length. References
The presence of cohesive sediment (clay) below
non-cohesive sand increases the preservation Alexander, R. M. 1985. Mechanics of posture and gait
potential of tracks (in the non-cohesive sedi- in some large dinosaurs. Zoological Journal of the
ment) at the junction between the two. aasd
122 P. L. MANNING

ALLEN, J. R. L. 1989. Short paper: Fossil vertebrate in birds and humans. Journal of the Zoological
tracks and indenter mechanics. Journal of the Society, London, 224, 127-147.
Geological Society, London, 146, 600-602. GATESY, S. M., MIDDLETON, K. M., JENKINS, F. A. &
ALLEN, J. R. L. 1997. Subfossil mammalian tracks SHUBIN, S. H. 1999. Three-dimensional preservation
(Flandrian) in the Severn Estuary, SW Britain: of foot movements in Triassic theropod dinosaurs.
mechanics of formation, preservation and distri- Nature, 399, 141-144.
bution. Philosophical Transactions of the Royal GILLETTE, D. D. & LOCKLEY, M. G. (eds) 1989. Dino-
Society, London, Series B, 352, 381-518. saur Tracks and Traces. Cambridge University
ATKINSON, J. H. & BRANSBY, P.L. 1978. The Mechanics Press, Cambridge.
of Soils: An Introduction Critical State Soil HAUBOLD, H. H. 1971. Ichnia amphibiorum et repti-
Mechanics. McGraw-Hill, London. liorum fossilium. In: KUHN, O. (ed.) Handbuch
BASAN, P. B. 1979. Trace fossil nomenclature: the der Palaoherpetologie, Part 18. Gustav Fisher,
developing picture. Palaeogeography, Palaeo- Stuttgart, 1-7.
climatology, Palaeoecology, 28, 143-167. HAUBOLD, H. H. 1984. Saurierfahrten. Wittenberg
BETTS, R. P., FRANKS, C. I. & DUCKWORTH, T. 1991. Lutherstadt, Germany: Ziemsen 231pp.
Foot pressure studies: normal and pathologic HITCHCOCK, E. 1858. Ichnology of New England. A
gait analyses. Part II: Biomechanics of the foot Report on the Sandstone of the Connecticut
and ankle. In: JAHSS, M. H. (ed.) Disorders of the Valley, Especially its Fossil Footmarks. W. White,
Foot and Ankle-Medical and Surgical Manage- Boston, MA [reprinted 1974 by Arno Press, New
ment, 2nd edition, W B Saunders, Philadelphia, York].
Volume 1, Chapter 18, 484-519. JOHNSON, R. E. & OSTROM, J. H. 1995. The forelimb of
BIRD, R. T. 1944. Did brontosaurs ever walk on land? Torosaurus and an analysis of the posture and gait
Natural History, 53, 60-67. of ceratopsian dinosaurs. In: THOMASON, J. J. (ed.)
BOUSSINESQ, J. 1883. Application des potentials a I 'etude Functional Vertebrate Morphology in Vertebrate
de I'equilibre et du mouvement des solides elastiques. Paleontology. Cambridge University Press, Cam-
Gauthier-Villars, Paris. bridge, 205-218.
BRAJA, M. D. 1994. Principles of Geotechnical Engineer- KARAFIATH, L. L. & NOWATZKI, E. A. 1978. Soil
ing. PWS, Boston, MA. asdfasdfsadfasdfad
BRITISH STANDARDS INSTITUTION, 1986. BS8004, Code Transglobal Technical Publications, Aedermanns-
of Practice for Foundations, BSI, London. dorf.
BUCKLAND, W. 1828. Note sur les traces de tortues LAMBE, T. W. 1961. Soil Mechanics. Wiley, New York.
observees dans les gres rouge. Annals de Sciences LEONARDI, G. 1987. Glossary and Manual of Tetrapod
Natural, Paris, 13, 85-86. Footprint Ichnology. Ministerio das minas e
CLARK, J. & ALEXANDER, R. M. 1975. Mechanics of energia, Departamento National da Produ9ao
running by quail (Coturnix). Journal of Zoology Mineral, Brasil.
London, 176, 87-113. LINGEN, G. J. & ANDREWS, P. B. 1969. Hoof-print
COOMBS, W. P. 1980. Swimming ability of carnivorous structures in beach sand. Journal of Sedimentary
dinosaurs. Science, 207, 1198-1200. Petrology, 39, 350-357.
CRAIG, R. F. 1992. Soil Mechanics. Chapman & Hall, LOCKLEY, M. G. 1991. Tracking Dinosaurs. Cambridge
London. University Press, Cambridge.
DEMATHIEU, G. R. 1987. Thickness of the footprint LOCKLEY, M. G., MATSUKAWA, M. & OBATA, I. 1989.
reliefs and its significance: research on the distribu- Dinosaur tracks and radial cracks: unusual foot-
tion of weights upon the autopodia. In: LEONARDI, print features. Bulletin of the National Science
G. (ed.) Glossary and Manual of Tetrapod Foot- Museum, Tokyo, 15, 151-160.
print Palaeoichnology, Department Nacional du LOCKLEY, M. G., HUNT, A. P. & MEYER, C. A. 1994.
Producao Mineral, Brasil, 60-61. Vertebrate tracks and the ichnofacies concept:
DODD, J. R. & STANTON, R. J. 1990. Paleoecology: implications for palaeoecology and palichno-
Concepts and Applications. Wiley, New York. stratigraphy. In: Donovan, S. (ed.) Palaeobiology
FARLOW, J. O. 1989. Ostrich footprints and trackways: of Trace Fossils. Belhaven Press, New York,
implications for dinosaur ichnology. In: GILLETTE, 241-268.
D. D. & LOCKLEY, M. G. (ed.) Dinosaur Tracks MclLROY, D. 2004. In: MC!LROY, D. (ed.) 2004. The
and Traces. Cambridge University Press, Cam- Application of Ichnology to Palaeoenvironmental
bridge, 243-248. and Stratigraphic Analysis. Geological Society,
FARLOW, J. O. 1992. Sauropod tracks and track- London, Special Publications, 228, 3-27.
makers: integrating the ichnological and skeletal McKEE, E. D. 1947. Experiments on the development
records. Zubia, 10, 89-138. of tracks in fine cross-bedded sand. Journal of
FARLOW, J. O. & LOCKLEY, M. G. 1993. An osteometric Sedimentary Petrology, 17, 23-28.
approach to the identification of the makers of McMAHON, T. A. 1984. Muscles, Reflexes, and Loco-
Early Mesozoic tridactyl dinosaur footprints. In: motion. Princeton University Press, Princeton.
LUCAS, S. G. & MORALES, M. (eds) The Nonmarine NORMAN, D. B. 1986. On the anatomy of Iguanodon
Triassic. New Mexico Museum of Natural History asdasdfsadfasdfsdafasdfasdfasdfasdfasdfasdfasdfdasfadf
and Science Bulletins, 3, 123-131. Bulletin de I'lnstitut Royale des Sciences
GATESY, S. M. & BIEWENER, A. A. 1991. Bipedal Naturelles de Belgique: Sciences de la Terre, 56,
locomotion: effects of speed, size and limb posture 281-372.
CONTROLS ON PRESERVED TRACK MORPHOLOGY 123

PADIAN, K. & OLSEN, P. E. 1984. Footprints of the National Monument, California: their context,
Komodo Monitor and the trackways of fossil classification and preservation. Palaeogeography,
reptiles. Copeia, 3, 662-671. Palaeoclimatology, Palaeoecology, 57, 285-331.
PEABODY, F. E. 1948. Reptile and amphibian trackways SEILACHER, A. 1964. Sedimentological classification
from the Lower Triassic Moenkopi Formation of and nomenclature of trace fossils. Sedimentology,
Arizona and Utah. University of California Publi- 3, 240-252.
cations, Bulletin of the Department of Geological SEILACHER, A. 1967. Bathymetry of trace fossils.
Sciences, 27, 295-468. Marine Geologist, 5, 413-428.
PRANDTL, L. 1920. Uber die Harte plastischer Korper. SWARTZ, S. M. & BIEWENER, A. A. 1992. Shape and
Nachricten von der Koniglichen Gesellschaft der scaling. In: BIEWENER, A. A. (ed.) Biomechanics
Wissenschaften zu Gottingen (Mathematisch - Structures and Systems: A Practical Approach.
physikalische Klasse aus dem jahre 1920), Berlin. Oriel Press, Oxford, 21-43.
PRANDTL, L. 1921. Uber die Eindringungsfestigkeit SMITH, M. J. 1981. Soil Mechanics. Longman Scientific
(Harte) plastischer Baustoffe und die Festigkeit & Technical, London.
von schneiden. Zeitscrift fur angerwandte Mathe- THULBORN, R. A. 1990. Dinosaur Tracks. Chapman &
matik und Mechanik, 1, 15-20. Hall, London.
REYNOLDS, R. E. 1989. Dinosaur trackways in the THULBORN, R. A. & WADE, M. 1989. A Footprint as a
Lower Jurassic Aztec Sandstone of California. In history of movement. In: GILLETTE, D. D. &
GILLETTE, D. D. & LOCKLEY, M. G. (eds) Dinosaur LOCKLEY, M. G. (eds) Dinosaur Tracks and
Tracks and Traces. Cambridge University Press, Traces. Cambridge University Press, Cambridge,
Cambridge, 285-292. 51-56.
ROMANO, M. & WHYTE, M. A. 1996. Fossils explained TUCKER, M. E. & BURCHETTE, T. P. 1977. Triassic dino-
16: Trace fossils 3 - dinosaur tracks. Geology saur footprints from South Wales: Their context
Today, 12, 75-79. and preservation. Palaeogeography, Palaeoclima-
ROMANO, M & WHYTE, M. A. 2003. Jurassic dinosaur tology, Palaeoecology, 22, 195-208.
tracks and trackways of the Cleveland Basin, WEISHAMPEL, D. D., DODSON, P. & OSMOLSKA, H. (eds)
Yorkshire: preservation, diversity and distribu- 1990. The Dinosauria. University of California
tion. Proceedings of the Yorkshire Geological Press, California.
Society, 54, 185-215. WHYTE, M. A. & ROMANO, M. 1981. A footprint in the
ROMER, A. S. 1923. The pelvic musculature of saur- sand of time. Journal of the University of Sheffield
ischian dinosaurs. Bulletin of the American Geological Society, 7.6, 323-330.
Museum of Natural History, 48, 605-617. WHYTE, M. A. & ROMANO, M. 1993. Footprints of a
SARJEANT, W. A. S. 1974. A history and bibliography of sauropod dinosaur from the middle Jurassic of
the study of fossil vertebrate footprints in the Yorkshire. Proceedings of the Geologists' Associa-
British Isles. Palaeogeography, Palaeoclimatology, tion, 104, 195-199.
Palaeoecology, 16, 265-378. WHYTE, M. A. & ROMANO, M. 1994a. Probable sauro-
SARJEANT, W. A. S. 1990. A name for the trace of an act: pod footprints from the Middle Jurassic of
approaches to the nomenclature and classification Yorkshire, England. Gaia, 10, 15-26.
of fossil vertebrate footprints. In: CARPENTER, K. WHYTE, M. A. & ROMANO, M. 1994b. Lower Middle
& CURRIE, J. C. (edsj Dinosaur Systematics: Per- Jurassic sequences between Whitby and Saltwick.
spectives and approaches. Cambridge University In: SCRUTTON, C. (ed.) Yorkshire Rocks and
Press, Cambridge, 299-307. Landscape: A Field Guide. Ellenbank Press,
SCRIVNER, P. J. & BOTTJER, D. J. 1986. Neogcne avian Mayport, 158-164.
and mammalian tracks from Death Valley
This page intentionally left blank
Phanerozoic history of deep-sea trace fossils
ALFRED UCHMAN

Institute of Geological Sciences, Jagiellonian University, Oleandry 2a,


30-063 Krakow, Poland (e-mail: fred@ing.uj.edu.pl)

Abstract: The Phanerozoic diversity of deep-sea trace fossils, based on 151 flysch formations,
displays distinct, non-linear changes through the Phanerozoic, with peaks in the Ordovician-
Early Silurian and Early Carboniferous, lowered in the Permian-older Late Jurassic, a peak
in the Tithonian-Aptian, lowered in the Albian, and the maximum in the Eocene. The
contribution of graphoglyptids in trace fossil assemblages rises gradually up to the end of
the Cretaceous, shows a peak in the Palaeocene-Eocene but a depression in the Oligocene.
All the changes were probably influenced by competition for food and food supply,
bottom water temperatures and oxygenation, and frequency of flysch habitats. There is no
clear influence of major biotic crises (Ordovician/Silurian, Cretaceous/Tertiary, Palaeo-
cene/Eocene) on the diversity of deep trace fossils, except the Eocene/Oligocene crisis.

It is obvious that environmental conditions can Trace fossil diversity: previous studies and
affect evolutionary processes. Vice versa, in methods
some cases organisms can influence environ-
ments. These environment-organism relation- There is no consensus on changes of diversity of
ships are reconstructed from the fossil record, deep-sea trace fossils through geological time.
which in the case of deep-sea invertebrate benthic Crimes (1974) documented a sharp increase in
animals is full of gaps. These gaps can be the number of ichnogenera in Cretaceous-
partially filled by studying trace fossils. In this Tertiary flysch deposits, but suggested that this
paper, the Phanerozoic diversity of deep-sea could be an artefact partly caused by 'more
trace fossils, stratigraphic ranges of selected detailed studies' on formations of this age.
ichnotaxa, general composition of trace fossil According to Seilacher (1974, 1976), the diversity
assemblages (contributions of some behavioural of deep-sea trace fossils increased through the
groups and some ichnogenera) and contribution Phanerozoic with a rapid acceleration in the
of graphoglyptids in trace fossil assemblages are Cretaceous. Subsequently, Seilacher (1977b)
considered. evoked a gradual increase of diversity of 'flysch
Attempts at this sort of study have been ichnocoenoses' through the Phanerozoic, but
undertaken since the 1970s, but reconsidera- later (Seilacher 1978) proposed a 'mid-Cretac-
tion is necessary because of the huge amount eous diversity burst' (see also Frey & Seilacher
of new ichnological data becoming available 1980). These views were tested by McCann
during the last decade. Seilacher (1974, (1990), who concluded that the increase of
1977b, 1978) based his work on 16 selected diversity was neither gradual in most of the
formations. McCann (1990) used 34 forma- Phanerozoic nor rapid in the Cretaceous.
tions, but lumped some units together inappro- According to Orr (2001), the diversity of deep-
priately - for instance the Jurassic-Tertiary sea trace fossil assemblages displayed differences
flysch of the Polish Carpathians, which in particular periods of the Palaeozoic, with a
includes several formations deposited in differ- distinct increase from the Cambrian to Ordo-
ent basins (Ksia_zkiewicz 1977). Orr (2001) vician, a low peak in the Ordovician, and further
considered 50 formations within the Ordo- increase in the Carboniferous. The diversity of
vician-Carboniferous interval. In this paper, graphoglyptid trace fossils, which are the most
151 flysch formations are considered for the characteristic component of the deep-sea
entire Phanerozoic. Flysch is understood as a Nereites ichnofacies (Seilacher 1967; Frey &
facies, of which nine jfeatures were defined by Seilacher 1980), was analysed by Uchman
Dzulynski & Walton (1965). (2003). It was low from the Cambrian to the
The illustrated specimens are housed in the Middle Jurassic, with a low peak in the Ordo-
Institute of Geological Sciences, Jagiellonian vician followed by a drop in the Silurian. In the
University, Krakow, Poland (acronym 154P), Late Jurassic, after the late Palaeozoic low, the
and in the Institute of Palaeontology, Wurzburg graphoglyptid diversity started to increase
University, Wurzburg, Germany (acronym PIW gradually and markedly in the Late Cretaceous,
1998IV). probably during the Turonian. It dropped
From: MC!LROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 125-139. 0305-8719/04/S15.00 © The Geological Society
of London.
Fig. 1. The diversity diagram and diversity curve of deep-sea trace fossils through the Phanerozoic. The dashed line shows the questionable depressions in the curve.
The numbers from 1 to 151 refer to formations according to a list in the author's database (housed in the Geological Society of London). The timescale which is based
on Gradstein & Ogg (1999) can be obtained from the Society Library or the British Library Document Supply Centre, Boston Spa, Wetherby, West Yorkshire
LS23 7BQ, UK as Supplementary Publication No. SUP 18209 (60 pages). It is also available online at http://www.geolsoc.org.uk/SUP18209.
Fig. 2. Diagram of the proportion of graphoglyptids in trace fossil assemblages that contain 10 or more ichnotaxa. The curve shows average values. The numbers refer
to formations according to a list in the author's database. The timescale is based on Gradstein & Ogg (1999).
128 A. UCHMAN

again in the Palaeocene, and subsequently (1974) recognized, has been partially filled by
increased to a maximum in the Eocene. The subsequent research (e.g. Yang 1986, 1988;
Oligocene was marked by a sharp decrease in Kozur et al 1996; Tchoumatchenco & Uchman
graphoglyptid diversity and frequency, followed 1999), but the ichnology of deep-sea formations
by an increase in the Miocene (Uchman 2003). of this age is still poorly known, as is also true
There is also no uniform method for estimat- for the Upper Cambrian and the Middle-Upper
ing trends in diversity. Seilacher (1974, 1977b, Devonian. The largest number of considered
1978) selected 16 well-understood deep-sea formations (46%) derive from the Upper Cretac-
formations and considered their number of eous-Eocene, representing a mere 12% of
ichnospecies. McCann (1990) analysed the num- Phanerozoic time. Only 8% of the formations
bers of both ichnospecies and of ichnogenera. cover the Permian-Jurassic, representing 27%
Orr (2001) considered only ichnogenera in Cam- of the Phanerozoic. Post-Miocene deep-sea for-
brian-Carboniferous formations. Apart from the mations are only marginally considered because
number of ichnogenera in a particular formation, they are underrepresented in the fossil record:
he used the 'average diversity calculated by most of them are still covered by the sea, and
period' as a parameter. This parameter is rather trace fossil (mostly graphoglyptid) diversity
questionable, because many formations cross tends to be grossly underestimated in cores.
period boundaries, and it is difficult to tell Obviously, the higher the number of formations
which ichnogenera should be considered in in a given period, the more objective the assess-
which period. The diversity can change very ment of diversity, as can be judged from statistics
much within periods, and the average values rules.
can mask the changes. Moreover, there is The proportion of graphoglyptid ichnogenera
always under-representation of trace fossil asso- to total number of ichnogenera was considered
ciations of low diversity, which do not attract for formations that contain 10 or more ichno-
the attention of ichnologists, and it is unclear genera. This limitation was applied because in
how such data might skew the results. smaller trace fossil assemblages each grapho-
In this paper, the total number of ichnogenera glyptid taxon changes this parameter excessively.
in each of the 151 formations (database housed The percentage values were plotted against the
in the Geological Society of London) is plotted Phanerozoic timescale of Gradstein & Ogg
against geological time (Fig. 1), as in Orr (1999) (Fig. 2). Potential construction of a
(2001). Consideration at the ichnogeneric level curve of the contribution of graphoglyptids on
is more convenient and objective than at the the basis of the highest values in a given
ichnospecies level, because of many unresolved period was rejected as too subjective, because
ichnotaxonomic problems at ichnospecific level. some authors seem to omit 'less attractive', com-
The data are taken from the literature, own monly simple, non-graphoglyptid ichnotaxa.
observations, and unpublished data from flysch Instead, a curve of the average values of the
deposits of Turkey, Greece, the Crimea, the percentage of graphoglyptids was constructed
Balkans and the Polish Carpathians. Deep-sea (Fig. 2).
deposits are not limited to flysch facies, but the
other facies display much less diversity, mostly
because of their lower preservational potential Diversity through time
(e.g. Wetzel 1984). The data were revised with
respect to ichnotaxonomy in a consistent way, During the Phanerozoic the diversity fluctuated,
through application of concepts previously and displays characteristic anomalies (Fig. 1).
developed by the author (Uchman 1995, 1998, There was a distinct increase of diversity from
1999, 2001). The considered formations differ the Cambrian to Ordovician. It continued to
not only in thickness, stratigraphic range, rise during the Early Silurian, when it reached
facies, and exposure, but also in the amount of an early Palaeozoic peak in the Aberystwyth
attention by researchers. This problem is, how- Grits Formation of Wales, UK (no 17 in Figs
ever, difficult to avoid. Abundance can be 1, 2) (Crimes & Crossley 1991). Diversity
measured by the number of formations in dropped slightly in the Early Devonian. There
which a given trace fossil occurs (Uchman 2003). is only one Middle-Upper Devonian formation
A curve of changes in diversity was con- among the available data, which shows much
structed on the basis of the most diverse trace less diverse trace fossil assemblages than the
fossil associations (one association for one older formations. The data by Orr (2001) for
formation) in any given period. The selected this period do not point to a diversity higher
formations are not uniformly distributed in than eight ichnogenera, but these data are not
time. The Permian-Jurassic gap, which Seilacher available for the author of this paper. It is
DEEP SEA ENVIRONMENTS AND EVOLUTION 129

unclear whether this is a virtual or apparent drop the Coniacian up to the Palaeozoic and the
of diversity resulting from environmental Tithonian-Aptian levels and rose further to a
changes or from under-representation of data Phanerozoic maximum during the Eocene
from flysch formations of this age, but the [Beloveza Formation, Polish Carpathians (no
latter possibility seems very likely. The next 122 in Figs 1, 2; Ksia_zkiewicz 1977; Uchman
peak of diversity occurred in the Early Carboni- 1992) and the Zumaya flysch (no 111 in Figs 1,
ferous Culm deposits of the Czech Republic (no 2; Leszczynski & Seilacher 1991 and references
31 in Figs 1, 2) (Lang et al 1979; Pek & Zapletal therein)]. Diversity decreased after the Eocene.
1990 and references therein). Diversity dropped
again in the Late Carboniferous and reached a
depression in the Permian and most of the Stratigraphic range of selected ichnotaxa and
Triassic. It rose during the Jurassic to the levels composition of trace fossil assemblages
of the Palaeozoic peaks in the Tithonian-
Aptian. There is a questionable depression in Stratigraphic ranges of ichnotaxa can be used as a
the younger Middle Jurassic and older Late Stratigraphic tool. The Cruziana ichnostrati-
Jurassic that probably results from too low a graphy of Gondwana (Seilacher 1970, 1994) is a
number of formations of this age. The Albian good example that is very useful in Lower Palaeo-
shows a distinct depression in the diversity zoic shallow-marine siliciclastics. In deep-sea
curve. Diversity rose from the Cenomanian to trace fossil assemblages there is no group of

Table 1. Occurrences and Stratigraphic ranges of Dictyodora ispp

Ichnotaxa Occurrence References

'Dictyodora' simplex Seilacher Lower Cambrian, Salt Range, Pakistan Seilacher 1955
Ordovician, Stara Planina, Bulgaria Acenolaza & Yanev 2001
Dictyodora tennis (M'Coy) Middle Ordovician-Lower Silurian, New Pickerill 1980; Pickerill et al
Brunswick, Canada 1987;
Lower Llandovery, Gala and Penkill Benton & Trewin 1980
groups, Hawick Rocks, UK
Ordovician, Barrancos, Portugal Neto de Carvalho 2001
Dictyodora cf. tennis (M'Coy) Lower-Middle Ordovician, Upper Hovin Hanken et al. unpublished
Group, Norway,
Dictyodora scotica (M'Coy) Lower Llandovery, Gala and Penkill Benton & Trewin 1980
groups, Hawick Rocks, UK
Middle Ordovician-Lower Silurian, Pickerill 1980; Pickerill et al.
Metapedia Group, New Brunswick, 1987
Canada
Llandovery, Devil's Bridge Formation, Orr 1995
Neuadd Fawr, Welsh Basin, UK
Lower Silurian, Waterville Formation, Orr & Pickerill 1995
Maine, USA
Dictyodora zimmermanni Hundt ?Llandeilo-Caradoc, Hauptquarzit, Benton 1982
Thuringia, Germany
Lower Ordovician, Skiddaw Group, UK Orr 1996
Lower Ordovician, Manx Group, UK Orr & Howe 1999
Dictyodora ?zimmermanni Hundt Lower-Middle Ordovician, Upper Hovin Hanken et al. unpublished
Group, Norway
Dictyodora silurica Yang Silurian, Qinling, China Yang & Hu 1992
Dictyodora liebeana (Geinitz) Lower Carboniferous, Culm, Thuringia, Benton 1982 and references
Germany therein
Lower Carboniferous, Culm, Moravia and Lang et al. 1979; Pek &
Silesia, Czech Republic Zapletal 1990 and
references therein
Lower Carboniferous, Culm, Stepanek & Geyer 1989
Frankenwalds, Germany
Lower Carboniferous, East Menorca, Orr l994;Onetal. 1996
Spain
Lower Carboniferous, Culm, Minorca, Llompart & Wieczorek
Spain 1997
130 A. UCHMAN

trace fossils having similar value, but certainly


some attempts can be made. Most ichnotaxa
show wide stratigraphic ranges, but some of
them show characteristic evolutionary trends
(e.g. Seilacher 1977a, Uchman 2003). In some
instances, abundance can also be used for this
purpose.
Thus, for example, in deep-sea facies the ich-
nogenus Dictyodora ranges from the Ordovician
to the Carboniferous, though its ichnospecies
display narrower ranges and could, with caution,
be used in stratigraphy (Table 1, Fig. 3). Nereites
irregularis (Schafhautl) (Fig. 4a) (formerly
Helminthoida labyrinthica Heer) ranges from
the Turonian, questionably from the Aptian.
Phycosiphon incertum Fischer-Ooster (Fig. 4c)
occurs in deep-sea facies from the Late Devo-
nian. Scolicia (Fig. 4b) and Ophiomorpha (Fig.
5b) occur from the Tithonian, but became more
common only since the Turonian (Tchoumatch-
enco & Uchman 2001). Cladichnus fischeri
(Heer) (Fig. 6) ranges from the Coniacian to
the Eocene, and Rotundusichnium zumayense
(Gomez de Llarena) (Fig. 5a) from the Maas-
trichtian to the Eocene. Most graphoglyptids
have their first occurrences in the Upper
Cretaceous or Palaeogene (Uchman 2003). For
instance, Glockerichnus alata (Seilacher) (Fig.
7a) occurs only in the Eocene. These facts can
be used as an accessory tool for stratigraphy.
However, some of graphoglyptids have long
breaks in their stratigraphic record, e.g. Hel-
minthorhaphe flexuosa Uchman (Fig. 4d) from
the Cambrian to the Late Cretaceous. It is very
probable that this trace fossil was made by
animals having different stratigraphic ranges
(Elvis ichnotaxon, cf. Erwin & Droser 1993).
The composition of flysch trace fossil assem-
blages changes with time. Lower Cambrian
flysch assemblages are poorly diversified, and
dominated by simple forms (Planolites, Palaeo-
phycus) with significant contribution of arthro-
pod traces and the so-called 'shallow-marine'
ichnotaxa (Skolithos, Monocraterion, Phycodes).
They display a very low proportion of pascichnia
and agrichnia (Orr 2001). However, the Middle
Cambrian (age according to Pickerill, personal
communication) Meguma Group (Nova Scotia,
Canada) deep-sea deposits already contain 17
ichnogenera (Pickerill & Williams 1989). A
significant break took place after the Middle
Cambrian, when the agrichnial makers of
Fig. 3. Stratigraphic ranges and abundance of
graphoglyptids migrated from shallow-marine selected ichnotaxa. For Dictyodora see Table 1.
environments to the deep sea (Crimes et al. Range of Cladichnus fischeri (Heer) based on Uchman
1992; Crimes & Fedonkin 1994), probably at or (1998, 1999). Ranges of the remaining ichnotaxa are
near the Cambrian-Ordovician boundary. The based on the database housed in the Geological
colonization of the deep sea by graphoglyptid Society of London. The question mark indicates that
producers initiated the development of the the determination is questionable.
DEEP SEA ENVIRONMENTS AND EVOLUTION 131

Fig. 4. Selected deep-sea trace fossils, (a) Nereites irregularis (Schafhautl) and Chondrites intricatus
(Brongniart) (in the left corner), Ofterschwanger Beds (upper Albian-upper Cenomanian), Obsermaiselstein,
Rhenodanubian Flysch, Germany, PIW 1998IV3. (b) Scolicia isp., Zarzecze Beds (lower Eocene), Konina,
Magura Nappe, Polish Carpathians, 144P10. (c) Phycosiphon incertum Fischer-Ooster, Bleichernhorn Series
(Maastrichtian), road-cut between Schnifis and Thuringerberg, Rhenodanubian Flysch, Austria, PIW
1998IV11A. (d) Helminthorhaphe flexuosa Uchman, Beloveza Beds (Eocene), Slopnice, Magura Nappe,
Polish Carpathians, 144P11. Scale bars - 1 cm.

Fig. 5. Other selected deep-sea trace fossils, (a) Rotundusichnium zumayensis (Gomez de Llarena), Sievering
Beds (Maastrichtian-Palaeocene), Miihlberg, Vienna Woods, Rhenodanubian Flysch, Austria, 145P9, Scale bar
jkasdhkjlasdhkjalksdjfhjkasdfhalskdjfhlaskjdfhljkdasfhadsjklfhadjksfhladjksfhsdjklfhsdjkfhsdjkfhhf
photograph, scale - 10cm.
132 A. UCHMAN

Fig. 6. Cladichnus fischeri (Heer), Zementmergel Series (Maastrichtian), Kalkgraben quarry, Rhenodanubian
Flysch, Germany, PIW 1998IV120, scale in mm.

archetypal Nereites ichnofacies of Seilacher


(1967) (Orr 2001).
Ordovician-Carboniferous trace fossil assem-
blages are relatively similar in that they contain
characteristic trace fossils such as Dictyodora,
Nereites (but not Nereites irregularis (Schaf-
hautl)), Neonereites (included in Nereites by
many authors; Rindsberg 1994; Uchman 1995),
Protovirgularia, Megagrapton and Squamodict-
yon. Oldhamia ispp. are present in some muddy
facies from the Neoproterozoic to the Early
Carboniferous (Seilacher 1997). In younger
Palaeozoic strata, Phycosiphon, Lophoctenium
and Spirodesmos can be added to the list.
Permian-Middle Jurassic trace fossil assem-
blages are less distinctive. They commonly
contain Chondrites, Lophoctenium, Phycosiphon,
Nereites, Megagrapton and Agrichnium, Desmo-
grapton pamiricus (Yialov) (Fig. 7b) is a rare
but characteristic graphoglyptid in Triassic-
Middle Jurassic flysch. In Lower Cretaceous
trace fossil assemblages it is difficult to select
characteristic ichnotaxa; however, Helminthopsis
is quite common, and Belorhaphe zickzack (Heer)
and Glockerichnus glockeri (Ksi^zkiewicz) are
Fig. 7. Other stratigraphically useful deep-sea trace relatively common at that time.
fossils, (a) Glockerichnus alata (Seilacher), Eocene Upper Cretaceous-Neogene trace fossil
flysch of the Carman Prealps, Italy, based on a field assemblages commonly contain a different suite
photograph, (b) Desmograpton pamiricus (Vialov), of graphoglyptids co-occurring with Scolicia,
Tavrida Flysch (Upper Triassic-Lower Jurassic), Nereites irregularis (Schafhautl), Phycosiphon
Crimea, Ukraine, based on a field photograph. and Zoophycos. In sandstone-rich facies, Ophio-
DEEP SEA ENVIRONMENTS AND EVOLUTION 133

morpha rudis (Ksi^zkiewicz) and O. annulata previous views (Crimes 1974; Crimes & Droser
(Ksi^zkiewicz) are common. 1992), the low diversity of the Cambrian deep-
sea trace fossil assemblages was a reflection of
poor oxygenation of bottom waters and insuffi-
Contribution of graphoglyptids cient supply of organic detritus. Orr (2001)
challenged this view, suggesting that increased
Graphoglyptids form probably the most impor- competition for ecospace and/or resources in
tant component of deep-sea trace fossil assem- shallow-water environments pressed trace-
blages from the Ordovician. Their producers makers to migrate to the deep sea.
are well adapted to relatively stable oligotrophic Diversity from the Ordovician to the Early
conditions (Seilacher 1977b; Miller 1991; Carboniferous remains at a relative high, with
Uchman 2003), and they are typical indicators two peaks in the Ordovician-Early Silurian and
of a K-selected strategy, indicating stable envir- Early Carboniferous (Fig. 1). However, the
onments (Ekdale 1985). The contribution of possible Middle-Late Devonian decrease of
graphoglyptids in their palaeoecological context diversity remains uncertain because sufficient
has been considered in Alpine flysch (e.g. Tunis data are lacking. The distinct decrease of diver-
& Uchman 1996). Orr (2001) considered agri- sity after the Early Carboniferous and especially
chnia (mostly graphoglyptids) and pascichnia during the Permian (by about 50%) can be
together for the Cambrian-Carboniferous deep- related to the formation of Pangea and lowering
sea formations, and concluded that their con- of temperature of deep-sea waters as a result of
tribution to trace fossil assemblages in the the Gondwanan glaciations. Formation of a
Cambrian was distinctly lower than in younger single supercontinent, Pangea, and one supero-
periods. cean, Panthalassa, shortened active margins
The curve constructed herein of the proportion and the abundance of related flysch deposits. It
of graphoglyptid ichnogenera to the total is probable that the decreased habitat area low-
number of ichnogenera (Fig. 2) shows an ered the diversity of its inhabitants. This crisis
increase of the value since the Early Cambrian also influenced the composition of trace fossil
to Ordovician from about 10% to about 20%. assemblages. Dictyodora, Spirodesmos and
The value then grows slowly and gradually Oldhamia disappeared from the fossil record.
through the rest of the Palaeozoic and the Meso- Seilacher (1974) supported a suggestion pro-
zoic up to about 25%, reaching a maximum in posed by Wolf (1960), who regarded the
the Palaeocene and Eocene of about 40% and evolution of the abyssal benthos to be strongly
then dropping in the Oligocene below 35%, influenced by temperature changes of bottom
only to rise again in the Miocene. waters. Glacial intervals in the Quaternary
show reduced benthic species diversity, including
the deep sea (Cronin & Raymo, 1997). Seilacher
Discussion (1974), however, having a much smaller dataset
than presented herein, concluded that the
The curves of diversity and contribution of Upper Palaeozoic glaciations had no influence
graphoglyptids, stratigraphic ranges of some on the diversity of trace fossils. Most probably
ichnotaxa and main changes in composition of reduction of bottom temperatures was an impor-
trace fossil assemblages imply evolutionary tant stress factor, which favoured opportunistic
changes of trace fossils. The changes were related trace-makers, and a corresponding lowering of
to environmental and palaeoecological events ichnofaunal diversity. Moreover, drying of the
and changes. Their potential controls are dis- climate in the Permian reduced vegetation and
cussed below. probably decreased phytodetrital input - a possi-
ble but indirect food source - to the deep sea via
turbiditic currents.
Palaeozoic There is no clear evidence of influence in the
ichnological dataset presented herein of the
There is a distinct rise of diversity and contribu- Ordovician/Silurian boundary crisis, which was
tion of graphoglyptids from Late Cambrian to caused by the Late Ordovician glaciation
Early Ordovician flysch (Crimes 1974). At that (Barnes et al 1996), on the diversity of deep-sea
time, graphoglyptids, which originated in shal- trace fossils in general (Fig. 1). However,
low-marine environments in the Early Cambrian, McCann (1990) noted that a drop of diversity
migrated to the deep sea (Crimes & Fedonkin occurred in the Welsh Basin at that time.
1994), probably at or near the Cambrian- Crimes et al. (1992, fig. 6) postulated a gradual
Ordovician boundary (Orr 2001). According to increase of all deep-sea trace fossils from the
134 A. UCHMAN

Late Precambrian (Vendian) to the Early Silur- benthic activity (e.g. Jacobs & Lindberg 1998
ian, but according to Orr (2001) the average and references therein).
diversity dropped slightly after the Ordovician.
Also, the number of graphoglyptid ichnotaxa
and the number of their first occurrences Late Cretaceous-Neogene
dropped after the Ordovician (Uchman 2003).
The Late Ordovician glaciation was much Following the Albian, the diversity of deep-sea
shorter and less extensive than that of the Late trace fossils increased quickly up to the Phanero-
Carboniferous and Permian. This could partially zoic optimum in the Eocene. Most probably this
explain the differences in their influence on the rapid growth was caused by further increase in
deep-sea temperatures and resulting trace fossil the competition for food and further improve-
diversity. ment of the deep-sea habitats, similarly to the
diversity of graphoglyptids (Uchman 2003).
Flysch deposition related to the active margins
Triassic-Early Cretaceous of the Tethys and circum-Pacific systems
became more common than in the Mesozoic.
There is no evidence of influence of the Permian/ Enlargement of habitats may have once again
Triassic boundary crisis on deep-sea trace fossils. enhanced the possibility for new adaptations.
It is very probable that diversity increased gradu- Frey & Seilacher (1980) suggested that the
ally during the Triassic to the older Late Jurassic divergence of continents resulted in the enlarge-
apart from the steps on the curve in the Middle/ ment of ocean basins, which in turn may have
Late Triassic (Fig. 1), which may be an artefact. resulted in increased speciation. Echinoid bur-
For this period a correspondingly small number rows (Scolicia) and large crustacean burrows
of flysch deposits are known, which moreover (Ophiomorpha) show a distinct increase in
occur mostly in central Asia and in some areas frequency at that time, up to an optimum in the
of the circum-Pacific tectonic belt. Many of Eocene. Also, small but efficient deposit-feeding
these await more detailed investigation, which, structures (Nereites irregularis} show similar
potentially, could modify the curve. Still, this trends (Fig. 3). Most graphoglyptids, whose
was a period with rather limited areas of flysch trace-makers are well adapted to competition
habitats. for food, exhibited their first occurrences or
The distinct increase in diversity within the became more frequent after the Late Cretaceous,
Tithonian-Aptian is recorded in the Carpathian and their frequency reached a maximum in the
Flysch of the Silesian Unit (Ksia_zkiewicz 1977). Eocene (Uchman 2003). In sum, new types of
The Tithonian deep-seafloor was colonized by trace fossil assemblages were established with
irregular echinoids producing Scolicia and common contributions by Scolicia, Ophiomor-
larger crustaceans producing Ophiomorpha pha, Nereites irregularis and diverse grapho-
(Tchoumatchenco & Uchman 2001). These glyptids. Older trace fossil assemblages,
efficient bioturbators strongly influenced the sea- especially those in the Triassic-Jurassic, are
floor, intensively ploughing it to ingest particles more similar to their Palaeozoic counterparts,
for food and perhaps improving its oxygenation but do not contain Dictyodora, as was
by irrigation of oxygenated water. As a result, previously noted by Seilacher (1978).
they increased the competition for food, which The Late Cretaceous changes may also be
favoured diversity of behavioural adaptations related to increased input of phytodetritus to
and consequently increased the diversity of the deep sea owing to the coeval evolution of
trace-makers (e.g. Wetzel 1991). Better oxygena- terrestrial angiosperms. Radiation of the grapho-
tion improved the habitats and enabled the glyptid trace-makers may be a response to the
expansion of other organisms into deeper tiers. increased flux of non-refractory organic matter,
These processes caused a minor revolution in mainly cellulose, into the oligotrophic deep-sea
the substrates like that of the major late Precam- environment, where it could be used by
brian revolution, i.e. the transition from the mat burrowers for microbial farming (Seilacher
grounds to mix grounds (Seilacher & Pfltiger 1974, 1977a, 1977b). More probably, however,
1994; Seilacher 1999). These processes were they were influenced by the coeval increase of
stopped to some extent by the Lower Cretaceous organic matter from plankton (Miller 1991;
anoxic events (e.g. Jenkyns 1980). In the Albian, Uchman 2003), because they were produced in
the diversity curve shows a depression. Many pelagic-hemipelagic mud deposited between
flysch facies of this age, at least within the turbiditic flows. Phytodetritus is typically utilized
Tethyan and Atlantic realm, were influenced by by the trace-makers of Scolicia and Ophio-
reduced oxygenation, which possibly reduced morpha, which burrow in order to exploit organic
DEEP SEA ENVIRONMENTS AND EVOLUTION 135

Fig. 8. The diversity curve of deep-sea trace fossils through the Phanerozoic (based on Fig. 1) in relation the
sea-level curve (after Hallam 1989), the mean global temperature curve (after Frakes 1979) and the global
diversity curve for marine genera (after Sepkoski 1995). G indicates major glaciations. Arrows indicate major
biotic crises. The timescale is based on Gradstein & Ogg (1999).
136 A. UCHMAN

matter deeply buried by turbiditic sediments and for food. Diversity of planktonic and benthic
maturated by microbes. foraminifera increased at the beginning of the
The Eocene optimum in diversity (Fig. 1) and Miocene (Thunell 1981; MacLeod et al. 2000),
frequency of deep-sea trace fossils is very dis- as did the diversity of the calcareous nanno-
tinct. Graphoglyptids at that time display the plankton (Bown et al. 1992).
largest contribution in trace fossil assemblages In general, the diversity curve of deep-sea ich-
(Fig. 2). In the Julian Prealps (Tunis & nogenera is similar to the curve of diversity of
Uchman 1996), Carpathians (Ksiajzkiewicz marine animal genera (Fig. 8) (Sepkoski 1995).
1977; Uchman 1998) and the Rhenodanubian However, most of the major biotic crises such
Flysch (Uchman 1999) the increase of grapho- as the Ordovician/Silurian, Frasnian/Famenian,
glyptid diversity had already begun in the late Triassic/Jurassic, and Cretaceous/Tertiary, did
Palaeocene and started to decrease in the latest not influence the diversity of deep trace fossils
middle Eocene. The high diversity is probably except for the lower rank Eocene/Oligocene
related to the advent of oligotrophic conditions crisis. There is no correlation with the mean
in the late Palaeocene-early Eocene (Tunis & global temperature curve (Frakes 1979) except
Uchman 1996), which is a widespread phenom- for the discussed decrease of temperature in the
enon (Hallock et al. 1991) related to global Late Carboniferous-Early Permian and after
warming (Savin et al. 1975; Boersma & Premoli the Eocene. There is also no direct correlation
Silva 1991). Global warming produced dense with sea-level (Hallam 1989).
saline waters that flowed to the deep sea and
caused a significant increase in deepwater tem-
peratures (e.g. Brass et al. 1982; Shackleton Conclusions
1986). These phenomena are rooted in global
tectonic and palaeoceanographic changes (Rea The diversity of deep-sea trace fossils displays
et al. 1990; Oberhansli 1996; Pickering 2000). distinct changes through the Phanerozoic,
Calcareous nannoplankton diversity displays a with peaks in the Ordovician-Early Silurian
second diversity peak in the middle Eocene and Early Carboniferous, a depression in
(Bown et al. 1992). The generic richness of the Permian-older Late Jurassic, a peak in
benthic foraminifera also shows a maximum in the Tithonian-Aptian, a depression in the
the middle Eocene (Lutetian) (MacLeod et al. Albian, and the maximum peak in the Eocene.
2000). The contribution of graphoglyptids rose
The Oligocene decrease in diversity and con- gradually up to the end of the Mesozoic,
tribution of graphoglyptids (Figs 1, 2) is very dis- with a peak in the Palaeocene-Eocene and a
tinct. After the Eocene no new graphoglyptid depression in the Oligocene.
ichnotaxa appear (Uchman 2003). This phenom- The diversity of deep-sea trace fossils and their
enon can be related to the Eocene/Oligocene evolutionary trends were influenced mostly by
boundary crisis, which also influenced plankton competition for food, bottom water tempera-
(Corliss 1979), including foraminifera, dino- tures, sediment oxygenation, and also, indir-
flagellates and nannoplankton, whose diversity ectly, by changes of the frequency of flysch
dropped drastically after the Eocene (Cavelier deposits (i.e. suitable deep-sea habitats).
et al. 1981). World temperatures dropped as There is no clear influence of the major biotic
well (Buchard 1978). The longitudinal Atlantic crises, such as the Ordovician/Silurian, Fras-
circulation and connection to the Arctic Ocean nian/Famenian, Triassic/Jurassic and Cretac-
was established (Prothero 1994). Cold polar eous/Tertiary, on the diversity of deep trace
water flowed in several deep-sea basins, such as fossils except for the lower rank Eocene/
in the Carpathians and in the Alps, with locally Oligocene crisis. However, the Ordovician/
anoxic conditions (Leszczynski 1997 and refer- Silurian, Cretaceous/Tertiary and Palaeo-
ences therein). cene/Eocene crises influenced graphoglyptid
Further improvement of the climatic condi- ichnodiversity and their relative abundance
tions in the Miocene (Savin et al. 1975) did not (Uchman 2003).
result in an increase of diversity of deep-sea Changes in the diversity of deep-sea trace
trace fossils (Fig. 1). Only the frequency of fossils show unpredictable dynamics, not
graphoglyptids (Uchman 2003) and their contri- simple linear changes through time. This
bution to trace fossil assemblages increased observation confirms that the time-stability
(Fig. 2). Benthic niches have remained filled hypothesis of Sanders (1968) is not valid.
with graphoglyptids since the Oligocene. Their
increased contribution in trace fossil assemblages This publication was supported by the Jagiellonian
indicates increased oligotrophy and competition University (funds DS/V/ING). Sponsors of the Lyell
DEEP SEA ENVIRONMENTS AND EVOLUTION 137

Meeting in 2003 supported the participation of AU in CRIMES, T. P, GARCIA HIDALGO, J. F. & POIRE, D. G.
the Lyell Meeting, London, 2003, where this paper 1992. Trace fossils from Arenig flysch sediments
was presented. A. K. Rindsberg (Alabama) improved of Eire and their bearing on the early colonisation
the English and provided helpful comments. Some of of deep seas. Ichnos, 2, 61-77.
the illustrated specimens were collected during tenure CRONIN, T. M. & RAYMO, M. E. 1997. Orbital forcing
of a grant from the Alexander von Humboldt Founda- of deep-sea benthic species diversity. Nature, 385,
tion. R. K. Pickerill (Fredericton), A. Wetzel (Basel) 624-627.
and D. Mcllroy (Wirral) critically reviewed the manu- DZULYNSKI, S. & WALTON, E. K. 1965. Sedimentary
script and proposed helpful suggestions and further lin- Features of Flysch and Greywackes. Elsevier,
guistic improvements. Amsterdam.
EKDALE, A. A. 1985. Paleoecology of the marine endo-
benthos. Palaeogeography, Palaeoclimatology,
References Palaeoecology, 50, 63-81.
ERWIN, D. H. & DROSER, M. L. 1993. Elvis taxa.
ACENOLAZA, F. G. & YANEV, S. 2001. El Ordovicico Palaios, 8, 623-624.
del sector occidental de Stara Planina (Montes FRAKES, L. A. 1979. Climates throughout Geological
Balcanes), Bulgaria: Icnofosiles e implicaciones Time. Elsevier, Amsterdam.
paleobiogeograficas. Museo Argentino de Ciencias FREY, R. W. & SEILACHER, A. 1980. Uniformity in
Naturales 'Bernardino Rivadavia', n.s., 3, 55-72. marine invertebrate ichnology. Lethaia, 13, 183-
BARNES, C. R., FORTEY, R. A. & WILLIAMS, S. H. 1996. 207.
The pattern of global bio-events during the GRADSTEIN, F. & OGG, J. 1999. Geological Time Scale:
Ordovician period. In: WALLISER, O. H. (ed.) Phanerozoic. Saga Petroleum, Malburk.
Global Events and Event Stratigraphy in the HALLAM, A. 1989. The case for sea-level change as a
Phanerozoic. Springer, Berlin, 139-172. dominant casual factor in mass extinction of
BENTON, M. J. 1982. Dictyodora and associated trace marine invertebrates. Philosophical Transaction
fossils from the Palaeozoic of Thuringia. Lethaia, of the Royal Society of London, B325, 437-455.
15, 115-132. HALLOCK, P., PREMOLI SILVA, I. & BOERESMA, A.
BENTON, M. J. & TREWIN, N. J. 1980. Dictyodora from 1991. Similarities between planktonic and larger
Silurian of Peeblesshire, Scotland. Palaeontology, foraminiferal evolutionary trends through Paleo-
23, 501-513. gene palaeoceanographic changes. Palaeogeogra-
BOERSMA, A. & PREMOLI SILVA, I. 1991. Distribution phy, Palaeoclimatology, Palaeoecology, 83, 49—64.
of Palaeogene planktonic foraminifera-analogies JACOBS, D. K. & LINDBERG, D. R. 1998. Oxygen
with the recent? Palaeogeography, Palaeoclimatol- and evolutionary patterns in the sea: Onshore/
ogy, Palaeoecology, 83, 29—48. offshore trends and recent recruitment of deep-
BOWN, P. R., BURNETT, J. A. & GALLAGHER, L. T. 1992. sea faunas. Proceedings of the National Academy
Calcareous nannoplankton evolution. Memoire di of Sciences of the United States of America, 95,
Scienze Geologiche, 43, 1—17. 9396-9401.
BRASS, G. W., SOUTHAM, J. R. & PETERSON, W. H. 1982. JENKYNS, H. C. 1980. Cretaceous anoxic events: from
Warm saline bottom water in the ancient ocean. continents to oceans. Journal of the Geological
Nature, 296, 620-623. Society of London, 137, 171-188.
BUCHARD, B. 1978. Oxygen isotope paleotemperatures KOZUR, H. W., KRAINER, K. & MOSTLER, H. 1996.
from the Tertiary period of the North Sea. Nature, Ichnology and sedimentology of the Early
275, 121-123. Permian deep-water deposits from the Lercara-
CAVELIER, C., CHATEAUNEUF, J.-J., POMEROL, C., Roccapalumba area (Western Sicily, Italy).
RABUSSEER, D., RENARD, M. & VERGNAUD- Fades, 34, 123-150.
GRAZZINI, C. 1981. The geological events at the KSIAZKIEWICZ, M. 1977. Trace fossils in the Flysch of
Eocene/Oligocene boundary. Palaeogeography, the Polish Carpathians. Palaeontologia Polonica,
Palaeoclimatology, Palaeoecology, 36, 223-248. 36, 1-208.
CORLISS, B. H. 1979. Response of deep-sea benthonic LANG, V., PEK, I. & ZAPLETAL, J. 1979. Ichnofosilie
Foraminifera to development of the psychro- kulmu jihovychodni casti Drahanske Vrchoviny
sphere near the Eocene/Oligocene boundary. (Trace fossils in Culm of the South-eastern part
Nature, 282, 63-65. of Drahanska Vrhovina - Highlands). Acta
CRIMES, T. P. 1974. Colonisation of the early ocean Universitatis Palackianae Olomucensis, Facultas
floor. Nature, 248, 328-330. Rerum Naturalium, 62, 57-96.
CRIMES, T. P. & CROSSLEY, J. D. 1991. A diverse ichno- LESZCZYNSKI, S. 1997. Origin of the Sub-Menilitie
fauna from Silurian flysch of the Aberystwyth Globigerina marl (Eocene-Oligocene transition)
Grits Formation, Wales. Geological Journal, 26, in the Polish Outer Carpathians. Annales Societa-
27-64. tis Geologorum Poloniae, 67, 367—472.
CRIMES, T. P. & DROSER, M. L. 1992. Trace fossils and LESZCZYNSKI, S. & SEILACHER, A. 1991. Ichnocoenoses
bioturbation: the other fossil record. Annual of a turbidite sole. Ichnos, 1, 293-303.
Review of Ecology & Systematics, 23, 339-360. LLOMPART, C. & WIECZOREK, J. 1997. Trace fossils
CRIMES, P. T. & FEDONKIN, M. A. 1994. Evolution from Culm facies of Minorca Island. Prace
and dispersal of deepsea traces. Palaios, 9, 74- Paristwowego Instytutu Geologicznego, 157, 99-
83. 101.
138 A. UCHMAN

MACLEOD, N., ORTIZ, N., FEFFERMAN, N., CLYDE, W., PICKERILL, R. K. & WILLIAMS, P. F. 1989. Deep bur-
SCHULTER, C. & MACLEAN, J. 2000. Phenotypic rowing in the early Palaeozoic deep-sea: examples
response of foraminifera to episodes of global from the Cambrian(?)-early Ordovician Meguma
environmental change. In: CULVER, S. J. & Group of Nova Scotia. Canadian Journal of
RAWSON, P. F. (eds) Biotic Response to Global Earth Sciences, 26, 1061-1068.
Change: The Last 145 Million Years. Cambridge PICKERILL, R. K., FYFFE, L. R. & FORBES, W. H. 1987.
University Press, Cambridge, 51-78. Late Ordovician- Early Silurian trace fossils from
McCANN, T. 1990. Distribution of Ordovician-Silurian the Metapedia Group, Tobique River, Western
ichnofossil assemblages in Wales: implications for New Brunswick, Canada. Maritime Sediments
Phanerozoic ichnofaunas. Lethaia, 23, 243-255. and Atlantic Geology, 23, 77-88.
MILLER, W. III. 1991. Paleoecology of graphoglyptids. PICKERING, K. T. 2000. The Cenozoic world. In:
Ichnos, 1, 305-312. CULVER, S. J. & RAWSON, P. F. (eds) Biotic
NETO DE CARVALHO, C. 2001. Implicagoes geometricas Response to Global Change: The Last 145 Million
na etologia do productor de Dictyodora tennis Years. Cambridge University Press, Cambridge,
(M'Coy) no Arenigiano de Barrancos (Portugal): 20-34.
custos e beneficios de ser um pioneiro. In: MELEN- PROTHERO, D. R. 1994. The Eocene-Oligocene Transi-
DEZ, G., HERRERA, Z., DELVENA, G. & AZANZA, B. tion: Paradise Lost. Columbia University Press,
(eds) Los Fosiles y la Paleogeografia. 17 Jornadas New York.
de la Sociedad Espanola de Paleontologia. REA, D. K., ZACHOS, J. C., OWEN, R. M. & GINGERICH,
Publicaciones del Seminario de Paleontologia de P. D. 1990. Global change at the Paleocene-
Zaragoza, 5, 369-378. Eocene boundary: climatic and evolutionary
OBERHANSLI, H. 1996. Klimatische und ozeanogra- consequences of tectonic events. Palaeogeography,
phishe Veranderungen im Eozan. Zeitschrift der Palaeoclimatology, Palaeoecology, 79, 117-128.
Deutschen Geologischen Gesellschaft, 147, 303-^13. RINDSBERG, A. K. 1994. Ichnology of the Upper Missis-
ORR, P. J. 1994. Trace fossil tiering within event beds sippian Hartselle Sandstone of Alabama, with
and preservation of frozen profiles: an example notes on other Carboniferous formations. Geologi-
from the Lower Carboniferous of Menorca. cal Survey of Alabama, Bulletin, 158, 1-107.
Palaios, 9, 202-210. SANDERS, H. L. 1968. Marine benthic diversity: a com-
ORR, P. J. 1995. A deep-marine ichnofaunal assem- parative study. American Naturalist, 100, 243-282.
blage from Llandovery strata of the Welsh Basin, SAVIN, S. M., DOUGLAS, R. G. & STEHLI, F. G. 1975.
west Wales, UK. Geological Magazine, 132, 267- Tertiary marine paleotemperatures. Geological
285. Society of America, Bulletin, 86, 1499-1510.
ORR, P. J. 1996. The ichnofauna of the Skiddaw Group SEILACHER, A. 1955. Spuren und Fazies im Unterkam-
(Early Ordovician) of the Lake District, England. brium. In: SCHINDEWOLF, O. H. & SEILACHER, A.
Geological Magazine, 133, 193-216. (eds) Beitrage zur Kenntnis des Kambriums in der
ORR, P. J. 2001. Colonization of the deep-marine envir- Salt Range (Pakistan). Akademie der Wissenschaf-
onment during the early Phanerozoic: the ichno- ten und der Literatur in Mainz, Abhandlungen der
faunal record. Geological Journal, 36, 265-278. mathematisch-naturwissenschaftlichen Klasse, 10,
ORR, P. J. & HOWE, M. P. A. 1999. Macrofauna and 373-399.
ichnofauna of the Manx Group (Early Ordovi- SEILACHER, A. 1967. Bathymetry of trace fossils.
cian), Isle of Man. In: WOODCOCK, N. H., Marine Geology, 5, 413^28.
QUIRK, D. G., FITCHES, W. R. & BARNES, R. P. SEILACHER, A. 1970. Cruziana stratigraphy of 'non-
(eds) In Sight of the Suture: the Palaeozoic Geology fossiliferous' Palaeozoic sandstones. In: CRIMES,
of the Isle of Man in its lapetus Ocean Context. T. P. & HARPER, J. C. (eds) Trace Fossils, Geologi-
Geological Society, London, Special Publications, cal Journal, Special Issue, 3, 447^76.
160, 33^4. SEILACHER, A. 1974. Flysch trace fossils: evolution of
ORR, P. J. & PICKERILL, R. K. 1995. Trace fossils from behavioural diversity in the deep-sea. Neues Jahr-
Early Silurian flysch of the Waterville Formation, buchfur Geologie und Paldontologie, Monatshefte,
Maine, USA. Northeastern Geology, 17, 394-414. 1974, 223-245.
ORR, P. J., BENTON, M. & TREWIN, N. H. 1996. Deep SEILACHER, A. 1976. Evolution von Spuren-Verge-
marine trace fossil assemblages from the Lower schellschaftungen. Zentrablatt fur Geologie und
Carboniferous of Menorca, Balearic Islands, Paldontologie, Teil 2, 1976, 396^02.
western Mediterranean. Geological Journal, 31, SEILACHER, A. 1977a. Pattern analysis of Paleodictyon
235-258. and related trace fossils. In: CRIMES, T. P. &
PEK, I. & ZAPLETAL, J. 1990. The importance of ichnol- HARPER, J. C. (eds) Trace Fossils 2, Geological
ogy in geologic studies of the eastern Bohemian Journal, Special Issue, 9, 289-334.
Massif (Lower Carboniferous), Czechoslovakia. SEILACHER, A. 1977b. Evolution of trace fossil commu-
Ichnos, 1, 147-149. nities in the deep sea. In: HALLAM, A. (ed.) Patterns
PICKERILL, R. K. 1980. Phanerozoic flysch trace fossil of Evolution. Elsevier, Amsterdam, 359-376.
diversity: observations based on an Ordovician SEILACHER, A. 1978. Evolution of trace fossil commu-
flysch ichnofauna from the Aroostook-Metapedia nities in the deep sea. Neues Jahrbuchfiir Geologie
Carbonate Belt of northern New Brunswick. und Paldontologie, Abhandlungen, 157, 251-255.
Canadian Journal of Earth Sciences, 17, 1259- SEILACHER, A. 1994. How valid is Cruziana stratigra-
1270. phy? Geologische Rundschau, 83, 752-758.
DEEP SEA ENVIRONMENTS AND EVOLUTION 139

SEILACHER, A. 1997. Fossil Art. Royal Tyrell Museum UCHMAN, A. 1995. Taxonomy and palaeoecology of
of Palaeontology, Drumheller. flysch trace fossils: the Marnoso-arenacea Forma-
SEILACHER, A. 1999. Biomat-related lifestyles in the tion and associated facies (Miocene, Northern
Precambrian. Palaios, 14, 86-93. Apennines, Italy). Beringeria, 15, 3-115.
SEILACHER, A. & PFLUGER, F. 1994. From biomats to UCHMAN, A. 1998. Taxonomy and ethology of flysch
benthic agriculture: a biohistoric revolution. In: trace fossils: a revision of the Marian Ksi^zkiewicz
KRUMBEIN, W. E., PATERSON, D. M. & STAL, L. J. collection and studies of complementary material.
(eds) Biostabilization of Sediments. Bibliotheks- Annals Societatis Geologorum Poloniae, 68, 105-
und Informationssystem der Universitat Olden- 218.
burg, Oldenburg, 97-105. UCHMAN, A. 1999. Ichnology of the Rhenodanubian
SEPKOSKI, J. J. JR. 1995. Patterns of Phanerozoic Flysch (Lower Cretaceous-Eocene) in Austria
extinctions: a perspective from global data bases. and Germany. Beringeria, 25, 65-171.
In: WALLISER, O.H. (ed.) Global Events and Event UCHMAN, A. 2001. Eocene flysch trace fossils from the
Stratigraphy. Springer, Berlin, 35-51. Hecho Group of the Pyrenees, northern Spain.
SHACKLETON, N. J. 1986. Paleogene stable isotope Beringeria, 28, 3-41.
events. Palaeogeography, Palaeoclimatology, UCHMAN, A. 2003. Trends in diversity, frequency and
Palaeoecology, 57, 91-102. complexity of graphoglyptid trace fossils: evolu-
STEPANEK, J. & GEYER, G. 1989. Spurenfossilien aus tionary and palaeoenvironmental aspects. Palaeo-
dem Kulm (Unterkarbon) des Frankenwaldes. geography, Palaeoclimatology, Palaeoecology, 192,
Beringeria, 1, 1-55. 123-142.
TCHOUMATCHENCO, P. & UcHMAN, A. 1999. Lower and WETZEL, A. 1984. Bioturbation in deep-sea fine-grained
Middle Jurassic flysch trace fossils from the sediments: influence of sediment texture, turbidite
eastern Stara Planina Mountains, Bulgaria: a frequency and rates of environmental changes. In:
contribution to the evolution of Mesozoic ichno- STOW, D. A. V. & PIPER, D. J. W. (eds) Fine
diversity. Neues Jahrbuchfur Geologie und Paldon- Grained Sediments: Deep-Water Processes and
tologie, Abhandlungen, 213, 169-199. Facies. Geological Society, Special Publications,
TCHOUMATCHENCO, P. & UCHMAN, A. 2001. The oldest 15, 597-608.
deep-sea Ophiomorpha and Scolicia and associated WETZEL, A. 1991. Ecologic interpretation of deep-sea
trace fossils from the Upper Jurassic-Lower trace fossil communities. Palaeogeography,
Cretaceous deep-water turbidite deposits of SW Palaeoclimatology, Palaeoecology, 85, 47-69.
Bulgaria. Palaeogeography, Palaeoclimatology, WOLF, T. 1960. The hadal community: an introduction.
Palaeoecology, 169, 85-99. Deep-Sea Research, 6, 95-124.
THUNELL, R. C. 1981. Cenozoic palaeotemperature YANG, S. 1986. Turbidite flysch trace fossils from
changes and planktonic foraminiferal speciation. China and their paleoecology and paleo-
Nature, 289, 670-672. environments. Chinese Paleontological Society.
TUNIS, G. & UCHMAN, A. 1996. Trace fossil and facies 13th—14th Annual Meeting, Selected papers.
changes in the Upper Cretaceous-Middle Eocene Anjing Science and Technology Publishing
flysch deposits of the Julian Prealps (Italy and Company, 143-161.
Slovenia): consequences of regional and world- YANG, S. 1988. Permian-Triassic flysch trace fossils
wide changes. Ichnos, 4, 169-190. from the Guoluo and Yushu regions, Qinghai.
UCHMAN, A. 1992. Skamienialosci sladowe w eocen- Acta Sedimentologica Sinica, 6, 1-12.
skim cienko- i sredniolawicowym fliszu strefy YANG, S. & Hu, Y. 1992. Silurian trace fossils from the
bystrzyckiej plaszczowiny magurskiej (Trace western part of West Qinling. Geoscience, Journal
fossils of the Eocene thin- and medium-bedded of the Graduate School, China University of
flysch of the Bystrica Zone of the Magura Nappe Geoscience, 6, 385-391.
in Poland). Przeglqd Geologiczny, 40, 430^35.
This page intentionally left blank
A re-evaluation of the relationship between trace fossils and dysoxia

KATE D. MARTIN

Millbank Lodge, Kirkton of Mary culler, Aberdeen AB12 5FS

Abstract: Geochemical and palaeontological methods are used to determine the oxygenation
histories of Jurassic sequences at Ravenscar, North Yorkshire, and Lyme Regis, Dorset. The
ichnology of these sequences is compared with interpreted oxygen levels, allowing current
models of oxygen-related trace fossil occurrence to be tested. These case studies support
pre-existing models of trace fossil occurrence in demonstrating that burrow diversity,
diameters and depth of infaunal tiering increase with increasing oxygen levels. The case
studies suggest that trace fossil ethologies may not always be a reliable indicator of
palaeo-oxygenation: in some cases, substrate consistency may have a greater influence over
ethology than oxygen levels. Chondrites is confirmed as a common constituent of dysoxic
settings; however, other trace types may also be indicative of such settings.

Palaeontological studies in dysoxic environments have not been linked to independent evidence
have been motivated by the importance of such for oxygenation. It is thus possible that ichnoco-
environments in the formation of petroleum enoses interpreted as dysaerobic may have been
source rocks (e.g. Savrda et al. 1984), their possi- controlled by factors other than oxygenation.
ble use as global transgression indicators (e.g. This work therefore adopts a slightly different
DeMaison & Moore 1980), as analogues for the approach from many previous studies of dys-
Proterozoic evolution of life (e.g. Rhoads & aerobic trace fossils. Two Jurassic case studies
Morse 1971) and, more recently, as indicators are presented. In each of these, oxygenation
of likely anthropogenic effects on the marine histories have been determined using a combina-
biota (e.g. Oschmann 1991; Tyson & Pearson tion of geochemical and palaeontological
1991). methods. The ichnological succession was then
Studies of trace fossils in dysoxic environments compared with the palaeo-oxygenation histories.
can be considered as particularly important This method allows some of the assumptions
owing to the reduced diversity of calcareous behind the current oxygen-related trace fossil
forms in such settings (e.g. Rhoads & Morse models to be tested.
1971), resulting in a relative paucity of body
fossils. A number of models have been developed
that attempt to use trace fossils to assess oxygen Current models of oxygen-related trace fossil
gradients within dysoxic strata, incorporating
information on the specific ichnogenera present occurrence
in addition to more general observations on
The case studies presented here will be used to
depth and extent of bioturbation, burrow size
demonstrate that some of the assumptions
and burrow morphology (e.g. Bromley &
Ekdale 1984; Savrda & Bottjer 1986; Ekdale & made in current oxygen-related trace fossil
Mason 1988). Although these models are models may not always apply. A brief review of
the current models is therefore appropriate.
supported by numerous examples of dysaerobic
ichnocoenoses, other studies have suggested
that they do not always accurately predict the
characteristics of such ichnocoenoses (e.g. Savrda and Bottjer model: tiering and
Hudson & Martill 1991; Wignall 1991). This oxygen-related ichnocoenoses
suggests that there is a need to evaluate
dysaerobic ichnocoenoses during different Bromley & Ekdale (1984,1986) demonstrated how
periods of the Phanerozoic. the cross-cutting relationships of trace fossils
The oxygen-related trace fossil models cur- could be used to determine tiering profiles in the
rently in use have been based primarily on rock record, the deepest traces cross-cutting all
modern oxygen-restricted environments, deter- shallower ones, and the shallowest traces being
mining the characteristics of the incipient traces cross-cut by all other trace types. The determina-
and using these to enable recognition of ancient tion of tiering profiles in various Mesozoic trace
examples of dysaerobic trace fossils. The majority fossil associations allowed Bromley & Ekdale
of studies of ancient dysaerobic ichnocoenoses (1984) to infer that the vertical partitioning of
From'. MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 141-156. 0305-8719/04/S15.00 © The Geological Society
of London.
142 K. D. MARTIN

changing degrees of bottom water oxygenation:


burrow diversity, diameters, and penetration
depth. With declining oxygen levels, biodiversity
is generally considered to decrease (e.g. Diaz &
Rosenberg 1995). Maximum burrow diameters
are also seen to decrease under lower oxygen con-
ditions (e.g. Savrda et al. 1984). The loss of larger
Fig. 1. Cartoon of the anatomy of an idealized infaunal forms under such conditions tends to
indistinctly burrowed bed. result in reduced irrigation and aeration of the
sediment. A consequent rise in the position of
the redox potential discontinuity (RPD) causes
the infauna was controlled primarily by porewater the remaining burrows to be emplaced at higher
oxygenation. This recognition led to the develop- levels within the sediment than they would be
ment of a trace fossil model for identifying changes under better-oxygenated conditions, thus redu-
in oxygen levels within dysoxic strata (Savrda & cing the vertical extent of the burrows (e.g.
Bottjer 1986, 1987, 1989a, 1989b), derived from Bromley & Ekdale 1984). These three criteria
late Mesozoic and younger examples. were used by Savrda & Bottjer to characterize
Trace fossil preservation can be considered as units deposited under similar oxygen levels, and
a continuum between two end members: dis- were referred to as oxygen related ichnocoenosis
tinctly burrowed and indistinctly burrowed (ORI) units.
(burrow mottled) beds (Savrda & Bottjer 1991). Once determined, the authors used the
Both end members have well developed 'piped changing sequence of ORI units to recognize
zones' (Fig. 1) - areas where burrows were oxygenation and deoxygenation events, also
emplaced into underlying laminated strata, determining in each case whether the change in
resulting in the preservation of discrete trace oxygen levels had been gradual or rapid. Gradual
fossils. Distinctly burrowed beds also have well- deoxygenation results in a progressive loss of
preserved traces within the primary strata (very ichnogenera, with a consequent simplification
bioturbated sediments that have passed through of the ichnofabric, whereas rapid deoxygenation
the surface-mixed layer). Savrda & Bottjer can be recognized by the presence of 'frozen'
(1986, 1987) developed and applied tiering profiles (Fig. 2). Determination of the nature
models to distinctly burrowed beds to determine and extent of changes in oxygen allows the recon-
their oxygenation histories. struction of a relative oxygenation curve for the
Savrda & Bottjer's model was based on three strata under investigation.
fundamental factors that have been shown in Savrda & Bottjer (1989a) considered that the
some modern environments to change with tiering model can accurately be applied only to

Fig. 2. Tiering patterns predicted for gradual and rapid deoxygenation events in distinctly burrowed beds.
From Wignall (1994).
TRACE FOSSILS AND DYSOXIA 143

distinctly burrowed beds, and proposed a


method for determining relative durations and
magnitudes of oxygenation episodes within
indistinctly burrowed beds of dysoxic strata.
Short-term oxygenation events (from several
days to hundreds of years) are inferred to pro-
duce relatively thin primary strata, with ichno-
genera showing simple vertical segregation
without extensive cross-cutting relationships. In
contrast, protracted oxygenation events (from
hundreds to millions of years) are recognized
by thicker primary strata and extensive cross-
cutting relationships due to the upward migra-
tion of the infaunal associations through the
oxygenation event (see Mcllroy 2004). Magni-
tudes of palaeo-oxygenation are recognized
using the diversity, size and depth relationships
of piped zone burrows: increases in these factors
are taken to reflect increases in levels of palaeo-
oxygenation.
Consideration of Savrda & Bottjer's model
indicates that it may be complicated by the diffi-
culties of recognizing ancient tiering relation-
ships in the fossil record, particularly where the
earliest emplaced traces have been completely
obscured by later ones (Fig. 3). A similar
problem may be identified in situations where
frequent oxygenation events of relatively low
magnitude allow bioturbation sufficient to dis-
turb any laminated strata deposited within

Fig. 4. Cartoon showing the potential for frequent


low-magnitude oxygenation events to allow
bioturbation sufficient to obliterate evidence of
intervening anoxic periods, (a) Oxygenation event
produces an indistinctly burrowed bed. (b) Return to
anoxic conditions; deposition of laminated strata, (c)
Oxygenation event allows bioturbation sufficient to
disturb laminated strata, (d, e) as (b) and (c).

Fig. 3. Cartoon showing the potential for early traces intervening anoxic periods (Fig. 4). This could
to be obliterated by later ones: (a) initial feasibly result in a totally bioturbated sediment
emplacement of piped zone traces; (b, c) production fabric, which would be interpreted as reflecting
of further traces obscures those produced by the continuous oxygenation. Ichnocoenoses in
earliest colonists. which tiering has been controlled by factors
144 K. D. MARTIN

other than oxygen levels may also be difficult to silts and sands; below storm wavebase to upper
identify. continental slope or equivalent areas free from
turbidity flows' (Frey 1975, p. 17). More recently,
the Zoophycos ichnofacies has been interpreted
Ekdale and Mason model as characteristic of stressed, quiet-water environ-
ments, especially those experiencing anoxia
Ekdale & Mason (1988) documented their obser- (Pemberton et al. 1992), over a relatively wide
vations on characteristic late Palaeozoic and range of water depths. Ichnofossils considered
modern deepwater trace fossils, suggesting that characteristic of the Zoophycos ichnofacies are
certain morphological (and therefore beha- typically present at low diversities, though indivi-
vioural) types are typical of low-oxygen environ- dual traces can be abundant. The traces are
ments. typically horizontal or slightly inclined spreiten
Domichnia-dominated aerobic biofacies are structures (Pemberton et al. 1992), including
seen to be succeeded by pascichnial associations Zoophycos, Phycosiphon and Spirophyton (Frey
(typically including Scalarituba, Spirophyton & Pemberton 1984) and Chondrites (Bromley &
and Phycosiphori) where porewaters become dys- Asgaard 1991).
oxic and bottom waters either dysoxic or anoxic. Bromley (1990) also identified a series of recur-
Endostratal pascichnia are found (according to ring trace fossil associations, based on etho-
Ekdale & Mason 1988) in dysoxic but not logical characteristics of the trace fossils rather
anoxic sediments and are rare in oxic sediments. than their environmental setting. These trace
This is attributed to the producing organisms fossil associations are described as ichnoguilds
requiring some interstitial oxygen, as no surface and defined as 'a group of ichnospecies that
connection is maintained, but being limited in express a similar sort of behaviour, belong to
oxic sediments by the paucity of appropriate the same trophic group and occupy a similar
food. Lower dysoxic environments (settings tier or location within the substrate' (Bromley
where anoxic interstitial waters are overlain by 1996, p. 345). Bromley's ichnoguilds were defined
dysoxic waters at the SWI) are typified by using data from Cretaceous Chalk ichnofacies,
fodinichnia-dominated associations. Ekdale & yet one of the ichnoguilds can be seen as applic-
Mason (1988) state that such traces can poten- able to dysaerobic biofacies in general. The
tially penetrate to depths of several decimetres Chondrites-Zoophycos ichnoguild is considered
or even metres, and this may well be true in to comprise mainly deep-tier, deposit-feeding
better-oxygenated environments; however, in (or possibly chemosymbiotic - Bromley 1996)
dysoxic settings they are normally emplaced at structures produced by non-vagile organisms
shallower (centimetre to decimetre) levels (e.g. that are the dominant traces in ichnofabrics
Smith et al 2000). The maintenance of open from oxygen-restricted environments, and are
burrows by the deposit-feeding organisms that also characteristic of the deepest tiers in settings
produce fodinichnial traces is inferred as with better oxygenated bottom-water (Bromley
enabling them to exploit the rich organic matter 1990). The dominant ichnogenera belonging to
content of the sediment while circulating dysoxic this ichnoguild are several size classes of Zoophy-
water through the burrow, for respiration. cos and Chondrites. Savrda (1992) also includes
Ekdale & Mason (1988) consider Chondrites Teichichnus and Trichichnus.
and Zoophycos to be characteristic of such envir- Bromley & Ekdale's (1984) paper has perhaps
onments. been the most widely followed of all work con-
cerning dysaerobic trace fossils. Bromley &
Ekdale identified Chondrites as 'a trace fossil
Zoophycos ichnofacies and Zoophycos- indicator of anoxia in sediments', citing examples
Chondrites ichnoguild of Chondrites occurring as the sole trace fossil in
strata interpreted as having been deposited under
Nine recurring associations of contemporaneous dysoxic bottom-water conditions. The presence
environmentally related traces (ichnofacies) were of Chondrites (often in association with a range
recognized by Seilacher (1967). These ichnofacies of other trace fossils) has subsequently been
were originally interpreted as reflecting bathyme- used erroneously by many authors as evidence
try, but they are now recognized as reflecting a for dysoxic bottom-water conditions (e.g. Jarvis
wide range of environmental parameters, related et al. 1988). However, Bromley & Ekdale
only indirectly to water depth (Frey et al. 1990). (1984) and Bromley & Asgaard (1991) emphasize
Seilacher's Zoophycos ichnofacies was first the point that, where Chondrites is present
thought to be typical of 'sublittoral to bathyal; in completely bioturbated sediments, it does
quiet-water, offshore type conditions; impure not indicate oxygen-deficient seafloors, but
TRACE FOSSILS AND DYSOXIA 145

exploitation of organic matter in deeper tiers of pyrite framboid size-frequency distributions


within the sediment where porewaters may be were performed on samples from a number of
reducing. the case studies investigated. The analyses were
carried out on carbon-coated polished blocks
and polished thin sections using scanning elec-
Methods used to determine oxygen levels tron microscopy.

Both palaeontological and geochemical param-


eters can be used as a proxy for palaeo-oxygena- Oxygen-restricted biofacies
tion. Geochemical techniques are not subject to
the same biases of preservation or sampling as The principal of using species richness to deter-
palaeontological methods, though they can be mine oxygen gradients within dysoxic strata
influenced by the effects of time-averaging. Use was recognized by Rhoads & Morse (1971) and
of both geochemical and palaeontological data- Byers (1977). This method was developed by
sets in evaluating levels of palaeo-oxygenation Wignall & Hallam (1991) in their oxygen-
therefore provides a more robust framework restricted biofacies (ORB) scheme. Wignall &
against which to compare trace fossil data. Hallam used shelly macrofaunal and lithological
Gamma-ray spectrometry, pyrite framboid data from British Jurassic black shales as a basis
analyses and the assignment of the benthic for six biofacies that distinguish between anoxic,
macrofauna to oxygen-restricted biofacies are lower- and upper-dysoxic environments. The
used herein to interpret oxygenation. macrofauna of the case studies presented here
have been used to classify the sections according
to the ORB scheme, thus providing a measure of
Gamma-ray spectrometry palaeo-oxygenation against which to compare
the trace fossil data.
Gamma-ray spectrometry has been employed in
the evaluation of sedimentary rocks since the
1950s, significant works including those of Toarcian of Ravenscar, North Yorkshire
Adams & Weaver (Adams & Weaver 1958;
Adams et al 1959). Adams & Weaver (1958) The early Toarcian in Britain is characterized by
recognized that the U and Th contents of fine- a number of small-scale extinction events (Little
grained elastics varied according to the deposi- & Benton 1995; Hallam 1996) that may have
tional redox conditions. More recently this been caused by an oceanic anoxic event (OAE)
concept has been developed by Myers & Wignall (Jenkyns 1988). The exaratum Subzone (Fig. 5)
(1987) and Wignall & Myers (1988), who showed is thought to represent the maximum develop-
that Th/U ratios can be used to distinguish ment of this anoxic event (Saelen et al. 1996);
between anoxic, dysoxic and oxic depositional subsequent subzones record the recovery from
settings. Th/U ratios of 3, 2 and 0.5 are taken the event.
as the boundary between dysoxic and oxic Coastal sections in North Yorkshire provide
strata in shales, marls and limestones respectively the best record of the British Toarcian and
(Wignall & Myers 1988; Martin 2001). Data were have consequently been intensively studied (e.g.
collected according to the criteria of Myers & Dean 1954; Howarth 1962; Knox 1984). How-
Wignall (1987). ever, the ichnology of this interval has received
little attention and is therefore investigated
herein.
Pyrite framboid size—frequency analyses Fieldwork was carried out on wave-cut plat-
form and cliff exposures on the foreshore
Pyrite is an extremely common authigenic between Old Peak [NZ 980 025] and Blea Wyke
mineral in mudrocks. It is typically present as a [NZ991 016] near the village of Ravenscar,
number of different morphologies, including North Yorkshire. Approximately 90 m of strata,
framboids, clusters, aggregates of bladed crystals from the falciferum to the levesquei ammonite
and equant crystals. Recent work (Wilkins et al. zone (Fig. 5), were logged (Figs 6a, b), beginning
1996; Wignall & Newton 1998) has shown that at bed x of Howarth (1962). Parts of the section
the size distributions of pyrite framboids in were inaccessible, however, preventing the collec-
modern environments reflect the depositional tion of data. Gamma-ray spectrometry was
oxygen conditions, and that this can be used to carried out at ~1 m intervals, and the occurrence
accurately reconstruct the palaeoredox condi- of hard-shelled fauna was noted throughout the
tions of ancient clastic environments. Analyses sequence. These data provide Th/U ratios and
146 K. D. MARTIN

Fig. 5. North Yorkshire Toarcian lithostratigraphy, informal nomenclature and ammonite zonation. Shaded
box shows the position of the sequence studied. Compiled from data in Knox (1984) and Howarth (1992).

ORB with which to interpret the depositional Bositra buchii} occur as autochthonous shell
oxygen levels. pavements on some bedding planes. The abun-
dance of this epibenthic fauna varies within the
Jet Rock Member, decreasing upwards from
Evidence for oxygenation the middle of bed xii to the top of bed xiv,
where the fauna is sparse. The lower part of the
Evidence for oxygenation is provided using Th/U member is therefore assigned to ORB 4 and the
ratios and ORB. Within the lower part of the upper part to ORB 3, indicating that episodically
Jet Rock Member, Th/U values under 3 indicate dysoxic conditions prevailed throughout the
the presence of authigenic U, and therefore deposition of the member, and that periods of
oxygen-restricted depositional conditions. improved oxygenation were of longer duration
Above the middle of bed xii Th/U values range in the lower part.
between 3 and 8, indicating oxic depositional The lower 3 m of the Alum Shale Member con-
conditions. tain the same benthic faunal assemblage as the
The Jet Rock Member contains a common Jet Rock Member, though with greater abun-
nektonic fauna of ammonites, fish debris and dance and therefore interpreted as belonging to
belemnites. Isolated crinoid ossicles occur from ORB 4. A gap in the section of approximately
bed xix upwards. Epibenthic bivalves (Pseudo- 4m occurs at this point; during this interval
mytiloides dubius, Meleagrinella substriata and Pseudomytiloides dubius disappears and the
TRACE FOSSILS AND DYSOXIA 147

hrace fossils within the Jet Rock Member and Alhale Member of the Toarcian
of Ravenscar, North Yorkshire. Bed numbers are those of Howarth (1962); logs for the inaccessible parts of
the sequence are compiled from observations of cliff exposures and data in Knox (1984).

infaunal bivalve Dacryomya ovum appears. The scattered throughout the sediment rather than
benthic fauna remains the same up to bed xxvii, restricted to particular bedding planes: thus
after which point there is another gap in expo- conditions are interpreted as having been persis-
sure within which the infaunal bivalve Pleuromya tently lower dysoxic.
costata appears. The last occurrence of Meleagri- The lack of accessible exposure of the Peak
nella substriata is recorded in bed xxxii; the tri- Mudstone Member and most of the Fox Cliff
gonid Vaugonia pulchella appears 4.5m above Siltstone Member has meant that ORB cannot
bed xliii. The benthic assemblage of B. buchii, be determined for these units. Where the expo-
D. ovum, P. costata and V. pulchella continues sure resumes, towards the top of the Fox Cliff
at least up to bed 37, after which there is a Siltstone Member, the fauna is sparse and
~23 m gap in exposure. The fauna of the Alum poorly preserved. Shelly fragments are abun-
Shale Member is assigned to ORB 4 and dant, but the recognizable fauna consists only
indicates that the member was deposited under of abraded specimens of belemnites, the trigonid
lower dysoxic conditions. The faunas are Vaugonia pulchella and serpulids.
148 K. D. MARTIN

Fig. 6. (b) Distribution of trace fossils within the Alum Shale Member, Peak Mudstone Member, Fox Cliff
Siltstone Member and Grey Sandstone Member of the Toarcian of Ravenscar, North Yorkshire. Roman
numerals are bed numbers of Howarth (1962), Arabic numerals are bed numbers of Dean (1954).

The combined data from Th/U ratios and the determination of ORB; however, Th/U
ORB indicate that the Jet Rock Member was ratios give evidence for deposition under oxic
deposited under episodically dysoxic conditions, conditions.
with periods of improved oxygenation being The entire sequence is interpreted as recording
of longer duration in the lower than the upper a gradual increase in palaeo-oxygenation follow-
part of the member. The Alum Shale Member ing the early Toarcian OAE. This broadly agrees
was deposited under persistent lower dysoxic with the interpretation of Saelen et al. (1996),
conditions. Determination of palaeo- based on their analyses of stable O and C
oxygenation in the Peak Mudstone Member isotopes from belemnite rostra. However, the
and lower Fox Cliff Siltstone Member has not evidence from this study indicates that the
been possible owing to the lack of accessible Alum Shale Member was deposited under
exposure. Poor preservation of the hard-shelled lower dysoxic conditions, whereas Saelen et al.
fauna within the top of the Fox Cliff Siltstone (1996) consider that the water column was oxy-
Member and the Grey Shale Member precludes genated by the top of the Jet Rock Member.
TRACE FOSSILS AND DYSOXIA 149

Trace fossils Burrow widths range between 2mm and


10 mm; preserved lengths can exceed 28cm.
The distribution of trace fossils is shown in Fig- The burrows are partially pyritized. Where
ures 6a and b. Eight trace types are recognized; pyrite is present the burrow margin is irregu-
however, only four of these are assignable to lar; where pyrite is absent the margin is
recognized ichnogenera. The four remaining smooth. These traces were probably formed
trace types are described briefly below. by pascichnial activity.
Trace type 1 are elongate, straight to sinuous, Distinct trace fossils, of type 4, first appear
non-branching, approximately horizontal within bed xx (Fig. 6a). The 3m of overlying
traces with a diffuse pyrite fill. Widths range strata contain no distinct traces, but from bed
from 4mm to 6mm; preserved lengths from xxiv trace types 2 and 3 occur and increase in
2cm to 3cm. Their morphology suggests a abundance over ~2m, after which point there
pascichnial origin. is a ~6m gap in exposure. On the resumption
Trace type 2 are horizontally oriented, non- of exposure, in bed xxxii, trace types 2 and 3
branching, straight to slightly sinuous burrows are present. Type 3 traces were not recorded
with a subcircular cross-section and a dense between beds xxxiv and xlii; however, type 4
pyrite fill. Burrow diameters are constant traces were recorded in bed xxxv, and type 1
within individual traces; between traces they traces in beds xxxvi and xxxvii. Possible exam-
vary from 1 mm to 3 mm. Preserved burrow ples of Chondrites are present within bed xxxviii:
lengths reach a maximum of 5 cm. Their mor- these traces are predominantly horizontal and
phology suggests that they were the product of display sparse low-angle branching. From bed
pascichnial activity. xliv 5 m of strata contain only trace types 2 and
Trace type 3 are simple, vertically oriented 3, which decrease in abundance upwards until
burrows with a subcircular cross-section of they disappear (Fig. 6b). Small Chondrites
1-3 mm diameter and a dense pyrite fill. (~lmm diameter) are present within bed 42;
Their morphology indicates a domichnial above this point there is a gap in exposure of
origin. 20m. The exposure resumes towards the top
Trace type 4 comprises horizontal to obliquely of the Fox Cliff Siltstone Member, revealing
oriented, straight to curved, elongate traces traces of types 2 and 3 co-occurring with Rhizo-
without wall or internal structures (Fig. 7). corallium in a background of burrow-mottled

Fig. 7. Trace type 4. Bed xx, Alum Shale Member, Toarcian, Ravenscar.
150 K. D. MARTIN

Fig. 8. Cartoon of trace fossil occurrence with relation to oxygenation in the Toarcian of Ravenscar.

sediments. Type 2 and 3 traces disappear before from lower dysoxic to oxic depositional environ-
the top of the member, just beneath the level at ments (Fig. 8). The diversity of distinct traces
which Chondrites reappears. The Grey Sand- increases in parallel with the increase in oxygena-
stone Member is characterized by completely tion, and maximum burrow diameters show a
bioturbated sediments, with Chondrites and similar general increasing trend. This supports
Rhizocorallium, from bed 71 Thalassinoides, the model of Savrda and Bottjer (1986, 1987).
and, at the very top of the studied section, Taeni- Ekdale and Mason's (1988) model predicts
dium. that an increase in oxygenation will produce a
The first traces to appear (type 4) have maxi- sequence of fodinichnia-, pascichnia- and then
mum diameters of 6mm. Following a gap in domichnia-dominated ichnological associations.
trace fossil occurrence, maximum diameters All three behavioural categories are represented
increase from 1 mm in bed xxiv to 2 mm in bed in the Ravenscar section, but, in contrast to
xxvi. Maximum diameters of 3mm occur over Ekdale and Mason's model, a sequence of
13.5m of strata from bed xxxii, with the excep- pascichnia-dominated assemblages followed by
tion of a single occurrence of trace type 4 in domichnia-dominated assemblages parallels the
bed xxxv, which has a diameter of 10mm. Iso- increase in oxygenation. The observations from
lated examples of Chondrites at the top of the this study do not therefore support Ekdale &
Alum Shale Member record maximum diameters Mason's model. Wignall (1991) reported ichno-
of 1 mm. The maximum trace diameter recorded coenoses from the Upper Jurassic Kimmeridge
in the upper part of the Fox Cliff Siltstone Clay in which a pascichnia-dominated associa-
Member is 10 mm; this increases to 25mm tion of trace fossils occurred under lower
within the Grey Sandstone Member. oxygen levels than other ethological types.
Pascichnia (type 4 traces) are the first traces to Wignall suggested that this assemblage was con-
appear in the Alum Shale Member. Pascichnia trolled by substrate consistency, inferring that
dominate throughout the member, though soupy substrates favour the production of
domichnia and, occasionally, fodinichnia (Chon- traces that did not require the maintenance of a
drites) are also present. The Fox Cliff Siltstone connection to the sediment-water interface (i.e.
and Grey Sandstone Members are dominated pascichnia). The Alum Shale Member has some
by domichnia, though pascichnia and fodin- evidence for soft substrate conditions - ammo-
ichnia also occur. nites within bed xx of the member were found
in vertical and oblique orientations in addition
to the more usual horizontal orientation. The
Comparison with current oxygen-related occurrences of pascichnia-dominated ichnocoe-
trace fossil models noses in this study may therefore support
WignalFs (1991) conclusions.
The trace fossils of the Whitby Mudstone For- The identification of Chondrites as the sole
mation and Blea Wyke Sandstone Formation occurring trace under dysoxic conditions
at Ravenscar were formed during a transition (Bromley & Ekdale 1984) is not supported by
TRACE FOSSILS AND DYSOXIA 151

the Ravenscar case study. Although Chondrites The shales generally contain a low-diversity
occurs within the strata deposited under lower allochthonous faunal assemblage made up of
dysoxic conditions, it is preceded by four differ- fish scales, echinoid spines, bivalve fragments
ent trace types. and, occasionally, ammonites. Large scours, up
to 60cm in length and 6cm deep, occur at the
base of the shales of bed 6; these are infilled
Blue Lias of Lyme Regis, Dorset with calcareous material containing abundant
fragmentary fauna, including Ostrea, other
The Blue Lias is a widely used descriptive term bivalves, rhynchonellids and crinoids.
that refers to successions of alternating shales The marls contain a body fossil assemblage
and limestones deposited over much of north- that normally includes Gryphaea, Ostrea, a
west Europe (Callomon & Cope 1995) during small unidentified bivalve, fish scales and frag-
the latest Triassic and early Jurassic (Hettan- ments of bivalves. The marls of bed 5 addition-
gian-Sinemurian). The sediments of the Blue ally contain abundant Calcirhynchia calcaria.
Lias are typically developed in repeating deposi- The limestones contain a more diverse fauna
tional cycles (Fig. 9). These cycles are not always than the marls; this is probably the result of
complete; some cycles pass from shale to marl early diagenetic cementation of the limestones
and back to shale, whereas others lack the pale allowing better preservation of the fauna rather
marls and limestones. Milankovitch cyclicity, than the original faunal communities being
particularly obliquity (House 1985), has been more diverse. The limestones normally preserve
suggested as the controlling mechanism behind many of the following: C. calcaria, Gryphaea,
these rhythms. The origin of the limestones (pri- Ostrea, small unidentified bivalves, the gastro-
mary versus diagenetic) has previously been the pod ?Pleurotemaria, ammonites, crinoids and
subject of debate; they are now widely accepted echinoid spines. Bed 7 is characterized by parti-
as being diagenetic (e.g. Hallam 1986; Bottrell cularly abundant crinoid debris. Bed 13 is
& Raiswell 1990). highly distinctive, containing abundant very
The section at Lyme Regis was measured large ammonites and nautiloids throughout.
approximately 1 km southwest of The Cobb, at Additionally, a layer of very abundant mixed
SY 3285 9110. The studied section lies within crinoid and bivalve debris is present towards
the conybeari Subzone of the bucklandi Zone the top of the bed; these shell fragments are occa-
(Sinemurian). The base of the section measured sionally concentrated within the chambers of
here corresponds to bed 21 of Lang (1924). ammonoids. At the top of bed 13, Gryphaea is
very abundant, and there are large pieces of
wood with encrusting oysters.
Evidence for oxygenation Changes in ORB through the sequence are pre-
dominantly coincident with changes in substrate
The Blue Lias at Lyme Regis contains a relatively from shales to marls. The shale of bed 3 is devoid
diverse fauna, including well-preserved skeletons of all fauna and is thus assigned to ORB 1, indi-
of marine reptiles (e.g. Martill 1995). cating that the water column was anoxic. The
shales of beds 9 and 15 contain nektonic forms
and small fragments of bivalves, which are
considered to be allochthonous: thus they are
assigned to ORB 2, reflecting deposition in
conditions of anoxia with a habitable water
column. Beds 6, 11 and 12 contain shales with
rare occurrences of the benthic form C. calcaria;
these are assigned to ORB 3 and reflect episodi-
cally dysoxic conditions.
The marl-limestone units generally contain
common benthic forms, including thick-shelled
bivalves (e.g. Gryphaea, Ostrea) and crinoids.
These forms are indicative of oxic depositional
conditions.
Analysis of the size-frequency distribution of
pyrite framboids was carried out on a shale
Fig. 9. Idealized sedimentary cycle within the Blue sample from bed 3. The sample contained abun-
Lias. Based on field logging within the conybeari dant small framboids, giving a size-frequency
Subzone, Lyme Regis, Dorset. distribution characteristic of deposition under
152 K. D. MARTIN

euxinic conditions. The marls and limestones did an abrupt break in the presence of hard-shelled
not contain sufficient pyrite to be analysed using fauna at the top of the limestone-marl units.
this method. This implies that the decrease in oxygen levels
Pyrite framboid and ORB data thus indicate was rapid. However, the erosive bases of the
that conditions were persistently anoxic during shales mean that an undetermined amount of
the deposition of the shales of bed 3. The rest sediments and their associated fauna have been
of the shale beds are interpreted as having been removed, complicating the interpretation.
deposited under conditions of episodic dysoxia.
Oxic conditions are considered to have occurred
during the deposition of the marl-limestone Trace fossils
units. The improvement in oxygenation is con-
sidered to have occurred relatively rapidly, as a The ranges of the ichnogenera recognized within
diverse hard-shelled fauna is usually present the sequence are shown in Figure 10. The traces
close to the base of the marls. There is typically can be divided into two groups, based on their

Fig. 10. Ichnological data from the Lyme Regis section. Behavioural types: Dom., domichnia; Fod.,
fodinichnia; Pasc., Pascichina. Oxygen levels: 1, anoxic/episodically dysoxic; 2, lower dysoxic; 3, upper dysoxic;
4, oxic.
TRACE FOSSILS AND DYSOXIA 153

occurrence relative to the interpreted oxygen The transition from laminated shales up into
levels. The first includes those traces that are bioturbated marls represents the change from
present only within the marl-limestone units; anoxic/episodically dysoxic conditions to oxic
the second comprises traces that are present at conditions that prevailed during the deposition
the transition from shales to marls. of the marl-limestone units. The shale/marl
All of the trace fossils recognized occur within boundaries can be categorised as one of two
the marl-limestone units (Fig. 10). Large hori- types. In the first type (bed 3) the shale passes
zontal expanses of the beds on the wave-cut directly into homogenised marl lacking distinct
platforms provide good exposures of the associa- traces; this marl probably represents sediment
tions. This allows recognition of the cross-cutting that was originally laminated but was disturbed
relationships between the ichnogenera, and by organisms within the surface-mixed layer
shows that complex, tiered ichnocoenoses were following the increase in oxygenation. In the
developed (Fig. 11) (see discussion of ichno- second type, the bed boundary region contains
coenoses in Mcllroy 2004). The upper tier was a piped zone of distinct traces emplaced within
occupied by organisms that thoroughly biotur- the tops of the laminated strata. Piped zones
bated the substrate but did not leave distinct are present within beds 9, 11, 12 and 15. The
traces. The middle tier included the producers piped zone of bed 9 contains Chondrites, that of
of Arenicolites, Thalassinoides, Diplo crater ion, bed 11 Planolites, that of bed 12 Chondrites and
Palaeophycus and Taenidium, whereas the deep- Thalassinoides, and that of bed 15 Chondrites
est tier contained the organisms responsible for and Diplocraterion. Although the Thalassinoides
Chondrites and Planolites. of bed 12 and the Diplo crater ion of bed 15
Trace fossil assemblages within the marl- occur within the piped zones, cross-cutting
limestone units are moderately diverse, contain- relationships indicate that they were emplaced
ing between two and eight distinct ichnogenera. later than the piped-zone Chondrites, and also
The beds with the lowest diversity tend to be later than other traces, which were emplaced
those in which the recognizable traces are within the (oxic) marl-limestone units.
poorly preserved members of the deepest tier: The Sinemurian sequence at Lyme Regis
thus the apparent low diversities may be a section reflects an environment that was subject
function of preservation. Maximum burrow to long-term (thousands of years) fluctuations
diameters within the units are relatively large in oxygen levels, from anoxic or episodically dys-
(Fig. 10), varying between 8mm and 40mm. oxic during the deposition of the laminated
Penetration depths are difficult to measure shales to oxic during the deposition of the
owing to the lack of vertical exposure; an marl-limestone units. Trace fossils were not
example of Diplo crater ion from bed 4 had a emplaced during the deposition of either the
minimum penetration depth of 7cm. Most of anoxic or episodically dysoxic shales. The initial
the units contain fodinichnial, domichnial and colonists under reoxygenation gradients were
pascichnial traces (Fig. 10); a few lack domich- small organisms that disturbed the shale laminae
nial traces, whereas others lack pascichnia. but did not produce distinct traces. Chondrites

Fig. 11. Cartoon of ideal tiered community developed within marl-limestone units. Based on field logging
within the conybeari Subzone, Lyme Regis, Dorset.
154 K. D. MARTIN

and Planolites were the first distinct traces to be References


emplaced and penetrated between 4cm and
10cm into the underlying shales - deeper than ADAMS, J. & WEAVER, R. 1958. Thorium and uranium
the initial colonists. These observations indicate ratios as indicators of sedimentary processes:
examples of concept of geochemical fades. Amer-
that penetration depths, burrow diameters and ican Association of Petroleum Geologists Bulletin,
trace diversity increase with increasing oxygena- 42, 387^30.
tion, conforming to the predictions of Savrda & ADAMS, J., OSMOND, J. & ROGERS, J. 1959. The
Bottjer (1986, 1987). The occurrence of Chon- geochemistry of uranium and thorium. Physical
drites as one of the first distinct traces to be Chemistry of the Earth, 3, 229-328.
emplaced within the reoxygenation gradient sup- BOTTRELL, S. & RAISWELL, R. 1990. Primary versus
ports the observations of Bromley & Ekdale diagenetic origin of Blue Lias rhythms (Dorset,
(1984); however the co-occurrence of Planolites UK): evidence from sulphur geochemistry. Terra
shows that other trace types may also be charac- Nova, 1, 451^56.
teristic of restricted oxygen conditions. Both BROMLEY, R. 1990. Trace Fossils. Chapman & Hall,
London.
fodinichnia and pascichnia occurred under the BROMLEY, R. 1996. Trace Fossils: Biology, Taphonomy
lowest oxygen conditions; this does not conform and Applications. Chapman & Hall, London.
to the model of Ekdale & Mason (1988). BROMLEY, R. & ASGAARD, U. 1991. Ichnofacies: a
mixture of taphofacies and biofacies. Lethaia, 24,
152-163.
BROMLEY, R. & EKDALE, A. 1984. Chondrites: a trace
Conclusions fossil indicator of anoxia in sediments. Science,
224, 872-874.
The two Jurassic case studies presented here use BROMLEY, R. & EKDALE, A. 1986. Composite ichnofab-
geochemical and palaeontological data to inter- rics and tiering of burrows. Geological Magazine,
pret palaeo-oxygen levels, allowing the occur- 123, 59-65.
rence of trace fossils relative to oxygenation to BYERS, C. 1977. Biofacies patterns in euxinic basins:
be investigated. Savrda & Bottjer's (1986, 1987) a general model. In: COOK, H. & ENOS, P. (eds)
models of oxygen-related trace occurrences, Deep- Water Clastic Environments, Society of Eco-
based on changes in trace diversity, diameters nomic Palaeontologists and Mineralogists, Special
and penetration depths, applied to both of the Publication, 25, 5-17.
case studies. This supports their assertions that CALLOMON, J. & COPE, J. 1995. The Jurassic geology
of Dorset. In: TAYLOR, D. (ed.) Geology of the
oxygen levels have a strong control over these British Jurassic. The Geological Society, London,
parameters, and that they can be used to 51-103.
distinguish between strata deposited under differ- DEAN, W. 1954. Notes on part of the Upper Lias
ent oxygen conditions at different temporal succession at Blea Wyke, Yorkshire. Proceedings
scales. Ekdale & Mason's (1998) behaviourally of the Yorkshire Geological Society, 29, 161-179.
based model was found not to apply in either DEMAISON, G. & MOORE, G. 1980. Anoxic environ-
case study, implying that factors other than ments and oil source bed genesis. Bulletin of the
oxygenation - possibly substrate stability - may American Association of Petroleum Geologists, 64,
control the occurrence of the trace-producing 1179-1209.
DIAZ, R. J. & ROSENBERG, R. 1995. Marine benthic
organisms. hypoxia: a review of its ecological effects and the
Chondrites occurred under conditions of behavioural responses of benthic macrofauna.
bottom-water dysoxia in the Toarcian at Raven- Oceanography and Marine Biology: An Annual
scar, and was present both in piped zones in Review, 33, 245-303.
reoxygenation gradients and as a deep-tier EKDALE, A. & MASON, T. 1988. Characteristic trace
trace in oxic sediments from the Sinemurian of fossil assemblages in oxygen-poor sedimentary
Lyme Regis. These observations support those environments. Geology, 16, 720-723.
of Bromley & Ekdale (1984). However, other FREY, R. 1975. The realm of ichnology: its strengths
trace fossils were found to co-occur with Chon- and limitations. In: FREY, R. (ed.) The Study of
drites in dysoxic settings, or to be present at Trace Fossils. Springer-Verlag, Berlin, 18-38.
FREY, R. & PEMBERTON, S. G.I984. Trace fossil facies
lower oxygen levels: thus other ichnogenera models. In: WALKER, R. (ed.) Facies Models.
may also be characteristic of sediment dysoxia/ Geological Association of Canada, Geoscience
anoxia. Canada Reprint Series, 1, 189-207.
FREY, R., PEMBERTON, S. G. & SAUNDERS, T. 1990. Ich-
This work was funded by NERC as part of a PhD thesis nofacies and bathymetry: a passive relationship.
and carried out at the University of Leeds under the Journal of Palaeontology, 64, 155-158.
supervision of P. Wignall to whom thanks are due. P. HALLAM, A. 1986. Origin of minor limestone-shale
Doyle, D. Mcllroy and C. R. C. Paul provided helpful cycles: climatically induced or diagenetic? Geology,
reviews 14, 609-612.
TRACE FOSSILS AND DYSOXIA 155

HALLAM, A. 1996. Recovery of the marine fauna in allostratigraphic significance. In: WALKER, R. &
Europe after the end-Triassic and early Toarcian JAMES, N. (eds) Facies Models: Response to Sea
mass extinctions. In: HART, M. (ed.) Biotic Recov- Level Change. Geological Association of Canada,
ery from Mass Extinction Events. Geological Ottawa, 42-72.
Society, London, Special Publications, 102, 231- RHOADS, D. & MORSE, J. 1971. Evolutionary and eco-
236. logic significance of oxygen deficient marine
HOUSE, M. 1985. A new approach to an absolute time- basins. Lethaia, 4, 413^28.
scale from measurements of orbital cycles and SAELEN, G., DOYLE, P. & TALBOT, M. 1996. Stable-
sedimentary microrhythms. Nature, 315, 721-725. isotope analyses of belemnite rostra from the
HOWARTH, M. 1962. The Jet Rock Series and the Alum Whitby Mudstone Formation, England: surface
Shale Series of the Yorkshire Coast. Proceedings of water conditions during the deposition of a
the Yorkshire Geological Society, 33, 381^22. marine black shale. Palaios, 11, 97-117.
HOWARTH, M. 1992. The ammonite family Hildocerati- SAVRDA, C. 1992. Trace fossils and benthic oxygena-
dae in the Lower Jurassic of Britain. Monograph of tion. In: MAPLES, C. & WEST, R. (eds) Trace
the Palaeontological Society, 1. Fossils. Palaeontological Society, Knoxville,
HUDSON, J. & MARTILL, D. 1991. The lower Oxford Short courses in Palaeontology, 5, 172-196.
Clay: production and preservation of organic SAVRDA, C. & BOTTJER, D. 1986. Trace fossil model for
matter in the Callovian (Jurassic) of central Eng- reconstruction of palaeo-oxygenation in bottom
land. In: TYSON, R. & PEARSON, T. (eds) Modern waters. Geology, 14, 3-6.
and Ancient Continental Shelf Anoxia. Geological SAVRDA, C. & BOTTJER, D. 1987. Trace fossils as indica-
Society, London, Special Publications, 58, 363-379. tors of bottom water redox conditions in ancient
JAR vis, I., CARSON, G. ET AL. 1988. Microfossil assem- marine environments. In: BOTTJER, D. (ed.) New
blages and the Cenomanian-Turonian (late Concepts in the Use ofBiogenic Sedimentary Struc-
Cretaceous) oceanic anoxic event: new data from tures for Palaeoenvironmental Interpretation.
Dover, England. Cretaceous Research, 9, 3-103. Society of Economic Palaeontologists and Miner-
JENKYNS, H. 1988. The early Toarcian (Jurassic) anoxic alogists, Pacific Section, Los Angeles, 52, 3-26.
event: stratigraphic, sedimentary and geochemical SAVRDA, C. & BOTTJER, D. 1989a. Anatomy and implica-
evidence. American Journal of Science, 288, 101— tions of bioturbated beds in 'black shale' sequences:
151. examples from the Jurassic Posidonienscheifer
KNOX, R. B. 1984. Lithostratigraphy and depositional (southern Germany). Palaios, 4, 330-342.
history of the late Toarcian sequence at Raven- SAVRDA, C. & BOTTJER, D. 1989b. Development of a
scar, Yorkshire. Proceedings of the Yorkshire trace fossil model for the reconstruction of
Geological Society, 45, 99-108. palaeo-bottom-water redox conditions: evaluation
LANG, W. 1924. The Blue Lias of the Dorset and Devon and application to Upper Cretaceous Niobrara
coasts. Proceedings of the Geologists' Association, Formation, Colorado. Palaeogeography, Palaeo-
35, 169-185. climatology, Palaeoecology, 74, 49-74.
LITTLE, C. & BENTON, M. 1995. The early Jurassic mass SAVRDA, C. & BOTTJER, D. 1991. Oxygen related bio-
extinction: a global long-term event. Geology, 23, facies in marine strata: an overview and update.
495^98. In: TYSON, R. & PEARSON, T. (eds) Modern and
MARTILL, D. 1995. An ichthyosaur with preserved soft Ancient Continental Shelf Anoxia. Geological
tissue from the Sinemurian of southern England. Society, London, Special Publications, 58, 201-220.
Palaeontology, 38, 897-903. SAVRDA, C., BOTTJER, D. & GORSLINE, D. 1984. Develop-
MARTIN, K. D. 2001. Secular trends in dysaerobic trace ment of a comprehensive oxygen-deficient marine
fossils. PhD thesis, University of Leeds. biofacies model: evidence from Santa Monica, San
MclLROY, D. In: MC!LROY, D. (ed.) Some ichnological Pedro and Santa Barbara Basins, California Con-
concepts, methodologies, applications and frontiers. tinental Borderland. Bulletin of the American
Geological Society, London, Special Publications, Association of Petroleum Geologists, 68, 1179-1192.
228, 3-27. SEILACHER, A. 1967. Bathymetry of trace fossils.
MYERS, K. & WIGNALL, P. 1987. Understanding Marine Geology, 4, 413^28.
Jurassic organic-rich mudrocks: new concepts SMITH, C. R., LEVIN, L. A., HOOVER, D. J., McMuR-
using gamma-ray spectrometry and palaeo- TRY, G. & GAGE, J. D. 2000. Variations in biotur-
ecology: examples from the Kimmeridge Clay of bation across the oxygen minimum zone in the
Dorset and the Jet Rock of Yorkshire. In: northwest Arabian Sea. Deep Sea Research II,
LEGGETT, J. & ZUFFA, G. (eds) Marine Clastic 47, 227-257.
Sedimentology. Graham & Trotman, London, TYSON, R. & PEARSON, T. 1991. Modern and ancient
172-189. continental shelf anoxia: an overview. In: TYSON,
OSCHMANN, W. 1991. Anaerobic-poikiloaerobic- R. & PEARSON, T. (eds) Modern and Ancient Conti-
aerobic: a new facies zonation for modern and nental Shelf Anoxia. Geological Society, London,
ancient neritic redox facies. In: EINSELE, G., Special Publications, 58, 1-24.
RICKEN, W. & SEILACHER, A. (eds) Cycles and WIGNALL, P. 1991. Dysaerobic trace fossils and ichno-
Events in Stratigraphy. Springer-Verlag, Berlin, fabrics in the Upper Jurassic Kimmeridge Clay of
565-571 southern England. Palaios, 6, 264-270.
PEMBERTON, S. G., MAC£ACHERN, J. & FREY, R. 1992. WIGNALL, P. 1994. Black Shales. Oxford University
Trace fossil facies models: environmental and Press, Oxford.
156 K. D. MARTIN

WIGNALL, P. & HALLAM, A. 1991. Biofacies, strati- WIGNALL, P. & NEWTON, R. 1998. Pyrite framboid dia-
graphic distribution and depositional models of meter as a measure of oxygen deficiency in ancient
British onshore Jurassic black shales. In: TYSON, mudrocks. American Journal of Science, 298, 537-
R. & PEARSON, T. (eds) Modern and Ancient 552.
Continental Shelf Anoxia. Geological Society, WILKINS, R., BARNES, H. & BRANTLEY, S. 1996. The
London, Special Publications, 58, 291-309. size distribution of framboidal pyrite in modern
WIGNALL, P. & MYERS, K. 1988, Interpreting benthic sediments: an indicator of redox conditions.
oxygen levels in mudrocks: a new approach. Geochimica et Cosmochimica Acta, 60, 3897-
Geology, 16, 452-455. 3912.
Ichnology of Carboniferous tide-influenced environments and
tidal flat variability in the North American Midcontinent
M. GABRIELA MANGANO & LUIS A. BUATOIS

Conicel-Insugeo, Casilla de correo 1, correo central, 4000 San Miguel de Tucumdn, Argentina
(e-mail: ichnolog@ infovia.com.ar)

Abstract: Trace fossils are sensitive indicators of environmental fluctuations and, accordingly,
ichnological studies have the potential to improve facies characterization of marginal-marine
systems. Carboniferous intertidal deposits in eastern Kansas and western Missouri accumu-
lated under contrasting palaeoenvironmental conditions, ranging from the open shoreline to
fluvio-estuarine transitions. Comparative analysis of these exposures illustrates lateral varia-
tions in trace-fossil content and allows characterization of the intertidal ichnofaunas developed
in three sub-environments: open marine, restricted bays and fluvio-estuarine transitions. Open-
marine tidal flat ichnofaunas are characterized by (1) high ichnodiversity, (2) marine ichnotaxa
produced by both euryhaline and stenohaline forms, (3) the presence of both infaunal and
epifaunal traces, (4) the presence of simple and complex structures produced by presumed
trophic generalists and specialists respectively, (5) dominance of horizontal trace fossils of
the Cruziana ichnofacies, (6) presence of multispecific associations, (7) high density, and (8)
wide size range. This ichnofauna is present in heterolithic deposits and reflects the activity of
a biota that inhabited tidal flats dominated by normal-marine salinities and connected directly
to the open sea. Restricted-bay ichnofaunas display (1) low ichnodiversity, (2) ichnotaxa com-
monly found in marine environments, but produced by euryhaline organisms, (3) dominance of
infaunal traces rather than epifaunal trails, (4) simple structures produced by opportunistic
trophic generalists, (5) a combination of vertical and horizontal trace fossils from the Skolithos
and Cruziana ichnofacies, (6) the presence of monospecific associations, (7) variable density,
and (8) small size. This assemblage occurs in heterolithic facies and records the activity of a
brackish-water benthic fauna inhabiting intertidal areas of estuarine basins and embayments.
Fluvio-estuarine ichnofaunas are characterized by (1) moderate to relatively high diversity, (2)
forms typically present in continental environments, (3) the dominance of surface trails and
absence of burrows, (4) temporary structures produced by a mobile deposit-feeding fauna,
(5) a mixture of trace fossils belonging to the Scoyenia and Mermia ichnofacies, (6) moderate
density of individual ichnotaxa, (7) absence of monospecific suites, and (8) small size. This
assemblage occurs in tidal rhythmites and records the activity of a typical freshwater/terrestrial
benthos inhabiting tidal flats that were developed in the most proximal zone of the inner
estuary under freshwater conditions. Through integration of ichnological and sedimentological
data, conventional sedimentological interpretations of marginal-marine depositional systems
can be refined and enhanced.

Tidal flats are complex depositional environments energy, and substrate types and consistency (e.g.
that are extremely variable, depending on coastal Reise 1985; Mangano et al 2002a). Although
physiography, climate, dominant sedimentary numerous studies have focused on the animal-
processes and tidal regimes, among other param- sediment interactions in modern tidal flats (see
eters. They develop in a wide spectrum of coastal review in Mangano et al. 2002a), relatively few
environments, such as open coasts, bays, lagoons authors have attempted to apply observations
and estuaries. In this study, tidal flats are consid- on modern animal-sediment interactions to the
ered as that region extending between spring high- study of ancient tidal flats (e.g. Goodwin &
and spring low-tide levels, grading landward into Anderson 1974; Miller & Knox 1985; Wescott &
the supratidal marshes and seaward into the sub- Utgaard 1987; Simpson 1991).
tidal region (after van Straaten 1954). In contrast, Late Palaeozoic eustatic sea-level rises caused
some authors also include flat to gently sloping the development of extensive epicontinental
upper-subtidal areas and lower-supratidal areas seas over the cratonic USA Midcontinent
in the definition of the 'tidal flat' (Reineck 1967; (Moore 1964; Heckel 1977; Ross & Ross 1987;
Weimer et al. 1981; Mcllroy 2004). Tidal flats Watney et al. 1989). The development of exten-
are dissected by genetically related meandering sive Carboniferous-Permiantide-influenced
tidal creeks. Intertidal areas represent rigorous coastlines resulted in the formation of tidal-flat
environments, where marine organisms experi- areas that are preserved within equatorial carbo-
ence extreme changes in temperature, duration nate/siliciclastic cyclothems (Mangano et al.
of subaerial exposure, salinity, hydrodynamic 2002a). Alternating sandstone and mudstone

From: MC!LROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 157-178. 0305-8719/04/S15.00 © The Geological Society
of London.
158 M. G. MANGANO & L. A. BUATOIS

Fig. 1. Distribution of outcrops of the Marmaton, Kansas City, Pleasanton, Lansing, Douglas, and Shawnee
Groups and the localities studied. Localities are numbered as follows: 1, Bandera Quarry (Bourbon County);
2, quarry 1 mi southeast of Eudora, near Coleman Creek (Douglas County); 3, road-cut along the Kansas
Turnpike US Highway 70, 1 km west of the intersection with Kansas Highway 7 (Wyandotte County);
4, road-cut along Kansas Highway 24, 1.5 km west of the intersection with Kansas Highway 7 (Leavenworth
County); 5, site south of the Kansas City International Airport (Platte County, Missouri); 6, road-cuts on
Kansas Highway 166, 3km west of Peru (Chautauqua County); 7, Toronto Lake (Woodson County);
8, Buildex Quarry (Franklin County); 9, Lonestar spillway (Douglas County); 10, Waverly trace fossil locality,
7 km west of the town of Waverly (Coffey County); 11, road-cut along Country Road 6, 4 km south of Stull
(Douglas County); 12, road-cut 1 km west of Kanwaka (Douglas County); 13, outcrop east of Perry Lake,
2.5km northeast of Lakewood Hills (Jefferson County).

layers in these intertidal deposits preserve a vari- Carboniferous tidal flats formed in open-
ety of biogenic structures that provide valuable marine shorelines with ichnofaunas of restricted
insights into the palaeoecological and deposi- bays and the fluvio-estuarine transition.
tional dynamics of these ancient shorelines.
Carboniferous intertidal deposits in eastern
Kansas and western Missouri (Fig. 1) accumu- An integrated sedimentological and
lated under palaeoenvironmental conditions ichnological model of tide-influenced tidal flat
that ranged from open shoreline to the fluvio- and subtidal sandbar complexes
estuarine transition. The intertidal deposits ana-
lysed in this paper occur in several Carboniferous Our present knowledge of the ichnology of tide-
units of eastern Kansas and western Missouri influenced shallow-marine environments lags
(Fig. 2). Comparative analysis of these exposures behind that of wave-dominated settings. Integra-
illustrates lateral variations in trace-fossil con- tion of ichnological and sedimentological obser-
tent, and allows characterization of intertidal vations has led to the establishment of the
ichnofaunas that are developed in three sub- 'shoreface model' (MacEachern & Pemberton
environments: open marine, restricted bay, and 1992; MacEachern et al 1999; Pemberton et al.
the fluvio-estuarine transition. The aims of this 2001). Although a comparable model for tide-
paper are: (1) to propose an integrated ichno- influenced shorelines is still lacking, we have out-
logical and sedimentological model of intertidal lined in previous contributions some of the most
flat and subtidal sandbar complexes and (2) to relevant characteristics of ichnofaunas from tidal
characterize and compare the ichnofaunas of flat and subtidal sandbar complexes (Mangano
CARBONIFEROUS BRACKISH-WATER ICHNOFAUNAS 159

Fig. 2. Pennsylvanian stratigraphy of Kansas and western Missouri, outlining the lithostratigraphic units
analysed. Asterisks indicate the stratigraphic position of the different ichnofaunas. Stratigraphy based on
Moore et al. (1951).
160 M. G. MANGANO & L. A. BUATOIS

Well sorted, fine-grained sandstones with large-scale


cross stratification (eolian dunes). Dominance of dwelling
and locomotion traces of invertebrates (crabs and insects) and
vertebrates. Low ichnodiversity.

Intensely rooted mudstones and dwelling traces of invertebrates


(crabs).
Massive or parallel laminated mudstones and rare sandstone
interbeds. Lenticular bedding. Inclined heterolithic stratification in
small to medium-sized tidal creeks. Dominance of horizontal, poorly
defined, feeding traces of deposit feeders. Low ichnodiversity.
Mudstones and very fine-grained sandstones with wavy and flaser
bedding. Inclined heterolithic stratification in medium-sized tidal
creeks. Dominance of horizontal feeding traces of deposit feeders,
detritus feeders and grazers; great variety of ethological categories
and moderate ichnodiversity.
High variability depending on tidal regime. Current ripple cross-
laminated fine- to very fine-grained sandstones with mudstone
partings in micro- to mesotidal settings. Medium to fine-grained
sandstones with planar, trough and herringbone cross-bedding and
upper-flow parallel lamination in meso- to macrotidal regimes.
Inclined heterolithic stratification in large-sized tidal creeks.
Dominance of horizontal feeding traces of deposit feeders, detritus
feeders and grazers, great variety of ethological categories and high
to moderate ichnodiversity in micro- to mesotidal sand flats. Low-
ichnodiversity assemblages of vertical dwelling and escape traces of
passive predators and suspension feeders in macrotidal sandflats.
Medium to fine-grained sandstones with planar, trough and
herringbone cross-bedding. Dominance of vertical dwelling and
escape traces of passive predators and suspension feeders along
reactivation surfaces. Low ichnodiversity.

Fig. 3. Integrated sedimentological and ichnological model of intertidal flats and subtidal sandbar complexes
(modified from Buatois & Mangano 2000).

& Buatois 1999; Buatois & Mangano 2000; (sand and mud) flat, and an upper-intertidal
Buatois et al. 2002a; Mangano et al 2002a). In mudflat. Landward of the mudflat, supratidal
this section we shall expand these observations salt marshes may be present, and the subtidal
to propose an integrated ichnological and sedi- zone occurs seawrard of the sandflat. Tide-
mentological model for some tide-influenced dominated parasequences are, consequently,
environments (Fig. 3). This model is based not upward fining (Van Wagoner et al. 1990).
only on observations in the Late Palaeozoic of The supratidal area may be vegetated, forming
the North American Midcontinent, but also in salt marshes where sedimentary fabric is
a wide spectrum of tidal deposits in Argentina, obliterated by root traces. Animal traces include
ranging in age from the Cambrian to the Mio- representatives of the Psilonichnus ichnofacies
cene. (Frey & Pemberton 1987). Supratidal marshes
Based on the integration of observations from grade landwards into a wide variety of terrestrial
modern and ancient deposits, Klein (1971, 1977) environments characterized by different trace-
proposed a facies model for tide-dominated fossil assemblages that are mostly included
shorelines that includes a supratidal region, the in the Scoyenia and Coprinisphaera ichno-
upper-, middle- and lower-intertidal zones, and facies (Buatois & Mangano 1995; Genise et al.
the subtidal area (see summary in Dalrymple 2000).
1992). Because tidal energy increases seaward, The upper zones of the tidal flat, referred to as
tidal flats generally coarsen seaward, in contrast the mudflat, are dominated by deposition of fine-
to wave-dominated shorelines, which coarsen grained suspended sediment. Mudflats consist
landward. Therefore a typical tidal-flat profile of laminated mudstone with rare siltstone and
in a landward direction consists of a lower- sandstone interbeds and interlaminae. Lenticular
intertidal sandflat, a middle-intertidal mixed bedding is the dominant bedding style. Scarcity of
CARBONIFEROUS BRACKISH-WATER ICHNOFAUNAS 161

sandstone layers limits trace fossil preservation in Weimer et al. 1981; de Mowbray 1983; Thomas
mudflat deposits, and an indistinct, mottled et al. 1987; Dalrymple 1992). Inclined hetero-
texture is common. Interface trace fossils of the lithic stratification is the typical structure
Cruziana ichnofacies can, however, be preserved formed by this tidally influenced lateral accretion
in the rare sandstone intercalations (e.g. Man- (Thomas et al. 1987). In the upper muddy zones
gano et al. 2002a). Middle-intertidal areas of the tidal flats channels are small to medium
(mixed flat) are typified by sedimentation from size, but in the lower sandy areas they tend to
traction alternating with fallout from suspension. coalesce, forming wider, deeper channels (Dal-
Heterolithic bedding is typical, mostly repre- rymple 1992). The degree of bioturbation is
sented by flaser and wavy bedding. Elements of lower in the point bars than in tidal flats, most
the Cruziana ichnofacies are characteristic of likely reflecting higher rates of sedimentation
the mixed flat. Alternation of sandstone and along unstable channel margins (cf. Gingras
mudstone layers enhances preservation of hori- et al. 1999; Mangano et al. 2002a).
zontal interface traces, such as those that typify The subtidal zone is characterized by maxi-
the Cruziana ichnofacies. The lower zones of mum energy with high current velocities
the tidal flat, referred to as the sandflat, are (Dalrymple 1992). As a result, large-scale bed-
characterized by bedload traction of sand-sized forms, such as dunes, migrate across the subtidal
sediment. As is the case for the lower shoreface areas, forming sandbars (compound dunes) and
in wave-dominated shorelines (MacEachern & ridges. Wilson (1982, 1986) noted that, in
Pemberton 1992), the sandflat is the most variable modern subtidal regions, few benthic species
intertidal zone in terms of both sedimentary are able to survive in zones of actively migrating
facies and trace fossil content. Whereas the bedforms. Accordingly, faunal diversity
character of facies in the lower shoreface varies, increases toward areas with smaller bedforms
depending on the intensity and frequency of and in the outer regions where dunes are replaced
storms (see MacEachern & Pemberton 1992; by small ripples and interbeds and interlaminae
Pemberton et al. 2001), that of the lower tidal of mud. Studies of marine benthic ecology also
flat is controlled mainly by tidal range (Buatois demonstrate that suspension feeders are the
et al. 2002a). In a macro tidal regime, character- dominant trophic type in high-energy subtidal
ized by high current speeds, migration of large- environments (Wilson 1982). As in the case of
scale bedforms, such as two-dimensional and macrotidal sandflats, vertical trace fossils of the
three-dimensional dunes (Dalrymple 1992), pro- Skolithos ichnofacies are the dominant
duces thick cross-bedsets. Macrotidal estuaries elements, commonly extending down into the
may display upper-flow regime horizontal sediment from reactivation surfaces (e.g. Pollard
planar parallel lamination and rare current rip- et al. 1993). Subtidal sandbar sands grade
ples (Dalrymple et al 1990; Dalrymple 1992). seaward into outer shelf muds, commonly char-
As a consequence, the lower intertidal zone of acterized by the Zoophycos ichnofacies.
macrotidal shorelines is very difficult to distin- To summarize, the ichnofacies gradient in tide-
guish from sub tidal areas. High-energy, rapidly influenced shorelines is opposite to that in wave-
migrating bedforms generally preclude the estab- dominated shoreface environments. As overall
lishment of a mobile epifaunal and/or shallow tidal energy increases from supratidal to subtidal
infaunal biota, inhibiting development of the settings, the Skolithos ichnofacies tends to occur
Cruziana ichnofacies. Bioturbation is restricted seaward of the Cruziana ichnofacies (Mangano
to vertical elements of the Skolithos ichnofacies, et al 2002a) (Fig. 3). This shoreward decrease
typically reflecting colonization windows along of energy parallels a decrease in oxygenation,
reactivation surfaces (Pollard et al. 1993; Mac- sand content, amount of organic particles in
Eachern & Pemberton 1994; Mangano et al. suspension, and mobility of the substrate. This
1996). Conversely, in microtidal to mesotidal gradient is consistent with information from
regimes, migrating bedforms are commonly modern tide-influenced environments, where
represented by small current ripples - producing the highest faunal diversity is present around
cross-lamination - under low-energy conditions. mid-tide level (Beukema 1976). Modern ana-
High-diversity assemblages of the Cruziana logues of the Cruziana ichnofacies are common
ichnofacies are characteristic of this type of in tidal flat deposits (e.g. Bajard 1966; Howard
sandflat. & Dorjes 1972; Swinbanks & Murray 1981;
Tidal flat deposits are typically dissected by a Ghare & Badve 1984; Frey et al 1987; Gingras
network of meandering tidal channels and et al 1999). The preservational potential of
creeks that migrate across the intertidal zone, biogenic structures in intertidal settings is, how-
producing lateral accretion of bedsets in point ever, highly variable, with a clear bias in the
bars (Reineck 1958; Bridges & Leeder 1976; fossil record towards deeper-tier structures,
162 M. G. MANGANO & L. A. BUATOIS

particularly in post-Palaeozoic examples (see the slip face of duneforms. These sandstone
Mangano et al. 2002a for discussion). deposits are interpreted as a subtidal sandbar
complex that probably was formed in the lower
zone of a tide-dominated estuary.
Stratigraphic and depositional setting Evidence of estuarine incision at the base of
the Rock Lake Shale Member was presented by
Intertidal ichnofaunas analysed in this paper were Feldman et al. (1993), who analysed the Garnett
recorded from exposures of the Desmoinesian palaeovalley from outcrops in east-central
Bandera Shale Formation (Marmaton Group), Kansas. The valley is incised into the underlying
the Missourian Rock Lake Shale Member (Stan- Stanton Limestone and Vilas Shale, and is
ton Formation, Lansing Group), the Virgilian oriented northwest to southeast. They estimated
Stranger Formation (Douglas Group), the Virgi- the estuarine valley to be 9.8m deep and 300m
lian Lawrence Shale Formation (Douglas wide. Outcrops in northeast Kansas and north-
Group), and the Virgilian Stull Shale Member west Missouri also include tidal flat deposits
(Kanwaka Shale Formation, Shawnee Group). that were formed along a brackish-water,
These units outcrop in narrow belts across east restricted embayment (Russell 1972, 1974;
Kansas and west Missouri (Fig. 1). The analysed Hakes 1976, 1977, 1985; Mangano et al. 1999).
siliciclastic intervals are separated by laterally con- The Douglas Group represents deposition
tinuous transgressive limestone units (e.g. Haskell within two estuarine valleys, namely the Tonga-
Limestone, Toronto Limestone, Spring Branch noxie and Ireland palaeovalleys (Archer et al.
Limestone) that serve as marker beds. This situa- 1994a). The Tonganoxie sequence records the
tion allows spatio-temporal comparisons to be infill of a northeast to southwest oriented estuar-
effected with a degree of accuracy that typically ine valley, incised during the latest Missourian
is otherwise possible only in modern environ- sea-level fall and filled during the subsequent
ments. These deposits, however, contain biogenic transgression in the earliest Virgilian (Lanier
structures that have already passed through the 1993; Lanier et al. 1993; Gibling et al. 1993;
taphonomic filter - that is, the fossilization barrier Archer et al. 1994a; Feldman et al. 1995; Buatois
- and are thus better analogues to other ancient et al. 1998a). The Tonganoxie palaeovalley is
tide-influenced depositional systems. Integration incised into the underlying Weston Shale as
of sedimentological, sequence-stratigraphic, well as units of the Lansing Group and is about
palaeoecological, and ichnological evidence with 41 m deep, 11 km wide, and 240km long (Feld-
palaeogeographic reconstructions indicates that man et al. 1995). Palaeocurrent studies indicate
these units record, for the most part, deposition the existence of southwesterly flowing rivers
in embayments and estuaries, opening to the sea and ebb-dominated tidal dunes (Gibling et al.
towards the south (Archer et al. 1994a; Mangano 1993). Northwesterly palaeocurrents are evident
et al. 2002a). Tidal flats were formed in different higher in the sequence and reveal a reversal in
geomorphological positions, such as fluvio-estuar- sediment transport, reflecting the establishment
ine transitions, restricted bays and open-marine of a flood-dominated tidal system during the sub-
environments. sequent transgression (Gibling et al. 1993). The
Outcrop and subsurface data indicate that the Ireland palaeovalley is incised into the Stranger
Bandera Shale Formation was deposited in tide- Formation; although its morphology is not well
dominated embayments (Brownfield et al. 1998). known, the overall succession appears similar
These authors suggested that sand was trans- to that of the Tonganoxie palaeovalley (Archer
ported to the basin by fluvial systems incised et al. 1994a). Tidal flat deposits recorded from
during a sea-level fall and reworked by marine these units accumulated in different geomorpho-
processes during the subsequent transgression. logical settings. The succession of the Tonga-
The Bandera Shale ichnofauna is recorded from noxie Sandstone Member at Buildex Quarry
Bandera Quarry (Bourbon County, Southeast (Franklin County, eastern Kansas) consists, for
Kansas). The succession at this locality consists the most part, of planar-laminated coarse-
of interbedded sandstone and shale and large- grained siltstone deposited on an upper tidal
scale cross-stratified sandstone (Brownfield et al. flat, close to or at the fluvial-estuarine transition
1998). The sandstone and shale are rhythmically of a macrotidal estuarine palaeovalley (Lanier
interbedded and display flaser and wavy bedding, et al. 1993; Buatois et al. 1997). Outcrops of the
mudstone rip-up clasts and flat-topped ripples, Lawrence Formation at Lonestar spillway
suggesting tidal processes in an intertidal envir- (Douglas County, Northeast Kansas), on the
onment. The large-scale cross-stratified sand- other hand, consist of mudstone and very fine-
stone has sinuous crested dunes preserved at grained sandstone displaying flaser, wavy and
the top. Current ripples are locally preserved on lenticular bedding, and record deposition in the
CARBONIFEROUS BRACKISH-WATER ICHNOFAUNAS 163

intertidal to subtidal zones of the middle estuary 2002a). Further to the southwest in Coffey
(Archer et al, 1994a). Outcrops of the Ireland County, tidal flats were formed outside the
Sandstone exposed in Toronto Lake (Woodson embayment in direct connection with the open
County, East Kansas) consist of laterally persis- sea (Mangano et al. 2002a).
tent, parallel-laminated sandstone that probably
formed in upper-flow regime sandflats of the
lower estuary (Archer et al. 1994a). Finally, a Ichnofaunas and tidal flat variability
succession of the 'Stalnaker' Sandstone exposed
in road-cuts on Kansas Highway 166, 3km Comparative analysis of these Carboniferous
west of Peru (Chautauqua County, Southeast exposures illustrates lateral variations in trace-
Kansas), consists of fine-grained sandstone and fossil contents and allows characterization of
mudstone with abundant trace fossils and body three different types of intertidal ichnofauna,
fossils. These deposits are thought to have been which correspond to three different sub-environ-
formed in tidal flats outside the estuarine embay- ments: open marine, restricted bays, and fluvio-
ment, directly connected with the open sea estuarine transitions. Although our study is
(Archer et al. 1994a; Mangano & Buatois 1997). based on outcrops in Kansas and Missouri, we
The Stull Shale Member exhibits lateral provide comparisons to show that similar ichno-
changes of facies from restricted deposits faunas are recognized in other late Palaeozoic
formed in an embayment in northeastern areas of the American Midcontinent and the
Kansas to open-marine intervals to the south- Appalachians.
west (Mangano & Buatois 1997; Mangano et al.
2002a). Ichnological and sedimentological
studies of outcrops of the Stull Shale in Douglas Open-marine intertidal ichnofaunas
and Jefferson Counties indicate marginal-
marine, brackish-water conditions (Hakes 1976, Tidal flats may form on open coasts outside
1985; Mangano & Buatois 1997; Mangano et al. embayments. Examples of ichnofaunas from

Fig. 4. Physical sedimentary structures of tidal-flat deposits as seen in bedding-plane views, (a) Flat-topped
ripples. Stull Shale. Waverly trace-fossil site. Lens cap is 5.5cm in diameter, (b) Relict troughs and wrinkle
marks. Stull Shale. Waverly trace-fossil site. Lens cap is 5.5cm in diameter, (c) Ladderback ripples and
flat-topped ripples. Note associated Gyrochorte isp. Stalnaker Sandstone. Road-cut on Kansas Highway 166.
Coin is 2.4cm across, (d) Dendritic runnel marks. Tonganoxie Sandstone. Buildex Quarry. Bar= 1 cm.
(e) Foam marks. Tonganoxie Sandstone. Buildex Quarry. Bar= 1 cm. (f) Raindrop impact structures on an
otherwise rippled surface. Tonganoxie Sandstone. Buildex Quarry. Bar — 1 cm.
164 M. G. MANGANO & L. A. BUATOIS

such complexes have been analysed from two body fossils (e.g. bivalves, brachiopods, crinoids)
main outcrops in Kansas: the Waverly trace- are also common.
fossil site in Coffey County (Stull Shale Ichnofaunas present in these tidal flat deposits
Member), and road-cuts on Kansas Highway in are extremely rich, including resting traces
Chautauqua County ('Stalnaker' Sandstone). (Asteriacites lumbricalis, Lockeia ornata, Lockeia
The former was studied in detail by Mangano siliquaria, Rusophycus isp.), locomotion traces
et al. (2002a), and the latter was briefly referred (Cruziana problematica, Cruziana isp., Curvo-
to in a number of publications (e.g. Archer lithus simplex, Curvolithus multiplex, Gyrochorte
1993; Archer et al. 1994b; Mangano & Buatois isp., Protovirgularia rugosa), grazing traces
1997; Mangano et al. 2002b). (Nereites cambriensis, Nereites imbricata, Ner-
These interbedded fine- to very fine-grained eites jacksoni, Nereites missouriensis, Planolites
sandstones and siltstones contain a wealth of beverleyensis, Psammichnites grumula, Psam-
physical and biogenic structures. Physical michnites implexus, Psammichnites plummeri),
sedimentary structures include asymmetric to feeding traces (Chondrites isp., Halopoa isp.,
symmetric ripples, flat-topped ripples (Fig. 4a), Parahaentzschelinia ardelia, Phycodes palmatus,
wrinkle marks (Fig. 4b), relict troughs (Fig. Phycodes isp., Phycosiphon incertum, Rhizocoral-
4b), ladder-back ripples (Fig. 4c), interference lium irregulare, Teichichnus rectus, Trichophycus
ripples, gutter casts, flute marks, load casts, and isp.) and dwelling traces (Arenicolites isp., Con-
sand volcanoes. Flaser, wavy and lenticular ichnus conicus, Diplo crater ion isp., Palaeophycus
bedding are the dominant bedding styles. The tubularis, Pentichnus pratti, Rosselia socialis,
association clearly indicates deposition in the Skolithos isp., Solemyatuba isp.). Associated
intertidal zone. There is significant evidence for orthomyalinid bivalve shells have been exten-
wave influence in these open-marine tidal sively affected by bioerosion, displaying borings
facies. Facies analysis suggests sedimentation in by acrothoracican barnacles, polydorid worms
sandflats, mixed flats, and mudflats, dissected (ichnogenus Caulostrepsis), and ctenostomatid
by intertidal channels (Mangano et al. 2002a). bryozoans (Baker 1995).
Trace fossils are particularly abundant in the Open-marine tidal flat ichnofaunas are charac-
sandflat deposits (Figs 5, 6 and 7). Marine terized by:

Fig. 5. Idealized reconstruction of open-marine tidal-flat ichnofaunas showing the most typical ichnotaxa.
CARBONIFEROUS BRACKISH-WATER ICHNOFAUNAS 165

Fig. 6. Base of a sandstone bed showing typical occurrence of intertidal ichnofaunas in the open-marine
setting. Surfaces are totally covered by trace fossils, and ichnodiversity is very high. Cs, Curvolithus simplex;
Cm, Curvolithus multiplex; Ls, Lockeia siliquaria; D, Diplo crater ion isp.; Al, Asteriacites lumbricalis; Cp,
Cruziana problematical Pb, Protovirgularia bidirectionalis. Note subparallel orientation of specimens of
Protovirgularia bidirectionalis. Stull Shale. Waverly trace-fossil site.

high ichnodiversity; Heterogeneous distribution of biogenic struc-


marine ichnotaxa produced by both euryha- tures is remarkable at both small and large
line and stenohaline organisms; scales (Mangano et al. 2002a). At the larger
the presence of both infaunal and epifaunal scale, zonational distribution is expressed along
traces; the entire tidal range; on a smaller scale (i.e.
the presence of simple and complex structures within each sub-environment), spatial segrega-
produced by presumed trophic generalists and tion of species may reflect distinct microhabitats
specialists respectively; and partitioning of energy resources. Detailed
dominance of horizontal trace fossils of the analysis of bivalve trace fossils at the Waverly
Cruziana ichnofacies; site by Mangano et al. (1998) reveals the exis-
the presence of multispecific associations; tence of palimpsest surfaces. These surfaces
high density; were formed by a succession of colonization
wide size ranges. events, including an initial colonization event,
erosive scouring and subsequent sedimentation
Most trace fossils are interface traces, and thus and eventual re-colonization. This succession of
the degree of bioturbation is relatively low. events produced time-averaged surfaces that
Bedding planes are, however, generally covered record the work of several communities (ichno-
by trace fossils, forming multispecific assem- coenoses) of burrowing bivalves. The presence
blages (Fig. 6). Dominant trace-makers are of time-averaged associations may have artifi-
attributed to bivalves, ophiuroids and annelids. cially exaggerated ichnodiversity; the work of
166 M. G. MANGANO & L. A. BUATOIS

Fig. 7. Selected trace fossils from open-marine intertidal deposits, (a) Asteriacites lumbricalis, showing
horizontal repetition. Base of sandstone bed. Stull Shale. Waverly trace-fossil site. Coin is 1.4cm across,
(b) Protovirgularia bidirectionalis, displaying V-shaped markings with opposite directions meeting at a central
point. Note that the direction of movement is from the centre to the ends. Base of sandstone bed. Stull Shale.
Waverly trace-fossil site. Bar = 1 cm. (c) Nereites missouriensis with well-developed lateral lobes. Top of
sandstone bed. Stull Shale. Waverly trace-fossil site. Bar= 1 cm. (d) Rusophycus isp. (R) and Cruziana isp. (C)
preserved on the base of a sandstone bed. Stalnaker Sandstone. Roadcut along Kansas Highway 166.
Bar = 1 cm. (e) Specimen of Lockeia ornata connected with chevron locomotion traces (Protovirgularia rugosa).
Chevron orientation indicates that the animal exited the resting structure. Base of sandstone bed. Stull Shale.
Waverly trace-fossil site, (f) Curvolithus simplex preserved on the top of a sandstone bed. Stull Shale. Waverly
trace-fossil site. Lens cap is 5.5cm in diameter, (g) Solemyatuba isp. Cross-section view. Stalnaker Sandstone.
Roadcut along Kansas Highway 166. Bar = 1 cm. (h) Psammichnites implexus preserved on the top of a rippled
sandstone and showing guided meanders on ripple troughs. Stull Shale. Waverly trace-fossil site. Bar= 1 cm.
(i) Psammichnites grumula with well-developed holes along a median line and prominent levees on both sides of
the trace. Base of sandstone bed. Stull Shale. Waverly trace-fossil site. Bar = 1 cm.

successive communities may be expressed at a inhabiting open-marine intertidal areas experi-


single bedding plane. ence less stress than those developed in estuaries
These ichnofaunas reflect the activity of biotas and brackish bays, characterized by steep
that inhabited tidal flats dominated by normal or salinity gradients combined with fluctuating
near-normal marine salinities. Benthic faunas temperature, oxygen and turbidity, among
CARBONIFEROUS BRACKISH-WATER ICHNOFAUNAS 167

other variables (Wightman et al. 1987; Pember- Lower Pennnsylvanian Fentress Formation in
ton & Wightman 1992; MacEachern & Pember- eastern Tennessee (Miller & Knox 1985), the
ton 1994; Pemberton et al. 2001). This fact is Lower Pennsylvanian Caseyville and Tradewater
reflected by the high diversity of the open- formations of southern Illinois (Devera 1989),
marine tidal flat ichnofaunas and by the presence the Middle Pennsylvanian Kanawha Formation
of a Cruziana ichnofacies. in West Virginia (Martino 1989, 1996), and the
Upper Pennsylvanian Tecumseh Shale Member
in Kansas (Hakes 1976). Similar ichnofaunas
Restricted-bay intertidal ichnofaunas have been documented in the United Kingdom,
but have been placed within a deltaic context
Restricted-bay intertidal deposits accumulate in (Eagar et al. 1985; Pollard 1988; Buckman 1992).
incised valleys, lagoons and embayments. Exam- The assemblages occur in heterolithic facies
ples of late Palaeozoic ichnofaunas from consisting of very fine-grained sandstone and
restricted-bay environments are widespread in mudstone displaying flaser, wavy and lenticular
Kansas and Missouri, and include those bedding. Flat-topped ripples and wrinkle marks
recorded in the Bandera Quarry (Bourbon are relatively common. Beds are commonly
County, Bandera Shale), several outcrops of the stacked, forming fining-upward parasequences
Rock Lake Shale Member in northwest Missouri that reflect tidal-flat progradation. Sand- and
and northeast Kansas (Leavenworth and Wyan- mud-filled intertidal channels typically cut the
dotte Counties; see Mangano et al. 1999), succes- tidal-flat deposits. Coal beds are present in
sions of the Douglas Group in Lonestar spillway most outcrops. In contrast to tidal flats from
(Douglas County) and Toronto Lake (Woodson open shorelines, there is less evidence for wave
County), and several outcrops of the Stull Shale influence in these restricted tidal facies. Marine
Member in Douglas and Jefferson Counties (see body fossils are notably absent. In fact, some of
Mangano et al. 2002a). Some of these ichnofau- these deposits were originally thought to have
nas have been discussed by Bandel (1967b), formed in continental environments, but ichno-
Hakes (1976, 1977, 1985), Mangano & Buatois logical evidence (i.e. the presence of marine ich-
(1997) and Mangano et al. (1999, 2002a). Other notaxa such as Asteriacites and Psammichnites)
examples of Carboniferous restricted-bay ichno- indicates marine influence (Hakes 1976, 1977,
faunas in the United States include those 1985).
reported from the Upper Mississippian Hartselle Ichnofaunas from restricted-bay intertidal
Sandstone of Alabama (Rindsberg 1994), the deposits are not diverse (Figs 8, 9). With the

Fig. 8. Idealized reconstruction of restricted-bay, tidal-flat ichnofaunas showing the most typical ichnotaxa.
168 M. G. MANGANO & L. A. BUATOIS

Fig. 9. Selected trace fossils from restricted-bay intertidal deposits. All bars = 1 cm. (a) Psammichnites
plummeri. Base of sandstone bed. Ireland Sandstone. Toronto Lake, (b) Lingulichnus isp. Base of sandstone
bed. Rock Lake Shale. Quarry near Coleman Creek, SE of Eudora. (c) Gyrochorte isp. (arrow). Top of
sandstone bed. Bandera Shale. Bandera Quarry, (d) Teichichnus rectus. Stull Shale. Roadcut along Country
Road 6, south of Stull (locality 8 of Hakes 1976). (e) Asteriacites lumbricalis with V-shaped striations
ornamenting the arms. Note the presence of two partially preserved, shallower impressions toward the upper
left documenting vertical repetition. The escape strategy is also recorded by an elite bivalve trace (upper left).
Base of sandstone bed. Rock Lake Shale. Site south of the Kansas City International Airport, (f) Dense
assemblage of small specimens of Lockeia siliquaria. Base of sandstone bed. Rock Lake Shale. Quarry near
Coleman Creek, SE of Eudora. (g) Palaeophycus tubularis (Pt) and small Protovirgularia (P). Top of sandstone
bed. Stull Shale Member. Roadcut west of Kanwaka (locality 5 of Hakes 1976).

exception of Lingulichnus, no ichnotaxa were lumbricalis, Lockeia siliquarid), locomotion


found to occur exclusively in these deposits; traces (Gyrochorte isp., Protovirgularia rugosa),
several components of the open-marine assem- grazing traces (Nereites isp., Psammichnites
blage, including resting traces (Asteriacites implexus, Psammichnites plummeri), feeding
CARBONIFEROUS BRACKISH-WATER ICHNOFAUNAS 169

traces (Teichichnus rectus) and dwelling traces conditions, brackish-water faunas are less diverse
(Diplo crater ion isp., Lingulichnus isp., Palaeo- than their marine and freshwater equivalents
phycus tubular-is), were present. (e.g. Croghan 1983; Hudson 1990; Pickerill &
These ichnofaunas display: sdgfsdfgsdfgsdfgsdfgafgsdfgsdfg
should not be equated with biodiversity (see
low ichnodiversity;
sdfgsdfgsdfgsdfgsdfgsdfgsdfgfdgfsdfgdfgsdfddddd
ichnotaxa commonly found in marine environ-
may provide some rough information on general
ments, but produced by presumed euryhaline
trends in species richness if used with caution.
organisms;
Notably, the reduction in faunal diversity experi-
a dominance of infaunal traces rather than
enced by brackish-water faunas is reflected by a
epifaunal trails;
parallel decrease in ichnodiversity.
simple structures attributed to opportunistic
Restricted-bay ichnofaunas record the activity
trophic generalists;
of a brackish-water benthos inhabiting intertidal
a combination of vertical and horizontal trace
areas of estuarine basins and embayments. As
fossils from the Skolithos and Cruziana ichno-
such, they essentially display the diagnostic fea-
facies;
tures of brackish-water trace-fossil assemblages,
monospecific associations;
as documented by Wightman et al. (1987), Pem-
variable abundances;
berton & Wightman (1992), MacEachern &
small sizes.
Pemberton (1994) and Pemberton et al. (2001).
The abundance of trace fossils in the restricted Minor differences were discussed by Buatois
bay deposits is highly variable. The presence of et al. (2002b), who suggested that Pennsylvanian
thick packages of restricted bay deposits with estuarine ichnofaunas differ from their post-
only a few trace fossils is not unusual. However, Palaeozoic counterparts in having lower ichno-
high-density assemblages may be common diversity, a lower degree of bioturbation, scarce
locally as well. Monospecific suites of Psammich- crustacean burrows, apparent lack of firmground
nites (Fig. 9a) or Lingulichnus (Fig. 9b) are very suites, and an absence of specialized morpho-
common. Association of bivalve (Lockeia-Proto- logic adaptations designed to protect organisms
virgularid) and ophiuroid (Asteriacites) trace from salinity fluctuations. Interestingly, fresh-
fossils are very common as well (Fig. 9e). In con- water organisms tend to disappear rapidly, even
trast to the conventional view of Asteriacites as a with slight increases in salinity, whereas marine
normal-marine salinity indicator, Mangano et al. organisms experience a more gradual decrease
(1999) demonstrated that this ichnotaxon is in number under dilution of normal-marine
widespread in brackish-water facies of the salinity (Pemberton & Wightman 1992). Accord-
American midcontinent. This is consistent with ingly, brackish-water ichnofaunas typically
evidence from modern environments, which resemble impoverished marine ichnofaunas
shows that some echinoderms, and particularly rather than a mixture of freshwater and marine
asterozoans, are able to inhabit brackish-water ichnotaxa (Pemberton & Wightman 1992).
environments (e.g. Binyon 1972; Stancyk 1973;
Turner 1980; Pagett 1980, 1981). It has been
noted that reduced size is one of the most notable Fluvio-estuarine intertidal ichnofaunas
features of brackish-water assemblages (Hakes
1976, 1985). This is in agreement with studies In estuaries, tidal influence commonly extends
of marine benthic ecology, which documented further landward than the saltwater intrusion,
reduced size in brackish-water faunas, particu- particularly in macrotidal systems (Fairbridge
larly ophiuroids, bivalves and worms (Remane 1980; Allen 1991; Dalrymple et al. 1992; Buatois
& Schlieper 1971; Spaargaren 1979, 1995; et al. 1997). Accordingly, tidal flats may develop
Gingras et al. 1999). In contrast, crustaceans in the uppermost zone of an estuary, under fresh-
usually produce large burrows in brackish- water conditions. Ecological conditions in these
water environments (Gingras et al. 1999). It has fluvio-estuarine transitional settings are remark-
further been postulated that size reduction in ably different from those in tidal flats formed
response to salinity occurs either as a morpho- along embayment margins or in direct connec-
logic adaptation or as a result of population tion with the open sea (Buatois et al. 1997).
dynamics (Gingras et al. 1999). In the first case, Our characterization of fluvio-estuarine inter-
decreasing size allows the organism to increase tidal ichnofaunas is based on outcrops and
its surface area to mass ratio to control osmotic cores of the Tonganoxie Sandstone Member at
transfer. In the second case, large populations the Buildex Quarry (Franklin County, Kansas).
of small forms that attain full growth result in The Buildex ichnofauna was analysed in detail
the same biomass. As a consequence of stressful in a series of papers (Bandel 1967a; Buatois
170 M. G. MANGANO & L. A. BUATOIS

et al 1997, 1998a, 1998b, 1998c; Mangano et al. 1993). Facies are stacked in symmetrical cycles
1997). Other examples of virtually identical that each display a gradual increase and sub-
ichnofaunas in similar depositional settings are sequent decrease in bed thickness. Climbing
present in the Lower Pennsylvanian Whetstone ripples, mudstone drapes and soft-sediment
Beds, Mansfield Formation, in Indiana (Archer deformation structures, such as convolute lami-
& Maples 1984, Archer 1993; Archer et al nation and micro-faults, are common. Bedding-
1994b), the Middle Pennsylvanian Pottsville surface structures are abundant, including tool
Formation of Alabama (Cincosaurus and Haplo- marks, drainage or seepage rill marks, runnel
tichnus assemblages of Rindsberg 1990), and the marks (Fig. 4d), runoff washouts, foam marks
Middle Pennsylvanian Crane succession of the (Fig. 4e), raindrop impressions (Fig. 4f), falling-
Mansfield Formation in Indiana (Kvale & Barn- water marks, and wrinkle marks. Preservation
hill 1994; Mangano et al. 2001). of autochthonous upright plants is common.
At the Buildex Quarry, the Tonganoxie Sand- Recurrent thickness fluctuations indicate that
stone Member overlies the Ottawa Coal and con- these deposits must be regarded as tidal rhyth-
sists of very fine-grained sandstone, siltstone, and mites, with thicker strata representing deposition
mudstone. Outcrop information was integrated near spring tide and thinner ones recording con-
with subsurface data derived from 10 drilled ditions during neap tide periods (Lanier et al.
cores recovered from the quarry. The deposits 1993). Some of the bedding-surface structures
consist of laterally persistent, normally graded (e.g. raindrop impressions, rill marks, runnel
and parallel-laminated siltstone beds that are marks) indicate brief periods of subaerial expo-
replaced upward by a channelized siltstone sure. The Buildex Quarry succession is inter-
body and a thicker-bedded, planar-stratified preted to have been deposited in tidal flats close
siltstone that is capped by a thin, pervasively to or at the fluvio-estuarine transition of a
rooted, silt-rich coal (Lanier 1993; Lanier et al. macrotidal estuarine valley (Lanier et al. 1993;

Fig. 10. Idealized reconstruction of fluvio-estuarine tidal-flat ichnofaunas showing the most typical ichnotaxa.
CARBONIFEROUS BRACKISH-WATER ICHNOFAUNAS 171

Buatois et al. 1997). Animal trace fossils are dominated by arthropod trackways (e.g. Den-
exclusively preserved on bedding planes (Figs droidichnites irregulare, Diplichnites gouldi,
10, 11); root traces are the only biogenic struc- Diplopodichnus bifurcus, Kouphichnium isp.,
tures seen in cross-section. Accordingly, the Mirandaichnium famatinense, Stiallia pilosa,
core expression of such an assemblage is one of Stiaria intermedia) and grazing traces (e.g.
parallel-laminated deposits with minimal or no Gordia indianaensis, Helminthoidichnites tenuis,
bioturbation (Buatois et al. 1998a). Helminthopsis hieroglyphica). Subsurface-feeding
Ichnofaunas from fluvio-estuarine intertidal traces (Circulichnus montanus, Treptichnus bifur-
environments are relatively diverse. They are cus, T. pollardi), apterygote insect resting and

Fig. 11. Selected trace fossils from fluvio-estuarine transition intertidal deposits. All photos are bases of
siltstone beds from the Tonganoxie Sandstone Member at the Buildex Quarry. All bars = 1 cm. (a)
Tonganoxichnus buildexensis. (b) Stiallia pilosa (Sp) and Treptichnus bifurcus (Tb). Note that the trackway is
cross-cutting Treptichnus bifurcus on the left, (c) Stiaria intermedia (Si) and Tonganoxichnus ottawensis (To), (d)
Helminthopsis hieroglyphica. (e) Gordia indianaensis cross-cut by a series of thin, long tool marks, (f) Undichna
britannica. (g) Diplichnites gouldi.
172 M. G. MANGANO & L. A. BUATOIS

Fig. 12. Relationship of species diversity, trace fossil assemblages, and salinity levels (modified from Remane &
Schlieper 1971; Wightman et al. 1987; Buatois et al. 1997). Open-marine intertidal areas display the maximum
ichnodiversity and are characterized by a Cruziana ichnofacies. The reduction in faunal diversity experienced by
brackish-water faunas is reflected by a parallel decrease in ichnodiversity, illustrated by the presence of a
depauperate Cruziana and Skolithos ichnofacies. The relatively high ichnodiversity of the fluvio-estuarine
transition may be understood as recording the secondary peak in diversity typically associated with the activity
of freshwater/terrestrial biotas along a salinity gradient (mixed Scoyenia and Mermia ichnofacies in tidal
rhythmites).

feeding traces (Tonganoxichnus buildexensis, T. gradient (Fig. 12). This is consistent with the
ottawensis), fish traces (Undichna britannica, U. taxonomic composition of these ichnofaunas,
simplicitas) and tetrapod trackways are also which consist entirely of ichnotaxa commonly,
common. though not exclusively, recorded from continen-
These ichnofaunas are characterized by: tal settings. No ichnotaxa indicative of marine
influence have been detected. Arthropods are
moderate to relatively high diversity;
the dominant trace-makers. In fact, the fluvio-
forms typically present in continental environ-
estuarine ichnofaunas seem to display a mixture
ments;
of elements of the continental Scoyenia and
the dominance of surface trails and absence of
Mermia ichnofacies (Buatois & Mangano 1995,
burrows;
1998). Undoubtedly, taphonomy played a role
temporary structures produced by mobile
in increasing the ichnodiversity. Suppressed
deposit-feeding fauna;
erosion during rising tides allowed excellent pre-
a mixture of trace fossils belonging to the
servation of delicate surficial structures (Archer
Scoyenia and Mermia ichnofacies;
et al. 1994b). Additionally, the absence of perva-
moderate density of individual ichnotaxa;
sive burrowers in such palaeoenvironments
the absence of monospecific suites;
improves the preservation potential of such sur-
small size.
face traces because such activity by infaunal
Interestingly, these ichnofaunas are in marked organisms commonly leads to the destruction
contrast to those recorded from brackish-water of the uppermost tiers (Bromley 1996; Buatois
settings. The relatively high ichnodiversity may et al 1997).
be understood as recording the secondary peak The occurrence of assemblages dominated by
in diversity typically associated with the activity trackways and trails in tidal rhythmites has
of freshwater/terrestrial biotas along a salinity puzzled researchers for some time (e.g. Archer
CARBONIFEROUS BRACKISH-WATER ICHNOFAUNAS 173

& Maples 1984; Rindsberg 1990; Archer 1993; freshwater benthos inhabiting this zone do not
Archer et al 1994b) and, accordingly, a have the special adaptations necessary to survive
number of clastic successions have been regarded in the brackish environment. Whereas fully
as either marine or non-marine. Although the marine ichnofaunas gradually decrease in diver-
presence of these continental ichnofaunas in sity into brackish-water settings, ichnofaunas
tidal rhythmites makes it difficult to reconcile from fluvio-estuarine transitions do not inter-
the sedimentological and ichnological evidence, grade with those from brackish water. This
understanding the ecological conditions existing ichnological signal reflects the fact that fresh-
in fluvio-estuarine transitions of macrotidal water organisms disappear rapidly with even
estuaries helps to solve this apparent paradox slight increases in salinity, whereas marine
(see discussion in Buatois et al. 1997). The organisms experience a more gradual decrease
assemblages documented herein occur in tidal under dilution of normal-marine salinity.
rhythmites, and record the activity of typical
freshwater/terrestrial biotas inhabiting tidal
flats developed in the most proximal zone of Stratigraphic implications
the inner estuary under freshwater conditions.
The presence of these mixed freshwater/ The ichnofaunas characterized in this study indi-
terrestrial ichnofaunas in tidal rhythmites cate not only certain depositional zones within
corresponds to the zone situated between the coastal tide-influenced environments, but also
maximum landward limit of tidal action and particular depositional stages. Incised valleys
the seaward salinity limit. Therefore recognition may begin to fill during a lowstand, but sediments
of these ichnofaunas helps to delineate fluvio- typically accumulate during the subsequent sea-
estuarine transitions with great precision. The level rise (Zaitlin et al. 1994). If lowstand,

Fig. 13. Palaeoenvironmental distribution of the intertidal trace-fossil assemblages analysed.


174 M. G. MANGANO & L. A. BUATOIS

coarse-grained sediments are deposited and constructed only on the basis of sedimentological
preserved, they are replaced vertically by finer- evidence.
grained facies of the transgressive systems tract.
Commonly, during a lowstand the valley acts as We thank Sigma Delta Epsilon, the Paleontological
a bypass zone, or lowstand deposits are eroded Society, the Mid-America Paleontological Society
and reworked during the subsequent transgres- (MAP Award) and the Antorchas Foundation for
providing financial support. Research for this paper
sion (MacEachern & Pemberton 1994). In such was undertaken during tenure at the Kansas Geological
cases, transgressive deposits directly overlie the Survey and funded by an external grant from the
sequence boundary, resulting in the formation Argentinean Research Council (CONICET) awarded
of co-planar surfaces (MacEachern et al. 1992). to both authors. We are very grateful to several
Scoyenia-Mermia intertidal ichnofaunas colleagues who supplied valuable information on
characterize not only deposition in the proximal Pennsylvanian facies and trace fossils, including S.
portion of the inner estuary, but also the basal Beaty, P. Daniel, B. Lanier, R. Leisler, C. Maples, T.
transgressive deposits immediately overlying the Stanley and R. West. R. West is also thanked for his
co-planar surface (Buatois et al. 1998a). In this valuable comments on an early draft. J. MacEachern
and M. Gingras provided careful reviews that consider-
specific depositional setting and at this particular ably improved the paper. We thank D. Mcllroy for the
stage of estuarine valley evolution, freshwater invitation to participate in the Lyell Meeting, 2003. We
conditions coexist with tidal influence. As trans- are also grateful to D. Ruiz Holgado and M. P. Jimenez
gression proceeds, backstepping brackish-water for the drawings.
deposits accumulate. The ichnologic signature
of such a change in depositional conditions is
reflected in the upward replacement of a mixed References
Scoyenia and Mermia ichnofacies by a mixed
ALLEN, G. P. 1991. Sedimentary processes and facies in
depauperate Skolithos and Cruziana ichnofacies. the Gironde estuary: a model for macrotidal
In cores, this change is evidenced by the transi- estuarine systems. In: SMITH, D. G., REINSON,
tion from parallel-laminated deposits with mini- G. E., ZAITLIN, B. A. & RAHMANI, R. A. (eds)
mal or no bioturbation to deposits displaying Clastic Tidal Sedimentology. Canadian Society of
bioturbation due to the activity of infaunal Petroleum Geologists, Memoirs, 16, 219-226.
organisms. If transgression proceeds, a diverse ARCHER, A. W. 1993. Reappraisal of Pennsylvanian
Cruziana ichnofacies eventually becomes estab- trace-fossil assemblages in the Eastern Interior
lished, recording the passage to normal-marine Coal Basin, USA. In: ARCHER, A. W., FELDMAN,
salinity conditions. H. R. & LANIER, W. P. (eds) Incised Paleovalleys
of the Douglas Group in Northeastern Kansas.
Field Guide and Related Contributions. Kansas
Geological Survey, Open-File Reports, Lawrence,
Concluding remarks 93-24, 5-1 to 5-14.
ARCHER, A. W. & MAPLES, C. G. 1984. Trace-fossil dis-
Our comparative study of a number of Carboni- tribution across a marine-to-nonmarine gradient
ferous intertidal ichnofaunas in the North Amer- in the Pennsylvanian of southwestern Indiana.
ican Midcontinent emphasizes the importance of Journal of Paleontology, 58, 448-466.
ichnology in the delineation of marginal-marine ARCHER, A. W., LANIER, W. P. & FELDMAN, H. R.
environments and allows characterization of 1994a. Stratigraphy and depositional history
three different types of intertidal trace fossil within incised-paleovalley fills and related facies,
Douglas Group (Missourian/Virgilian; Upper
assemblages. These three ichnofaunas clearly Carboniferous) of Kansas, USA. In: DALRYMPLE,
correlate with three different sub-environments R., BOYD, R. & ZAITLIN, B. A. (eds) Incised
within coastal, tide-influenced depositional sys- Valley Systems: Origin and Sedimentary
tems: open marine, restricted bays, and fluvio- Sequences. Society of Economic Paleontologists
estuarine transitions (Fig. 13). Most of these and Mineralogists, Special Publications, Tulsa,
tidal-flat deposits cannot be distinguished on Oklahoma, 51, 175-190.
the basis of physical sedimentology alone, ARCHER, A. W., FELDMAN, H. R., KVALE, E. P. &
because they have virtually the same set of LANIER, W. P. 1994b. Comparison of drier- to
physical sedimentary structures. We stress that wetter-interval estuarine roof facies in the Eastern
although lithofacies distribution is, for the most and Western Interior coal basins, USA. Palaeo-
geography, Palaeoclimatology, Palaeoecology,
part, salinity-independent, the distribution of 106, 171-185.
the benthos and therefore biogenic structures BAJARD, J. 1966. Figures et structures sedimentaires
is not. Therefore ichnological studies of dans la zone intertidale de la partie orientale
marginal-marine depositional systems can de la Baie du Mont-Saint-Michel. Revue de Geo-
provide the resolution necessary to refine facies graphie physique et de Geologie Dynamique, 8,
and sequence-stratigraphic models previously 39-111.
CARBONIFEROUS BRACKISH-WATER ICHNOFAUNAS 175

BAKER, J. 1995. Quantitative assessment of bioerosion the ichnogenus Beaconichnus and additional
and encrustation of Orthomyalina from shell beds examples from the Carboniferous of Kansas,
of the Stull Shale Member (Kanwaka Shale, USA. Ichnos, 5, 287-302.
Upper Pennsylvanian, Virgilian) of eastern Kansas. BUATOIS, L. A., MANGANO, M. G. & ACENOLAZA, F. G.
MS thesis, University of Kansas, Lawrence. 2002a. Trazas fosiles: Senates de comportamiento
BANDEL, K. 1967a. Trace fossils from two Upper Penn- en el registro estratigrdfico. Museo Paleontologico
sylvanian sandstones in Kansas. The University of Egidio Feruglio, Trelew.
Kansas Paleontological Contributions, 18, 1-13. BUATOIS, L. A., MANGANO, M. G., ALISSA, A. & CARR,
BANDEL, K. 1967b. Isopod and limulid marks and trails T. R. 2002b. Sequence stratigraphic and sedimen-
in Tonganoxie Sandstone (Upper Pennsylvanian) tologic significance of biogenic structures from a
of Kansas. The University of Kansas Paleonto- late Paleozoic reservoir, Morrow Sandstone, sub-
logical Contributions, 19, 1-10. surface of Southwest Kansas, USA. Sedimentary
BEUKEMA, J. J. 1976. Biomass and species richness of Geology, 152, 99-132.
the macro-benthic animals living on the tidal BUCKMAN, J. O. 1992. Palaeoenvironment of a Lower
flats of the Dutch Wadden Sea. Netherlands Carboniferous sandstone succession northwest
Journal of Sea Research, 10, 236—261. Ireland: ichnological and sedimentological studies.
BINYON, J. 1972. Physiology ofEchinoderms. Pergamon In: PARNELL, J. (ed.) Basins on the Atlantic
Press, Oxford. Seaboard: Petroleum Sedimentology and Basin
BRIDGES, P. H. & LEEDER, M. R. 1976. Sedimentary Evolution. Geological Society, London, Special
model for intertidal mudflat channels, with Publications, 62, 217-241.
examples from the Solway Firth, Scotland. Sedi- CROGHAN, P. C. 1983. Osmotic regulation and the
mentology, 23, 533-552. evolution of brackish- and fresh-water Faunas.
BROMLEY, R. G. 1996. Trace Fossils: Biology, Journal of the Geological Society, London, 140,
Taphonomy and Applications. Chapman & Hall, 39-46.
London. DALRYMPLE, R. W. 1992. Tidal depositional systems.
BROWNFIELD, R. L., BRENNER, R. L. & POPE, J. R. 1998. In: WALKER, R. G. & JAMES, N. P. (eds) Facies
Distribution of the Bandera Shale of the Marma- Models: Response to Sea Level Change. Geological
ton Group, Middle Pennsylvanian of Southeastern Association of Canada, Ontario, 195-218.
Kansas. Current Research in Earth Sciences, 241, DALRYMPLE, R. W., KNIGHT, R. J., ZAITLIN, B. A. &
29-41. MIDDLETON, G. V. 1990. Dynamics and facies
BUATOIS, L. A. & MANGANO, M. G. 1995. The paleoen- model of a macrotidal sand-bar complex, Cobe-
vironmental and paleoecological significance of quid Bay-Salmon River estuary (Bay of Fundy).
the lacustrine Mermia ichnofacies: an archetypical Sedimentology, 37, 577-612.
subaqueous nonmarine trace fossil assemblage. DALRYMPLE, R. W., ZAITLIN, B. A. & BOYD, R. 1992.
Ichnos,4, 151-161. Estuarine facies models: conceptual basis and
BUATOIS, L. A. & MANGANO, M.G. 1998. Trace fossil stratigraphic implications. Journal of Sedimentary
analysis of lacustrine facies and basins. Palaeogeo- Petrology, 62, 1130-1146.
graphy, Palaeoclimatology, Palaeoecology, 140, DE MOWBRAY, T. 1983. The genesis of lateral accretion
367-382. deposits in recent intertidal mudflat channels,
BUATOIS, L.A. & MANGANO, M.G. 2000. Icnologia: Solway Firth, Scotland. Sedimentology, 30, 425-
Aplicaciones de la icnologia en prospeccion de 435.
hidrocarburos y caracterizacion de reservorios. DEVERA, J. A. 1989. Ichnofossil assemblages and
Bole tin de Informaciones Petroleras, 62, 64-85. associated lithofacies of the Lower Pennsylvanian
BUATOIS, L. A., MANGANO, M. G., MAPLES, C. G. & (Caseyville and Tradewater Formations), south-
LANIER, W. P. 1997. The paradox of nonmarine ern Illinois. In: COBB, J. D. (coordinator) Geology
ichnofaunas in tidal rhythmites: Integrating of the Lower Pennsylvanian in Kentucky, Indiana,
sedimentologic and ichnologic data from the late and Illinois. Illinois Basin Studies, Indiana Geo-
Carboniferous of eastern Kansas, USA. Palaios, logical Survey, Bloomington, 1, 57-83.
12,467-481. EAGAR, R. M. C., BAINES, J. G., COLLINSON, J. D.,
BUATOIS, L. A., MANGANO, M. G., MAPLES, C. G. & HARDY, P. G., OKOLO, S. A. & POLLARD, J. E.
LANIER, W. P. 1998a. Allostratigraphic and sedi- 1985. Trace fossil assemblages and their occur-
mentologic applications of trace fossils to the rence in Silesian (Mid-Carboniferous) deltaic
study of incised estuarine valleys: an example sediments of the Central Pennine Basin, England.
from the Virgilian Tonganoxie sandstone of In: CURRAN, H. A. (ed.) Biogenic Structures:
eastern Kansas. Current Research in Earth Their Use in Interpreting Depositional Environ-
Sciences, 241, 1-27. ments. Society of Economic Paleontologists and
BUATOIS, L. A., MANGANO, M. G., MAPLES, C. G. & Mineralogists, Special Publications, Tulsa, Okla-
LANIER, W. P. 1998b. Ichnology of an Upper homa, 35, 99-149.
Carboniferous fluvio-estuarine paleovalley: the FAIRBRIDGE, R. W. 1980. The estuary: its definition and
Tonganoxie Sandstone, Buildex Quarry, eastern geodynamic cycle. In: OLAUSSON, E. & CATO, I.
Kansas, USA. Journal of Paleontology, 72, 152- (eds) Chemistry and Biogeochemistry of Estuaries.
180. Wiley, Bath, 1-35.
BUATOIS, L. A., MANGANO, M. G., MAPLES, C. G. & FELDMAN, H. R., ARCHER, A. W., WEST, R. R.,
LANIER, W. P. 1998c. Taxonomic reassessment of MAPLES, C. G., CUNNINGHAM, C. R., SCHULTZE,
176 M. G. MANGANO & L. A. BUATOIS

H. P. & LANIER, W. P. 1993. The Garnett and Mineralogists, Special Publications, Tulsa, Okla-
Hamilton Paleovalleys and their relationship to homa, 35, 21-35.
the Douglas Group Paleovalleys. In: ARCHER, HECKEL, P. H. 1977. Origin of phosphatic black-shale
A. W., FELDMAN, H. R. & LANIER, W. P. (eds) facies in Pennsylvanian cyclothems of midconti-
Incised Paleovalleys of the Douglas Group in North- nent North America. American Association of
eastern Kansas. Field Guide and Related Contribu- Petroleum Geologists, Bulletin, 61, 1045-1068.
tions. Kansas Geological Survey, Open-File HOWARD, J. D. & DORIES, J. 1972. Animal-sediment
Reports, Lawrence, 93-24, 12-1 to 12-10. relationships in two beach-related tidal flats;
FELDMAN, H. R., GIBLING, M. R., ARCHER, A. W., Sapelo Island, Georgia. Journal of Sedimentary
WIGHTMAN, W. G. & LANIER, W. P. 1995. Petrology, 42, 608-623.
Stratigraphic architecture of the Tonganoxie HUDSON, J. D. 1990. Salinity from faunal analysis and
Paleovalley Fill (Lower Virgilian) in Northeastern geochemistry. In: BRIGGS, D. E. G. & CROWTHER,
Kansas. American Association of Petroleum P. R. (eds) Palaeobiology: A Synthesis. Blackwell
Geologists, Bulletin, 79, 1019-1043. Scientific, London, 406-410.
FREY, R. W. & PEMBERTON, S. G. 1987. The Psilonich- KLEIN, G. DE V. 1971. A sedimentary model for deter-
nus ichnocoenose, and its relationship to adjacent mining paleotidal range. Geological Society of
marine and nonmarine ichnocoenoses along the America, Bulletin, 82, 2585-2592.
Georgia coast. Bulletin of Canadian Petroleum KLEIN, G. DE V. 1977. Clastic Tidal Facies. CEPCO,
Geology, 35, 333-357. Champaign, IL.
FREY, R. W., HOWARD, J. D. & HONG, J. S. 1987. KVALE, E. P. & BARNHILL, M. L. 1994. Evolution of
Prevalent Lebensspuren on a modern macrotidal Lower Pennsylvanian estuarine facies within two
flat, Inchon, Korea: ethological and environ- adjacent paleovalleys, Illinois Basin, Indiana. In:
mental significance. Palaios, 2, 571-593. DALRYMPLE, R., BOYD, R. & ZAITLIN, B. A. (eds)
GENISE, J. F., MANGANO, M. G., BUATOIS, L. A., LAZA, Incised Valley Systems: Origin and Sedimentary
J. & VERDE, M. 2000. Insect trace fossil associa- Sequences. Society of Economic Paleontologists
tions in paleosols: the Coprinisphaera ichnofacies. and Mineralogists, Special Publications, Tulsa,
Palaios, 15, 33-48. Oklahoma, 51, 191-207.
GHARE, M. A. & BADVE, R.M. 1984. Observations on LANIER, W. P. 1993. Bedform sedimentology of the
ichnoactivity from the intertidal environment, Lonestar Spillway and Buidex Quarry stops. In:
west coast of Raigad District, Maharashtra. ARCHER, A. W., FELDMAN, H. R. & LANIER, W.
Biovigyanam, 10, 173-178. P. (eds) Incised Paleovalleys of the Douglas Group
GIBLING, M. R., FELDMAN, H. R., ARCHER, A. W. & in northeastern Kansas. Field Guide and Related
LANIER, W. P. 1993. Sedimentology, stratigraphy, Contributions. Kansas Geological Survey, Open-
and paleoflow patterns of the Tonganoxie Sand- File Reports, Lawrence, 93-24, 4-1 to 4-10.
stone Member and related strata in northeast LANIER, W. P., FELDMAN, H. R. & ARCHER, A. W. 1993:
Kansas and southwest Missouri. In: ARCHER, Tidal sedimentation from a fluvial to estuarine
A. W., FELDMAN, H. R. & LANIER, W. P. (eds) transition, Douglas Group, Missourian-Virgilian,
Incised Paleovalleys of the Douglas Group in Kansas. Journal of Sedimentary Petrology, 63,
northeastern Kansas. Field Guide and Related Con- 860-873.
tributions. Kansas Geological Survey, Open-File MACEACHERN, J. A. & PEMBERTON, G. 1992. Ichnologi-
Reports, Lawrence, 93-24, 3-1 to 3-39. cal aspects of Cretaceous shoreface successions
GINGRAS, M. K., PEMBERTON, S. G., SAUNDERS, T. & and shoreface variability in the Western Interior
CLIFTON, H. E. 1999. The ichnology of Modern seaway of North America. In: PEMBERTON, S. G.
and Pleistocene brackish-water deposits at (ed.) Applications of Ichnology to Petroleum
Willapa Bay, Washington: Variability in estuarine Exploration. Society of Economic Paleontologists
settings. Palaios, 14, 352-374. and Mineralogists, Core Workshops, Tulsa, Okla-
GOODWIN, P. W. & ANDERSON, E. J. 1974. Associated homa, 17, 57-84.
physical and biogenic structures in environmental MAC£ACHERN, J. A. & PEMBERTON, S. G. 1994. Ichno-
subdivision of a Cambrian tidal sand body. logical aspects of incised valley fill systems from
Journal of Geology, 82, 779-794. the Viking Formation of the Western Canada
HAKES, W. G. 1976. Trace fossils and depositional Sedimentary Basin, Alberta, Canada. In: DALRYM-
environment of four clastic units, Upper Pennsyl- PLE, R., BOYD, R. & ZAITLIN, B. A. (eds) Incised
vanian megacyclothems, northeast Kansas. The Valley Systems: Origin and Sedimentary
University of Kansas Paleontological Contribu- Sequences. Society of Economic Paleontologists
tions, 63, 1-46. and Mineralogists, Special Publications, Tulsa,
HAKES, W. G. 1977. Trace fossils in Late Pennsylvanian Oklahoma, 51, 129-157.
cyclothems, Kansas. In: CRIMES, T. P. & HARPER, MACEACHERN, J. A., RAYCHAUDHURI, I. & PEMBERTON,
J. C. (eds) Trace Fossils 2. Geological Journal, S. G. 1992. Stratigraphic applications of the Glos-
Special Issue, 9, 209-226. sifungites ichnofacies: delineating discontinuities
HAKES, W. G. 1985. Trace fossils from brackish-marine in the rock record. In: PEMBERTON, S. G. (ed.)
shales, Upper Pennsylvanian of Kansas, USA. In: Applications of Ichnology to Petroleum Explora-
CURRAN, H. A. (ed.) Biogenic Structures: Their tion. Society of Economic Paleontologists and
Use in Interpreting Depositional Environments. Mineralogists, Core Workshops, Tulsa, Okla-
Society of Economic Paleontologists and homa, 17, 169-198.
CARBONIFEROUS BRACKISH-WATER ICHNOFAUNAS 177

MACEACHERN, J. A., ZAITLIN, B. A. & PEMBERTON, S. Pennsylvanian coal-bearing sequence, northern


G. 1999. A sharp-based sandstone of the Viking Cumberland Plateau, Tennessee, USA. In:
Formation, Joffre Field, Alberta, Canada: criteria CURRAN, H. A. (ed.) Biogenic Structures: Their
for recognition of transgressively incised shoreface Use in Interpreting Depositional Environments.
complexes. Journal of Sedimentary Research Society of Economic Paleontologists and Mineral-
Section B, 69, 876-892. ogists, Special Publications, Tulsa, Oklahoma, 35,
MANGANO, M. G. & BUATOIS, L. A. 1997. Analisis icno- 67-97.
logico comparative de planicies mareales carboni- MOORE, R. C. 1964. Paleoecological aspects of Kansas
feras del este de Kansas. Memorias ler Congreso Pennsylvanian and Permian cyclothems. In:
Latinoamericano de Sedimentologia, 2, 1-6. MERRIAM, D. F. (ed.) Symposium on Cyclic Sedi-
MANGANO, M. G. & BUATOIS, L. A. 1999. Ichnofacies mentation. Kansas Geological Survey, Bulletin,
models in Early Paleozoic tide-dominated 169, 287-380.
quartzites: onshore-offshore gradients and the MOORE, R. C., FRYE, J. C., JEWETT, J. M., WALLACE, L.
classic Seilacherian paradigm. Acta Universitatis & O'CONNOR, H. G. 1951. The Kansas Rock
Carolinae, 43, 151-154. Column. Kansas Geological Survey, Bulletin, 89.
MANGANO, M. G., BUATOIS, L. A. & ACENOLAZA, G. F. PAGETT, R. M. 1980. Tolerance to brackish water by
1996. Trace fossils and sedimentary facies from an Ophiuroids with special reference to a Scottish
Early Ordovician tide-dominated shelf (Santa sea Loch, Loch Etive. In: JANGOUX, M. (ed.)
Rosita Formation, northwest Argentina): implica- Echinoderms: Present and Past. Proceedings of
tions for ichnofacies models of shallow marine the European Colloquium on Echinoderms, Balk-
successions. Ichnos, 5, 53-88. ema, Rotterdam, 223-229.
MANGANO, M. G., BUATOIS, L. A., MAPLES, C. G. & PAGETT, R. M. 1981. The penetration of brackish-water
LANIER, W. P. 1997. Tonganoxichnus, a new by the echinodermata. In: JONES, N. V. & WOLFF,
insect trace fossil from the Upper Carboniferous W. J. (eds) Feeding and Survival Strategies of
of eastern Kansas, USA. Lethaia, 30, 113-125. Estuarine Organisms. Plenum Press, New York,
MANGANO, M. G., BUATOIS, L. A., WEST, R. R. & 135-151.
MAPLES, C. G. 1998. Contrasting behavioral and PEMBERTON, S. G. & WIGHTMAN, D. M. 1992. Ichno-
feeding strategies recorded by tidal-flat bivalve logical characteristics of brackish water deposits.
trace fossils from the Upper Carboniferous of east- In: PEMBERTON, S. G. (ed.) Applications of Ichnol-
ern Kansas. Palaios, 13, 335-351. ogy to Petroleum Exploration. Society of Eco-
MANGANO, M. G., BUATOIS, L. A., WEST, R. R. & nomic Paleontologists and Mineralogists, Core
MAPLES, C. G. 1999. The origin and paleoecologic Workshops, Tulsa, Oklahoma, 17, 141-167.
significance of the trace fossil Asteriacites in the PEMBERTON, S. G., SPILA, M., PULHAM, A. J., SAUN-
Pennsylvanian of Kansas and Missouri. Lethaia, DERS, T., MACEACHERN, J. A., ROBBINS, D. &
32, 17-30. SINCLAIR, I. K. 2001. Ichnology & Sedimentology
MANGANO, M. G., LABANDEIRA, C., KVALE, E. & of Shallow to Marginal Marine Systems. Ben
BUATOIS, L. A. 2001. The insect trace fossil Tonga- Nevis & Avalon Reservoirs, Jeanne d'Arc Basin.
noxichnus from the Middle Pennsylvanian of Geological Association of Canada, Short Course
Indiana: paleobiologic and paleoenvironmental Notes, St. John's, Newfoundland, 15, 1-343.
implications. Ichnos, 8, 165-175. PICKERILL, R. K. & BRENCHLEY, P. J. 1991. Benthic
MANGANO, M. G., BUATOIS, L. A., WEST, R. R. & macrofossils as paleoenvironmental indicators in
MAPLES, C. G. 2002a. Ichnology of an equatorial marine siliciclastic facies. Geoscience Canada, 18,
tidal flat: the Stull Shale Member at Waverly, east- 119-138.
ern Kansas. Bulletin of the Kansas Geological POLLARD, J. E. 1988. Trace fossils in coal-bearing
Survey, 245, 1-130. sequences. Journal of the Geological Society,
MANGANO, M. G., BUATOIS, L. A. & RINDSBERG, A. K. London, 145, 339-350.
2002b. Carboniferous Psammichnites: systematic POLLARD, J. E., GOLDRING, R. & BUCK, S. G. 1993.
re-evaluation, taphonomy and autecology. Ichnofabrics containing Ophiomorpha: signifi-
Ichnos, 9, 1-22. cance in shallow-water facies interpretation.
MARTINO, R. L. 1989. Trace fossils from marginal Journal of the Geological Society, London, 150,
marine facies of the Kanawa Formation (Middle 149-164.
Pennsylvanian), West Virginia. Journal of Paleon- REINECK, H.-E. 1958. Longitudinale schragschicht in
tology, 63, 389^03. Watt. Geologische Rundschau, 47, 73-82.
MARTINO, R. L. 1996. Stratigraphy and depositional REINECK, H.-E. 1967. Layered sediments of tidal flat
environments of the Kanawha Formation beaches, and shelf bottoms of the North Sea. In:
(Middle Pennsylvanian), southern West Virginia, LAUFF, G. H. (ed.) Estuaries. American Associa-
USA. International Journal of Coal Geology, 31, tion for the Advancement of Science, Special
217-248. Publications, Washington D.C., 83, 191-206.
MclLROY, D. 2004. In: MC!LROY, D. (ed.) 2004. The REISE, K. 1985. Tidal Flat Ecology: An experimental
Application of Ichnology to Palaeoenvironmental Approach to Species Interactions. Ecological
and Stratigraphic Analysis, Geological Society, Studies, 54, Springer, Berlin/Heidelberg.
London, Special Publications, 228, 237-272. REMANE, A. & SCHLDEPER, C. 1971. Biology of Brackish
MILLER, M. F. & KNOX, L. W. 1985. Biogenic struc- Water (2nd rev. edn). Wiley Interscience, New
tures and depositional environments of a Lower York.
178 M. G. MANGANO & L. A. BUATOIS

RINDSBERG, A. K. 1990. Freshwater to marine trace TURNER, R. L. 1980. Salinity tolerance of the brackish-
fossils of the Mary Lee Coal zone and overlying water echinoderm Ophiophragmus filograneus
strata (Westphalian A), Pottsville Formation of (Ophiuroidea). Marine Ecology Progress Series,
northern Alabama. In: GASTALDO, R. A., DEMKO, 2, 249-256.
T. M. & Liu, Y. (eds) Carboniferous Coastal VAN WAGONER, J. C., MITCHUM, R. M., CAMPION,
Environments and Paleocommunities of the Mary K. M. & RAHMANIAN, V. D. 1990. Siliciclastic
Lee Coal zone, Marion and Walker Counties, Ala- sequences, stratigraphy in well logs, cores, and
bama: A Guidebook for Field Trip VI, Southeastern outcrops. American Association of Petroleum
Section, Geological Society of America. Geological Geologists, Methods in Exploration Series, 7, 1-55.
Survey of Alabama, Tuscaloosa, 82-95. WATNEY, W. L., FRENCH, J. & FRANSEEN, E. K. 1989.
RINDSBERG, A. K. 1994. Ichnology of the Upper Missis- Sequence Stratigraphic Interpretations and Model-
sippian Hartselle Sandstone of Alabama, with ing of Cyclothems in the Upper Pennsylvanian
notes on other Carboniferous formations. Geologi- (Missourian) Lansing and Kansas City Groups in
cal Survey of Alabama Bulletin, 158, 1-107. Eastern Kansas. Kansas Geological Society, 41st
Ross, C. A. & Ross, J. R. P. 1987. Late Paleozoic Sea Annual Field Trip, Lawrence.
Levels and Depositional Sequences. In: Ross, C. A. WEIMER, R. J., HOWARD, J. D. & LINDSAY, D. R. 1981.
& HAMAN, D. (eds) Timing and Depositional Tidal flats and associated tidal channels. In:
History of Eustatic Sequences: Constraints on SCHOLLE, P. A. & SPEARING, D. (eds) Sandstone
Seismic Stratigraphy. Cushman Foundation for Depositional Environments. American Association
Foraminiferal Research, Special Publications, of Petroleum Geologists, Memoirs, Tulsa, Okla-
Tulsa, Oklahoma, 24, 137-168. homa, 31, 191-245.
RUSSELL, J. L. 1972. Depositional environment of the WESCOTT, W. A. & UTGAARD, J. E. 1987. An Upper
Rock Lake Shale. Geological Society of America, Mississippian trace-fossil assemblage from the
Abstracts with Programs, 4, 293. Tar Springs Sandstone, southern Illinois. Journal
RUSSELL, J. L. 1974. Comparison of two late Paleozoic of Paleontology, 61, 231-241.
red shales of the Midcontinent region. PhD thesis, WIGHTMAN, D. M., PEMBERTON, S. G. & SINGH, C.
University of Nebraska. 1987. Depositional modelling of the Upper Mann-
SEGERSTRALE, S. 1957. Baltic Sea. Geological Society of ville (Lower Cretaceous), east-central Alberta:
America, Memoirs, 67, 1-32. implications for the recognition of brackish
SPAARGAREN, D. H. 1979. Marine and brackish-water water deposits. In: TILLMAN, R. W. & WEBER,
animals. In: MALLOIY, G. M. O. (ed.) Comparative K. J. (eds) Reservoir Sedimentology. Society of
Physiology of Osmoregulation in Animals. Aca- Economic Paleontologists and Mineralogists,
demic Press, Edinburgh, 83-116. Special Publications, Tulsa, Oklahoma, 40, 189-
SPAARGAREN, D. H. 1995. A functional model for 220.
describing the responses to salinity stress in WILSON, J. B. 1982. Shelly faunas associated with tem-
aquatic animals. Comparative Biochemistry and perate offshore tidal deposits. In: STRIDE, A. H.
Physiology, 111, 501-506. (ed.) Offshore Tidal Sands: Processes and Deposits.
STANCYK, S. E. 1973. Development of Ophiolepis Chapman & Hall, New York, 126-171.
elegans (Echinodermata: Ophiuroidea) and its WILSON, J. B. 1986. Faunas of tidal current and wave-
implications in the estuarine environment. dominated continental shelves and their use in the
Marine Biology, 21, 7-12. recognition of storm deposits. In: KNIGHT, R. J. &
STRAATEN, L. M. J. VAN. 1954. Composition and struc- McLEAN, J. R. (eds) Shelf Sands and Sandstone
ture of recent marine sediments in the Nether- Reservoirs. Canadian Society of Petroleum Geolo-
lands. Leidse Geologische Mededelingen, 19,1—110. gists, Memoirs, Ontario, 11, 313-326.
SWINBANKS, D. D. & MURRAY, J. W. 1981. Biosedi- ZAITLIN, B. A., DALRYMPLE, R. W. & BOYD, R. 1994.
mentological zonation of Boundary Bay tidal The Stratigraphic organization of incised-valley
flats, Fraser River Delta, British Columbia. Sedi- systems associated with relative sea-level changes.
mentology, 28, 201-237. In: DALRYMPLE, R., BOYD, R. & ZAITLIN, B. A.
THOMAS, R. G., SMITH, D. G., WOOD, J. M., VISSER, J., (eds) Incised Valley Systems: Origin and Sedimen-
CAVERLEY-RANGE, E. A. & KOSTER, E. H. 1987. tary Sequences. Society of Economic Paleontolo-
Inclined heterolithic stratification: terminology, gists and Mineralogists, Special Publications,
description, interpretation and significance. Tulsa, Oklahoma, 51, 45-60.
Sedimentary Geology, 53, 123-179.
Differentiation of estuarine and offshore marine deposits using
integrated ichnology and sedimentology: Permian Pebbley Beach
Formation, Sydney Basin, Australia

KERRIE L. BANN1, CHRISTOPHER R. FIELDING2, JAMES A. MAcEACHERN3


& STUART C. TYE4

1
1Department of Earth and Atmospheric Sciences, University of Alberta, Edmonton,
Alberta, CanadaT6G 2E3 (e-mail: kbann@ualberta.ca)
2
2DepartmentofGeosciences, 214 Bessey Hall, University of Nebraska-Lincoln,
NE 68588-0340, USA (e-mail:cfieldmg2@unl.edu)
3Department of Earth Sciences, Simon Fraser University, Burnaby,
British Columbia, Canada V5A 1S6 (e-mail: jmaceach@sfu.ca)
4
4Husky Energy, 707 8th Ave Calgary, SW Alberta, Canada T2P 3G7
(e-mail: stuart.tye@huskyenergy.ca)

Abstract: This study integrates ichnology and sedimentology to refine the palaeoenviron-
mental and sequence stratigraphic interpretations of the Early Permian Pebbley Beach
Formation, in the southern Sydney Basin, Australia. This succession has been interpreted
previously to reflect entirely inner to outer shelf and slope environments of deposition.
Detailed analysis of the formation reveals ichnological and sedimentological characteristics
that contradict a fully marine interpretation. Instead, the interval reflects the vertical super-
position and lateral juxtaposition of brackish-water and fully marine units. Marine facies
comprise: (1) thoroughly bioturbated muddy siltstone (lower offshore); (2) thoroughly bio-
turbated sandy siltstone (upper offshore); (3) interbedded bioturbated sandy siltstone and
laminated sandstone (delta-influenced offshore transition); (4) thoroughly bioturbated
muddy sandstone (distal lower shoreface); (5) interbedded laminated sandstone, bioturbated
muddy sandstone and dark claystone (delta-influenced lower shoreface); and (6) bioturbated,
laterally variable sandstones (transgressive sand sheets). Estuarine facies comprise: (1)
channelized heterolithic sandstone-mudstone (active estuarine channels); (2) sheet-like
heterolithic sandstone-mudstone (active estuarine basins); and (3) laminated mudstone
(abandoned estuarine channels and basins). The interpreted fully marine deposits contain
ichnological suites that exhibit moderate to intense bioturbation, high diversities (31 ichno-
species belonging to 20 ichnogenera), uniform distributions of ichnogenera, and significant
numbers of structures reflecting specialized feeding/grazing behaviours. In marked contrast,
interpreted estuarine (brackish-water) deposits contain impoverished ichnological suites
(9 ichnogenera), show variable but significantly reduced degrees of bioturbation intensity,
pronounced variability in ichnogenera distributions and the predominance of a few, simple
forms representing simple feeding strategies of resilient trophic generalists. The new analysis
allows the recognition of a series of highly top-truncated and condensed sequences (cycles of
relative sea-level fall and physical rise), which can be physically correlated over several
kilometres. Sequence boundaries typically cut down through shoreface sandstones to directly
overlie offshore facies, leading to an interface with little apparent lithological contrast. In the
absence of laterally continuous exposure, these surfaces may be recognized by careful
ichnofacies evaluation. Thus the re-evaluation presented herein has facilitated a more
realistic sequence stratigraphic analysis of the Pebbley Beach Formation.

Although facies models for indented coastal stratigraphic analysis provides a useful way of
systems are well-established (e.g. Dalrymple discriminating between lithologically similar
et al. 1992), there is nonetheless a shortage of facies. Literature on the ichnology of brackish-
reliable diagnostic criteria to allow discrimina- water environments is limited, particularly for
tion between fine-grained facies of coastal successions of late Palaeozoic age (but see
origin and those of marine shelf origin. The Buatois et al. 2002). Here, we describe a succes-
possibility therefore exists for confusion between sion of Early Permian age that contains both
facies that may be superficially similar but which offshore marine and estuarine facies that are
record widely separated environments of lithologically similar, but which can be separated
deposition. The integration of ichnology into successfully using a combination of ichnology

From: MC!LROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 179-211. 0305-8719/04/S15.00 © The Geological Society
of London.
180 K. L. BANN ET AL.

and sedimentology. From this fades analysis we of brackish-water trace fossil assemblages of
present a sequence stratigraphic model for the Permian age.
succession, highlighting the usefulness of ichnol- The Pebbley Beach Formation crops out
ogy and providing new data on the composition along the coast of southern New South Wales,

Fig. 1. Location maps: (a) position of Sydney-Bowen Basin; (b) southern Sydney Basin; (c) Pebbley Beach
Formation study area. Measured section localities are marked by bullet points.
ESTUARINE VS FULLY MARINE DEPOSITS 181

Australia, between Point Upright in the south excellent late Palaeozoic comparators to the
and Clear Point in the north (Fig. 1). This Cretaceous and Holocene estuarine trace fossil
paper focuses on the middle and upper portions models.
of the formation, which are well exposed in sea
cliffs at Point Upright, Mill Point and Clear
Point. A previous facies scheme for the Pebbley Geologic setting
Beach Formation (Gostin & Herbert 1973)
proposed a shallow marine to broadly coastal The Sydney Basin comprises the southernmost
environment of deposition; in contrast, Eyles portion of the larger, north-south elongate
et al. (1998) concluded an entirely inner to Sydney-Gunnedah-Bowen Basin (Fig. la), of
outer shelf and slope environment of deposition Late Carboniferous to Triassic age, one of a
for the interval. In that study, Eyles et al. series of contiguous basins that formed along
(1998) regarded a series of channel features the Panthalassan margin of Gondwana. The
prominent in the middle and upper parts of the basin has a complex, multiphase history. Late
unit as the product of shoreface erosion and Carboniferous (Pennsylvanian) to Early Permian
deposition in a shallow marine setting. The rifting in a back-arc environment was followed
ichnological characteristics of the various facies by middle Permian thermal subsidence (Schieb-
compel us, however, to dispute this interpreta- ner 1974; Battersby 1981; Murray 1990). The
tion, as well as the assertion that the succession basin then evolved into a retro-arc foreland
is entirely of fully marine origin. basin during the Late Permian to Middle
The brackish-water and fully marine units of Triassic, adjacent to the New England Fold
the Pebbley Beach Formation are lithologically Belt (Holcombe et al. 1997).
and (to a lesser extent) sedimentologically simi- The Pebbley Beach Formation forms a part of
lar, contributing to the confusion regarding the the Early Permian (Sakmarian to Artinskian:
depositional setting. Although there are primary Briggs 1998) succession in the southernmost
sedimentological features that occur persistently portion of the exposed, onshore Sydney Basin
within the various facies of both depositional sys- (Fig. 2). It overlies the shallow marine Wasp
tems, the ichnological differences are profound. Head Formation and is in turn overlain by the
The differences in bioturbation intensity, uni- shallow marine Snapper Point Formation
form versus sporadic distribution of bioturba- (Gostin & Herbert 1973; Tye et al. 1996). The
tion and, most importantly, the trace fossil Wasp Head Formation forms a part of the
assemblages themselves permit the reliable differ- Talaterang Group, which includes the Clyde
entiation of brackish-water (estuarine) deposits Coal Measures and represents deposition
from fully marine offshore and lower shoreface during the initial extensional phase of basin
intervals. The recognition of these disparate formation (Tye et al. 1996). The Pebbley Beach
depositional environments has a profound Formation comprises the basal unit of the Shoal-
impact upon the high-resolution sequence haven Group (Gostin & Herbert 1973), which
stratigraphic interpretation of the succession. was deposited late in the extensional phase and
The integration of ichnology with the sedimento- during the ensuing phase of passive thermal sub-
logical facies model has led to the identification sidence. The fully marine and nearshore coastal
of sequence boundaries, marine flooding surfaces deposits of the Pebbley Beach Formation can
and transgressive surfaces of erosion, demon- be correlated lithostratigraphically with the
strating that the Pebbley Beach Formation has fluvial Yadboro Conglomerate to the immediate
a complex sequence-stratigraphic architecture. west (Tye et al. 1996). The interpreted palaeo-
The recognition of brackish-water deposits geography for this period involves a north-
based on ichnological characteristics is princi- south-elongate shoreline that was at most times
pally derived from studies of: (1) the Holocene somewhere to the immediate west of the expo-
from the North Sea (e.g. Schafer 1962; Reineck sures described herein, with sediment dispersal
et al. 1967, 1968; Hertweck 1970) and the from continental drainage systems from west to
Georgia coast of the US (e.g. Frey & Howard east (Tye et al. 1996).
1972; Howard & Frey 1973, 1975; Howard et al. During Early Permian time, eastern Australia
1975); and (2) Cretaceous units of the Western was located along the Panthallassan margin of
Interior Seaway of North America (e.g. Pember- the Gondwana landmass, connected to Antarc-
ton et al. 1982; Ekdale et al. 1984; Howard & tica and adjacent to the palaeo-Pacific Ocean
Frey 1984; Wightman et al. 1987; Beynon et al. (Veevers & Powell 1987). The South Pole was
1988; Ranger & Pemberton 1992; MacEachern near the eastern edge of Antarctica (Crowell &
& Pemberton 1994). The trace fossil assemblages Frakes 1975), close to eastern Australia, suggest-
of the Pebbley Beach Formation serve as ing that the latitude of the southern Sydney
182 K. L. BANN ET AL.

Fig. 2. Stratigraphy of the southern Sydney Basin (modified from Tye et al. 1996). YCM, Yurrunga Coal
Measures; N-m, non-marine; Art., Artinskian; Kun., Kungurian.

Basin was similar to that of the present-day Ross by proximity to a contemporaneous delta
Ice Shelf. The presence of (1) glendonites (a system), to coastal environments that are broadly
pseudomorph after the mineral ikaite which estuarine (brackish water) in character.
iforms at temperatures less than 5 °C, and from The use of the terms 'upper offshore' and
which cold climate conditions have been 'lower offshore' follows the usage of Mac-
invoked extensively in the Permian of eastern Eachern & Pemberton (1992) and Pemberton &
Australia: Kaplan 1979; Carr et al. 1989), (2) MacEachern (1995, 1997), modified from
large, exotic outsized clasts, and (3) the distinc- Howard (1971, 1972), Howard & Reineck
tive coldwater Eurydesma fauna (Runnegar (1981), Howard & Frey (1984), Vossler &
1979; Dickens 1984) provides convincing Pemberton (1989) and Frey (1990). The upper
evidence for persistent cold climate throughout offshore lies below fair-weather wavebase
the deposition of the Pebbley Beach Formation. but adjacent to (and grading into) the lower
The cold period was associated with a wide- shoreface. The proximity to the lower shoreface
spread Gondwana glaciation that began in Late results in the upper offshore receiving significant
Carboniferous time and persisted until the amounts of sand and silt with clay, producing
Early Permian (Dickens 1984; Veevers & characteristic sandy mudstone and sandy
Powell 1987). siltstone facies. The lower offshore grades
basinward into the shelf. The lower offshore
receives more silt than sand, with clay
Facies analysis deposition predominant during fair-weather
conditions, resulting in the production of silty
Nine facies are defined here, based on lithology, mudstone.
primary sedimentary structures and ichnology.
Measurement of bioturbation intensity (BI) in
the field follows the scheme illustrated in Figs 3 Marine facies
and 4 (adapted after Reineck 1963; Taylor &
Goldring 1993), and trace fossils are listed in Facies 1: Thoroughly bioturbated siltstone
decreasing order of abundance. These facies
record a depositional gradient from lower off- Sedimentology
shore environments in an open marine setting, This facies comprises siltstone and silty mud-
through shoreface environments (locally affected stone with few if any preserved sedimentary
ESTUARINE VS FULLY MARINE DEPOSITS 183

Fig. 3. Legend of symbols used in the graphic logs.

Fig. 4. Bioturbation intensity (BI). Measurement of the intensity of bioturbation in the Pebbley Beach
Formation, modified from Bann (1998), originally adapted and modified after Reineck (1963) and Taylor &
Goldring (1993).
184 K. L. BANN ET AL.

Fig. 5. Fades 1 and 2. (a) Heavily bioturbated (BI5) Fades 2, capped by a marine flooding surface (FS),
overlain by bioturbated (BI 4-5) Fades 1. The marine flooding surface is overlain by a thin, gritty sandstone
layer. Arrows mark Diplocraterion habichi (D), Phycosiphon (Ph) and Chondrites (Ch). Fig. 15, 15.5m. (b)
Large, tabular glendonite (Gl) in Facies 1 (lower offshore), Fig. 15, 16m. (c) Thoroughly bioturbated (BI4-5)
Facies 2. A thin, remnant tempestite showing wavy parallel lamination is present in the centre of the photo.
The trace fossil assemblage reflects the archetypal Cruziana ichnofacies, and comprises Chondrites (Ch),
Teichichnus (T), Zoophycos (Z), Phycosiphon (Ph), Planolites (P), Asterosoma (As) and Palaeophycus (Pa).
Fig. 14, 17-18.3m. (d) Thoroughly bioturbated (BI4-5) Facies 2 containing a remnant parallel-laminated
tempestite. The trace fossil suite reflects the archetypal Cruziana ichnofacies, manifest as Planolites (P),
Chondrites (Ch), Rosselia (Ro), Palaeophycus (Pa), Asterosoma (As), Zoophycos (Z), Helminthopsis (H),
Phycosiphon (Ph) and fugichnia (fu). Fig. 14, 17-18.3m. (e) Intensely bioturbated (BI5) Facies 1 with large,
outsized, exotic clast. Fig. 1, Depot Beach.
ESTUARINE VS FULLY MARINE DEPOSITS 185

structures (Fig. 5a). Interstitial sand (very fine- to MacEachern 1997). The presence of abundant,
fine-grained) is rare. Also rare are thin (<1 cm), large, exotic clasts within these fine-grained
sharp-based, very fine- to fine-grained sandstone deposits (Fig. 5e) has been used by previous
beds that contain low-angle undulatory parallel workers (Gostin & Herbert 1973; Eyles et al
lamination, possible small-scale hummocky 1998) as evidence for persistent delivery of ice-
cross-stratification (HCS), combined flow-ripple rafted debris. Glendonites are also strong
and current-ripple cross-lamination. Glendonites evidence for very cold climatic conditions.
(Fig. 5b), outsized clasts ranging from 5mm to This facies reflects deposition in a lower off-
greater than 1 m in diameter, carbonaceous detri- shore, open marine environment, affected by
tus, pyrite staining and wood fragments are very cold climatic conditions, but only rarely by
locally common. severe storms.

Ichnology
Facies 1 is characterized by more or less per- Facies 2: Thoroughly bioturbated sandy
vasive bioturbation, varying from BI4 to BI6, siltstone
but typically BI5. Bioturbation is locally spora-
dic, decreasing in intensity in the thin sandstone Sedimentology
beds. Most ichnogenera are uniformly distribu- This facies comprises thoroughly bioturbated
ted throughout the facies, with less common siltstone and sandy siltstone units with rare but
elements sporadically distributed. upwardly increasing numbers of discrete, thin
The trace fossil assemblage is dominated by (0.5-2 cm), very fine- to fine-grained sandstone
Phycosiphon, Planolites, Rosselia socialis, lesser beds. Sandstone units display remnant low-
Rosselia rotatus and small tightly curved Taeni- angle, undulatory, parallel lamination and
dium (type A). Common but subordinate ele- lesser wave-ripple lamination (Fig. 5c, d). Inter-
ments comprise Zoophycos (Fig. 5c), Chondrites stitial sand is pervasively distributed throughout
(Fig. 5d), Teichichnus, Palaeophycus tubularis the siltstone units, Synaeresis cracks are rare and
and Palaeophycus heberti. Uncommon elements are associated with decreased levels of bioturba-
consist of Helminthopsis and Asterosoma. The tion. Dispersed outsized clasts, clast clusters,
thin sandstone beds also locally contain Diplo- allochthonous logs, soft-sediment deformation
craterion habichi, rare Skolithos and fugichnia. structures and glendonites are locally abundant.
Phycosiphon occurs as both robust and diminu-
tive elements, whereas Rosselia, Zoophycos, Ichnology
Diplocraterion, Skolithos and Asterosoma are Facies 2 is characterized by more or less perva-
diminutive. The assemblage is interpreted to sive bioturbation, with intensities that vary
reflect a diverse, distal expression of the Cruziana from BI 5 to BI 6, and typically BI 6. Most ichno-
ichnofacies. The introduction of some Skolithos genera are uniformly distributed throughout the
ichnofacies elements is attributed to the sporadic facies, though elements associated with inter-
emplacement of sand beds during storms. calated sandstone beds are more sporadically
distributed. Uncommon elements are also spora-
Interpretation dically distributed. The trace fossil suite is
The fine-grained, bioturbated lithology of Facies dominated by Rosselia socialis, Zoophycos, Phy-
1 indicates a standing-water depositional envir- cosiphon, Planolites, Palaeophycus tubularis,
onment beyond the reach of most currents or Palaeophycus heberti, Teichichnus and Rhizo-
waves. The presence of marine invertebrate corallium irregulare. Subordinate elements com-
fossils points to a quiet-water, open marine set- prise Diplocraterion habichi, small Taenidium
ting. The thin sandstone beds record exceptional (type A), Rosselia rotatus, Skolithos, Bergaueria
storms and are characteristic of distal tempestites and fugichnia. Rosselia and Zoophycos occur as
that have been partially biogenically reworked by diminutive elements. The assemblage reflects
small, simple burrow types during fair-weather two phases of infaunal colonization. The main
periods. Facies 1 contains moderately diverse assemblage is interpreted to reflect a diverse
trace assemblages produced by deposit-feeding expression of the archetypal Cruziana ichno-
and grazing/foraging behaviours, typical of facies. The presence of Skolithos ichnofacies
open marine environments lying well below elements (e.g. Diplocraterion, Skolithos and Ber-
fair-weather wavebase. The preservation gaueria) reflects infaunal tempestite colonization.
potential of distal tempestites is high, owing to
emplacement well below fair-weather wavebase Interpretation
where physical processes are not competent to Facies 2 is considerably more variable than
modify them (Wheatcroft 1990; Pemberton & Facies 1, owing to the greater numbers of
Fig. 6. Fades 3. (a) Weakly to moderately bioturbated (BI2), interbedded sandstone, sandy siltstone, and mudstone with tempestites with low-angle, undulatory,
parallel lamination. Some sandstone beds show wave ripple lamination and normal grading. The trace fossil suite comprises a diverse expression of the archetypal
Cruziana ichnofacies, manifest by Phycosiphon (Ph), Chondrites (Ch), Planolites (P), Diplocraterion (D) and fugichnia (fu). Fig. 1, Mill Point, (b) Bedding plane view of
silty sandstone tempestite showing a BI of 3. The trace fossil suite comprises the archetypal Cruziana ichnofacies, and consists of Psammichnites (Ps), Phycosiphon (Ph),
Taenidium (type A, Ts) and Planolites (P). Fig. 1, Mill point, (c) Weakly to moderately bioturbated (BI 1-3) sandstone, sandy siltstone, and mudstone with wave
ripples and lesser low-angle, undulatory, parallel lamination. Mudstone interbeds locally contain very rare, diminutive synaeresis cracks (sy). The trace fossil suite
comprises the archetypal Cruziana ichnofacies, and contains Palaeophycus (Pa), Planolites (P), Chondrites (Ch), Phycosiphon (Ph), Taenidium (type A, Ts),
Diplocraterion (D) and fugichnia. Fig. 1, Mill Point, (d) Close-up, slightly to the right of the lower part of photo (c), showing more intensely bioturbated (BI3)
sandstone and clayey siltstones. The sandstones display remnant low-angle, undulatory, parallel lamination and lesser wave ripple lamination. The trace fossil suite
consists of Chondrites (Ch), Phycosiphon (Ph), Psammichnites (Ps), Palaeophycus (Pa), Planolites (P), Taenidium (type A, Ts) and Diplocraterion (D). Some diminutive
fugichnia are visible locally. Fig. 1, Mill Point.
ESTUARINE VS FULLY MARINE DEPOSITS 187

tempestites. Remnant, low-angle, undulatory, lamination, wave and combined flow-ripples are
parallel lamination is interpreted as HCS. The commonly preserved (Fig. 6c, d). Sandstone
mudstone and silty sandstone beds, coupled beds are locally draped with unbioturbated,
with the high ichnological diversities and bio- dark claystone containing organic detritus and
turbation intensities, suggest deposition in an locally abundant and generally shallow synaer-
environment that experienced lengthy periods esis cracks. Dispersed pebbles, clast clusters,
of fair-weather near, but below fair-weather allochthonous logs and shelly lags are locally
wavebase. The diverse trace fossil assemblage, common.
dominated by traces produced by deposit-
feeding and, to a lesser extent, grazing/foraging Ichnology
behaviours, is characteristic of the archetypical Facies 3 is characterized by persistent though
Cruziana ichnofacies and reflects quiescent largely sporadically distributed bioturbation.
conditions in environments lying between storm Bioturbation intensities vary from BIO to BI4
and fair-weather wavebase. Vertical burrows and typically BI3-4. Most ichnogenera are
that represent the dwellings of suspension- uniformly distributed throughout the facies,
feeding organisms (e.g. elements of the Skolithos though elements associated with intercalated
ichnofacies) correspond to the opportunistic laminated sandstone beds are more sporadically
colonization of the storm beds. More intense distributed. Rosselia and Zoophycos constitute
reworking of tempestites is the product of: exceptions, typically increasing in abundance
and size within some intervals. Uncommon
greater vertical penetration by more robust
elements are also sporadically distributed.
deposit feeders; The trace fossil assemblage can be subdivided
increased sizes and abundances of vertical
into one associated with the bioturbated sandy
burrows; and
siltstone, and one reflecting infaunal colonization
increased time between storm events, facilitat-
of the laminated sandstone units. The suite asso-
ing complete colonization and development of
ciated with the sandy siltstone is dominated by
deeply penetrating structures.
Rosselia socialis, Teichichnus, Palaeophycus tubu-
This facies is interpreted to reflect deposition in laris, Palaeophycus heberti, Phycosiphon and
an upper offshore environment. Planolites. Common but subordinate elements
Outsized clasts are interpreted to have been comprise Chondrites, Zoophycos, Rhizocorallium
introduced by ice rafting. Decreased levels of bio- irregulare, Diplocraterion habichi, Taenidium
turbation associated with rare, locally occurring (type A) and Conichnus. Helminthopsis, Skolithos
soft-sediment deformation and synaeresis and Cylindrichnus are uncommon.
cracks (Plummer & Gostin 1981; but see Pratt The laminated sandstone beds contain an
1998) may indicate a combination of increased assemblage dominated by Diplocraterion habichi,
sedimentation rates, sporadic deposition, fluctu- Diplocraterion parallelum, very large Rhizocoral-
ating salinity levels, periods of reduced oxy- lium irregulare (type A), Phycosiphon and Taeni-
genation and soupy substrates (Coates & dium (type A). Common but subordinate
MacEachern 1999, 2000). elements include Palaeophycus tubularis, Macar-
onichnus isp., Planolites, Rosselia socialis, Rosse-
lia rotatus, Teichichnus, fugichnia, Skolithos and
Facies 3: Interbedded bioturbated sandy Palaeophycus heberti. Uncommon elements
siltstone and laminated sandstone include Chondrites, Psammichnites, Rhizocoral-
lium irregulare, Zoophycos, Cylindrichnus and
Sedimentology Asterosoma. Diplocraterion parallelum and Rhi-
This facies is dominated by sparsely to moder- zocorallium irregulare (type A) are commonly
ately bioturbated sandy siltstone beds inter- large.
calated with thin (generally <10cm), low-angle
undulatory, parallel-laminated, very fine- to Interpretation
fine-grained sandstone beds, and thin (<5cm), Facies 3 records a marine depositional environ-
dark mudstone beds (Fig. 6a, b). Sandy siltstone ment that was periodically affected by storms,
beds are commonly truncated from above, and leading to the superimposition of sharp-based
their bases are indistinct owing to biogenic sand lenses and tabular beds as distal tempestites
mixing across the interface with underlying sand- over muds. Interbedded muds settled from sus-
stone units. Laminated sandstone beds are gener- pension or were distributed by gentle currents,
ally erosionally based and display laminated to and must therefore record periods of more
burrowed bedding, colloquially referred to as subdued (fair) weather. Facies 3 records a fair-
'Lam-Scram'. Relict HCS, low-angle planar weather resident trace fossil suite occupying
188 K. L. BANN ET AL.

sandy siltstones, erosionally truncated and over- reflect minor deltaic influence on the depositional
lain by sandstones with HCS, locally with a basal environment. Deltaic settings are associated with
lag, and commonly containing escape structures. increased levels of fluvial discharge and large
The fair-weather suite, dominated by the struc- amounts of associated organic detritus and clay
tures of deposit-feeding and grazing/foraging material, transported basinward during and
organisms, represents a diverse expression of immediately following storm events. During
the archetypal Cruziana ichnofacies. In contrast, post-storm conditions, this mud mantles the
the trace fossil suite in the sandstone beds is tops of storm beds, shielding them from opportu-
dominated by vertical burrows of opportunistic nistic colonization, particularly by suspension
suspension feeders, resilient surface detritus feeders. High organic contents in the mud
feeders and passive carnivores and is indicative result in its rapid oxidation, and resulting
of the Skolithos ichnofacies. This recurring oxygen depletion at the seafloor: as a conse-
juxtaposition of the Cruziana and Skolithos ich- quence, many organisms are unable to colonize
nofacies is the basis for recognizing the mixed these substrates (e.g. Leithold 1989; Raychaud-
Skolithos-Cruziana ichnofacies (Howard & huri & Pemberton 1992; Saunders et al. 1994;
Frey 1984; Pemberton & Frey 1984; MacEachern Coates & MacEachern 1999, 2000; Bann &
& Pemberton 1992; Pemberton & MacEachern Fielding 2004). The common association of
1997). The overall mixed, diverse Skolithos- synaeresis cracks with the dark mudstone
Cruziana ichnofacies suite is complex and is drapes suggests a close linkage between the
characteristic of interbedded fair-weather units storm events and concomitant heightened preci-
and tempestites deposited within the upper off- pitation, increased surface runoff and rapid
shore, lying close to but below fair-weather fluvial discharge into the basin following storm
wavebase in a moderately storm-influenced abatement.
setting. This zone reflects the transition from The heterolithic character of this facies and, in
the lower shoreface to the offshore/shelf (e.g. off- particular, the presence of the dark claystones
shore transition; cf. Howard & Reineck 1981). with synaeresis cracks suggests that the setting
The storm assemblage represents the activities lies along depositional strike, but down-
of opportunistic organisms re-colonizing the sub- longshore drift, of a contemporaneous delta
strate following storm disruption (Pemberton & complex (e.g. Coates & MacEachern 1999,
Frey 1984; Vossler & Pemberton 1988; Pember- 2000; Bhattacharya & Willis 2001; Bann &
ton & MacEachern 1997). The tops of the Fielding 2004).
sandstone units contain trace fossils that repre-
sent initial opportunistic colonization. The sand-
stones grade upward into sandy siltstones Facies 4: Thoroughly bioturbated muddy
containing a fair-weather resident trace fossil sandstone
suite that indicates a return to quiescent condi-
tions following storm abatement (Pemberton Sedimentology
et al. 1992a, 1992b; Pemberton & MacEachern This facies comprises intensely bioturbated, 90-
1997). Storm activity presumably disrupted 180cm thick muddy sandstone units, inter-
bottom fauna and probably resulted in mass bedded with minor, thin (<15cm), erosionally
stranding and the transportation of organisms based, fine- to medium-grained sandstone beds.
to other environments (Rees et al. 1977; Dobbs Some sandstone beds display low-angle, undula-
& Vozarik 1983; Butman 1987). The large- tory, parallel lamination, interpreted as HCS and
diameter living tubes of Diplo crater ion wave ripple cross-lamination. Shell fragments,
parallelum and large Rhizocorallium irregulare dispersed pebbles and granules, carbonaceous
(type A) are commonly filled with cleaner and/ detritus and allochthonous logs are locally
or coarser sediment within some assemblages of common.
the mixed Skolithos-Cruziana ichnofacies. The
coarse infills of these structures are interpreted Ichnology
as 'tubular tempestites' (Wanless et al. 1988; Facies 4 is characterized by pervasive bioturba-
Tedesco & Wanless 1991), and provide good tion, with intensities that vary from BIS to
evidence for the prior existence of storm beds. BI6, typically BIS (Fig. 7a, b). Bioturbation
The diversities of the trace fossil assemblages intensity is more or less uniform. Most ichno-
generally decrease in response to increases in genera are uniformly distributed.
storm intensity and/or frequency, and with The trace fossil assemblage is dominated by
increasing deltaic influence. Thin, unburrowed Rosselia socialis, Phycosiphon, Planolites, Diplo-
claystone drapes with synaeresis cracks occurring craterion habichi, Rhizocorallium irregulare,
at the tops of tempestites at some localities may Teichichnus, Rhizocorallium irregulare (type A),
ESTUARINE VS FULLY MARINE DEPOSITS 189

Fig. 7. (a) Thoroughly bioturbated (BI5) silty sandstone with clay laminae (Fades 4). Proximal expression of
the Cruziana ichnofacies including Phycosiphon (Ph), Palaeophycus (Pa), Chondrites (Ch), Teichichnus,
Diplocraterion (D), Helminthopsis (H) and Planolites. Fig. 15, 11-13m. (b) Thoroughly bioturbated (BI5) silty
sandstone with clay laminae (Facies 4). Proximal expression of the Cruziana ichnofacies including Palaeophycus
(Pa), Phycosiphon, Planolites (P), Teichichnus (T), Chondrites (Ch), Rhizocorallium (Rh) and Psammichnites
(Ps). Shell debris (sh) is locally present. Fig. 15, 11-13m. (c) Moderately to intensely bioturbated (BI3-5)
sandstone (Facies 5) with Rosselia rotatus (Ro) containing Teichichnus-\ike expressions of the living tube (Te).
Fig. 1, south Pebbley Beach, (d) Sandstone (Facies 5) with abundant Teichichnus (Te) that represent the lower
tubular portion of Rosselia socialis and R. rotatus. Fig. 1, Mill Point, (e) Plan view of the same bed figured in
(d) showing a cross-section through the associated Rosselia mud balls (Ro).

Macaronichnus isp., Palaeophycus tubular is and Interpretation


Palaeophycus heberti. Common but subordinate The dominance of sharp-based sandstone beds in
elements are Taenidium (type A), larger, less sin- Facies 4 together with the sedimentary structure
uous Taenidium (type B), sandy, robust IZoo- and presence of marine fossils indicates a shallow
phycos, Skolithos, fugichnia, Psammichnites and marine environment of deposition frequently
Chondrites, affected by storms. Rare, thin, sharp-based
190 K. L. BANN ET AL.

Fig. 8. Fades 5. (a) Weakly bioturbated (BI1-2) HCS and wave rippled sandstone, siltstone and thin, dark
claystone. The trace fossil suite comprises Diplocraterion habichi (D), Planolites (P), Rosselia (R) and fugichnia
(fu), corresponding to a proximal expression of the Cruziana ichnofacies. Fig. 14, 14.5-16.2m. (b) Plan view of
Diplocraterion habichi (D) in a tempestite. Fig. 1, Mill Point, (c) Moderately well bioturbated (BI4) sandstone
showing remnant low-angle, undulatory, parallel lamination and clay interlaminae. The trace fossil suite reflects
a proximal expression of the Cruziana ichnofacies, and comprises Planolites, Rosselia (R), Phycosiphon,
Chondrites, Helminthopsis, Teichichnus (Te), Diplocraterion and Palaeophycus. Fig. 1, Mill Point, (d) Very large
Rhizocorallium irregulare (Rh) and Diplocraterion habichi (D) in a tempestite bed. Fig. 1, Mill Point,
(e) Non-burrowed to weakly bioturbated (BIO-1), HCS and wave-rippled sandstone and dark claystone.
ESTUARINE VS FULLY MARINE DEPOSITS 191

HCS sandstone beds represent the deposits of are dark, organic-rich (carbonaceous detritus),
particularly severe storms. The trace fossil generally silt-poor, and contain common to
suite, comprising a diverse mixture of robust, abundant, short synaeresis cracks.
complex deposit- and detritus-feeding structures,
is a proximal expression of the Cruziana ichno- Ichnology
facies. Facies 4 is interpreted to reflect deposition This facies is characterized by highly variable
in a well-oxygenated, open marine setting at or bioturbation intensities, ranging from BIO to
just above fair-weather wavebase, consistent BI4. Laminated sandstone beds, interpreted
with the distal lower shoreface. The intensity as tempestites, range from BIO to BIS, and
and uniformity of burrowing, and the scarcity typically BI2. Bioturbated muddy sandstone
of preserved primary sedimentary structures, beds range from BI3 to BI4, typically BI4. Inter-
suggest considerable time breaks between vening thin claystone beds are generally non-
storms, during which the fair-weather fauna burrowed but rarely have a BI of 1. Bioturbation
reworked the substrate. The presence of consid- is sporadically distributed in the facies and 'Lam-
erable numbers of vertical burrows, such as Scram' bedding is common.
Diplocraterion habichi in some localities (Fig. The trace fossil assemblage is dominated by
7a), suggests that storm beds were rapidly colo- Rosselia socialis, R. rotatus and Rosselia (type
nized by opportunistic, suspension-feeding A) that exhibits a lateral shift of the mud-ball
organisms, prior to their re-colonization and and contains lateral spreiten (Figs 7c-e, 8a, c,
thorough reworking by the resident fair-weather h; Bann 1998). All forms of Rosselia may occur
community. with or without Teichichnus-\ikQ expressions of
the dwelling tube. Phycosiphon (Fig. 9a), Diplo-
craterion habichi (Fig. 8b), Rhizocorallium irregu-
Facies 5: Interbedded laminated sandstone, lare (type A, Fig. 8d), Diplocraterion parallelum
bioturbated muddy sandstone and dark (Fig. 9c, d), Macaronichnus isp. and fugichnia
clay stone are also abundant. Common but subordinate
elements comprise Palaeophycus tubularis, Taeni-
Sedimentology dium (type A), Skolithos, Conichnus (Fig. 9e),
This facies is composed principally of tabular to Psammichnites (Fig. 9f) and sandy, robust IZoo-
lensoidal sandstone beds, typically 0.2-0.5m phycos (Fig. 8f, g). Uncommon elements include
thick. These beds can be erosionally amalga- Lingulichnus (Fig. 9h), Chondrites, Macaron-
mated, or interbedded with bioturbated muddy ichnus segregatis, Taenidium (type B) and Cylin-
sandstone, or in some cases separated by thin drichnus. Fragmented Rosselia mud balls are
(0.2-6.0 cm), dark claystone partings. A charac- present within the tempestites, recording ero-
teristic feature of this facies is the presence of sional truncation, reworking and deposition as
interbedded lenses or thin beds of very coarse- clasts. The claystone interbeds contain small,
grained to granular/pebbly sandstone, the tops rare Planolites (Fig. 8e).
of which form sets of large (height 2-5 cm, wave- The assemblage, overall, is interpreted to
length 15-20 cm), symmetrical, gravelly ripples. reflect a diverse expression of the proximal
Some of these beds are internally composed of Cruziana ichnofacies (Fig. 9b).
a single set of unidirectional cross-stratification.
Most other sandstone beds show one or more Interpretation
of the following: well-preserved HCS, low-angle The strongly heterolithic character of Facies 5
planar cross-stratification, small symmetrical reflects marked variations in physical energy
wave ripples and combined flow ripple cross- and sedimentation rates. Most of the physical
lamination. The muddy sandstone beds resemble structures reflect storm deposition, particularly
those of Facies 4. Shelly debris along with dis- HCS, low-angle planar cross-stratification, com-
persed pebbles and granules and carbonaceous bined-flow and wave ripples. The laminated
detritus are locally common. Claystone interbeds sandstone units record episodic colonization

Carbonaceous detritus occurs within the sandstone beds. The trace fossil suite consists of diminutive Planolites
(P), Chondrites (Ch) and Phycosiphon (Ph). Note that, despite the presence of Phycosiphon, small synaeresis
cracks (sy) are present in the claystone. Fig. 14, 18.4-19 m. (f) Bedding plane view of a tentatively identified
Zoophycos (Z) comparable to that shown in cross-section in photo (g). The structure occurs in a silty HCS
sandstone bed interpreted as a tempestite. Fig. 15, 12.9-14.9 m. (g) Weakly bioturbated (BI2), HCS sandstone,
siltstone and dark claystone with a bedding plane expression of the structures displayed in photo (f). The
structure (a cone-like, sand-filled depression) is tentatively identified as Zoophycos (Z). Fig. 14, 14.5-16.2m.
(h) Example of a complex Rosselia rotatus (R). Fig. 1, south Pebbly Beach.
192 K. L. BANN ET AL.

Fig. 9. Trace fossils in Fades 4 and 5. (a) Phycosiphon (Ph) in a tempestite. (b). Idealized graphic
representation of characteristic trace fossils in lower shoreface deposits in the Pebbley Beach Formation. The
distal lower shoreface is characterized by a diverse suite of structures produced by complex deposit- and
detritus-feeding behaviours and specialized grazing behaviours. The vertical burrows of opportunistic
suspension feeders are associated with rare, thin tempestites. The overall trace fossil assemblage represents a
proximal expression of the Cruziana ichnofacies. The delta-influenced proximal lower shoreface is characterized
by interbedded tempestites, bioturbated fair-weather deposits and thin, unburrowed claystone beds with
synaeresis cracks. The trace fossil assemblage comprises a diverse mixture of robust, complex detritus- and
deposit-feeding structures, abundant fugichnia and the burrows of opportunistic suspension feeders, and
ESTUARINE VS FULLY MARINE DEPOSITS 193

Fig. 10. (a) Fades 4, bioturbated (BI5) silty sandstone erosionally truncated by a transgressive surface of
erosion (TSE) and overlain by trough cross-bedded, medium-grained sandstones of Facies 6. The cross-bedded
sandstone shows a BI of 2, and contains moderate numbers of Diplocraterion habichi (Dh), reflecting the
Skolithos ichnofacies. Fig. 1, Mill Point, (b) Moderately bioturbated (BI 3-4) sandstone (Facies 6). The unit
contains abundant, elongate Diplocraterion habichi (Dh) and reflects the Skolithos ichnofacies. Fig. 15,
18.4-19m.

following rapid sand emplacement, with higher environment downdrift of a contemporaneous


proportions of fugichnia, lesser numbers of delta complex.
trace fossils, but otherwise ichnological suites
comparable to the muddy sandstone beds. In
contrast, the claystone interbeds show highly Facies 6: Bioturbated, laterally variable
reduced bioturbation intensities and restricted sandstone facies
suites of the Cruziana ichnofacies, probably
reflecting stressful environmental conditions. Sedimentology
Facies 5, overall containing a diverse Cruziana This facies is composed of sandstone beds, 40-
ichnofacies, is consistent with sedimentation 100cm thick, with minor granule-, pebble- and
within the lower shoreface. The unburrowed shell-rich horizons. Most beds have erosional
claystone interbeds are interpreted as post- basal contacts (Fig. lOa). Facies 6 also shows
storm mud drapes, and are believed to be asso- considerable lateral variability at the outcrop
ciated with heightened precipitation, increased level. Individual sandstone beds are, in some
surface runoff at the coast, and rapid fluvial cases, intensely bioturbated, whereas others
discharge through distributaries from nearby show HCS and trough cross-bedding (Fig. lOa).
delta lobes (as described in Facies 3). Such Cross-sets show sigmoidal foresets with mud-
delta lobes presumably lie along depositional stone partings on foresets. Both small-scale and
strike and updrift of the facies. Gravelly large, gravelly, symmetrical wave ripples (as
symmetrical-rippled beds are believed to record for Facies 5) are also common. Some coarser-
periods of minimal sediment supply, allowing grained sandstone beds comprise a single cross-
winnowing of sediment by waves and concentra- set, whereas others preserve co-sets of sigmoidal
tion of the coarse fraction. Facies 5 is interpreted cross-beds. This facies differs from Facies 4 and 5
as the product of sediment accumulation in in that it displays a greater range of grain
an open marine proximal lower shoreface sizes, a broader range of physical sedimentary

represents a slightly more diverse expression of the proximal Cruziana ichnofacies. The deltaic influences on the
depositional environment force the trace fossil assemblage to remain Cruziana in expression rather than
shifting to a distal expression of the Skolithos ichnofacies, as would be expected in a non-delta-influenced
proximal lower shoreface environment. Rosselia socialis (Rs), R. rotatus (Rr), Rosselia (type A, Rm),
Phycosiphon (P), Planolites (PI), Diplocraterion habichi (Dh), D. parallelum (Dp), Rhizocorallium irregulare
(Rh), Teichichnus (T), Macaronichnus isp. (M), M. segregatis (Ms), Palaeophycus tubularis (Pt), P. heberti (Ph),
Taenidium (type A, Ts), sandy, robust ?Zoophycos (Z), Skolithos (S), Psammichnites (Ps), Chondrites (Ch),
Conichnus (C), Lingulichnus (L), Taenidium (type B, Ta), Cylindrichnus (Cy), fugichnia (fu), Lam-Scram (LS),
synaeresis cracks (sy), truncated Rosselia mud balls (tR). (c) Plan view of large Diplocraterion parallelum (D).
(d) Vertical section through a large Diplocraterion parallelum (D). (e) Vertical section through a sand-filled
Conichnus (C). (f) Plan view of Psammichnites (Ps). (g) Vertical section through Diplocraterion habichi (D).
(h) Plan view of Lingulichnus (Li). Examples c-h are from Mill Point.
194 K. L. BANN ET AL.

structures, and greater variety in the intensity a variety of water depths. The well-preserved
and character of bioturbation. Palaeocurrent cross-bedding and other structures suggest shal-
data from cross-beds indicate a predominantly low, regularly agitated water above fair-weather
west to northwest sediment dispersal direction. wavebase for the most part. The overall trace
Shell material occurs as dispersed fragments fossil assemblage is dominated by burrows of
and disarticulated valves, or concentrated opportunistic suspension-feeding organisms,
pebbly lags largely composed of robust articu- with subordinate detritus- and deposit feeders,
lated Eurydesma hobartense. Other taxa present and is interpreted to reflect a distal expression
include shallowly burrowing bivalves such as of the Skolithos ichnofacies. The firmground
Megadesmus, Pyramus, Schizodus and Stutch- assemblages subtending from underlying erosion
buria, vagrant epifaunal forms including Aviculo- surfaces are characteristic of the Glossifungites
pecten and Peruvispira, the bellerophont ichnofacies, and indicate infaunal colonization
Warthia, the spiriferid brachiopod Ingelarella of firm but unlithified substrates during periods
and a biplicate species of the terebratuloid Gille- of depositional hiatus. Eurydesma hobar tense is
dia (Runnegar 1979). Carbonaceous detritus, interpreted as an opportunistic species that flour-
including allochthonous logs, and large (up to ished on silt-free, current-swept, sublittoral
2m diameter), ovoid concretions are locally substrates (Runnegar 1979). Units that directly
common. overlie Facies 6 (typically the fine-grained
marine Facies 1, 2 and 3) tend to reflect deposi-
Ichnology tion in significantly deeper (more distal) deposi-
This facies is characterized by highly variable tional environments than those that underlie it.
bioturbation intensities, ranging from BI2-5. This suggests that the erosion surfaces beneath
Cross-bedded sandstone units tend to be less Facies 6 units represent transgressive surfaces
thoroughly bioturbated (BI2-3), with sporadi- of erosion (TSE). This facies is therefore inter-
cally distributed burrows. The trace fossil suite preted to represent transgressive sand sheets
consists of Diplocraterion parallelum, D. habichi that underwent winnowing and concomitant
(Fig. lOa, b), Rosselia socialis, Macaronichnus concentration of coarse-grained detritus and
isp. and Skolithos. shelly debris during periods of rising relative
The pebbly, shelly lags and pebbly sandstone sea-level. The composite nature and lateral
beds range from BI3—5, and typically BI5. The variability of this facies suggests that it reflects
trace fossil assemblage is dominated by Diplocra- deposition in various well-oxygenated, sedi-
terion habichi. Common but subordinate forms ment-starved, shallow marine settings. Predomi-
include Diplocraterion parallelum, Phycosiphon, nantly westward sediment dispersal direction is
Rosselia socialis, Planolites, Teichichnus and also consistent with deposition under trans-
sandy, robust IZoophycos. gressive conditions.
Some sandstone beds are intensely bioturbated
(BI4-5), with a diverse trace fossil assemblage
dominated by Diplocraterion habichi, D. paralle- Estuarine facies
lum and Rhizocorallium irregulare (type A).
Common but subordinate elements include Facies 7: Channelized, heterolithic
fugichnia, Phycosiphon, Rosselia socialis, R. rota- sandstone-mudstone
tus (with or without Teichichnus-like expressions
of the dwelling tube), Teichichnus, Planolites, Sedimentology
Palaeophycus tubularis, Macaronichnus isp. and This facies comprises interlaminated and thinly
Skolithos. interbedded siltstone and fine-grained sandstone
Most occurrences of this facies directly overlie beds, confined to channelized bodies up to 6m
erosional discontinuities, associated with firm- thick and typically 100-250 m wide, though com-
ground trace fossil assemblages. These assem- posite bodies (Facies 7/9: see below) may reach
blages comprise vertical, unlined, and passively several hundred metres in apparent width.
infilled domichnia that penetrate into the under- These bodies incise through all other facies,
lying facies and cross-cut the underlying trace including other Facies 7 units. Successions
fossil assemblage. pinch out laterally into thin (a few cm), laterally
persistent beds of siltstone with fine-grained
Interpretation sandstone laminae (which are representatives of
The presence of marine invertebrates again sug- Facies 8). Channel bases are typically marked
gests an open marine environment of deposition by thin, laterally discontinuous but persistent
for Facies 6, but the range of sedimentary struc- lags of well-rounded granules and pebbles, with
tures and lithologies points towards deposition in common coalified log casts (some petrified) and
ESTUARINE VS FULLY MARINE DEPOSITS 195

other carbonaceous plant debris. Lags pass least moderately sinuous. The predominance of
upward abruptly into the heterolithic facies linsen, lenticular and wavy bedding indicates
noted above, which typically show little system- that subequal proportions of sand and mud
atic upward change in lithology. The basal one were transported through the channels by uni-
metre or so of the channel bodies are locally directional flows of modest strength, and the
composed of trough cross-bedded, medium- to resulting bedforms were later modified by low-
very coarse-grained sandstone. Palaeocurrent energy waves in some cases. The bipolar distri-
measurements display a bipolar distribution of bution of ripple migration directions, and the
bedform migration directions. The heterolithic presence of mud drapes and rhythmic sand-
facies typically display 50:50 sand:silt ratios, mud couplets, confirm that tidal currents were
with individual beds ranging up to 5cm thick. active in the channels. The low-diversity, low-
The facies shows linsen, lenticular and wavy abundance trace fossil assemblage is extremely
bedding (Fig. 11 a, c). Sandstone beds contain a impoverished and represents a very restricted
spectrum of interlamination structures ranging expression of the mixed Skolithos-Cruziana ich-
from linsen (pinstripe), through lenticular and nofacies that is characteristic of inshore coastal
wavy bedding to flaser bedding, Ripple cross- environments of deposition. These features
lamination is predominantly unidirectional with contrast strongly with the characteristics of the
some evidence of wave modification (symmetri- other heterolithic facies interpreted to have
cal sand drapes over unidirectional cross- formed in offshore and offshore transition envir-
lamination sets. Furthermore, the migration onments.
directions of current ripples define a markedly Bipolar current flow, suggesting reversing tidal
bipolar distribution in most outcrops. Some flows in otherwise low-energy settings, is charac-
ripple sets are overlain by a continuous mud teristic of inshore settings, and uncommon in the
drape (e.g. Fig. 11 a) and rhythmically inter- offshore; tidal flow in distal settings is typically
laminated sand-mud couplets are also common rotary rather than reversing. The impoverished
(Fig. lib). Some intervals show convolute ichnological suites are typical of such estuarine
bedding and other soft-sediment deformation settings, which are subjected to highly variable
structures, including synaeresis cracks (Fig. and generally reduced salinities. The abundant
11 a). Another characteristic feature of this and persistent synaeresis cracks also support a
facies is inclined heterolithic stratification brackish-water interpretation. Facies 7 is there-
(IHS). At the southern end of Point Upright, a fore interpreted to reflect active fill of laterally
channel with IHS also shows flat lying, discor- migrating, estuarine channels.
dant stratification (intrasets) between through-
going inclined bedding surfaces. Palaeocurrent
data indicate broadly westward and eastward Facies 8: Sheet-like, heterolithic sandstone—
sediment dispersal. mudstone
Ichnology Sedimentology
This facies is characterized by very low bioturba- This facies resembles Facies 7 except that it
tion intensities, ranging from BIO-1. Trace occurs as tabular units, rather than being con-
fossils are sporadically distributed and occur in fined to channels, it lacks IHS, it contains less
very low numbers. Diversity of ichnogenera is plant debris, and it lacks both the large log
extremely low. Sandier expressions of the hetero- casts and the coarse lags seen in Facies 7.
lithic facies are largely devoid of bioturbation. Many of the ripple-scale structures show
There are no ichnological differences between evidence of wave modification in the form of
intervals that clearly form inclined heterolithic symmetrical drapes and bidirectional cross-
stratification (IHS) and those that do not. The lamination superimposed on unidirectional sets.
assemblage consists of Planolites (Fig. lib, c), Micro-HCS is common, suggesting the increased
rare fugichnia and rare Skolithos and is incidence of combined flow processes. Palaeo-
extremely impoverished. currents are generally bimodal and broadly
bipolar, but this reflects wave-modified current
Interpretation ripple cross-lamination as well as a bipolar distri-
Facies 7 is interpreted to reflect fine-grained bution of current ripple directions. Mud drapes
deposition within estuarine channels. IHS is and rhythmic sand-mud couplets are also
interpreted as lateral accretion surfaces asso- common (Fig. lid). Synaeresis cracks and soft-
ciated with tidally modified channel flow sediment deformation structures are generally
(Howard et al 1975; Thomas et al 1987; Shanley less common than in Facies 7 but are locally
et al. 1992), indicating that the channels were at common (Fig. lie).
196 K. L. BANN ET AL.

Fig. 11. (a) Fades 7. Non-burrowed (BIO) wavy bedding with oppositely oriented current ripples, and thick
mudstone drapes. Synaeresis cracks (sy) are locally developed within some mudstone interbeds. Fig. 15,
20.2-21 m. (b) Facies 7. Very weakly bioturbated (BI1) wavy bedded sandstone and mudstone. Unit shows
undulatory parallel drapes of silt and clay as well as oppositely oriented ripples. Diminutive Planolites (PI) is
the only visible trace fossil. Synaeresis cracks occur locally. Fig. 15, 20.2-21 m. (c) Facies 7. Sandstone-
dominated wavy bedding, showing current ripple lamination, some combined flow ripple lamination and
parallel-laminated drapes. Bioturbation intensities reach BI 1-2, with some evidence of soft-sediment
deformation in the lower part of the photo. The assemblage consists of Planolites (PI) and exceedingly rare,
diminutive Skolithos. Synaeresis cracks are common. Fig. 15. 22.2-25m. (d) Facies 8. A flaser-bedded interval
of reactivated current ripples and combined flow ripples, with some evidence of aggradation. The facies shows
low bioturbation intensities (BI 1), manifest by isolated, diminutive Planolites (PI). Small synaeresis cracks (sy)
are locally developed. Fig. 14, 5.5-6.6m. (e) Facies 8. Lenticular- and linsen-bedded sandstone and mudstone.
The unit shows very low bioturbation intensities (BI 1), consisting of diminutive Planolites (PI). Synaeresis
cracks (sy) are common. Fig.l, north Mill Point, (f) Facies 8. Mudstone-dominated interval with siltstone and
sandstone ripples and rare synaeresis cracks (sy). The unit shows moderate bioturbation intensities (BI 3),
consisting of Planolites (PI), Teichichnus (T) and Cylindrichnus (Cy), and reflects a low-diversity expression of
the Cruziana ichnofacies. Fig. 1, south Mill Point.
ESTUARINE VS FULLY MARINE DEPOSITS 197

Fig. 12. Fades 8. (a) Sandstone dominated, wavy to flaser bedding abruptly overlying Fades 1 mudstones,
marking the position of a sequence boundary (SB). The overlying heterolithic unit shows remnant wave,
combined flow and rarer current ripple lamination. This interval is moderately to abundantly bioturbated
(BI3-4), and displays a suite comprising Palaeophycus (Pa), Planolites (P), Rosselia (Ro), Teichichnus (Te) and
unnamed equilibrium-adjustment burrows (e-a) reflecting a low-diversity expression of the Cruziana
ichnofacies. Note that some of the equilibrium-adjustment burrows subtend across the sequence boundary,
reflecting a palimpsest softground omission suite. Fig. 1, north Mill Point, (b) Bedding plane view showing
well-developed equilibrium-adjustment burrows (e-a), Planolites (P) and Taenidium (type B, Ta). Bioturbation
intensity is BI3. Fig. 15, 1-4m.

Ichnology that deposition occurred in a permanently sub-


Fades 8 displays highly variable bioturbation aqueous setting (i.e. not an intertidal flat).
intensities ranging from BI 0 to BI 4. Trace fossils Reversing, tidal currents are indicated by less
are sporadically distributed, and some ichnogen- common, opposed current ripples, mud drapes
era are confined to local outcrop areas, though and rhythmic sand-mud couplets.
most are persistent within the facies. Suites are The trace fossil assemblage, although moder-
locally moderately diverse, containing Planolites ately diverse, is sporadically distributed and is
(Fig. llf), Skolithos, fugichnia, Psammichnites, characterized by structures that reflect simple
Teichichnus, Conichnus, unnamed equilibrium- deposit-feeding behaviours, with less intercalated
adjustment structures (Fig. 12a, b), 1 Rosselia suspension-feeding behaviours that are typical of
socialis, possible Diplo crater ion parallelum, opportunistic suites. The sporadic presence of
Taenidium (type B, Fig. 12 b), Siphonichnus, IZoophycos at specific horizons within the
Cylindrichnus, and large, sandy, cone-shaped facies suggests short-lived, episodic development
IZoophycos. of fully or near-fully marine conditions in an
otherwise brackish-water setting. The suite
Interpretation corresponds to a highly stressed expression of
This facies is dominated by wave ripples, wave- the Cruziana ichnofacies, with moderately
modified current ripples and combined flow diverse expressions of the mixed Skolithos-
ripples (micro-HCS). There is no evidence to sup- Cruziana ichnofacies sporadically distributed
port periods of subaerial exposure, suggesting throughout. The local abundance of synaeresis
198 K. L. BANN ET AL.

Fig. 13. (a) Soft-sediment deformed inclined heterolithic stratification (IHS) of Facies 7 (lower third of photo),
capped by dark mudstone of Facies 9. The mudstones are truncated by a transgressive surface of erosion (TSE)
and overlain by bioturbated (BI4) silty sandstone of Facies 5. The TSE is ichnologically demarcated by a
firmground trace fossil assemblage that consists of unlined, robust, passively filled domichnia. The assemblage
contains Diplocraterion habichi (Dh), Skolithos and Planolites and represents the Glossifungites ichnofacies.
The photographed interval is approximately 50cm high. Fig. 1, north Point Upright, equivalent to Fig. 14,
14-14.5m. (b). Finely laminated dark silty mudstone (Facies 9). The facies is non-burrowed (BIO). Fig. 14,
11.5-14.4m. (c). Dark, non-burrowed (BIO) mudstone erosionally truncated by a transgressive surface of
erosion (TSE) and overlain by bioturbated (BI4) Facies 6. The TSE is ichnologically demarcated by a
Glossifungites ichnofacies. Diplocraterion habichi (Dh). Fig. 15, 4.3m. (d) Plan view of robust, unlined, sharply
outlined Diplocraterion habichi (Dh) that represent the Glossifungites ichnofacies. The burrow infill is
composed of coarse-grained sand from the overlying unit, and the structures stand out clearly from the
mudstone host facies. Fig. 1, south Pebbly Beach. (E) Robust, sharply outlined, passively filled Arenicolites (Ar)
subtending from a TSE into an underlying sandstone bed. This burrow represents colonization of a palimpsest
softground during transgression. Fig. 1, north of Clear Point.
ESTUARINE VS FULLY MARINE DEPOSITS 199

cracks associated with the facies is also consistent accumulation in estuarine channel and basin
with generally brackish-water conditions. The environments, in which tidal currents played an
facies is interpreted to reflect deposition in pro- important role and waves were generally sub-
tected, wave-influenced and periodically tidally dued. This implies a highly indented coastal
influenced brackish-water basins that formed planform, with perhaps large funnel-shaped
laterally adjacent to the estuarine channels of estuaries crossed by sinuous channels that were
Facies 7. flanked by (or drained into) shallow-water
basins. The lack of high-energy wave structures
in the estuarine facies suggests that estuaries
Facies 9: Laminated mudstone may have been barred, and this notion is also
favoured by the presence of Facies 6, recording
Sedimentology transgressive complexes that may have origi-
This facies consists of laminated to blocky, nated as estuary-mouth barriers that migrated
medium to dark grey claystone and clayey silt- inboard during transgressions (cf. Dalrymple
stone, occurring either as channel fill or as et al. 1992). A summary of the ichnological
tabular units (Fig. 13a, b). The mud-filled differences between the estuarine and offshore
channel deposits are lateral equivalents to facies is illustrated in Figure 16a-c.
Facies 7 and form part of the channel complexes Similar arrays of facies to those recorded
described above, with a fill geometry that is here, and comparable interpreted depositional
broadly form-concordant with the basal erosion settings, are presented by Shanmugam et al.
surface. Tabular mudstone units typically overlie (2000), Beets et al. (2003) and Takano &
Facies 8. Thin (<3cm thick) sandstone and Waseda (2003).
siltstone layers with linsen (pinstripe) lamination
are persistent, and locally normally graded.
Small pyrite concretions and coaly plant debris Sequence stratigraphy of the Pebbley Beach
are locally present, but rare. At Point Upright Formation
the mudstone facies occurs within a channel
with a well-developed basal clast layer. The vertical stacking patterns of facies in the
Pebbley Beach Formation pose some consider-
Ichnology able challenges to sequence stratigraphic inter-
Bioturbation is absent (BIO) except where the pretation (Fig. 16d, e, Fig. 17). Nevertheless, a
mudstone is truncated by a discontinuity surface series of unconformity-bounded, cyclical stratal
(Fig. 13c). packages can be recognized from the vertical
arrangement of facies. In the part of the Pebbley
Interpretation Beach Formation under consideration, these
The complete absence of bioturbation suggests cycles are < 10 m thick, and thus could represent
anaerobic and/or reduced salinity conditions. high-frequency (fourth- to sixth-order) para-
Robust, passively filled domichnia that cross- sequences (Van Wagoner et al. 1988; see Naish
cut the mudstone are characteristic of the Glossi- & Kamp 1997), intermediate-scale packages
fungites ichnofacies and indicate depositional (e.g. Swift et al. 2003), or true sequences
hiatus and colonization of a firm but unlithified formed in a low-accommodation setting (e.g.
substrate (see Fig. 13d, e for other examples of Kidwell 1997; Fielding et al 2000). Given the
omission suites and Glossifungites ichnofacies long period apparently recorded by the Pebbley
in the Pebbley Beach Formation). Facies 9 is Beach Formation (c. 14 Ma, according to
interpreted to reflect deposition in abandoned Briggs 1998), it seems likely that the cycles
estuarine channels and basins. under consideration are true sequences (third-
order or lower).
Regardless of the differing timeframes over
Summary of depositional environment which these cycles may have formed, many of
the examples cited above show a condensed char-
The middle and upper parts of the Pebbley Beach acter, with little preservation of lowstand systems
Formation are interpreted to record a range of tracts and considerable erosional truncation of
coastal and nearshore marine depositional envir- the highstand systems tract. This pattern is also
onments (Figs 14, 15). Open marine facies (1-5) evident in the Pebbley Beach Formation, where
record lower offshore to lower shoreface water most sequences show evidence of significant
depths, with shallower shoreface and shoreline erosion during the falling limb of relative sea-
facies largely absent (see Sequence stratigraphy level cycles. The greatest degree of erosion is
below). Coastal facies (7-9) record sediment associated with the incision of the estuarine
200 K. L. BANN ET AL.

Fig. 14. Graphic log of the Pebbley Beach Formation at Point Upright.

channels. The amount of section removed varies facies from previous cycles adds to the challenge
locally, in some cases an entire sequence appar- of developing a realistic and useful sequence
ently removed and the estuarine channel fill stratigraphic model.
juxtaposed above facies from an earlier deposi- The integration of ichnofacies analysis and
tional cycle. This juxtaposition of fine-grained sedimentology has provided a powerful tool for
coastal deposits onto lithologically similar the reliable discrimination between lithologically
ESTUARINE VS FULLY MARINE DEPOSITS 201

Fig. 15. Graphic log of the Pebbley Beach Formation at Clear Point.

similar but nonetheless disparate environments, suites, and in particular the Glossifungites ichno-
Ichnology in particular has proven to be invalu- facies, has facilitated the delineation and genetic
able in the discrimination of fully marine and interpretation of key surfaces, notably sequence
brackish estuarine deposits. In addition, the boundaries and transgressive surfaces (Fig.
recognition of substrate-controlled trace fossil 18a-e). Surfaces recognized in this study have
202 K. L. BANN ET AL.

Fig. 16. Summary diagram of the ichnological signature of brackish-water versus open marine deposits in the
Pebbley Beach Formation showing the differences in bioturbation intensity, uniformity versus sporadic
distribution of bioturbation and trace fossil assemblages. Block diagrams show idealized representation of the
characteristic trace fossil assemblage in (a) fine-grained fully marine and (b) estuarine deposits, (c) Trace fossil
assemblage key and indication of typical bioturbation intensity and uniform versus sporadic distribution.
See Figure 4 for an explanation of bioturbation intensity (BI). (d) Vertical view through heterolithic interval
illustrating the minimal differences in lithology between the offshore facies and the estuarine channel fill,
(e) Close-up view of the juxtaposition of heterolithic estuarine deposits onto heterolithic offshore units.
This surface is very subtle and in some instances marked by an omission suite of Diplocraterion habichi (D).
Fig. 15, 18-23 m.
Fig. 17. Photomosaic of IHS estuarine channel fill of Fades 7 (IHS) truncating underlying delta-influenced lower shoreface sandstone (LSF) and offshore transition
siltstones and sandstones of Facies 3 (OST). The base of the channel fill is interpreted as a transgressively modified sequence boundary (FS/SB). The estuarine channel
is capped by dark mudstone of Facies 9. The estuarine complex is truncated by a transgressive surface of erosion (TSE), overlain by heterolithic lower shoreface
sandstones and siltstones (LSF), which is truncated by another TSE and overlain by lower offshore deposits (LOS). Approximately 150 m long and 22 m high.
204 K. L. BANN ET AL.

Fig. 18. Discontinuity surfaces, (a) Thoroughly bioturbated (BI5) muddy siltstone (Facies 1, lo), overlain by
non-bioturbated, coarse-grained, channel base of Facies 7 (ec). The boundary between the two units is
ichnologically demarcated by Skolithos (Sk) of the Glossifungites ichnofacies, and represents an amalgamated
sequence boundary/flooding surface (i.e. FS/SB). The offshore siltstone contains a distal expression of the
Cruziana ichnofacies, dominated by diminutive Rosselia (Ro), Phycosiphon, diminutive Zoophycos (Z),
Helminthopsis, Teichichnus (Te) and Planolites (P). Fig. 15, 21.1-21.85m. (b) Thoroughly bioturbated (BI5),
muddy siltstone (Facies 1, lo), overlain by moderately bioturbated, wavy to lenticular bedded sandstone and
mudstone (eb, Facies 8). The boundary between the units contains grit-filled Conichnus (C) interpreted to
constitute part of the Glossifungites ichnofacies, and represents a FS/SB. Fig. 1, north Mill Point,
(c) Thoroughly bioturbated (BI 5), muddy siltstone (Facies 1, lo), overlain by moderately bioturbated, wavy to
lenticular bedded sandstone and mudstone (eb, Facies 8) with equilibrium adjustment structures (ea)
protruding across the sequence boundary. Fig. 1, north Mill Point, (d) Lenticular to wavy bedded sandstone
and silty mudstone (Facies 7, ec), overlain by thoroughly bioturbated (BI 5) muddy siltstone (Facies 1, the unit
seen in the basal half of photo A, lo). The boundary between the two units is marked by a thoroughly
bioturbated (BI 5) pebbly sandstone horizon that hosts a palimpsest suite of Diplocraterion habichi (D) and is
ESTUARINE VS FULLY MARINE DEPOSITS 205

been physically traced over the extent of cliff out- Skolithos also occur locally (Fig. 18a, b). The
crops and can be shown to be continuous over sandstone is overlain by heterolithic sandstone
the distance from Point Upright to Clear Point and siltstone. At other localities the sandstone
(c. 5km straight line distance, considerably is absent, and the erosive channel bases are
further around the coastline; Fig. 1). Over the directly overlain by either heterolithic fill or
vertical interval considered, eight sequences can dark grey siltstone (Facies 9). The heterolithic
be recognized. Of these, two (Sequences 2 and deposits probably represent backfilling of the
6) are not readily divisible into systems tracts, estuarine channels during ensuing sea-level rise.
but rather seem to record a stack of coarsening- The presence of firmground suites of trace fossils
upward parasequences. The other sequences, along the sequence boundaries adds strength to
however, display the highly condensed and trun- the argument for channel filling during rising
cated architecture described above. Sequences 4 sea-level (transgressive systems tract) as,
and 5 are highly complex, recording multiple although subaerial exposure and/or erosion
generations of channel incision at Mill Point during lowstand may generate widespread
(Fig. 18f), and may indeed each represent more dewatered or firm substrates, such surfaces are
than one sequence. Furthermore, these two unlikely to have become colonized unless they
sequences become amalgamated as they are were subsequently exposed to marine or
traced southward towards and along Point marginal marine conditions (Pemberton &
Upright (Fig. 19). MacEachern 1995). Sequence boundaries in the
Pebbley Beach Formation are therefore
amalgamated with marine flooding surfaces (i.e.
Sequence boundaries FS/SB).

Erosion surfaces at the base of the estuarine


facies are interpreted to be sequence boundaries. Transgressive surfaces
These discontinuities represent a significant
basinward shift of facies, and they are generated Discontinuity surfaces across which a significant
during lowstand in relative sea-level. deepening in depositional environment can be
In the Pebbley Beach Formation, heterolithic demonstrated are abundant in the Pebbley
estuarine channel and basin deposits commonly Beach Formation, and are interpreted as
directly overlie fine-grained or heterolithic off- transgressive surfaces. Transgressive surfaces
shore and offshore transition deposits. The occur either as largely non-erosive, low-energy
boundary between the two facies is locally marine flooding surfaces (FS) or as low-relief,
subtle, involving no significant change in lithol- high-energy transgressive surfaces of erosion
ogy and only modest changes in physical sedi- (TSE). Flooding surfaces in the Pebbley Beach
mentary structures (Fig. 16d). Nevertheless, Formation occur as sharp contacts across
profound changes in bioturbation intensity, which there is evidence of an increase in
sporadic versus uniform of bioturbation, and water depth. These surfaces are mantled locally
details of the trace fossil assemblages themselves, with dispersed granules and shelly material.
all serve to indicate significant changes in deposi- The surfaces also commonly host palimpsest
tional environment. softground suites of Diplocraterion habichi,
Estuarine channel bases locally contain which protrude down into underlying facies
coarse-grained, trough cross-bedded sandstone and cross-cut the original resident trace fossil
fill. Glossifungites ichnofacies assemblages and assemblage.
palimpsest softground suites consisting of Diplo- The degree of biogenic reworking of flooding
craterion habichi, Conichnus and subordinate surfaces varies locally. Where muddy siltstone

interpreted as a transgressive surface of erosion (TSE). Fig. 15, 20.8-21.3m. Lens cap is 5cm across,
(e) Transgressive surface of erosion veneered with shelly pebbly lag. This surface has abundant Diplocraterion
habichi (D) of the Glossifungites ichnofacies descending into the underlying, thoroughly bioturbated (BI5)
Facies 3. Fig. 15, l l . l m . (f). Outcrop view of the complex nature of the stacking patterns in the Pebbley Beach
Formation (sequences 3-6). Lenticular to wavy bedded Facies 8 (EB) are overlain (to the left of the photo) by
HCS Facies 5 (LSF). The boundary between these two basal units is interpreted as a transgressive surface of
erosion (TSE). These facies are truncated by lenticular to wavy bedded Facies 7 showing IHS and occupying a
channel. The base of this channel is interpreted as an FS/SB that amalgamates to the right with the underlying
TSE. The channel fill is truncated by a TSE that is overlain by burrowed silty sandstone (Facies 4). This is
truncated by another channel to the left that wedges out along strike to the north (right) and is truncated by
burrowed silty sandstone (Facies 4). See Figure 19, Mill Point south end.
Fig. 19. Proposed sequence stratigraphic framework of the Pebbley Beach Formation showing lateral continuity of facies.
ESTUARINE VS FULLY MARINE DEPOSITS 207

fades (lower offshore deposits) overlie intensely Implications of the sequence stratigraphic
bioturbated sandy siltstone fades (upper off- interpretation
shore deposits), the FS is generally not visibly
disturbed by the diminutive surface-grazing As suggested above, the thin and condensed
trace-makers that are characteristic of the lower sequences recognized in the upper Pebbley
offshore environment. However, where intensely Beach Formation indicate a continental margin
burrowed upper offshore sandy siltstones overlie environment where sediment accumulation in
sandy lower shoreface deposits, the FS is the nearshore realm was limited by accommoda-
generally more gradational in expression, largely tion. Sequence boundaries may be recognized
as a result of the complete reworking of the with care from a combination of lithological
contact by the comparatively more robust and ichnological criteria, but apart from local
deposit feeders that are abundant in the upper coarse-grained ?alluvial sandstones preserved at
offshore. the base of some channels, no lowstand systems
Transgressive surfaces of erosion in the tract deposits are evident. Transgressive systems
Pebbley Beach Formation occur as low-relief, tract facies are well preserved though typically
extensive discontinuity surfaces that show thin, and highstand systems tracts are typically
evidence of excavation by wave and current erosionally truncated by the overlying sequence
processes, associated with erosional shoreface boundary. This has led to a highly complex
retreat during transgression ('ravinement': Swift stratigraphic architecture with a dominance of
1975; Arnott 1995). These surfaces are generally heterolithic sandstone-mudstone facies, and no
veneered by a pebbly lag locally built into large thick sandstone bodies preserved (Fig. 19).
(height 3-5 cm, wavelength 15-20 cm), symmetri- One corollary of this architecture that would
cal ripples. The lags can also contain abundant not have been evident from previous analyses
shell material (Fig. 18e). Carbonaceous detritus of the unit is that significant volumes of shore-
including allochthonous logs is also locally face sand are missing from these sequences. We
common. suggest that, during falling sea-level, much of
Most TSE in the Pebbley Beach Formation the record of the previous coastal progradation
host passively filled, palimpsest suites of trace (perhaps 10-20 m vertical interval of dominantly
fossils that subtend from the discontinuity sand) must have been exported into greater off-
surface and cross-cut the underlying trace fossil shore, to the east of the exposures. Thus,
suite. Where the underlying units consist of although the exposed succession is of little
finer-grained mudstone and siltstone, the direct interest to hydrocarbon exploration
palimpsest suites reflect the Glossifungites owing to a lack of viable reservoir, it may provide
ichnofacies. In contrast, where the underlying vectors towards more substantial reservoir devel-
stratum is sandstone, a softground palimpsest opment down-palaeoslope to the east.
ichnological suite is more typically developed.
Suites of the Glossifungites ichnofacies in
the Pebbley Beach Formation consist of vertical Conclusions
to subvertical domichnia, produced by oppor-
tunistic, predominantly suspension-feeding The Pebbley Beach Formation in the southern
organisms during hiatus (erosional and/or non- Sydney Basin contains a mixture of:
depositional). The burrows are generally sharp- fully marine, fine-grained, moderately to inten-
walled, robust and unlined, reflecting the firm sely bioturbated offshore deposits;
but unlithified nature of the substrate at the moderately to intensely bioturbated sandy
time of colonization and burrow excavation. shoreface successions;
The passive nature of the burrow fills indicate weakly to moderately bioturbated, interlami-
that the structures remained open after the nated sandy and silty delta-influenced shore-
inhabitants had vacated them, and they were face successions; and
subsequently filled with sediment from the weakly to moderately bioturbated, generally
successive depositional event. The most abun- mostly fine-grained estuarine deposits.
dant element of the Glossifungites ichnofacies
is densely spaced Diplocraterion habichi, with Marine deposits contain abundant and diverse
subordinate Skolithos and Conichnus. trace fossil suites. Offshore deposits in the
Palimpsest softground suites in the Pebbley Pebbley Beach Formation contain highly diverse
Beach Formation are also dominated by Diplo- trace fossil assemblages that comprise a complex
craterion habichi. Diplocraterion parallelum and mixture of structures produced by deposit-
the lower, Teichichnus-like tubes of Rosselia are feeding and grazing/foraging behaviours and
subordinate elements. reflect the archetypal Cruziana ichnofacies.
208 K. L. BANN ET AL.

Vertical burrows that represent the dwellings of The middle and upper parts of the Pebbley
predominantly suspension-feeding organisms Beach Formation have been divided into a
(e.g. elements of the Skolithos ichnofacies) series of sequences, each reflecting a cycle of
correspond to the opportunistic colonization of fall and rise in relative sea-level. The sequences
distal tempestites. Intensely burrowed sandy are thin, condensed and top-truncated, but are
shoreface intervals in the Pebbley Beach Forma- nonetheless similar to some other published
tion contain trace fossil assemblages that contain examples of continental margin successions accu-
diverse mixtures of robust, complex deposit- and mulated under low-accommodation conditions.
detritus-feeding structures. The assemblages The basal sequence boundary for each sequence
reflect proximal expressions of the Cruziana is demarcated by a channelized erosion surface,
ichnofacies and are characteristic of well- in some cases with a coarse sandstone recording
oxygenated, open marine settings at or immedi- the lowstand systems tract. The overlying estuar-
ately above fair-weather wavebase. ine channel and/or basin deposits, transgressive
Delta-influenced shoreface successions are surface and fining-upward marine facies record
strongly heterolithic and contain sporadically the transgressive systems tract, and fine-grained
distributed, diverse trace fossil suites that to coarsening-upward nearshore marine facies
record proximal expressions of the Cruziana are erosionally truncated at the top by the next
ichnofacies. Unburrowed clay stone interbeds sequence boundary.
reflect heightened precipitation, increased sur- Key discontinuity surfaces in the Pebbley
face runoff at the coast, and enhanced fluvial dis- Beach Formation are generally ichnologically
charge through distributaries of nearby delta demarcated by palimpsest omission trace fossil
lobes that lie updrift and along depositional suites. Firmground assemblages reflecting the
strike. In general, the fully marine successions Glossifungites ichnofacies occur at the base of
in the Pebbley Beach Formation contain trace estuarine channel and basin facies, suggesting
fossil suites that display moderate to intense bio- that sequence boundaries in the Pebbley Beach
turbation, are characterized by a high diversity of Formation are amalgamated with marine flood-
forms, contain significant numbers of structures ing surfaces (i.e. FS/SB). Transgressive surfaces
reflecting specialized feeding/grazing behaviours, of erosion in the Pebbley Beach Formation are
and display uniform ichnogenera distributions, also demarcated by firmground suites consisting
all characteristics of equilibrium communities of vertical to sub vertical domichnia, produced by
within fully marine environments. opportunistic, predominantly suspension-feed-
Estuarine deposits in the Pebbley Beach ing organisms during periods of depositional
Formation contain extremely impoverished hiatus (erosional and/or non-depositional).
ichnological suites. In general, the facies show The re-evaluation of the middle and upper
variable but significantly reduced degrees of bio- Pebbley Beach Formation presented herein pro-
turbation intensity, pronounced variability in the vides a vivid example of the value of ichnology
distribution of individual ichnogenera, and the to stratigraphic analysis of nearshore marine to
dominance of a few, simple forms. The dominant coastal facies successions. The resulting sequence
elements represent simple feeding strategies of stratigraphic model is consistent with all field
resilient trophic generalists. data, and has predictive capabilities that may
Estuarine active channel deposits are sparsely be useful in the search for hydrocarbons in this
burrowed by trace fossil suites comprising less under-explored basin.
than three ichnogenera, and generally only one.
The low-diversity, low-abundance mixed Funding was supplied by a University of Queensland
Skolithos-Cruziana ichnofacies assemblages Research Fellowship to KLB and by a research grant
and the abundance of synaeresis cracks in these awarded to KLB and CRF by Oil Company of Austra-
lia Ltd and SANTOS Ltd. JAM was funded by
units reflect fluctuating salinity levels, episodic NSERC Operating Grant #184293. Professors Brian
deposition, and variability in substrate consis- Jones and Tony Wright are gratefully acknowledged
tency. Estuarine basin deposits contain slightly for insights into the Pebbley Beach Formation. R.
more diverse trace fossil assemblages, and some Higgs and R. McNaughton are thanked for their
suites are moderately diverse locally. The suites reviews.
can be regarded as impoverished marine
assemblages, and reflect a low-diversity expres-
sion of the mixed Skolithos-Cruziana ichno- References
facies. Estuarine abandonment deposits reflect
hostile bottom-water conditions (e.g. anaerobic ARNOTT, R. W. C. 1995. The parasequence definition:
and/or reduced salinity) that precluded are transgressive deposits inadequately addressed?
colonization. Journal of Sedimentary Research, B65, 1-6.
ESTUARINE VS FULLY MARINE DEPOSITS 209

BANN, K. L. 1998. Ichnology and sequence stratigraphy COATES, L. & MACEACHERN, J. A. 2000. Integrating
of the Early Permian Pebbley Beach Formation and ichnology and sedimentology to differentiate
Snapper Point Formation in the southern Sydney between river-dominated deltas, wave-dominated
Basin. PhD thesis. University of Wollongong. deltas and shorefaces: examples from the Cretac-
BANN, K. L. & FIELDING, C. R. 2004. An integrated eous of Western Canada. Geological Society of
ichnological and sedimentological comparison of America, Cordilleran Section, 96th Annual Meet-
non-deltaic shoreface and subaqueous delta ing, Vancouver, British Columbia, V.32, p. A7.
deposits in Permian reservoir units of Australia. CROWELL, J. C. & FRAKES, L. A. 1975. The Late
In: MclLROY, D. (ed.) The Application of Ichnology Palaeozoic Glaciation. In: CAMPBELL, K. S. W.
to Palaeoenvironmental and Stratigraphic Analysis. (ed.) Gondwana Geology. Australian National
Geological Society, London, Special Publications, University Press, Canberra, 313-331.
228, 273-307. DALRYMPLE, R. W., ZAITLIN, B. A. & BOYD, R. 1992.
BATTERSBY, D. G. 1981. New discoveries in the Surat/ Estuarine facies models: conceptual basis and
Bowen Basin. Australian Petroleum Exploration Stratigraphic implications. Journal of Sedimentary
Association Bulletin, 21, 39-44. Petrology, 62, 1130-1146.
BEETS, D. J., DE GROOT, T. A. M. & DAVIES, H. A. DICKENS, J. M. 1984. Evolution and climate in the Upper
2003. Holocene tidal back-barrier development Palaeozoic. In: BRENCHLEY P. (ed.) Fossils and
at decelerating rise: a 5 millennia record, exposed Climate, John Wiley & Sons, Chichester, 317-327.
in the western Netherlands. Sedimentary Geology, DOBBS, F. C & VOZARIK, J. M. 1983. Immediate effects
158, 117-144. of a storm on coastal ichnofauna. Marine Ecology
BEYNON B. M., PEMBERTON S. G, BELL, D. A. & LOGAN. Progress Series, 11, 273-279.
C. A. 1988. Environmental implications of ichno- EKDALE, A. A., BROMLEY, R. G. & PEMBERTON, S. G.
fossils from the Lower Cretaceous Grand Rapids 1984. Ichnology: Trace Fossils in Sedimentology
Formation, Cold Lake Oil Sands Deposit. In: and Stratigraphy. Society of Economic Paleon-
JAMES, D. P. & LECKIE, D. A. (eds) Sequences, tologists and Mineralogists, Short Course, Tulsa,
Stratigraphy, Sedimentology: Surface and Subsur- Oklahoma, 15.
face. Canadian Society of Petroleum Geologists, EYLES, C. H., EYLES, N. & GOSTIN, V. A. 1998. Facies
Memoirs, Calgary, Alberta, 15, 275-290. and allostratigraphy of high-latitude glacially
BHATTACHARYA, J. P. & WILLIS, B. J. 2001. Lowstand influenced marine strata of the Early Permian
deltas in the Frontier Formation, Powder River southern Sydney Basin, Australia. Sedimentology,
Basin, Wyoming, implications for sequence strati- 45, 121-161.
graphic models. American Association of Petro- FIELDING, C. R., NAISH, T. R., WOOLFE, K. J. &
leum Geologists Bulletin, 85, 261-294. LAVELLE, M. A. 2000. Facies analysis and strati-
BRIGGS, D. J. C. 1998. Permian Productidina and graphy of CRP-2/2A, Victoria Land Basin,
Stropholosiidina from the Sydney-Bowen Basin Antarctica. Terra Antarctica, 1, 323-338.
and New England Orogen: systematics and biostra- FREY, R. W. 1990, Trace fossils and hummocky cross-
tigraphic significance. Association of Australasian stratification, Upper Cretaceous of Utah. Palaios,
Palaeontologists, Memoirs, Sydney, Australia, 19. 5, 203-218.
BUATOIS, L. A., MANGANO, M. G., ALISSA, A. & CARR, FREY, R. W. & HOWARD, J. D. 1972. Georgia coastal
T. R. 2002. Sequence Stratigraphic and sedimento- region, Sapelo Island, USA. Sedimentology and
logic significance of biogenic structures from a late biology, VI. Radiographic study of sedimentary
Paleozoic marginal-to open-marine reservoir, structures made by beach and offshore animals
Morrow Sandstone, subsurface of southwest in aquaria. Senckenbergiana Maritima, 4, 169—182.
Kansas, USA. Sedimentary Geology, 152, 99-132. GOSTIN, V. A. & HERBERT, C. 1973. Stratigraphy of the
BUTMAN, C. A. 1987. Larval settlement of soft- Upper Carboniferous and Lower Permian
sediment invertebrates: the spatial scales of pattern sequence, southern Sydney Basin. Journal of the
explained by active habitat selection and the Geological Society of Australia, 20, 49-70.
emerging role of hydrodynamical processes. HERTWECK, G. 1970. Die Bewohner des Wattensmeeres
Oceanography and Marine Biology, Annual in ihren Auwirkungen das sediment. In: REINECK,
Review, 25, 113-165. H. E. (ed.) Das Watt, Ablagerungs- und Lebens-
CARR, P. F, JONES, B. G. & MIDDLETON, R. G. raum. Kramer, Frankfurt am Main, 106-130.
1989. Precursor and formation of glendonites HOLCOMBE, R. J., STEPHENS, C. J. et al. 1997. Tectonic
in the Sydney Basin. Australian Mineralogist, 4, evolution of the northern New England Fold Belt:
3-12. the Permian-Triassic Hunter-Bowen event. In:
CoAXES, L. & MACEACHERN, J. A. 1999. The ichno- ASHLEY, P.M. & FLOOD, P.G. (eds) Tectonics and
logical signature of wave- and river-dominated Metallogenesis of the New England Orogen. Geo-
deltas: Dun vegan and basal Belly River Forma- logical Society of Australia, Special Publications,
tions, West-Central Alberta. In: WRATHALL, B., Sydney, Australia, 19, 52-65.
JOHNSTON, G., ARTS, A., Rozsw, L, ZONNEVELD, HOWARD, J. D. 1971. Comparison of the beach to-to-
J.-P., ARCURI, D. & MCLELLAN, S. (eds) Digging offshore sequence in modern and ancient sedi-
Deeper: Finding a Better Bottom Line, Canadian ments. In: HOWARD, J. D., VALENTINE, J. W. &
Society of Petroleum Geologists and Petroleum WARME, J.E. (eds) Recent Advances in Paleoecology
Society 1999 Core Conference Paper, Calgary, and Ichnology. American Geological Institute,
Alberta, p. 99-114. Short Course Lecture Notes, Virginia, 148-183.
210 K. L. BANN ET AL.

HOWARD, J. D. 1972. Trace fossils as a criteria for NAISH, T. R. & KAMP, P. J. J. 1997. Sequence stratigra-
recognizing shorelines in the stratigraphic record. phy of sixth-order (41 k.y.) Pliocene-Pleistocene
In: RIGBY, J. K. & HAMBLIN, W. K. (eds) Recognition cyclothems, Wanganui Basin, New Zealand: a
of Ancient Sedimentary Environments. Society of case for the regressive systems tract. Geological
Economic Paleontologists and Mineralogists, Society of America Bulletin, 109, 978-999.
Special Publications, Tulsa, Oklahoma, 16,215-225. PEMBERTON, S. G. & FREY, R. W. 1984. Ichnology of
HOWARD, J. D. & FREY, R. W. 1973. Characteristic storm-influenced shallow marine sequence: Car-
physical and biological sedimentary structures in dium Formation (Upper Cretaceous) at Seebe,
Georgia estuaries. American Association of Petro- Alberta. In: STOTT, D. F. & GLASS, D. J. (eds)
leum Geologists Bulletin, 62, 1169-1184. The Mesozoic of Middle North America. Canadian
HOWARD, J. D. & FREY, R. W. 1975. Estuaries of the Society of Petroleum Geologists Memoirs,
Georgia Coast, USA: Sedimentology and biology. Calgary, Alberta, 9, 281-304.
II. Regional animal-sediment characteristics of PEMBERTON, S. G. & MACEACHERN, J. A. 1995. The
Georgia estuaries. Senckenbergiana Maritima, 7, sequence stratigraphic significance of trace fossils:
33-103. examples from the Cretaceous foreland basin of
HOWARD, J. D. & FREY, R. W. 1984. Characteristic Alberta, Canada. In: VAN WAGONER, J. C. & BER-
trace fossils in nearshore to offshore sequences, TRAM, G. (eds) Sequence Stratigraphy of Foreland
Upper Cretaceous of east-central Utah. Canadian Basin Deposits: Outcrop and Subsurface Examples
Journal of Earth Sciences, 21, 200-219. from the Cretaceous of North America. American
HOWARD, J. D. & REINECK, H.-E. 1981. Depositional Association of Petroleum Geologists, Memoirs,
facies of a high energy beach-to-offshore sequence: Tulsa, Oklahoma, 64, 429^75.
comparison with low-energy sequence. American PEMBERTON, S. G. & MACEACHERN, J. A. 1997. The ich-
Association of Petroleum Geologists Bulletin, 65, nological signature of storm deposits: the use of
807-830. trace fossils in event stratigraphy. In: BRETT,
HOWARD, J. D., ELDERS, C. A. & HEINBOKEL, J. F. C. E. (ed.) Paleontological Event Horizons: Eco-
1975. Estuaries of the Georgia Coast, USA: logical and Evolutionary Implications. Columbia
Sedimentology and biology. V. Animal-sediment University Press, New York, 73-109.
relationships in estuarine point bar deposits, PEMBERTON, S. G., FLACH, P. D. & MOSSOP, G. D. 1982.
Ogeechee River-Ossabaw Sound. Senckenbergiana Trace fossils from the Athabasca oil sands,
Maritima, 7, 181-203. Alberta, Canada. Science, III, 825-827.
KAPLAN, M. E. 1979. Calcite pseudomorphs (pseudo- PEMBERTON, S. G., MACEACHERN, J. A. & FREY, R. W.
gaylussite, jarrowite, thinolite, glendonite, genno- 1992a. Trace fossil facies models environmental
ishi, White Sea Hornlets) in sedimentary rocks: and allostratigraphic significance. In: WALKER,
origins of the pseudomorphs. Lithology and R. G. & JAMES, N. P. (eds) Facies Models: Response
Mineral Resources, 14, 623-636. to Sea Level Change. Geological Association of
KIDWELL, S. M. 1997. Anatomy of extremely thin Canada, St John's Newfoundland, 47-72.
marine sequences landward of a passive-margin PEMBERTON, S. G., MACEACHERN, J. A. & RANGER, M.
hinge zone: Neogene Calvert Cliffs succession, J. 1992b. Ichnology and event stratigraphy: the use
Maryland, USA. Journal of Sedimentary Research, of trace fossils in recognizing tempestites. In: PEM-
67, 322-340. BERTON, S. G. (ed.) Applications of Ichnology to
LEITHOLD, E. L. 1989. Depositional processes on an Petroleum Exploration: A Core Workshop. Society
ancient and modern muddy shelf, northern of Economic Paleontologists and Mineralogists,
California. Sedimentology, 36, 179-202. Core Workshops, Tulsa, Oklahoma, 17, 85-118.
MACEACHERN, J. A. & PEMBERTON, S. G. 1992. Ichno- PLUMMER, P. S. & GOSTIN, V. A. 1981. Shrinkage
logical aspects of Cretaceous shoreface successions cracks: desiccation or synaeresis. Journal of
and shoreface variability in the Western Interior Sedimentary Petrology, 51, 1147-1156.
Seaway of North America. In: PEMBERTON, S. G. PRATT, B. R. 1998. Syneresis cracks: subaqueous
(ed.) Applications of Ichnology to Petroleum shrinkage in argillaceous sediments caused by
Exploration, a Core Workshop. Society of earthquake-induced dewatering. Sedimentary
Economic Paleontologists and Mineralogists, Geology, 117, 1-10.
Core Workshops, Tulsa, Oklahoma, 17, 57-84. RANGER, M. J. & PEMBERTON, S. G. 1992. The sedimen-
MACEACHERN, J. A. & PEMBERTON, S. G. 1994. Ichno- tology and ichnology of estuarine point bars in
logical aspects of incised valley fill systems from the McMurray Formation of the Athabasca Oil
the Viking Formation of the Western Canada Sands Deposit, northeastern Alberta, Canada.
Sedimentary Basin, Alberta, Canada. In: BOYD, In: PEMBERTON, S. G. (ed.) Applications of Ichnol-
R., DALRYMPLE, R. W. & ZAITLIN, B. A. (eds) ogy to Petroleum Exploration: A Core Workshop.
Incised Valley Systems: Origin and Sedimentary Society of Economic Paleontologists and Mineral-
Sequences. Society of Economic Paleontologists ogists, Core Workshops, Tulsa, Oklahoma, 17,
and Mineralogists, Special Publications, Tulsa, 401-421.
Oklahoma, 51, 129-157. RAYCHAUDHURI, I. & PEMBERTON, S. G. 1992. Ichno-
MURRAY, C. G. 1990. Tectonic evolution and metallo- logic and sedimentologic characteristics of open
geny of the Bo wen Basin. In: Bowen Basin Sympo- marine to storm dominated restricted marine set-
sium, Geological Society of Australia, Queensland tings within the Viking/Bow Island Formations,
Division, Proceedings, 201-212. south central Alberta. In: PEMBERTON, S. G. (ed.)
ESTUARINE VS FULLY MARINE DEPOSITS 211

Applications of Ichnology to Petroleum Explora- bay to fluvial basin: the Eocene Ishikari Group,
tion: A Core Workshop. Society of Economic Ishikari Coal Field, Hokkaido, Japan. Sedimen-
Paleontologists and Mineralogists, Core Work- tary Geology, 160, 131-158.
shops, Tulsa, Oklahoma, 17, 119-139. TAYLOR, A. M. & GOLDRING, R. 1993. Description
REES, E. I. S., NICHOLAIDOU, A. & LASKERIDOU, P. and analysis of bioturbation and ichnofabric.
1977. The effects of storms on the dynamics of Journal of the Geological Society, London. 150,
shallow water benthic associations. In: KEEGAN, 141-148.
B. F, CEIDIGH, P. O. & BOADEN, P. J. (eds) Biology TEDESCO, L. P. & WANLESS, H. R. 1991. Generation of
of Benthic Organisms. Pergamon Press, Oxford, sedimentary fabrics and facies by repetitive exca-
465-474. vation and storm infilling of burrow networks,
REINECK, H.-E. 1963. Sedimentgefuge im Bereichder Holocene of south Florida and Caicos Platform,
sudlichen Nordsee. Abhandlungen der Senckenber- British West Indies. Palaios, 6, 326-343.
gische Naturforschende Gesellschaft, 505. THOMAS, R. G., SMITH, D. G., WOOD, J. M., VISSER, J.,
REINECK, H.-E. & CHENG, Y. M. 1978. Sedimento- CALVERLEY-RANGE, E. A. & KOSTER, E. H. 1987.
logische und faunistische Untersuchungen an Inclined heterolithic stratification: terminology,
Watten in Taiwan. I. Aktuogeologissche Untersu- description, interpretation and significance.
chungen, Senckenbergiana Maritima, 10, 85-115. Sedimentary Geology, 53, 123-179.
REINECK, H.-E, GUTMANN, W. F. & HERTWECK, G. TYE, S. C., FIELDING, C. R. & JONES, B. G. 1996.
1967. Das Schlickgebeit sudlich Helgoland als Stratigraphy and sedimentology of the Talaterang
Beispiel rezenter Schelfablagerungen. Seckenbergi- and Shoalhaven Groups of the southernmost
ana Lethaea, 48, 219-275. Sydney Basin. Australian Journal of Earth
REINECK, H.-E., DORJES, J., GADOW, S. & HERTWECK, G. Sciences, 43, 57-69.
1968. Sedimentologie, fauenzonierung und fazie- VAN WAGONER, J. C., POSAMENTIER, H. W., MITCHUM,
sabfolge vor der Ostkkiiste der inneren Deutschen R. M., VAIL, P. R., SARG, J. F., LOUTIT, T. S. &
Bucht. Senckenbergiana Lethaea, 49, 261-309. HARDENBOL, J. 1988. An overview of the
RUNNEGAR, B. 1979. Ecology of Eurydesma and the fundamentals of sequence stratigraphy and key
Eurydesma fauna, Permian of eastern Australia. definitions. In: WILGUS, C. K., HASTINGS, B. S.,
Alcheringa, 3, 261-285. POSAMENTIER, H. W., VAN WAGONER, J. C.,
SAUNDERS, T. D. A., MACEACHERN, J. A. & PEMBER- Ross, C. A. & KENDALL, C. G. ST C. (eds) Sea-
TON, S. G. 1994. Cadotte Member Sandstone: Level Changes: An Integrated Approach. Society
Progradation in a boreal basin prone to winter of Economic Paleontologists and Mineralogists,
storms. In: PEMBERTON, S. G., JAMES, D. P. & Special Publication, Tulsa, Oklahoma, 42, 39-45.
WIGHTMAN, M. (eds) Mannville Core Conference. VEEVERS, J. J. & POWELL, C. McA. 1987. Late glacial
Canadian Society of Petroleum Geologists, episodes in Gondwanaland reflected in transgres-
Calgary, Alberta, 331-349. sive-regressive depositional sequences in Euramer-
SCHAFER, W. 1956. Wirkungen der Benthos-Organis- ica. Geological Society of America Bulletin, 98,
men auf den jungen Schichtverband. Sencken- 475-487.
bergiana Lethaea, 37, 183-263. VOSSLER, S. M. & PEMBERTON, S. G. 1988. Skolithos in
SCHAFER, W. 1962. Aktuo-Palaontologie nach Studien in the upper Cretaceous Cardium Formation: an ich-
der Nordsee. Kramer, Frankfurt am Main. nofossil example of opportunistic ecology.
SCHIEBNER, E. 1974. A plate tectonic model of the Lethaia, 21, 351-362.
Palaeozoic history of New South Wales. Journal VOSSLER, S. M. & PEMBERTON, S. G. 1989. Ichnology
of the Geological Society of Australia, 20, 405^426. and paleoecology of offshore siliciclastic deposits
SHANLEY, K. W., MCCABE, P. J. & HETTINGER, R. D. in the Cardium Formation (Turonian, Alberta,
1992. Tidal influence in Cretaceous fluvial strata Canada). Palaeogeography, Palaeoclimatology,
from Utah, USA. Sedimentology, 39, 905-930. Palaeoecology, 74, 217-229.
SHANMUGAM, G., POFFENBERGER, M. & TORO ALAVA, J. WANLESS, H. R., TEDESCO, L. P. & TYRRELL, K. M.
2000. Tide-dominated estuarine facies in the 1988. Production of subtidal tubular and surficial
Hollin and Napo ('T' and 'U') Formations (Cre- tempestites by Hurricane Kate, Caicos Platform,
taceous), Sacha Field, Oriente Basin, Ecuador. British West Indies. Journal of Sedimentary Petrol-
American Association of Petroleum Geologists ogy, 58, 739-750.
Bulletin, 84, 652-682. WIGHTMAN, D. M., PEMBERTON, S. G. & SINGH, C. 1987.
SWIFT, D. J. P. 1975. Barrier-island genesis: evidence Depositional modelling of the Upper Mannville
from the central Atlantic shelf, eastern USA. (Lower Cretaceous), central Alberta: implications
Sedimentary Geology, 14, 1-43. for the recognition of brackish water deposits. In:
SWIFT, D. J. P., PARSONS, B. S., FOYLE, A. & OERTEL, TILLMAN, R. W. & WEBER, K. J. (eds) Reservoir
G. F. 2003. Between beds and sequences: strati- Sedimentology. Society of Economic Paleontolo-
graphic organization at intermediate scales in the gists and Mineralogists, Special Publication,
Quaternary of the Virginia coast, USA. Sedimen- Tulsa, Oklahoma, 40, 189-220.
tology, 50, 81-111. WHEATCROFT, R. A. 1990. Preservation potential of
TAKANO, O. & WASEDA, A. 2003. Sequence strati- sedimentary event layers. Geology, 18, 843-845.
graphic architecture of a differentially subsiding
This page intentionally left blank
Palaeoecology of the Bright Angel Shale in the eastern Grand Canyon,
Arizona, USA, incorporating sedimentological, ichnological and
palynological data
CHRISTOPHER T. BALDWIN1, P. K. STROTHER2, J. H. BECK2 & EBEN ROSE3
1
Department of Geography & Geology, Sam Houston State University,
Huntsville, Texas 77341, USA
2
Palaeobotanical Laboratory, Weston Observatory of Boston College, Department of
Geology & Geophysics, 381 Concord Road, Weston, Massachusetts 02493, USA
3
Department of Geology & Geophysics, Yale University, New Haven,
Connecticut 06520, USA

Abstract: The Middle Cambrian Bright Angel Shale in the eastern Grand Canyon contains a
depauperate normal marine fauna, but trace fossils and palynomorphs are abundant
throughout the formation. Conventional interpretations place the depositional setting of
this shale below wavebase as the distal component of a shelfal transgression, but the palyno-
logical signature in the mudstones of the Bright Angel Shale indicates a freshwater source to
these muds. Examination of several sections in the vicinity of Proterozoic monadnocks and
the integration of sedimentological, ichnological and palynological observations yield a more
robust model for the palaeoecology of the Bright Angel Shale. Initial correspondence
between organic matter content in mudstones and feeding type and intensity (as indicated
by traces) is consistent with an estuarine setting for this deposit. The level of organic activity
preserved in these sediments indicates that the carbon flux into shallow marine settings due to
terrestrial runoff was substantial by middle Cambrian (Glossopleura biozone) time.

Although the Grand Canyon of Arizona is continues to pervade most introductory texts
known as a geological wonder, modern research that deal with the classical Cambrian transgres-
on the palaeoecology and depositional environ- sion of Laurentia, resulting in overly simplistic
ments of some of its significant strata is lacking. transgressive models.
The middle Cambrian Bright Angel Shale (BAS), The abundance of trace fossils in some sections
situated between the underlying Tapeats Sand- of the BAS in the eastern Grand Canyon is
stone and the overlying marly Muav Limestone impressive (Schuchert 1918). These sediments
(these three together form the Tonto Group), have preserved a snapshot of an active ecosystem
has long been considered to be the distal that persisted throughout the entire depositional
'fining-up/cleaning-up' clastic facies of the basal sequence of the BAS and on upwards into the
Cambrian Sauk transgression. The classic work Muav Limestone. In addition to its rich ichno-
of McKee & Resser (1945) certainly left the fauna, we show below that the BAS also contains
impression that the BAS contains abundant a rich, sporadically distributed palynoflora (see
fossils of marine invertebrates, particularly tri- also Strother & Beck 2000). These organic-
lobites, that lend support to this interpretation. walled microfossils include spores of probable
But in the eastern Grand Canyon body fossils land plants and terrestrial algae in addition to
are generally quite rare, and with few exceptions cuticle fragments and probable egg cases of
are relegated to specific coarser sandy beds. metazoans. Significantly, the lack of acantho-
Within the olive-green mudstones that dominate morphic acritarchs in the microflora points to
the BAS, body fossils are limited to rare Lingu- minimal marine influence during the deposition
lella and rare impressions of small trilobites. of the primary muds and silts. The recovered
Given this paucity of palaeontological evidence, microflora indicates a distinctive freshwater
contemporary palaeoenvironmental interpreta- provenance to portions of the BAS sequence.
tion of the BAS, and hence of the Sauk cratonic The non-marine microfloral signature charac-
sequence of this part of North America, has teristic of pelitic lithologies appears in stark
relied more on sedimentological and lithofacies contrast to that of the 'marine' lithological and
criteria than on biological data or on combina- ichnological signatures derived from the inter-
tions of these. It is this restricted interpretation calated coarser beds. This is contrary to the
based on physical sedimentological criteria that standard interpretation of clastic deposition

From: MC!LROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 213-236. 0305-8719/04/S15.00 © The Geological Society
of London.
214 C. T. BALDWIN ET AL.

applied to the section by McKee, wherein finer observations supporting the shallow marine
grains were supposed to be the more marine nature of Tonto deposition. He emphasized the
distal representation of proximal sands. It local character of basal Tapeats deposition,
should be noted also that many of the trace fos- observing that much of the Tapeats sediment
sils, although preserved in convex hyporelief at was derived locally from immediate underlying
the base of sandstones, were in fact produced Proterozoic basement rocks. He concluded that
on or in muds such that the apparently para- the Tapeats was a deltaic deposit, 'on the land
doxical juxtaposition of marine and non-marine side rather than under the influence of the invad-
indicators is very intimately intertwined. It is ing sea' (Schuchert 1918, p. 366). He argued that
this intimacy that allows us to refine the palaeoe- the greenish shales that begin in the upper
cological setting for the BAS by combining the Tapeats and persist through the BAS into the
preserved elements of trace fossils with their Muav were derived from lengthy rivers draining
potential food. Thus we are herein concerned a long eroded 'Precambrian' peneplain. He inter-
with some of the gastronomic drivers that stimu- preted the Bright Angel Shale to be 'very shallow,
lated trace-makers to behave in certain different covering wide flats near to shore'. The overlying
and distinctive ways - and to leave preservable Muav was likewise interpreted to be a 'shallow-
traces of their behaviour in a complex sequence water, near-shore, marine deposit'. Schuchert
of rocks. considered the abundant glauconite as evidence
It is now possible to integrate ichnological and of organic activity, and he commented that the
palynological data within McKee's original extent of burrowing in the Muav was the greatest
palaeogeographic setting to create a composite he had ever seen in any Palaeozoic section.
picture of deposition and ecology of the BAS in Schuchert's observations overlapped with the
the eastern Grand Canyon. Trophic structure work of Noble (1910, 1922), who did not specu-
ascertained from ichnofabric analysis is made late about the primary depositional settings, but
more robust when integrated with organic did mention that boulders up to 30 ft in diameter
maceral analysis. In essence, these sediments were to be found in Tapeats sediments adjacent
have preserved the traces of feeding behaviour to Proterozoic basement highs that poke through
(and abundance of metazoan consumers) while the basal Cambrian sediments. Noble (1910)
simultaneously preserving an index of food type likened the palaeotopography to the contem-
and abundance in the non-bioturbated portions porary Laurentian shield, citing the isolated
of beds and the presumably indigestible residues monadnocks of resistant Baraboo Quartzite in
(cf. Mcllroy & Logan 1999) in the bioturbated Wisconsin as a specific analogue. In the Grand
portions. This integrated approach to facies inter- Canyon, resistant monadnocks of Neoprotero-
pretation, employing lithofacies, ichnofacies and zoic Shinumo Quartzite were gradually onlapped
palynofacies criteria - giving as much weight as by the muds of the transgressing Sauk Sea.
possible to the cryptic palaeoenvironmental signa- McKee & Resser (1945) expanded on these
ture of the mud rock components - produces a ideas, focusing on the Cheops section, which
more convincing palaeoenvironmental interpreta- they referred to as 'Tapeats Bay'. They measured
tion. The result is an environmental model that current directions in this embayment and
intrinsically implies greater temporal and spatial determined that the predominant flow direction
complexity than that derived from any single was nearly perpendicular to regional flow (conti-
facies criterion or from macroscopic and sand- nental runoff) (Fig. 2).
dominated ichno/litho-facies combinations. Sedimentation immediately adjacent to the
Proterozoic 'islands' can be quite spectacular; it
is possible today to stand on debris slumps of
Prior studies on the palaeoecology of the Tapeats Sandstone age and view the cliffs from
Tonto Group which the 500 Ma old breccias were derived. All
previous authors have remarked on the fact
The Tonto Group (Fig. 1) of the Grand Canyon that the sediments in the basal units adjacent to
region is composed of three intergradational the monadnocks were formed in situ and were
formations - in ascending order, Tapeats not significantly transported. But even though
Sandstone, Bright Angel Shale, Muav Limestone McKee & Resser (1945) recognized both the
- that form a prominent, largely planar bench- embayed and proximal nature of these sedi-
like feature directly above the relatively narrow ments, they still envisioned only a marine origin
inner canyon that is cut into various Proterozoic for them.
rocks. Reinterpretation by Wanless (1973) of shal-
Schuchert (1918) spent only five days studying low-water deposition for the overlying Muav
the Tonto Group, but he made some critical Limestone was extended down-section into the
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 215

Fig. 1. Stratigraphic context of the Bright Angel Shale (BAS) in the early Palaeozoic sequence of the Grand
Canyon of Arizona, USA. The Tonto Group onlaps, oversteps and drapes various components of the
Proterozoic substrate, which locally forms a distinctive monadnock unconformity surface.

'ironstone' beds of the BAS. He concluded that deposition in the Lower Cambrian Zabriskie
the BAS 'ironstones' are preserved laterites Quartzite and Wood Canyon Formation, which
developed subaerially on deflation surfaces. are lithological (though not allostratigraphic)
Wanless identified dololaminite sequences in equivalents of the Tapeats Sandstone to the
the upper Muav Limestone of the extreme west in the southern Great Basin. Depositional
western Grand Canyon that he argued were environments of Lower Palaeozoic sheet sand-
analogous to modern tidal flat deposits of stones are notoriously difficult to interpret
Andros Island. This reinterpretation undermined (Haddox & Dott 1990; Myrow et al 2003). How-
a basic tenet of the McKee & Resser (1945) ever, the prevalence of well-preserved channels,
model, i.e. that the Muav Limestone, especially overbank deposits and extensive lateral accretion
of the western Grand Canyon, represents the surfaces within the 'unfossiliferous' Tapeats
deepest and most distal offshore deposits of a Sandstone brings fluvial braidplain to the fore-
relatively uncomplicated eastward transgression. front among possible palaeoenvironments, an
To some extent it was a return to Schuchert's interpretation that compares favourably with
original interpretation of the Muav. the high bedload fluvial 'combing' model pro-
Perhaps all of the Tonto Group was deposited posed by Todd & Went (1991) for the Cambrian
in a much shallower-water setting than McKee & Aldernay Sandstone Formation of the Channel
Resser (1945) had originally suggested. Hereford Islands and the Devonian Glashabeg Formation
(1977) postulated fluvial processes within the of Ireland. Middleton & Elliott (2003) have
underlying Tapeats Sandstone of central Ari- suggested that the Tapeats Sandstone channels
zona. Similarly, Fedo & Cooper (2001) described represent tidal deposition in a subtidal setting,
fluvial influences in distal braidplain settings for but the architectural style of the Tapeats
216 C. T. BALDWIN ET AL.

Fig. 2. Location map showing interpreted palaeogeography for the eastern Grand Canyon. Shaded regions are
outcropping Precambrian monadnocks; underlying diagram depicts the reconstructed topography plus the
generalised sediment transport directions. Sample localities of measured sections shown (see text for location
names).

Sandstone contrasts sharply with models of Methods


multi-storey submarine channels as described
by Peakall et al (2000). In the eastern Grand Canyon there is an approxi-
Previous workers (Schuchert 1918; McKee & mately 60 km stretch of outcrop associated with
Resser 1945; Seilacher 1970; Elliott & Martin clastic deposition adjacent to and abutting
1987) noted the abundance of trace fossils in directly onto Proterozoic monadnocks that
the Tonto Group. All considered the sections were subaerially exposed during mid Cambrian
to be fully marine. Elliott & Martin (1987) time (Fig. 2). We have examined sections in
discussed exclusively shallow shelf, storm-influ- four drainage regions of this area: Thunder
enced ichnocoenoses in the BAS. Such inter- River (TC), Demaray Point, Cheops Pyramid
pretations were heavily influenced by the (CP) and Red Canyon (RC). Two sections were
prevailing notion that the Cruziana ichnofacies examined in detail at Sumner Butte (SB), which
as originally proposed by Seilacher (1967) is are halfway between the topographic highs at
essentially shelf al in character. To some extent Demaray Point and Cheops Pyramid, a total
there has been a positive feedback loop distance of 7 km. The first of these is a complete
reinforcing deeper-water interpretations of the section through the BAS, a thickness of close to
Cruziana ichnofacies, as many traces are found 100m (Figs 3, 5). In a lateral gully, a second
interbedded with mudstones (barren of body partial section was measured near the base of
fossils) that themselves are cited as evidence of the BAS. This subsection was informally referred
distal (mid-to-outer) shelf deposition. But, in to as Teichichnus Gully' (Fig. 4).
the BAS, there is truly a mixture of trace fossil Samples for palynology were taken from shale
types. As described elsewhere (e.g. Droser 1991; intervals in measured sections at Sumner Butte
Pemberton et al. 1992), vertical burrows typical with an average sampling frequency of one
of Skolithos ichnofacies occur in the same beds every 7m. Elsewhere, spot samples were taken
as horizontal traces related to the Cruziana during logging. Organic residues were extracted
ichnofacies, and in the BAS these mixtures of by treatment in hydrofluoric acid and oxidized
ichnofacies components are juxtaposed with either in Schultze's reagent or in nitric acid.
sedimentological and palynological indicators Further separation was achieved by flotation in
that suggest water depths much less than zinc chloride (s.g, = 2), by sieving, or both.
middle to outer shelf. Trace fossil distributions and ichnofabric indices
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 217

Fig. 3. A graphic log of the Sumner Bute section through the Bright Angel Shale, 'm' indicates various forms
of red and iron-rich sandstone units including those previously referred to by McKee as 'Magenta Beds'. Note
the increase in thickness of these 'm' beds up the section. Details of representative 'm' sequences in (A) are
shown in: (B) 18-23m; (C) 50.5-53 m. Facies interpretations summarized in Table 1 are included on the
detailed sequence logs.

(ii) (Droser & Bottjer 1986) were noted in the Canyon in combination with McKee's more
field as sections were logged. At Sumner Butte general work. This classification draws attention
34 trace fossil assemblages were recorded from to the conversion and overprinting of various
outcrop and from localized float. heterolithic lithofacies (Facies 1) into variously
configured facies (Facies 2) that represent differ-
ent forms and amounts of biogenic overprinting
Results of the original lithofacies. An additional pair of
glauconitic and other iron-rich lithofacies
Lithofacies (Facies 3 and 4) are distinct from the ambient,
heterolithic lithofacies. Brief descriptions and
Lithofacies associations in the BAS were origin- labels for the four facies are summarized in
ally described by Noble (1922) and well charac- Table 1.
terized by McKee & Resser (1945). A summary The sediments of the BAS are dominated by
(Table 1) is presented here, based on recent light olive grey (5Y 5/2) to dark greenish grey
field descriptions from the eastern Grand (5Gy 4/1) mudstones, siltstones and shales (Figs
218 C. T. BALDWIN ET AL.

Fig. 4. A section from Teichichnus Gully' (Fig. 3) showing the complex intermixing of feeding strategies
within the framework of the lithostratigraphy: (a) lithostratigraphy; (b) guild type (ichnofacies); (c) composite
facies type; (d) qualitative trace fossil designations. Palynological samples are designated by the boxed T1-T4.

5b, 6a, b). These mudrocks in field exposure dis- planes, especially within the lower transition
play varying degrees of platyness and lamination zone beds.
(Figs 6a, b, 7a), and typically include scattered The mudstone successions are punctuated by
glauconite grains throughout. Earlier authors sandstone beds of varying thickness, as noted
noted the abundance of micaceous bedding in the facies and section descriptions (Figs 3,
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 219

Fig. 5. Distinctive iron-rich sandstone beds ('Magenta Beds') forming regionally mappable horizons with the
Bright Angel Shale, (a) Two cliff-forming indurated iron cemented red sandstones (Facies 4) separated by a
poorly exposed interval of heterolithic Facies 1 and 2. The more prominent cliff in centre view is composed of a
glauconitic sandstone that becomes progressively more indurated and red. It culminates in a vertically
burrowed, wave worked surface, (b) The upper portion of the Sumner Butte section shows the repeated stacks
of glauconitic and red iron-rich horizons. Elevations within the section (see Fig. 3) are shown on the outcrop.

4, 5). These sandstone beds are typically appar- at Sumner Butte impart a dark green colour to
ently well laminated (ii 1-2) (Figs 6b, 7a) and the section (Figs 5, 6).
contain what appears to be muscovite, but The distinctive liver-coloured ironstone hori-
upon closer field and petrographic inspection zons ('Magenta Beds' of McKee & Resser
are found to be weathered glauconites. In these 1945) split and coalesce over several kilometres
finer-grained sandstones, peloidal glauconites in the eastern Grand Canyon in regionally trace-
do not dominate the texture, but they do able composite beds (Fig. 5). Phosphatized and
appear as a pervasive component. Peloidal variously eroded inarticulate brachiopod, tri-
glauconites can make up a substantial fraction lobite, echinoderm and hyolithid fragments are
of the coarser ironstones of Facies 3 and 4, and locally concentrated in irregular patches within
glauconite-rich beds in the middle of the section and particularly on the tops of these poorly
220 C. T. BALDWIN ET AL.

Table 1. Summary of fades designations

Fades label Observations/notes

Fades 1 (ii < 1) Primary depositional lithofacies


Fla Parallel-laminated shales/siltstones
Fib Interlaminated planar shales and fine sandstones
Flc Flaser and lenticular shales and sandstones
Fid Discrete, thin sandstone stringers in heterolithic
shales/siltstones/sandstones
Fie Flat-bedded sandstones
Flf Cross-bedded/ripple bedded sandstones
Facies 2 (ii > 1) Biogenic overprint of lithofacies
F2a Tunnelled siltstone/sandstone [reworked Fla or Fib]
F2b Discrete tunnelled sandstone stringers in tunnelled
heterolithics [reworked Fid]
F2c Tunnelled heterolithics [reworked Fib or Flc]
F2d Tunnelled sandstones [reworked Fie or Flf]
Facies 3 (ii 1-5) Glauconitic
F3a Glauconitic heterolithics May represent precursors to Facies 4 or incomplete
development of estuarine bars
F3b Glauconitic cross-bedded sandstones
F3c Glauconitic wave rippled sandstones
F3d Glauconitic sandstones
Facies 4 (ii 1-5) Red/iron = Magenta Beds
F4a Red sandstones Bar/estuarine sand build-ups
F4b Red ironstones
Further combinations to facies listed above combined as subscript descriptor (e.g. F3bBF):
DT Red mudstone draped top Iron-rich mud drape from suspension
BT Burrowed top Colonized omission surface
RT Wave-rippled top Switch from current to oscillatory flows
BF Burrowed foresets Burrows 'hanging' below foreset surfaces
Fe Iron-rich pellets Hematite/glaucony

ii = Ichnofabric Index (Droser & Bottjer 1986)

sorted arkosic ironstones. Magenta beds are typi- laminated at a centimetre scale with fine-grained
cally medium to coarse quartz sandstones, parallel-laminated or ripple-laminated mottled (ii
cemented by haematite, but with varying 0-1) quartz sandstones (Facies 1). These grade
amounts of peloidal glauconite of similar size upward into more heavily bioturbated (ii 2-5)
and lesser amounts of lithic fragments, feldspar siltstones (ii 2-4) (Facies 2) that are capped with
and shell fragments. They range in thickness coarse, dark-green or magenta-coloured, cross-
from a centimetre or so up to perhaps 2m, but bedded ironstones (Facies 3 and 4) typically with
typically form beds 2-5 dm thick (Figs 5a, 6a). scoured bases (Figs 5a, 6a) and with very variable
The distribution of Facies 4 (Red Ironstone ichnofabrics (ii 1-4 locally). At Sumner Butte,
facies) is shown for a relatively thick, perfectly haematite staining tends to increase as one
exposed section of the BAS at Sumner Butte moves up-section into the Muav Limestone
(Fig. 3a). The sequential stacking of the different above. These units higher in the section (approxi-
combinations of all facies is shown in examples mately 30-70 m) are ultimately erosively capped
of magenta bed sequences from a number of by decimetre-scale amalgamated and trough and
sample sites in the eastern Grand Canyon (Figs tabular cross-bedded and herringbone cross-
3, 4). stratified coarse-grained ironstones, or variously
The BAS is characterized by repeated coarsen- haematitic coarse-grained arkosic sandstones of
ing-upward sequences, typically 1.5-5.Om thick Facies 4 (Figs 5, 6a).
but ranging up to l l m thick, which are typically Underlying the distinctive magenta beds are
marked in the field by ferruginous sandstone dark green glauconitic interbedded sandstones,
caps. Several of these cycles are visible in Fig. siltstones and shales (Figs 5, 6). Many glauconi-
6a. They begin with non- to low-bioturbated tic sandstones are bidirectionally cross-bedded
(ii 1-3, 0-2) green shales and mudstones inter- with 180° bipolar set. At a number of different
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 221

Fig. 6. Outcrop at 30m, Sumner Butte section: (a) sharp lower contact of red sandstone facies (Facies 4) with
heterolithic Facies 1 and 2; (b) close-up of the heterolithic facies, demonstrating the intertwined nature of the
Palaeophycus burrowing within what was once a muddy substrate, and displaying tunnelled sandstone stringers
within sharply laminated heterolithic units of Facies la and Ib.

locations vertical U-shaped and straight burrows these frequently define diffuse foresets. At the
extend downwards from sharply defined foreset top of these units that underlie the distinct
surfaces for a few centimetres, implying a reddened beds the cross-bedded sands give way
distinctly episodic sedimentation. Rusty red- to burrowed and destratified (in part ii 5-6)
brown, iron-rich silt and clay 'biscuits' are units of sand/shale lenticles (Fig. 6b), plus bio-
present along with clear quartz granules, and genie pads composed of various Cruziana-formmg
222 C. T. BALDWIN ET AL.

Fig. 7. Intrastratal burrows (possibly 'burrowed Cruziantf or Palaeophycus) at 32 m in the Sumner Butte
section, sample GC98-13: (a) cross-section showing the unburrowed centre of the bed; (b) lower bedding plane
view; (c) lower bedding plane view in outcrop, showing the almost complete coverage of the active surface near
its active margin.

subunits ('dig marks' of Baldwin 1976, p. 137) forms represent a different set of burrowers or
and various superimposed Arenicolites, Phycodes, burrowing taking place under different physical
Teichichnus and indeterminate burrow fills. Also conditions or at significantly different times. The
present in this fades are rare but recurrent large traces, which contrast so markedly with
Teichichnus and lArenicolites that, in comparison the uniform tier depth of smaller ('normal')
with specimens from this and coeval sections, forms (Fig. 8), appear to represent exploratory
seem particularly large and cut down through probings, whereas the normal forms exploited a
multiple laminae and/or multiple sandstone 'discovered' persistent food source.
beds. It should be noted that these larger forms Numerous examples of the tops of the distinc-
are larger only in the sense of the extended tier tive 'Magenta Beds' are further distinguished by:
depth, and that in all other morphological
respects they are identical to the shallower dense packing of Skolithos and Arenicolites
forms. It therefore seems unlikely that the larger (ii locally 4) (Fig. 9a) that cross-cut multiple
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 223

Fig. 8. Dense packing of Teichichnus and Phycodes typical of the underside of sandstone stringers within Facies
2b in the lower 2m of ''Teichichnus Gully' exposures (see Fig. 4a).

foresets and sometimes penetrate units of includes Palaeophycus and Planolites. This
climbing ripple cross-lamination; group is often found in association with
individual species and superimposed multiple suspension feeders (Figs lOc, 11 a, b, d). This
species of epichnial grooves; guild contains evidence of shallow-tier bur-
interference and wave ripples; rowing and possibly surficial furrowing or
thin sheet-like or isopachous dark red mud sediment feeding right at the sediment water
drapes that cover these ripples. interface (Figs lib, d).
Other distinctive green glauconitic heterolithic
The beds and layers demonstrating sediment-
units occur above and below the arkoses, where
feeding strategies are ubiquitous. They vary
they exhibit almost no penetrative bioturbation
from individual lamina-like simple horizontal
or contain small Phy codes/Teichichnus-like
burrows (Fig. 6a, 7a, b) to galleries that may be
spreiten bearing walls that 'hang' from the base
either discrete and separate, or overlapping
of some sharply defined sandstone beds (cf.
(Fig. 7c). Beds subjected to sediment feeding
Droser et al 2002a, 2002b) and are confined to
can be completely disrupted (ii 4-5) (F3 b/c
the next adjacent underlying mudstone layer.
in Fig. 5a). Sediment feeding is common in
heterolithic units and also in sandstone beds in
the range of 5-25 cm thick. Hypichnial traces
Ichnoguilds are visually dominant in these sandstone field
exposures, with sediment-feeding galleries
At a very coarse scale, trace fossils were grouped
marking the lower interface of most discrete
into three associations or ichnoguilds (sensu
sandstones (of any size) where a variety of tier-
Bromley 1990, p. 211) based on deduced general
like layers extend down into the mudstone
feeding strategies:
below (Figs 6, 9a). However, it should be
Sediment feeding, as composed of mostly hori- noted that a significant number of burrows
zontal furrowing and burrowing ichnogenera extend through the overlying sandstone beds,
such as Cruziana, Palaeophycus, Phycodes, and that sediment browsing of the tops of
Rusophycus (including 'dig marks' and other the same beds may be equally prevalent. An
components of obvious arthropodan origin) example of this is illustrated in Figs 9a, b,
and Teichichnus (Figs 7, 8, lOd). which reveal a similar trace density at the top
Filter I suspension feeding, composed of vertical and bottom of the same slab. This situation
and 'IT burrows assignable to Arenicolites, somewhat contrasts with the Droser et al.
Diplocraterion, Monocraterion and Skolithos (2002a) observations of earlier Neoproterozoic-
(Figs 9, lib, d). Cambrian ichnofabrics, where 'unattached'
Surface feeders are a mix of trail-formers, forms predominate.
some of which produce positive relief casts Suspension feeders are recognized by the pre-
on the upper surfaces of sandstone beds and sence of vertical burrows. Skolithos communities
224 C. T. BALDWIN ET AL.

Fig. 9. Burrow forms within iron-rich sandstone horizons (including fully developed Magenta Beds) in
transition beds near the base of Cheops Pyramid: (a) completely tunnelled sand beds passing upwards into
horizons of discrete Skolithos and various 'U' burrows on and within foreset surfaces; (b, c) various densities of
packing of sprieten-bearing U burrows on the upper surfaces of iron cemented sandstones.

are common, particularly in the basal transition influx of Bright Angel muds was quite favourable
zone (Fig. 9). On exposed upper surfaces in to filter/suspension feeding. Filter feeding is
thinner sandstone beds, it is not uncommon to exclusively associated with sand beds, and
see both Monocraterion and paired holes mostly with thick, frequently multi-storey units
corresponding to Diplocraterion (Fig. lib, d). of sandstone that may or may not include
The basal section and parts of the underlying varieties of cross-bedding (Fig. 9). Some thin
Tapeats Sandstone contain many different units in which the primary depositional texture
versions of the 'dumbbell-shaped' impressions has been obliterated by vertical burrows (ii 5-6)
that mark the various levels of exhumation of occur infrequently throughout the section,
U-shaped burrows (Fig. 9b, c). It seems likely indicating minor hiatuses (see Mcllroy 2004;
that much of the Tonto habitat prior to the Pemberton et al 2004).
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 225

Fig. 10. Sharply defined biogenic marks (mostly trilobite scratches and dig marks) from both the upper and
lower interfaces of sandstones from Facies 2: (a) hypichnial Dimorphichnus and Monomorphichnus wipes on the
base of a thin fine sandstone; (b) epichnial grooves and paired raised welts on the upper surface of a fine
sandstone; (c) hypichnial casts and fills on a wave-rippled sandstone; (d) hypichnial casts of Cruziana and other
arthropod digging marks.

Surface traces are numerous throughout the addition to the surficial cover of Palaeophycus,
BAS. They may be true furrows or the imprints and therefore may represent pre-depositional
of the lower surfaces of exhumed burrow fills and post-depositional suites. Surface furrow
(Figs lOb, 11 a, c), and they are often preserved traces (i.e. grooves excavated into the top surfaces
as positive casts (e.g. Fig. lib, c). Beds such as of detail-retaining cohesive muds and sands) such
those in Figure lib and d appear to retain as Monomorphichnus (Fig. lOa), Cruziana and
evidence of two guild types, as they contain a Rusophycus (Baldwin 1977) show exceptional
spatial distribution of both burrow pairs in detail of preservation, with no sign of sand-mud
226 C. T. BALDWIN ET AL.

Fig. 11. Upper Surface traces, (a) Wrinkle marks ('interference "tadpole" ripples' of McKee & Resser 1945),
which include spatulate, tablespoon-sized depressions and a few trails. These sedimentary structures have been
proposed as shallow water indicators and also to have been formed by adhesion from the binding properties of
microbial cover (Hagadorn 1997). (b) Detail of the bedding surface in (d), showing the tops of paired vertical
tubes and the casts of trails formed on rippled beds, Sumner Butte section 11 m. (c) Upper surface traces on
thin sandstone bed, again demonstrating the ability of formerly wet sand to retain its shape after the
subsequent influx of mud. (d) Upper bedding surface of a rippled sandstone bed at Sumner Butte section 11 m;
outline demarcates enlargement in (b).

substrate intermixing (cf. Droser et al. 2002a, As with most field (versus core) descriptions
2002b). Burrowing forms (e.g. Arenicolites, Pla- or utilizations of trace fossils there is a strong
nolites and Teichichnus) typically increase up tendency to consider the sandier facies; it is
section over several metres (Fig. 4b, c) with com- after all sands that tend to cast and ultimately
mensurate destruction of laminae and increasing preserve and expose trace fossils as physical,
presence of glauconitic peloids. The resultant discrete entities, whereas the muds are poorly
texture is silty and highly bioturbated (ii 4—5), preserved. In the BAS, cast and filled structures
comparable to the 'green crumbly siltstone' predominate but - given the variety and inter-
facies of McKee & Resser (1945) (Fig. 5). mixing of original lithotopes, particularly as
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 227

represented by the heterolithic fades (Fades 2c, Type A represents primary-sourced organic
but also 2a and 2b, Table 1) - we are reasonably matter that is dominated by cryptospores and
confident that we have observed a broad non-marine algal cell clusters. Metazoan frag-
distribution of traces and feeding strategies ments are rare or missing, but overall preser-
in both sandstones and other finer-grained vation is excellent. Type B contains a more
lithologies. mixed assemblage that includes thinner-walled
cells and cell clusters with relatively fewer
thicker-walled (cryptospore) forms. This assem-
Palynofacies blage contains noticeable fragments of metazoan
origin (cuticles and structural elements), and may
Palynofacies studies are typically limited to a contain larger leiospheres that could be pro-
determination of the 'marine' or 'non-marine' toctist or metazoan derived. Preservation is
character of an assemblage. This is achieved by mixed. Type C represents a degraded assemblage
assessing the non-marine fraction in a typically that contains significant metazoan remains
mixed assemblage. In Devonian and younger dominating primary photosynthetic biota and
strata the provenance of palynological elements algal debris.
can be quite obvious, particularly for land An additional distinctive assemblage was
plants where spores and pollen grains fit into recovered immediately adjacent to Proterozoic
well-known plant classifications. This makes the highgrounds in Red Canyon drainage. These
characterization of palynofacies relatively clear. samples are characterized by excellent preserva-
Prior to the Devonian, we are dependent upon tion of large tissue fragments and 'leiospheres'
the cryptospore record to provide the 'terrestrial' in addition to pervasive non-marine crypto-
index that is used in palynofacies construction. spores and terrestrially derived algal cell clusters.
In this way, terrestrial palynofacies have been In addition, mats of filaments, similar to the
recognized as far back as the lower Ordovician tissues that comprise the Silurian terrestrial
(Wellman et al 2003). plant Nematothallus, are found in the Red
With the BAS assemblages, the primary hurdle Canyon samples. However, as this assemblage
to a basic palynofacies assessment is that no is not characteristic of palynomorphs from
recognizable marine elements are present in the Sumner Butte, where trace fossil data were col-
sequences. Acanthomorphic acritarchs, which lected, it will not be discussed further.
are characteristic of marine palynomorphs from Figure 12 illustrates the primary components
this time (Servais et al. 1996), are entirely missing of the most degraded palynofacies type C. Photo-
from the BAS, as are zonal taxa of any sort. This synthetic debris includes degraded clusters of
leads to an assertion that the assemblage is of cells (Fig. 12a-c, f, 1, m) and a few cryptospore-
non-marine or estuarine origin (Strother & like specimens such as tetrads (Fig. 12k) and
Beck 2000). Such an interpretation is in consort dyads (Fig. 12d, h). The preservation of this
with older conclusions about tidal features and material is generally poor, and the recovery
the 'embayments' of the eastern Grand Canyon from this sample, T-l, was very low. A cuticular
proposed by McKee & Resser (1945). The fragment (Fig. 12q) is the best preserved of any
Middle Cambrian palynomorphs recovered of the organic pieces recovered. Just higher in
from the BAS do not match any younger assem- the section at Teichichnus Gully' is a slightly
blages exactly, but in overall character they are better preserved assemblage (sample T-2,
similar to assemblages from known paralic Fig. 13), which can still be considered a type C
sequences (e.g. the Tucsarora Formation in palynofacies. Smooth well-preserved cuticle
Pennsylvania: Strother & Traverse 1979; John- (Fig. 13t) is similar to that seen in the prior
son 1985). sample. However, this assemblage is more
Although there is little doubt from the palyno- diverse in its cuticular remains, including some
logical perspective that the BAS assemblages are pseudo-cellular cuticles reminiscent of non-
heavily influenced by estuarine and/or non- marine debris found in younger rocks (Fig. 13g,
marine elements, without a taxonomic basis q). Most of the spore-like cells and clusters in
from which to proceed it is not yet possible to this group are physically damaged, presumably
generate a quantitative assessment of assemblage as a result of physical reworking of the sediment,
heterogeneity and hence palynofacies distribu- although it is still possible to recognize both thin-
tion. It is possible to broadly classify palyno- walled, 'leiosphere-like' cells (Fig. 13a, b, p)
morphs and palynodebris from the BAS, and and thicker-walled components (Fig. 13c, d).
such an attempt leads to sorting into 'fades' Again, as in sample T-l, the recovery in terms
based both on their elemental composition and of absolute abundance of organic matter was
on their quality of preservation (degradation). poor.
228 C. T. BALDWIN ET AL.

Fig. 12. Palynofacies Type C assemblage, sample T-1: (a) degraded cell cluster; (b) polyad cluster, genus V;
(c) degraded cell cluster; (d) cryptospore dyad; (e) envelope-enclosed cluster; (f) broken cryptospore tetrads and
polyad; (g) degraded dyad pair; (h) well-preserved minute dyad; (i) poorly preserved thin-walled dyad;
(j) degraded tetrad; (k) degraded tetrad; (1) polyad; (m) small cluster of minute cells; (n) degraded and broken
large 'leiosphere'; (o) degraded fragment; (p) degraded thick cuticle; (q) large thin cuticle fragment.

Fig. 13. Palynofacies type C assemblage, sample T-2: (a) thin-walled 'leiosphere'; (b) thin-walled folded
'leiosphere'; (c) large broken dyad; (d) large broken tetrad; (e) distorted dyad; (f) moderately well-preserved cell
cluster; (g) degraded reticulate tissue fragment; (h) smaller distorted dyad; (i) envelope enclosed spore-like body;
(j) broken enclosed spore-like body; (k) small enclosed monad; (1) damaged large enclosed monad; (m) cluster of
polyads, genus V; (n) degraded cluster of polyads, genus V; (o) degraded cluster of thin-walled cells; (p) large
'leiosphere' with broken wall; (q) thick reticulate cuticle forming a pseudocellular pattern; (r) cuticle? fragment;
(s) small tissue fragment with embedded tubular structures; (t) large folded fragment of smooth cuticle.
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 229

Fig. 14. Palynofacies type B assemblage, sample GC97-23: (a) polyad cluster; (b) poly ad cluster; (c) irregular
tetrad; (d) polyad cluster; (e) thin-walled dyads; (f) attached, thin-walled cluster; (g) planar cryptospore tetrad
with contact thickenings; (h) irregular to planar tetrad with envelope; (i) loose clump of small dyads and
monads; (j) loosely associated dyad and monad; (k) small cryptospores loosely associated; (1) attached dyad
pair - note thickened contact regions and envelope; (m) pair of attached cryptospore dyads; (n) large irregular
cluster; (o) amorphous cuticle fragment; (p) amorphous cuticle fragment; (q) amorphous cuticle fragment; (r)
structural metazoan fragment; spine?

An assemblage of palynomorphs from hetero- primary (i.e. photosynthetically derived) and


lithic fades F2b was recovered at 26m in the secondary (metazoan) organisms.
Sumner Butte section (roughly equivalent to the An assemblage from shales resting immedi-
shale seen in Fig. 6a). This assemblage is inter- ately on a sandstone surface colonized by trace
mediate between the degraded type C and the fossils of presumed filter feeders in Teichichnus
pristine type A samples, and is designated a Gully' (T-4) is the model for a type A palyno-
type B assemblage (Fig. 14). It contains the facies. This sample (Fig. 15) is well preserved,
ubiquitous photosynthetically derived elements without signs of physical damage, and contains
of spore-like tetrads and dyads (Fig. 14g-m), in no metazoan debris. The cryptospores include
addition to a particularly distinctive form, symmetric tetrads (Fig. 15a, b, j) and dyads
'genus V (Fig. 14a-f). Unlike the more pristine (Fig. 15d, f, o, q). Some of these components
type A palynofacies assemblage, however, bits can be seen in their more degraded forms in
of possible metazoan tissue are preserved assemblage type A. For example, small dyads
(Fig.l4o-r). Therefore, significantly, in spite of (Fig. 15o), clusters of small spherical cells
the moderate burrowing present in this hetero- (Fig. 15n) and irregular clusters (Fig. 15m)
lithic sample, preservation is quite good, and appear to have degraded equivalents in the type
the shales seem to contain remains of both C assemblages. This seems reasonable given the
230 C. T. BALDWIN ET AL.

Fig. 15. Palynofacies type A assemblage, sample T-4: (a) tetrahedral cryptospore tetrad; (b) tetrahedral
cryptospore tetrad showing clearly the nature of the thickened contact rings that characterize these forms; (c) a
more irregular form; (d) thin-walled cryptospore dyad; (e) large dyad; (f) small envelope-enclosed dyad; (g)
compact cryptospore tetrad with thin envelope; (h) more or less planar tetrad; (i) large thin-walled attached
cells; (j) well-formed spore/cryptospore tetrad; (k) linear spore cluster; (1) monad cluster; (m) poly ad; (n) cluster
of smaller cells; (o) small dyad; (p) loosely attached cell pair; (q) envelope-enclosed dyad; (r) planar tetrad of
envelope-enclosed cells; (s) thin-walled cells forming an open planar tetrad.

assumption that assemblage type A represents deposits, and that open furrow traces (e.g.
the signature of the primary photosynthetic Rusophycus) were preserved by rapid suspension
input into this ecosystem. Palynomorph abun- settling from waning post-event flows. But sand-
dance is high in this assemblage - probably 103 shale laminae may also occur in any number of
times that seen in the type C assemblage. settings, with a subtle variety of distinctive fea-
tures. Stochastic or episodic sheet sands depos-
ited during storm events should result in
Discussion grading, extensive scouring, and possibly diffuse
sand-to-mud transitions (B-C beds of the
Distal storm versus estuarine models of Bouma sequence) (Seilacher 1991; Seilacher &
deposition Aigner 1991; Walker 1992; Walker & Flint
1992). However, typical BAS sandstone inter-
Elliott & Martin (1987) proposed that the sand- beds have sharp upper and lower contacts (Fig.
shale interbeds of the BAS represent shelf storm 6) with well-preserved traces on both surfaces.
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 231

For example, detailed sole markings such as sandy beds attests to this source. Release of the
Monomorphichnus (Fig. lOa) may occur on the pulses of sand was perhaps storm controlled or
same thin sandstone interbeds whose tops are storm influenced, but there is little evidence
covered with pre- and post-depositional suites that storm mechanisms were the principal sedi-
including Planolites and others (Fig. lOa, b). mentological drivers actually within the estuarine
Some exposures of sand-shale interbeds show domain. Capping sandstones may have been
cyclic thickening and thinning of sandstone initiated by preferential erosion of softground
layers strongly suggestive of tidal bundles substrates (the green crumbly siltstone of
(Davis et al. 1998). Tidal signatures abound in McKee & Resser 1945; Facies 3 herein), but
the BAS, including herringbone cross-stratifica- tides and currents rather than waves acted as
tion showing time/tidal asymmetry (Klein 1970) the primary sediment-moving agent. Bar sands
within the highly scoured and sometimes amalga- on occasion built up sufficiently that their tops
mated glauconitic sandstones and ironstones that perhaps became emergent for finite periods
are most likely the product of very high hydro- of time, but were subsequently abandoned,
dynamic energy in very shallow water. The drowned and became moribund in the terminol-
coarsening-up to ironstone-cap facies successions ogy of North Sea sand ridges (Stride et al. 1982).
of the Bright Angel Shale are comparable to those In combination with palynological and ichno-
described by Kim & Lee (2000) from the Ordo- logical data, the basic sedimentology of the
vician Dongjeom Formation of Korea, and may BAS does not lend strong support to a storm-
represent repeated submergence and emergence influenced subtidal depositional setting as
of metre-scale intertidal to sub tidal cycles. promoted by Middleton & Elliott (2003 and
These, sometimes amalgamated, ironstones references therein). The non-marine organic
exhibit certain characteristics typical of con- signature of the mud rocks simply does not
densed sections (Loutit et aL 1988), including a support the predicted uniformity or homogeneity
concentration of coarse grains, extensive haema- of such an exclusively marine setting. The mud
tite cement, and what appear to be fossil stein- rocks in the BAS tell a very different input
kerns and boxstones. The tops of these beds story, and an estuarine model better explains
were reworked to some extent by both currents the whole package of partitioned and non-
and waves. They are capped by what appears uniform facies and sequences.
to be a phase of wave reworking, subsequent Deposition of the BAS in the eastern Grand
abandonment or still-stand linked to coloniza- Canyon occurred within the context of an estuar-
tion by filter feeders and some sediment grazers, ine to paralic system for which modern North
and subsequent draping with suspension fallout Atlantic storm-influenced shelf and inner shelf
iron-rich muds. Top surfaces of magenta beds analogues are inappropriate. Certainly the classi-
may therefore represent periods of maximum cal Sauk transgression of Sloss (1963) is very real,
flooding or abandonment. In this context the but it is also very complex, not only in terms of its
parasequences that form the BAS may represent often acknowledged diachronous properties but
repeated sand build-ups that punctuated a also in terms of preserved environments of
muddy, tidally influenced, estuary - but an estu- deposition. Combined biological and sedimento-
ary that was not incised in the often used modern logical criteria indicate that in the eastern Grand
sense (cf. Dalrymple et al. 1994). Even though Canyon the transgression manifests itself as a
these sequences comprise a transgressive package heterolithic set of beds representative not of a
that required some 100m of accommodation shelf but of coastal plain compartments situated
space, this was not a simple proximal-to-distal between subdued structural highs. Sediment type
fill sequence. The shales of the Bright Angel are and supply, not distance from shore, was the
not substantially bathymetrically different from most important factor in determining the resul-
either the Tapeats Sandstone clastic 'carapace' tant rock sequence. The most convenient label
below or the ccleaned-up' Muav Limestone to apply to these compartments is estuaries.
above.
This low-relief estuarine system was subjected
to the migration of bar-like sand bodies, whose
movement was driven by normal lateral sedimen- Trophic reconstruction based on integrated
tation and migration of estuarine sand bars or palynofacies and ichnofacies
from episodes during which pulses of coarse sedi-
ment were brought into or released into the estu- Palynomorph assemblages represent residual
ary from more offshore, marine environments. organic remains trapped in muds after burial.
The concentration of nearly every potentially Given the overwhelming abundance and sheer
marine invertebrate element into allochthonous volume of burrowing activity in the BAS, it
232 C. T. BALDWIN ET AL.

seems reasonable to assume that the organic of phytoplankton and other fallout from the
content of the Bright Angel mudstones could overlying column, but this seems to us to be
represent a subsample of the primary food highly unlikely given the complete absence of
sources driving this middle Cambrian ecosystem. other recognizable marine palynomorphs. Addi-
The preservation of refractory organic matter tionally, there is no evidence to suggest that this
in a depositional system is affected by a myriad inferred estuarine-like environment was charac-
of factors, including: terized by anything approaching an unmixed
water wedge with buoyant fresh water sitting
primary abundance of organic material from
above a saltwater unit below. Such an arrange-
autochthonous and allochthonous sources;
ment would surely have again left at least some
in situ physical chemistry of the depositional
mud rocks with a marine palynomorph signa-
environment - in this case, the Eh as it affects
ture. This is not the case anywhere that we
on oxidative degradation of refractory organic
have sampled the BAS. Thus organic-rich
matter;
muds, including some of terrestrial derivation,
microbial degradation of organic matter,
form the background and perhaps ambient
which may be syndepositional or early post-
motif to sedimentation in the BAS environment.
depositional in character;
The depositional environment was not likely
maceration by metazoan feeding.
anaerobic because accumulations of amorphous
The recovery of preserved refractory organic organic matter are rare: there are no black
matter in shales indicates that decay and degra- shales here. Instead, currents and surface
dation was incomplete or arrested at some mixing were supplying plenty of oxygen to the
point in time during the burial process. When respiring animals that produced the trace fossils.
organic matter is completely missing, we cannot Shallow surface waters above both muds and
reconstruct its prior history, except to propose sands must have been well oxygenated, either
that the sources were originally absent, the by atmospheric diffusion or by well-mixed terres-
organic matter was completely recycled within trial runoff. Preserved organic matter in the
the primary ecosystem, or post-depositional Bright Angel shales consists of particulate refrac-
physical chemical processes were driven to tory materials - principally spores, cell clusters
completion. and cuticular fragments. This is very unlike the
In the case of the BAS, however, the immedi- microgranulate amorphous organic (kerogen)
ate presence of traces indicates that food was matter typically extracted from anaerobic shales.
abundant in the primary system. Thus at least These sediments have preserved traces that
we can rule out that primary organic matter have developed at an ecological timescale that
was lacking. Palynomorphs and associated can be interpreted as successional transitions
microdebris also record indicators of both micro- from grazed surfacial microbial mats developed
bial and chemical degradation. Microbial degra- on low-water-content muds to fully bioturbated
dation is indicated by patterned wall destruction, and more vertically burrowed, high-water-
essentially holes that form in spore walls or content muds. The tops of BAS sequences are
permeate other tissues. The formation of pyrite, capped with both vertical Skolithos-facies
as small crystals, also occurs within spore walls, burrows immediately underlain by traces of
leaving an unambiguous indicator of reducing Cruziana, Palaeophycus, Teichichnus and Ruso-
chemical conditions prior to cessation of active phycus burrowing/furrowing traces. The sands
chemistry after burial. The quality of spore wall that overlie the heavily burrowed muddy tops
preservation may vary from perfectly smooth to became bioturbated by vertical Arenicolites,
granular and imperfect. Such variation is difficult Diplo crater ion and Skolithos, some of which
to assess with respect to its timing in the overall may have housed suspension feeders, and also
sequence of organic burial and degradation; by a variety of grazers.
however, when well-preserved and imperfectly The preservation detail seen in the BAS sand-
preserved spores co-occur in a sample, it stones of various thickness indicates that the
does indicate varying degrees of early stages of primary sands were bound and stabilized
degradation. enough to retain fine structure. Presumably,
All samples contain a background component there was an abundance of microbial exudate
of spherical to subspherical clusters of cells, and/or binding meiofauna that provided this
most of which appear to be the resting cysts of stabilization of the substrate. The formation of
terrestrial (freshwater) algae. These remains micro-reticulate 'wrinkle marks' ('interference
represent allochthonous non-marine material "tadpole" ripple marks' of McKee & Resser
brought into the system from fluvial sources. 1945, plate 12c), which occur high in the section
They also could be the autochthonous remains at Sumner Butte (Fig. 1 la), has been ascribed to
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 233

the binding strength of near-surface microbial the most degraded palynomorphs, and contain
mats (Hagadorn 1997). Modern analogues of metazoan fragments in high relative abundances.
this form occur in muddy supratidal environ- The mixed thin sandstone/shale beds (ii 2-4) are
ments on Padre Island in Texas, although they of intermediate quality of spore preservation.
can range into subtidal habitats as well (Haga- And shales basal to sequence-topping horizons
dorn 1997). The presence of grazing trails on that contain traces produced by active filter fee-
numerous similar surfaces (Fig. 9b) is a clear ders appear to contain the highest percentages
indication that nutrient was available at the of cryptospore tetrads and dyads, forms that
sand surface, reinforcing the notion that food have the highest degree of affinity with subaerial
sources were readily abundant. It seems reason- land plants.
able to speculate that some of that food was in It appears that the distinctive iron-rich facies
the form of either microbial mats or a meiofauna of the BAS can be considered as somewhat
(cf. Mcllroy & Logan 1999) whose polysacchar- exotic but regionally extensive aerobic facies
ide-rich extracellular exudates aided in binding that repeatedly punctuated a muddy estuarine
sediment. shoreline. The muds were characterized by high
Shales immediately overlying these grazed levels of organic input, but the high degree of
and burrowed upper surfaces contain a type A trace-making activity indicates that oxygen
palynoflora. These are assemblages dominated supply was not generally depleted. Both glauco-
by well-preserved spore and spore clusters with nitic and haematitic sands appear to represent
only minimal metazoan and/or proctoctist the ingress of tidal and other bar sand build-
remains. These represent primary (phototrophic) ups into an otherwise quiet and moderately
sources of organic matter that were brought into low-relief, muddy estuarine setting.
place with source muds, although it is possible
that estuarine aquatic algae may have contribu-
ted to the preserved organic content. Regardless Conclusions
of exact source, suspension feeders capable of
living in muddy waters would have had ample In the absence of palaeontological data from
food in this environment. Sample T-4 in which to infer depositional settings through uni-
'Teichichnus Gully' is of this type, occurring formitarian methodology, it is extremely difficult
low in the section near the transition zone in a for geologists to infer or reconstruct palaeo-
part of the section dominated by dense Skolithos environments with any degree of certainty.
and U-shaped burrow colonization. Although that seems to have been the case for
the Bright Angel Shale, in fact there is a wealth
of palaeontological information preserved both
Ecosystem reconstruction incorporating in the trace fossils and in the microscopic organic
multiple sources of data remains recovered from the ubiquitous shales of
the formation. When viewed together, the over-
When viewed together, the descriptive sequences whelming weight of evidence indicates that the
of lithofacies, ichnoguilds and palynofacies can depositional setting was estuarine and not fully
be placed into a partially constrained palaeo- marine.
geographic setting to create a robust depositional Marine invertebrates are almost exclusively
model for the Bright Angel system. At the outset, restricted to allochthonous ('Magenta Beds'),
the sheer volume of traces indicates an exception- but the in situ Lingulella species, which is the
ally rich ecosystem, possibly one that was fed by only invertebrate of note preserved in the shale
heterogeneous sources, and inhabited by a matrix, has long been considered a very shal-
variety of different kinds of organism. This low-water indicator. Rudwick (1970, p. 158)
ecosystem appears to have been fed by a rich ter- indicates that Lingulides may well have tolerated
restrial organic runoff derived from freshwater brackish conditions, so that an estuarine setting
algal and probably subaerial plant sources. for Lingulella is by no means out of the question.
Therefore it remains unlikely that the physical This fits the proposed model. Trace fossils asso-
setting for the BAS deposition included waters ciated with the Cruziana ichnofacies of Seilacher
of normal marine salinity and certainly not of are generally considered to be shelfal. But this
the type that we would expect to be present in interpretation can be modified or overturned in
any domain to which the epithet 'shelf might the light of other evidence. Other researchers
be applied. have discovered this ichnofacies in demonstrably
The preservation of both trace fossils and deltaic environments (Legg 1985) and Cruziana/
palynomorphs supports this interpretation. Skolithos associations in estuarine settings.
Excessively bioturbated beds (ii 5-6) contain The BAS preserves thin sandstone beds with
234 C. T. BALDWIN ET AL.

Mono crater ion I Skolithos on the tops and Cruzi- Society (Strother & Baldwin); American Chemical
ana on the bottoms of the same beds. These Society (Strother); Sam Houston State University
ichnotaxa may impart different environmental Office of Research (Baldwin); AAPG Grant-in-Aid,
meaning on paper, but in this case they were Grand Canyon Association, SEPM, Friday Lunch
Club (Rose). We also appreciate the valuable guidance
formed in the same bathymetric setting. Given provided by our reviewers: Mary L. Droser, M. Moczy-
the complete lack of a marine signal in the dlowska Vidal and Duncan Mcllroy.
palynological assemblages extracted from the
enclosing muds, that ecosystem was probably
estuarine in nature. References
The intimate admixture of burrowing activity
and organic deposition has the potential to BALDWIN, C. T. 1976. The trace fossil stratigraphy of
tightly characterize and partially reconstruct some shallow marine Cambro-Ordovician rocks
the trophic structure of this ancient ecosystem. from Brittany, NW Spain and the UK. PhD
Even though the analysis presented here is a thesis, Liverpool University.
BALDWIN, C. T. 1977. Internal structures of trilobite
work in progress, crude correlations among trace fossils indicative of an open surface furrow
palynomorph assemblage composition, sample origin. Palaeogeography, Palaeoclimatology,
preservation quality, and trace fossil activity Palaeoecology, 21, 273-284.
have been noted. In the most actively biotur- BROMLEY, R. G. 1990. Trace Fossils: Biology and
bated sediments, preserved structural organic Taphonomy. Special Topics in Palaeontology, 3,
matter is most degraded, and metazoan remains Unwin, London.
(e.g. cuticle fragments) form a significant com- DALRYMPLE, R. W., BOYD, R. & ZAITLIN, B. A. (eds)
ponent of the overall recovery. The opposite is 1994. Incised-Valley Systems: Origin and Sedimen-
seen in muds deposited near the base of sedimen- tary Sequences. Society of Economic Paleontolo-
tary packages that rest directly on beds with gists and Mineralogists, Tulsa, Oklahoma, Special
Publications, 51.
grazed surfaces and vertical burrows. These DAVIS, R. A. JR, ALEXANDER, C. R. & HENRY, V. J.
surface-grazing and filter-feeding trace fossil 1998. Tidal sedimentology: historical background
communities were inundated by muds that and current contributions. In: ALEXANDER, C. R.,
retain a primary signature of fallout of photo- DAVIS, R. A. & HENRY, V. J. (eds) Tidalites:
synthetic food sources. Ichnofacies characterized Processes & Products, Society of Economic
by mixed sandstone/shale interbeds with a Paleontologists and Mineralogists, Tulsa, Okla-
moderate degree of burrowing contain an inter- homa, Special Publications, 61, 1-4.
mediate palynoflora, reflecting both primary DROSER, M. L. 1991. Ichnofabric of the Paleozoic
input (photosynthetic organism) and secondary Skolithos ichnofacies and the nature and distribu-
remains (autochthonous fossils). tion of Skolithos piperock. Palaios, 6, 316-325.
DROSER, M. L. & BOTTJER, D. J. 1986. A semi-
The great abundance and diversity of traces in quantitative field classification of ichnofabrics.
the BAS have yet to be quantified; yet they are Journal of Sedimentary Petrology, 56, 316—325.
truly vast. The influx of carbon needed to fuel DROSER, M. L, JENSEN, S. & GEHLING, J. G. 2002a.
these animal communities was substantial, and Trace fossils and substrates of the terminal Proter-
would have been comparable to that seen in ozoic-Cambrian transition: implications for the
modern estuaries and other shallow marine record of early bilaterians and sediment mixing.
settings that support up to 100% mixing of Proceedings of the National Academy of Sciences,
burrowed substrate. Recovery of palynomorphs 99, 12572-12576.
reveals the source of at least part of the primary, DROSER, M. L, JENSEN, S., GEHLING, J. G., MYROW,
P. M. & NARBONNE, G. M. 2002b. Lowermost
photosynthetic organisms that drove carbon pro- Cambrian Ichnofabrics from Chapel Island
duction in this system. Ironically, these are not Formation, Newfoundland: implications for
marine algae, but what appears to be a mix of Cambrian Substrates. Palaios, 17, 3-15.
non-marine algae and ancestral land plants that ELLIOTT, D. K. & MARTIN, D. L. 1987. A new trace
produced resistant-walled spores (cryptospores fossil from the Cambrian Bright Angel Shale,
sensu Strother & Beck 2000). Thus the ecosystem Grand Canyon, Arizona. Journal of Paleontology,
under study in the BAS represents an important 61, 641-648.
first step in characterizing and quantifying the FEDO, C. M. & COOPER, J. D. 2001. Sedimentology
earliest known example of carbon dynamics at and sequence stratigraphy of Neoproterozoic and
Cambrian units across a craton-margin hinge
the terrestrial/marine interface. zone, southeastern California, and implications
for the early evolution of the Cordilleran margin.
For fieldwork assistance we thank C. Lenk, B. Haley, Sedimentary Geology, 141-142, 501-522.
L. Marschall, J. Strother and T. McNulty; for Geo- FEDO, C. M. & PRAVE, A. R. 1991. Extensive Cambrian
chemistry: K. Harrison. Funding support for the braidplain sedimentation: insights from the
project includes the following: National Geographic southwestern USA Cordillera. In: COOPER, J. D.
PALAEOECOLOGY OF THE BRIGHT ANGEL SHALE 235

& STEVENS, C. H. (eds) Paleozoic Paleogeography NOBLE, L. F. 1910. Contributions to the geology of
of the Western United States-H, Vol 1. The the Grand Canyon, Arizona; the geology of the
Pacific Section Society of Economics Paleontolo- Shinumo area. American Journal of Science, ser
gists and Mineralogists, Los Angeles, Book 67, 4, 29, 369-528.
227-236. NOBLE, L. F. 1922. A section of the Paleozoic forma-
HADDOX, C. A. & DOTT, R. H. JR. 1990. Cambrian tions of the Grand Canyon, at the Bass trail.
deposits in northern Michigan. Journal of Sedi- USGS Professional Papers, 131-B, 23-73.
mentary Petrology, 60, 697-716. PEAKALL, J., MCCAFFREY, B. & KNELLER, B. 2000.
HAGADORN, J. W. 1997. Wrinkle structures: microbially A process model for the evolution, morphology,
mediated sedimentary structures common in and architecture of sinuous submarine channels.
subtidal siliciclastic settings at the Proterozoic- Journal of Sedimentary Research, 70, 434^448.
Phanerozoic transition. Geology, 25, 1047-1050. PEMBERTON, G. S., MACEACHERN, J. A. & FREY, R. W.
HEREFORD, R. 1977. Deposition of the Tapeats Sand- 1992. Trace fossil facies models: environmental
stone (Cambrian) in central Arizona. Geological and allostratigraphic significance. In: WALKER,
Society of America Bulletin, 88, 199-211. R. G. & JAMES, N. (eds) Facies Models and
JOHNSON, N. G. 1985. Early Silurian palynomorphs Sea Level Change (3rd edn). Geological Asso-
from the Tuscarora Formation in central Pennsyl- ciation of Canada, St. Johns, Newfoundland,
vania and their paleobotanical and geological sig- 47-72.
nificance. Review of Palaeobotany and Palynology, PEMBERTON, G. S., MACEACHERN, J. A. & SAUNDERS, T.
45, 307-360. 2004. Stratigraphic applications of substrate-
KIM, Y. & LEE, Y. I. 2000. Ironstones and green specific ichnofacies: delineating discontinuities
marine clays in the Dongjeom Formation (Early in the rock record. In: MclLROY, D. (ed.) The
Ordovician) of Korea. Sedimentary Geology, 130, Application of Ichnology to Palaeoenvironmental
65-80. and Stratigraphic Analysis. Geological Society,
KLEIN, G. De V. 1970. Depositional and dispersal London, Special Publications, 228, 29-62.
dynamics of intertidal sand bars. Journal of RUDWICK, M. J. S. 1970. Living and Fossil Brachiopods.
Sedimentary Petrology, 40, 1095-1127. Hutchinson, London.
LEGG, I. C. 1985. Trace fossils from a Middle Cambrian SCHUCHERT, C. 1918. The Cambrian of the Grand
deltaic sequence, north Spain. In: CURRAN, H. A. Canyon of Arizona. American Journal of Science,
(ed.) Biogenic structures: their use in interpreting ser 4, 45, 362-369.
depositional environments. SEPM Special Publica- SEILACHER, A. 1967. Bathymetry of trace fossils.
tion, 35, 151-165. Marine Geology, 5, 413-428.
LOUTIT, T. S., HARDBENOL, J., VAIL, P. R. & Ross, G. R. SEILACHER, A. 1970. Cruziana stratigraphy of non-
1988. Condensed sections: the key to age dating fossiliferous Palaeozoic sandstones. In: CRIMES,
and correlation of continental margin sequences. T. P. & HARPER, J. C. (eds) Trace Fossils. Geo-
In: WILGUS, C. K., HASTINGS, B. S., KENDALL, logical Journal, Special Issue 3, 447-476.
C. G. ST C., POSAMENTIER, H. W., Ross, C. A. & SEILACHER, A. 1991. Events and their signatures. In:
VAN WAGONER, J. C. (eds) Sea Level Changes: ElNSELE, G., RlCKEN, W. & SEILACHER, A. (eds)
An Integrated Approach. Society of Economic Cycles and Events in Stratigraphy. Springer,
Paleontologists and Mineralogists, Tulsa, Okla- Berlin, 222-226.
homa, Special Publications, 42, 183-213. SEILACHER, A. & AIGNER, T. 1991. Storm deposition
MclLROY, D. 2004. Some ichnological concepts, at the bed, facies, and basin scale: the geological
methodologies, applications and frontiers. In: perspective. In: EINSELE, G., RICKEN, W. &
MclLROY, D. (ed.) The Application of Ichnology SEILACHER, A. (eds) Cycles and Events in Stratigra-
to Palaeoenvironmental and Stratigraphic Analysis. phy. Springer, Berlin, 249-267.
Geological Society, London, Special Publications, SERVAIS, T., MOLYNEUX, S. G., BROCKE, R., FATKA, O.
228, 3-27. & LE HERISSE, A. 1996 Value and meaning of
MclLROY, D. & LOGAN, G. A. 1999. The impact of the term acritarch. Ada Universitatis Carolinae -
bioturbation on infaunal ecology and evolution Geologica, 40, 631-643.
during the Proterozoic-Cambrian transition. SLOSS, L. L. 1963. Sequence in the cratonic interior of
Palaios, 14, 58-72. North America. Geological Society of America
McKEE, E. D. & RESSER, C. E. 1945. Cambrian History Bulletin, 74, 93-114.
of the Grand Canyon Region. Carnegie Institute STRIDE, A. H., BELDERSON, R. H., KENYON, N. H. &
Publication, Washington DC, 563. JOHNSON, M. A. 1982. Offshore tidal deposits:
MIDDLETON, L. T. & ELLIOTT, D. K. 2003. Tonto sand sheets and sand bank facies. In: STRIDE,
Group. In: BELTS, S. S. & MORALES, M. (eds) A. H. (ed.) Offshore Tidal Sands, Process and
Grand Canyon Geology (2nd edn). Oxford Deposits. Chapman & Hall, Edinburgh, 95-125.
University Press, Oxford, 90-106. STROTHER, P. K. & BECK, J. H. 2000. Spore-like
MYROW, P. M., TAYLOR, J. F., MILLER, J. F., ETHING- microfossils from Middle Cambrian strata:
TON, R. L., RlPPERDAN, R. L & ALLEN, J. 2003. expanding the meaning of the term cryptospore.
Fallen arches: dispelling myths concerning In: HARLEY, M. M., MORTON, C. M. & BLACK-
Cambrian and Ordovician paleogeography of the MORE, S. (eds) Pollen and Spores: Morphology
Rocky Mountain region. Geological Society of and Biology. Royal Botanic Gardens, Kew, 413-
America Bulletin, 115, 695-713. 424.
236 C. T. BALDWIN ET AL.

STROTHER, P. K. & TRAVERSE, A. 1979. Plant micro- Models: Response to Sea Level Change. Geological
fossils from Llandoverian and Wenlockian rocks Society of Canada, St. Johns, Newfoundland,
of Pennsylvania. Palynology, 3, 1-21. 239-263.
TAYLOR, K. G. & CURTIS, C. D. 1995. Stability and WALKER, R. G. & PLINT, A. G. 1992. Wave- and storm-
facies associations of early diagenetic mineral dominated shallow marine systems. In: WALKER,
assemblages: an example from a Jurassic iron- R. G. & JAMES, N. P. (eds) Facies Models:
stone-mudstone succession, UK. Journal of Sedi- Response to Sea Level Change. Geological Society
mentary Research, A65, 358-368. of Canada, St. Johns, Newfoundland, 219-238.
TODD, S. P. & WENT, D. J. 1991. Lateral migration of WANLESS, H. R. JR. 1973. Cambrian of the Grand
sand-bed rivers: examples from the Devonian Canyon: a re-evaluation of the depositional
Glashabeg Formation, SW Ireland and the environments. PhD dissertation, Johns Hopkins
Cambrian Alderney Sandstone Formation, Chan- University.
nel Islands. Sedimentology, 38, 997-1020. WELLMAN, C. H., OSTERLOSS, P. L. & MOHIUDDIN, U.
WALKER, R. G. 1992. Turbidites and submarine fans. 2003. Fragments of the earliest land plants.
In: WALKER, R. G. & JAMES, N. P. (eds) Facies Nature, 425, 282-285.
Ichnofabrics and sedimentary fades of a tide-dominated delta: Jurassic
He Formation of Kristin Field, Haltenbanken, Offshore Mid-Norway

DUNCAN McILROY

Sedimentology and Internet Solutions Ltd, 29 Proctor Road, Hoylake,


Wirral, CH47 4BB, UK (e-mail: dmc@duncanmcilroy.com)

Abstract: Tide-dominated deltas are poorly known from the stratigraphic record and are
notoriously complex, owing to the wide spectrum of facies encountered and their spatial/
temporal variability. The tide-dominated deltaic palaeoenvironment combines the ecological
harshness of brackish-water settings with complex tidal channel/tidal-flat type facies architec-
ture on the delta top, in association with more classic deltaic facies-stacking patterns. The He
Formation is interpreted herein as a tide-dominated delta deposited in a microtidal setting.
Its palaeoenvironments are interpreted based on a combination of ichnology, ichnofabric
analysis and sedimentology. Ichnofabric stacking patterns are used to elucidate the internal
architecture of the notoriously problematic aggradational multi-storey tidal channel units.
The tide-dominated deltas of the He Formation have a distinctive ichnological signature
that may be used to characterize tide-dominated deltas. In comparison to typical river-
dominated deltas the Skolithos ichnofacies is less well developed and ichnodiversity is lower
than expected in wave-dominated deltas. The ichnofabric model presented has potential to be
used, with modification, in other tide-dominated deltaic settings.

Introduction from geochemical, palaeontological and ichnolo-


gical studies (as appropriate).
The ichnology of tide-dominated brackish-water The major challenge with understanding
settings is complex, owing to the wide range of petroleum reservoirs in tide-dominated deposi-
environmental conditions experienced by the tional settings is confidence in distinguishing
infauna. In most case studies investigating the between different facies at a suitable resolution
ichnology of ancient tidal deposits, the palaeo- to reduce uncertainty in reservoir geology.
environment is that of a drowned river valley, i.e. Tide-dominated deltas contain both deltaic and
an estuary (e.g. Goldring et al. 1978; MacEachern brackish-water facies, similar to estuaries, and
1989; Wrightman et al 1987; Mattison et al 1989; therefore pose special problems. The resolution
Pattison 1992; Pemberton & Wrightman 1992; of facies analysis can be significantly improved
Pemberton et al 1992; Buatois et al 1998; Buatois in highly heterogeneous reservoir facies, such as
& Mangano 2004). All tidal systems comprise a characterize tidal deposits, through detailed
complex mosaic of environments in which a ichnological study using either a bespoke assem-
range of physical and chemical conditions affect blage or an ichnofabric approach such as pre-
infaunal (trace-making) organisms. Understand- sented herein. It is noted that in some (mostly
ing of detailed trace fossil distributions within Palaeozoic) cases the ichnology of tidal environ-
such complex tidal settings is commonly restricted ments can constitute almost entirely bedding
to the observation that tidal (particularly estuar- parallel traces (e.g. Buatois et al 1998; Buatois
ine) facies are characterized by a mixture of the & Mangano 2004), which leave very little ichno-
Cruziana and Skolithos ichnofacies (e.g. Pember- fabric when seen in vertical cross-section and
ton et al 2001). It is noted, however, that a similar restrict the use of core analysis to the limited
admixture of the Cruziana and Skolithos ichno- bedding surfaces between core segments.
facies can characterize lower shoreface event
beds (Bromley & Asgaard 1991; Frey & Goldring
1992). The relative importance of various hydro- Objectives
dynamic regimes (e.g. waves versus tides) allows
definition of a system as being 'influenced' or The primary objectives of this work were thus to
'dominated' by a particular depositional process. assess the sedimentology and ichnology of the He
The recognition of the character of the predomi- Formation on Kristin Field through sedimentary
nant hydrodynamic regime is integral to the con- logging at a 1:50 scale, and to provide ichnologi-
struction of realistic facies models, and is ideally cally constrained facies models and a framework
based on a combination of diagnostic sedimentary for integration with other in-house datasets (e.g.
structures along with palaeoenvironmental data M0rk et al 1997; Stilling 2000; Greger et al
From: MC!LROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 237-272. 0305-8719/04/S15.00 © The Geological Society
of London.
238 D. McILROY

Fig. 1. Location of the Kristin Field in relation to other fields in the Haltenbanken area, offshore Mid-
Norway.

2001) and as input to stochastic reservoir models Sedimentology and stratigraphy of the He
of this important reservoir interval. Formation
The He Formation is a shallow-marginal marine
Studied core succession of Aalenian age that forms part of
the Fangst Group (Fig. 2). It is widespread in
This study focused on the He Formation in the distribution on Haltenbanken and is generally
four cored wells from Kristin Field (6406/2-5; between 80m and 90m in thickness. The for-
6506/11-6; 6406/2-3-T3; 6406/2-5A-T2) (Fig. 1). mation is strongly progradational at its base,
Previous in-house studies on the field (M0rk comprising an upward-coarsening delta front
et al 1997; Stilling 2000; Greger et al 2001) succession that is in gradational contact with
were not available to the author, and this work the underlying Ror Formation. This is overlain
is independent of their content. by a broadly aggradational delta top succession
The scale of the current study was such that with tidal channels that cut into and rework the
bed-by-bed sedimentological and ichnological adjacent tidal flats. The upper portion of the lie
observations could be made. Study of the He Formation records a gradual flooding of the
Formation on Sm0rbukk Field (see Fig. 1) con- system and is dominated by subtidal flat and
firms that the ichnofabric scheme proposed prodelta deposits (see Fig. 3). Good-quality
herein could be applied with slight modification sandstone reservoirs in the He Formation are
to adjacent fields. largely reliant upon the presence of grain-coating
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 239

communities of organisms. Although description


of ichnological assemblages is a worthwhile
exercise in itself, it becomes a much more power-
ful tool when the ichnological assemblage is
deconstructed into its component parts using
ichnofabric analysis (see Taylor & Goldring
1993; Taylor et al. 2003; Mcllroy 2004). This
can lead to a greater understanding of conditions
before, during and after deposition of a bed, and
is especially useful in the recognition of surfaces
of Stratigraphic importance (cf. Taylor &
Gawthorpe 1993; Taylor & Goldring 1993;
Taylor et al. 2003; Mcllroy 2004). Once charac-
terized, ichnofabrics are conventionally named,
e.g. PlanoliteslAsterosoma ichnofabric, with the
eponymous trace(s) being either highly visible
component(s) of the ichnofabric or its most
characteristic component(s).

Combining ichnological and sedimentological


Fig. 2. Stratigraphic units of the Triassic and approaches
Jurassic on Haltenbanken; major reservoir intervals
shaded grey. Trace fossils are best considered as biogenic
(animal/plant-made) sedimentary structures
that contribute to the sediment texture as seen
chlorite, which inhibits quartz cementation (see in core/outcrop. Their study is therefore as
summary in Ehrenberg 1993; Ehrenberg et al much a branch of sedimentology as it is a subdis-
1998). In order to assess the importance of cipline of palaeontology. Like physical sedimen-
palaeoenvironment in controlling the distribu- tary structures, trace fossils can yield important
tion of chlorite-coated grains a refined palaeo- information about the environment at the time
environmental/facies model was constructed for of sedimentation.
comparison with other datasets. Facies are defined using physical sedimentary
structures (to understand the depositing current)
along with trace and body fossils (Reading 1996).
Ichnofabric description Further refinement of this approach can be
achieved by studying the trace fossil assemblage
This study has focused upon the ichnology of the in more detail. For example, a trough cross-
He Formation within the context of broader- bedded tidally influenced channel facies will
scale sedimentological studies on Haltenbanken. contain similar sedimentary structures (at core
This integrated approach to reservoir description scale) along its entire length (may be hundreds
is fundamental to rigorous facies analysis, in of kilometres), but the distribution of trace
which palaeontological/palaeoenvironmental fossil assemblages/ichnofabrics, as well as being
information is considered alongside evidence affected by current strength, will also be influ-
from physical sedimentary structures. Most enced by the salinity gradient. In this way a
descriptions of sediments are taken with refer- single sedimentary facies can be subdivided, on
ence to beds, within which observations of physi- a combination of ichnological and sedimentolo-
cal and biogenic sedimentary structures are gical grounds, into its component parts. This is
made. This necessitates sedimentary logging at important for the recognition of subtle stacking
at least 1:50 scale to capture most individual patterns in apparently aggradational successions
beds. Indeed, the most easily described unit in [e.g. stacked tidal channels (herein) and shore-
ichnological studies is an ichnological assem- faces (Martin & Pollard 1996)]. Such variations
blage, which comprises all the trace fossils in stacking patterns are the fundamental data
found within a bed (see Mcllroy 2004). The that sedimentologists/sequence stratigraphers
term 'ichnological assemblage' does not discrimi- use to predict inter-well facies architectures by
nate between the relative ages of the component application of Walther's law (Walther 1894),
trace fossils, and may represent time-averaged and in this respect sedimentary facies, ichno-
burrowing resulting from several successive fabrics, ichnological assemblages and archetypal
240 D. McILROY

Fig. 3. Summary stratigraphic section through the He Formation as expressed on Kristin Field, combined from
features of all four cored wells.
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 241

ichnofacies can be treated in exactly the same Ichnofabrics of the He Formation


manner.
What follows is an account of the ichnofabric
types documented from the He Formation. It
Ichnology of the He Formation results from many individual observations of
trace fossils from the four wells studied. The
Trace fossils of the He Formation ichnofabric scheme is by necessity fairly exten-
sive owing to the large total ichnodiversity and
The He Formation contains a diverse assemblage sedimentological/palaeoenvironmental disparity
of trace fossils that show a strong palaeoenviron- that characterizes the He Formation (see
mental control on their distribution. These below).
palaeoenvironmental controls on trace fossils The 25 ichnofabrics are ordered alphabetically
allow highly refined facies analysis of their host with respect to their abbreviated ichnofabric
sediments when combined with sedimentological title to ease comparison with the reservoir
interpretations. As the most fundamental unit of descriptions and the sedimentological descrip-
stratigraphy is that of facies, improved facies tions below, and are integrated into the con-
characterization is integral to the construction ceptual model (see sedimentology section
of robust conceptual facies models, sequence below). In all cases, ichnofabric description is
stratigraphic models and the petroleum geo- supplemented with sedimentological implica-
logical applications thereof. tions and distribution in sedimentary facies.
The diverse assemblage of ichnofossils docu- Ichnofabric constituent diagrams for all ichno-
mented from the He Formation described fabrics are presented in Appendix Figs 1 and 2
during this study (Table 1) indicates a shallow using a modified version of the ICD method
marine depositional setting with no diagnosti- proposed by Taylor & Goldring (1993) (see
cally non-marine ichnotaxa represented. Mcllroy 2004).

Table 1. Trace fossils found in the uppermost Ror and He Formations and their facies distributions (see Table 2)

Trace fossil Palaeoenvironmental range in the He

Arenicolites isp. PDMB, TC, IDMBTC


Asterosoma isp. PDMB, STF, OTDC, DMB
Bergaueria ?perata STF
Chondrites isp. TC, IDMBTC, PDMB, PPD, DMB
Diplocraterion parallelum DMB, DDMB
Diplocraterion polyupsilon DMB, DDMB
Gyrochorte isp. DMB
Gyrolithes polonicus TC
Lockeia amygdaloides ITF, STF, TC, PDMB, DMB, DDMB, ATC
Monocraterion isp. TC, IDMBTC
Ophiomorpha nodosa PDMB, IDMBTC, TC
Ophiomorpha irregularis PDMB, TC, IDMBTC, DMB
Palaeophycus tubularis STF, ATC, PDMB, DMB, PPD, DPD
Palaeophycus heberti PPD, DDMB, DMB
Phoebichnus trochoides PPD, DPD
Phycosiphon incertinum DPD, PPD, DDMB, DMB, TC, STF, OTDC
Planolites beverleyensis DPD, PPD, DDMC, DMB, PDMB, OTDC, STF, ITF, ITMF, IDMBTC, TC, ATC
Rhizocorallium irregularis DPD, PPD, CB
Rosselia rotatus STF, DMB, ATC
Siphonichnus isp. DMB, ATC
Skolithos verticalis DDMB, DMB, PMB, IDMBTC, TC
Taenidium serpentium PPD
Taenidium isp. PPD, PPD, PMB
Teichichnus rectus PPD, PPD, PMB, STF, TC
Teichichnus zigzag PPD
Schaubcylindrichnus isp. PPD
Thalassinoides suevicus PPD, PPD, STF, ATC, TC

Trace fossils are identified from cross-sections in core, and identifications are tentative to a variable degree.
242 D. McILROY

Fig. 4. (a) Chond/Skol ichnofabric showing colonization from the slightly erosive base of a sandstone. The
traces figured are thinly lined small Skolithos (sk) and cross-sections of the horizontal portion of Chondrites
(ch), which is typically low-density. Note the shallow-tier burrow mottling, (b) Diplo ichnofabric showing
colonization of the upper surface of a parallel-laminated sandstone (DMB facies) by abundant small
Diplocraterion (di) and overlain by a heavily bioturbated mudstone with Planolites (pi) and Thalassinoides (th).
(c) Gyro/esc ichnofabric showing a sparsely bioturbated sand-rich succession with abundant clay-draped ripple
cross-laminations and Gyrochorte (gy). (d) Detail of P. herbi ichnofabric with abundant Palaeophycus heberti
(he) showing the diagnostic thick-walled burrows in longitudinal and horizontal cross-section. The burrows are
dark brown coloration due to sideritization during diagenesis. Width of core 10cm; all cores displayed with
oldest strata bottom left.

Chondrites I Skolithos ichnofabric (Chond/Skol) typically composed entirely of shallow-tier


(Fig. 21a) traces, sometimes with discernible Planolites
The Chondrites /Skolithos ichnofabric found in isp. (Fig. 4a)
association with coarse tidal channel sandstones Associated ichnofabrics include the Thalassi-
is - like the Trichichnus ichnofabric described noides /Chondrites (Thal/Chond) ichnofabrics,
below - primarily a post-depositional coloniza- which may even be interbedded within the same
tion ichnofabric. However, unlike the Trichich- channel-fill succession.
nus ichnofabric, the fluid mud deposits that
directly overlie the Chond/Skol horizons are
commonly heavily bioturbated (see Phyco, PI/ Diplocraterion ichnofabric (Diplo) (Fig. 2la)
Tae, Pl/Phyco, Pl/Thal below). The ichnofabric The Diplocraterion ichnofabric is a post-deposi-
is a secondary ichnofabric that normally over- tional ichnofabric composed of low to moderate
prints a burrow-mottled ichnofabric, which is intensities of bioturbation and small burrow
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 243

width (3—5mm). The ichnological assemblage with the eponymous trace fossils are Phycosiphon
is generally monotypic (Fig. 4b), but may be and more rarely Asterosoma isp.
overprinted by trace fossils in the overlying
mudstone unit. The Diplocraterion-bearing Palaeophycus heberti ichnofabric (P. herbi)
sandstones are commonly overlain by moder- (Fig. 21a)
ately to highly bioturbated sandy heterolithic Trace fossil assemblages composed exclusively of
facies, especially PljTeich, PI I Thai and PI/ Palaeophycus heberti are uncommon in the lie
Phyco ichnofabrics. Formation, but may be of great stratigraphic
The Diplocraterion ichnofabric is rather significance. Ichnofabrics containing the trace
restricted in sedimentological range, being con- fossil P. heberti are normally found in prodelta
fined to the central to distal portion of distribu- and distal mouth-bar heteroliths (P. herbi/
tary mouth-bar facies. The Diplocraterion show Schau ichnofabric), but the trace fossil itself is
particular affinity for the parallel-laminated generally completely absent from the more
sandstones that are inferred to be deposited proximal heterolithic facies such as tidal flats
from a buoyant sediment plume by suspension and tidal channels.
fallout (see Brettle et al 2002). The Diplocrater- A single horizon of mudstone completely bio-
ion are exclusively protrusive in nature (i.e. the turbated by P. heberti within a channel fill succes-
latest formed U-tube is the deepest and sand- sion (Fig. 4d) could therefore be interpreted as
infilled). The trace fossils are also uniformly reflecting a marked (albeit temporary) shift in
small, which is taken to support the inter- palaeoenvironmental conditions. However,
pretation of a single phase of opportunistic given that the facies above and below this
colonization with the traces of immature organ- horizon are similar, the change in conditions
isms being preserved (cf. Frey & Goldring that allowed development of the intense P.
1992). Dwarfism in suboptimal conditions is heberti ichnofabrics are likely to be of local
also possible, though such an interpretation is rather than regional importance. Either a minor
considered unlikely given the inferred sedimento- flooding event or a phase of abandonment
logical setting (delta front). could be responsible for such a horizon. This
is an excellent example of a case where ichno-
Gyrochorte/escape burrow ichnofabric logical study can supplement sedimentology to
(Gyro/esc) (Fig. 2la) produce improved palaeoenvironmental inter-
Ichnofabrics with abundant Gyrochorte in asso- pretations.
ciation with v-shaped escape burrows (Fig. 4c)
are common only in distributary mouth-bar Palaeophycus hebertijSchaubcylindrichnus
facies of the He Formation. The Gyrochorte/ ichnofabric (P. herbi/Schau) (Fig. 2la)
escape burrow ichnofabric is characteristic of Ichnofabrics containing the thickly lined tubular
ripple cross-laminated sandstones with moderate trace fossils Palaeophycus heberti and Schaub-
to high intensities of bioturbation. Sedimentary cylindrichnus isp. are common in the upper Ror
horizons with the equilibrium-style Gyro/esc to lower He Formation, and are commonly
ichnofabric tend to have low ichnodiversity. A associated with intense bioturbation of hetero-
post-depositional suite may be present, which lithic sands and muds (from 60% to 100%;
typically constitutes either the Pl/smAst or Fig. 5ai,ii). The traces are typically predomi-
Diplo ichnofabrics in depositionally active nantly horizontal with some vertical pipes repre-
mouth-bar settings. Alternatively, the Gyro/esc sented (cf. Frey et al. 1990) and are associated
ichnofabric may be interbedded with the diverse with current ripples, minor wave ripples and
ichnological assemblages of subtidal flats (Ross/ synaeresis cracks. In some cases, the current
Asf) during longer hiatal periods in which ripples can be demonstrated to show mud cou-
subtidal flats are generated by reworking during plets deposited at the ebb and flood slacks. The
delta lobe abandonment. two eponymous traces are typically the earliest
The burrows themselves record the life-action component of the ichnofabric, and comprise
of organisms living in areas of high sedimen- the primary ichnocoenosis. As cross-cutting
tation rates and well adapted to escape from relationships between the two are relatively
burial under sandbeds. The v-shaped escape common - neither is systematically preceded by
burrows in this ichnofabric are probably created the other - they are thus considered contempora-
by bivalve molluscs (in bedding plane expression neous. The primary ichnocoenosis is typically
the trace fossil is Lockeia amygdaloides). Gyro- reworked by numerous small Phycosiphon isp.
chorte burrows are more difficult to interpret, that pervade the whole ichnofabric and vary in
with their distinctive 'm'-shaped morphology in proportion from 10% to 50% as seen in vertical
transverse cross-section. Commonly associated section.
244 D. McILROY

Fig. 5. (ai) Complex ichnofabrics of the lower He Formation with early Palaeophycus heberti (he),
Schaubcylindrichnus (sc) and Thalassinoides (th) that are re-burrowed by a later Phycosiphon (pc) ichnocoenosis.
(aii) Detail of a vertically emplaced specimen of Schaubcylindrichnus (sc). Burrow diameter is 6mm. (b) The
Ophio ichnofabric containing Planolites (pi) and Ophiomorpha (op) colonizes the top of a flat-laminated
medium-grained sandstone with an earlier ichnofabric comprising escape burrows (lo) and Skolithos (sk)
burrows, (c) Ophio/Tae ichnofabrics in sand-rich heterolithic facies with intense bioturbation by Ophiomorpha
irregularis (op), Planolites (pi) Palaeophycus (ph) and Taenidium (ta). (d) OphiojChond ichnofabrics with large
Ophiomorpha nodosa (on) and Chondrites (ch) overprinting an earlier ichnofabric with escape burrows (lo).
Width of core 10cm; all cores displayed with oldest strata bottom left.

Ophiomorpha ichnofabric (Ophio) (Fig. 2la) and Planolites. Cessation of sedimentation is


Within the He Formation there are numerous attributed to autocyclic lobe switching or
examples of upward-fining successions from seasonal changes in sediment supply/sediment
parallel-laminated sandstones with mud couplets discharge rather than regional flooding by allo-
into ripple cross-laminated sandstones, approxi- cyclic means.
mately 10-30 cm thick, with draped ripple
cross-laminations also showing mud couplets Ophiomorpha/Taenidium ichnofabric
(PDMB and DMB). Trace fossils within the (OphiojTae) (Fig. 21a)
parallel-laminated sandstones are restricted to Ichnofabrics with the thinly mud/organic-lined
small escape burrows, rare vertical Skolithos Ophiomorpha irregularis and the meniscate
burrows and horizontal unlined Planolites bur- backfilled Taenidium are common in proximal
rows. A syndepositional PljSkol or P//esc suite distributary mouth-bar settings in the He Forma-
may be cut by this post-depositional Ophio- tion (Fig. 5c) along with Planolites and Palaeo-
morpha (Ophio) suite, which consists of phycus. The bioturbation in this facies is
shallow-tier Planolites and mid- to deep-tier typically intense (60-80%). The diversity of the
thick-walled Ophiomorpha irregularis burrows assemblage is comparatively low, but may be
(Fig. 5b). This ichnolfabric can therefore be supplemented by Planolites and occasionally
separated into bioturbation that occurred at Diplocraterion; indeed it would seem that the
the same time as deposition of sand from a Diplo and Ophio I Tae ichnofabrics are closely
decelerating sediment plume, and by post- related. The palaeoenvironmental conditions
depositional colonization by large Ophiomorpha experienced by this ichnofauna appear to be
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 245

good for both suspension- and deposit-feeding Phycosiphon ichnofabric (Phyco) (Fig. 2la)
organisms. The depositional environment is This heavily bioturbated mudstone facies
probably a high-energy setting with some nutri- consists of an ichnofabric composed entirely of
tion provided within the sediment, associated Phycosiphon burrows. It is found inter bedded
with thin mud laminations. with thick cross-bedded upward-fining tidal
channel sandstones (TC) in thick mudstones
Ophiomorpha/Chondrites ichnofabric interpreted as fluid-mud deposits. Phycosiphon
(OphiojChond) (Fig. 2la) burrows have a thick halo of siltstone surround-
Coarse-grained upward-fining sandstone succes- ing the mud-filled burrow core and do not cross-
sions with thick micaceous drapes are typical of cut themselves (Fig. 6b). The monotypic nature
the inter-distributary mouth-bar tidal channels of this ichnofabric suggests opportunistic coloni-
(IDMBTC) and some of the tidal channels zation of fluid mud deposits.
(TC) in the He Formation. In some of these a dis- From studying the facies distribution of
tinctive and easily recognizable ichnofabric is Phycosiphon in the He Formation it is clear that
developed, which is identified by the presence the ichnogenus is not tolerant of low salinity.
of large deep-tier Ophiomorpha nodosa burrows Phycosiphon is generally absent in tidal channel
that have thick pellet-lined walls to both their facies, which would have had brackish waters.
horizontal and vertical shafts. The Ophiomorpha The exception is this ichnofabric (Fig. 6b),
nodosa are associated with, but commonly where the Phycosiphon colonize thick fluid mud
overprinted by, similarly deep-tier Chondrites. deposits. The fluid muds are inferred to have
The OphiojChond ichnofabric may overprint an been deposited in the turbidity maximum and
earlier Ophio ichnofabric with Ophiomorpha irre- are intimately associated with the mixing zone
gularis owing to changing palaeoenvironmental (see discussion in the sedimentology section
conditions (see Fig. 5d). This ichnofabric is also above). It may therefore be the case that this
associated with the ChondjSkol ichnofabrics ichnofabric is developed opportunistically when
typical of some tidal channels (TC) in the He the turbidity maximum retreats landward (for
Formation, possibly as a result of minor hiatuses example during low discharge periods such as
in intra-channel deposition. There are similarities summer). Tidal channels deposits with Phycosi-
to the estuarine Ophiomorpha ichnofabrics of phon-bioturbated fluid muds are thus interpreted
Pollard et al. (1993), though the authors did to be formed at the distal (i.e. marine) end of the
not specifically discuss deltaic occurrences of tidal channel system or in channels subject to
Ophiomorpha. anomalous conditions such as those that prevail
during a marine flooding event or periods of
Phoebichnus ichnofabric (Phoe) (Fig. 2la) anomalously low discharge.
Ichnofabrics containing Phoebichnus are' un-
common in the He Formation on Kristin Field Planolites ichnofabric (Planol) (Fig. 2la)
but are abundant in the upper portion of the Planolites ichnofabrics (Planol} are composed
upper Ror Formation, where they are typically exclusively of large sand-filled Planolites, are
found at the top of high-order shoaling cycles. common in the channelized facies of the lie
Phoebichnus ichnofabrics are typically intensely Formation (TC), and are intimately associated
bioturbated and comprise a diverse ichnofauna with fluid mud deposits as described in the
composed entirely of fully marine ichnotaxa sedimentology section above.
(Fig. 6a). Dominant among these are Thalassi- The intensity of bioturbation is highly variable
noides, Phycosiphon, Planolites and Chondrites. (typically between 60% and 80%), and is made
The depositional environment in which the up of a large sand-filled species of Planolites,
ichnofabric is developed is inferred to be a which is probably P. beverlyensis (Fig. 6c). The
normal marine setting with moderate to low mudstones, where preserved between the
rates of sedimentation in the proximal to distal burrows, are characteristically well laminated
prodelta (PPD-DPD). with either parallel laminations or small ripple
The significance of documenting offshore cross-laminations defined by silt or sandstone
ichnofacies genetically related to reservoir units stringers. Colonization occurred subsequent to
is that it allows us to recognize the full scope of deposition and followed early lithification of
biogenic fabrics, such that when they are inter- the fluid mud, as the burrows commonly
bedded with reservoir facies we can estimate the have very sharp margins and can be almost
amount of flooding represented. This improves uncompressed. This suggests that the mud-
understanding of the flooding events and their stones were deposited in a hostile environment
likely impact on facies architecture. without bioturbators, before conditions (perhaps
246 D. McILROY

Fig. 6. (a) Phoebichnus ichnofabric showing intense bioturbation and high ichnodiversity typical of proximal
to distal prodelta settings. Traces figured include Phoebichnus (po), Thalassinoides (th), Phycosiphon (pc) and
Chondrites (ch). (b) Phycosiphon ichnofabric from a 3 cm thick fluid mud deposit within a tidal channel
succession. The trace has very little contrast between the fill and halo and is difficult to photograph, (c)
Succession including two fluid mud deposits showing bioturbation of firm fluid mud deposits. The lower is a
monotypic Planol ichnofabric with large idiomorphic Planolites (pi), whereas the upper mudstone contains
Planolites (pi) and Thalassinoides (th), and is an example of the PIj Thai ichnofabric. The upper sand is an
example of the OphiojChond ichnofabric. The upper sandstone bed contains Ophiomorpha (on) and Chondrites
(ch) of the (OphiojChond) ichnofabric. (d) Ichnofabric composed of v-shaped escape traces (lo; probably
produced by bivalve molluscs) and unlined horizontal tubular burrows of Planolites (pi). The primary P//esc
ichnofabric is overprinted by a Pl/smAst ichnofabric. Width of core 10cm; all cores displayed with oldest strata
bottom left.

salinity or turbidity) ameliorated to allow a low-diversity trace-fossil assemblage composed


colonization. entirely of escape burrows and Planolites (Fig.
6d) are common throughout the He Formation.
Planolites/escape burrow ichnofabric (P//esc) The low ichnodiversity may be a function of
(Fig. 21a) slightly reduced salinity but is mostly the result
Primary equilibrium-style ichnofabrics com- of high rates of sedimentation. The trace fossils
posed of ripple cross-laminated sandstone with are the result of the equilibrium fauna of the
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 247

fades that they are found in, i.e. they are that colonizes laminated unbioturbated muds
syndepositional and may be overprinted by as described above, but is also present in
post-depositional ichnofabrics (Fig. 6d). medium-grained sandstones.
In the present case, initial colonization of the
Planolites/Phycosiphon ichnofabric (PljPhyco) fluid mud by a Planolites (Planol) ichnofabric
(Fig. 21b) occurs soon after deposition by subhorizontal
This composite ichnofabric is found in associa- Planolites-making organisms that created open
tion with heterolithic fluid mud deposits within burrows that subsequently became sand-filled.
upward-fining channel sandstones (TC), in The early Planolites ichnofabric was then re-
which the fluid muds are initially colonized by burrowed by Phycosiphon that reworked the
a Planolites ichnocoenosis (Planol ichnofabric) Planolites burrows and the intervening (semi-
that is subsequently re-burrowed by a low-den- lithified) fluid mud deposits. This ichnofabric is
sity Phycosiphon ichnocoenosis (low intensity comparatively rare and may represent a transi-
Phyco ichnofabric) (Fig. 7 a). The latter ichnocoe- tional environment in which the fluid muds at
nosis is similar to the Phycosiphon ichnofabric the proximal end of the turbidity maximum

Fig. 7. (a) Intra-channel heterolith comprising small Planolites (pi) and a low-density Phycosiphon (pc) trace
fossils, interbedded with thick, coarse-grained, trough cross-bedded sandstones, (b) Pl/Pph ichnofabric in a
clean, well-sorted sandstone overlying a fluid mud deposit with a Sipho ichnofabric with Siphonites (si), and cut
by an erosive-based sandstone with climbing ripples. The Pl/Pph ichnofabric is dominated by Planolites (pi)
and Palaeophycus (ph), which in this example has destroyed all primary laminations and is thus purely a
biogenic fabric, (ci) Cross-bedded sandstone showing neap-spring cyclicity and a v-shaped escape burrow
centred upon a vertical burrow and attributed to Skolithos (sk) (Monocraterion morphology), (cii) Bedding
plane view of a vertical Skolithos burrow showing the concentric cone-in-cone laminations diagnostic of
Skolithos (sk) (Monocraterion). (d) Detail of Pl/Schau ichnofabric developed in a heterolithic ripple cross-
laminated sandstone. Trace fossils present are Planolites (pi) and Schaubcylindrichnus (sc). Width of core 10cm;
all cores displayed with oldest strata bottom left.
248 D. McILROY

zone are initially colonized by the normal chan- in low ichnodiversity and low intensity of
nel fauna (producing the Planolites ichnofabric bioturbation.
described above) during landward penetration
of the salt wedge - perhaps during periods of Planolites I Schaubcylindrichnus ichnofabric
decreased flow (Planolites ichnofabric described (Pl/Schau) (Fig. 21b)
above) - and then, with further decrease in Ichnofabrics in which the major contributors to
discharge, the euryhaline Phycosiphon trace- the fabric are Planolites and Schaubcylindrichnus
maker was able to colonize the sediment. are comparatively rare in the He Formation.
They may be found in heterolithic facies com-
Planolites I Palaeophycus ichnofabric (Pl/Pph) posed of interbedded ripple cross-laminated
The predominant component of this ichnofabric sandstone and burrow-mottled mudstone. Bio-
is a burrow-mottled background bioturbation turbation intensity is comparatively intense,
with poorly defined horizontal burrows without constituting 50-70% of the sedimentary fabric.
linings - referred to Planolites isp. - along with Ichnodiversity is moderate and includes Plano-
a lower proportion of Palaeophycus burrows lites, Phycosiphon, Schaubcylindrichnus and
that are defined by a thin clay lining to the rarely Palaeophycus heberti (Fig. 7d). The
outer margin of the burrow (Fig. 7b). The low ripples in this facies are asymmetrical and clay-
ichnological diversity but high intensity of bio- draped with a sparse, syndepositional escape
turbation (80-100%) is typical of a stressed burrow ichnofabric. The moderate ichnodiver-
sedimentary environment with low rates of sedi- sity and moderate intensity of bioturbation sug-
mentation. The burrows that make up the ichno- gest an equable palaeoenvironment.
fabric are both likely to have been made by The inferred palaeoenvironmental setting for
deposit-feeding organisms, which suggests a this ichnofabric is thus one of a distal distribu-
high primary organic content. As this ichnofacies tary mouth-bar (DDMB) in which the effluent
is interbedded with more classic mouth-bar facies currents are fairly weak, and near-normal
it is interpreted along with them as a delta front marine conditions prevail.
deposit. The apparently low rates of sedimenta-
tion in coarse-grained sandstones may suggest Planolites/Thalassinoides ichnofabric (PI/Thai)
colonization of temporarily abandoned parts of (Fig. 21b)
the delta front, with diversity perhaps being The PII Thai ichnofabric is found in association
restricted by lowered salinity or simply very with tidal channel complexes in the He Forma-
high faunal densities - i.e. amensalism. tion, and is formed in firmground mudstones.
The fabric is composed mainly of Planolites
Planolites I Skolithos ichnofabric (Pl/Skol) ispp. including large (P. beverleyensis) and
(Fig. 21b) small (P. montanus), along with large, slightly
Coarse- to medium-grained sandstones with oval, sand-filled burrows with a laminated
thick micaceous mud-drapes are common in the draft-infill interpreted as shallow-tier firmground
transitional zone between tidal channel (TC) Thalassinoides ?suevicus (Fig. 8a). In some cases a
and inter distributary mouth-bar tidal channel late Phycosiphon-dominated (Pl/Phyco) ichno-
facies (IDMBTC). The sandstones in these set- fabric can overprint this primary ichnofabric.
tings are commonly rapidly deposited and show This ichnofabric is characteristic of the outer
a primary P//esc ichnofabric (as described reaches of the tidal channels (Fig. 12), and is
above) that is overprinted by, or contempora- often associated with Chond/Skol- and Trich-
neous with, a Planolites/Skolithos ichnofabric. bioturbated sandstones (Fig. 4a & 9d).
The Pl/Skol ichnofabric is a syndepositional
fabric in which vertical Skolithos burrows and Planolites ITaenidium ichnofabric (Pl/Tae)
Planolites burrows were contemporaneous with (Fig. 21b)
deposition. This is evidenced in some cases by Interbeds of 2-3 cm thick silty mudstone may
the presence of equilibration behaviour of Sko- rarely be found in association with proximal
lithos trace-makers producing Monocraterion distributary mouth-bar facies (PDMB). These
(Fig. 7ci, ii). The interpretation of this facies as mudstones are interpreted as fluid mud deposits,
occurring in the distal (seaward) part of the which have been formed right out at the distri-
tidal channel is supported by the observation butary mouth through mixing of seawater and
that it is commonly overprinted by the Ophio- clay-rich freshwater. The fact that this is a
morpha ichnofabric (see also Fig. 5b). The comparatively rare ichnofacies suggests that the
palaeoenvironmental conditions inferred for turbidity maximum - in which these mudstone
this setting are slightly reduced salinity and deposits are formed - is rarely forced out as far
high rates of sedimentation, which is reflected as the delta front by high fluvial discharge.
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 249

Fig. 8. (a) Thick mud with abundant sand-filled Planolites (pi) and ?Thalassinoides (th), constituting the
PljThal ichnofabric. (b) PljTae ichnofabric in a thin fluid mud deposit showing moderately intense bioturbation
by Planolites (pi), Palaeophycus (ph) and Teichichnus (te). (c) Pl/smAst ichnofabric with small Asterosoma isp.
(sm) in a ripple cross-laminated sandstone with escape traces, (d) Rosselia/Asterosoma ichnofabric dominated
by Rosselia rotatus (ro) with the diagnostic meniscate fill of the subvertical pipe. Width of core 10cm; all cores
displayed with oldest strata bottom left.

These mudstones, once deposited, are typically finely laminated thin concentric walls and a
nutrient-rich and thus, after deposition, are comparatively wide central sand-filled burrow
heavily bioturbated by Planolites and meniscate (3-5 mm in diameter) (Fig. 8c) and are mor-
burrows in the equable conditions of the delta phologically distinct from the more classic
front by an ichnofauna comparable to that of morphologies of the Asterosoma found in the
the PljPph ichnofabric in adjacent sandstones Rosselia/Asterosoma ichnofabric described
(Fig. 8b). below (Fig. 8d). Accessory traces also described
The ichnofauna itself comprises an opportunis- from this ichnofacies include rare Teichichnus,
tic colonization ichnofauna of Planolites, Palaeo- which is otherwise recorded only from the PI/
phycus, Teichichnus and Taenidium. Although Tae ichnofabric (described above) in fluid mud
this is not a diverse or complex ichnofauna, its deposits deposited near the distributary mouth.
intensity does reflect equable conditions. The The absence of Rosselia distinguishes this from
range of traces demonstrating a deposit-feeding the Ross/Ast ichnofabric, which is an important
mode of life of the majority of the infauna component of subtidal flat facies.
distinguishes it from other associated PDMB
ichnofabrics. Rosselia/Asterosoma ichnofabric (Ross/Ast)
(Fig. 21b)
PlanolitesI small Asterosoma ichnofabric Variably sand-rich current-rippled heterolithic
(Pl/smAst) (Fig. 21b) sandstones are common in the He Formation
The primary ichnocoenosis comprises an intense and can be found as part of upward-coarsening
monotypic fabric generated by Planolites cf. packages that do not pass into mouth-bar
montanus and escape burrows, which is of vari- facies with tidal channels but rather into inter-
able intensity (ranging between 5% and 90%). tidal flat deposits in complete uninterrupted
The Planolites are small in comparison with the successions. The ichnofauna of these subtidal
P. beverlyensis that make up the idiomorphic flats is more diverse than that of similarly hetero-
Planol ichnofabric. The shallow-tier Planolites lithic distributary mouth-bar facies (DMB). The
can commonly be demonstrated to be cut by ichnofabric is ubiquitously composed of a
small, poorly formed Asterosoma isp. that have primary equilibrium escape-burrowed fabric in
250 D. McILROY

the ripple cross-laminated sandstones, which is Siphonichnus ichnofabric (Sipho) (Fig. 21b)
overprinted by a colonization fabric of mid-tier Sipho ichnofabrics are characterized by intensely
Asterosoma and shallow-tier Rosselia rotatus bioturbated horizons (80-100%) with low-
(Fig. 8d), sometimes with accessory Arenicolites. diversity associations of small Siphonites of
Both Asterosoma and Rosselia are interpreted as around 5-8 mm diameter, commonly with Plano-
mud-lined burrows of detritus-feeding organisms lites. The eponymous trace fossil is a vertical
and dominate the ichnofabric. Interestingly, the trace with a meniscate-lined tube wall in which
only occurrences of Rosselia in the He Formation the meniscae are orientated convex down (Figs
are in demonstrably subtidal settings, supporting 7b, 9a). The original description of this trace
the empirical observations of the author based attributed these meniscae to protrusive burrow-
on the ichnology of the similarly tide-dominated ing into the sediment through the life and
Lajas and Tilje Formations that Rosselia is a growth of a bivalve (Stanistreet et al. 1980),
good indicator of subtidal settings, particularly though the present material differs somewhat
subtidal flats. from the type material in being thinner, deeper

Fig. 9. (a) Vertical Siphonichnus (si) originating from a mud-rich abandonment horizon (Sipho ichnofabric)
and cutting through it. Notice also the large 'crab burrow' disturbing the horizon at the bottom of the
photograph, (b) ThaljChond ichnofabric with small Thalassinoides (th) developed from a burrow-mottled
horizon in a tidally bundled trough cross-bedded tidal channel sandstone with rare Planolites (pi) burrows,
(c) Intensely bioturbated DDMB/OTDC facies comprising heterolithic ripple cross-laminated sandstones and
mudstones with large Thalassinoides (th) along with Phycosiphon (pc), overprinting an earlier ichnofabric with
escape burrows (lo), Planolites (pi) and Asterosoma (as), (d) Sparsely bioturbated cross-bedded sandstones with
isolated mud-filled Trichichnus (tr). Width of core 10cm; all cores displayed with oldest strata bottom left.
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 251

and more curved. The ichnofabric itself is Trichichnus ichnofabric (Trich) (Fig. 21b)
uncommon in the He Formation and, where This ichnofabric is found exclusively in asso-
present, is commonly found to be associated ciation with beds of coarse to very coarse sand-
with facies shifts attributable to sudden stone with granule/pebble lags in upward-fining
reduction in sediment supply. Two examples successions 4-6 m in thickness. These sands
of such a scenario are tidal channel abandon- may also contain 1-10 cm thick beds of homo-
ment and delta lobe abandonment: cases of geneous, unbioturbated mudstone layers. The
both are suggested to have occurred in the He ichnofabric itself is composed entirely of mud-
Formation, and are both probably autocylic in filled Trichichnus burrows with colonization
nature. surfaces at sand-sand contacts. Levels of bio-
Owing to the hiatal nature of the surfaces asso- turbation are typically low, seldom exceeding
ciated with Siphonichnus, bioturbation intensity 10% of the sedimentary fabric (Fig. 9d). The
tends to be great and ichnodiversity typically paucity of bioturbation and the monotypic
increases markedly from the same surface nature of the assemblage suggest that environ-
upward into the abandonment facies. mental conditions during deposition of this
facies were particularly harsh, and allowed colo-
Thalassinoides I Chondrites ichnofabric nization only during restricted periods.
(ThaljChond) (Fig. 21b) The broad-scale upward-fining packages in
Ichnofabrics composed of small, deep-tier Thalas- which this facies is found suggest that it was
sinoides and penecontemporaneous Chondrites deposited in a tidal channel that - given the evi-
are unusual components of tidal channel facies dence for mixing of fresh- and saltwater provided
in the He Formation. They are normally post- by the presence of fluid muds - was strongly
depositional ichnofabrics associated with the influenced by freshwater and would have been
upper surface of coarse-grained cross-bedded variably brackish during the tidal cycle (see
sandstones with tidal bundling (see Fig. 9b). below). Thus the major controls on the infauna
The presence of Thalassinoides in tidal channel are likely to have been current strength and
sandstones is unusual. Its close relative Ophio- either periodically or permanently reduced sali-
morpha is more common in this setting as its nity. These factors served to exclude other
pelletal wall lining is an adaptation for life in trace-making organisms from this facies, with
mobile substrates. The presence of Thalassinoides the Trichichnus utilizing rare colonization win-
could therefore be taken to indicate that the dows afforded by amelioration of environmental
sandstones containing this ichnofabric may conditions. The absence of bioturbation in the
have undergone early diagenesis or were nutrient-rich fluid mud deposits highlights this
otherwise particularly firm at the time of as a particularly hostile palaeoenvironment,
burrowing. The well-developed shallow-tier even harsher than that of the PIj Thai ichnofabric
burrow-mottling, found as part of the same described above.
ichnofabric, would appear to support the infer-
ence of low sedimentation rates and a hiatus Problematic ichnofabric (Probl)
prior to development of the ThaljChond ichno- The top of the He Formation is characterized by
fabric. This ichnofabric is typical of tidal channel a distinctive ichnofabric, containing abundant
facies in the He and is commonly associated with Thalassinoides burrows, that is reworked by a
the ChondjSkol ichnofabric. small, unnamed, mud-filled trace of variable
morphology that commonly shows numerous
Thalassinoides I Phycosiphon ichnofabric upward-directed projections from a curved sub-
(ThaljPhyco) (Fig. 21b) vertical stem. Other traces include resting traces
Thalassinoides I Phycosiphon ichnofabrics are Rhizocorallium irregulare and Teichichnus isp.
typically intensely bioturbated with a diverse (Fig. 10). The mud-rich nature of this facies
ichnofauna dominated by the Thalassinoides invites comparison with the central basin of
and Phycosiphon along with Palaeophycus a wave-dominated estuary (Fig. 11), but this
heberti, Schaubcylindrichnus and Palaeophycus remains somewhat tentative, as is discussed in
isp. (Fig. 9c). The ichnofabric is found in hetero- more detail in the sedimentology section below.
lithic ripple cross-laminated sands interpreted to
have been deposited in the distal distributary
mouth-bar to proximal prodelta region. The Sedimentology of the lie Formation on
intense bioturbation and high ichnodiversity Kristin Field
identify this ichnofabric as characterizing a
fully marine palaeoenvironment with good Sedimentary facies of the He Formation can be
living conditions for the infauna. grouped according to broad-scale sedimentary
252 D. McILROY

almost identical, but their relative volumetric


importance differs between the two systems.
What is remarkable is that, despite the low
tidal range, sedimentation of the He Formation
on Kristin Field is still dominated by tidally
modulated flow. This is taken to indicate that
the basin was sheltered from wave action and
probably had a comparatively small width in
the direction of the prevailing winds.
Through sedimentological assessment of the
He Formation carried out during this study, a
conceptual model for the spatial distribution of
the observed sedimentary facies has been con-
structed (Fig. 12). It must be noted, however,
that the diagram represents a freeze-frame and
does not truly represent the extent of sedimen-
tary facies in ancient deposits (e.g. meander
belts extend below the tidal flats). The diagram
does not account for the relative preservation
potential of these facies. For example, in a tidal
deltaic setting the tidal channels will - with
progradation - cannibalize their own distribu-
tary mouth-bars (Fig. 13), resulting in a delta
top sand sheet of amalgamated tidal channels
(cf. Monahan et al. 1993).
The major sedimentary facies associations
documented from the He Formation are tidally
influenced fluvial/tidal channels (tidal currents
being suppressed by riverine outflow), tide-domi-
nated intertidal to subtidal flats, and distributary
mouth-bar through delta front/prodelta to off-
shore (Fig. 12). These facies associations can be
broken down further through careful sedi-
mentary analysis and further subdivided by a
combined sedimentological and ichnological
Fig. 10. Possible estuarine facies with abundant approach (see Fig. 14).
bioturbation by Thalassinoides (th), Rhizocorallium
(rh) and an unidentified small feathery mud-filled The full range of sedimentary facies distin-
burrow (fy). The core is 10cm wide. guished during this study is presented below
(see Table 2).

environment and subdivided with respect to


sedimentary facies, which in turn can be split 1. Delta front palaeoenvironments
into subfacies based on a combination of their
sedimentology and ichnology. Tidal range can The delta front includes the most distal facies
be estimated where continuous successions represented in the He Formation on Kristin
from subtidal to supratidal facies can be deter- Field, with the highest ichnodiversity and some
mined (i.e. the stratigraphic thickness from sub- of the highest intensities of bioturbation
to supratidal approximates tidal range) (Klein recorded. Sedimentation of these deltaic facies
1971). In the case of the He Formation on Kristin occurs though rapid deposition of sediment at
Field intertidal facies are scarce but, where they the interface between the fluvial and marine
are present, the thickness of intertidal facies is realm. The most distal facies represented are
seldom more than 2-3 m, suggesting a microtidal strongly heterolithic and variable in sandstone
regime. content. Superficially, this facies association
The He Formation comprises a diverse array of resembles the offshore transition zone of classic
sedimentary palaeoenvironments similar to those wave-dominated systems, but the sediments are
described from the Jurassic Lajas Formation of devoid of the diagnostic hummocky cross-strati-
the Neuquen Basin, Argentina (Mcllroy et al. fication (HCS) (cf. Dott & Bourgeois 1982), and
1999). The range of sedimentary facies is contain indicators of tidal processes such as mud
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 253

Fig. 11. Idealized model for distribution of fades in a wave-dominated estuary (redrawn from Dalrymple et al.
1992). This depositional setting is only one of several protected back-barrier scenarios; determining the most
likely will require a regional perspective beyond the scope of this work.

couplets and thinly laminated beds showing tidal Formation contain a low-diversity ichno-
bundling. coenoses indicative of stressed conditions. The
Distinguishing between shoreface and deltaic variable salinity and high sedimentation rates
environmental settings is critical for realistic reser- encourage opportunistic colonization by organ-
voir modelling of the He Formation (see also Bann isms with a generalist feeding strategy and
& Fielding 2004). Shoreface settings should con- trace-making organisms with a tolerance for
tain extensive coast-parallel reservoir sandstones, salinity stress. This environmental diversity also
whereas a tide-dominated delta will contain accounts for the disparity of ichnofabrics found
numerous smaller sandstone bodies oriented per- in this facies.
pendicular to the coastline, with the concomitant Ichnofabrics associated with this sedimentary
implications for production planning. facies are Chond/Skol, Ophio, OphiojTae, Ophioj
Chond, Pl/Skol, Pl/Tae, Pl/Pph and Planol, as
la. Proximal distributary mouth-bar fades described in the ichnofabrics section above.
(PDMB)
This sedimentary facies is characterized by paral- Ib. Central distributary mouth-bar facies (DMB)
lel-laminated to trough cross-bedded medium- to Medium- to fine-grained sandstones with clay
coarse-grained sandstones, commonly associated drapes, ripple cross-laminations and abundant
with micaceous mudstone horizons up to 10cm escape burrows are characteristic of this facies
thick (Fig. 15a). Bioturbation is irregular in (Fig. 15b). Climbing ripples and occasionally
distribution and is commonly either intense or herringbone cross-lamination are recorded from
absent. The most common trace fossils are this facies, which - like PDMB - is largely
Skolithos, Ophiomorpha, Chondrites, Trichichnus, sand-dominated with occasional thin (and some-
Piano lit es, Palaeophycus and Taenidium. times highly micaceous) mudstone horizons. The
In distributary mouth-bar settings, currents are ichnofauna is typically more diverse than in
ebb-dominated and salinities may be extremely proximal mouth-bar settings (PMB), and bio-
variable, dependent on effluent discharge. Sedi- turbation is more constant in its intensity at
mentation rates are commonly high owing to the 30-50%.
rapid deceleration of ebb currents upon meeting In the central part of distributary mouth-bars,
the effectively stationary marine waters, and floc- deceleration of the current may allow deposition
culation of clays is common (e.g. Gibbs 1977; of both the tractional and suspended load (e.g.
Martinius et al. 2001). The net effect of these Wright 1977). The sediment plume emitted
palaeoenvironmental conditions is that facies from the feeder channel can be either buoyant
deposited in such environments in the He or dense depending on suspended load and
Fig. 12. Conceptual fades model for the relative positions of fades in the tide-dominated deltas of the He Formation on Kristin Field. Acronyms are facies
abbreviations as used in the text.
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 255

Fig. 13. Model to explain the sharp juxtaposition of tidal channel fades (lie Formation) upon mouth-bar
fades (Ile-Ror Formation) in relation to the interplay between available accommodation space and channel
incision. Diagrams a-d and e-h are in time-stratigraphic order with the oldest at the base, (a-d) Show
autocyclic development of a delta front in a low accommodation space setting with a sharp grain-size change
expected between delta front and feeder channel as seen in the tide-dominated deltas at the base of the He
Formation, (e-h) Represents a similar, but high accommodation, setting in which more of the delta front is
preserved as seen in the Lajas Formation, Argentina (Mcllroy unpublished).

salinity contrast between the outflow and marine bioturbated than the intervening mudstones
waters, and is invariably tidally modulated to and siltstones. These heteroliths have Gyro/esc,
some degree (see discussion in Brettle et al. P//esc, Diplo, P. herbijSchau and Pl/Schau
2002). The more distal position of this fades rela- ichnofabrics.
tive to the PDMB is inferred based on the smaller The parallel-laminated sandstone horizons are
size of bedforms and higher ichnodiversity. The interpreted as resulting from deposition by sus-
increased ichnodiversity in this facies probably pension fallout from a decelerating buoyant
reflects the more normal marine salinities found effluent plume, which from the presence of
further out from the distributary channel mud-couplets can be see to be tidally modulated
mouth. In addition, rates of sedimentation are and deposited in a subtidal setting (cf. Brettle
likely to be lower and more episodic (than in et al. 2002). In contrast, the ripple cross-
DMB), which encourages colonization (Leving- laminated sandstones indicate deposition from
ton 1970; Grassle & Grassle 1974; Frey & traction, showing that the depositing flow was
Goldring 1992). highly variable in character (Fig. 15b).
Ichnofabrics associated with this sedimentary It must be emphasized that thinly interlami-
facies are P//esc, Gyro/Qsc, Pl/smAst and Sipho nated sandstones with clay layers can look similar
(following lobe switching) as described above. to hummocky cross-stratification (HCS) in core
because bedding surfaces in both are commonly
Ic. Distal distributary mouth-bar facies (DDMB) very low amplitude and/or long wavelength (cf.
This is a heterolithic facies composed of thinly Dott & Bourgeois 1982; Brettle et al 2002). The
interbedded fine-grained ripple cross-laminated physical sedimentological processes responsible
and parallel-laminated sandstones interbedded for these two bedform styles are quite distinct,
with bioturbated siltstones and mudstones (Fig. however, and allow discrimination between
15c). The proportion of sandstone in this facies wave- and tide-dominated deltaic settings.
is defined as being less than 50%. Bioturbation Hummocky cross-stratification is not exclusive
is moderate to intense and ichnodiversity is to wave-dominated depositional systems; indeed
typically comparatively great (60-90%). The HCS can occur in estuarine and lagoonal settings
parallel and ripple cross-laminated sandstones during exceptional events. However, its general
both contain mud-couplets and are much less absence in offshore heterolithic facies of the He
Fig. 14. Conceptual fades model for the relative positions of fades and ichnofabrics in the tide-dominated deltas of the He Formation on Kristin Field. Acronyms are
ichnofabric abbreviations as used in the text.
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 257

Table 2. Palaeoenvironments and fades of the He Formation along with fades


abbreviations as used in the text

Inferred environment Fades Abbreviation


of deposition

Deltaic Central distributary mouth-bar DMB


Distal distributary mouth-bar DDMB
Proximal prodelta PPD
Proximal distributary mouth-bar PDMB
Anomalous massive bedded sandstone
Tidal flats Intertidal mud flats ITMF
Intertidal sand flats ITF
Subtidal flats STF
Offshore tide-dominated coastline OTDC
Channels Strongly fluvially influence channels FIC
Tidal channels TC
Inter-distributary mouth-bar channels IDMBTC
Abandoned tidal channel ATC
? Estuarine ?Central basin CB
?Bayhead delta BHD

Formation on Kristin Field precludes the pre- by tidal sedimentation and may be subject to
sence of a normal wave-dominated shoreface. subaerial exposure during part of the tidal
In the present case, the strong tidal signature in cycle. Herein, in line with some earlier authors
the rest of the overlying lie Formation, and the (Reineck 1967; Klein 1971; Weimer et al. 1982),
lack of typical mid-lower shoreface facies with this is extended to cover the subtidal portion of
HCS, are taken as evidence that sedimentation a sedimentological continuum that extends
occurred in the distal parts of a tide-dominated below the reach of even the spring tides. In
delta similar to the Fly River Delta (Alongi most normal marine systems storm-dominated
1991; Harris et al. 1993) and the lower Lajas sedimentation would be expected in this subtidal
Formation (Mcllroy unpublished). regime; however, in the He - and in similar tide-
Characteristic ichnofabrics of this distal distri- dominated systems such as the Tilje Formation
butary mouth-bar sedimentological setting in the (Martinius et al. 2001) and the Lajas Formation
He Formation on Kristin Field include Diplo, P. (Mcllroy et al. 1999) - wave energy is subordi-
herbi, PljSchau and P. herbijSchau. nate to tidal energy even in the shallow subtidal
realm (Fig. 16). The volumetric predominance
Id. Proximal prodelta (PPD) of subtidal facies over intertidal/supratidal
The proximal prodelta is defined herein as the facies in the He Formation on Kristin Field is
region in which biogenic reworking of sediment taken to indicate a small tidal range as discussed
through bioturbation destroys most of the (Fig. 16). This of course assumes incremental
sedimentary laminations. Grain size is typically increases in accommodation space rather than
silt-very fine sand, and the sediment tends to be gradual accommodation space generation,
mud-rich, with mud being mixed into the sedi- which could then be filled by aggrading intertidal
ment by bioturbation. Intensity of bioturbation flats. For example, in a microtidal setting, a 10m
is typically great (90-100%) and ichnodiversity flooding event would result in 10 m of accommo-
is high (Fig. 6a). Ichnofabrics in this setting are dation space, of which approximately the upper
complex and include Phoe and Thal/Phyco as 2m would be filled with intertidal facies and
described above. The occasional thin, fine- the remainder would be composed of subtidal
grained sandstone horizons that are preserved sediments (Fig. 16). In addition, the upper
comprise ripple cross-laminated sandstones portion of a succession is most likely to be subse-
with clay-draped foresets. quently eroded during flooding ravinement or
channel erosion.

2. Tidal flat palaeoenvironments 2a. Intertidal mudflats (ITMF)


This uncommon He facies is composed of inter-
Tidal flats are normally defined as that part of bedded mudstones and fine- to medium-grained
the marine depositional system that is influenced ripple cross-laminated sandstones with neither
258 D. McILROY

Fig. 15. Sedimentary facies of the deltaic facies association: (a) proximal distributary mouth-bar facies
(PDMB) with rip-up clasts and fluid muds at the base and fining upward to parallel-laminated micaceous
sands and silts; (b) central distributary mouth-bar facies (DMB) showing typical transitions from low-angle
cross-laminated sands and muds showing tidal bundling into wave and current ripple cross-laminated
sandstones; (c) distal distributary mouth-bar facies (DDMB) showing intense bioturbation of fine sands with
remnants of ripple cross-laminations and thin fluid mud deposits. Width of core 10cm; all cores displayed with
oldest strata bottom left.

bioturbation nor rootlets (Fig. 17a). The ripples The rarity of this facies in the He Formation is
are primarily oscillation ripples that are inferred probably a function of a variety of parameters
to have been formed by wave action in shallow discussed below, including the inferred micro-
water at states of high tide. This facies is tidal range. The propensity for mudflats to be
known from only two decimetre-thick horizons eroded by both auto- and allocyclic processes
in the He on Kristin field; in both cases it is exacerbates their infrequency. Erosion is due to
found at the top of a blocky or upward-fining either their association with tidal channels or
succession of intertidal sandflat facies in which their position at the top of parasequences
bioturbation, grain size and bed thickness grade where they stand to be eroded either during or
into this facies. after relative sea-level rise (Mcllroy et al 1999).
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 259

Fig. 16. Idealized prograding tidal flat successions in tide-dominated macrotidal settings, tide-dominated
microtidal settings and wave- and tide-influenced macrotidal settings. The variable grain-size profiles and facies
associations can be used to help determine depositional setting.

2b. Intertidal sand flats (ITF) is typically of low intensity and low diversity
This facies comprises thin successions of fine- or (Fig. 17b). Occasional thick clay beds, up to
medium-grained sandstone with trough cross- 5 cm in thickness, are also documented.
bedding, ripple cross-lamination (both wave The range of sedimentary structures found in
and current ripples), and rarer planar-laminated these sandstones is indicative of variable, but
beds. Grain size within a given intertidal sandflat generally high, flow regimes, and they grade
succession is generally uniform but may show into channel-margin deposits. The abundance
upward-fining trends up to 2m thick, passing of escape traces related to the bivalve trace
either into subtidal flats or into intertidal mud- fossil Lockeia is also taken as supportive evi-
flats depending on stratigraphic setting. The dence. The typical intertidal sandflat ichnofacies
foresets of trough cross-beds are commonly, of the He Formation is P//esc with Trich, Sipho
but not always, clay draped, and bioturbation and Chond/Skol sometimes present (described
260 D. McILROY

Fig. 17. Sedimentary facies of the tidal flat fades association: (a) intertidal mudflat facies (ITMF) with parallel
and ripple cross-laminated interbedded mudstone and sandstone; (b) intertidal sandflat facies (ITF) showing a
mineralogically immature sandstone with parallel (upper flow regime planar beds) and trough cross-bedding
interpreted to form adjacent to tidal channels; (c) subtidal flat facies (STF) showing the typical heterolithic
ripple cross-laminated sandstones with clay-draped ripple cross-laminae and abundant diverse bioturbation.
Width of core 10cm; all cores displayed with oldest strata bottom left.

above). However, this facies is also commonly of beds typical of tidal sandflats. The restricted
completely unbioturbated, when it is character- ichnodiversity is probably due mainly to
ized solely on physical sedimentary structures temperature stress on the intertidal flats during
and the rather aggradational stacking patterns low tides, as discussed (cf. Johnson 1965).
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 261

2c. Subtidalflats (STF) controlling (though weak) environmental stres-


Sub tidal flat fades are typified by thinly bedded, ses in both geomorphological settings. Ichnofab-
very fine to medium-grained sandstones, inter- rics documented from the base of prograding
bedded with siltstones and mudstones. The subtidal flat successions include Phoe, P. herbij
sandstones are 2—5cm thick and are wave and Schau and Thal/Phyco.
current ripple cross-laminated, commonly with
mud-couplets on their foresets and abundant
escape burrows. The upper surfaces of sandstone 3. Tidal channel palaeoenvironments
beds are characterized by opportunistic styles of
colonization with ichnofabrics rich in mud-lined The lower-middle portion of the He Formation
burrows, especially Rosselia and Asterosoma, on Kristin Field comprises numerous stacked
that colonize the upper surface following sand upward-fining, coarse- to very coarse-grained,
deposition (Fig. 17c). trough cross-bedded sandstones (Fig. 3) with
The presence of mud couplets and the com- anomalous mudstones/siltstones. These
paratively high ichnodiversity attest to the subti- upward-fining successions have erosional bases
dal setting of this ichnofacies, and the absence of with coarse pebbly lags, sometimes with wood-
HCS precludes deposition in a storm-dominated clasts, and are interpreted as channel sandstones
offshore transition zone (Dott & Bourgeois deposited in meander belts on the delta top. In
1982). The ichnodiversity is greater than is some cases, tidal signatures can be demonstrated
typical of He Formation sandy intertidal flat in the form of tidal bundling and mud-couplets
settings on Kristin Field. The sand-mud propor- (cf. Allen 1981), but more commonly in the He
tion is much lower, and ichnodiversity higher, Formation their tidal origin is betrayed by the
than the lithologically similar distal distributary presence of thick mudstones interpreted as fluid
mouth-bar facies described above. mud deposits (Allen et al 1980; Wolanski &
The ichnofabrics associated with He Forma- Gibbs 1995) and the presence of some marine
tion subtidal sandflat facies are RossjAst, PI/ trace fossils. Fluid mud deposits are interpreted
esc, Pl/Tae and Planol as having formed within the turbidity maximum
of a tidal channel in brackish conditions by rapid
2d. Offshore tide-dominated coastline (OTDC) flocculation (cf. Allen et al 1980; Wolanksi &
This facies is characterized by intense biotur- Gibbs 1995).
bation with rare thin ripple cross-laminated Tidal signatures are not, however, ubiquitous
horizons of fine-grained sandstone. Ichnodiver- within such channel sandstones, and some
sity is high and ichnofabrics are typically rich intervals of sandstone show no physical sedi-
in Phycosiphon (Fig. 17d). mentological evidence that sedimentation was
Although it is clear that, away from the major tidally influenced. This is particularly true of
axes of progradation (deltas), intertidal flats and the most coarse-grained facies within the He
subtidal flats with their distinctive sedimentology Formation. One of the challenges, therefore, is
form the normal prograding coastline (Fig. 12), the detailed characterization of these sandstones
no pre-established model is in existence for how in an attempt to be able to resolve facies-stacking
such facies change offshore in a tide-dominated patterns within multi-storey aggradational flu-
setting (see Fig. 16) and grade laterally into vial/tidal channel sandstones.
prodelta environments. This facies represents Tidal channels can show a range of sedimen-
the equivalent of the offshore transition zone of tary structures characteristic of flow within a
storm-dominated settings. Distinguishing this channel, but few of these as seen in core are diag-
facies from the proximal prodelta is difficult on nostic of high-resolution proximal-distal trends
both ichnological and sedimentological grounds. (Howard & Frey 1973; Frey & Howard 1986).
The complete absence of parallel/low angle lami- According to idealized facies models of marginal
nated (suspension fallout) distal distributary marine tidal channels (e.g. Dalrymple et al.
mouth-bar sandstones is one possible criterion 1992), channels are typically meandering at
(cf. Fig. 15b) but is unlikely to be practical their proximal end, becoming straighter towards
owing to the high intensity of bioturbation in the sea - from which some interpretations about
normal marine settings (see also Bann & Fielding likely facies architecture could be extrapolated.
2004). However, the rheology of the background delta
Ichnofabrics associated with offshore tidal top sediment can strongly dictate the geomor-
coastlines are likely to be similar to those of phology of tidal channel deposits. By analogy
prodelta settings within the same formation. with fluvial systems, channels would be expected
Salinity stress, sedimentation rates and palaeo- to have narrower meander belts in mud-rich
oxygenation are likely to have been the background facies (e.g. Fenies & Faugeres
262 D. McILROY

1998). It should also be noted that the influence simply as tidal channels are described and
of tidal currents can often be felt large distances classified based on combined study of their
into the delta plain through the slowing of physical and biogenic sedimentary structures
riverine flow during spring tides in a macrotidal (Fig. 12).
regime. The landward limit of such tidal effects
is known as the tidal reach and influences the 3a. Fluvially influenced channels (FIC)
sedimentology of tidal channels far beyond FIC are composed of very coarse- to coarse-
penetration of the salt wedge (e.g. Harris et al. grained sandstone facies with thick trough
1993). The penetration of the salt wedge is a cross-bedded sandstone beds, gravel lags with
function of discharge volume versus angle of woody debris, which may be rounded into
the delta plain and tidal range. wood clasts. Neither the foresets nor toesets of
Ichnological characterization of the channel bedforms are mud draped. The sandstones are
sandstones and their contained mudstones/ entirely unbioturbated and show only unidirec-
heteroliths enables identification of a diverse tional palaeocurrents.
array of tidal channel ichnofacies that occur Sandstones of this facies are most easily inter-
along the gradient of hydrodynamic and chemi- preted as proximal tidal channel facies with a
cal conditions found in the transition from fluvial strong fluvial influence. The absence of fluid
through brackish to marine conditions within mud deposits and diagnostically marine trace
such channels. This is due to the sensitivity fossils in these coarse sandstones suggests
of burrowing infaunal organisms to salinity highly reduced salinity or even freshwater condi-
changes (Milne 1940; Dorjes & Howard 1975). tions at the channel floor and deposition land-
With these factors in mind, sharp-based ward of the turbidity maximum (see Schubel
upward-fining successions typically described 1968). The fact that the coarsest sediments in

Fig. 18. Sedimentary facies of the tidal channel facies association, (a) Tidal channel facies (TC) showing
typical trough cross-bedding and thick fluid mud deposits in between bedsets and low intensities of
bioturbation. (b) Modern channel-margin tidal flats of the River Dee Estuary, Wirral, UK, showing partially
reworked fluid-mud deposits formed in the summer of 2001 and both semi-lithified by October 2001. (c) The
same deposits were highly reworked by November 2001 and were common as clasts associated with ebb-tidal
dunes, (d) Succession of cross-bedded tidal channel facies with bioturbated fluid mud deposits overlain by
highly bioturbated abandoned tidal channel facies (ATC). Width of core 10cm; all cores displayed with oldest
strata bottom left.
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 263

the He Formation on Kristin Field are to be


found in this fades also supports its inferred
proximality.

3b. Tidal channels (TC)


This fades comprises medium- to coarse-grained
sandstones organized in upward-fining sedimen-
tary packages with erosional bases and lags of
extra-basinal pebbles, rounded mud clasts and
plant debris. Sandstone beds are generally
trough cross-bedded with clay- or organic
debris-draped foresets. The clay drapes may be
organized into demonstrable tidal bundles,
sometimes with superimposed ripple cross-
laminations. Sandstones may also be interbedded
with 1-10 cm thick mudstone or rippled muddy
siltstone horizons (Fig. 18a). Bioturbation is
typically of low intensity and low diversity,
with a predominance of vertical burrows and
escape burrows in sandstone beds. The ichno-
faunal assemblage of the associated mudstones
is highly variable, showing colonization while
soupground or softground conditions prevailed
(e.g. Phyco, P. herbi and Pl/Phyco ichnofabrics),
or representing firmground assemblages of open
sand-filled burrows (e.g. Planol, PI/Thai ichno-
fabrics) as discussed above (Figs 19, 20).
The presence of such thick mudstones in
association with coarse-grained sandstones is
apparently anomalous. However, sedimentation
of faecal pellets, and rapid flocculation in the
turbidity maximum of brackish environments,
coincident with decreased current velocities
associated with states of high and low tide, can
allow thick layers of mud to accumulate (e.g.
Kranck 1981; Edelvang & Austen 1997). These
water-rich muds, known as fluid muds, can be
up to 1m thick in the Fly River depositional
system (Wolanski & Gibbs 1995) and are
common in most estuaries to some degree. These
fluid mud deposits become rapidly dewatered
and compacted in intertidal settings and undergo
early lithification over a period of weeks (unpub-
lished observations from the Dee Estuary
Cheshire, UK). Erosion within tidal channels
either at the base or on channel margins can
rework such semi-lithified muds into mud clasts
that are commonly seen in the toesets of dune-
forms at the base of tidal channels but also
occasionally on channel-margin tidal flats (Fig.
18b, c).
The ichnology of the tidal channel sandstones
as described above is invaluable in distinguishing
between tidal channel facies and aids recognition
of tidal channel stacking patterns, which are Fig. 19. Conceptual model showing the distribution
notoriously difficult in multi-storey channel of tidal channel ichnofabrics from both the fluid
deposits. Ichnofabrics recognized in the He mud deposits and channel sandstones. Symbols as in
Formation tidal channels are the Trich, Chond, Fig. 3.
Fig. 20. Vertical cross-sections showing multi-storey tidal channel successions and their ichnofabric stacking patterns with interpretation. Based on a composite
database from Kristin Field. Facies hatchings as in Fig. 12.
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 265

P//esc, Thal/Chond, Chond/Skol, Planol, PI/ The ichnofabrics documented from this facies
Phyco, PI I Thai and Phyco ichnofabrics described are transitional between those from proximal
in the previous section (see also Figs 19, 20). delta front mouth-bars and those from more
proximal tidal channels. It is clear that there is
3c. Abandoned tidal channels (ATC) significant palaeoenvironmental stress on the
In the He Formation some upward-fining suc- trace-making infauna of this facies, as evidenced
cessions with erosional bases overlain by trough by the low ichnodiversity and low intensity
cross-bedded sandstones, commonly with of bioturbation (0-30%). The upward-fining
coarse gravel lags and thick mudstone laminae/ nature of the sedimentary packages suggests
drapes 1-5 cm in thickness, are capped by more deposition in a channel setting in distributary
clay-rich facies (Fig. 18d). The sand: mud ratio channel thalwegs between laterally and down-
is generally less than 60% sand, but in some stream accreting longitudinal bars. This is also
cases much higher (approximately 80% sand): borne out by the high flow regime, as indicated
in such cases the sand-rich channel fill is ichno- by trough cross-bedding and planar lamination.
logically rather similar to subtidal flat facies but Lithological and ichnological similarities with
is distinguished by the more variable grain size proximal mouth-bar facies and tidal channel
and slightly lower ichnodiversity. facies suggest a transitional location at the distri-
The sedimentological character of the lower butary mouth, where channel forms are present
portion of an abandoned channel typically between the emergent bars at the delta front.
concords with one of the types of active tidal
channel described above. The marked increase
in intensity of bioturbation associated with 4. ?Estuarine palaeoenvironments
the abandonment event is thought to reflect
avulsion at the proximal end of the tidal channel. The uppermost beds of the He Formation on
Following abandonment, channels are influ- Kristin field comprise an unusual association of
enced primarily by marine waters that are much facies that is not seen at lower stratigraphic
less hostile to animal life, and, as a result, a levels. It is also seen on adjacent fields (e.g.
diverse ichnofauna can develop. This is in Smorbukk). The facies present are inferred to
marked contrast to the salinity stress and high be broadly estuarine in character in that they
sedimentation rates that are typical of more show evidence of harsh palaeoenvironmental
active fluvially connected tidal channels. A simi- conditions - as evidenced by the ichnology as
lar case is currently present in the Fly River described below. In addition the facies typically
Delta, in which the abandoned portion of the overlie a coarse-grained lag that may be inter-
delta contains relict channels that support a preted as either a ravinement lag or an erosional
dominantly marine ichnofauna (Alongi 1991). lag caused by incision during relative sea-level
Ichnofabrics commonly found in association fall. No evidence was available to reliably
with the post-abandonment phase of channel determine which is the case. The facies associa-
fill are Sipho, Phoe, ThaljPhyco, Ross/Ast, tion as a whole is dominated by wave-generated
Planol and Pl/Skol, as described above. bedforms, and suggests that the whole deposi-
tional system may have changed to one in
3d. Inter-distributary mouth-bar tidal channels which wave action was no longer suppressed.
(IDMBTC) The broad-scale depositional setting envisaged
Medium- to fine-grained upward-fining sand- for the facies described below is one of a back-
stone successions with trough cross-bedding barrier environment with significant salinity
and ripple cross-laminations or planar cross- stress, predominantly low current strength and
laminations characterize IDMBTC facies. Fore- perhaps hypoxia, as is common in, but not
sets of most bedforms are draped in highly exclusive to, protected microtidal estuarine
micaceous clay (Fig. 18e). These thick micaceous settings (e.g. Diaz & Rosenburg 1995). A candi-
horizons may be up to 10cm thick in extreme date depositional environment is represented in
cases and are usually unbioturbated or show Fig. 11.
only a Planol ichnofabric (described below)
with large sand-filled Planolites cf. beverleyensis. 4a. ?Central basin (CB)
The foresets of bedforms are largely unbiotur- The uppermost few metres of the He Formation
bated but may show opportunistic colonization are characterized by an unusual sedimentary
ichnofabrics including Chond/Skol, Ophioj facies (Fig. 10), as developed on Kristin,
Chond, Pl/smAst and equilibrium colonization Mikkel, Smorbukk and Sm0rbukk S0r fields
as PI/esc ichnofabrics (see discussion of coloniza- (see Fig. 1) at the transition to the overlying
tion in Mcllroy 2004). Not Formation (Fig. 2). The succession is
266 D. McILROY

Fig. 21. (a) Example ichnofabric constituent diagrams of idealised ichnofabrics from the He Formation on
Kristin Field.
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 267

Fig. 21. (b) Example ichnofabric constituent diagrams of idealised ichnofabrics from the He Formation on
Kristin Field.
268 D. McILROY

broadly upward coarsening and overlies a hori- delta) or as storm wash-over deposits (cf.
zon with evidence for erosion in another Kristin Andrews 1970; Hubbard & Barwis 1976;
well, but is marked by coal deposition in other Carter 1978).
parts of Haltenbanken (Midgard Field; Fig. 1).
The surface itself is typically overlain by a 4b. ?Bayhead delta (BHD)
coarse lag of granule- to pebble-sized extra- Upward-coarsening mud-rich sandstones may
basinal clasts, commonly in a muddy matrix. overlie the mudstones of facies 4a (CB) at the
Alternatively, this surface may be a palimpsest top of the He Formation. The sandstones are
colonization surface with Pl/smAst ichnofabrics largely unbioturbated and show symmetrical
being overprinted by Phyco and P. herbi ichno- oscillation-rippled surfaces. Core recovery is
fabrics. poor in this interval, however, and interpretation
The sediments above the coarse basal lag are is based on well-log signatures and comparison
typically dark, mud-rich and heavily bioturbated with adjacent fields.
by Thalassinoides and a problematic feather-like The dearth of bioturbation and coarse grain
mud-lined vertical burrow (Probl ichnofabric; size of this facies may be attributed to deposition
Fig. 10). There are some thick unbioturbated in a proximal estuarine setting, in which a strong
mudstone horizons, a high proportion of detrital freshwater influence may have discouraged the
organic matter and thin stringers of very coarse infauna (see review in Mangano & Buatois
sandstone. 2004). Given the preferred interpretation of the
The lower bounding surface of this facies underlying sediments as having been formed in a
is erosional on a regional scale and shows a central basin setting, such an upward-coarsening
succession of ichnofabrics that appear to demon- fluvially influenced succession may be interpreted
strate a flooding event. The coarse sedimentary as a bayhead delta.
layer itself could be interpreted as either a
channel lag formed during sea-level fall or a
ravinement lag formed by winnowing during Anomalous massive bedded sandstone facies
sea-level rise. At present this is not possible to
determine through lack of a more regional The last of the facies described from the He
perspective. Formation is an anomalous facies comprising
The overlying sediments are highly biotur- stacked massive sandstones. The unbioturbated
bated (70-90%), but ichnodiversity is low, sug- sandstones are medium to coarse grained and
gesting hostile conditions ideally suited to a few of remarkably even grain size. They occasionally
specialist species (Levington 1970) and probably contain faint clay drapes or thick muddy ripple
due to low salinity and/or oxygenation in cross-laminated heteroliths.
conjunction with low sedimentation rates and This facies is commonly associated with the
palimpsesting of identical ichnocoenoses due to IDMBTC facies and as such is probably
aggradation (see Mcllroy 2004). The trace fossils deposited at the distributary mouth by rapid
recorded are Thalassinoides and Rhizocorallium, deceleration of currents with high suspended
which are normally considered to be marine in concentrations of well-sorted sediment. Given
nature, and the feather-like trace (Fig. 10). Few that the sands are completely unbioturbated
sedimentary structures are present, but the and without sedimentary structures, objective
marked contrast in grain size is suggestive of a interpretation cannot extend beyond extrapola-
normally quiescent environment that is subject tion from common association with other
to periodic events capable of transporting large facies, to which the anomalous facies is presumed
grain sizes. to be genetically related.
The sedimentological setting that best fits the
range of sedimentary and chemical conditions
is that of a protected central basin environment Application of ichnofabric stacking patterns
(cf. Zaitlin & Schultz 1984, 1990). The central
basin of a wave-dominated estuary is influ- In attempting to resolve the interrelationships of
enced by marine waters but protected from facies the known environmental tolerances of
strong currents by the presence of what is trace fossils and community structure (described
effectively a standing body of water con- as ichnofabric) have been combined with
strained on the seaward side by a barrier bar. information from sedimentary structures and
Occasional storm events may account for the common association of facies (ichnofabrics)
thin but very coarse sand stringers present in using Walther's law. Once a model for the distri-
this facies, either through increased transport bution of both sedimentary facies and ichno-
from the fluvial system (through the bayhead fabrics has been constructed (Figs 12, 14), the
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 269

vertical stacking of facies and their ichnofabrics similar scheme to be built from the available
in core can be used to assess changes in relative dataset.
sea-level. The ichnology of tide-dominated deltas in
During the course of this study special focus general remain poorly known; however, compar-
was centred on the ichnology of centimetre- ison of these data with data from wave- and
thick mudstone horizons within tidal channel river-dominated deltas (Gingras et al. 1998)
successions. This enabled production of a suggests that: (1) the Skolithos ichnofacies is
detailed model for the spatial distribution of better developed in river-dominated deltas; and
tidal channel ichnofabrics (Fig. 19). This concep- (2) ichnodiversity is lower than in typical wave-
tual model for the distribution of tidal channel dominated deltas. Tide-dominated deltas would
ichnofabrics can be used to interpret stacking seem therefore to represent an intermediate con-
patterns (Fig. 20). This technique is particularly dition between the wave- and river-dominated
pertinent to the He Formation, owing to the end members that probably results from an
homogeneity of the lower (petroleum-bearing) increased marine influence in delta front to
tidal channel-rich interval (Fig. 3). delta top settings.
Throughout this study, the main depositional
environments have been determined using a com- This study was supported by Kristin License and
bined sedimentological/stratigraphical approach, Statoil. Logistical support and technical discussions
and further refined using ichnology. As discussed with K. Dale Grov, C. Elfenbein (Statoil), C. Brostr0m
above, tidal sediments may be deposited in a (Kristin License, Statoil) & T. Laerdal (University of
diverse suite of environments with unique Bergen) are recognized with thanks. Critical appraisal
of the manuscript by A. Martinius and R. Twitchett
physiochemical conditions that affect the distri- is gratefully acknowledged. In addition, Kristin License
bution of the infauna. The range of palaeo- are thanked for permission to publish these findings.
environments in which a given trace fossil is
found is an invaluable tool for the study of
problematic facies, and greatly aids facies References
characterization.
ALLEN, J. R. 1981. Lower Cretaceous tides revealed by
cross bedding with mud drapes. Nature, 289, 579-
Conclusion 581.
ALLEN, G. P., SALOMON, J. C., BASSOULLET, P., Du
The He Formation is a widespread reservoir PENHOAT, Y. & DE GRANDPRE, C. 1980. Effects
interval in the Haltenbanken area offshore of tides on mixing and suspended sediment trans-
port in macrotidal estuaries. Sedimentary Geology,
Mid-Norway with excellent, though complex, 26, 69-90.
sandstone reservoir intervals that are pre- ALONGI, D. M. 1991. The role of intertidal mudbanks
dominantly in tidal channel and tidal flat facies. in the diagenesis and export of dissolved and
Challenges vary from field to field but are in particulate materials from the Fly Delta, Papua
many cases improved by high-quality facies New Guinea. Journal of Experimental Marine
characterization. Biology and Ecology, 149, 81-107.
This study has aimed to incorporate all avail- ANDREWS, P. B. 1970. Facies and Genesis of a Hurricane
able sedimentological and ichnological evidence Washover fan, St Joseph Island, Central Texas
to produce a refined understanding of facies Coast. Bureau of Economic Geology, University
distributions. The facies models and ichnofabric of Texas, Austin, Texas, Report Investigations, 67.
BANN, K. L. & FIELDING, C. 2004. An integrated
distribution models presented herein allow ichnological and sedimentological comparison of
assessment of facies-stacking patterns that are non-deltaic shoreface and subaqueous delta
the fundamental basis of sequence stratigraphic deposits in Permian reservoir units of Australia.
analysis and prediction of facies architecture in In: MclLROY, D. (ed.) The Application of Ichnology
inter-well areas. This bespoke sedimentological/ to Palaeoenvironmental and Stratigraphic Analysis.
ichnofabric approach has been demonstrated to Geological Society, London, Special Publications,
be highly effective in characterizing the complex 228,273-310.
tidal-deltaic facies of the He Formation on BRETTLE, M. J., MC!LROY, D., DAVIES, S. J., ELLIOTT,
Kristin Field. Preliminary work with the T. & WATERS, C. N. 2002. Identifying cryptic
adjacent Sm0rbukk field has demonstrated tidal influences within deltaic successions: an
example from the Marsdenian (Namurian)
that, with little modification, the same scheme interval of the Pennine Basin, UK. Journal of the
can be applied on a semi-regional basis. How- Geological Society, London, 159, 379-391.
ever, in different basins of different stratigraphic BROMLEY, R. G. & ASGAARD, U. 1991. Ichnofacies: a
ages the palaeoenvironmental tolerance of trace- mixture of taphofacies and biofacies. Lethaia, 24,
making organisms is likely to differ and require a 153-163.
270 D. McILROY

BUATOIS, L. A. & MANGANO, G.M. 2004. Animal- GIBBS, R. J. 1977. Distribution and transport of
substrate interactions in freshwater environments: suspended particulate material of the Amazon
applications of ichnology in facies and sequence River in the Ocean. In: WILEY, M. (ed.) Estuarine
stratigraphic analysis of fluvio-lacustrine succes- Processes II: Circulation, Sediments and Transfer
sions. In: MclLROY, D. (ed.) The Application of of Material in the Estuary. Academic Press,
Ichnology to Palaeoenvironmental and Strati- London, 35-47.
graphic Analysis, Geological Society, London, GlNGRAS, M. K., MACEACHERN, J. A. & PEMBERTON,
Special Publications, 228, 311-333. S. G. 1998. A comparative analysis of the ichnol-
BUATOIS, L. A., MANGANO, G. M., MAPLES, C. G. & ogy of wave- and river-dominated allomembers
LANIER, W. P. 1998. Allostratigraphic and sedi- of the Upper Cretaceous Dun vegan Formation.
mentologic applications of trace fossils to the Bulletin of Canadian Petroleum Geology, 46, 51-73.
study of incised estuarine valleys: an example GOLDRING, R., BOSENCE, D. W. J. & BLAKE, T. 1978.
from the Virgilian Tonganoxie Sandstone Estuarine sedimentation in the Eocene of southern
Member of eastern Kansas. Current Research in England. Sedimentology, 25, 861-876.
Earth Sciences, 241, 1-27. GRASSLE, J. F. & GRASSLE, J. P. 1974. Opportunistic life
CARTER, C. H. 1978. A regressive and barrier-protected histories and genetic systems in marine benthic
deposit: depositional environments and geographic polychaetes. Journal of Marine Research, 32,
setting of the late Tertiary Cohansey Sand. Journal 253-284.
of Sedimentary Petrology, 48, 933-950. GREGER, J. A, NEDRELID, T. et al. 2001. PL134BJ199
DALRYMPLE, R. W., ZAITLIN, B. A. & BOYD, R. 1992. Kristin Field Plan for Development and Operation:
Estuarine facies models: conceptual basis and Supporting Document — Geology & Geophysics.
stratigraphic implications. Journal of Sedimentary Statoil Internal Report [confidential].
Petrology, 62, 1130-1146. HARRIS, P. T., BAKER, E. K., COLE, A. R. & SHORT, S. A.
DIAZ, R. J. & ROSENBURG, R. 1005. Marine benthic 1993. A preliminary study of sedimentation in the
hypoxia: a review of its ecological effects and the tidally dominated Fly River Delta, Gulf of Papua.
behavioural responses of benthic macrofauna. Continental Shelf Research, 13, 441^472.
Oceanography and Marine Biology Annual HOWARD, J. D. & FREY, R. W. 1973. Characteristic
Review, 33, 245-303. physical and biogenic sedimentary structures in
DOTT, R. H. & BOURGEOIS, J. 1982. Hummocky strati- Georgia estuaries. American Association of Petro-
fication: significance of its variable bedding leum Geologists Bulletin, 57, 1169-1184.
sequences. Geological Society of America Bulletin, HUBBARD, D. K. & BARWIS, J. H. 1976.Discussion of
93, 663-680. tidal inlet sand deposits: examples from the
DORIES, J. & HOWARD, J. D. 1975. Fluvial-marine South Carolina coast. In: HAYES, M. O. & KANA,
transition indicators in an estuarine environment, A. (eds) Terriginous Clastic Depositional Environ-
Ogeechee River - Ossabaw Sound. Estuaries of ments: Some Modern Examples. American Asso-
the Georgia Coast, USA: Sedimentology and ciation of Petroleum Geologists Field Course.
biology. Senckenbergiana Maritima, 1, 137-179. University of South Carolina Technical Report
EDELVANG, K. & AUSTEN, I. 1997. The temporal varia- 11-CRD, 128-142.
tion of floes and fecal pellets in a tidal channel. JOHNSON, R. G. 1965. Temperature variation in the
Estuarine, Coastal and Shelf Science, 44, 361-367. infaunal environment of a sand flat. Limnology
EHRENBERG, S. N. 1993. Preservation of anomalously and Oceanography, 10, 114-120.
high porosity in deeply buried sandstones by grain KLEIN, G. DE V. 1971. A sedimentary model for deter-
coating chlorites: examples from the Norwegian mining paleotidal range. Geological Society of
Continental Shelf. The American Association of America, Bulletin, 82, 2585-2592.
Petroleum Geologists Bulletin, 77, 1260-1286. KRANCK, K. 1981. Particulate matter grain-size
EHRENBERG, S. N., DALLAND, A., NADEAU, P. H., characteristics and flocculation in a partially
MEARNS, E. W. & AMUNDSEN, H. E. F. 1998. mixed estuary. Sedimentology, 28, 107-114.
Origin of chlorite enrichment and neodymium LEVINGTON, J. S. 1970. The paleoecological significance
isotopic anomalies in Haltenbanken sandstones. of opportunistic species. Lethaia, 3, 69-78.
Marine and Petroleum Geology, 15, 403-425. MACEACHERN, J. A. 1989. Estuarine channel deposition
FENIES, H. & FAUGERES, J. C. 1998. Facies and geo- within the Lower Cretaceous Waseca Formation,
metry of tidal channel fill deposits (Arcachon Upper Mannville Group, Lloydminster area,
Lagoon, SW France). Marine Geology, 150, 131- Saskatchewan. In: REINSON, G. E. (ed.) Modern
148. and Ancient Examples of Clastic Tidal Deposits: A
FREY, R. W. & COLORING, R. 1992. Marine event beds Core and Peel Workshop. Canadian Society of Pet-
and recolonization surfaces as revealed by trace roleum Geologists, Second International Research
fossil analysis. Geological Magazine, 129, 325-335. Symposium on Clastic Tidal Deposits, 50-59.
FREY, R. W. & HOWARD, J. D. 1986. Perspectives: MANGANO, G. M. & BUATOIS, L. A. 2004. Ichnology
Mesotidal estuarine sequences: a perspective of Carboniferous tide-influenced environments
from the Georgia Bight. Journal of Sedimentary and tidal flat variability in the North American
Petrology, 56, 911-924. Midcontinent. In: MC!LROY, D. (ed.) The Applica-
FREY, R. W., PEMBERTON, S. G. & SAUNDERS, T. D. A. tion of Ichnology to Palaeoenvironmental and
1990. Ichnofacies and bathymetry: a passive Stratigraphic Analysis. Geological Society,
relationship. Journal of Paleontology, 64, 155-158. London, Special Publications, 228, 157-178.
A JURASSIC TIDE-DOMINATED DELTA, NORWAY 271

MARTIN, M. A. & POLLARD, J. E. 1996. The role of Ichnology to Petroleum Exploration: A Core Work-
trace fossil (ichnofabric) analysis in the develop- shop. Society for Sedimentary Geology, Tulsa,
ment of depositional models for the Upper Oklahoma, SEPM Core Workshops, 17, 141-167.
Jurassic Fulmar Formation of the Kittiwake PEMBERTON, S. G., REINSON, G. E. & MACEACHERN,
Field (Quadrant 21 UKCS). In: HURST, A. (ed.) J. A. 1992. Comparative ichnological analysis of
Geology of the Number Group: Central Graben Late Albian estuarine valley fill and shelf-shore-
and Moray Firth, UKCS. Geological Society, face deposits, Crystal Viking Field, Alberta. In:
London, Special Publications, 114, 163-183. PEMBERTON, S.G. (ed.) Applications of Ichnology
MARTINIUS, A. W., KAAS, I., N^ss, A., HELGESEN, G., to Petroleum Exploration: A Core Workshop.
KJAEREFJORD, J. M. & LEITH, D. A. 2001. Sedi- Society for Sedimentary Geology, Tulsa, Okla-
mentology of the heterolithic and tide-dominated homa, SEPM Core Workshops, 17, 291-317.
Tilje Formation (Early Jurassic, Halten Terrace, PEMBERTON, S. G., SPILA, M., PULHAM, A. J., SAUN-
offshore mid-Norway). In: MARTINSEN, O. J. & DERS, T., MACEACHERN, J. A., ROBBINS, D. &
DREYER, T. (eds) Sedimentary Environments SINCLAIR, I. 2001. Ichnology and Sedimentology
Offshore Norway: Palaeozoic to Recent. NPF of Shallow to Marginal Marine Systems. Ben
Special Publications, 10, Elsevier Science, Amster- Nevis and Avalon Reservoir, Jeanne d'Arc Basin.
dam, 103-144. Geological Association of Canada, Short Course
MATTISON, B. W., Fox, A. J. & PEMBERTON, S. G. 1989. Notes, St. John's, Newfoundland, 15.
Sedimentologic paleontologic and ichnologic POLLARD, J. E., COLORING, R. & BUCK, S.G. 1993.
criteria for the recognition of ancient estuarine Ichnofabrics containing Ophiomorpha: signifi-
deposits: an example from the Lower Cretaceous cance in shallow-water facies interpretation.
McMurray Formation in the Athabasca Oil Journal of the Geological Society, 150, 149-164.
Sands area of northeastern Alberta. In: REINSON, READING, H. G. 1996. Introduction. In: READING, H. G.
G. E. (ed.) Modern and Ancient Examples of (ed.) Sedimentary Environments: Processes, Facies
Clastic Tidal Deposits: A Core and Peel Workshop. and Stratigraphy. Blackwell Science, Oxford, 1-4.
Canadian Society of Petroleum Geologists, REINECK, H. E. 1967. Layered sediments of tidal flat
Second International Research Symposium on beaches and shelf bottoms of the North Sea. In:
Clastic Tidal Deposits, 66-67. LAUFF, G. H. (ed.) Estuaries. American Associa-
MclLROY, D. 2004 A review of ichnological concepts, tion for the Advancement of Science, Washington
methodologies, applications and frontiers. In: DC, Special Publications, 83, 191-206.
MclLROY, D. (ed.) The Application of Ichnology SCHUBEL, J. R. 1968. Turbidity maximum of the north-
to Palaeoenvironmental and Stratigraphic Analysis. ern Chesapeake Bay. Science, 161, 1013-1015.
Geological Society, London, Special Publications, STANISTREET, I. G., LE BLANC SMITH, G. & CADLE, A. B.
228, 3-27. 1980. Trace fossils as sedimentological and palaeo-
MclLROY, D., FLINT, S. S. & HOWELL, J. A. 1999. environmental indices in the Ecca Group (Lower
Applications of high resolution sequence strati- Permian) of the Transvaal. Transactions of the
graphy to reservoir prediction and flow unit Geological Society of South Africa, 83, 333-344.
definition in aggradational tidal systems In: STILLING, J. 2000. lie Formation, Western Haltenbanken
HENTZ, T. (ed.) Advanced Reservoir Characteriza- area, Norwegian Sea. Statoil Internal Report
tion for the Twenty-first Century. GCSSEPM, [confidential].
Houston, Texas, 19, 121-132. TAYLOR, A. M. & GOLDRING, R. 1993. Description and
MILNE, A. 1940. The ecology of the Tamar Estuary, IV: analysis of bioturbation and ichnofabric. Journal
the distribution of fauna and flora on buoys. of the Geological Society, London, 150, 141-148.
Journal of the Marine Biologists Association, UK, TAYLOR, A. M. & GAWTHORPE, R. L. 1993. Application
24, 69-87. of sequence stratigraphy and trace fossil analysis
MONAHAN, P. A., LUTERNAUER, J. L. & BARRIE, J. V. to reservoir description: examples from the
1993. A delta topset sheet sand and modern Jurassic of the North Sea. Petroleum Geology of
sedimentary processes in the Fraser River delta, Northwest Europe: Proceedings of the 4th Con-
British Columbia. Current Research, Part A; Geo- ference, 317-335.
logical Survey of Canada Paper, 93-1 A, 263-272. TAYLOR, A. M., COLORING, G. & GOWLAND, S. 2003.
M0RK, G., EVENSEN, A.I. et al. 1997. Discovery Analysis and application of ichnofabrics. Earth
Report Kristin. Saga Petroleum Internal Report Science Reviews, 60, 227-259.
[confidential]. WALTHER, J. 1894. Einleitung in die Geologie als
PATTISON, S. A. J. 1992. Recognition and interpretation Historische Wissenschaft, Bd. 3. Lithogenesis der
of estuarine mudstones (central basin mudstones) Gegenwart. G. Fischer Verlag, Jena, 535-1055.
in the tripartite valley-fill deposits of the Viking WEIMER, R. J., HOWARD, J. D. & LINDSAY, D. R. 1982.
Formation, Central Alberta. In: PEMBERTON, Tidal flats. In: SCHOLLE, P. A. & SPEARING, D.
S. G. (ed.) Applications of Ichnology to Petroleum (eds) Sandstone Depositional Environments.
Exploration: A Core Workshop. Society for American Association of Petroleum Geologists,
Sedimentary Geology, Tulsa, Oklahoma, SEPM Memoirs, Tulsa, Oklahoma, 31, 191-245.
Core Workshops, 17, 223-249. WOLANSKI, E. & GIBBS, R. J. 1995. Flocculation of
PEMBERTON, S. G. & WRIGHTMAN, D. M. 1992. Ichno- suspended sediment in the Fly River estuary,
logical characteristics of brackish water deposits. Papua New Guinea. Journal of Coastal Research,
In: PEMBERTON, S. G. (ed.) Applications of 11, 794-762.
272 D. McILROY

WRIGHT, L. D. 1977. Sediment transport and deposi- ZAITLIN, B. A. & SCHULTZ, B. C. 1984. An estuarine-
tion at river mouths: a synthesis. Geological embayment fill model from the Lower Cretaceous
Society of America Bulletin, 88, 857-868. Mannville Group, west-central Saskatchewan. In:
WRIGHTMAN, D. M., PEMBERTON, S. G. & SINGH, C. STOTT, D. F. & GLASS, D. J. (eds) Mesozoic of
1987. Depositional modelling of the Upper Middle North America. Canadian Society of
Mannville (Lower Cretaceous), central Alberta. Petroleum Geologists, Memoirs, 9, 455—469.
Implications for the recognition of brackish water ZAITLIN, B. A. & SCHULTZ, B. C. 1990. Wave-
deposits. In: TILLMAN, R. W. & WEBER, K. J. influenced estuarine sand body, Senlac heavy oil
(eds) Reservoir Sedimentology. Society of Economic pool, Saskatchewan, Canada. In: BARWIS, J. H.
Paleontologists and Mineralogists, Special Publica- (ed.) Sandstone Petroleum Reservoirs. Springer-
tions, Tulsa, Oklahoma, 40, 189-220. Verlag, New York, 363-387.
An integrated ichnological and sedimentological comparison of
non-deltaic shoreface and subaqueous delta deposits in
Permian reservoir units of Australia
KERRIE L. BANN1 & CHRISTOPHER R. FIELDING2
1
Department of Earth and Atmospheric Sciences, University of Alberta,
Edmonton, Alberta, Canada T6G 2E3 (e-mail: kbann@ualberta.ca)
Department of Geosciences, 214 Bessey Hall, University of Nebraska-Lincoln,
NE 68588-0340, USA (e-mail: cfielding2@unl.edu)

Abstract: Permian sequences in the Denison Trough of Queensland, eastern Australia, are
productive conventional gas reservoirs. Previous attempts to interpret the reservoir bodies
in terms of depositional environments have relied largely on sedimentary facies analysis
and palynology. Sequences have been re-evaluated from a detailed, integrated ichnological
and sedimentological perspective, resulting in a significant increase in the precision and
resolution of the palaeoenvironmental interpretation. Various marine, coastal and upper
delta plain deposits are recognized. Ichnological signatures have facilitated the differentiation
of subaqueous delta deposits from those deposited in non-deltaic offshore and shoreface
environments. Delta front facies are further subdivided by integrating ichnological and sedi-
mentological data. Permian offshore and shoreface successions in the Denison Trough contain
ichnological signatures that exhibit high diversities (25 ichnogenera comprising 32 ichno-
species), moderate to intense levels of bioturbation, uniformity of burrowing, and a wide vari-
ety of structures representing specialized feeding strategies. Examples from an additional
reservoir dataset, the Tern Formation from the offshore Bonaparte Basin in north Western
Australia, also clearly demonstrate the ichnological complexity of Permian shoreface succes-
sions. In contrast, Permian deltaic deposits contain ichnological signatures that reflect stressed
environmental conditions. Assemblage diversity is reduced (16 ichnospecies), bioturbation
intensity is significantly reduced, and uniformity of burrowing is sporadic.

A delta is generally understood to be a discrete An accurate genetic model of deltaic (and


shoreline protuberance formed where a river shoreface) successions is required to refine
enters an ocean or other large body of water exploration techniques and optimize production
and supplies sediment more rapidly than it can and field development for these types of reser-
be redistributed by basinal processes (Elliott voir. Coastal and deltaic deposits, in general,
1986). In tide-dominated settings the shoreline form similar upward-coarsening and shallowing
may be strongly embayed rather than protuber- successions, so that the specific type of deposi-
ant, but sediment is nonetheless supplied more tional environment and relative importance of
rapidly than can be redistributed, and the fluvial, wave and tidal processes cannot be
system still progrades, irrespective of shoreline analysed on the basis of well logs alone (Allen
morphology (Dalrymple 1999). Ancient deposits & Chambers 1998). In many cases, diagnosis of
are difficult or even impossible to characterize deltaic deposits has relied exclusively on physical
as 'deltaic' on the basis of a single core or outcrop sedimentary structures such as current-generated
section; a reconstructed plan view of the system structures, unimodal or dispersive palaeocurrent
or good three-dimensional control of facies distributions, and the broad cross-sectional and
patterns is necessary (Bhattacharya & Walker plan geometry of a unit.
1992). Numerous researchers have recognized that
Ancient deltas are economically important trace fossils record animal responses to subtle
owing to their association with coal, oil and gas changes in environmental parameters such as
reserves. Deltaic reservoirs generally consist of sedimentological, hydraulic, chemical and
an amalgamation of several different sandstone bathymetric regimes (Seilacher 1964; Frey &
facies, e.g. distributary channels and mouth Pemberton 1985; Pemberton et al 1992). The
bars, and therefore form multilateral and multi- interpretation of sedimentary rocks and palaeo-
storey systems comprising a wide variety of environments is therefore enhanced considerably
facies types and reservoir characteristics (Cole- by incorporating ichnology (Seilacher 1955).
man & Wright 1975; Allen & Chambers 1998). This is particularly true in the evaluation of
From: MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 273-310. 0305-8719/04/S 15.00 © The Geological Society
of London.
274 K. L. BANN & C. R. FIELDING

shallow marine successions, where distinction along the western margin of the Bowen Basin
between deltaic and non-deltaic deposits is (Fig. 1). These units are extensive along deposi-
challenging and potentially contentious. The tional strike, in a north-south direction, but
ultimate objective of this study is to portray the fine, thin and pinch out eastward into the
facies implications of various trace fossil assem- basin. Palaeocurrent data from these units indi-
blages and then to establish the differences cate persistently eastward sediment dispersal
between deltaic and non-deltaic shoreface succes- into the basin, from source areas located in
sions. Marine ichnofacies represent archetypal earlier Palaeozoic fold belt strata to the west
facies models based upon recurrent ichnological (Fig. 1). The coarse-grained formations are sepa-
assemblages (Seilacher 1967, 1978; Frey & rated from each other by intervals of fine-grained
Pemberton 1984, 1985). An integrated approach shale, rich in marine invertebrate fossils, inter-
combining several potential environmental preted to record predominantly offshore marine
indicators, ichnology, sedimentology and palae- and shelf conditions. The succession can there-
ontology has significantly enhanced the palaeo- fore be regarded as a series of progradational
environmental interpretations in this study of cycles separated by marine shale intervals
Permian successions from Australia. (Fig. 2a). Many of the sandstone-dominated
The Permian succession of the Denison units of the Denison Trough succession are
Trough of central Queensland, Australia, con- prospective for hydrocarbons, and several pro-
tains a number of shallow marine and deltaic ducing gas fields in the area deliver from reser-
units. The depositional context of several strati- voirs within these formations.
graphic units is uncertain and insufficient infor- The oldest stratigraphic unit recognized in the
mation exists on the plan and cross-sectional Denison Trough - the Reids Dome Beds - com-
geometry of these units to firmly establish a prises a locally thick (up to > 1 km) succession of
deltaic origin. In this paper we interpret the siliciclastics and some thick coal seams, and is
ichnology of formations that have been indepen- interpreted as the mainly fluvial and lacustrine
dently established as deltaic and of shallow deposits of technically active rift basins. The
marine in origin. An additional dataset from a contact with the overlying Cattle Creek Forma-
basin 3000km to the west (Tern Formation, tion is known to be strongly diachronous. In
Bonaparte Basin, Western Australia) is also the northern Denison Trough, where the Cattle
presented as it further demonstrates the diverse Creek Formation is thickest, five alternating
ichnological characteristics of Permian shoreface mudrock and sandstone-dominated members
successions. We then apply the ichnological and have been recognized, and are interpreted as off-
sedimentological criteria to the diagnosis of a shore marine and broadly coastal respectively
formation, the context of which is complex and (Fielding & Lang 1988, Fig. 2a). Overlying the
cryptic, in order to demonstrate the value of uppermost mudrock unit of the Cattle Creek
ichnology in this process. Formation, the Sirius Mudstone Member, is
the Lower Aldebaran Sandstone. This unit is
largely confined to extensional sub-basins, but
Geological setting isopachs show the gradually diminishing influ-
ence of the extensional topography. The Lower
The Denison Trough forms the southwestern part Aldebaran Sandstone was interpreted by Baker
of the Bowen Basin of east-central Queensland, (1991) as a deltaic deposit. The Upper Aldebaran
Australia (Fig. 1). The Trough exists as a concen- Sandstone unconformably overlies the Lower
tration of extensional grabens and half-grabens Aldebaran Sandstone, the angularity of the rela-
formed during a period of limited though wide- tionship being apparent only from seismic reflec-
spread crustal extension across eastern Australia tion data. Unlike the underlying unit, the Upper
in the Late Carboniferous to Early Permian Aldebaran Sandstone is not confined to the early
(Holcombe et al 1997; Fielding et al 2000, extensional lows, but rather overlaps basement
2001). Subsequent extension-related and thermal highs and extends considerably further eastward
sag-related subsidence formed the broader into the basin than older units. The Upper Alde-
feature known as the Bowen Basin, which was baran Sandstone shows a prominent arcuate
then further modified by foreland loading asso- bulge into the basin in plan geometry, and was
ciated with the long-lived Hunter-Bowen con- interpreted by Baker (1991) and subsequent
tractional event (Fielding et al 2000, 2001). The workers as the deposit of a large, eastward-
Denison Trough therefore forms part of the prograding, mixed-influence delta.
cratonic (western) margin of the Bowen Basin. The top of the Upper Aldebaran Sandstone
During late Early to Late Permian times, a is marked by a marine flooding surface, and
series of coarse clastic formations were deposited overlain by the Freitag Formation. The lower,
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 275

Fig. 1. Location maps of the Denison Trough.

mainly fine-grained part of this unit was depos- could represent the southern margin of a large
ited in an offshore and shelf environment, delta complex. The top of the Freitag Formation
whereas the upper half consists of a number of is represented by a marine flooding surface of
progradational parasequences that (in the inter-regional extent, traceable throughout the
north) include coastal facies. The unit thickens length of the Bowen-Gunnedah-Sydney Basin
significantly to the north, into an area of limited System (Fielding et al. 2001). In the Denison
subsurface data, and the possibility exists that Trough, this surface marks the base of the
the Freitag Formation in the Denison Trough Ingelara Formation, a laterally extensive
276 K. L. BANN & C. R. FIELDING

Fig. 2. (a) Stratigraphic column of the Denison Trough succession, (b) Legend of symbols to Figures 11-14.

marine mudrock interval. The Ingelara Forma- by John & Fielding (1993) to be of mostly non-
tion passes upward into the sandstone- deltaic shallow marine to coastal plain origin,
dominated Catherine Sandstone and its lateral with a small number of modest deltaic depo-
equivalent the German Creek Formation to the centres; however, integrated sedimentological
north. The Catherine Sandstone was interpreted and ichnological analysis suggests that a
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 277

number of the shallow marine intervals of the previously underutilized ichnological database,
Catherine Sandstone are of deltaic origin. The has led to a substantial revision of the facies
German Creek Formation to the north, which scheme originally outlined and illustrated by
hosts economically important coal deposits, has Fielding et al. (1996). A summary of the new
been interpreted as the deposits of a large, east- facies scheme is given in Table 1. Eight facies
ward-dispersing, mixed-influence delta complex associations (A to H) are recognized, each
(Falkner & Fielding 1993), with some parts of containing a number of component facies, and
the succession recording deposition in shallow representing depositional environments ranging
shoreface and foreshore environments (Bann & from shelf to coastal plain. Association H
Fielding 2001). facies (upper delta plain interdistributary lakes
The overlying Peawaddy Formation is another and mires) are outside the area of focus of this
broadly coarsening-upward unit that is inter- paper.
preted as a further prograding delta system. A complete spectrum of shallow marine
However, in contrast to underlying units, the deposits is preserved, ranging from shelfal
volcanic lithic Peawaddy Formation shows through offshore and offshore transition to
evidence of axial dispersal southward into the shoreface and foreshore environments. Offshore
basin, and thus the section in the Denison terminology used in this paper follows the work
Trough represents only a distal margin of this of Howard (1971, 1972), Howard & Reineck
system. Furthermore, the plan shape of the (1981), Howard & Frey (1984), Vossler &
Peawaddy Formation sandstone facies indicates Pemberton (1989) and Frey (1990), and subse-
an elongate (birdsfoot) morphology, and quently modified by MacEachern & Pemberton
together with a general lack of tidal indicators (1992) and Pemberton & MacEachern (1995,
suggests a fluvially dominated delta style. The 1997). A brief summary is given in Bann et al.
succeeding Bandanna Formation is of a similar (2004).
sedimentary character, and ranges up to the Most of the shallow marine facies in the
latest Permian coal measures, in the Denison Denison Trough have all the hallmarks of
Trough. wave-dominated successions (Johnson & Bald-
The formations considered herein represent win 1996). Additionally, a separate suite of
the late extensional to passive thermal subsidence facies has been defined to accommodate shallow
phases of the basin's development. All coarse- marine facies with somewhat different character,
grained formations are derived from the craton believed to record deltaic environments (Associa-
to the west of the Bowen Basin, and all were tion E: Table 1). Association E includes facies
accumulated under similar environmental condi- interpreted to record prodelta, through distal
tions. The climate is interpreted to have been delta front, proximal delta front into mouth
cool temperate and then humid in the aftermath bar and river mouth settings (Table 1). The inter-
of the late Palaeozoic Gondwanan glaciation pretation of these facies follows an integrated
(Crowell & Frakes 1971, 1975; Veevers & approach based on their context within vertical
Powell 1987; McLoughlin 1993), with some sequences, their lithology, cross-sectional geome-
floating ice still rafting outsized debris into the try and contained physical and biogenic sedimen-
marine basin. Previous work (Fielding & Lang tary structures.
1988; Baker 1991; Falkner & Fielding 1993) has Deltas occur in a wide variety of shapes and
interpreted deltaic depositional environments forms, depending on the types and energy of
for some of the coarse-grained units, largely on the coastal processes and the volume and grain
the basis of gross plan shape of formations, and size of the fluvial sediment influx (Coleman
the interpretation of thick sandstones as distri- 1981; Reading 1986; Bhattacharya & Walker
butary channel and mouth bar deposits. These 1992). Protuberant deltas generally comprise
criteria are in some cases difficult to apply, or three major depositional environments: the
may be misleading. Ichnofacies analysis provides delta plain, the delta front, and the prodelta.
a powerful additional criterion with which to Each of these environments forms a depositional
identify deltaic and deltaically influenced system that consists of a distinct suite of litholo-
marine environments. gies, facies and reservoir potential (Coleman &
Wright 1975; Allen & Chambers 1998). The
delta plain comprises the subaerial part of the
Facies analysis delta and will not be discussed in any detail
herein. The subaqueous portion of the delta
A re-evaluation of depositional environments of exists in the shallow marine zone (or lacustrine
Permian formations in the Denison Trough and zone not discussed here), below the low tide
adjacent areas, incorporating the extensive and mark (subtidal), and fringes the lower part of
Table 1.

Facies Lithology Primary Structures Ichnofacies Interpretation

Al: Laminated moderately to Claystones and siltstones, laminated, Lamination, local glendonites, Zoophycos ichnofacies - distal Shelf
thoroughly bioturbated moderately to intensely bioturbated, local outsized clasts Cruziana ichnofacies
claystone and siltstone dispersed coarser-grained sediment, locally
fossiliferous
A2: Moderately to thoroughly Siltstone or claystone with minor Local lamination, local linsen and Distal Cruziana ichnofacies Lower offshore
bioturbated muddy siltstone interlaminated and admixed sandstone, lenticular bedding, rare micro HCS
moderately to thoroughly bioturbated, and small-scale ripple cross-
locally fossiliferous lamination, local glendonites
Bl: Bioturbated sandy Siltstone with interlaminated and admixed Linsen, lenticular and wavy Diverse archetypal Cruziana Upper offshore
siltstone sandstone, intensely to thoroughly bedding, current ripple cross- ichnofacies with mixed
bioturbated, minor discrete sandstone beds lamination, micro hummocky-cross SkolithosI Cruziana ichnofacies
<3 cm thick, locally fossiliferous stratification (HCS) in thin discrete sandstone beds
B2: Interlaminated, Sandy siltstone and interlaminated, As for fades Bl, plus small-scale Diverse archetypal Cruziana Offshore transition
bioturbated sandy siltstone commonly discrete and sharp-bounded HCS ichnofacies with Skolithos
and laminated sandstone sandstone beds < 10 cm in thickness, locally ichnofacies in the sandstone
fossiliferous beds
CIA: Thoroughly bioturbated Intensely bioturbated muddy sandstone Low-angle, undulatory parallel Diverse proximal Cruziana Distal lower
muddy sandstone units with rare, discrete, sharp-bounded lamination (HCS), oscillation ichnofacies with Skolithos shoreface
sandstone layers < 15 cm in thickness, minor ripple lamination ichnofacies in the clean
dispersed gravel, locally fossiliferous sandstone beds
C1B: Interbedded bioturbated Interbedded moderately to thoroughly Thicker sandstones show local Distal Skolithos ichnofacies Proximal lower
muddy sandstone and bioturbated muddy to clean sandstone and basal and top surface gravel lags, shoreface
laminated sandstone sharp-bounded, tabular, sparsely to flat and low-angle lamination,
moderately bioturbated clean sandstone abundant HCS, symmetrical
beds < 1 m thick, dispersed gravel and thin ripples, interference ripples, large
conglomerate beds gravelly symmetrical ripples
C2: Interbedded massive and Thickly interbedded sandstone in sharp- Similar structures to facies C1B, Archetypal Skolithos Middle shoreface
laminated sandstone bounded, sparsely to moderately HCS is large-scale ichnofacies with uncommon
bioturbated tabular beds and composite elements from the proximal
units < 1.5 m thick, minor thin conglomerate Cruziana ichnofacies
beds and dispersed gravel
C3: Amalgamated laminated Clean, well-sorted sandstone in Basal and top surface gravel lags, Skolithos ichnofacies, typically Upper shoreface
and cross-bedded sandstone amalgamated tabular beds and composite multi-directional trough cross- impoverished. Pervasive
units, un- to moderately bioturbated, minor bedding, low-angle bidirectional Macaronichnus segregatis
thin conglomerate beds and dispersed gravel cross-bedding, HCS (typically assemblage common
swale dominated), internal erosion
surfaces
Dl: Flat-laminated sandstone Well-sorted, sparsely to unbioturbated Flat and low-angle or vague Impoverished Skolithos Foreshore
quartzose sandstone, tabular beds and lamination typical, some low-angle ichnofacies
composite units, minor gravel seaward-dipping tabular cross-
bedding
D2: Cross-bedded, coarse- Coarse- to very coarse-grained sandstone Cross-bedding in sets <0.3 m thick, Skolithos ichnofacies Tidal inlet
grained sandstone and conglomerate in erosionally-based often bipolar (heringbone),
(channelized) beds <3m thick, some fining sigmoidal foresets, local mud
upward, variable levels of bioturbation, drapes (some bundled), minor flat
local shell debris lamination, internal erosion
surfaces
El: Unbioturbated gravelly Sandstone, sharply-based but typically Abundant cross-bedding, internal No bioturbation observed River mouth
sandstone coarsening upward, some dispersed gravel, erosion surfaces, less common flat- (expansion of
intraformational clasts, plant debris, coaly lamination and ripple cross- outflows)
traces lamination, very rare HCS
E2: Amalgamated tabular Medium-grained, well-sorted, largely Dominated by HCS (typically Sporadically distributed, Mouth bar
sandstone unbioturbated amalgamated sandstone swale-dominated), flat and low- stressed proximal Cruziana
units with tabular internal and external angle lamination, common internal ichnofacies
geometry and some coarsening upward, erosion surfaces
common plant debris and coaly traces
E3: Interbedded tabular Thickly interbedded tabular intervals of Thicker sandstone beds show basal Sporadically distributed, Proximal delta front
sandstone and moderately and top surface gravel lags, flat and
sporadically bioturbated sandstone in sharp- stressed expression of the
bioturbated sandstone bounded beds and composite units <1.5m low-angle lamination, abundant proximal Cruziana ichnofacies
thick, common plant debris and HCS, local onshore-directed cross-
carbonaceous detritus, minor dispersed bedding, internal erosion surfaces,
gravel and conglomerate beds, rythmic some bed tops covered by
bedding style symmetrical ripples, interference
ripples or gravelly symmetrical
ripples
E4: Interbedded sandstone, Thinly interbedded and interlaminated Dominated by small-scale HCS, Sporadically distributed, Distal delta front
bioturbated silty sandstone sandstone, siltstone and claystone with lam-scram common, also planar stressed Cruziana ichnofacies,
and claystone common discrete sharp-bounded sandstone parallel to low-angle cross-bedding, very rare Skolithos ichnofacies
beds, common plant material and soft-sediment deformation
carbonaceous detritus, dispersed gravel, structures, pin-stripe (linsen) to
locally fossiliferous lenticular, wavy and flaser bedding,
ripple cross-lamination,
carbonaceous detritus and wood
material
Table 1. Continued

Facies Lithology Primary Structures Ichnofacies Interpretation

E5: Interlaminated siltstone, Siltstone with interlaminated and admixed Linsen, lenticular and wavy Sporadically distributed, Prodelta
claystone and silty sandstone sandstone, sporadically sparsely to intensely bedding, current ripple cross- stressed expression of the distal
bioturbated, locally fossiliferous, abundant lamination, micro HCS, common Cruziana ichnofacies
carbonaceous material soft-sediment deformation and
synaeresis cracks
Fl: Biopolar cross-bedded Erosionally-based, thick (typically 8-30 m), Trough cross-bedding dominates, Locally common Skolithos Lower delta plain
sandstone channelized, fining upwards sandstone sets <2 m thick, large-scale/low- ichnofacies distributary channel
bodies, some muddy sandstone beds, minor angle (epsilon) cross-bedding
siltstone partings (some show bundling), (herringbone), flat-lamination,
some gravel, locally moderately bioturbated ripple cross-lamination, local soft-
sediment deformation structures
F2: Trough cross-bedded As for facies Fl but no bioturbation, no As for facies Fl but no bipolar None Upper delta plain
sandstone bundling of fine-grained partings cross-bedding distributary channel
Gl: Sparsely to moderately Interlaminated and thinnly interbedded Linsen, lenticular and wavy Mixed Skolithos I Cruziana Sandy bays, lagoons
bioturbated, interbedded sandstone and siltstone (beds <10cm), bedding, micro HCS, current and ichnofacies, size and diversity and flats
sandstone and siltstone dispersed gravel, bioturbation is variable, wave-generated ripples, restricted
common plant debris and coaly traces, sand symmetrical, ladder and
content 10-80% interference ripples, local soft-
sediment deformation and
synaeresis cracks
G2: Sparsely bioturbated Siltstone and or sandy siltstone, <10% As for facies Gl Mixed Skolithos I Cruziana Muddy bays,
mudstone interlaminated sandstone in beds <10cm ichnofacies, size and diversity lagoons and flats
thick, bioturbation variable but generally restricted
sparse, common plant debris and coaly
traces, rare dispersed gravel
G3: Laminated mudstone Laminated siltstone and claystone, very rare Parallel lamination, rare soft- Very rare, size and diversity Isolated lagoon
bioturbation, common plant debris and sediment deformation structures restricted Cruziana ichnofacies
coaly traces
G4: Carbonaceous shale/coal Carbonaceous siltstone, very rare sandy Parallel lamination No ichnofacies, sparse rootlets Coastal plain mire
detritus, coal
G5: Sparsely bioturbated Sandstone, sharp-bounded bed <50cm Flat and low-angle lamination, Skolithos ichnofacies Storm washover
sandstone thick, local gravel, variable bioturbation, minor cross-bedding
tabular or channelized cross-section
G6: Sparsely to moderately Sandstone, composite units with minor Cross-bedding, less common flat- Skolithos ichnofacies Flood tidal delta
bioturbated sandstone siltstone partings, some show overall lamination and ripple cross-
coarsening or thickening upward, lamination, rare HCS, common
bioturbation variable symmetrical ripples
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 281

the delta plain. The proximal portion of the invertebrate fossils and glendonites (pseudo-
subaqueous delta (termed the delta front) can morphs after the mineral ikaite CaCO3.6H2O:
in turn be subdivided into proximal and distal Kaplan 1979) are common. Ankeritic and side-
components. The more seaward or distal ritic carbonate-cements occur locally.
portion, the prodelta, is where suspended fluvial
silt and clay settles out. Ichnology
Sand is transported to the subaqueous delta Bioturbation, where discernible, is uniformly dis-
through distributaries and accumulates at the tributed and intense (BI4—6). Characteristic ich-
distributary mouth, as a direct result of the nogenera include pervasive, small Phycosiphon,
decrease in current velocity, to form mouth bar abundant Helminthopsis, common Chondrites,
deposits. Mouth bar deposits form good reser- Helminthoida, and Zoophycos.
voirs, although their size, geometry and internal
facies patterns vary according to the degree of Facies interpretation
fluvial/wave/tide influence and sediment input The very fine-grained nature and intensity of
(Coleman & Wright 1975; Allen & Chambers biogenic reworking of this facies indicates slow,
1998; Brettle et al 2002). Mouth bar deposits continuous deposition under quiescent, fully
have been previously identified in the Denison marine conditions. The trace fossil suite repre-
Trough from mapping of both vertical succes- sents grazing and foraging behaviours, with the
sions and lateral facies relationships (Falkner & traces of deposit feeders less common. This
Fielding 1993; Fielding et al 1996). Opencut assemblage is indicative of the Zoophycos ichno-
coalmine highwalls containing mouth bar depos- facies or, in some cases, a distal expression of
its at Gregory and Oaky Creek mines were the Cruziana ichnofacies, and is characteristic
shown by Falkner & Fielding (1993) to pass of quiescent marine shelf environments, well
laterally into distributary channel facies. below maximum storm wavebase. Although
Delta front and prodelta deposits in the grazers and foragers may have been abundant,
Denison Trough show a greater abundance of upon compaction and dewatering their traces
current-generated sedimentary structures relative are not easily preserved (MacEachern & Pember-
to their non-deltaic offshore and shoreface ton 1992). Inhabitants of soupy sediments and
counterparts. These, however, are not sufficiently watery softgrounds tend to cause diffusive turbu-
distinctive on a purely physical sedimentological lence during passage through the substrate and
basis to confidently differentiate them from non- produce a structureless fabric (Bromley 1996).
deltaic offshore and shoreface deposits. Laterally extensive horizons of outsized clasts
This paper focuses on ichnofacies analysis of within shelf deposits are convincing evidence
the offshore/shoreface and subaqueous delta for ice rafting. Glendonites also provide evidence
deposits, in order to facilitate a more robust dis- for very cold climatic conditions as ikaite is
tinction between these two palaeoenvironments. unstable at temperatures above 5°C, at which
Measurement of bioturbation intensity follows point it decomposes to calcium carbonate
the scheme outlined in Bann et al. (2004), (Shearman & Smith 1985; Jansen et al. 1987).
adapted after Reineck (1963) and Taylor & Ikaite has been recorded in several modern,
Goldring (1993). cold climate localities including Antarctica
(Suess et al. 1982), Zaire deep-sea fan (Jansen
et al. 1987) and Alaska (Kennedy et al. 1987).
Non-deltaic offshore and shoreface deposits Glendonites are relatively common in the Per-
mian of Eastern Australia and have been
A summary of the characteristics of the 23 recorded from several localities (Carr et al.
component facies is given in Table 1. 1989; Bann 1998; Bann et al. 2004).

Fades Al: Laminated, moderately to Facies A2: Moderately to thoroughly


thoroughly bioturbated claystone and bioturbated silty muds tone
siltstone
Sedimentology
Sedimentology The siltstone and silty mudstone of this facies
This facies comprises flat/parallel-laminated, contain minor proportions of interlaminated or
light to dark grey, fine siltstone or claystone (more commonly) admixed very fine- to fine-
with rare scattered sand and/or gravel clasts grained sandstone with few preserved sedi-
(Fig. 3a, also see Fig. lib). Laterally extensive mentary structures. Flat/parallel lamination
horizons containing outsized clasts, marine imparted by grain-size variations, pinstripe
282 K. L. BANK & C. R. FIELDING

Fig. 3. (a) Fades Al. GSQ Springsure-18, 220m. (b) Fades A2. Black arrows indicate Phycosiphon; white
arrows indicate scattered outsized clasts. Five cent piece is 10mm in diameter, Catherine Sandstone, Arcturus
Core -6:2. (c-g) Facies B. (c) Pervasive Phycosiphon (Ph), Facies B2, Rolleston Core -12:1, Freitag Formation.
(d) A section through the basal portion of a Rosselia mud-ball (R), Phycosiphon (Ph), Facies B2, Rolleston
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 283

(linsen) and lenticular bedding occur locally. indicates deposition in a lower offshore, open
Rare (<lcm thick), sharp-based, very fine- to marine environment, affected by very cold climatic
fine-grained sandstone layers contain low-angle conditions, but only rarely by severe storms.
undulatory parallel lamination, possible small-
scale hummocky cross-stratification (HCS),
small symmetrical wave ripples, combined flow Facies Bl: Bioturbated sandy silts tone
and current ripple cross-lamination. Scattered
outsized clasts (Fig. 3b), glendonites, marine fos- Sedimentology
sils, carbonaceous detritus, pyrite staining and This facies is characterized by heavily biotur-
carbonate cements (typically ankerite/siderite) bated siltstone with minor to moderate amounts
are locally common. of interlaminated and interstitial very fine- to
fine-grained sandstone (Fig. 3e). Minor discrete,
Ichnology sharp-bounded sandstone beds <5cm thick
Bioturbation is generally pervasive, varying from occur locally and display remnant low-angle
BI4 to BI6, although it is locally sporadic, undulatory parallel lamination, laminated to
decreasing in intensity in association with the scrambled bedding ('lam-scram', Fig. 3e) and
presence of the rare, thin sandstone layers. Char- rarer oscillation ripple lamination. Dispersed
acteristic ichnogenera are uniformly distributed outsized clasts, fossiliferous layers, carbonate-
throughout the facies, and include abundant cements (typically ankerite/siderite) and glendo-
Phycosiphon (Fig. 3b), Helminthopsis, Planolites nites occur locally.
and Teichichnus. Less common elements are
sporadically distributed, and include Chondrites, Ichnology
Zoophycos, Palaeophycus tubularis and Palaeo- Bioturbation in this facies is characteristically
phycus heberti. Uncommon elements consist of uniformly distributed and intense, varying from
Asterosoma, fugichnia and ?Thalassinoides. BI5 to BI6, and typically BI6. The trace fossil
suite (Fig. 3e) is dominated by Rosselia socialis,
Facies interpretation Zoophycos, Phycosiphon, Planolites, Palaeo-
The moderately diverse trace fossil assemblage phycus heberti, Teichichnus and Rhizocorallium
reflects the reworking of the substrate by diminu- irregulare. Subordinate elements are Asterosoma,
tive deposit-feeding and small grazing/foraging Helminthoida, Chondrites, Palaeophycus tubu-
organisms, and is typical of open marine envir- laris, Macaronichnus simplicatus, Diplocraterion
onments lying below storm wavebase. The rare, habichi and fugichnia. The dominant ichno-
thin sandstone beds are characteristic of distal genera are generally uniformly distributed
tempestites that have been partially reworked throughout the facies, but subordinate elements
by the inhabitants of small, simple burrow occur sporadically.
types that show minimal vertical penetration of
the bedding. The preservation potential of these Facies interpretation
distal tempestites is high, owing to their emplace- This facies differs from Facies A in that it con-
ment below fair-weather wavebase, where physi- tains a greater number of sandstone beds with
cal processes are generally not capable of remnant, low-angle undulatory parallel lamina-
modifying them (Wheatcroft 1990; Pemberton tion (interpreted as HCS and associated with evi-
& MacEachern 1997). Scattered outsized clasts dence of tempestites). The intensely bioturbated,
are indicative of ice rafting. muddy and silty sandstone beds suggest deposi-
The assemblage is interpreted to reflect a distal tion in an environment that experienced lengthy
expression of the Cruziana ichnofacies and periods near, but below, fair-weather wavebase.

Core -12:1, Freitag Formation, (e) Interbedded intensely bioturbated sandy siltstone and very fine- to fine-
grained, thin, sharp bounded laminated sandstone beds that display remnant low-angle undulatory parallel
lamination and 'lam-scram' bedding (LS). Note: above this interval of 'lam-scram' is another interval, and a
Rosselia (Rl) truncated by a tempestite displays a readjustment of the burrow in an upward direction (R2).
The fair-weather assemblage represents a diverse expression of the archetypal Cruziana ichnofacies. Visible
structures here include Palaeophycus heberti (Pa), Phycosiphon (Ph) and Chondrites (Ch). Elements of the
Skolithos ichnofacies include Diplocraterion habichi (Dh). Facies Bl, Tern Formation, (f) Large truncated
Rosselia (R) overprinting a pervasively bioturbated mottled background texture (BI 5) comprising abundant
Planolites (P), Chondrites (Ch) and Phycosiphon (Ph). Facies B2, Tern Formation, (g) Planolites (P),
Phycosiphon (Ph) and Asterosoma (A) representing components of the diverse Cruziana ichnofacies. Facies B2,
Tern Formation.
284 K. L. BANN & C. R. FIELDING

The highly diverse trace fossil suite, dominated Planolites (Fig. 3f). Common but subordinate
by the structures of deposit-feeding and, to a forms include Rosselia rotatus, Palaeophycus
lesser extent, grazing/foraging behaviours is tubularis, Palaeophycus heberti, Asterosoma
characteristic of the archetypical Cruziana ichno- (Fig. 3g), Chondrites, Diplocraterion habichi,
facies and reflects quiescent conditions in upper small Taenidium, Skolithos, Cylindrichnus and
offshore environments lying between storm and ?Thalassinoides.
fair-weather wavebase. Vertical burrows (i.e. The cleaner, well-laminated sandstone beds
elements of the Skolithos ichnofacies) such as are less intensely bioturbated and contain a
Diplocraterion habichi suggest that opportunistic 'lam-scram' appearance, with the BI ranging
suspension-feeding organisms colonized storm from 0 at the base to 4 upwards. The trace
beds. Intensely reworked tempestites are the fossil assemblage is dominated by Diplocraterion
product of greater vertical penetration by more habichi, Diplocraterion parallelum, large Rhizo-
robust deposit feeders, increased size and abun- corallium irregulare (Type A), Phycosiphon,
dance of vertical burrows and increased time fugichnia and Taenidium 'synyphes\ Common
between storm events that facilitates complete but subordinate elements comprise Palaeophycus
colonization and development of deeply pene- tubularis, Macaronichnus isp., Planolites, Rosse-
trating structures (cf. Frey & Goldring 1992). lia socialis, Rosselia rotatus, Teichichnus, Sko-
lithos and Palaeophycus heberti. Uncommon
elements include Psammichnites, Conichnus, Rhi-
Fades B2: Inter laminated, bioturbated sandy zocorallium irregulare Type B, Cylindrichnus and
siltstone and laminated sandstone Arenicolites.

Sedimentology Facies interpretation


This heavily bioturbated facies consists of inter- The palaeoenvironment represented by this
laminated and thinly interbedded siltstone and facies records fair-weather sandy siltstones,
very fine- to fine-grained sandstone. Sandy silt- erosionally truncated and overlain by HCS tem-
stone beds are commonly truncated, and bases pestites, locally with a basal lag, and commonly
are indistinct owing to biogenic mixing across containing escape structures. The diverse trace
the interface with underlying sandstones. Lami- fossil assemblage, dominated by the structures
nated sandstone beds are generally discrete, ero- of deposit-feeding and grazing/foraging organ-
sionally based, 5 cm), and contain 'laminated to isms, represents a diverse expression of the arche-
scrambled' bedding profiles. Outsized clasts and typal Cruziana ichnofacies and reflects infaunal
shelly fossil debris are locally common. Pinstripe reworking of the substrate during fair-weather.
(linsen) to lenticular, wavy and flaser bedding, The trace fossil assemblage associated with
internally comprising ripple cross-lamination, HCS sandstone units is dominated by vertical
ripple form sets, flat lamination and micro HCS burrows of opportunistic suspension-feeders,
(ripple scale) are common. Small-scale HCS resilient surface detritus-feeders and passive
and soft-sediment deformation structures occur carnivores, and is indicative of the Skolithos
locally. ichnofacies. This recurring juxtaposition of
Cruziana and Skolithos ichnofacies (mixed
Ichnology Skolithos-Cruziana ichnofacies: Howard &
This facies is characterized by persistent though Frey 1984; Pemberton & Frey 1984; MacEachern
locally sporadically distributed bioturbation & Pemberton 1992; Pemberton & MacEachern
with intensities varying from BIO to BI5 and 1997) is characteristic of interbedded fair-weather
typically 3-5. Most ichnogenera are uniformly units and tempestites deposited within the upper
distributed throughout the facies, though ele- offshore, lying close to but below fair-weather
ments associated with intercalated laminated wavebase in a moderately storm-influenced
sandstone beds are more sporadically distribu- setting. This zone reflects the transition from
ted. The trace fossil suite can be subdivided the lower shoreface to the offshore/shelf (e.g. off-
into one associated with the sandy siltstone, shore transition: cf. Howard & Reineck 1981).
and one reflecting infaunal colonization of the
laminated sandstone units.
The sandy siltstone beds are intensely bio- Facies CIA: Thoroughly bioturbated muddy
turbated (BI4-5) and dominated by Rosselia sandstone
socialis, Phycosiphon (Fig. 3c, d), Teichichnus,
Helminthopsis, Zoophycos, very large and small Sedimentology
varieties of Rhizocorallium irregulare (Rhizocor- This facies consists of intensely bioturbated
allium type A is very large, type B is small) and muddy sandstone units with rare, thin
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 285

(< 15 cm), discrete, sharply bounded (erosionally moderately to weakly storm-influenced distal
based), fine- to medium-grained sandstone beds. lower shoreface.
Sandstone beds locally contain low-angle, un-
dulatory parallel lamination (HCS), oscillation
ripple lamination, shelly debris, dispersed peb- Facies C1B: Interbedded bioturbated muddy
bles and granules and carbonaceous detritus. sandstone and laminated sandstone
Ichnology Sedimentology
Bioturbation in this facies is pervasive (BI5-6, This facies is composed of thinly to thickly
Fig. 4), with hazy, indiscrete bioturbate textures interbedded, tabular intervals of bioturbated
dominating (Fig. 4c-e). Sandstone interbeds, muddy sandstone and fine- to medium-grained
though present, are largely biogenically sandstone, in discrete, tabular to lensoidal beds,
reworked. Most ichnogenera are uniformly dis- typically 0.2-0.5 m in thickness (Fig. 5a). The
tributed throughout the facies. muddy sandstone beds are similar to units
The trace fossil suite is dominated by Rosselia within Facies CIA. Thicker sandstone beds
socialis (Figs lid, 12d), Rosselia rotatus, Phyco- generally display erosional bases, and units are
siphon (Fig. 4a, c), Diplocraterion habichi, Tei- commonly amalgamated. Minor scattered
chichnus (Fig. 4h), Rhizocorallium irregulare gravel and discrete thin conglomerate beds and
Type B (Fig. 4b, g-h), Rhizocorallium irregulare single clast layers are locally common. Siltstone
Type A, Zoophycos (Fig. 4a), Diplocraterion and thinly interbedded siltstone/sandstone inter-
parallelum (Fig. 4f), Macaronichnus isp. (Fig. vals occur locally and contain pinstripe (linsen)
4c), Planolites (Fig. 4j), Palaeophycus tubularis and lenticular bedding. Thicker sandstone beds
(Fig. 4c) and Palaeophycus heberti (Fig. 4b, d). show gravel lags (basal and/or top surface), flat
Common but subordinate elements are and low-angle lamination, and HCS are gener-
Conichnus, Taenidium isp., Taenidium, Skolithos, ally laminated to burrowed or sparsely to non-
Cylindrichnus (Fig. 4e), Siphonichnus, Schaub- bio turbated. Bed tops contain symmetrical
cylindrichnus, Asterosoma, fugichnia, Psamm- ripples, interference ripples or coarse-grained
ichnites, Scolicia, Helminthopsis (Fig. 4a) and symmetrical ripples. Shelly debris, dispersed
Chondrites (Fig. 4i). pebbles and granules, soft-sediment deformation
structures and carbonaceous detritus are locally
Facies interpretation common.
Intensity and uniformity of bioturbation in the
muddy sandstones and the lack of primary sedi- Ichnology
mentary structures indicate either limited storm The interbedded, bioturbated muddy sandstone
influence on the environment or considerable and laminated sandstone units are characterized
time between storms, during which the fair- by highly variable bioturbation intensities,
weather assemblage reworked the substrate. ranging from BI0 to BI5. Bioturbated sandstone
The fair-weather trace fossil suite, dominated beds range from BI 3 to BI 5, whereas the lami-
by a very diverse mixture of robust, complex nated sandstone units have bioturbation intensi-
structures produced by deposit- and detritus- ties between BIO and BI4. Bioturbation and
feeding organisms, is characteristic of a diverse trace fossil distribution is more or less persistent
proximal expression of the Cruziana ichnofacies. within the muddy sandstone beds but sporadi-
Considerable numbers of vertical structures, cally distributed in the laminated sandstone
such as Diplocraterion parallelum, reflect a layers, where 'lam-scram' bedding predominates.
component of the Skolithos ichnofacies and sug- The trace fossil assemblage in the muddy
gest that storm beds were rapidly colonized by sandstone beds is dominated by robust Rosselia
opportunistic, suspension-feeding organisms, socialis (Figs 5a, b, 12e), Rosselia rotatus, Diplo-
prior to their recolonization and thorough craterion habichi, Diplocraterion parallelum (Figs
reworking by the resident fair-weather commu- 5b, lie), Teichichnus, Phycosiphon (Fig. 5a-c),
nity (cf. Frey & Goldring 1992). Locally occur- Palaeophycus tubularis (Fig. 5b-c), Planolites
ring massive sandstone beds with indistinct (Fig. 5a-c) and Rhizocorallium irregulare types
bioturbate textures suggest that an infaunal com- A (Fig. 5b) and B. Common but subordinate ele-
munity occupied a soupy substrate with high ments include Macaronichnus isp., Arenicolites,
porewater content. Rare, thin, sharp-based Cylindrichnus and Skolithos. The trace fossil
HCS sandstone beds represent tempestites. suite in the laminated sandstone beds is domi-
These features indicate deposition in a well- nated by Diplocraterion habichi and fugichnia
oxygenated, open marine setting at or just (Fig. 5c). Diplocraterion parallelum (Fig. 5d),
above fair-weather wavebase consistent with a Skolithos (Fig. 5d), Rhizocorallium jenense,
286 K. L. BANN & C. R. FIELDING

Fig. 4. Characteristic ichnofossils in Facies CIA. (a-d) Intensely reworked muddy sandstone with a mottled
bioturbate texture (BI5-6), Tern Formation. Bioturbation has occurred in several phases, with mottled
background textures overprinted by larger more discrete structures such as IZoophycos (Z?) and Rhizocorallium
irregulare type B (Rh). These structures have been reworked by small gazing and foraging organisms
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 287

Arenicolites, Cylindrichnus and Conichnus repre- Ichnology


sent subordinate elements. Bioturbation is variable, ranging from BI0 to a
very rare maximum of 5. The average BI is 1-3,
Fades interpretation many beds have little or no bioturbation pre-
The heterogeneity of this fades demonstrates served, and 'lam-scram' profiles are common.
variations in physical energy and sedimentation The trace fossil assemblage is less diverse than
rates. Most of the physical structures reflect in Facies C1B. The suite is dominated by
storm deposition, particularly HCS, low-angle robust vertical structures such as Diplo crater ion
planar cross-stratification, and small- to large- parallelum, long deeply penetrating Diplocration
scale waning-stage combined flow ripples. Non- habichi (Fig. 6c), Skolithos and Arenicolites.
burrowed siltstone interbeds are interpreted to Common but subordinate elements comprise
reflect post-storm draping of tempestites during Macaronichnus segregatis, robust vertical mud-
waning flow conditions, rather than fair-weather filled structures such as Rosselia socialis, Cylin-
deposition. Alternations between episodic tem- drichnus, and Parahaentzschelinia surlyki (Fig.
pestite emplacement and thorough bioturbation 6a, b). Fine conglomerate beds are generally
of the interbedded muddy sandstone record unbioturbated but locally contain Skolithos,
changes in sedimentation rates and storm influ- Arenicolites and Diplocraterion habichi.
ence. The abundance of fugichnia in the tempes-
tites supports episodically high sedimentation Facies interpretation
rates and erosional amalgamation of the beds. This facies is characterized by sedimentary
The bioturbated sandstone beds contain a rela- structures that reflect deposition in a high-
tively diverse trace fossil suite that records a energy, wave-dominated environment. The
distal expression of the Skolithos ichnofacies, thick amalgamated units of SCS, HCS and low-
consistent with sedimentation within the more angle planar cross-stratified sandstones are
proximal portion of the lower shoreface. characteristic of storm deposits. The thickest
amalgamated units indicate deposition in a
storm-dominated setting above storm wavebase.
Fades C2: Interbedded massive and More heavily bioturbated units represent the
laminated sandstone biogenic reworking of storm-deposited sand
during fair-weather periods in a well-oxygenated
Sedimentology open marine environment above fair-weather
This facies consists of thickly interbedded, tabu- wavebase. The absence of muddy interbeds also
lar intervals of both clean, massive sandstone reflects deposition above fair-weather wavebase.
and laminated fine- to medium-grained sand- The trace fossil suite is dominated by burrows
stone. Gravel, clast layers and discrete thin of suspension-feeding organisms, and represents
conglomerate beds are common locally. Thick the Skolithos ichnofacies. The general lack of
sandstone beds have erosional bases, gravel lags complex, horizontal burrows formed by
(basal and/or top surface), flat and low-angle deposit-feeding organisms indicates high storm
planar cross-stratification, large-scale HCS and intensity and/or frequency, and reflects a paucity
swaley cross-stratification (SCS). Bed tops are of interstitial food material for infaunal
covered by symmetrical ripples, interference deposit feeders (MacEachern & Pemberton
ripples or symmetrical gravel ripples locally. 1992). Vertical, robust, mud ball (Rosselia-type)

interpreted as Helminthopsis (H) and Phycosiphon (Ph). The mottled background texture represents the
repeated reworking of successive storm and fair-weather units during fair-weather periods in a distal lower
shoreface environment. The trace fossil suite comprises a mixture of subvertical/vertical structures, representing
elements of the Skolithos ichnofacies, such as Palaeophycus tubularis (Pt), Diplocraterion parallelum (Dp) and
Skolithos (S), and the structures of a diverse range of grazing/foraging, deposit- and detritus-feeding organisms
and passive carnivores representing a diverse proximal expression of the Cruziana ichnofacies. Structures visible
here include Phycosiphon (Ph), Planolites (P), Palaeophycus heberti (Pa), Chondrites (Ch), Rosselia (R),
Schaubcylindrichnus (Sc) and Macaronichnus sp (M). (e) Thoroughly bioturbated silty sandstone with a
mottled bioturbate texture overprinted by long thin Cylindrichnus (Cy), Catherine Sandstone, Arcturus
Core -6:3. (f) Heavily bioturbated sandstone with Diplocraterion parallelum (Dp), GSQ Springsure Core -17.
(g) An example of the top portion of a Rhizocorallium irregulare type B (Rh), a structure that is very common
in Permian shoreface facies. Complex structures such as this are often difficult to identify in core, GSQ
Springsure Core -17. (h) Teichichnus (Te), GSQ Springsure Core -17. (i) Pervasively bioturbated sandstone, the
bioturbate texture comprising tubular structures such as Phycosiphon (Ph), Planolites (P) and Palaeophycus
tubularis (Pt), Tern Formation, (j) Pervasively bioturbated mudstone bed, Planolites (P), Tern Formation.
288 K. L. BANN & C. R. FIELDING

structures reflect the domiciles and associated Pemberton et al (2001) also considered Rosselia
waste disposal activities of resilient detritus- to be typical of the Skolithos ichnofacies. The
feeding organisms. These structures are physical characteristics and dominance of the
common components of the Skolithos ichno- Skolithos ichnofacies suggest a middle shoreface
facies in Permian deposits from Australia. environment.

Fig. 5. Characteristic bedding styles and ichnofossils in Facies Clb. (a-c) Interbedded laminated clean
sandstone and heavily bioturbated muddy sandstone, Tern Formation. The trace fossil assemblage is
dominated by a relatively diverse mixture of vertical and horizontal structures such as Diplocraterion parallelum
(Dp) and Skolithos (S), robust heavily lined Rosselia socialis (R), Phycosiphon (Ph), large Planolites (P), large
Palaeophycus tubularis (Pt), Chondrites (Ch), and possible large Rhizocorallium irregulare type A (Rh).
Small pervasive Helminthopsis (H) and Phycosiphon (Ph) have reworked the muddy linings of other larger
structures. Fugichnia are common in the laminated sandstone beds (f). (d) Clean sandstone with examples
of Skolithos ichnofacies elements, Diplocraterion parallelum (Dp) and Skolithos (S), Lower Aldebaran
Sandstone, Arcturus Core -6:4. (e) Bioturbated sandstone with Skolithos, Staircase sandstone, GSQ Core
Springsure -16.
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 289

Fig. 6. (a-c) Outcrop examples of Fades C2, German Creek Formation, Bundoora Dam, Queensland,
(a, b) Large, vertical, mud-filled burrows interpreted as Parahaentzschelinia surlyki. (c) Long curved
Diplocraterion habichi. (d) Tiering profile for Facies C3. Shallower tiers are occupied by the subvertical to
vertical dwelling structures of passive carnivores and opportunistic suspension-feeding organisms. Ichnotaxa
include Palaeophycus tubularis (Pt), Diplocraterion habichi (Dh), Conichnus (C), Rhizocoralium jenense (Rj),
Arenicolites (A), Diplocraterion parallelum (Dp), Skolithos (S) and very robust, vertical Cylindrichnus (Cy).
The deepest tier is occupied by Macaronichnus segregatis (Ms). This trace fossil suite represents the Skolithos
ichnofacies. (e) Outcrop example of pervasive Macaronichnus segregatis in Facies C3, Freitag Formation,
Fairbairn Dam, Queensland.
290 K. L. BANN & C. R. FIELDING

Fades C3: Amalgamated laminated and deeply burrowing suspension-feeding organisms


cross-bedded sandstone that were able to withstand the continually
migrating bed-forms. The Macaronichnus trace-
Sedimentology maker is most common in deposits from very
Thick amalgamated tabular beds and composite high-energy environments, typically from
units of clean, moderately to well-sorted, quart- around and above the upper shoreface-foreshore
zose, generally medium-grained sandstone are contact (Saunders & Pemberton 1986; Saunders
typical of this facies along with scattered gravel et al. 1994). The preservation potential of Macar-
and discrete beds and single clast layers of con- onichnus segregatis was high, despite the fact that
glomerate. Beds show basal and internal erosion it represents the activities of a deposit feeder,
surfaces, gravel lags (basal and/or top surface), owing to the deep-tier position that it occupied
trough cross-stratification and low-angle bidirec- (Saunders & Pemberton 1986; Pemberton et al.
tional planar cross-bedded sets. Low-amplitude/ 2001). In the intertidal and innermost surf zone
long-wavelength and swale-dominated HCS are of wave-exposed beaches, oxygenated surface
locally preserved. waters can circulate several metres into the
sand, well below the reach of surface wave
Ichnology disturbance (Pemberton et al. 2001). In addition,
Trace fossils are locally common but rarely large volumes of dissolved and fine particulate
abundant, and assemblage diversity is generally organic matter, most of which is mineralized at
low. Most units have little or no bioturbation, depth, is filtered through the porous sandy
with the BI ranging from 0 to a very rare maxi- system of the beach (McLachlan et al. 1984).
mum of 4. The average BI is 0-2. Where the This phenomenon creates a second habitat at
degree of bioturbation is high, the trace fossil depth, separate from the physically controlled
assemblage is usually dominated by one ichno- habitat at the surface. The deeper habitat is
species. Lam-scram profiles are preserved locally stable, predictable, and is constantly replenished
but are uncommon. The trace fossil suite is with food material (Pemberton et al. 2001).
characterized by vertical structures and pervasive Saunders & Pemberton (1986) suggested that
Macaronichnus segregatis (Fig. 6d-e). Different the Macaronichnus organism fed on microorgan-
ichnotaxa are locally more prevalent, but the isms several metres below the sediment/water
most common forms overall include Macaronich- interface.
nus segregatis, Skolithos, Conichnus, Arenicolites
and Diplocraterion parallelum. Diplo crater ion
habichi, Rhizocorallium jenense, Palaeophycus Deltaic deposits
tubularis and vertical, robust Cylindrichnus are
locally common but subordinate overall. A Facies El: Unbioturbatedgravelly sandstone
general lack of silt in burrow fills and linings
produces a pervasive bioturbation that is not Sedimentology
always obvious, but can be seen clearly in some This facies is characterized by sharp-based,
examples (Fig. 6e). coarsening-upward intervals of unbioturbated,
fine- to coarse-grained sandstone. Scattered
Facies interpretation gravel, intraformational clasts, plant debris and
Sedimentary structures in this facies reflect associated coaly fragments are common. The
deposition by shore-parallel wave-driven cur- dominant visible structure is high-angle trough
rents interacting with currents that are generated cross-bedding in sets typically <0.5m thick.
by translatory flow and plunging waves (Mac- Internal erosion surfaces, flat parallel-bedded
Eachern & Pemberton 1992). Such structures cross-lamination and ripple cross-lamination
are typical of upper shoreface deposits (Reinson are preserved locally. Hummocky cross-stratifi-
1984). Storm deposits in the upper shoreface cation is rare.
occur as ridge and runnel systems rather than
major depositional events, as they are in the Ichnology
lower and middle shoreface, and indicate erosion No trace fossils have been recognized in this
of the beach face and transport of sediments to facies.
the more distal parts of the shoreface (Mac-
Eachern & Pemberton 1992). Facies interpretation
The trace fossil suite represents an impover- Lack of bioturbation in this facies suggests an
ished expression of the Skolithos ichnofacies. inhospitable depositional environment. Migrat-
Most of the trace fossils are heavily lined, vertical ing large-scale bedforms present a problem for
to subvertical, and represent the domiciles of infaunal colonization, so that only deeply
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 291

penetrating trace fossils might be expected (Mac- lining; rather the burrow mantle reflects
Eachern 2001). The lack of vertical burrows, selective particle ingestion with dark micaceous
which reflect the domiciles of opportunistic sus- material discarded to the outer edge of the
pension feeders, may reflect rapid and/or alter- structure. This type of specialized, grain-selective
nating sedimentation rates, salinity fluctuations, deposit feeding is also associated with other
increased turbidity levels and rapidly shifting Macaronichnus species (Saunders & Pemberton
bedforms in a river mouth environment. Bio- 1986; Saunders et al. 1994; Pemberton et al.
genie structures produced in such high-energy 2001).
environments stand little chance of preservation
(Bromley 1996). Facies interpretation
The well-sorted, medium-grained nature of this
facies and the abundance of large-scale, swale-
Fades E2 Amalgamated, tabular sandstone dominated HCS and flat parallel and low-angle
cross-lamination suggest deposition in a wave-
Sedimentology dominated marine environment above fair-
This facies comprises amalgamated intervals of weather wavebase. An abundance of internal
generally medium-grained, well-sorted, quart- erosion surfaces supports this interpretation.
zose sandstone with rare gravel. Sandstone The well-sorted tabular sand bodies suggest
bodies are tabular (Fig. 7a), with generally that the depositional environment was exposed
tabular internal architecture also, and locally dis- to continuous reworking (i.e. by river outflow
play overall coarsening-upward trends and abun- currents and sea waves). The facies is character-
dant internal erosion surfaces. Low amplitude istic of wave-reworked mouth bar deposits, the
but long-wavelength, swale-dominated forms of finer fraction continuously removed and swept
HCS, flat parallel and low-angle lamination are out to sea where it is then deposited in more
common. distal delta front and prodeltaic settings. The
paucity of vertical burrows and the low-diversity
Ichnology trace fossil suite (dominated by pervasive
Trace fossils are mostly absent but, where they Macaronichnus isp.) represents an ichnological-
are present, bioturbation is of low intensity and assemblage composed of the sporadically
is very sporadically distributed. The trace fossil distributed burrows of passive carnivores and
suite is of very low-diversity and is usually deposit-feeding organisms. The assemblage
dominated by Macaronichnus isp. (Figs 8a, b, reflects a stressed but proximal expression of
14d) that contains a diverse and complex range the Cruziana ichnofacies, suggesting that for
of burrow infill (passive, active and meniscate the most part the sandy substrate was inhospita-
forms, Fig. 9c). Teichichnus, Palaeophycus tubu- ble to infaunal organisms and especially so to
laris (Fig. 8b) and Planolites (Fig. 8a) are very suspension feeders. The physical sedimentary
rare. structures, vertical facies succession and ichno-
At first glance the burrows interpreted herein logical characteristics indicate deposition in a
as Macaronichnus isp. appear to have the charac- mixed river- and wave-influenced mouth bar
teristics of more than one ichnogenus (Palaeo- environment.
phycus, Planolites and Macaronichnus isp.).
Closer inspection, however, indicates that the
tubular burrows actually represent different Facies E3: Interbedded tabular sandstone
components of the same biogenic structure. The and moderately bioturbated sandstone
cross-sectional shape and size of the burrows is
consistent, but the linings and infill of the struc- Sedimentology
tures are quite variable (Fig. 9b-c). Whereas This facies consists of thickly interbedded,
some parts of the burrow system appear to tabular intervals of fine- and medium-grained
have been lined and then remained open (thus sandstone in sharp-bounded (erosionally based)
being passively infilled), there is evidence that beds and composite units <1.5m thick inter-
other parts are actively filled and are thought to bedded with bioturbated sandstone. Inter-
represent backfilled or waste-stuffed chambers. laminated siltstone/fine-grained sandstone beds,
The composite morphology of this structure scattered gravel, discrete thin conglomerate
suggests a time lag across the mantle/infill beds and single clasts occur locally.
discontinuity (conformant reprobing/branching) The thick sandstone beds contain basal and/or
in the form of meniscate infill, local collapse etc. top surface gravel lags, flat parallel and low-
(T. Saunders personal communication 2003). angle lamination and HCS. Onshore-directed
Many examples do not contain an obvious cross-bedding is rare. Locally, bed tops are
292 K. L. BANN & C. R. FIELDING

Fig. 7. (a) Outcrop view of Fades E2, Crocker Sandstone, Capricorn Highway Blackwater to Emerald,
Queensland, (b-d) Facies E3, Lower Aldebaran Sandstone, (b) outcrop view, (c, d) equivalent core from GSQ
Springsure -16 (see sedimentological log, Fig. 13). (b) Macaronichnus isp. with burrow crossover marked by
arrow. Cattle Creek Nianda, Queensland, (c) Sandstone units with low-angle lamination and Rosselia mud-ball
rip-up clast (R), Phycosiphon isp. (Ph) and Macaronichnus isp. (M). (d) Amalgamated low-angle laminated
sandstone with Macaronichnus isp. (M) and truncated fugichnia (f).
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 293

Fig. 8. (a, b) Facies E2. Medium-grained sandstone with Macaronichnus isp. (M), Palaeophycus tubularis (Pt)
and Planolites (P). (c-e) Facies E3. (c) Outcrop view of pervasive Macaronichnus isp. individual burrows are
clearly discernible and have a micaceous mantle (M). Lower Aldebaran Sandstone, Cattle Creek Nianda,
Queensland, (d, e) Soft-sediment deformation structures in core. Also visible is the sporadic nature of
bioturbated intervals characteristic of this and the other deltaic facies. An interval of moderately bioturbated
silty sandstone containing Macaronichnus isp. (M) and Phycosiphon (Ph) is visible in (e). Catherine Sandstone,
GSQ Springsure core -18.

covered by symmetrical ripples, interference Ichnology


ripples or coarse-grained symmetrical ripples. This facies is characterised by a sporadically dis-
Soft-sediment deformation structures (Fig. tributed, moderate to low-diversity trace fossil
8d, e) and carbonaceous detritus are common. assemblage. Bioturbation intensities are highly
294 K. L. BANN & C. R. FIELDING

Fig. 9. Fades E4. (a) Discrete laminated clean sandstone beds and interbedded bioturbated siltstone and
sandstone visible in outcrop. Lower Aldebaran Sandstone, Cattle Creek Nianda, Queensland, (b) Macaronichnus
isp. (M) forming a pervasive bioturbate texture. Note the overprinting of individual burrow cross-sections.
Aldebaran Sandstone, Arcturus Core -6:4. (c) Macaronichnus isp. (M) in core, showing variation in burrow fill
(one of the unusual characteristics of this ichnospecies). Freitag Formation, GSQ Springsure -17.
(d) Carbonaceous detritus (ca) and wood fragments (w) in core, Freitag Formation, GSQ Springsure -17.
(e) Distal delta front deposits in core showing interbedded low-angle cross-bedded sandstone (LA) and
bioturbated interlaminated siltstone and sandstone with a very low-diversity trace fossil suite comprising
Macaronichnus isp. (M), Planolites (P) and Phycosiphon (Ph). Lower Catherine Sandstone, GSQ Springsure -18.

variable throughout the fades, and range from 0 facies E2. Corkscrew-shaped varieties are locally
to 4. Bioturbated intervals are thin (<20 cm), and common (Fig. 14c). This type of morphology
are interbedded with non-bioturbated laminated would ordinarily be classified as Gyrolithes. Phy-
sandstone intervals. Overall the trace fossil suite cosiphon (Fig. 7c), Planolites, Rosselia socialis
is dominated by Macaronichnus isp. (Figs 7b-d, and Cylindrichnus are rare subordinate elements
8c). In this facies these burrows display an even that occur locally. Rosselia mud balls, preserved
wider variety of preservational forms than in as rip-up clasts (Fig. 7c), and fugichnia are
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 295

common in some laminated sandstone beds (Fig. lamination. Ripple form sets, flat parallel lamina-
7d). There is a noticeable paucity of vertical bur- tion and micro-hummocky cross-stratification
rows, but Diplocraterion habichi occurs locally. (ripple scale) are common in these intervals.
Claystone beds generally occur as thin (<3cm)
Fades interpretation drapes on the tops of sandstone beds, are rarely
This facies is broadly similar to Facies C2 and C3 bioturbated (Fig. 13b), and locally contain
in that it comprises mainly amalgamated lami- synaeresis cracks. Carbonaceous detritus and
nated sandstone bedsets interbedded with biotur- woody material are abundant throughout this
bated sandstone beds. These features indicate facies (Fig. 9d). The facies passes conformably
that the depositional environment was character- upwards into proximal delta front and mouth
ized by breaking waves, surf zone conditions and bar deposits (Figs 13, 14). Many intervals show
ultimately wave swash, typical of facies deposited rhythmic interbedding of sand and silt/clay.
above fair-weather wavebase in conditions similar
to that of the middle to upper shoreface. There is Ichnology
significant evidence for storm wave reworking of Bioturbation in this facies is sporadically distrib-
the substrate such as large-scale HCS (amplitude uted, and many intervals are not burrowed (Fig.
0.3-0.5m, wavelength up to 20m), internal ero- lOa, c). Bioturbation intensities are variable, with
sion surfaces and the excavation and re-deposition the overall BI ranging from 0 to 5. The trace
of Rosselia mud balls as rip-up clasts. Ichno- fossil suite is dominated by large Macaronichnus
logically, this facies differs from shoreface deposits isp. (Figs 9b, c, lOa, c, d, e, 13b, 14b) and Phyco-
in subtle ways. The trace fossil suite is similar to siphon (Figs lOa, d, 14b), which occurs as both
that from mouth-bar deposits, but bioturbated very robust and diminutive forms. These two
intervals here are thicker and more abundant, trace fossil types occur in sporadically distribu-
and intensities are higher. The trace fossil assem- ted but pervasively bioturbated horizons (Fig.
blage represents a stressed but proximal expres- 13b). Fugichnia comprise a common but sub-
sion of the Cruziana ichnofacies. Integrated with ordinate element of the assemblage. Teichichnus,
the sedimentological characteristics and vertical Rosselia socialis, Rosselia rotatus (Fig. lOb),
facies succession, this ichnofacies suggests deposi- Chondrites (Fig. 13b), Rhizocorallium irregular e,
tion in a wave-influenced proximal delta front Asterosoma, Psammichnites, Planolites (Fig. 9e)
environment. The delta front, in general, usually and Zoophycos occur locally but are rare. Tei-
forms a relatively steep slope, and high sedi- chichnus, Rosselia, Asterosoma and Zoophycos
mentation rates and slope instability result in occur as diminutive forms. Vertical burrows
penecontemporaneous deformation, slumping such as small Diplocraterion habichi occur in
and gravity movements, producing soft-sediment sparse quantities.
deformation structures such as those seen in
Figure 8d-e. Facies interpretation
The interbedded laminated sandstones and bio-
turbated intervals clearly display characteristics
Facies E4: Interbedded sandstone, similar to those of a storm-dominated lower
bioturbated silly sandstone and clay stone shoreface deposit. Examples include HCS,
planar parallel to low-angle cross-bedding, lam-
Sedimentology scram bedding and heavily bioturbated intervals
This facies comprises discrete, sharp-bounded, that reflect periods of quiescence associated with
laminated clean sandstone beds, bioturbated fair-weather. Ichnologically, however, this facies
sandstones, interlaminated and thinly inter- differs subtly from lower shoreface deposits. The
bedded siltstone and very fine- to fine-grained trace fossil suite is reduced in diversity, indivi-
sandstones and thin claystone beds (Fig. 9a). Out- dual ichnogenera differ in size as compared
sized clasts and shelly fossil debris are common with their non-deltaic shoreface counterparts,
locally. Small-scale HCS is the dominant primary the uniformity of burrowing is sporadic, degree
structure in the laminated sandstones, and many of bioturbation is more variable and generally
beds display 'lam-scram' bedding and common less intense, and there is a noticeable paucity of
fugichnia. The coarsest grained sandstone beds vertical burrows. The trace fossil suite represents
contain planar parallel cross-bedding or low- a depauperate expression of the Cruziana ichno-
angle cross-bedding. The interlaminated and facies, which indicates that the depositional
interbedded sandstone and siltstone intervals environment was physically or chemically hostile
contain common soft-sediment deformation to infaunal organisms.
structures, pinstripe (linsen) to lenticular, wavy The non-bioturbated, carbonaceous claystone
and flaser bedding with internal ripple cross- layers represent post-storm drapes on top of
296 K. L. BANN & C. R. FIELDING

Fig. 10. Facies E4. (a) Sporadic distribution of bioturbated intervals and intervals that lack any evidence of
burrowing activity. The trace fossil suite is of very low diversity and comprises Macaronichnus isp. (M) and
Phycosiphon (Ph). Lower Catherine Sandstone, GSQ Core Springsure -18. (b) Slender Rosselia rotatus (R) in
low-angle cross-bedded sandstone that shows little evidence of other burrow types. The base of this bed sharply
truncates the underlying non-bioturbated, laminated silty sandstone unit. Freitag Formation, GSQ Core
Springsure -17. (c) Sharp-based laminated sandstone that truncates a bioturbated interval with Macaronichnus
isp. (M) and Phycosiphon (Ph). Freitag Formation, GSQ Core Springsure -17. (d) Characteristic trace fossil
suite in Permian distal delta front deposits, sporadically distributed large Macaronichnus isp. (M) and robust
Phycosiphon (Ph). Freitag Formation, Rolleston Core -11:2. (e) Pervasively burrowed silty sandstone interval
within distal delta front facies. The trace fossil suite is of very low diversity and comprises Macaronichnus isp.
(M) that displays burrow overlap and a variety of burrow infill, Teichichnus (T) and Phycosiphon (Ph).
Aldebaran Sandstone, Arcturus Core -6:4.
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 297

tempest! tes. Similar clay stone drapes have been with persistent environmental fluctuations. The
described from delta front deposits by Saunders fine-grained, interlaminated nature of the facies
et al (1994), Gingras et al (1998) and Bann records fair-weather deposition in a relatively
et al. (2004). quiescent environment, below fair-weather wave-
Physical structures and ichnological character- base. The sparse, thin sandstone beds represent
istics similar to those in this facies are known distal tempestites. The association of synaeresis
from wave-dominated deltas in the Upper cracks with the claystone layers suggests
Cretaceous Dunvegan and Basal Belly River fluctuating salinity levels associated with
Formations in North America (Gingras et al. heightened precipitation and increased fluvial
1998; Coates & MacEachern 1999). This facies discharge during storm events. The facies has
reflects deposition in the distal delta front of a similar physical characteristics to offshore
wave-influenced delta. deposits, but the vertical facies succession and
stressed ichnological signature suggest deposi-
tion in a prodeltaic environment.
Facies E5: Interlaminated siltstone,
clay stone and silty sandstone
Case study 1: The Lower Aldebaran
Sedimentology Sandstone
This facies underlies Facies E3 and E4 (Figs 13a,
14a) and consists of mainly siltstone and clay- The Lower Aldebaran Sandstone is recognized as
stone with minor to moderate proportions of a mixed-influence (wave-tide-fluvial-interactive)
interlaminated and admixed very fine- to fine- delta deposit (Baker 1991). This diagnosis is
grained sandstone. Discrete, sharp-bounded based on the presence of erosively-based, cross-
sandstone beds <5cm thick are rare. Inter- bedded sandstone deposits (Facies Fl, distribu-
lamination structures range from pinstripe tary channel), the radiating pattern of palaeo-
(linsen) to lenticular and wavy bedding, intern- current directions, and the arcuate plan shape
ally comprising ripple cross-lamination, ripple of the body (Baker 1991; Fielding et al. 1995,
form sets, flat lamination and micro-hummocky 2000). This study identified a significant propor-
cross-stratification (ripple scale). Scattered out- tion of Facies E components, further supporting
sized clasts, marine body fossils and shell debris a deltaic interpretation for the Lower Aldebaran
are rare. Abundant carbonaceous detritus, soft- Sandstone.
sediment deformation structures and carbonate The unit overlies the uppermost mudstone
cements (typically ankerite/siderite) occur member of the Cattle Creek Formation, the
locally. The claystone beds also contain sparse, Sirius Mudstone Member (Figs 2, 9a, 13). In out-
small synaeresis cracks. crop, the contact is defined on the basis of expo-
sure character: the Sirius Mudstone Member
Ichnology forms deeply gullied, but flat terrain, whereas
As for the sandier deltaic deposits described the overlying Lower Aldebaran Sandstone forms
above, this facies is characterized by variable hilly country with small sandstone cliff exposures.
levels of bioturbation. The degree of reworking The contact is easily traced from aerial photo-
varies from BIO to BI5. The trace fossil suite graphs. In many subsurface cores and surface
(Fig. 15a) is dominated by small Macaronichnus exposures, such as in Cattle Creek, the contact
isp. and Phycosiphon. Diminutive Asterosoma, marks a significant and abrupt change in grain
Planolites, Chondrites, fugichnia and Helmin- size and depositional setting, and is interpreted
thopsis are relatively common but subordinate. as a sequence boundary. In other outcrops and
Teichichnus and Zoophycos comprise uncommon cores however, the contact is often cryptic, with
elements of the assemblage, both occurring as a continuous coarsening upward trend from one
diminutive forms. Vertical structures are gener- unit into the other (e.g. Fig. 13a).
ally absent. The characteristic sedimentary succession of the
Lower Aldebaran Sandstone is seen in Figure 13a.
Facies interpretation The interval consists of a coarsening-upward
The trace fossil suite in this facies is dominated facies succession, showing a transition from the
by a relatively diverse but diminutive and spora- muddier prodelta deposits of the upper Sirius
dically distributed mixture of structures, pro- Mudstone Member into the sandier delta front
duced by grazing/foraging and deposit-feeding and mouth-bar facies of the Lower Aldebaran
behaviours. This assemblage represents a Sandstone. The ichnological signature of this
stressed distal expression of the Cruziana ichno- succession is one of a fully marine trace
facies, and indicates fully marine conditions fossil assemblage that experienced significant
298 K. L. BANN & C. R. FIELDING

Fig. 11. (a) Sedimentological section of the contact between the Moorooloo Mudstone Member and the
overlying Staircase Sandstone Member (both components of the Cattle Creek Formation), from GSQ Core
Springsure -16. The section comprises a coarsening and shallowing upwards succession. The bioturbation is
intense (BI5-6) and uniformly distributed throughout the interval. Diminishing BI readings in the upper half
of the succession represent 'lam-scram' bedding in tempestites. The overall trace fossil suite is diverse and
contains a mixture of structures produced by the activities of grazing/foraging behaviours, simple and complex
deposit-feeding strategies and detritus-feeding organisms. Vertical structures become common in the distal
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 299

environmental and possibly chemical stresses and could not be positively associated with a
(Fig. 13b). In comparison with the ichnological delta unless good three-dimensional control was
signature of non-deltaic shoreface successions available (Bhattacharya & Walker 1992). How-
the trace fossil suites are reduced in diversity, ever, in the Permian successions in this study,
individual ichnogenera differ in size, uniformity ichnological signatures have provided a powerful
of burrowing is sporadic, the degree of bioturba- additional means in the recognition and identifi-
tion is more variable and significantly less intense, cation of deltaic depositional environments.
and there is a distinct paucity of vertical burrows.
A section of core (Fig. 13b) from around the
340m mark on the core in Figure 13a represents Case study 2: The Freitag Formation
distal delta front deposits. The trace fossil suite
is dominated by Macaronichnus isp. and Phycosi- The enigmatic Freitag Formation overlies the
phon. The Macaronichnus isp. displays a variety coarse-grained Upper Aldebaran Sandstone in
of morphology and burrow fills. The large exam- the northern Denison Trough (Fig. 2a). An accu-
ples in the basal half of the core distinctly suggest rate plan shape is indicated by isopachs, but the
a Gyrolithes-type spiral morphology, whereas the northern extent of the unit is difficult to define
smaller examples in the upper part of the core owing to sparse data. The arcuate, eastward-
appear to maintain a more consistently horizon- bulging plan shape, though incomplete, is sug-
tal nature, possibly suggesting that burrow gestive of a delta. Previous work by Fielding &
construction (in the larger example) took place McLoughlin (1992) on a large surface exposure
during increased sedimentation rates. at Fairbairn Dam in the northernmost Denison
The deltaic succession passes up into inter- Trough suggested a non-deltaic coastal platform,
bedded muddy and sandy interdistributary bay although the exposed section included a thick
fill deposits with highly reduced trace fossil erosionally based cross-bedded sandstone body
suites characteristic of Permian opportunistic, (Facies Fl) interpreted as a major distributary
brackish-water trace fossil assemblages (see channel deposit. This study, and other recent
Bann et al. 2004). The lower delta plain interval work by Bann & Fielding (2001) and Trueman
is overlain by sandy, cross-bedded distributary (2002), suggested that the Freitag Formation
channel deposits. This succession (Figs 11 a, contains a complex array of progradational
12a) displays many of the characteristics of cycles, some representing deltas and others
deposits formed in prograding wave-dominated coastal platform.
shoreface settings (Curray et al. 1969; Fielding The base of the unit is a well-defined flooding
1989; Boyd et al. 1992; MacEachern & surface, with fine-grained facies passing upward
Pemberton 1992; Walker & Plint 1992 and refer- into sandstone-dominated facies. A series of
ences therein; Bann et al. 2003). Prograding broadly coarsening-upward parasequences 5-
successions are also characteristically deposited 40m thick are recognized here (Fig. 14a) from
during progradation of delta lobes (Elliott the cored stratigraphic drill-hole GSQ Spring-
1986; Coleman & Wright 1975; Bhattacharya & sure-18. A complete coarsening-upward parase-
Walker 1992; Gingras et al. 1998; Coates & quence, bounded by interpreted transgressive
MacEachern 1999). Until recently, successions surfaces of erosion, is visible in Figure 14a.
such as this could be interpreted only as a pro- The succession is physically and sedimento-
grading wave- or storm-dominated shoreface logically very similar to the prograding shoreface

lower shoreface and increase upwards to be abundant in the proximal lower shoreface deposits, (b-e) Changing
upwards through the prograding shoreface succession, (b) Facies Al, shelf al mudstone, in core. Some vague
primary lamination is visible; trace fossil types are difficult to ascertain. Ingelara Formation, GSQ Core
Springsure -17. (c) Characteristic upper offshore deposits. Bioturbation is intense and persistent throughout
the facies, and the trace fossil assemblage is dominated by structures produced by grazing/foraging organisms
such as Zoophycos (Z) and Phycosiphon (Ph), and complex deposit-feeding behaviours such as Asterosoma (A).
Ingelara Formation, GSQ Core Springsure -17. (d) Intensely bioturbated distal lower shoreface sandstone
dominated by the basal tubes of Rosselia (R). Rosselia is one of the most abundant trace fossil types in
Permian shoreface deposits, and represents an element of both the Cruziana and Skolithos ichnofacies.
In this example Rosselia represents a component of a proximal expression of the Cruziana ichnofacies.
Freitag Formation, GSQ Core Springsure -17. (e) Characteristic proximal lower shoreface deposits.
A passively filled Diplocraterion parallelum (Dp) represents an element of the Skolithos ichnofacies and
overprints a mottled background texture that reflects biogenic reworking during fair-weather,
Tern Formation.
300 K. L. BANN & C. R. FIELDING

Fig. 12. (a) Sedimentological section of a characteristic prograding shoreface succession, from the Permian
Tern Formation, Bonaparte Basin, Australia. The bioturbation intensity (BI) decreases upwards in relation to
coarsening and shallowing. Diminishing BI readings in the proximal lower shoreface deposits represent 'lam-
scram' bedding associated with the biogenic reworking of the upper portions of tempestites. The succession is
truncated by an erosion surface that hosts a firmground assemblage of trace fossils and represents the
Glossifungites ichnofacies. The trace fossil assemblages in the offshore to offshore transition consist of a very
diverse mixture of structures produced by grazing/foraging, detritus and deposit feeders. The suite reflects the
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 301

successions illustrated in Figures 11 and 12, sug- numbers of structures reflecting specialized feed-
gesting a similarity in depositional processes and ing/grazing behaviours. Shelf and offshore
bathymetry. However, when the ichnological deposits contain diverse expressions of the
characteristics are compared there are significant Zoophycos, distal Cruziana and archetypal
differences. Bioturbation intensity (BI, see Fig. Cruziana ichnofacies. In the more proximal
2b) in the Freitag Formation section (Fig. 14a) shoreface successions, diverse assemblages com-
is reduced, and uniformity of burrowing is posed of the robust burrows of deposit- and
sporadic. In addition, the trace fossil assemblage detritus-feeders are mixed with a diverse array
diversity is significantly reduced in comparison of sub vertical to vertical structures that represent
with the trace fossil suites in the bathymetrically the burrows and activities of carnivores, scaven-
equivalent (offshore and offshore transition) gers and opportunistic suspension-feeders. This
deposits in Figures 11 and 12. Figure 14b-d illus- complex assemblage represents a diverse proxi-
trates the characteristics of the deltaic facies. mal expression of the Cruziana ichnofacies
Figure 14b shows an example of a distal delta mixed with a significant proportion of elements
front interval from the Freitag Formation that from the Skolithos ichnofacies. This suite reflects
is pervasively bioturbated by Macaronichnus a well-oxygenated fully marine environment that
isp. in the basal portion of the photo and by Phy- evidently provided a plentiful supply of food
cosiphon in the upper part. An example of prox- material and stable conditions under which
imal delta front deposits from the upper Freitag mature infaunal communities were able to
Formation in GSQ Core Springsure-17 is illu- thrive, even in the more proximal environments
strated in Figure 14c. Once again bioturbation that show evidence of strong and repetitive
is sporadic, and the trace fossil assemblage is storm influence.
dominated by Macaronichnus isp. and Phycosi- In contrast, the ichnological signature of the
phon. The unusual spiral-like morphology seen Permian subaqueous delta deposits displays
in this (Macaronichnus isp.) example is typical low-diversity trace fossil assemblages (16 ichno-
of the morphology of Gyrolithes. This morphol- species overall) that are generally dominated by
ogy may reflect an escape mechanism to avoid one or two ichnotaxa. Bioturbation is sporadi-
burial and suffocation from emplacement of the cally distributed and significantly reduced in
overlying sandy tempestite. Clean sandstones overall intensity. The assemblage represents a
interpreted as mouth-bar deposits are dominated stressed expression of the Cruziana ichnofacies.
by Macaronichnus, and display a paucity of ver- Elements of the Skolithos ichnofacies are largely
tical structures (Fig. 14d). absent except for rare Diplocraterion habichi in
the proximal to distal delta front.
The ichnological differences between Permian
Discussion and conclusions prodelta and offshore deposits are illustrated in
Figure 15. In prodeltaic intervals (Fig. 15a)
Permian offshore and shoreface deposits from primary lamination generally dominates over
reservoirs in Australia contain ichnological sig- biogenic reworking. The trace fossil suite is rela-
natures characterized by moderate to intense tively diverse, but individual ichnospecies are
bioturbation, high assemblage diversities (25 reduced in size and are sporadically distributed
ichnogenera comprising 32 ichnospecies), uni- throughout the interval. Bioturbated horizons
form lateral (and to a lesser extant vertical) are usually dominated by Phycosiphon and
distribution of bioturbation, and significant Macaronichnus isp. The thin sandstone layers

activities of a mature fair-weather community of organisms with specialized feeding strategies, and represents a
diverse expression of the distal to archetypal Cruziana ichnofacies. Vertical structures that reflect the influence
of suspension-feeding organisms represent elements of the Skolithos ichnofacies and indicate opportunistic
colonization associated with storm deposition. These structures increase in abundance upwards through the
succession and dominate the trace fossil suite from the proximal lower shoreface through the middle shoreface
and up into the upper shoreface deposits, (b-f) Succession illustrated in (a) showing the characteristics of facies
upwards through the core from (b) upper offshore to (f) upper shoreface, and in particular the change in
morphology of Rosselia with increasing wave energy, (b) Permian Rosselia is generally diminutive, slender and
vertical to inclined in offshore deposits, (c, d) In offshore transition to distal lower shoreface deposits it is
generally truncated but very robust, and displays a variety of complex transitional forms, (e) In the proximal
lower shoreface to upper shoreface (and even into the foreshore in some localities) Permian Rosselia is long,
vertical, heavily lined but slender in nature as compared with specimens from more distal shoreface deposits,
(f) Clean, well-sorted upper shoreface deposits with rare Skolithos.
302 K. L. BANN & C. R. FIELDING

Fig. 13. Sedimentological section of an upward-coarsening and shallowing deltaic succession from the top of
the Sirius Mudstone Member (Cattle Creek Formation) and overlying Lower Aldebaran Sandstone, GSQ Core
Springsure -16. The contact between the two formations is interpreted as a sequence boundary. Bioturbation is
sporadic throughout this succession. The overall trace fossil assemblage is reduced in diversity, especially when
compared with the trace fossil suites in the sections illustrated in Figures 11 and 12, and represents a stressed
expression of the Cruziana ichnofacies. Elements of the Skolithos ichnofacies are conspicuous in their absence
from the sandier facies, where the trace fossil suite is of very low diversity and is dominated by the unusual
tubular burrows interpreted as Macaronichnus isp. (b) This portion of core (from 340m depth on the
sedimentological section) shows the characteristics of Permian distal delta front deposits. The facies contains
interlaminated and thinly interbedded siltstone and very fine- to fine-grained sandstones with low-angle
cross-bedding, and thin claystone beds. The trace fossil suite is dominated by Macaronichnus isp. (M) and
Phycosiphon (Ph). The thin claystone drapes are rarely bioturbated but here contain Chondrites (Ch).
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 303

Fig. 14. (a) Sedimentological section of a prograding deltaic succession from the lower Freitag Formation,
GSQ Core, Springsure -18. Bioturbation in this succession is sporadically distributed and reduced in diversity.
The suite is dominated by Phycosiphon and Macaronichnus isp. and contains rare Planolites, Chondrites,
Teichichnus and Zoophycos. The trace fossil suite in the proximal delta front is also dominated by
Macaronichnus isp. and lacks the vertical burrows of suspension-feeding organisms. The mouth-bar deposits
are unbioturbated. This succession is interpreted as a mixed river/wave-influenced delta package.
(b-d) Characteristics of coarsening-upwards deltaic facies from distal delta front to mouth bar deposits,
(b) An intense bioturbate texture from a distal delta front interval dominated by Macaronichnus isp. and
Phycosiphon (Ph). Freitag Formation, GSQ Core, Springsure -17. (c) A sparsely bioturbated proximal delta
front interval with Macaronichnus isp. and a structure that displays the characteristics of Gyrolithes. Upper
Freitag Formation, GSQ Core, Springsure -17. (d) Macaronichnus isp. in clean sandy mouth-bar deposits,
Freitag Formation, GSQ Core, Springsure -17.
304 K. L. BANN & C. R. FIELDING

Fig. 15. Graphic representation of the differences between prodeltaic and offshore deposits in core, (a)
Prodeltaic deposits contain interbedded siltstone and claystone with minor, interlaminated very fine- to fine-
grained sandstone layers with synaeresis cracks (sy) and soft-sediment deformation (ss). Bioturbation is
sporadically distributed, the assemblage diversity is reduced, and individual ichnospecies are reduced in size.
The trace fossil suit is characteristically dominated by Phycosiphon (Ph) and Macaronichnus isp. (M).
Asterosoma (A), Chondrites (Ch), Helminthopsis (H), Teichichnus (T) and Zoophycos (Z) occur as diminutive,
subordinate elements. Fugichia (f) are common. The trace fossil suite represents a stressed distal expression of
the Cruziana ichnofacies. In contrast, offshore deposits (b) are characterized by uniformly distributed intense
bioturbation comprising a very diverse mixture of structures produced by grazing/foraging, detritus- and
deposit-feeding activities. Ichnospecies include Phycosiphon (Ph), Rosselia (R), Rhizocorallium irregulare (Rh),
Planolites (P), Palaeophycus heberti (Pa), Asterosoma (A), Chondrites (Ch), Helminthopsis (H), Teichichnus (T),
Helminthoida (He), Macaronichnus isp. (M) and Zoophycos (Z). Thin sandstone beds contain remnant parallel
and low-angle lamination, and are associated with elements of the Skolithos ichnofacies such as Diplocraterion
habichi (Dh) and Palaeophycus tubularis (Pt). This assemblage represents fair-weather deposits containing a
very diverse expression of the archetypal Cruziana ichnofacies and minor influence from the Skolithos
ichnofacies associated with tempestites.
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 305

commonly contain soft-sediment deformation opportunistic colonization, particularly by sus-


structures (ss), synaeresis cracks (sy) and fugich- pension feeders (MacEachern 1994; Saunders
nia (f). In contrast, Figure 15b is a diagrammatic et al. 1994; Gingras et al. 1998). In addition,
representation of the characteristics of Permian high levels of organic detritus in the muds may
non-deltaic offshore deposits. Bioturbation have produced rapid oxidization and oxygen
intensity is high (BI5-6), it is uniformly distribu- depletion at the seafloor, which would poten-
ted, and the trace fossil suite is very diverse. Thin tially inhibit most organisms from colonizing
sandstone layers (tempestites) have been largely the substrate (Coates & MacEachern 1999,
reworked by biogenic activity. A high proportion 2000). The apparent absence of the vertical
of the facies has been completely homogenised burrows that would generally characterize an
by the repeated reworking during fair-weather opportunistic storm suite may also reflect high
by a diverse infaunal assemblage of grazing/ sedimentation rates, and increased levels of sedi-
foraging and specialized deposit-feeding ment reworking in the delta front environment
organisms. (Gingras et al. 1998). Internal erosion surfaces,
Ichnologically, shoreface deposits and delta lam-scram bedding and the truncation and re-
front deposits differ in significant ways (Fig. deposition of Rosselia mud balls as rip-up clasts
16). Lower shoreface deposits contain a diverse provide evidence that storm induced exhumation
fair-weather assemblage (Fig. 16a, b) that repre- produced the amalgamation of units and the
sents a proximal expression of the Cruziana possible removal of previously established
ichnofacies, and clearly reflects an environment infaunal communities.
with plentiful food supplies and long periods of Stressed, low-diversity trace fossil assemblages
stable fully marine conditions. Tempestites in such as recorded from the Permian deltaic depos-
the lower shoreface (Fig. 16c) contain a relatively its reflect environments that do not provide opti-
diverse assemblage of vertical traces that repre- mum living conditions for infaunal organisms.
sents a distal expression of the Skolithos ichno- Hence most of the inhabitants are opportunistic
facies, suggesting that a variety of organisms species that flourish periodically in environments
were able to colonize the substrate even during that are, at other times, too inhospitable for even
periods of repeated storm influence. In contrast, the most resilient of trophic generalists.
trace fossil assemblages in distal delta front In general these Permian deltaic facies are
deposits (Fig. 16d-e) reflect the activities of a characterized by physical structures and ichnolo-
stressed infaunal community only rarely inhab- gical signatures that indicate rapid sedimentation
ited by diminutive grazing/foraging organisms rates, high degrees of salinity variation, a mixture
and deposit-feeders and dominated by opportu- of current and wave processes, and large quanti-
nistic deposit-feeding organisms. The vertical ties of organic detritus. This evidence includes:
burrows of opportunistic suspension-feeders are
the low-diversity, sporadically distributed trace
very rarely present.
fossil assemblage that represents a stressed
The overall paucity of evidence of infaunal
expression of the Cruziana ichnofacies;
suspension-feeders in the deltaic deposits may
common fugichnia but a noticeable paucity of
reflect high levels of water turbidity, caused by
vertical burrows, even in the more proximal
river discharge into the basin. High levels of
facies where opportunistic suspension-feeders
suspended sediment within the water column
would ordinarily be expected;
produce unfavourable conditions for suspen-
thick coarsening-upward facies successions;
sion-feeding organisms by interfering with their
thick organic-rich prodelta siltstone intervals;
filter-feeding apparatus and reducing the percen-
common soft-sediment deformation struc-
tage of food per unit of material ingested
tures;
(Moslow & Pemberton 1988; Buatois & Angri-
abundant organic detritus and siderite
man 1992; Gingras et al. 1998; Coates & Mac-
cements;
Eachern 1999). Large quantities of suspended
organic claystone drapes on tempestites with
fine sediment and organic detritus are associated
common synaeresis cracks; and
with heightened precipitation, increased surface
abundant wave-generated sedimentary struc-
runoff at the coast, and elevated fluvial discharge
tures.
in deltaic settings. Freshwater input and thermal
variations during increased fluvial discharge Such evidence strongly supports the interpreta-
would be additional environmental stresses in tion of this deltaic facies association as that of
the deltaic environment (cf. also Mcllroy 2004). prograding, mixed river- and wave-influenced
The thin claystone drapes on tempestites sug- delta lobes, and clearly differentiates this
gest that the storm beds were rapidly covered facies from non-deltaic offshore and shoreface
with organic-rich mud, shielding them from deposits.
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 307

Funding for this project was provided by the University BANN, K. L., FIELDING, C. R., MAC£ACHERN, J. A. &
of Queensland in the form of a Post-Doctoral Research TYE, S. C. 2004. Differentiation of estuarine and
Fellowship to KLB. Oil Company of Australia Ltd and offshore marine deposits using integrated and sedi-
Santos Ltd are also gratefully acknowledged for their mentology: Permian Pebbley Beach Formation,
generous funding of this research. Staff from the Sydney Basin, Australia. In: MC!LROY, D. (ed.)
Department of Mines, Zillmere, are thanked for The Application of Ichnology to Palaeoenvironmen-
laying out of core. M. Wilkinson, J. Trueman and T. tal and Stratigraphic Analysis. Geological Society,
Grimison are thanked for field assistance. Santos Ltd London, Special Publications, 228, 179-211.
is also thanked for granting permission to publish BHATTACHARYA, J. & WALKER, R. G. 1992. Deltas. In:
material from the Tern Formation. J. Lerette is WALKER, R. G. & JAMES, N. P. (eds) Fades
thanked for useful editorial comments. M. Gregory Models, Response to Sea Level Change. Geological
and D. Mcllroy are thanked for their helpful reviews. Association of Canada, St. John's, Newfound-
land, 157-158.
BOYD, R., DALRYMPLE, R. & ZAITLIN, B. A. 1992. Clas-
References sification of clastic coastal depositional environ-
ments. Sedimentary Geology, 80, 139-150.
ALLEN, G. P. & CHAMBERS, J. L. C. 1998. Sedimentation BRETTLE, M. J., MC!LROY, D., DAVIES, S. J., ELLIOTT,
in the Modern and Miocene Mahakam Delta. Indo- T. & WATERS, C. N. 2002. Identifying cryptic
nesian Petroleum Association, Jakarta, Indonesia. tidal influences within deltaic successions: an
BAKER, J. C. 1991. Diagenesis and reservoir quality of example from the Marsdenian (Namurian) inter-
the Aldebaran Sandstone, Denison Trough, east- val of the Pennine Basin, UK. Journal of the
central Queensland, Australia. Sedimentology, 38, Geological Society, London, 159, 379-391.
819-838. BROMLEY, R. G. 1996. Trace Fossils: Biology, Taphon-
BANN, K. L. 1998. Ichnology and sequence stratigraphy omy and Applications (2nd edn). Chapman &
of the Early Permian Pebbley Beach Formation and Hall, London.
Snapper Point Formation in the southern Sydney BUATOIS, L. A. & ANGRIMAN, O. L. 1992. The ichnol-
Basin. PhD thesis, University of Wollongong. ogy of a submarine braided channel complex: the
BANN, K. L. & FIELDING, C. R. 2001. Applications of Whiskey Bay Formation of James Ross Island,
ichnology to hydrocarbon exploration: examples Antarctica. Palaeogeography, Palaeoclimatology,
from the Permian of eastern Australia. In: HILL, Palaeoecology, 94, 119-140.
K. C. & BERNECKER, T. (eds) Eastern Australasian CARR, P. F, JONES, B. G. & MIDDLETON, R. G. 1989.
Basins Symposium 2001, Petroleum Exploration Precursor and formation of glendonites in the
Society of Australia Special Publication, Aust- Sydney Basin. Australian Mineralogist, 4, 3-12.
IMM, Melbourne, 603-612. COATES, L. & MACEACHERN, J. A. 1999. The ichno-
BANN, K. L., KLOSS, O., WOOD, G., LANG, S., KASSAN, logical signature of wave- and river-dominated
J. & BENSON, J. 2004. Palaeoenvironments and deltas: Dun vegan and basal Belly River Forma-
depositional history of the Tern Field, Bonaparte tions, West-Central Alberta. In: WRATHALL, B.,
Basin. In: ELLIS, G. & GORTERS, J. (eds) Timor JOHNSTON, G., ARTS, A., Rozsw, L, ZONNEVELD,
Sea Petroleum Geoscience, Bonaparte Basin & J-P., ARCURI, D. & MCLELLAN, S. (eds) Digging
Surrounds. Petroleum Exploration Society of Deeper, Finding a Better Bottom Line. CSPG
Australia, Special Publication, Perth, Australia, and Petroleum Society 1999 Core Conference
in press. Paper, Calgary, Alberta, p. 99-114.

Fig. 16. Comparison of the ichnological signatures of Permian lower and middle shoreface deposits with delta
front deposits, (a) Graphic representation of ichnospecies abundance in lower shoreface, distal delta front,
middle shoreface and proximal delta front deposits, (b) Tiering profile of the fair-weather assemblage for
Permian lower shoreface deposits. The suite reflects a very diverse mixture of grazing/foraging structures,
detritus-feeding and complex deposit-feeding structures, and represents a diverse proximal expression of the
Cruziana ichnofacies. (c) Tiering profile for event beds in Permian lower shoreface deposits, reflecting a
relatively diverse mixture of structures produced by opportunistic suspension-feeders and robust detritus-
feeders, which represents a distal expression of the Skolithos ichnofacies. (d) Tiering profile representing the
trace fossil signature from the most environmentally stable portions of distal delta front deposits. The trace
fossil suite contains a relatively diverse, but diminutive and sporadically distributed, assemblage of structures
produced by grazing/foraging and deposit-feeding behaviours. All of the ichnospecies illustrated are rare but
tend to occur in association with each other, and are interpreted as representing the least stressed component of
the deltaic infaunal assemblage. The assemblage represents a stressed expression of the Cruziana ichnofacies
and contrasts markedly with the fair-weather assemblage from the non-deltaic lower shoreface deposits,
(e) Representation of the tiering profile of distal delta front deposits that were colonized only by trophic
generalists. The assemblage is dominated by Macaronichnus isp. and Phycosiphon. All other vertical structures
are absent except for rare examples of small Diplocraterion habichi. This trace fossil suite comprises a low-
diversity mixture of opportunistic ichnospecies, and represents a stressed proximal expression of the Cruziana
ichnofacies.
308 K. L. BANN & C. R. FIELDING

COATES, L. & MACEACHERN, J. A. 2000. Integrating ich- FIELDING, C. R., KASSAN, J. & DRAPER, J. J. 1996.
nology and sedimentology to differentiate between Geology of the Bowen and Surat Basins, eastern
river-dominated deltas, wave-dominated deltas, Queensland. Australasian Sedimentologists Group
and shorefaces: examples from the Cretaceous of Field Guide Series, Geological Society of Austra-
Western Canada. Geological Society of America, lia, Sydney, 8, 1-126.
Cordillleran Section, 96th Annual Meeting. Van- FIELDING, C. R., SLIWA, R., HOLCOMBE, R. J. &
couver, British Columbia, 32, p. A7. KASSAN, J. 2000. A new palaeogeographic synth-
COLEMAN, J. M. 1981. Deltas: Processes of Deposition esis of the Bowen Basin of central Queensland.
and Models for Exploration. Burgress Publishing In: BEESTON, J. (ed.) Bowen Basin Symposium
Company, Minneapolis, MN. 2000, 22-24 October 2000, Geological Society of
COLEMAN, J. M. & WRIGHT, L. D. 1975. Modern river Australia Coal Geology Group & Bowen Basin
deltas: variability of processes and sand bodies. Geologists Group, Brisbane, 287-302.
In: BROUSSARD, M. H. (ed.) Deltas: Models FIELDING, C. R., SLIWA, R., HOLCOMBE, R. J. &
for Exploration. Houston Geological Society, 99- JONES, A. T. 2001. A new palaeogeographic
149. synthesis for the Bowen, Gunnedah and Sydney
CROWELL, J. C. & FRAKES, L. A. 1971. Late Palaeozoic Basins of eastern Australia. In: HILL, K. C. &
glaciation of Australia. Journal of the Geological BERNECKER, T. (eds) Eastern Australasian Basins
Society of Australia, 17, 115-155. Symposium, Petroleum Exploration Society of
CROWELL, J. C. & FRAKES, L. A. 1975. The Late Australia, Special Publications, Australasian
Palaeozoic Glaciation. In: CAMPBELL K. S. W. Institute of Mining & Metallurgy, Melbourne,
(ed.) Gondwana Geology. Australian National 269-278.
University Press, Canberra, 313-331. FREY, R. W. 1990. Trace fossils and hummocky cross-
CURRAY, J. R., EMMAL, F. J. & CRAMPTON, P. J. S. stratification, Upper Cretaceous of Utah. Palaios,
1969. Holocene history of a strand plain, lagoonal 5, 203-218.
coast, Nayarit, Mexico. In: CASTANARES, A. A. & FREY, R. W. & COLORING, R. 1992. Marine event beds
PHLEGER, F. B. (eds) Coastal Lagoons: A Sympo- and recolonization surfaces as revealed by trace
sium. Universidad Nacional Autonoma, Mexico, fossil analysis. Geological Magazine, 129, 325-335.
63-100. FREY, R. W. & PEMBERTON, S. G. 1984. Trace fossils
DALRYMPLE, R. W. 1999. Tide-dominated deltas: do facies models. In: WALKER, R. G. (ed.) Facies
they exist or are they all estuaries? American Asso- Models, 2nd edition. Geoscience Canada (Reprint
ciation of Petroleum Geologists Annual Meeting, Series, 1), Toronto, Ontario, 189-287.
San Antonio, Texas, A29-30. FREY, R. W. & PEMBERTON, S. G. 1985. Biogenic struc-
ELLIOT, T. 1986. Deltas: In: READING, H. G. (ed.) tures in outcrops and cores: 1. Approaches to ich-
Sedimentary Environments and Fades. Blackwell nology. Bulletin of Canadian Petroleum Geology,
Scientific Publications, Oxford, 113-154. 33,72-115.
FALKNER, A. J. & FIELDING, C. R. 1993. Geometrical GINGRAS, M. K., MACEACHERN, J. A. & PEMBERTON,
facies analysis of a mixed-influence deltaic S. G. 1998. A comparative analysis of the ich-
system: the Late Permian German Creek Forma- nology of wave- and river-dominated allomembers
tion, Bowen Basin, Australia. In: MARZO, M. & of the Upper Cretaceous Dun vegan Formation.
PUIGDEFABREGAS, C. (eds) Alluvial Sedimentation. Bulletin of Canadian Petroleum Geology, 46, 51-
International Association of Sedimentologists, 73.
Special Publication, Oxford, 17, 195-209. HOLCOMBE, R. J., STEPHENS, C. J. et al. 1997. Tectonic
FIELDING, C. R. 1989. A tide- and wave-moulded shelf evolution of the Northern New England Fold Belt:
sequence from the Permian of the south-west the Permian-Triassic Hunter-Bowen event. In:
Bowen Basin, Queensland, Australia. Australian ASHLEY, P. A. & FLOOD, P. G. (eds) Tectonics
Journal of Earth Sciences, 36, 29-40. and Metallogenesis of the New England Orogen,
FIELDING, C. R. & LANG, S. C. 1988. A facies analysis Geological Society of Australia, Special Publica-
of the Staircase Sandstone Member (Early tion, Sydney, 19, 52-65.
Permian) in the Springsure area, southwestern HOWARD, J. D. 1971. Comparison of the beach-to-off-
Bowen Basin. Geological Society of Australia shore sequence in modern and ancient sediments.
Abstracts Series, 21, 137-138. In: HOWARD, J. D., VALENTINE, J. W. & WARME,
FIELDING, C. R. & MCLOUGHLIN, S. 1992. Sedimentol- J. E. (eds) Recent Advances in Paleoecology and
ogy and palynostratigraphy of Permian rocks Ichnology. American Geological Institute, Short
exposed at Fairbairn Dam, central Queensland, Course Lecture Notes, Alexandria, Virginia,
Australia. Australian Journal of Earth Sciences, 148-183.
39, 631-649. HOWARD, J. D. 1972. Trace fossils as a criteria for
FIELDING, C. R., STEPHENS, C. J., KASSAN, J. & recognizing shorelines in the stratigraphic record.
HOLCOMBE, R. J. 1995. Revised palaeogeographic Society of Economic Paleontologists and Mineralo-
maps for the Bowen Basin, central Queensland. gists, Special Publications, Tulsa, Oklahoma, 16,
In: FOLLINGTON, I. L., BEESTON, J. W. & HAMIL- 215-225.
TON, L. H. (eds) Bowen Basin Symposium 1995 HOWARD, J. D. & FREY, R. W. 1984. Characteristic
Proceedings, Mackay, Qld, 1-3 October, 1995, trace fossils in nearshore to offshore sequences,
Geological Society of Australia Coal Geology Upper Cretaceous of east-central Utah. Canadian
Group, 7-15. Journal of Earth Sciences, 21, 200-219.
ICHNOLOGY OF SHOREFACE VS DELTAIC FACIES 309

HOWARD, J. D. & REINECK, H.-E. 1981. Depositional front sandstone sequences. Canadian Society of
fades of a high energy beach-to-offshore sequence: Petroleum Geologists Memoir, 15, 373-386.
comparison with low-energy sequence. American PEMBERTON, S. G. & FREY, R. W. 1984. Ichnology
Association of Petroleum Geologists Bulletin, 65, of storm-influenced shallow marine sequence:
807-830. Cardium Formation (Upper Cretaceous) at
JANSEN, J. H. F., WOENSDREGT, C. F., KOOIESTRA, M. J. Seebe, Alberta. In: STOTT, D. F. & GLASS, D. J.
& VAN DER GAAST, S. J. 1987. Ikaite pseudo- (eds) The Mesozoic of Middle North America.
morphs in Zaire Deep Sea Fan: intermediate Canadian Society of Petroleum Geologists
between calcite and porous calcite. Geology, 15, Memoir, Calgary, Alberta, 9, 281-304.
245-248. PEMBERTON, S. G. & MACEACHERN, J. A. 1995. The
JOHN, B. H. & FIELDING, C. R. 1993. Reservoir poten- sequence stratigraphic significance of trace fossils:
tial of the Catherine Sandstone, Denison Trough, examples from the Cretaceous foreland basin of
east-central Queensland. Australian. Petroleum Alberta, Canada. In: VAN WAGONER, J. C. & BER-
Exploration Association Journal, 33, 176-187. TRAM, G. (eds) Sequence Stratigraphy of Foreland
JOHNSON, H. D. & BALDWIN, C. T. 1996. Shallow clastic Basin Deposits: Outcrop and Subsurface Examples
seas. In: READING, H. G. (ed.) Sedimentary Environ- from the Cretaceous of North America. American
ments: Processes, Fades and Stratigraphy. Black- Association of Petroleum Geologists, Memoirs,
well Scientific Publications, Oxford, 232-280. Tulsa, Oklahoma, 64, 429^75.
KAPLAN, M. E. 1979. Calcite pseudomorphs (pseudo- PEMBERTON, S. G. & MACEACHERN, J. A. 1997. The
gaylussite, jarowite, thinolite, glendonite, genno- ichnological signature of storm deposits: the use
ishi, White Sea Hornlets) in sedimentary rocks: of trace fossils in event stratigraphy. In: BRETT,
origins of the pseudomorphs. Lithology and C. E. (ed.) Paleontological Event Horizons: Eco-
Mineral Resources, 14, 623-636. logical and Evolutionary Implications. Columbia
KENNEDY, G. L., HOPKINS, D. M. & PICKTHORN, W. J. University Press, New York, 73-109.
1987. Ikaite, the glendonite precursor, in estuarine PEMBERTON, S. G. MACEACHERN, J. A. & FREY, R. W.
sediments at Barrow, Artie, Alaska. Geological 1992. Trace fossil facies models environmental and
Society of America, Abstracts with Programmes, allostratigraphic significance. In: WALKER, R. G.
19, 725. & JAMES, N. P. (eds) Facies Models: Response to
MACEACHERN, J. A. 1994. Integrated ichnological-sedi- Sea Level Change. Geological Association of
mentological models: applications to the sequence Canada, St John's, Newfoundland, 47-72.
stratigraphic and palaeoenvironmental interpreta- PEMBERTON, S. G., SPILA, M., PULHAM, A. J., SAUN-
tion of the Viking and Peace River Formations, DERS, T., MACEACHERN, J. A., ROBBINS, D. &
West-Central Alberta. PhD thesis. University of SINCLAIR, I. 2001. Ichnology and Sedimentology
Alberta. of Shallow to Marginal Marine Systems. Ben
MACEACHERN, J. A. 2001. Ichnology and sedimentol- Nevis and Avalon Reservoir, Jeanne d'Arc Basin.
ogy in a sequence stratigraphic framework: inte- Geological Association of Canada, Short Course
grated facies models for subsurface analysis. Notes, St John's, Newfoundland, 15.
Rocky Mountain Association of Geologists, Short READING, H. G. (ed.) 1986. Sedimentary Environments
Courses, Denver, Colorado, 1. and Facies. Blackwell Scientific Publications,
MACEACHERN, J. A. & PEMBERTON, S. G. 1992. Ichno- Oxford, 113-154.
logical aspects of Cretaceous shoreface successions REINECK, H.-E. 1963. Sedimentgefuge im Bereich der
and shoreface variability in the Western Interior sudlichen Nordsee. Abhandlungen der Senckenber-
Seaway of North America. In: PEMBERTON, S. G. gische Naturforschende Gesellschaft, 505, 138p.
(ed.) Applications of Ichnology to Petroleum REINSON, G. E. 1984. Barrier-island and associated
Exploration: A Core Workshop. Society of Eco- strand plain systems. In: WALKER, R. G. (ed.),
nomic Paleontologists and Mineralogists, Core Facies Models (2nd edn). Geological Association
Workshops, Tulsa, Oklahoma, 17, 57-84. of Canada, Geoscience Canada Reprint Series,
MclLROY, D. 2004. Ichnology and facies model of a Kitchener, Ontario, 1, 119-140.
tide-dominated delta: Jurassic upper Ror and He SAUNDERS, T. & PEMBERTON, S. G. 1986. Trace Fossils
Formations of Kristin Field, Halten Terrace, Off- and Sedimentology of the Appaloosa Sandstone:
shore Mid-Norway. In: MclLROY, D. (ed.) The Bearpaw-Horseshoe Canyon Formation Transition,
Application of Ichnology to Palaeoenvironmental Dorothy, Alberta. Canadian Society of Petroleum
and Stratigraphic Analysis. Geological Society, Geologists Fieldtrip Guide Book, Calgary,
London, Special Publications, 228, 237-272. Alberta.
McLACHLAN, A., COCKCROFT, A. C. & MALAN, D. E. SAUNDERS, T. D. A., MACEACHERN, J. A. & PEMBER-
1984. Benthic faunal response to a high energy TON, S. G. 1994. Cadotte Member Sandstone: pro-
gradient. Marine Ecology Progress Series, 16, gradation in a boreal basin prone to winter storms.
51-63. In: PEMBERTON, S. G., JAMES, D. P. & WIGHTMAN,
MCLOUGHLIN, S. 1993. Plant fossil distributions in M. (eds) Mannville Core Conference, Canadian
some Australian Permian non-marine sediments. Society of Petroleum Geologists, Calgary,
Sedimentary Geology, 85, 601-619. Alberta, 331-349.
MOSLOW, T. F. & PEMBERTON, S. G. 1988. An inte- SEILACHER, A. 1955. Ecological significance of fossil
grated approach to the sedimentological analysis tracks and trails. Geological Society of America
of some Lower Cretaceous shoreface and delta Bulletin, 66, 1663.
310 K. L. BANK & C. R. FIELDING

SEILACHER, A. 1964. Biogenic sedimentary structures. TRUEMAN, J. D. 2002. Stratigraphy and sedimentology
In: IMBRIE, J. & NEWELL, N. (eds). Approaches to of the Burdekin Delta, Queensland and comparisons
Paleoecology. Wiley, New York, 296-316. with Permian coastal fades in the Denison Trough,
SEILACHER, A. 1967. Bathymetry of trace fossils. SW Bowen Basin, Australia. Unpublished PhD
Marine Geology, 5, 189-200. thesis, University of Queensland, Brisbane.
SEILACHER, A. 1978. Use of trace fossils for recognizing VEEVERS, J. J. & POWELL, C. McA. 1987. Late glacial
depositional environments. In: BASAN, P. (ed.) episodes in Gondwanaland reflected in trans-
Trace Fossil Concepts. Society of Economic gressive-regressive depositional sequences in Eura-
Palaeontologists and Mineralogists, Short merica. Geological Society of America Bulletin, 98,
Courses, Tulsa, Oklahoma, 5, 185-201. 475^87.
SHEARMAN, D. J. & SMITH, A. M. 1985. Ikiate, the VOSSLER, S. M. & PEMBERTON, S. G. 1989. Ichnology
parent material of jarrowite-type pseudomorphs. and paleoecology of offshore siliciclastic deposits
Proceedings of the Geologists' Association, 96, in the Cardium Formation (Turonian, Alberta,
421-452. Canada). Palaeogeography, Palaeoclimatology,
SUESS, E., BALZER, W., HESSE, K. F., MULLER, P. J., Palaeoecology, 74, 217-229.
UNGERER, C. A. & WEFER, G. 1982. Calcium car- WALKER, R. G. & FLINT, A. G. 1992. Wave- and storm-
bonate hexahydrate from organic-rich sediments dominated shallow marine systems. In: WALKER,
of the Antarctic shelf: precursors of glendonites. R. G. & JAMES, N. P. (eds). Fades Models
Science, 216, 1128-1130. Response to Sea Level Change. Geological Asso-
TAYLOR, A. M. & COLORING, R. 1993. Description ciation of Canada, St. John's, Newfoundland,
and analysis of bioturbation and ichnofabric. 219-239.
Journal of the Geological Society, London, 150, WHEATCROFT, R. A. 1990. Preservation potential of
141-148. sedimentary event layers. Geology, 18, 843-845.
Animal-substrate interactions in freshwater environments:
applications of ichnology in facies and sequence stratigraphic
analysis of fluvio-lacustrine successions
LUIS A. BUATOIS & M. GABRIELA MANGANO

Conicet-Insugeo, Casilla de correo 1, correo central, 4000 San Miguel de Tucumdn, Argentina
(e-mail: ichnolog@infovia.com.ar)

Abstract: At present, three continental archetypal ichnofacies are widely accepted: the
Scoyenia, Mermia and Coprinisphaera ichnofacies. The last is present in palaeosols, and
the first two occur in fluvio-lacustrine environments. Additionally, the Skolithos ichnofacies
may be present in relatively high-energy fluvio-lacustrine deposits. The ichnofauna from
active fluvial channels is characterised by low-diversity assemblages of simple vertical
burrows and escape traces, referred to the Skolithos ichnofacies. Abandoned or inactive
channel deposits characteristically contain low-diversity assemblages dominated by menis-
cate traces. Floodplain water bodies that experienced progressive drying (desiccated over-
bank deposits) may contain abundant arthropod and vertebrate trackways, backfilled
meniscate traces, ornamented burrows and bilobate traces with scratch marks, which allow
recognition of the Scoyenia ichnofacies. Floodplain water bodies that are filled by overbank
vertical accretion without experiencing desiccation (overfilled overbank deposits) include
simple grazing trails, locomotion trails and horizontal dwelling burrows, representing impo-
verished occurrences of the Mermia ichnofacies. Hydrologically closed lakes are very stress-
ful environments in which subaqueous ichnofaunas are rare. The richest ichnofaunas in
closed lakes are present at the lake margins, and record the activity of terrestrial rather
than aquatic faunas (Scoyenia ichnofacies). Hydrologically open lakes host relatively diverse
and abundant ichnofaunas, comprising the Scoyenia ichnofacies in low-energy, lake-margin
areas, and the Mermia ichnofacies in permanent subaqueous lacustrine zones. Sediments
deposited in relatively high-energy lacustrine environments, such as wave-dominated shore-
lines and delta mouth-bars, commonly are represented by the Skolithos ichnofacies.
Although continental trace fossils have not been extensively used in sequence stratigraphy,
they have potential for future integrated study. Softground trace fossils are commonly well
developed in overfilled lake basins and are useful to delineate parasequences and para-
sequence sets. In balanced-fill and underfilled lake basins, softground ichnofaunas are
poorly developed because of stressful conditions. In contrast, firmground suites are rare in
overfilled lake basins, but widespread in lowstand deposits of balanced-fill and underfilled
lake basins. Early lowstand amalgamated incised fluvial channels are usually unbioturbated,
but palaeosol ichnofaunas (e.g. the Coprinisphaera ichnofacies) may delineate sequence
boundaries in interfluve areas. Increasingly isolated fluvial channels encased in overbank
deposits develop during late lowstand, and lacustrine deposits accumulate during transgres-
sions, favouring preservation of biogenic structures.

The study of continental ichnofaunas has shown Archer 1989; Bromley & Asgaard 1991; Buatois
an explosive development during the last decade. & Mangano 1995a; Labandeira 1997, 1998,
This is revealed not only through careful docu- 2002; Buatois et al 1998; Genise et al 2000;
mentation of individual cases integrating ichno- Miller & Labandeira 2003).
logical, sedimentological and palaeobiological This paper focuses on the ichnology of fluvio-
data (e.g. Gierlowski-Kordesch 1991; Pickerill lacustrine depositional systems. Our emphasis
1992; Buatois & Mangano 1993; Genise & will be on the aquatic trace fossil assemblages
Bown 1994a, 1994b; MacNaughton & Pickerill of continental environments; fully terrestrial
1995; Metz 1995; Buatois et al. 1996a, 1997; ichnofaunas preserved in palaeosols are reviewed
Mangano et al. 1997, 2001), but also through by Genise et al. (2004). The objectives of this
renewed attempts to provide a conceptual back- paper are:
ground to the analysis of organism-substrate
interactions in non-marine successions b y t o review present knowledge of freshwater
means of definition of archetypal i c h n o f a c i e s i c h n o f a c i e s
and the evaluation of temporal and spatial to evaluate the utility of biogenic structures in
trends in trace fossil distribution (e.g. Maples & facies analysis by providing a summary of the

From: MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 311-333. 0305-8719/04/$ 15.00 © The Geological Society
of London.
312 L. A. BUATOIS & M. G. MANGANO

ichnology of fluvio-lacustrine environments; meniscate burrows, arthropod trackways and


and bilobed traces, as well as several physical sedi-
to explore the potential uses of trace fossils in mentary structures (e.g. desiccation cracks).
sequence stratigraphic analysis of continental Frey et al. (1984) noted that the Scoyenia ichno-
successions. facies subsequently was used as a catchall for all
assemblages of continental trace fossils. As noted
by these authors, the Scoyenia ichnofacies has
The ichnofacies model: its expansion into precise environmental implications because it
continental environments characterizes low-energy continental deposits
periodically exposed to air or inundated, inter-
Archetypal or Seilacherian ichnofacies are trace mediate between aquatic and non-aquatic (see
fossil assemblages that recur through long inter- also Frey & Pemberton 1984, 1987). The fact
vals of geologic time and are characteristic of a that the Scoyenia ichnofacies was only one of
given set of environmental conditions (Frey & the recurrent trace fossil assemblages of contin-
Pemberton 1984, 1985). The ichnofacies model, ental environments, and that continental envir-
originally put forward by Seilacher (1967), has onments are as diverse as marine settings, has
been expanded and refined in a series of papers. long been acknowledged by ichnologists. How-
Modifications include: definition of additional ever, it was not until recently that studies addres-
marine ichnofacies, such as the Psilonichnus sing the problem of recognizing additional
and Trypanites ichnofacies (Frey & Seilacher continental ichnofacies were published (e.g.
1980; Frey & Pemberton 1987); refinements in Smith et al. 1993; Buatois & Mangano 1995a;
the characterisation of previously defined ichno- Bromley 1996; Genise et al. 2000). At present,
facies, such as the Glossifungites ichnofacies three continental archetypal ichnofacies are
(Pemberton & Frey 1985; MacEachern et al. widely accepted: the Scoyenia, Mermia and
1992); further analysis of the hardground ichno- Coprinisphaera ichnofacies (Fig. 1; see review
facies (Bromley & Asgaard 1993; de Gibert et al, in Mcllroy et al. 2004). The last is present in
1998); and expansion of the model to cover con- palaeosols and is analysed in detail by Genise
tinental environments (Smith et al. 1993; Buatois et al. (2004); only the first two occur in fluvio-
& Mangano 1995a; Genise et al. 2000). In his lacustrine environments, and are therefore dis-
original model, Seilacher (1967) recognised only cussed herein. Bromley (1996) also tentatively
one ichnofacies for continental environments, proposed the Rusophycus ichnofacies for fluvial
the Scoyenia ichnofacies. He proposed the to shallow-lacustrine environments and the
Scoyenia ichnofacies for 'non-marine sands and Fuersichnus ichnofacies for lacustrine settings
shales, often red beds, with a distinctive below the fair-weather wavebase. The Ruso-
association of trace fossils' and referred to a phycus ichnofacies is dominated by locomotion
previous schematic illustration of this ichno- and resting traces of arthropods, and cannot be
fauna (Seilacher 1963, fig. 7), which included distinguished at present from the Scoyenia

Fig. 1. Ichnofacies model of continental environments.


NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 313

ichnofacies. The Fuersichnus ichnofacies is based localized high abundance.


on examples in which the eponymous ichnogenus
Among the meniscate traces, Scoyenia, Beaco-
occurs in non-marine settings. However, the
nites and Taenidium are typical components.
'type' examples suggested are from fluvial
Arthropod trackways are represented by a wide
deposits (MacNaughton & Pickerill 1995) and
variety of ichnotaxa, including Umfolozia, Mer-
ephemeral alluvial plain/sandflat facies (Gier-
ostomichnites, Diplichnites, Hexapodichnus, Per-
lowski-Kordesch 1991) rather than relatively
michnium and Acripes. Other horizontal traces
deep lacustrine settings. As currently defined,
include bilobate locomotion (Cruziana) and
the Fuersichnus ichnofacies cannot be distin-
resting structures (Rusophycus), simple forms
guished from the Scoyenia ichnofacies. High-
(Pianolites and Palaeophycus), sinuous crawling
energy continental environments, such as
traces (Cochlichnus) and banana-shaped traces
lacustrine wave-dominated shorelines, fluvial (Fuersichnus). Vertical burrows are represented
channels and delta mouth-bars, commonly
by Skolithos and Macanopsis. Short vertical bur-
contain simple vertical burrows (Skolithos), U-
rows are commonly referred to as Cylindricum
shaped vertical burrows (Arenicolites) and
(e.g. Pollard 1981). Crayfish burrows, included
escape structures (e.g. Bromley & Asgaard
in the ichnogenus Camborygma, are common
1979; Mangano et al. 1994). Such assemblages,
components of post-Palaeozoic examples (Wells
included by Bromley & Asgaard (1991) in the
1977; Hasiotis et al. 1993). Vertebrate tracks
Arenicolites ichnofacies, are hardly distinctive
may be abundant, including those produced by
from the marine Skolithos ichnofacies, so it is
amphibians (e.g. Limnopus, Anthichniuni), rep-
best to consider them as non-marine occurrences
tiles (e.g. Chirotherium, Rhynchosauroides)
of the Skolithos ichnofacies (Buatois & Mangano
including Dinosaur (e.g. Grallator, Eubrontes),
1995a, 1998).
mammals (e.g. Mylodontidichnum, Neomagather-
The Scoyenia ichnofacies (Fig. 2), as redefined
ichnum) and birds (e.g. Jindongornipes, Korea-
by Frey et al. (1984) and Buatois & Mangano
nornis) (Gand & Haubold 1984; Aramayo &
(1995a), is characterised by:
Bianco 1987a, 1987b, 1996; Fuglewicz et al.
horizontal meniscate backfilled traces pro- 1996; Lockley 1991). Although the Scoyenia
duced by mobile deposit feeders; ichnofacies is commonly represented by low-
locomotion traces, including both trackways diversity ichnocoenoses of meniscate trace fossils
and trails; (Frey et al. 1984), Palaeozoic examples usually
vertical domiciles; consist of moderate diverse assemblages of
a mixture of invertebrate (mostly arthropod), arthropod trackways (Buatois & Mangano
vertebrate and plant traces; 1995a). A narrower definition was recently pro-
low to moderate ichnodiversity; and posed by Bromley (1996), who considered the

Fig. 2. Schematic reconstruction of the Scoyenia ichnofacies. Based on Buatois et al. (2002).
314 L. A. BUATOIS & M. G. MANGANO

Fig. 3. Schematic reconstruction of the Mermia ichnofacies. Based on Buatois et al. (2002).

Scoyenia ichnofacies as a continental equivalent The Mermia ichnofacies (Fig. 3), proposed by
of the firmground Glossifungites ichnofacies of Buatois & Mangano (1995a), is characterised by:
the marine realm. The Scoyenia ichnofacies
dominance of horizontal to subhorizontal
may be subdivided into two distinct suites: one
grazing and feeding traces produced by
characterised by meniscate, backfilled structures
mobile deposit feeders;
without ornamentation (e.g. Taenidium, Beaco-
subordinate occurrence of locomotion traces;
nites) developed in a soft substrate, and the
relatively high to moderate ichnodiversity and
other typified by striated traces (e.g. Scoyenia,
abundance; and
Spongeliomorpha), cross-cutting the former and
low degree of specialisation of grazing pat-
developed in a firm substrate (Buatois et al.
terns.
1996b; Buatois & Mangano 2002). The resulting
palimpsest surfaces or composite ichnofabrics Typical components of this ichnofacies include a
reflect progressive desiccation of sediment. This variety of unspecialised grazing traces (e.g.
view was recently adopted by Savrda et al. Mermia, Gordia, Helminthopsis, Helminthoidich-
(2000), who documented an assemblage of back- nites and Cochlichnus), simple feeding structures
filled trace fossils without ornamentation (Taeni- (e.g. Treptichnus and Circulichnis\ locomotion
dium) in soft substrates of a fluvial environment. traces (e.g. Maculichna) and fish trails (e.g.
The Scoyenia ichnofacies occurs in low-energy Undichna). In a subsequent paper, Buatois &
deposits periodically exposed to air or periodi- Mangano (1998) explicitly stated that high
cally inundated, and intermediate between ichnodiversity does not necessarily equate with
aquatic and non-aquatic environments (see also species richness. These authors noted that
Frey & Pemberton 1984, 1987). Associated phy- different ichnogenera recorded in this ichnofacies
sical structures are indicative of periodic sub- might result from minor variations of behaviour
aerial exposure, such as desiccation cracks and of a very simple, unspecialised grazing pattern
raindrop imprints. In fluvial systems, this ichno- developed by a single trace-maker (e.g. Hel-
facies is present in floodplain deposits, covering a minthopsis, Helminthoidichnites, Gordia,
wide variety of sub-environments, such as ponds, Mermia).
levees and crevasse splays (Frey & Pemberton The Mermia ichnofacies characterizes fine-
1984, 1987; Frey et al. 1984; Buatois & Mangano grained sediments that occur in well-oxygenated,
1995a, 2002). In lacustrine complexes, the Scoye- low-energy, permanently subaqueous zones of
nia ichnofacies typically characterizes lake- lacustrine systems (Buatois & Mangano 1995a).
margin deposits, being present in both open Under anoxic conditions the ichnofacies is sup-
and closed lake basins, and in both ephemeral pressed. This ichnofacies is typical of open peren-
and perennial systems (Buatois & Mangano nial siliciclastic lacustrine systems, but has been
1998). The Scoyenia ichnofacies also occurs in recognised in carbonate lakes also (Buatois
wet interdunes (Buatois & Mangano 1996). et al. 2000a; de Gibert et al. 2000). The Mermia
NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 315

ichnofacies comprises sediments deposited in availability is of prime importance in controlling


fully lacustrine environments, extending from trace fossil distribution in continental environ-
shallow to deep bathymetric zones. There are ments, particularly in alluvial settings (Gier-
no archetypal trace fossil associations that lowski-Kordesch 1991). In turn, sediment water
clearly distinguish shallow from deep-lacustrine content strongly influences substrate degree of
settings, probably because of the wide variability consolidation. The role of substrate consolida-
of lacustrine systems. Because of this, Buatois & tion as controlling trace fossil preservation is
Mangano (1998) regarded the Mermia ichno- emphasised in this paper and, accordingly, the
facies as a continental equivalent of the Scoyenia and Mermia ichnofacies can be seen,
Cruziana, Zoophycos and Nereites ichnofacies at least in some sense, as taphofacies sensu
in the classical Seilacherian scheme. It is not Bromley & Asgaard (1991).
uncommon for trace fossil assemblages typically
recorded from relatively deep lacustrine areas to
be found in shallower zones. For example, Pick- Ichnology of fluvial systems
erill (1992) documented the Mermia ichnofacies
in lacustrine shoreface deposits, being deeper Trace fossils in fluvial successions are usually
zones of the lake characterised by anoxia. restricted to certain beds and depositional sur-
Buatois & Mangano (2002) expanded the faces. Fluvial ichnofaunas have been reported
environmental extent of the Mermia ichnofacies from two main sub-environments: channel and
to cover ichnofaunas produced in floodplain overbank settings (Buatois & Mangano 1996).
water bodies under subaqueous conditions.
This is supported by recent studies on the ichnol-
ogy of modern floodplains (Mikulas 2003). The Fluvial channel ichnofaunas
lower ichnodiversity of these floodplain ichno-
coenoses in comparison with lacustrine assem- Fluvial channels are characterised by high to
blages may reflect less stable conditions and the relatively high-energy, rapid fluctuations in
temporary nature of floodplain ponds. As rates of sedimentation and erosion, and coarser
stated in its original definition (Buatois & Man- grain sizes than those typically deposited in adja-
gano 1995a, table 2), the Mermia ichnofacies cent environments. Channels therefore represent
may occur in fjord settings under freshwater con- stressful environments for benthic organisms,
ditions owing to glacial melting (see discussion in and production and/or preservation of biogenic
Buatois & Mangano 2003). This is a common structures is usually precluded. Two main trace
situation in late Palaeozoic postglacial deposits fossil associations can be distinguished in fluvial
of Gondwana, where the Mermia ichnofacies channels, broadly corresponding to active and
developed in fjord environments affected by a abandoned channels.
strong discharge of freshwater due to melting The ichnofauna from active channels is char-
of the ice masses during deglaciation (Buatois acterised by low-diversity, typically monospeci-
et al 2001; Pazos 2002; Buatois & Mangano fic, suites of simple vertical burrows and escape
2003). traces (Fig. 4). Up to 30cm deep Skolithos,
It should be stressed that ichnofacies are not forming dense assemblages, were documented
indicators of particular sedimentary environ- in Permian channel-pebble conglomerate and
ments but reflect sets of environmental factors trough cross-bedded fine- to coarse-grained
instead. This is not exclusive to continental sandstone of Southern Victoria Land, Antarctica
ichnofacies; it is a lesson learnt from the history (Fitzgerald & Barrett 1986). Similar vertical
of marine ichnology (Buatois & Mangano burrows in fluvial channel facies have been
2002). As stated by Frey et al (1990), ichnofacies recorded in other Devonian and Permian units
are not intended to be simply indicative of of Antarctica (Bradshaw 1981; Zawiskie et al.
palaeobathymetry. For example, although typi- 1983; Woolfe 1990). Escape traces were recorded
cal of foreshore to upper-foreshore settings, the in accreting parallel-laminated sandstones
Skolithos ichnofacies may occur in more distal formed in channel bars (Sarkar & Chaudhuri
deposits, from lower-shoreface and offshore tem- 1992). The identity of the trace-maker and the
pestites to deep-marine turbidites (e.g. Crimes functional significance of the vertical burrows
1977; Pemberton & Frey 1984). Regardless of in fluvial channel facies are poorly understood.
the depositional environment involved, ichnofa- Fitzgerald & Barrett (1986) suggested that
cies development reflects a set of environmental 'worms' [sic] responded to a fluctuating water
conditions. The same certainly applies for conti- table in presumably ephemeral fluvial systems
nental ichnofacies that are essentially controlled in the same way that polychaete annelid worms
by a set of local environmental factors. Water adjust their burrows to changes in water level
Fig. 4. Taphonomic pathways of fluvial ichnofaunas, showing transitions between different channel and overbank trace-fossil assemblages. Substrate consolidation
plays a major role in controlling ichnofacies occurrence.
NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 317

in intertidal marginal marine environments. cases (Stanley & Fagerstrom 1974; Miller &
Burrowing crustaceans and other invertebrates Collinson 1994). Trace-producers are inferred
are able to adapt to sandy channels as far as to be behavioural generalists recording an oppor-
3000km upstream from the fluvial mouth tunistic strategy (Miller & Collinson 1994). Over-
(Zawiskie et al. 1983). The presence of vertical all, this kind of trace fossil association is similar
burrows in relatively high-energy deposits invites to those from overbank deposits. This is hardly
comparison with the Skolithos ichnofacies, and surprising, because abandoned channels lead to
suggests their functional interpretation as the formation of ponded areas, representing a
domiciles of suspension feeders. process of floodplain construction (abandoned-
Abandoned or inactive channel deposits are channel accretion of Miall 1996). A somewhat
characterised by low-diversity assemblages anomalous occurrence of fluvial channel ichno-
dominated by meniscate traces that have been faunas has been reported by MacNaughton &
assigned to Be aconites or Taenidium (Fig. 4) Pickerill (1995) from the Triassic of Eastern
(Keighley & Pickerill 1994; Goldring & Pollard Canada. In contrast to the usual situation,
1995). Accessory components include vertical to these channel deposits are more intensely bio-
inclined burrows (Skolithos) and simple horizon- turbated and display higher ichnodiversity than
tal burrows (Palaeophycus). Examples of this adjacent overbank deposits. These authors
type of ichnofauna have been documented suggested preferential colonization of stream
repeatedly in the literature (e.g. Allen & Williams channels within a fluvial system developed
1981; Graham & Pollard 1982; Bamford et al. under arid conditions with limited water supply.
1986; Sarkar & Chaudhuri 1992; Miller & Collin-
son, 1994; Miller 2000). This ichnofauna reflects
colonization of sandstone deposits after channel Overbank ichnofaunas
diversion ('abandonment') or, more rarely,
during periods of low discharge virtually charac- Overbank deposits usually host the most diverse
terised by non-deposition ('inactive'). Meniscate and abundant trace fossil assemblages in fluvial
burrows most likely reflect the activity of vagile depositional systems (e.g. Fordyce 1980; D'Ales-
organisms moving into the substrate in search sandro et al. 1987; Buatois et al. 1997; Buatois &
for food, revealing a combination of bypassing Mangano 2002). It is not unusual for the only
and ingestion. Vertical to inclined burrows have trace fossils in a fluvial succession to be preserved
several functions, including permanent domi- in fine-grained overbank deposits interbedded
ciles, semi-permanent shelters, nests and passage- within unbioturbated, stacked channel deposits.
ways. Sarkar & Chauchuri (1992) noted that, in The presence of distinct trace fossil-bearing
contrast to typical vertical burrows in marine pond deposits within an otherwise unfossilifer-
settings, linings are absent. This suggests that ous fluvial succession may thus be regarded as
construction of non-marine vertical burrows taphonomic and colonization windows (Buatois
was probably by insects, because insects do not et al. 1997). The presence of these trace fossil
usually produce discrete reinforcements to suites reflects minor breaks in sedimentation,
burrow walls. These unlined insect burrows representing local autocyclically induced hiatal
have commonly been referred to Cylindricum surfaces, but which are of limited lateral extent.
(Pollard & Lovell 1976). Some of these traces Conversely, during periods of rapid continuous
are probably not strictly dwelling structures, deposition invertebrate colonization is pre-
but temporary shelters in which insects live for cluded. Trace fossil distribution in fluvial settings
relatively short periods of time (Stanley & Fager- is largely a function of variability in stream
strom 1974). Some vertical burrows have basal discharge and the amount of time between
bulbous terminations, representing nests exca- depositional episodes (D'Alessandro et al.
vated and/or constructed by insects for breeding 1987). Taphonomic constraints are key to the
purposes, which are included in the ethological final shaping of fluvial ichnofaunas. Ratcliffe &
category calichnia (Genise & Bown 1994a). The Fagerstrom (1980) have shown that Holocene
presence of nesting structures in abandoned overbank deposits are very rich in invertebrate
channel deposits reflects the ability of insects to traces, but their ancient counterparts are consid-
colonize different substrate types (Genise et al. erably poorer, indicating the low preservation
2000). Additionally, vertical burrows may serve potential of these biogenic structures. Maples
as passageways connecting adjacent bioturbated & Archer (1989) analysed this problem further,
layers. Congruence of size distributions and suggesting that preservation of most of the very
common intergradations between the different shallow insect traces in floodplain deposits
morphotypes present in abandoned channel requires exceptional conditions, including depos-
deposits point to a common producer in most ition of fine-grained heterogeneous sediment,
318 L. A. BUATOIS & M. G. MANGANO

little or no reworking, and enough time between moderate, but vertebrate traces may be relatively
depositional events to allow colonization, but diverse. Associated physical structures are
not so much time that plant colonization obliter- indicative of periodic subaerial exposure (e.g.
ates animal traces (Fig. 4). Overbank settings desiccation cracks and raindrop imprints). Desic-
include a wide variety of deposits, such as flood- cated floodplain ichnofaunas represent an
plains, crevasse splays and levees. The ichnology example of the Scoyenia ichnofacies (Buatois &
of these deposits has recently been reviewed by Mangano 2002). Some of the trace fossils from
Buatois & Mangano (2002), who recognised this assemblage display features indicative of
two main recurrent trace fossil associations, firm substrates, such as striated walls in Scoyenia
which are herein referred to as the 'desiccated and Spongeliomorpha, sharp scratch marks in
overbank' and the 'overfilled overbank' associa- Tambia, Cruziana and Rusophycus, and well-
tions. defined appendage imprints in arthropod track-
Desiccated overbank deposits correspond to ways. In fact, desiccated floodplain ichnofaunas
floodplain water bodies that experienced pro- may be subdivided into two distinct suites: one
gressive drying (Fig. 4). Ichnofaunas from these 'pre-desiccation suite' characterised by menis-
deposits include abundant arthropod trackways cate, backfilled structures without ornamenta-
(e.g. Diplichnites, Trachomatichnus), vertebrate tion (e.g. Taenidium, Beaconites) developed in a
trackways (e.g. Limnopus, Hyloidichnus), back- soft substrate, and the second 'desiccation suite'
filled meniscate traces (e.g. Scoyenia, Taenidium), typified by striated traces (e.g. Scoyenia, Sponge-
ornamented burrows (e.g. Spongeliomorpha, liomorphd), cross-cutting the former and devel-
Tambid), and bilobate traces with scratch oped in a firm substrate (Fig. 5a, b) (Buatois
marks (Cruziana, Rusophycus). Vertical burrows, et al. 1996b). The resulting palimpsest ichno-
referred to Skolithos and Cylindricum, are acces- coenoses/assemblages reflect changing substrate
sory components. Insect and arachnid nesting conditions during progressive desiccation of
structures may also be present. Examples of floodplain sediments (Fig. 4). Desiccated over-
this type of association have been documented bank ichnofaunas tend to be dominant in
by numerous authors (e.g. Bromley & Asgaard distal overbank settings and/or arid to semiarid
1979, their Scoyenia and Rusophycus assem- settings.
blages; Biron & Dutuit 1981; Bracken & Picard Overfilled overbank deposits correspond to
1984; Squires & Advocate 1984; D'Alessandro floodplain water bodies that are filled by over-
et al 1987; Debriette & Gand 1990; Sarkar & bank vertical accretion without experiencing
Chaudhuri 1992; Smith 1993; Gand et al 1997; desiccation (Fig. 4). Ichnofaunas from overfilled
Kim & Paik 1997; Eberth et al 2000; Savrda overbank deposits are dominated by simple
et al 2000; Aramayo & Bocanegra 2003). Inver- grazing trails (e.g. Helminthopsis, Helminthoi-
tebrate ichnodiversity is low and only rarely dichnites), locomotion trails (e.g. Cochlichnus)

Fig. 5. Two suites of the Scoyenia ichnofacies in desiccated overbank deposits. Permian, La Colina Formation,
La Rioja province, western Argentina. All bars are 1 cm. (a) Beaconites barretti. Meniscate backfilled trace
fossils lacking striated walls, suggesting emplacement in a softground. (b) Firmground meniscate striated trace
fossils cross-cutting the softground suite.
NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 319

Fig. 6. Depauperate Mermia ichnofacies in overfilled overbank deposits. Permian, La Golondrina Formation,
Santa Cruz Province, southern Argentina. All bars are 1 cm. (a) Poorly defined sinusoidal trails of
Cochlichnus anguineus, showing intense deformation and presence of irregular levees. Sinusoidal trails
cross-cut by shafts of Ctenopholeus kutscheri. Morphologic features indicate emplacement in water-saturated
substrates, (b) Overlapping of well-defined specimens of Cochlichnus anguineus emplaced in a more cohesive
substrate.

and horizontal dwelling burrows (e.g. Cteno- floodplain basins support the establishment of
pholeus, Palaeophycus). Accessory components an aquatic biota. Floodplain ponds represent
include resting traces of bivalves or conchostra- suitable environments for the development of
cans (e.g. Lockeia). Tetrapod traces, arthropod low- to moderate-diversity suites of inverte-
trackways and bilobate traces with scratch brates, such as insects, crustaceans, nematodes,
marks are either absent or represent very minor nematomorphs, oligochaetes, molluscs, ostra-
components of the assemblage, in contrast to codes and conchostracans. The lower ichnodiver-
the desiccated overbank assemblage. Backfilled sity of these water bodies in comparison with
traces are typically absent. Examples of these their equivalents from lacustrine basins is prob-
ichnofaunas are also well documented (Turner ably an expression of the less stable conditions
1978; Fordyce 1980; Miller 1986; Pollard & and temporary nature of overbank environ-
Hardy 1991; Gluszek 1995; Buatois el al. 1997; ments. However, differences in stability and
Buatois & Mangano 2002). Ichnodiversity is temporal persistence between these two broad
thus generally low but may be moderate in environmental settings is irrelevant for opportu-
some cases. The assemblage is characterised by nistic organisms able to rapidly colonize stagnant
superficial to very shallow structures (Fig. waters in floodplain environments. Overfilled
6a, b). Physical structures indicative of subaerial overbank ichnofaunas tend to dominate in
exposure are absent. Additionally, morpho- proximal overbank settings and/or temperate
logical details are very poorly preserved (Fig. and humid settings.
6a), suggesting that the traces were formed in a
water-saturated substrate (e.g. Buatois et al.
1997). Features of these ichnofaunas reflect the Ichnology of lacustrine systems
subaqueous nature of the associated environ-
ment. Poorly preserved traces may be cross-cut Recent interest in lacustrine ichnology has paral-
by better-defined softground reflecting improv- leled renewed research into the sedimentology
ing taphonomic conditions due to increasing and stratigraphy of lake successions (e.g.
compaction (Figs 4, 6b). Overbank vertical Anadon et al. 1991; Gierlowski-Kordesch &
accretion with little associated erosion com- Kelts 1994, 2000). Lacustrine biogenic structures
monly allows good preservation of the ichno- probably have the highest preservation potential
fauna and precludes desiccation of the water of all continental trace fossils. Preservation of
bodies. Buatois & Mangano (2002) noted that, biogenic structures is particularly favoured in
although formed in floodplains, these ichno- low-energy settings. For example, alternation of
faunas lack most of the diagnostic features of very fine sand and mud deposited from under-
the Scoyenia ichnofacies, and suggested that flow currents is ideal for preservation of tiny
they represent examples of an impoverished surface to very shallow traces (Buatois & Man-
Mermia ichnofacies. Although overfilled over- gano 1995a, 1998). Underflow currents deposit
bank settings are of shorter lifespan than perma- sediments well below wavebase, where physical
nent lakes, freshwater bodies formed in reworking is rare. Low-density, fine-grained
320 L. A. BUATOIS & M. G. MANGANO

turbidity currents can usually create a similar Closed lake ichnofaunas


taphonomic scenario. In low-energy shoreline
areas, trace fossil preservation may commonly Closed lakes are characterised by high salinity
be linked to rapid influx of sand via non-erosive and rapidly fluctuating shorelines (Gore 1989).
sheet floods entering the lake (e.g. Zhang et al. This type of lake represents stressful environ-
1998). Lacustrine systems can be divided into mental conditions for the components of such
hydrologically open (i.e. with an outlet) and ecosystems. Accordingly, faunal diversity is typi-
hydrologically closed (i.e. without an outlet) cally extremely low, and biogenic structures are
(Gore 1989). few, if not completely absent. In some cases,

TAPHONOMIC PATHWAYS
LACUSTRINE ICHNOFAUNAS

Fig. 7. Taphonomic pathways of lacustrine ichnofaunas. Hydrologically open lakes contain more varied
softground ichnofacies. Vertical ichnofacies changes reflect lake regression. Hydrologically closed lakes display
limited softground ichnofacies, but abundant firmground suites in lake-margin deposits.
NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 321

Fig. 8. Densely superimposed trackways in marginal deposits of playa lakes. Permian? La Rioja Province,
western Argentina, (a) General view of a sandstone top exhibiting high density of arthropod trackways.
Coin is 1.6cm. (b) Close-up of the tracked surface. Bar is 1 cm.

trace fossils are restricted to certain beds and ichnofaunas. The Scoyenia ichnofacies is typi-
record short periods of reduced stress, such as cally present in low-energy, lake-margin areas,
periods of increased productivity or reduced whereas the Mermia ichnofacies characterizes
salinity (Price & McCann 1990). The trace permanent subaqueous lacustrine zones. Sedi-
fossil Beaconitesfiliformis, attributed to chirono- ments deposited in moderate to high-energy
mids, occurs in hypersaline lacustrine deposits lacustrine environments, such as wave-
(Uchman & Alvaro 2000). Notably, an ichno- dominated shorelines and delta mouth-bars, are
fauna of moderate diversity has been documen- commonly characterised by the Skolithos ichno-
ted in Neogene lacustrine evaporites of Spain facies (Fig. 9). These deposits contain simple
(Rodriguez-Aranda & Calvo 1998). The ichno-
fauna consists of plant, invertebrate traces of
chironomids, coleopterans and annelids, and
vertebrate traces of mammals and birds.
Undoubtedly, the richest ichnofaunas in closed
lake systems are present at the lake margins
and record the activity of terrestrial rather than
aquatic faunas (Fig. 7). These lake-margin trace
fossil assemblages are typical examples of the
Scoyenia ichnofacies, dominated by meniscate
backfilled traces, striated burrows and arthropod
trackways. Under appropriate taphonomic
conditions, omission surfaces totally covered by
trackways are preserved (Figs 7, 8a, b). Asso-
ciated physical structures indicate subaerial
exposure (e.g. desiccation cracks, raindrop
imprints). Examples of these ichnofaunas in
playa lake settings have been reported in a
number of studies (Bromley & Asgaard 1979;
Gierlowski-Kordesch 1991; Dam & Stemmerik
1994; Kozur & Lemone 1995; Clemmensen et al.
1998; Zhang et al. 1998; Schlirf et al. 2001).

Open lake ichnofaunas


Fig. 9. Elements of the Skolithos ichnofacies in high-
Open lakes are characterised by low salinity and energy continental deposits. Arenicolites isp. deltaic
relatively stable shorelines (Gore 1989). Such mouth-bar deposits. Triassic, Tanzhuang Formation,
lakes may host relatively diverse and abundant central China.
322 L. A. BUATOIS & M. G. MANGANO

vertical burrows (Skolithos), U-shaped vertical dominant component of the ichnofacies, with
burrows (Arenicolites) and escape structures arthropod trackways subordinate. Vertebrate
(e.g. Bromley & Asgaard 1979; Mangano et al. traces are represented by the fish trail Undichna
1994). (Anderson 1976; Higgs 1988; Turek 1989; Bua-
The distinction between lake-margin and fully tois & Mangano 1994; de Gibert et al. 1999),
subaqueous deposits can be established through and the amphibian trackways include Lunich-
careful ichnological analysis. Lake-margin nium and Gracilichnium (Turek 1989). Oxygen
deposits are characterised by meniscate, back- content, energy, food supply and substrate are
filled trace fossils and arthropod trackways of important controls on trace fossil distribution
the Scoyenia ichnofacies. Meniscate trace fossils in lacustrine systems. Oxygenation is commonly
are well documented from marginal lacustrine a limiting factor; in lakes with thermal stratifica-
facies (e.g. Daley 1968; Metz 1995, 1996; Hester tion the hypolimnion becomes anoxic/dysoxic,
& Lucas 2001), whereas assemblages dominated and bioturbation is absent. Turbidity and
by arthropod trackways have been recorded in underflow currents may provide oxygen to lake
the same setting (e.g. Pollard et al. 1982; Pollard bottoms, allowing the establishment of benthic
& Walker 1984; Walker 1985; Bandel & Quinzio- communities. Examples of permanently sub-
Sinn 1999; Cook & Bann 2000). Desiccation of aqueous freshwater ichnofaunas have been
marginal lacustrine deposits allows construction reported in various studies (Gibbard & Stuart
and preservation of striated meniscate traces 1974; Gibbard 1977; Gibbard & Dreimanis
and burrow galleries, such as Spongeliomorpha 1978; Walter 1985; Miller et al. 1991; Pickerill
(Metz 1993) in firmground substrates. More 1992; Buatois & Mangano 1993; Buatois et al.
rarely, bioerosion in stromatolites has been 1996a, 2000a; Walter & Suhr 1998; de Gibert
documented (Ekdale et al. 1989). Metz (1996) et al. 2000; Melchor et al. 2003; Melchor 2004).
noted that, in Triassic lacustrine deposits of the Ichnological analysis helps to distinguish
Newark Basin, elements of the Mermia ichno- between density underflows and turbidity cur-
facies are replaced by typical representatives of rents. Both processes commonly operate in
the Scoyenia ichnofacies during shoreline open lacustrine systems and are difficult to
regression. differentiate on sedimentological criteria alone.
Vertebrate trace fossils are extremely common Turbidites are deposited by episodic currents
in marginal lake facies (e.g. Olsen et al. 1978; that involves re-deposition of sediment initially
Lockley et al. 1986; Lim et al. 1989; Prince & emplaced under unstable conditions. Once
Lockley 1989). Avian trackways are also useful mobilised, a dense turbid fluid flows downslope
to delineate palaeoshorelines in lacustrine succes- to the basin-floor to deposit the turbidite. Under-
sions (Alonso 1985; Yang et al. 1995). A similar flow currents are a relatively continuous
situation is recorded by dinosaur trackways phenomenon that represents the uninterrupted
that occur at the top of prograding, shallowing- transport of river-borne sediment into the lake
upward lacustrine successions of the Jurassic basin, and are influenced by geostrophic effects
Morrison Formation of Colorado, recording (Pharo & Carmack 1979). Turbidites contain
the position of the ancient shoreline (Lockley trace fossils at bedding tops, recording coloniza-
et al. 1986; Prince & Lockley 1989). Dinosaur tion of opportunistic organisms after episodic
and fitosaur trackways are present in Triassic emplacement of the event bed (Buatois &
lacustrine mudflat facies of the northwest Mangano 1998). Underflow current deposits dis-
United States, associated with successive maxi- play distinctive suites of trace fossils in each
mum regressive phases of the lacustrine system lamina or lamina-set (e.g. Buatois & Mangano
(Olsen et al. 1978). Therefore, vertebrate track- 1995b; Melchor 2001), reflecting animal activity
ways help in delineation of cycles of expansion contemporaneous with sedimentation instead of
and contraction of water bodies (Lockley 1986, re-colonization after deposition of the turbidite
1989). Additionally, vertebrate trackways pro- bed (Buatois & Mangano 1998). The contrasting
vide valuable information on palaeoclimatic colonization styles allow differentiation between
conditions. For example, trackways of theropod near-continuous deposition from river-fed
dinosaur dominate over those of herbivorous density underflows and episodic sedimentation
dinosaur in semiarid lacustrine systems (Leo- from turbidity currents.
nardi 1989). Biogenic structures are also useful to distin-
In the permanent subaqueous zone of lakes, guish between marine and lacustrine turbidites,
under conditions of low energy and a high which are virtually indistinguishable in terms of
degree of environmental stability, the Mermia physical sedimentary structures. Deep-sea turbi-
ichnofacies develops. Feeding and grazing dites are characterised by a high diversity of
traces of detritus and deposit feeders are the ornate grazing traces and graphoglyptids that
NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 323

reflect highly specialised feeding strategies ichnological zonations may prove to be useful
recorded by the Nereites ichnofacies (Seilacher at the basin scale.
1967; Miller 1991; Uchman 1995, 2004). Deep
marine environments are typically more stable
than lacustrine systems and host a more diverse Applications of ichnology in continental
benthic fauna. In contrast, deep-lacustrine sequence stratigraphy
settings are characterised by non-specialised
grazing and feeding traces (Buatois & Mangano In comparison with their marine counterparts,
1998). Non-specialised feeding patterns are continental trace fossils have not been exten-
exemplified by the ichnogenus Mermia, which sively used in sequence stratigraphy. In marine
displays looping and common self-crossing, siliciclastic successions, biogenic structures aid
recording the repeated passage of the trace- in sequence stratigraphy in two main ways: (1)
maker across the same portion of sediment. identification of stratigraphic discontinuities
Such non-specialised trophic strategies probably using substrate-controlled ichnofacies, and (2)
relate to the abundance and accessibility of food recognition of palaeoenvironmental changes
in lacustrine systems (Buatois & Mangano through detailed documentation of vertical
1995a). The comparatively lower ichnodiversity changes in softground trace fossils (see recent
of lakes in comparison with deep marine settings reviews in Pemberton et al. 2001; Mcllroy 2004;
results from the more ephemeral nature of the Pemberton 2004). In particular, the firmground
former (Buatois & Mangano 1998). Structures Glossifungites ichnofacies develops in stable
referred to Paleodictyon in freshwater assem- and cohesive substrates, reflecting erosive exhu-
blages (Archer & Maples 1984; Pickerill 1992) mation of the substrate (MacEachern et al.
are remarkably simpler than those from marine 1992). The hardground Trypanites ichnofacies
turbidites. A feeding trace referred to Nereites and the woodground Teredolites ichnofacies
in lacustrine turbidites (Hu et al. 1998) lacks are used to a lesser extent (Pemberton et al.
the internal structure of this ichnogenus and 2001). Currently recognised allostratigraphic
only superficially resembles this marine form. surfaces are incised valley surfaces, incised sub-
Although most ichnological studies have marine canyon surfaces, regressive surfaces of
focused on siliciclastic systems, a few papers marine erosion, ravinement surfaces (transgres-
dealing with the ichnology of ancient carbonate sive) and co-planar surfaces of lowstand and
lakes have been published in recent years. Inver- transgressive erosion (MacEachern et al. 1992;
tebrate ichnofaunas in Cretaceous lacustrine Pemberton et al. 2001; Pemberton 2004). Soft-
lithographic limestones of Spain are restricted ground ichnofacies and ichnofabrics are mostly
and dominated by small, horizontal, shallow- used to detect sharp palaeoenvironmental
tier trace fossils of detritus and deposit feeders changes across discontinuity surfaces or gradual
(de Gibert et al. 2000; Buatois et al 2000a). palaeoenvironmental changes across parase-
Associated vertebrate ichnofaunas comprise quences. In the former case, they aid in the clas-
crocodilian trackways (Moratalla et al. 1995), sification of stratigraphic discontinuities; in the
pterosaurian tracks (Lockley et al. 1995) and latter, they allow palaeoenvironmental zonation
fish trails (de Gibert et al. 1999). Additionally, of parasequences and help in identifying trends
it has been noted that these lacustrine ichno- in parasequence stacking patterns.
faunas differ in taxonomic composition, The use of trace fossils in continental sequence
proportion of infauna and ethologic significance stratigraphy cannot be simply based on the
from trace fossil assemblages described from extrapolation of concepts established from the
marine lithographic limestones (Buatois et al. analysis of the marine stratigraphic record. Dis-
2000a). crepancies result from both differences in the
In large, deep lakes, depth-related trace fossil nature of continental versus marine ichnofaunas
zones can be established. For example, Walter and peculiarities of continental depositional
& Suhr (1998) documented a bathymetric systems with respect to the marine system. For
zonation in Pleistocene glacial lakes of Germany. example, substrate-controlled trace fossils in
In these lakes, shallow-lacustrine trace fossil continental successions only rarely indicate ero-
assemblages are dominated by arthropod track- sional exhumation; they are commonly related
ways (e.g. Warvichnium, Glaciichnium, Lusatich- to desiccation of water bodies (e.g. Buatois et al.
nium), whereas grazing trails are abundant 1996b). Moreover, continental firmgrounds form
in deeper zones (e.g. Cochlichnus, Gordia, rapidly under conditions of subaerial exposure,
Helminthoidichnites). Although there are no without implying a significant hiatus (Flirsich &
archetypal, recurrent ichnofacies that clearly dis- Mayr 1981). Additionally, some sequence strati-
tinguish shallow- from deep-lacustrine settings, graphic concepts (e.g. parasequence) are difficult
324 L. A. BUATOIS & M. G. MANGANO

Fig. 10. Trace fossil assemblages and lacustrine sequence stratigraphy: (a) overfilled lakes; (b) balanced-fill
lakes; (c) underfilled lakes. Stratal patterns illustrated after Bohacs et al. (2000).
NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 325

to apply in certain continental settings, such as hummocky cross-stratified sandstones deposited


fluvial depositional systems (Van Wagoner et al. by storm events, as well as fair-weather wave-
1990). Lacustrine systems also defy direct and combined-flow ripple cross-laminated sand-
application of the sequence stratigraphic model stone. Grazing trails of the Mermia ichnofacies
established for marine siliciclastic systems. As may occur forming colonization suites at the
noted by Bohacs et al. (2000), lakes differ from top of tempestites in such settings (e.g. Buatois
oceans in several ways, including the smaller & Mangano 1995b; Melchor et al. 2003). How-
volumes of sediment and water included in lacus- ever, assemblages are usually impoverished with
trine systems, the direct link between lake level respect to those of the more distal facies (Buatois
and sediment supply, and the fact that lake & Mangano 1998). Under conditions of continu-
shoreline migration may be due not only to ous wave agitation, elements of the Skolithos
progradation but also to withdrawal of water. ichnofacies may occur. Proximal facies include
Finally, in continental systems other allocyclic distributary channel, trough and tabular cross-
controls, such as tectonism and climate, play bedded sandstones that are commonly unbiotur-
major roles (Shanley & McCabe 1998). bated. Locally, escape traces and vertical - or
Bohacs et al. (2000) provided a workable more rarely horizontal - domiciles of suspension
sequence stratigraphic framework for the analy- feeders may be present, recording an occurrence
sis of lacustrine successions. These authors of the Skolithos ichnofacies (e.g. Melchor et al.
recognised three different types of lake basin: 2003). In the case of deep overfilled lake basins,
overfilled, balanced-fill and underfilled. Over- lake floor turbidite systems can develop. Turbi-
filled lake basins are characterised by rate of sedi- dite lobe successions are either progradational
ment/water exceeding potential accommodation. or aggradational. Ichnofaunas are relatively
Fluvio-lacustrine siliciclastic deposits that diverse; pre-depositional horizontal trace fossils
accumulate in hydrologically open lakes are the of the Mermia ichnofacies are common on the
most common, and parasequence development soles of thin-bedded turbidite sandstones (e.g.
is driven mainly by shoreline progradation and Buatois et al. 1996a, 2000b). Post-depositional
delta-channel avulsion. Balanced-fill lake basins suites occur either on the base or at the top of
occur when the rates of sediment/water supply turbidites and are commonly less diverse (Bua-
are in balance with potential accommodation. tois & Mangano 1998; Buatois et al. 1998).
Carbonate and siliciclastic facies can accumulate Land-plant-derived organic matter is the prime
in lakes that are alternatively hydrologically source of nutrients, favouring the development
open and closed. Successions record not only of a deposit-feeding benthic fauna in perma-
progradational parasequences, but also aggrada- nently subaqueous, low-energy zones. Firm-
tion of chemical sediments due to desiccation. ground suites are rare because such large lakes
Underfilled lake basins are characterised by usually do not experience desiccation.
rates of accommodation exceeding rate of Balanced-fill lake basins commonly contain
supply of sediment/water. In hydrologically abundant firmground trace fossils, but soft-
closed lakes deposition of evaporites dominates, ground assemblages are usually depauperate
and parasequences record vertical aggradation (Fig. lOb). During lowstands, shallow
due to desiccation. balanced-fill lakes are characterised by relatively
Softground trace fossils are commonly well thin aggradational parasequences due to desicca-
developed in overfilled lake basins (Fig. lOa), tion (Bohacs et al. 2000). Lowstand deposits
and are useful to delineate parasequences and contain abundant and widespread ichnofaunas
parasequence sets. Fluvial discharge into over- of the Scoyenia ichnofacies. Striated trace fossils,
filled lakes usually generates density currents such as Scoyenia and Spongeliomorpha, record-
that oxygenate lake bottoms, allowing the estab- ing the firmground suite of the Scoyenia ichno-
lishment of epifaunal and infaunal communities. facies, are extensively developed during lake
Additionally, hydrologically open lakes promote desiccation (e.g. Bromley & Asgaard 1979;
freshwater conditions and no stress due to hyper- Metz 1995; Clemmensen et al. 1998). Subsequent
salinity occurs, leading to the development of a flooding is associated with rapid influx of sand,
relatively diverse benthos. Upward-shallowing allowing preservation of biogenic structures. In
successions due to delta progradation are contrast, relatively thick aggradational para-
common. Distal facies commonly consist of sequence sets form in lake-floor turbidite systems
underflow current deposits that display elements in deep balanced-fill lakes during lowstands
of the Mermia ichnofacies in each lamina or (Bohacs et al. 2000). Under these conditions,
lamina-set (e.g. Buatois & Mangano 1995b; firmground trace fossils are absent. Lake hydrol-
Melchor 2001). Intermediate facies may contain ogy is closed during lowstands and salinity
wave-dominated delta-front deposits, including usually increases (Bohacs et al. 2000), therefore
326 L. A. BUATOIS & M. G. MANGANO

imposing a stress factor on the lake biota and experience rapid expansion and flash floods
impoverished softground ichnofaunas. Ichno- reach the basin, leading to deposition of sandy
faunas from turbidite systems in balanced-fill inundites. Trace fossil preservation is mostly
lakes are less abundant and diverse than those linked to rapid influx of sand via sheet floods
from overfilled lake turbidites. During transgres- entering into the lake (Zhang et al. 1998). Hyper-
sion, parasequences are relatively thick and salinity usually prevents the establishment of a
display retrogradational stacking patterns, subaqueous Mermia ichnofacies during trans-
while highstand parasequences are variable in gression and highstand. However, elements of
thickness and are either aggradational or progra- the Mermia ichnofacies may occur, albeit in
dational (Bohacs et al. 2000). Freshwater con- reduced numbers, in very shallow water thin
ditions are common during transgression, but deposits immediately above flooding surfaces at
dysaerobic conditions prevail, therefore produ- the base of parasequences. This assemblage is
cing a stress factor on the benthic biota. Trace abruptly replaced upward by the Scoyenia ichno-
fossils may occur locally in transgressive and facies reflecting lake regression (e.g. Metz 1996,
highstand carbonates. However, ichnodiversity 2000). Additionally, dwelling traces possibly
is low, and any trace fossils are almost exclusively produced by aquatic chironomid larvae may be
those produced by epifaunal organisms. This present (Rodriguez-Aranda & Calvo 1998;
dominance of surficial trace fossils and the Uchman & Alvaro 2000). Transgressive systems
paucity of infaunal traces result from brief tracts recorded by thin retrogradational parase-
periods of bottom-water oxygenation, but quence sets usually reflect drastic ichnofaunal
permanently anoxic interstitial waters that dis- changes, from terrestrial assemblages (Coprini-
courage infauna (e.g. Buatois et al. 2000a). A sphaera ichnofacies) to transitional terrestrial-
depauperate Mermia ichnofacies is present in subaqueous assemblages (Scoyenia ichnofacies)
these deposits. Scarcity of biogenic structures and salinity-tolerant subaqueous monospecific
due to oxygen depletion was also noted in trans- assemblages of Beaconites filiformis attributed
gressive and highstand siliciclastic deposits of to chironomids (Uchman & Alvaro 2000).
balanced-fill lakes (e.g. Olsen 1989; Mangano In alluvial settings, the sparse distribution of
et al. 1994, 2000; Metz 1995). Bioturbation is trace fossils primarily reflects changes in deposi-
occasionally present at the top of turbidite tional systems that may, in turn, be linked to sys-
sandstones, indicating increasing oxygenation tems tracts. Amalgamated and interconnected,
in the aftermath of turbidity currents (Mangano incised fluvial channels are commonly developed
et al. 1994, 2000). Additionally, elements of the in response to low rates of accommodation
Skolithos ichnofacies may occur in delta during early lowstands (Legarreta et al. 1993;
mouth-bars during highstand progradation of Legarreta & Uliana 1998; Shanley & McCabe
deltaic systems (Bromley & Asgaard 1979; 1998; Posamentier & Allen 1999). High-energy
Mangano et al. 1994, 2000). conditions associated with widespread, intense
The Scoyenia ichnofacies is widespread in erosion typically lead to extensive reworking of
underfilled lake basins, but the Mermia ichno- fluvial deposits, and high sedimentation rates
facies is commonly suppressed (Fig. lOc). prevent formation and/or preservation of bio-
Deposition during lowstands is restricted to eva- genic structures in fluvial channels. However,
porite accumulation in remnant pools developed palaeosols are commonly developed in interfluve
in the zones of maximum subsidence (Bohacs areas, and terrestrial trace fossils representing the
et al. 2000). Evaporite pools are among the work of social insects are very common, particu-
most stressful environments and, with few excep- larly in Cretaceous and younger strata (Genise
tions, lack biogenic structures. In the remaining et al. 2000; Genise et al. 2004). In particular,
zones, sediments that accumulated during the the Coprinisphaera ichnofacies (and other
previous highstand experience extreme desicca- palaeosol ichnofacies still unnamed) may deline-
tion during lowstand (Bohacs et al. 2000). The ate sequence boundaries.
Scoyenia ichnofacies is associated with desic- Increasingly isolated fluvial channels encased
cated substrates formed during lowstand condi- in overbank deposits develop as a result of
tions in underfilled lakes (e.g. Gierlowski- increasing fluvial accommodation during late
Kordesch 1991; Metz 1996, 2000). Density of lowstand. Eventually transgressive lacustrine
arthropod trackways may be high, forming and marsh deposits accumulate when rate of
tracked omission surfaces (e.g. Zhang et al. accommodation exceeds sediment supply (Legar-
1998). These omission surfaces are commonly reta et al. 1993; Posamentier & Allen 1999).
sequence boundaries expressed by co-planar Increased accommodation favours the preserva-
surfaces of lowstand and subsequent flooding. tion of biogenic structures. A trend is apparent
During pluvial periods, underfilled lakes towards the progressive replacement of vertical
NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 327

dwelling burrows and escape traces of the Sko- Although continental trace fossils have not
lithos ichnofacies in active channels by low- been extensively used in sequence stratigraphy,
diversity assemblages of meniscate traces in they have potential for further studies. Soft-
abandoned channels and both the softground ground trace fossils are commonly well devel-
and firmground suites of the Scoyenia ichno- oped in overfilled lake basins and are useful for
facies and even the subaqueous Mermia ichno- delineation of parasequences and parasequence
facies in overbank deposits. This trend is sets. Firmground suites are rare because these
reversed under increased sediment supply and lakes usually do not experience desiccation. In
decreased fluvial accommodation, leading to contrast, balanced-fill lake basins commonly
deltaic progradation and increased channeliza- contain abundant firmground trace fossils in
tion during highstand. lowstand deposits, but softground assemblages
are usually depauperate owing to oxygen-
depleted conditions. The Scoyenia ichnofacies is
Conclusions also widespread in desiccated lowstand deposits
of underfilled lake basins, but the Mermia ichno-
Fluvial ichnofaunas have been reported from facies is commonly suppressed owing to hypersa-
two main sub-environments: channel and over- linity. Early lowstand amalgamated incised
bank settings. The ichnofauna from active fluvial channels are usually unbioturbated
channels is characterised by low-diversity suites owing to high-energy conditions, widespread
of simple vertical burrows and escape traces, and intense erosion and high sedimentation
referred to the Skolithos ichnofacies. Abandoned rates. Palaeosol ichnofaunas (e.g. the Coprini-
or inactive channel deposits contain low- sphaera ichnofacies) delineate sequence bound-
diversity assemblages dominated by meniscate aries in interfluve areas. Increasingly isolated
traces. Accessory components include vertical fluvial channels encased in overbank deposits
to inclined burrows and simple horizontal develop during late lowstands, and lacustrine
burrows. Overbank deposits usually host the deposits accumulate during transgressions. This
most diverse and abundant trace fossil assem- is paralleled by the progressive replacement of
blages in fluvial environments. Floodplain vertical dwelling burrows and escape traces of
water bodies that experienced progressive the Skolithos ichnofacies in active channels by
drying (desiccated overbank deposits) host the low-diversity assemblages of meniscate traces in
Scoyenia ichnofacies, which includes abundant abandoned channels and both the softground
arthropod and vertebrate trackways, backfilled and firmground suites of the Scoyenia ichno-
meniscate traces, ornamented burrows and facies and even the subaqueous Mermia ichno-
bilobate traces with scratch marks. Floodplain facies in overbank deposits. This trend is
water bodies that are filled by overbank vertical reversed owing to deltaic progradation and
accretion without experiencing desiccation (over- increased channelization during highstand.
filled overbank deposits) host depauperate
examples of the Mermia ichnofacies, represented We thank the Antorchas Foundation and the National
by simple grazing trails, locomotion trails and Agency of Science and Technology for providing finan-
horizontal dwelling burrows. cial support. We are very grateful to D. Mcllroy for the
invitation to participate in the Lyell Meeting 2003. F.
Lacustrine biogenic structures probably have Fursich and S. Lucas provided useful reviews. We
the highest preservation potential of all continen- also thank M. Jimenez and D. Ruiz Holgado for the
tal trace fossils. Hydrologically closed lakes are drawings.
very stressful, and subaqueous ichnofaunas are
rare. The richest ichnofaunas in closed lakes
are present at the lake margins and record the References
activity of terrestrial rather than aquatic faunas
(e.g. arthropod trackways, striated traces), ALLEN, J. R. L. & WILLIAMS, B. P. J. 1981. Beaconites
recording the Scoyenia ichnofacies. Hydro- antarcticus: a giant channel-associated trace fossil
logically open lakes host relatively diverse and from the Lower Old Red Sandstone of South
abundant ichnofaunas. The Scoyenia ichnofacies Wales and the Welsh Borders. Geological Journal,
develops in low-energy, lake-margin areas, 166, 255-269.
whereas the Mermia ichnofacies is present in per- ALONSO, R. 1985. Icnitas de aves como control de
niveles boratiferos. Sociedad Cientifica del Nor-
manent subaqueous lacustrine zones. Sediments oeste Argentina, 1, 37-42.
deposited in relatively high-energy lacustrine ANADON, P., CABRERA, L. L. & KELTS, K. 1991. Lacus-
environments, such as wave-dominated shore- trine Fades Analysis. International Association
lines and delta mouth-bars, commonly are of Sedimentologists, Special Publications,
characterised by the Skolithos ichnofacies. Oxford, 13.
328 L. A. BUATOIS & M. G. MANGANO

ANDERSON, A. M. 1976. Fish trails from the early BRADSHAW, M. A. 1981. Palaeoenvironmental interpre-
Permian of South Africa. Palaeontology, 9, 397- tations and systematics of Devonian trace fossils
409. from the Taylor Group (Lower Beacon Super-
ARAMAYO, S. A. & BIANCO, T. M. 1987a. Hallazgo de group), Antarctica. New Zealand Journal of Geol-
una icnofauna continental (Pleistoceno tardio) en ogy and Geophysics, 24, 615-652.
la localidad de Pehuen-Co (partido de Coronel BROMLEY, R. G. 1996. Trace Fossils: Biology,
Rosales), provincia de Buenos Aires, Argentina. Taphonomy and Applications. Chapman & Hall,
Parte I: Edentata, Litopterna, Proboscidea. 4° Con- London.
greso Latinoamericano de Paleontologia, Adas, BROMLEY, R. G. & ASGAARD, U. 1979. Triassic fresh-
Santa Cruz de la Sierra, 1, 516-531. water ichnocoenoses from Carlsberg Fjord, East
ARAMAYO, S. A. & BIANCO, T. M. 1987b. Hallazgo de Greenland. Palaeogeography, Palaeoclimatology,
una icnofauna continental (Pleistoceno tardio) en Palaeoecology, 28, 39-80.
la localidad de Pehuen-Co (partido de Coronel BROMLEY, R. G. & ASGAARD, U. 1991. Ichnofacies: a
Rosales), provincia de Buenos Aires, Argentina. mixture of taphofacies and biofacies. Lethaia, 24,
Parte II: Carnivora, Artiodactyla y Aves. 4° Con- 153-163.
greso Latinoamericano de Paleontologia, Adas, BROMLEY, R. G. & ASGAARD, U. 1993. Endolithic
Santa Cruz de la Sierra, 1, 532-547. community replacement on Pliocene rocky coast.
ARAMAYO, S. A. & BIANCO, T. M. 1996. Edad y nuevos Ichnos, 2, 93-116.
hallazgos de icnitas de mamiferos y aves en el BUATOIS, L. A. & MANGANO, M. G. 1993. Trace fossils
yacimiento paleoicnologico de Pehuen-Co from a Carboniferous turbiditic lake: implications
(Pleistoceno Tardio), provincia de Buenos Aires, for the recognition of additional nonmarine ichno-
Argentina. In: MELCHOR, R. N. (ed.) Asociacion facies. Ichnos, 2, 237-258.
Paleontologica Argentina, Publicacion Especial, BUATOIS, L. A. & MANGANO, M. G. 1994. Pistas de
Buenos Aires, 4, 47-57. peces en el Carbonifero de la cuenca Paganzo
ARAMAYO, S. A. & BOCANEGRA, L. 2003. Icnofacies de (Argentina): Su significado estratigrafico y paleo-
Scoyenia en la Formation Rio Limay (Grupo ambiental. Ameghiniana, 31, 33-40.
Neuquen, Cretacico tardio) Provincia de Neu- BUATOIS, L. A. & MANGANO, M. G. 1995a. The
quen, Argentina. In: BUATOIS, L. A. & MANGANO, paleoenvironmental and paleoecological signifi-
M. G. (eds) Icnologia: Hacia una Convergencia cance of the lacustrine Mermia ichnofacies: an
entre Geologia y Biologia. Asociacion Paleonto- archetypical subaqueous nonmarine trace fossil
logica Argentina, Publicacion Especial, Buenos assemblage. Ichnos, 4, 151-161.
Aires, 9, 43^8. BUATOIS, L. A. & MANGANO, M. G. 1995b. Sedimen-
ARCHER, A. W. & MAPLES, C. G. 1984. Trace-fossil dis- tary dynamics and evolutionary history of a Late
tribution across a marine-to-nonmarine gradient Carboniferous Gondwanic lake in northwestern
in the Pennsylvanian of southwestern Indiana. Argentina. Sedimentology, 42, 415-436.
Journal of Paleontology, 58, 448-466. BUATOIS, L. A. & MANGANO, M. G. 1996. Icnologia de
BAMFORD, M. L. F., BRUCK, P. M., COOPER, M. A., ambientes continentales: Problemas y perspecti-
FORBES, W. H. & MACCARTHY, I. A. J. 1986. Bea- vas. In: MELCHOR, R. N. (ed.) Asociacion Paleon-
conites-type burrows from the Old Red Sandstone tologica Argentina, Publicacion Especial, Buenos
of Hook Head, Co. Wexford, Ireland. Proceedings Aires, 4, 5-30.
Geological Association, 97, 59-71. BUATOIS, L. A. & MANGANO, M. G. 1998. Trace fossil
BANDEL, K. & QUINZIO-SINN, L. A. 1999. Paleozoic analysis of lacustrine facies and basins. Palaeo-
trace fossil from the Cordillera Coastal near Con- geography, Palaeoclimatology, Palaeoecology,
ception, connected to a review of the Paleozoic 140, 367-382.
history of central Chile. Neues Jahrbuch fur BUATOIS, L. A. & MANGANO, M. G. 2002. Trace
Geologie und Paldontologie, Abhandlungen, 211, fossils from Carboniferous floodplain deposits in
171-200. western Argentina: implications for ichnofacies
BIRON, P. E. & DUTUIT, J.-M. 1981. Figurations sedi- models of continental environments. Palaeo-
mentaires et traces d'activite au sol dans le Trias geography, Palaeoclimatology, Palaeoecology,
de la formation d'Argana et de 1'Ourika 183, 71-86.
(Maroc). Bulletin Museum of Natural History, BUATOIS, L. A. & MANGANO, M. G. 2003. Caracteriza-
Paris, 4e ser., 3, 399^27. cion icnologica y paleoambiental de la localidad
BOHACS, K. M, CARROLL, A. R., NEAL, J. E. & MANKIE- tipo de Orchesteropus atavus, Huerta de Huachi,
wicz, P. J. 2000. Lake-basin type, source poten- provincia de San Juan, Argentina. Ameghiniana,
tial, and hydrocarbon character: an integrated 40, 53-70.
sequence-stratigraphic-geochemical framework. BUATOIS, L. A., MANGANO, M. G., Wu, X. & ZHANG,
In: GiERLOwsKi-KoRDESCH, E. & KELTS, K. (eds) G. 1996a. Trace fossils from Jurassic lacustrine
Lake Basins through Space and Time. American turbidites of the Anyao Formation (central
Association of Petroleum Geologists, Studies in China) and their environmental and evolutionary
Geology, Tulsa, Oklahoma, 46, 3-34. significance. Ichnos, 4, 287-303
BRACKEN, B. & PICARD, M. D. 1984. Trace fossils BUATOIS, L. A., MANGANO, M. G. & ACENOLAZA, F. G.
from Cretaceous/Tertiary North Horn Formation 1996b. Icnofaunas paleozoicas en sustratos firmes
in central Utah. Journal of Paleontology, 58, 477- no marinos: Evidencias del Permico de la cuenca
487. Paganzo. Ameghiniana, 33, 265-270.
NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 329

BUATOIS, L. A., JALFIN, G. & ACENOLAZA, F. G. 1997. Lodeve (sud du Massif central). Geologie de la
Permian nonmarine invertebrate trace fossils from France, 1, 19-32.
southern Patagonia, Argentina: ichnologic signa- EBERTH, D. A., BERMAN, D. S., SUMIDA, S. S. & HOPF,
tures of substrate consolidation and colonization H. 2000. Lower Permian terrestrial paleo-
sequences. Journal of Paleontology, 71, 324—336. environments and vertebrate paleoecology of the
BUATOIS, L. A., MANGANO, M. G., GENISE, J. F. & Tambach Basin (Thuringia, Central Germany):
TAYLOR, T. N. 1998. The ichnologic record of the Upland Holy Grail. Palaios, 15, 293-313.
the invertebrate invasion of nonmarine ecosys- EKDALE, A. A., BROWN, F. H. & FEIBEL, C. S. 1989.
tems: evolutionary trends in ecospace utilization, Nonmarine macroborings in Early Pleistocene
environmental expansion, and behavioral com- algal biolithites (Stromatolites) of the Turkana
plexity. Palaios, 13, 217-240. Basin, Northern Kenya. Palaios, 4, 389-396.
BUATOIS, L. A., MANGANO, M. G., FREGENAL- FITZGERALD, P. G. & BARRETT, P. J. 1986. Skolithos in
MARTINEZ, M. A. & DE GIBERT, J. M. 2000a. a Permian braided river deposit, southern Victoria
Short-term colonization trace-fossil assemblages Land, Antarctica. Palaeogeography, Palaeoclima-
in a carbonate lacustrine konservat-lagerstatte tology, Palaeoecology, 52, 237-247.
(Las Hoyas fossil site, Lower Cretaceous, FORDYCE, R. W. 1980. Trace fossils from Ohika
Cuenca, central Spain). Fades, 43, 145-156. Formation (Pororari Group, Lower Cretaceous),
BUATOIS, L. A., MANGANO, M. G., Wu, X. & ZHANG, G. lower Buller Gorge, Buller, New Zealand. Journal
2000b. Jurassic lake deposits from the Anyao For- of Geology and Geophysics, 23, 121-124.
mation, Central China. In: GIERLOWSKI-KORDESCH, FREY, R. W. & PEMBERTON, S. G. 1984. Trace fossil
E. & KELTS, K. (eds) Lake Basins through Space and facies models. In: WALKER, R. G. (ed.) Facies
Time. American Association of Petroleum Geolo- Models (2nd edn). Geoscience Canada, Reprint
gists, Studies in Geology, Tulsa, Oklahoma, 46, Series, Ontario, 1, 189-207.
189-194. FREY, R. W. & PEMBERTON, S. G. 1985. Biogenic struc-
BUATOIS, L. A., MANGANO, M. G. & NETTO, R. G. tures in outcrops and cores. I. Approaches to ich-
2001. Paleoecosistemas anactualisticos vinculados nology. Bulletin of Canadian Petroleum Geology,
a la glaciacion gondwanica: evidencias en el 33,72-115.
Paleozoico superior del oeste de Argentina y sur FREY, R. W. & PEMBERTON, S. G. 1987. The Psilonich-
de Brasil. Segundo Simposio Argentina del Paleo- nus ichnocoenose, and its relationship to adjacent
zoico Superior, Resumenes, Trelew, p. 3. marine and nonmarine ichnocoenoses along the
BUATOIS, L. A., MANGANO, M. G. & ACENOLAZA, F. G. Georgia coast. Bulletin of Canadian Petroleum
2002. Trazasfosiles: Senales de comportamiento en Geology, 35, 333-357.
el registro estratigrdfico. Museo Paleontologico FREY, R. W. & SEILACHER, A. 1980. Uniformity in marine
Egidio Feruglio, Edition Especial, 2. invertebrate ichnology. Lethaia, 13, 183-207.
CLEMMENSEN, L. B., KENT, D. V. & JENKINS, F. A. JR. FREY, R. W., PEMBERTON, S. G. & FAGERSTROM, J. A.
1998. A Late Triassic lake system in East Green- 1984. Morphological, ethological, and environ-
land: facies, depositional cycles and paleoclimates. mental significance of the ichnogenera Scoyenia
Palaeogeography, Palaeoclimatology, Palaeoecol- and Ancorichnus. Journal of Paleontology, 58,
ogy, 140, 135-159. 511-528.
COOK, A. G. & BANN, K. 2000. Trace fossils from the FREY, R. W., PEMBERTON, S. G. & SAUNDERS, T. D. A.
Upper Carboniferous Jericho Formation, Central 1990. Ichnofacies and bathymetry: a passive rela-
Queensland. Memoirs of the Queensland Museum, tionship. Journal of Paleontology, 64, 155-158.
45, 235-251. FUGLEWICZ, R., PTASZYNSKI, T. & RDZANEK, K. 1996.
CRIMES, T. P. 1977. Trace fossils of an eocene deep-sea Lower Triassic footprints from the Swietokrzyskie
sand fan. In: CRIMES, T. P. & HARPER, J. C. (eds) (Holy Cross) Mountains, Poland. Ada Paleontolo-
Trace Fossils 2. Geological Journal Special Issue, gica Polonica, 35, 109-164.
9, 71-90. FURSICH, F. T. & MAYR, H. 1981. Non-marine Rhizo-
D'ALESSANDRO, A., EKDALE, A. A. & PlCARD, M. D. corallium (trace fossil) from the Upper Freshwater
1987. Trace fossils in fluvial deposits of the Molasse (Upper Miocene) of southern Germany.
Duchesne River Formation (Eocene), Uinta Neues Jahrbuch fur Geologie und Palaontologie,
Basin, Utah. Palaeogeography, Palaeoclimatology, Monashefte, 1981, 321-333.
Palaeoecology, 61, 285-301. GAND, G. & HAUBOLD, H. 1984. Traces de Vertebres du
DALEY, B. 1968. Sedimentary structures from a non- Permien du Bassin de Saint-Affrique (Description,
marine horizon in the Bembridge Marls (Oligo- datation, comparaison avec delles du bassin de
cene) of the Isle of Wight, Hampshire, England. Lodeve). Geologie Mediterraneenne, 4, 321-348.
Journal of Sedimentary Petrology, 38, 114-127. GAND, G., KERP, H., PARSONS, C. & MARTINEZ
DAM, G. & STEMMERIK, L. 1994. East Greenland lacus- GARciA, E. 1997. Palaeoenvironmental and strati-
trine complexes. In: GIERLOWSKI-KORDESH, E. & graphic aspects of animal traces and plant remains
KELTS, K. (eds) Global Geological Record of Lake in Spanish Permian red beds (Pena Sagra, Cantab-
Basins, 1. Cambridge University Press, Cambridge. rian Mountains, Spain). Geobios, 30, 295-318.
DEBRIETTE, P. & GAND, G. 1990. Consequences strati- GENISE, J. F. & BOWN, T. M. 1994a. New Miocene
graphiques et paleoenvironnementales de nou- scarabeid and hymenopterous nests and Early
velles observations paleontologiques dans le Miocene (Santacrucian) paleoenvironments,
Permien de la partie occidentale du bassin de Patagonian Argentina. Ichnos, 3, 107-117.
330 L. A. BUATOIS & M. G. MANGANO

GENISE, J. F. & BOWN, T. M. 1994b. New trace fossils sequences: the Newark Supergroup (Triassic-
of termites (Insecta: Isoptera) from the Late Jurassic), eastern North America. Palaeogeogra-
Eocene-Early Miocene of Egypt, and the recon- phy, Palaeoclimatology, Palaeoecology, 70, 29-51.
struction of ancient isopteran social behavior. GRAHAM, J. R. & POLLARD, J. E. 1982. Occurrence of
Ichnos, 3, 155-183. the trace fossil Beaconites antarcticus in the
GENISE, J. F., MANGANO, M. G., BUATOIS, L. A., LAZA, Lower Carboniferous fluviatile rocks of County
J. & VERDE, M. 2000. Insect trace fossil associa- Mayo, Ireland. Palaeogeography, Palaeoclimatol-
tions in paleosols: the Coprinisphaera ichnofacies. ogy, Palaeoecology, 38, 257-268.
Palaios, 15, 33-48. HASIOTIS, S. T., MITCHELL, C. E. & DUBIEL, R. F. 1993.
GENISE, J. F., BELLOSI, E. S. & GONZALEZ, M. G. 2004. Application of morphologic burrow interpreta-
An approach to the description and interpretation tions to discern continental burrow architects:
of ichnofabrics in palaeosols. In: MC!LROY, D. lungfish or crayfish? Ichnos, 2, 315-333.
(ed.) The Application of Ichnology to Palaeoenvir- HESTER, P. M. & LUCAS, S. G. 2001. Lacustrine
onmental and Stratigraphic Analysis. Geological depositional environments of the Upper Triassic
Society, London, Special Publications, 228, 355- Redonda Formation, East-Central New Mexico.
382. New Mexico Geological Society Guidebook, 52nd
GIBBARD, P. L. 1977. Fossil tracks from varved Field Conference, 153-168.
sediments near Lammi, south Finland. Bulletin HIGGS, R. 1988. Fish trails in the Upper Carboniferous
Geological Society of Finland, 49, 53-57. of south-west England. Palaeontology, 31, 55-272.
GIBBARD, P. L. & DREIMANIS, A. 1978. Trace fossils Hu, B., WANG, G. & COLORING, R. 1998. Nereites
from late Pleistocene glacial lake sediments in (or Neonereites) from Lower Jurassic lacustrine
southwestern Otario, Canada. Canadian Journal turbidites of Henan, Central China. Ichnos, 6,
of Earth Sciences, 15, 1967-1976. 203-209.
GIBBARD, P. L. & STUART, A. J. 1974. Trace KEIGHLEY, D. G. & PICKERILL, R. K. 1994. The
fossils from proglacial lake sediments. Boreas, 3, ichnogenus Beaconites and its distinction from
69-74. Ancorichnus and Taenidium. Palaeontology, 37,
DE GIBERT, J. M., MARTINELL, J. & DOMENECH, R. 305-337.
1998. Entobia ichnofacies in fossil rocky shores, KIM, J. Y & PAIK, I. S. 1997. Nonmarine Diplocraterion
Lower Pliocene, Northwestern Mediterranean. luniforme (Blanckenhorn 1916) from the Hasan-
Palaios, 13, 476^87. dong Formation (Cretaceous) of the Jinju area,
DE GIBERT, J. M., BUATOIS, L. A., FREGENAL-MARTI- Korea. Ichnos, 5, 131-138.
NEZ, M. A., MANGANO, M. G., ORTEGA, F., KOZUR, H. W. & LEMONE, D. V. 1995. New terrestrial
POYATO-ARIZA, F. J. & WENZ, S. 1999. The fish arthropod trackways from the Abo Member
trace fossil Undichna from the Cretaceous of (Sterlitamakian, late Sakmarian, late Wolf-
Spain: taphonomic and palaeoenvironmental campian) of the Shalem Colony section, Robledo
implications for the ichnogenus. Palaeontology, Mountains, New Mexico. In: LUCAS, S. G. &
42, 409^27. HECKERT, A. B. (eds) Early Permian Footprints
DE GIBERT, J. M., FREGENAL-MARTINEZ, M. A., and Fades. New Mexico Museum of Natural
BUATOIS, L. A & MANGANO, M. G. 2000. Trace History and Science, Bulletins, 6, 107-113.
fossils and their palaeoecological significance in LABANDEIRA, C. C. 1997. Early history of arthropod
Lower Cretaceous lacustrine conservation depos- and vascular plant associations. Annual Review
its, El Montsec, Spain. Palaeogeography, Palaeo- of Earth and Planetary Sciences, 26, 329-377.
climatology, Palaeoecology, 156, 89-101. LABANDEIRA, C. C. 1998. Plant-insect associations
GIERLOWSKI-KORDESCH, E. 1991. Ichnology of an from the fossil record. Geotimes.
ephemeral lacustrine/alluvial plain system: Juras- LABANDEIRA, C. C. 2002. The history of associations
sic East Berlin Formation, Hartford Basin, USA. between plants and animals. In: HERRERA, C. &
Ichnos, 1, 221-232. PELLMYR, O. (eds) History of Plant-Animal Inter-
GIERLOWSKI-KORDESCH, E. & KELTS, K. 1994. Global actions. Blackwell Science, Oxford, 26-74 & 248-
Geological Record of Lake Basins 1. Cambridge 261.
University Press, Cambridge. LEGARRETA, L. & ULIANA, M. A. 1998. Anatomy of
GIERLOWSKI-KORDESCH, E. & KELTS, K. 2000. Lake hinterland depositional sequences: Upper Cretac-
Basins Through Space and Time. American Asso- eous fluvial strata, Neuquen Basin, West-Central
ciation of Petroleum Geologists, Studies in Argentina. In: SHANLEY, K. W. & McCABE, P. J.
Geology, Tulsa, Oklahoma, 46. (eds) Relative Role of Eustasy, Climate, and
GLUSZEK, A. 1995. Invertebrate trace fossils in the Tectonism in Continental Rocks. SEPM Special
continental deposits of an Upper Carboniferous Publications, Tulsa, Oklahoma, 9, 83-92.
coal-bearing succession, Upper Silesia, Poland. LEGARRETA, L., ULIANA, M. A., LAROTONDA, C. A. &
Studia Geologica Polonica, 108, 171-202. MECONI, G. R. 1993. Approaches to nonmarine
GOLDRING, R. & POLLARD, J. E. 1995. A re-evaluation sequence stratigraphy: theoretical models and
of Ophiomorpha burrows in the Wealden Group examples from Argentine basins. In: ESCHARD,
(Lower Cretaceous) of southern England. Cretac- R. & DOLIEZ, B. (eds) Subsurface Reservoir Char-
eous Research, 16, 665-680. acterisation from Outcrop Observations. Editions
GORE, P. J. W. 1989. Toward a model for open- and Technip, Paris, Collection Colloques et Semi-
closed-basin deposition in ancient lacustrine naires, 51, 125-145.
NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 331

LEONARDI, G. 1989. Inventory and statistics of the MANGANO, M. G., LABANDEIRA, C., KVALE, E. &
South American dinosaurian ichnofauna and its BUATOIS, L. A. 2001. The insect trace fossil
paleobiological interpretation. In: GILLETTE, Tonganoxichnus from the Middle Pennsylvanian
D. D. & LOCKLEY, M. G. (eds) Dinosaur Tracks of Indiana: paleobiologic and paleoenvironmental
and Traces. Cambridge University Press, New implications. Ichnos, 8, 165-175.
York, 165-178. MAPLES, C. G. & ARCHER, A. W. 1989. The potential
LIM, S. K., YANG, S. Y. & LOCKLEY, M. G. 1989. Large of Paleozoic nonmarine trace fossils for paleo-
dinosaur footprint assemblages from the Cretac- ecological interpretations. Palaeogeography,
eous Jindong Formation of southern Korea. In: Palaeoclimatology, Palaeoecology, 73, 185-195.
GILLETTE, D. D. & LOCKLEY, M. G. (eds) Dinosaur MclLROY, D. 2004. In: MC!LROY, D. (ed.) The Applica-
Tracks and Traces. Cambridge University Press, tion of Ichnology to Palaeoenvironmental and
New York, 333-336. Stratigraphic Analysis. Geological Society,
LOCKLEY, M. G. 1986. The paleobiological and London, Special Publications, 228, 3-27.
paleoenvironmental importance of dinosaur foot- MELCHOR, R. N. 2001. Icnologia y sedimentologia de
prints. Palaios, 1, 37^47. una sucesion lacustre influenciada por tormentas:
LOCKLEY, M. G. 1989. Summary and prospectus. In: Formation Los Rastros (Triasico), Talampaya,
GILLETTE, D.D. & LOCKLEY, M.G. (eds) Dinosaur La Rioja. IV Reunion Argentina de Icnologia y
Tracks and Traces. Cambridge University Press, Segunda Reunion de Icnologia del Mercosur, Resu-
New York, 441^47. menes, San Miguel de Tucuman, 56.
LOCKLEY, M. G. 1991. Tracking Dinosaur: A New Look MELCHOR, R. N. 2004. Trace fossil distribution in
at an Ancient World. Cambridge University Press, lacustrine deltas: examples from the Traissic rift
Cambridge. lakes of the Ishigualasto-Villa Union basin,
LOCKLEY, M. G., HOUCK, K. J. & PRINCE, N. K. 1986. Argentina. In: MC!LROY, D. (ed.) The Application
North America's largest dinosaur trackway site: of Ichnology to Palaeoenvironmental and Strati-
implications for Morrison Formation paleo- graphic Analysis. Geological Society, London,
ecology. Geological Society of America, Bulletin, Special Publications, 228, 335-354.
97, 1163-1176. MELCHOR, R. N., BELLOSI, E. & GENISE, J. F. 2003.
LOCKLEY, M. G., LOGUE, T. J., MORATALLA, J. J., Invertebrate and vertebrate trace fossils from a
HUNT, A. P., SCHULTZ, R. J. & ROBINSON, J. W. lacustrine delta: the Los Rastros Formation, Ischi-
1995. The fossil trackway Pteraichnus is ptero- gualasto Provincial Park, San Juan, Argentina. In:
saurian, not crocodolian: implications for the BUATOIS, L. A. & MANGANO, M. G. (eds) Icno-
global distribution of pterosaur tracks. Ichnos, 4, logia: Hacia una Convergencia entre Geologia y
7-20. Biologia. Asociacion Paleontologica Argentina,
MACEACHERN, J. A., RAYCHAUDHURI, I. & PEMBERTON, Publication Especial, Buenos Aires, 9, 17-33.
S. G. 1992. Stratigraphic applications of the METZ, R. 1993. A new ichnospecies of Spongeliomor-
Glossifungites Ichnofacies: delineating discontinu- pha from the Late Triassic of New Jersey. Ichnos,
ities in the rock record. In: PEMBERTON, S. G. (ed.) 2, 259-262.
Applications of Ichnology to Petroleum Explora- METZ, R. 1995. Ichnologic study of the Lockatong
tion. Society of Economic Paleontologists and Formation (Late Triassic), Newark Basin, south-
Mineralogists, Core Workshops, Tulsa, Okla- eastern Pennsylvania. Ichnos, 4, 43-51.
homa, 17, 169-198. METZ, R. 1996. Newark Basin ichnology: the Late
MACNAUGHTON, R. B. & PlCKERILL, R. K. 1995. Triassic Perkasie Member of the Passaic Forma-
Invertebrate ichnology of the nonmarine Lepreau tion, Sanatoga, Pennsylvania. Northeastern Geol-
Formation (Triassic), southern New Brunswick, ogy and Environmental Sciences, 18, 118-129.
Eastern Canada. Journal of Paleontology, 69, METZ, R. 2000. Triassic trace fossils from lacustrine
160-171. shoreline deposits of the Passaic Formation,
MANGANO, M. G., BUATOIS, L. A., Wu, X., SUN, J. & Douglassville, Pennsylvania. Ichnos, 7, 253-266.
ZHANG, G. 1994. Sedimentary facies, depositional MIALL, A. D. 1996. The Geology of Fluvial Deposits:
processes and climatic controls in a Triassic Sedimentary Facies, Basin Analysis, and Petroleum
lake, Tanzhuang Formation, western Henan Geology. Springer-Verlag, Berlin.
Province, China. Journal of Paleolimnology, 11, MIKULAS, R. 2003. The Mermia ichnofacies across the
41-65. fossilization barrier: a comparison of the Permian
MANGANO, M. G., BUATOIS, L. A., MAPLES, C. G. & Krkonoe Piedmont Basin (Czech Republic) and
LANIER, W. P. 1997. Tonganoxichnus, a new modern flood sediments at Prague. VII Inter-
insect trace fossil from the Upper Carboniferous national Ichnofabric Workshop, Abstracts, Basel,
of eastern Kansas, USA. Lethaia, 30, 113-125. 44-45.
MANGANO, M. G., BUATOIS, L. A., Wu X., SUN, J. & MILLER, G. D. 1986. The sediments and trace fossils of
ZHANG, G. 2000. Triassic lacustrine sedimentation the Rough Rock Group on Cracken Edge, Derby-
from the Tanzhuang Formation, Jiyuan-Yima shire. Mercian Geologist, 10, 189-202.
Basin, southeastern China. In: GIERLOWSKI- MILLER, M. F. 2000. Benthic aquatic ecosystems across
KORDESCH, E. & KELTS, K. (eds) Global Geological the Permian-Triassic transition: record from bio-
Record of Lake Basins. American Association of genie structures in fluvial sandstones, central
Petroleum Geologists, Studies in Geology, Tulsa, Transantarctic Mountains. Journal of African
Oklahoma, 46, 133-140. Earth Sciences, 31, 157-164.
332 L. A. BUATOIS & M. G. MANGANO

MILLER, M. F. & COLLINSON, J. W. 1994. Trace fossils POLLARD, J. E. 1981. A comparison between the
from Permian and Triassic sandy braided stream Triassic of Cheshire and south Germany. Palaeon-
deposits, central Transantarctic Mountains. tology, 24, 555-588.
Palaios, 9, 605-610. POLLARD, J. E. & HARDY, P. G. 1991. Trace fossils from
MILLER, M. F. & LABANDEIRA, C. C. 2003. Slow crawl the Westphalian D of Writhlington Geologica
across the salinity divide: delayed colonization of Nature Reserve, nr. Radstock, Avon. Geologists'
freshwater ecosystems by invertebrates. GSA Association Proceedings, 102, 169-178.
Today, 12, 4-10. POLLARD, J. E. & LOVELL, J. P. B. 1976. Trace fossils
MILLER, M. F., COLLINSON, J. W. & FRISCH, R. A. 1991. from the Permo-Triassic of Arran. Scottish
Depositional setting and history of a Permian post- Journal of Geology, 12, 209-225.
glacial black shale: Mackellar Formation, Central POLLARD, J. E. & WALKER, E. 1984. Reassessment of
Transantarctic Mountains. In: ULBRICH, H. & sediments and trace fossils from Old Red Sand-
ROCHA CAMPOS, A. C. (eds) Gondwana Seven: stone (Lower Devonian) of Dunure, Scotland,
Proceedings Seventh International Gondwana described by John Smith (1909). Geobios, 17,
Symposium, Sao Paulo, 201-215. 567-576.
MILLER, W. III. 1991. Paleoecology of graphoglyptids. POLLARD, J. E., STEEL, R. J. & UNDERSRUD, E. 1982.
Ichnos, 1, 305-312. Facies sequences and trace fossils in lacustrine/
MORATALLA, J. J., LOCKLEY, M. et al. 1995. A prelimin- fan-delta deposits, Hornelen Basin (M. Devo-
ary note on the first tetrapod trackways from the nian), western Norway. Sedimentary Geology, 32,
lithographic limestones of Las Hoyas (Lower 63-87.
Cretaceous, Spain). Geobios, 28, 777-782. POSAMENTIER, H. W. & ALLEN, G. P. 1999. Siliciclastic
OLSEN, P. E. 1989. Newark Basin, New Jersey. In: Sequence Stratigraphy: Concepts and Applications.
OLSEN, P. E., SCHLICHE, R. & GORE, P. J. (eds) SEPM Concepts in Sedimentology and Paleontol-
Tectonic, Depositional, and Palaeoecological ogy, Tulsa, Oklahoma, 7.
History of early Mesozoic Rift Basins, Eastern PRICE, S. & McCANN, T. M. 1990. Environmental
North America. International Geological Congress significance of Arenicolites ichnosp. in Pliocene
Field Trip Guidebooks, Washington DC, T-351, 2. lake deposits of southwest Turkey. Neues Jahrbuch
OLSEN, P. E., REMINGTON, C. L. & CORNET, B. 1978. fur Geologie und Palaontologie Monatshefte, 1990,
Cyclic change in Late Triassic lacustrine commu- 687-694.
nities. Science, 201, 729-732. PRINCE, N. K. & LOCKLEY, M. G. 1989. The sedimen-
PAZOS, P. J. 2002. Palaeoenvironmental framework of tology of the Purgatoire tracksite region, Morri-
the glacial-postglacial transition (Late Paleozoic) son Formation of southeastern Colorado. In:
in the Paganzo-Calingasta Basin (southern South GILLETTE, D. D. & LOCKLEY, M. G. (eds) Dinosaur
America) and the Great Karoo-Kalahari Basin Tracks and Traces. Cambridge University Press,
(southern Africa): ichnological implications. New York, 155-163.
Gondwana Research, 5, 619-640. RATCLIFFE, B. C. & FAGERSTROM, J. A. 1980.
PEMBERTON, S. G. & FREY, R. W. 1984. Ichnology of Invertebrate lebensspuren of Holocene flood-
storm-influenced shallow marine sequence: plains: their morphology, origin and paleoeco-
Cardium Formation (Upper Cretaceous) at logical significance. Journal of Paleontology, 54,
Seebe, Alberta. In: STOTT, D. F. & GLASS, D. J. 614-630.
(eds) The Mesozoic of Middle North America. RODRIGUEZ-ARANDA, J. P. & CALVO, J. P. 1998. Trace
Canadian Society of Petroleum Geologists, fossils and rhizoliths as a tool for sedimentological
Memoirs, Ontario, 9, 281-304. and palaeoenvironmental analysis of ancient
PEMBERTON, S. G. & FREY, R. W. 1985. The Glossi- continental evaporite successions. Palaeogeogra-
fungites ichnofacies: modern examples from the phy, Palaeoclimatology, Palaeoecology, 140, 383-
Georgia coast, USA. In: CURRAN, H. A. (ed.) 399.
Biogenic Structures: Their Use in Interpreting SARKAR, S. & CHAUDHURI, A. K. 1992. Trace fossils in
Depositional Environments. Society of Economic Middle Triassic fluvial redbeds, Pranhita-
Paleontologists and Mineralogists, Special Godavari Valley, south India. Ichnos, 2, 7-19.
Publications, Tulsa, Oklahoma, 35, 237-259. SAVRDA, C. E., BLANTON-HOOKS, A. D. et al. 2000.
PEMBERTON, S. G., SPILA, M., PULHAM, A. J., SAUN- Taenidium and associated ichnofossils in fluvial
DERS, T., MAC£ACHERN, J. A., ROBBINS, D. & deposits, Cretaceous Tuscaloosa Formation,
SINCLAIR, L K. 2001. Ichnology and Sedimentology Eastern Alabama, Southeastern USA. Ichnos, 7,
of Shallow to Marginal Marine Systems. Ben Nevis 223-242.
& Avalon Reservoirs, Jeanne d'Arc Basin. Geo- SCHLIRF, M., UCHMAN, A. & KtJMMEL, M. 2001.
logical Association of Canada, Short Course Upper Triassic (Keuper) non-marine trace
Notes, Quebec, 15, 1-343. fossils from the HaBberge area (Franconia,
PHARO, C. H. & CARMACK, E. C. 1979. Sedimentation south-eastern Germany). Palaontologische Zeits-
processes in a short residence-time intermontane chrift, 75, 71-96.
lake, Kamloops Lake, British Columbia. Sedimen- SEILACHER, A. 1963. Lebensspuren und Salinitatsfazies.
tology,26,523>-54\. Fortschritte in der Geologie Rheinland und West-
PICKERILL, R. K. 1992. Carboniferous nonmarine falens, 10, 81-94.
invertebrate ichnocoenoses from southern New SEILACHER, A. 1967. Bathymetry of trace fossils.
Brunswick, eastern Canada. Ichnos, 2, 21-35. Marine Geology, 5, 413-428.
NON-MARINE PALAEOENVIRONMENTS & ICHNOLOGY 333

SHANLEY, K. W. & McCABE, P. J. 1998. Relative role of Teruel Basin, NE Spain. Revista Espanola de
eustasy, climate, and tectonism in continental Paleontologia, 15, 203-218.
rocks: an introduction. In: SHANLEY, K. W. & VAN WAGONER, J. C., MITCHUM, R. M., CAMPION,
McCABE, P. J. (eds) Relative Role of Eustasy, K. M. & RAHMANIAN, V. D. 1990. Silicidastic
Climate, and Tectonism in Continental Rocks. Sequences, Stratigraphy in Well Logs, Cores, and
SEPM Special Publications, Tulsa, Oklahoma, 9, Outcrops. American Association of Petroleum
iii-iv. Geologists, Methods in Exploration Series,
SMITH, R. M. H. 1993. Sedimentology and ichnology of Tulsa, Oklahoma, 7.
floodplain paleosurfaces in the Beaufort Group WALKER, E. 1985. Arthropod ichnofauna of the Old
(Late Permian), Karoo Sequence, South Africa. Red Sandstone at Dunure and Montrose,
Palaios, 8, 339-357. Scotland. Transactions of the Royal Society of
SMITH, R. M. H., MASON, T. R. & WARD, J. D. 1993. Edinburgh: Earth Sciences, 76, 287-297.
Flash-flood sediments and ichnofacies of the WALTER, H. 1985. Zur ichnologie des Pleistozans von
Late Pleistocene Homeb Silts, Kuiseb River, Liebegast. Freiberger Forschungssheft, C 400 S,
Namibia. Sedimentary Geology, 85, 579-599. 101-116.
SQUIRES, R. L. & ADVOCATE, D. M. 1984. Meniscate WALTER, H. & SUHR, P. 1998. Lebesspuren aus kaltzei-
burrows from Miocene lacustrine fluvial-deposits, tlichen Bandersedimenten des Quartars. Abhan-
Diligencia Formation, Orocopia Mountains, dlungen des Staatlichen Museums fur Mineralogie
southern California. Journal of Paleontology, 58, und Geologie zu Dresden, 43/44, 311—328.
593-597. WELLS, R. F. 1977. Fresh water invertebrate living traces
STANLEY, K. O. & FAGERSTROM, J. A. 1974. Miocene of the Mississippi alluvial valley near Baton Rouge.
invertebrate trace fossils from a braided river MSc thesis, Lousiana State University.
environment, western Nebraska, USA. Palaeo- WOOLFE, K. J. 1990. Trace fossils as paleoenvironmen-
geography, Palaeoclimatology, Palaeoecology, 15, tal indicators in the Taylor Group (Devonian) of
63-82. Antarctica. Palaeogeography, Palaeoclimatology,
TUREK, V. 1989. Fish and amphibian trace fossils from Palaeoecology, 80, 301-310.
Westphalian Sediments of Bohemia. Palaeontol- YANG, S. Y., LOCKLEY, M. G., GREBEN, R., ERICKSON,
ogy, 32, 623-634. B. R. & LIM, S. K. 1995. Flamingo and duck-bird
TURNER, B. R. 1978. Trace fossils from the Upper tracks from the Late Cretaceous and Early
Triassic fluviatile Molteno Formation of the Tertiary: evidence and implications. Ichnos, 4,
Karoo (Gondwana) Supergroup, Lesotho. Journal 21-34.
of Paleontology, 52, 959-963. ZAWISKIE, J. M., COLLINSON, J. W. & HAMMER, W. R.
UCHMAN, A. 1995. Taxonomy and paleoecology of 1983. Trace fossils of the Permian-Triassic
flysch trace fossils: The Marnoso-arenacea Forma- Takrouna Formation, northern Victoria Land,
tion and associated facies (Miocene, Northern Antarctica. In: OLIVER, R. L., JAMES, P. R. &
Apennines, Italy). Beringeria, 15, 1-116. JAGO, J. B. (eds) Antarctic Earth Science. Austra-
UCHMAN, A. 2004. Phanerozoic history of deep-sea lian Academy of Science, Canberra, 215-220.
trace fossils. In: MC!LROY, D. (ed.) The Application ZHANG, G., BUATOIS, L. A., MANGANO, M. G. &
of Ichnology to Palaeoenvironmental and Strati- ACENOLAZA, F. G. 1998. Sedimentary facies and
graphic Analysis. Geological Society, London, environmental ichnology of a ?Permian playa-
Special Publications, 228, 125-139. lake complex in western Argentina. Palaeogeo-
UCHMAN, A. & ALVARO J. J. 2000. Non-marine inverte- graphy, Palaeoclimatology, Palaeoecology, 138,
brate trace fossils from the Tertiary Calatayud- 221-243.
This page intentionally left blank
Trace fossil distribution in lacustrine deltas: examples from the Triassic
rift lakes of the Ischigualasto-Villa Union basin, Argentina

RICARDO N. MELCHOR

Consejo Nacional de Investigaciones Cientificas y Tecnicas (CONICET) & Universidad


Nacional de La Pampa, Av. Uruguay 151, L6300CLB Santa Rosa, La Pampa, Argentina
(e-mail: rmelchor@exactas.unlpam.edu.ar)

Abstract: This paper reports six trace fossil assemblages from lacustrine deltas of the Triassic
Ischigualasto-Villa Union rift basin, northwest Argentina. They were recognized in three
correlated stratigraphic sections separated by about 100km, and come from river- and
wave-influenced deltas developed in low- and high-accommodation lacustrine basins.
Trace fossil assemblages correspond to delta front (six), delta plain (two) and marginal
lacustrine (one) facies associations. Each trace fossil assemblage is described, together with
a detailed lithofacies characterization of the trace fossil-bearing interval. They are analysed
in conjunction with previously described ichnological assemblages from partially correlative
sections of the same basin. Delta front facies contains a high-diversity assemblage (22 ichno-
taxa), including Cochlichnus (a ubiquitous form), Gordia, Helminthoidichnites, Helminthopsis,
Didymaulichnus, Diplichnites, Stiaria, Cruziana, Bifurculapes, Protichnites, Diplopodichnus,
Archaeonassa, Palaeophycus, Treptichnus, Rusophycus, Avolatichnium, 'rhomboidal traces',
'fusiform structures' and 'millimetre burrows'. Trace fossil assemblages from delta plain
facies are much less diverse (seven ichnotaxa), but display representatives of a greater variety
of ethological categories, including Rhynchosauroides, Skolithos, Palaeophycus, 'horseshoe-
shaped structures', escape trace and drab-haloed root traces. Marginal lacustrine deposits
of a river-dominated delta yielded a monospecific assemblage consisting of Cochlichnus
anguineus. Comparison of trace fossil assemblages in wave- and river-dominated lacustrine
deltas from the basin (mainly those of the delta front facies) revealed important differences
in ichnodiversity that can be useful in the discrimination between these lacustrine delta
types. Lacustrine delta deposits contain trace fossils that can be ascribed to three different
ichnofacies: a high-diversity occurrence of the Mermia ichnofacies in subaqueous delta
front sediments, a low-diversity occurrence of Mermia ichnofacies in subaqueous marginal
lacustrine facies, the Skolithos ichnofacies in high-energy upper delta front/lower delta
plain facies, and the Scoyenia ichnofacies in intermittently exposed upper delta plain
facies. The analysed trace fossil assemblages from delta front and marginal lacustrine settings
suggest environmental gradients within the Mermia ichnofacies.

The understanding of trace-fossil distribution in occurrences of the Skolithos ichnofacies (Man-


lacustrine basins has witnessed important gano et al. 1994; Buatois & Mangano 1995,
advances recently, through the identification of 1998; Melchor et al. 2003). Intermittently
the Mermia ichnofacies for fully subaqueous emergent shallow-lacustrine settings include
freshwater environments (Buatois & Mangano ichnofossils on softground and firmground
1995) and the improved documentation of trace (desiccated) substrates that are best ascribed to
fossil distributions in different lacustrine envir- the emended Scoyenia ichnofacies (Buatois &
onments, including evaporitic lacustrine basins Mangano 1995; Metz 1996; Melchor et al. 2003).
(Rodriguez-Aranda & Calvo 1998), shorelines At present, there is scarce documentation of
(e.g. Lockley et al. 1992, 1994; Metz 1996; the ichnofossil distribution in particular sedi-
Doyle et al. 2000; Kim et al. 2002) and floodplain mentary facies of freshwater deltas. Table 1
lakes (e.g. Buatois et al. 1997; Buatois & Man- contains a summary of the published ichnologic
gano 2002). The Mermia ichnofacies typifies information and trace fossil distribution in this
fine-grained sediments from well-oxygenated, environment. The examples of Permian post-
low-energy, permanently subaqueous zones of glacial sequences of South Africa, the Falkland
lacustrine systems including floodplain lakes Islands and probably Antarctica, where no
and the landward, freshwater part of fjords agreement about the salinity of the lake basin is
(Buatois & Mangano 1995, 1998, 2002, 2003). available (e.g. Kingsley 1981; Miller et al. 1991;
Trace fossils in high-energy settings of lacustrine Miller & Smail 1996; Seegers-Szablewski &
basins (e.g. wave-dominated shorelines, delta Isbell 1997; Trewinef al. 2002), must be regarded
mouth-bars) have been attributed to continental only as possible case studies. Some authors

From\ MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 335-354. 0305-8719/04/S15.00 © The Geological Society
of London.
Table 1. Trace fossil assemblages of lacustrine deltas and related environments

Age Unit/basin Country /region Depositional Ethology-diversity Ichnofossils Source


environment

Recent Atchafalaya Basin Mississippi delta Mouth-bar Unknown Roots traces and burrows Tye & Coleman
plain Delta front Unknown Significantly less bioturbation than (1989)
prodelta. Bioturbation decreases upward
Prodelta Unknown Highly bioturbated (>75% volume)
Recent Lake Manyara Tanzania Upper delta plain Moderate Seven kinds of mammalian track and one Cohen et al. (1991 ,
diversity, bird track type. Insect burrows 1993)
locomotion tracks
Pleistocene Saxony Germany Proximal glacio- Locomotion and Fine trails and burrows, including Walter & Suhr (1998)
lacustrine delta front grazing traces Cochlichnus
Distal glacio- Arthropod Warvichnium, Glaciichnium,
lacustrine delta front locomotion traces, Lusatichnium, fine trails and burrows
grazing traces
Open lacustrine Locomotion and Cochlichnus (dominant), fine trails
grazing traces
Miocene Lan Krabu Fm Thailand Mouth-bar unknown Bioturbation decreases toward the top Flint et al. (1989)
Lower mouth-bar to Unknown Isolated vertical burrows
prodelta
Open lacustrine Bioturbation s.L
Lower Jurassic Anyao Fm China Sublacustrine fan Moderate Cochlichnus, Helminthoidichnites, Buatois et al. (1996)
lobes diversity, feeding Helminthopsis, Monomorphichnus,
& grazing traces of Paracantoraphe, Vagorichnus,
deposit-feeding Tuber culichnus, 'tiny grazing trails',
organisms. 'irregularly branching burrows'
Burrows dominate
Lower Jurassic Portland Fm USA Sandstone delta lobes McDonald &
Bivalve escape traces, fish coprolites
(s.L) LeTorneau (1988)
Permian-Triassic Beaufort Gr South Africa Lacustrine shelf Arthropod trackways, burrows and trails Van Dijk et al. (1978)
of deposit-feeding organisms, coprolites
Upper prodelta/lower 'Scolicitf (= Archaeonassal}, Undichna,
delta front arthropod trackways, horizontal spreiten
structures, coprolites
Mouth-bar Rootlets and rare vertical burrows
Interdistributary bays Rootlets, arthropod trackways, 'minute
burrows', leaf mines
Permian Ripon Fm South Africa/ Delta-fed turbidites Low diversity, low Umfolozia, Undichna, 'small meandering Kingsley (1981)
Eastern Cape density, grazing trails'
Province locomotion and
grazing traces
Fort Brown Fm South Africa/ Prodelta/delta front Low diversity, Undichna, 'gastropod trails', 'unidentified Kingsley (1981)
Eastern Cape moderate density, burrows'
Province locomotion,
grazing and
?dwelling traces
Upper Permian Brenton Loch Falkland Islands Distal turbidite lobes Undichna ispp., Planolites, Trewin et al (2002)
(Cantera Mb) Diplocraterion, vertical and inclined
burrows with spreite
Proximal turbidite High diversity, Umfolozia, Koupichnium, Cochlichnus,
lobes high density, Haplotichnus , Helminthoidichnites,
locomotion traces Spirodesmos, Planolites, arthropod
dominant resting traces
Brenton Loch Delta slope Undichna ispp., Planolites, Spirodesmos,
(Saladero Mb) Diplocraterion
Lower Permian Mackellar Fm Transantarctic Turbidite fan lobes Surface grazing Mermia, Cochlichnus, Isopodichnus Miller et al (1991);
Mountains (= Cruziana) Miller & Smail (1996)
Delta front Low diversity Isopodichnus (— Cruziana) Seegers-Szablewski &
Isbell (1997)
Upper Carboniferous Pagoda Fm Transantarctic Shallow glacio- Very low diversity, Planolites, Palaeophycus, Isopodichnus Isbell et al (2001)
- Lower Permian Mountains lacustrine delta low to high density (= Cruziana)
feeding and
grazing traces
Upper Carboniferous Agua Colorada Argentina Delta-related Low diversity, Helminthoidichnites, Mermia Buatois & Mangano
Fm turbidite lobes. moderate density (1993a)
Episodic deposition surface grazing
trails
Delta-related High diversity and Gordia, Haplotichnus,
turbidite lobes. density, surface Helminthoidichnites, Helminthopsis,
Underflow currents grazing and Mermia, Or Chester opus, Undichna ispp.,
locomotion traces Treptichnus
Deep lacustrine. High diversity and Circulichnus, Cochlichnus, Gordia,
Muddy turbidity density, dominant Helminthoidichnites, Helminthopsis,
currents surface grazing Mermia, Or Chester opus, Treptichnus,
'string pits', 'rhomboidal traces'
Middle Devonian Hornelen Basin Norway Fan-delta (high- Siskemia, Merostomichnites, Pollard et al (1982);
sinuosity distributary Diplopodichnus, 'bilobed trails', 'ribbon Keighley & Pickerill
channels) trails', 'fine sinuous trails' (1996)
338 R. N. MELCHOR

consider that these sequences were related to a the same basin (Melchor et al. 2003) and
large lake (e.g. Kingsley 1981; Trewin et al. with case studies from the literature;
2002), and other researchers envisage the basin to ascribe the described trace fossil assem-
as a brackish sea (e.g. Visser 1993; Pazos 2002) blages to archetypical ichnofacies; and
or a sea with normal marine salinity (e.g. Stanis- to assess possible ichnological signatures of
treet et al. 1980; Johnson et al. 2001). In addition, different environments within lacustrine
the comparison of ichnofossil assemblages of deltas.
lacustrine deltas of different ages listed in Table
1 should consider the secular variations in the The described examples come from highstand
extent and depth of bioturbation and beha- deltas of flexural-margin and accommodation
vioural complexity recorded in the continental zone margin of the half-graben, including wave-
ichnofossil record (Buatois et al. 1998). dominated successions and river-dominated
The most diverse and best-documented trace deltas that prograded into either high-accom-
fossil assemblages in lacustrine deltas are modation anoxic or low-accommodation well-
delta-fed turbiditic lobes, followed by shal- oxygenated lacustrine basins.
lower-water upper delta plain and shoreline
assemblages (Table 1). On the other hand, there
are few records of trace fossil associations from Geological setting
lacustrine delta front settings. This scarcity of
ichnological studies in lacustrine deltas contrasts The Ischigualasto-Villa Union Basin from north-
with the extensive documentation of trace fossil west Argentina is one of the NW-SE-trending
distributions in marine deltas (e.g. Eagar et al. rifts developed on the west margin of south-
1985; Moslow & Pemberton 1988; Pollard 1988; western Gondwana during the Early Triassic
Coates & MacEachern 1999; Bann & Fielding (Uliana & Biddle 1988; Uliana et al. 1989;
2004; Mcllroy 2004). Pollard (1988) found Tankard et al. 1995; Fig. la). The basin fill is
some recurrent ichnocoenoses in inter- entirely continental and reaches a maximum
distributary bay and mouth-bar or crevasse thickness of approximately 4000 m (e.g. Milana
splay sediments of deltaic coal-bearing & Alcober 1994; Kokogian et al. 1999). The
sequences. Ichnological and sedimentological oldest deposits are the red-beds of the Talam-
features that distinguish Cretaceous river- and paya and Tarjados Formations, which are suc-
wave-dominated marine delta sequences from ceeded by thin volcaniclastic deposits of the
shorefaces have been proposed (Gingras et al. Chanares Formation and widespread lacustrine
1998; Coates & MacEachern 1999). Moslow & strata of the Ischichuca, Los Rastros and
Pemberton (1988) and Coates & MacEachern Lomas Blancas Formations (Fig. 2). The reader
(1999) noted that, in river-dominated delta is referred to Stipanicic & Bonaparte (1979),
successions, prodelta deposits are devoid of Lopez Gamundi et al. (1989) and Kokogian
bioturbation and delta front deposits display a et al. (1999) for further details on the stratigra-
low-density, moderate-diversity Cruziana assem- phy of the basin. Except for the lower part of
blage. Coates & MacEachern (1999) typified the Ischichuca Formation, which contains shal-
wave-dominated delta successions as having a low lacustrine non-deltaic deposits, the lacustrine
diverse, low-density, stressed Cruziana assem- succession is typically arranged in coarsening-
blage in prodelta sediments, and a moderately and shallowing-upward cycles (parasequences)
diverse, locally high-density mixed Skolithos- that record delta progradation (Lopez Gamundi
Cruziana assemblage in delta front deposits. In et al. 1989; Milana 1998; Bellosi et al. 2001;
the latter case, a marked decrease in the Melchor et al. 2003) (Fig. 3). The lacustrine
abundance and diversity of trace fossils is succession of the basin contains sediments of
attributed to higher-energy conditions (Coates different freshwater to saline palaeolakes that
& MacEachern 1999). Whether these relation- varied from shallow and well oxygenated (i.e.
ships can be extrapolated to the freshwater less than 10m deep) to moderately deep (up to
realm is unknown. 80m deep) and thermally stratified, the latter
The purposes of this paper are: with anoxic bottom waters (Milana 1998; Mel-
chor unpublished data). The deltaic lacustrine
to document the detailed stratigraphic distri- succession of the basin is envisaged to reflect
bution of trace fossils in lacustrine deltas of humid climatic conditions (e.g. Bonaparte
the Triassic Ischigualasto-Villa Union rift 1969) and was developed at tropical latitudes
basin of Argentina; (about 35-36°S after Prezzi et al. 2001). The
to compare these examples with the trace fossil freshwater nature of the lakes where the deltas
assemblages from shallow-shelf deltas from prograded is well documented by various
ICHNOLOGY OF LACUSTRINE DELTAS 339

Fig. 1. Location map. (a) Position of the study area in South America (left) and extension of Triassic rift
basins in northwest Argentina (right). The rectangle shows position of Fig. Ib. (b) Geologic map of the
Ischigualasto-Villa Union Basin showing the localities of studies 1-3. Modified from Stipanicic & Bonaparte
(1979).

independent lines of evidence: low carbon/sul- also in agreement with a freshwater lacustrine
phur ratios in the black prodelta shales, abun- setting.
dance of branchiopods (mainly conchostracans The studied localities are located in the north-
and notostracans), presence of the freshwater western (quebrada or canyon de Ischichuca,
algae Plaesiodictyon mosellanum Brenner & locality 1), eastern (Rio Gualo, Talampaya
Foster 1994 (Zavattieri & Melchor 1999), and Park, locality 2) and southeastern (La Torre,
the rarity or absence of evaporites. The recog- locality 3) area of the basin (Fig. Ib). All the
nized facies associations and the abundance of examples described in this paper come from the
plant, insect and palaeoniscid fish remains are lacustrine deltaic interval of the basin. The first
340 R. N. MELCHOR

Fig. 2. Stratigraphy of the Ischigualasto-Villa Union basin showing the relationships of the lacustrine units
and different formational names used in the three studied localities.

two localities are located in the flexural margin, Facies associations


and the third locality corresponds to an accom-
modation zone margin at the southern end of This section contains a brief and general descrip-
the half-graben. Sedimentological and stratigra- tion of the main facies associations recognized in
phical attributes suggest that these deltas are the deltaic lacustrine successions of the basin,
best compared with the tropical lacustrine including representative aspects of the described
deltas of the East African rift lakes, and in parti- deltaic lithofacies (Figs 4, 5). A more detailed
cular with those of low depositional slope (cf. lithofacies characterization of each trace fossil
Johnson et al, 1995). Detailed logging and strati- assemblage is given below, together with the
graphic analysis (including tracing of lacustrine description of the ichnofossils in Table 2. Para-
flooding and ravinement surfaces) permitted sequences range in thickness from 12 to 80m,
basinward correlation of the parasequences and the thickest ones are found at quebrada de
(Fig. 3) for over 100km. This correlation sug- Ischichuca. Both river-dominated and wave-
gests that the stacking pattern of the lacustrine dominated deltas were recognized. A further
deltas reflects lake-level changes. The three distinction is made on the basis of the nature of
sections are roughly stratigraphically equivalent, the body of water where these deltas prograded:
but have markedly different thickness: the thick- low-accommodation, shallow-water deltas were
est succession is located at the northernmost small and emplaced on a broad lacustrine shelf,
locality, and the thinnest succession is found at and high-accommodation deepwater deltas were
the southern-most locality (cf. Bossi 1971). composed by large and low sloping delta lobes
These differences are explained by contrasting (Table 2).
tectosedimentary regimes in a half-graben setting
(Melchor 2002), and are strongly linked with
the wedge-shaped stratal geometry that is Offshore lacustrine (OL)
typical of half-grabens (e.g. Leeder & Gawthorpe
1987). Invertebrate and plant trace fossils This facies association displays contrasting fea-
occur with different density and diversity in tures in the three analysed sections and appears
some parasequences, which are described below to lack evidence of bioturbation. At quebrada
(Fig. 3). de Ischichuca it is represented by thick (up to
ICHNOLOGY OF LACUSTRINE DELTAS 341

Fig. 3. Schematic stratigraphic sections at quebrada (canyon) de Ischichuca, Rio Gualo and La Torre areas
showing facies associations, correlation surfaces and trace fossil assemblages.
342 R. N. MELCHOR

Fig. 4. Field photos of delta front fades association, (a) Succession of offshore lacustrine (OL), delta front
(DF) and delta plain (DP) facies associations from wave-dominated delta at Rio Gualo; person for scale
(circled), (b) Siltstone-dominated, graded underflow deposits, (c) Hummocky cross-stratification from delta
front deposits at Rio Gualo (arrowed); circled hammer is 0.35m long, (d) Underflow plus overflow deposits
from delta front deposits of a river-dominated delta at quebrada de Ischichuca (staff divisions are 0.10m).
(e) Wave-rippled siltstones and sandstones from upper delta front deposits at La Torre.

42m) and monotonous successions of papery locality although fossils are less abundant.
black shales with as much as 3% total organic These successions are interpreted as reflecting
carbon content and no evidence of disruption deposition from suspension in a moderately
of the lamination (Fig. 5a, d). The shales have shallow lacustrine setting, with well-oxygenated
yielded varied fossil remains including fish, waters or temporary water stratification.
conchostracans, insects, plants and palyno-
morphs. These shales are interpreted to have
been deposited from suspension in a deep fresh- Delta front (DF)
water lake with anoxic bottom waters favoured
by thermal stratification. This facies association is composed of coarsening-
At Rio Gualo and La Torre (localities 2 and 3 and thickening-upward, thinly bedded, siltstone-
respectively) this facies association is represented dominated successions with minor sandstone
by olive grey or brown shales showing fine intercalations. These successions show their base
parallel lamination that compose intervals up transitional to offshore lacustrine shales, and
to 11 m thick (Fig. 4a). Massive or poorly lami- their top commonly is defined by a sharp contact
nated intervals and fine siltstone interbeds can with overlying distributary channel sandstones
occur. Fossil content is similar to the previous of the delta plain facies association (Fig. 4a).
ICHNOLOGY OF LACUSTRINE DELTAS 343

Fig. 5. Field photos of delta plain and marginal lacustrine fades associations from quebrada de Ischichuca
(locality 1, Fig. Ib). (a) Upper part of the second parasequence (Fig. 3) showing partially covered offshore
lacustrine and delta front deposits, capped by delta plain sediments. Arrow indicates the stratigraphic position
of Fig. 5b, c, (b) Detail of lateral accretion surfaces of interdistributary bay deposits sharply overlain by
coarser-grained channel sediments, (c) Detail of the previous figure showing wave-rippled siltstones interbedded
with carbonaceous mudstones. Lens cap is 5.8 cm wide, (d) Field view of the succession covered by Fig. 7a that
includes offshore lacustrine, delta front and marginal lacustrine facies associations. Note banded delta front
sediments in the lower left corner. OL, offshore lacustrine facies association; DF, delta front facies association;
DP, bay + channel = delta plain facies association; ML, marginal lacustrine facies association. TF-A to TF-C,
trace fossil assemblages.

Parallel-laminated, graded siltstones with laminae upward-fining cosets up to 9m thick (major


0.5-3 cm thick compose rhythmic intervals that channels, Fig. 5a), overlain by fine-grained
can reach several metres thick (Fig. 4b, d). Some sandstones, siltstones, tuffs, and occasional
laminated siltstone intervals display wave-ripple palaeosols (interdistributary deposits). The
structures (Fig. 4e). Interbedded sandstones can cross-bedded sandstones have erosive bases and
show parallel lamination, ripple cross-lamination commonly contain soft-sediment deformation
and hummocky cross-stratification (Fig. 4c). structures. Although uncommon, lateral accre-
Soft-sediment deformation structures are rare. tion surfaces can be associated with these
This facies association displays the highest diver- deposits (Fig. 5b, c). Trace fossils are scarce
sity and density of trace fossils. Delta front facies and commonly restricted to the uppermost
associations of river- and wave-dominated deltas finer-grained lithologies. This facies association
are distinguished by the common occurrence of represents active bedload deposition by moder-
wave and combined flow structures in the latter. ate-sinuosity fluvial channels in a low-gradient
setting, in which channels shifted position fre-
quently. Both major and secondary distributary
Delta plain (DP) channels can be recognized. The fine-grained
interval is interpreted as abandonment facies,
Delta plain deposits are essentially composed of which could be exposed subaerially and modified
cross-bedded distributary channel deposits and by pedogenic processes.
minor interdistributary bay sediments assigned
to crevasse channel, crevasse delta and levee
sub-environments. The delta plain is character- Marginal lacustrine (ML)
ized by fining-upward successions defined by
the presence of fine- to coarse-grained sandstones These littoral deposits are part of shallow shelf
with trough cross-stratification arranged in deltas at the top of the Ischichuca Formation
Table 2. Summary of trace fossil assemblages from lacustrine deltas of Ischigualasto-Villa Union basin organized by fades association

Fades Name Facies Delta Lake Ethological categories Diversi ty* Densitv
associations type* basin (BPBIJ3
Repichnia Pasichnia Domichnia Fodinichnia Cubichnia Fugichnia Plant
traces

Delta front TF-A Lower delta R-D Deep Undichna Cochlichnus Fine burrows Rusophycus 1 2
front Diplichnites
Underflow Stiaria
deposits Cruziana
TF-D Lower delta W-D Deep Bifurculapes. Cochlichnus Palaeophycus Treptichnus Rusophycus 18 3
front Diplichnites Gordia Avolatichnium
Underflow Protichnites Helminthoidichnites 'rhomboidal traces'
deposits Undichna ispp.
Cruziana
Diplopodichnus
Didymaulichnus
f Middle delta R-D Shelf Archaeonassa Cochlichnus Palaeophycus 7 2
front Undichna Gordia
Helminthoidichnites
Helminthopsis
TF-E Upper delta R-D Shelf Archaeonassa Cochlichnus 2 2
front
f
II Upper delta R-D shelf Cochlichnus Palaeophycus 3 2
front 'fusiform
structures'
Delta plain TF-B Minor R-D Deep Escape trace 1 2
channel
TF-F Channel R-D Shelf Root 1 2
traces
1
III Mouth-bar R-D Shelf Palaeophycus 2 2
Skolithos
iv* Channel R-D Shelf Rhynchosauroides Palaeophycus 'Horseshoe-shaped 4 2
ispp. structures'
Marginal TF-C Nearshore W-D Shelf Cochlichnus 1 2
lacustrine bars

*R-D, river-dominated; W-D, wave-dominated. ^Melchor et al. (2003). * Expressed as the number of ichnotaxa. ^Maximum bedding plane bioturbation index of Miller & Smail (1997).
ICHNOLOGY OF LACUSTRINE DELTAS 345

at quebrada de Ischichuca (Fig 5d). They are interval (Figs 4d, 7a). It is represented by a
characterized by thick siltstone-dominated suc- moderately diverse ichnofauna dominated by
cessions (up to 50 m thick) with secondary inter- locomotion traces (Cruziana problematica,
calations of wave-rippled or parallel-laminated, Undichna britannica, Diplichnites isp., Stiaria
fine-grained sandstone beds. They commonly isp.), although resting (Rusophycus stromnessi),
display sole marks and are associated with occa- grazing or locomotion (Cochlichnus anguineus)
sional isolated hummocky lenses (anisotropic and feeding structures (very thin, less than
hummocky cross-stratification of Midtgaard 1 mm in diameter, oblique burrows) were also
1996) and rhythmic graded heterolithic beds. recorded. The most abundant traces are Cruziana
These deposits are interpreted as products of and Rusophycus, which are found in the lower
sedimentation in a fully subaqueous, nearshore part of the interval along with most of the
lacustrine setting. Deposition is attributed to remaining ichnofossils. Arthropod locomotion
river-fed underflows and to oscillatory and com- traces and Cochlichnus are restricted to the
bined flows with occasional modification under upper part of the trace-bearing interval.
storm wave-generated oscillatory flows.
Trace fossil assemblage E (rived-dominated delta,
low-accommodation basin)
Trace fossil assemblages This assemblage was recorded at La Torre (local-
ity 3, Figs Ib, 3) within the Lomas Blancas
A total of six trace fossil assemblages were iden- Formation (Figs 3, 7c) and corresponds to
tified, which correspond to the following facies wave-rippled upper delta front facies. Delta
associations: delta front (three assemblages), front sediments at this locality are characterized
delta plain (two assemblages) and marginal by decimetre-thick beds that display a normal
lacustrine (one assemblage). They are described grading from grey tuffaceous siltstone to mud-
in detail below and summarized in Table 2. stone with parallel lamination. The laminated
This table also contains the trace fossil assem- muds grade upward to trace-bearing red silt-
blages of the correlative Los Rastros Formation stones with parallel lamination, wavy bedding
at Ischigualasto Park described by Melchor et al. and symmetrical ripples (Fig. 4e). Rare fine-
(2003). They are included for consideration in the grained sandstone beds display trough cross-
discussion because they are partially correlative bedding, convolute lamination and occasional
and represent similar environmental settings. recumbent folding (probable slump). This
Some of the typical trace fossils of the delta succession records deposition from underflow
front facies association are also illustrated currents and from settling associated with excep-
(Fig. 5). tional floods in a moderately shallow lake basin
(probably about 20-25 m deep as inferred from
the thickness of the parasequences and the
Delta front assemblages characteristics of associated offshore lacustrine
deposits).
These assemblages were recorded in the three The dominant ichnofossil of this assemblage is
analysed localities and are TF-A, TF-D and the sinusoidal burrow Cochlichnus anguineus,
TF-E, which correspond to the Ischichuca, Los which occur together with the trail Archaeonassa
Rastros and Lomas Blancas Formations, fossulata.
respectively (Figs 3, 7). They belong both to
river-dominated deltas, including deltas that Trace fossil assemblage D (wave-dominated delta,
prograded in high-accommodation (TF-A) and high-accommodation basin)
low- accommodation basins (TF-E), and to This assemblage occurs in two stacked para-
wave-dominated deltas that prograded in a sequences that are 32 and 50m thick, and crop
high-accommodation basin (TF-D). out at Rio Gualo (locality 2, Figs Ib, 3) and
belong to the Los Rastros Formation. Most of
Trace fossil assemblage A (river-dominated delta, the thickness of each parasequence represents
high-accommodation basin) siltstone-sandstone-dominated lower delta
TF-A from quebrada de Ischichuca (Figs 3, 7a) front facies. Trace-bearing facies at this locality
corresponds to lower delta plain facies and are essentially composed of laminated light-grey
occurs in silty underflow deposits. It was siltstones and minor sandstone interbeds (Figs
recorded from a 4m thick interval showing 4a, 7b). The sandstones display parallel lamina-
interbedded siltstone, mudstone and shale with tion, hummocky cross-stratification or wave
occasional convolute lamination that is replaced ripple cross-lamination. Thin (5-1 Ocm thick)
towards the top by a 10m thick rhythmite fine-grained sandstone/siltstone laminae with
346 R. N. MELCHOR

Fig. 6. Representative trace fossils from lacustrine delta front facies association, (a) Fish trail Undichna
britannica. (b) Grazing or locomotion trails Cochlichnus anguineus (C), Gordia marina (G), Helminthoidichnites
tenius (H). (c) Long specimens of Cruziana problematica (locomotion trace of arthropods), (d) Arthropod
locomotion and resting traces: Protichnites isp. (P), Cruziana isp. (Cr) and 'rhomboidal traces' (r). (e) Feeding
burrows: Treptichnus pollardi. (f) Bedding plane with moderately high bioturbation (BPBI — 3; Miller & Smail
1997). Most burrows can be assigned to Cochlichnus. Scale bar = 5cm.

normal grading and parallel lamination are (Fig. 6b, f), Gordia marina (Fig. 6b), Helminthoi-
common (Fig. 4b). Thick bedsets of hummocky dichnites tenius (Fig. 6b), Palaeophycus tubularis
cross-stratification are also recorded, but they and Treptichnus pollardi (Fig. 6e). Arthropod
are devoid of ichnofossils (Fig. 4c). Trace fossils ichnofossils are diverse and reveal locomotion
were recorded mostly on the tops but also on the (Cruziana problematica, Fig. 6c, Diplopodichnus
bases of finely laminated graded beds, which are biformis, Didymaulichnus lyelli, Bifurculapes
interpreted as underflow deposits (Fig. 4b). They isp., Diplichnites isp., Protichnites isp.) and rest-
represent semi-permanent lake floor sedimenta- ing/feeding activities (Rusophycus stromnessi,
tion in a well-oxygenated setting punctuated by Avolatichnium isp. and 'rhomboidal traces', Fig.
deposition of finer-grained sediments from 6d, comparable to those described by Buatois
settling (overflow deposits) and by sedimentation & Mangano 1993a). Fish trails are assigned
of coarse-grained sediments during high-energy to Undichna britannica (Fig. 6c), U. bina and U.
events (storms). cf. insolentia. The most common ichnotaxa
Trace fossils of both parasequences compose a are Cochlichnus anguineus (burrows that appear
high-diversity (18 ichnotaxa) and moderate- to with high densities at some stratigraphic
high-density assemblage (Figs 3, 7b) (Melchor levels), Cruziana problematica (and its tapho-
2001). It contains grazing/feeding burrows or nomic variants Diplopodichnus and Didymaul-
trails, locomotion and resting/feeding traces ichnus, see Keighley & Pickerill 1996), and
of arthropods, and fish trails (Figs 6a, b, 7b). Undichna britannica. Cochlichnus appears fre-
The first group includes Cochlichnus anguineus quently with deformed burrow walls. Besides
ICHNOLOGY OF LACUSTRINE DELTAS 347

Fig. 7. Detailed sedimentologic logs of selected examples of the trace fossil assemblages. See Fig. 3 for
stratigraphic position, (a) River-dominated deltaic and marginal lacustrine deposits from the Ischichuca
Formation, (b) Second parasequence of a wave-dominated delta from the Los Rastros Formation at Rio
Gualo. (c) Second parasequence of a river dominated delta from the Lomas Blancas Formation at La Torre.
348 R. N. MELCHOR

this compositional characterization, the upper Trace fossil assemblage F (river-dominated delta,
parasequence displays a preferred distribution low-accommodation basin)
of some ichnotaxa in delta front fades: the This assemblage occurs in distributary deposits
lower part (below the first medium-grained sand- of the top of the third parasequence from La
stone bed) contains a larger number of ichnotaxa Torre (locality 3, Figs Ib, 3) within the Lomas
than the upper part (Fig. 7b). TF-D contains the Blancas Formation (Fig. 7c) and is composed
higher density of burrowing (BPBI = 3 in the of root traces. Distributary channel deposits are
scheme of Miller & Smail 1997) of all analysed 5-6.5m thick, characterized by medium-grained
assemblages. sandstones with trough cross-bedding associated
with parallel-laminated or massive fine-grained
sandstones. The parasequence is capped by
Delta plain assemblages fine-grained, slightly reddened sandstone with
moderately abundant root traces. The delta
Two low-diversity and low-density ichnological plain interval of the overlying parasequence is
assemblages were recorded from delta plain similar, although it also contains heterolithic
settings of river-dominated deltas: TF-B from a deposits. They consist of fine-grained sandstone,
high-accommodation lake basin (Ischichuca siltstone and mudstones with abundant carbon-
Formation), and TF-F from a low-accommoda- aceous material, which display trough cross-bed-
tion lake basin (Lomas Blancas Formation). ding, parallel lamination and synsedimentary
microfaulting, and are succeeded by lateral
Trace fossil assemblage B (river-dominated delta, accretion deposits with abundant wavy and
high-accommodation basin) lenticular bedding. The coarse-grained sediments
This assemblage is represented by escape traces of the described parasequences reflect the progra-
in minor channels of the delta plain from dation of a delta lobe in lake waters with
quebrada de Ischichuca (locality 1, Fig. 3), common wave reworking. These sediments are
within the Ischichuca Formation. Traces occur sharply covered by channel deposits that were
in a 7m thick fining-upward cycle bounded by exposed subaerially, thus favouring the develop-
a sharp and erosive lower surface that includes ment of incipient soils. Heterolithic lateral accre-
(from bottom to top) planar cross-stratified sets tion deposits are related to sedimentation in a
with reactivation surfaces and a single through moderately sinuous distributary channel.
cross-bedded set (both showing soft-sediment Root traces are typically 0.01-0.8 cm in dia-
deformation structures) in medium-grained meter, up to 10 cm long, display occasional bifur-
sandstones, which are covered by fine-grained cations and have a conspicuous yellow-grey halo
heterolithic deposits. The latter comprises lateral with a maximum thickness of about 0.7cm.
accretion surfaces and wave-rippled sandstones, Former root cavities are filled with fibrous
and siltstones with climbing ripples and lenticu- silica, carbonate and remains of probable silici-
lar bedding, which are interbedded with dark fied plant tissue. These root traces are similar
plant-bearing mudstones (Fig. 5b, c). This cycle to the drab-haloed root traces of Retallack
is laterally correlative with crevasse deltas and (1983). Of the common origins postulated for
levee deposits. Escape traces have been recorded this type of root traces, both incipient water-
from the top of the trough cross-bedded set. logging and anaerobic decay of the organic
This cycle represents deposition in a minor dis- matter of the root during early burial are possible
tributary channel or a crevasse channel of the in this case (cf. Retallack 1983, 1990).
delta plain, as suggested by the sedimentological
attributes, the presence of common reactivation
surfaces, and lateral correlation with crevasse Marginal lacustrine assemblage
deltas and levee deposits. These channels were
dominated by lateral accretion onto point A single assemblage has been recorded from a
bars (lateral accretion surfaces), and suffered subaqueous marginal lacustrine setting of a
common stage changes (reactivation surfaces) river-dominated delta that prograded in a shallow
and probably frequent avulsions (thin fining- shelf (Ischichuca Formation).
upward cycles). It is possible that sedimentation
was rapid, thus promoting sediment instability Trace fossil assemblage C (river-dominated delta,
(soft-sediment deformation features) and transi- low-accommodation basin)
ent high sedimentation rates (escape structure). This assemblage was identified at quebrada de
Escape traces are composed of a central disrupted Ischichuca (locality 1, Figs Ib, 3) from the
zone (1cm wide and 8cm high) surrounded by homonymous formation. It is a monospecific
downward-deflected laminae. assemblage (Figs 5d, 7a), restricted to a 3m
ICHNOLOGY OF LACUSTRINE DELTAS 349

thick interval interpreted as subaqueous near- Trace fossil assemblages from the delta plain
shore lacustrine deposits laterally associated facies association are considerably less diverse
with shallow shelf deltas. than those from delta front facies, showing a
The deposits commonly are arranged in total of seven ichnotaxa that preferentially
coarsening-upward cycles composed of graded occur on delta tops associated with low-
heterolithic beds and laminated siltstones with accommodation basin states. In contrast, delta
sandstone interbeds, which are capped by plain facies related to high-accommodation lake
parallel-laminated and wave-rippled sandstones. basin states are almost devoid of ichnofossils,
Rare isolated sandstone hummocks (anisotropic limited to a single record of an escape trace in
hummocky cross-stratification) can occur in the TF-B. The low ichnodiversity contrasts with the
upper part of the cycles (Fig. 7a). The description large number of behavioural categories repre-
of the ML facies association contains further sented, including lacertoid vertebrate tracks
details. Trace fossils are represented by a low- to (Rhynchosauroides), dwelling burrows (Skolithos,
moderate-density, monospecific assemblage com- Palaeophycus), probable resting traces of arthro-
posed of Cochlichnus anguineus burrows. They pods ('horseshoe-shaped structures'), escape
are restricted to the tops of fine-grained sandstone trace and root traces (Table 2). Within these
beds showing symmetrical or interference ripples. assemblages there are also indicators of desic-
cated substrates, evidenced by the presence of
fine striations in Palaeophycus striatus from
Discussion trace fossil assemblage IV of the Los Rastros
Formation, Ischigualasto Park (Melchor et al.
Environmental and stratigraphic repartition 2003).
of trace fossils Ichnofossils from the studied Triassic lacus-
trine deltas are almost exclusively restricted to
The most diverse trace fossil assemblages are shallow penetrating traces on some bedding
found in the delta front facies association, planes. Most of the bedding planes with ichno-
which shows a fairly high ichnodiversity (22 fossils display less than 10% bioturbation
recorded ichnotaxa). Among these, the trace (BPBI = 2 of Miller & Smail 1997), with scarce
fossil assemblages corresponding to distal or examples reaching as much as 25% bioturbation
intermediate settings of the subaqueous delta (BPBI = 3) (Table 2). The restriction of burrow-
lobe contain the greater number of ichnotaxa. ing by benthic organisms to bedding planes and
This relationship holds for both river- and common low density of bioturbation has been
wave-dominated deltas, although the examples documented in other Late Palaeozoic and Trias-
from wave-dominated deltas of the Los Rastros sic lacustrine and fluvial successions (Buatois
Formation at Rio Gualo are by far the most et al. 1998; Miller et al. 2002). The single example
diverse assemblages (TF-D, Table 2). The of a shallow subaqueous marginal lacustrine
upper part of the delta front contains an trace fossil assemblage (TF-C) resembles those
impoverished assemblage, with ubiquitous found in upper delta front settings, especially
sinusoidal grazing or locomotion trails assigned because of the exclusive occurrence of simple
to Cochlichnus. This trace fossil is found in six grazing trails and the association with wave-
of the eight subaqueous assemblages, but com- ripple structures.
prises monospecific assemblages only in marginal In addition to tracing the differences in trace
lacustrine deposits (Table 2). Cochlichnus has fossil content along proximal to distal gradients
been recognized in a large variety of marine, within individual lacustrine deltas, sequence
transitional and continental environments, stratigraphic correlation allows comparison of
which range in age from Precambrian to Holo- laterally equivalent, but different, deltas. In
cene (Buatois et al. 1997). The apparent vertical particular, trace fossil assemblages B (locality
zonation found in TF-A & D could reflect parti- 1), D (second parasequence at locality 2) and
tioning of assemblages into proximal and distal E + F (locality 3) occur in parasequences that
delta front deposits (cf. Buatois & Mangano are bounded by correlative flooding surfaces
1993b). (Figs 3, 7). There is a striking difference between
Ethologically, the assemblages are dominated the almost lack of bioturbation in the high-
by locomotion and grazing traces with sub- accommodation rived-dominated delta of TF-
ordinate resting and feeding traces and rare B, the high diversity and high density of the
dwelling structures. Locomotion traces are high-accommodation wave-dominated deltas of
almost exclusively assigned to arthropods, TF-D, and the depleted ichnocoenoses of the
whereas the remaining repichnial ichnotaxa are low-accommodation, river-dominated delta of
ascribed to fishes or gastropods. TF-E and F.
350 R. N. MELCHOR

Controls on trace fossil distribution Buatois & Mangano 1993a, 1995). An important
difference is the dominance of locomotion and
In the permanently subaqueous delta front set- resting traces attributed to arthropods in these
tings, oxygenation, food supply, water turbidity, assemblages, instead of shallow surface-grazing
erosion and sedimentation rates influence trace trails that characterize the Mermia ichnofacies.
fossil distribution. It is well documented that As suggested by Buatois & Mangano (1998)
density currents, both discrete turbidity currents and Melchor et al. (2003), trace fossil assem-
and semi-permanent underflow currents, supply blages from shallow-lacustrine high-energy set-
oxygen and food to deep, subaqueous settings, tings are best ascribed to the Skolithos
thereby favouring the establishment of a diverse ichnofacies, and intermittently exposed lacus-
biota (e.g. Buatois et al. 1996; Buatois & trine shoreline deposits contain ichnofossils that
Mangano 1998). This is especially true for suggest assignation to the Scoyenia ichnofacies
oxygen-deficient bottom waters, as envisaged (Buatois and Mangano 1995, 2004). These three
for TF-A. Higher erosion and turbidity in ichnofacies are present in a single parasequence
river-dominated delta front environments may only in river-dominated low-accommodation
explain the low diversity and density of trace deltas at Ischigualasto Park locality (Fig. Ib).
fossils in these settings, in contrast to wave- The deeper or permanently subaqueous deltaic
dominated delta front settings. The later were successions contain assemblages that are less
emplaced in bottom waters with higher oxygena- variable. Nevertheless, as exemplified by the
tion than the deep anoxic and non-bioturbated apparent vertical zonation in TF-A & D (Fig.
intervals of the Ischichuca Formation lakes, 7a, b), it is possible that future studies will recog-
thus allowing the establishment of a resident nize characteristic and repetitive assemblages
fauna with less dependence on the supply of within the Mermia ichnofacies. Buatois &
oxygen and food from turbidity currents. Ero- Mangano (1996) have also noted an increase in
sion produced by storms may have destroyed the number of arthropod locomotion traces in
the trace fossils produced on previous underflow littoral lacustrine facies with relation to deep-
deposits. Thick amalgamated hummocky cross- offshore facies, which are dominated by sur-
stratified sandstones lack evidence of bio- face-grazing trails. The monospecific assemblage
turbation, which is interpreted as reflecting low from a subaqueous marginal lacustrine setting is
potential of preservation of shallow-penetrating regarded as an impoverished occurrence of the
traces (cf. Frey & Goldring 1992). In the upper Mermia ichnofacies.
part of delta front successions, where there are The compositional comparison with marine
indications for higher erosion and sedimentation deltaic sequences reveals significant differences.
rates, trace fossil assemblages are reduced in However, there are similar trends in freshwater
diversity and density, or are absent. Sedimento- and marine deltaic successions, with an upward
logic evidence suggests that the sinusoidal decrease of ichnodiversity and abundance in
burrow Cochlichnus is restricted to subaqueous shallower and more energetic settings. In
settings, and the frequent burrow-wall deforma- addition, a greater diversity and density of
tion observed in some specimens suggests that trace fossils is recorded from wave-dominated
they were produced in highly water-saturated settings (Table 2).
substrates (cf. Buatois et al. 1997).
Energy and substrate water content are
envisaged as the most important factors that Conclusion
controlled the formation and preservation of
ichnofossils in delta plain settings of the studied This study documents the composition and facies
examples. Actively filled fluvial channel deposits repartition of trace fossil assemblages from dif-
are devoid of traces, but trace fossils do occur in ferent locations of a Triassic lacustrine deltaic
intermittently exposed channel margin deposits succession. Trace fossils from all subaqueous
in the form of striated burrows, footprints and non-marine settings are assigned to the Mermia
root traces. ichnofacies. However, it is envisaged that addi-
tional detailed studies may allow further discri-
mination within this ichnofacies. This inference
Comparison with other trace fossil is supported by documentation of atypical
assemblages Mermia-type assemblages herein with moderate
to very high ichnodiversity and facies-dependent
The composition of trace fossil assemblages from vertical zonation of trace fossils (e.g. TF A & D).
delta front settings is comparable with typical Trace fossil assemblages from the Triassic
occurrences of the Mermia ichnofacies (e.g. lacustrine succession of Ischigualasto-Villa
ICHNOLOGY OF LACUSTRINE DELTAS 351

Union basin attain maximum diversity and BELLOSI, E., JALFIN, G., Bossi, G., MURUAGA, C.,
density in wave-dominated delta front fades, BOGGETTI, D. & CHEBLI, P. 2001. Ambientes
are absent in anoxic offshore lacustrine fades, sedimentarios en cuencas triasicas de Argentina.
and are scarce in delta plain facies. These differ- Boletin de Informaciones Petroleras, 68, 54—83.
BONAPARTE, J. F. 1969. Datos sobre la evolution
ences may aid in the stratigraphic analysis of paleoecologica en las formaciones Triasicas de
lacustrine successions and help to distinguish Ischigualasto-Villa Union (San Juan-La Rioja).
between wave- and river-dominated lacustrine Acta Geologica Lilloana, 10, 189-206.
deltas. Trace fossil assemblages in the analysed Bossi, G. E. 1971. Analisis de la Cuenca Ischigualasto-
lacustrine deltas mimic density/diversity patterns Ischichuca. Primer Congreso Hispano-Luso-
in marine deltas, although they are dominated by Americano de Geologia Economica, Madrid, 2,
non-marine ichnotaxa. 611-626.
This study contributes to the documentation BRENNER, W. & FOSTER, C. B. 1994. Chlorophycean
of the general trace fossil distribution in lacus- algae from the Triassic of Australia. Reviews of
Palaeobotany & Palynology, 80, 209-234.
trine deltas, from distal to proximal areas
BUATOIS, L. A. & MANGANO, M. G. 1993a. Trace fossils
(Tables 1, 2). Identified trends include: from a Carboniferous turbiditic lake: implications
the absence of traces in oxygen-deficient off- for the recognition of additional nonmarine
shore deposits; ichnofacies. Ichnos, 2, 237-258.
BUATOIS, L. A. & MANGANO, M. G. 1993b. The
the high diversity of simple grazing trails in paleoenvironmental and paleoecological signifi-
delta-fed turbidites (Buatois & Mangano cance of turbiditic lake ichnocoenoses from the
1993a; Buatois et al 1996; Trewin et al 2002); Late Carboniferous of the Paganzo Basin, Argen-
the high diversity and apparent dominance of tina. Comptes Rendus XII International Congress
arthropod traces in lower delta front deposits, Carboniferous-Permian, 2, 409^20.
which decrease in diversity toward the top of BUATOIS, L. A. & MANGANO, M. G. 1995. The paleo-
progradational successions; and environmental and paleoecologic significance of
the low diversity of ichnofossils representing the lacustrine Mermia ichnofacies: an archetypical
different ethologic categories in fine-grained, subaqueous nonmarine trace fossil assemblage.
occasionally exposed delta plain deposits. Ichnos, 4, 151-161.
BUATOIS, L. A. & MANGANO, M. G. 1996. Icnologia de
No diagnostic ichnological signature for identifi- ambientes continentales: problemas y perspectivas.
cation of key stratigraphic surfaces in lacustrine Asociacion Paleontologica Argentina, Publication
deltas was found, though improved ichnofacies Especial, Buenos Aires, 4, 5—30.
BUATOIS, L. A. & MANGANO, M. G. 1998. Trace fossil
characterization may enable ichnofacies-stacking analysis of lacustrine facies and basins. Palaeo-
patterns to be used in stratigraphic analysis, as is geography, Palaeoclimatology, Palaeoecology,
conventionally performed in marine successions. 140, 367-382.
In addition, identification of a diverse trace fossil BUATOIS, L. A. & MANGANO, M. G. 2002. Trace fossils
assemblage in deltaic lacustrine successions may from Carboniferous floodplain deposits in western
aid in recognition of distal delta lobe deposits Argentina: implications for ichnofacies models of
and thus help to locate potential reservoir facies. continental environments. Palaeogeography,
Palaeoecology, Palaeoecology, 183, 71-83.
Funding for this research was obtained from research BUATOIS, L. A. & MANGANO, M. G. 2003. Caracteriza-
grants PICT 6156 (ANPCyT) and PEI 157/98 (CONI- cion icnologica y paleoambiental de la localidad
CET), both from Argentina. The Universidad Nacional tipo de Orchesteropus atavus Frenguelli, Huerta
de La Pampa provided logistic support and partial de Huachi, provincia de San Juan, Argentina.
funding for fieldwork (project no. 136 of the Facultad Ameghiniana, 40, 53-70.
de Ciencias Exactas y Naturales). L. Buatois, A. BUATOIS, L. A & MANGANO, M. G. 2004. Animal-
Martin, D. Mcllroy and P. Pazos reviewed the manu- substrate interactions in freshwater environments:
script and made pertinent suggestions that improved applications of ichnology in facies and sequence
the paper. stratigraphic analysis of fluvio-lacustrine succes-
sions. In: MclLROY, D. (ed.) The Application of
Ichnology to Palaeoenvironmental and Strati-
References graphic Analysis. Geological Society, London,
Special Publications, 228, 311-333.
BANN, K. L. & FIELDING, C. R. 2004. An integrated BUATOIS, L. A., MANGANO, M. G., Wu, X. & ZHANG,
ichnological and sedimentological comparison of G. 1996. Trace fossils from Jurassic lacustrine
non-deltaic shoreface and subaqueous delta turbidites of the Anyao Formation (central
deposits in Permian reservoir units of Australia. China) and their environmental and evolutionary
In: MclLROY, D. (ed.) The Application oflchnology significance. Ichnos, 4, 287-303.
to Palaeoenvironmental and Stratigraphic Analysis. BUATOIS, L. A., JALFIN, G. & ACENOLAZA, F. G. 1997.
Geological Society, London, Special Publications, Permian nonmarine invertebrate trace fossils
228, 273-310. from southern Patagonia, Argentina: ichnologic
352 R. N. MELCHOR

signatures of substrate consolidation and coloni- systems, Tanqua Karoo, South Africa. Sedimen-
zation sequences. Journal of Paleontology, 71, tology, 46, 987-1023.
324-336. JOHNSON, T. C., WELLS, J. D. & SCHOLZ, C. A. 1995.
BUATOIS, L. A., MANGANO, M. G., GENISE, J. F. & Deltaic sedimentation in a modern rift lake. Geo-
TAYLOR, T. N. 1998. The ichnologic record of logical Society of America Bulletin, 107, 812-829.
the continental invertebrate invasion: evolutionary KEIGHLEY, D. G. & PICKERILL, R. K. 1996. Small
trends in environmental expansion, ecospace Cruziana, Rusophycus, and related ichnotaxa
utilization, and behavioral complexity. Palaios, from eastern Canada: the nomenclatural debate
13, 217-240. and systematic ichnology. Ichnos, 4, 261-285.
COAXES, L. & MACEACHERN, J. A. 1999. The ichnologi- KINGSLEY, C. S. 1981. A composite fan-delta-fluvial
cal signature of wave- and river-dominated deltas: model for the Ecca and Lower Beaufort Groups
Dun vegan and Basal Belly River formations, of Permian age in the Eastern Cape Province,
West-Central Alberta. In: WRATHALL, B., JOHN- South Africa. Transactions Geological Society
STON, G., ARTS, A., Rozsw, L., ZONNEVELD, J-P., South Africa, 84, 27^0.
ARCURI, D. & MCLELLAN, S. (eds) Digging KIM, J. Y., KIM, K.-S. & PICKERILL, R. K. 2002. Cretac-
Deeper, Finding a Better Bottom Line. CSPG & eous nonmarine trace fossils from the Hasandong
Petroleum Society Core Conference Paper, p. 99- and Jinju Formations of the Namhae Area,
114. Kyongsangnamdo, southeast Korea. Ichnos, 9,
COHEN, A. S., LOCKLEY, M., HALFPENNY, J. & MICHEL, 41-60.
A. E. 1991. Modern vertebrate track taphonomy KOKOGIAN, D. A., SPALLETTI, L. A. ET AL. 1999. Los
at Lake Manyara, Tanzania. Palaios, 6, 371-389. depositos continentales triasicos. Anales del
COHEN, A. S., LOCKLEY, M., HALFPENNY, J. & MICHEL, Instituto de Geologia y Recursos Minerales, 29,
A. E. 1993. Modern vertebrate tracks from Lake 377-398.
Manyara, Tanzania and their paleobiological LEEDER, M. R. & GAWTHORPE, R. L. 1987. Sedimentary
implications. Paleobiology, 19, 433^58. models for extensional tilt-block/half-graben
DOYLE, P., WOOD, J. L. & GEORGE, G. T. 2000. The basins. In: COWARD, M. P., DEWEY, J. F. &
shorebird ichnofacies: an example from the Mio- HANCOCK. P. L. (eds) Continental Extensional
cene of southern Spain. Geological Magazine, Tectonics. Geological Society, London, Special
137, 517-536. Publications, 28, 139-152.
EAGAR, R. M. C, BAINES, J. G., COLLINSON, J. D., LOCKLEY, M. G., YANG, S. Y., MATSUKAWA, M.,
HARDY, P. G., OKOLO, S. A. & POLLARD, J. E. FLEMING, F. & LIM, S. K. 1992. The track record
1985. Trace fossil assemblages and their occur- of Mesozoic birds: evidence and implications.
rence in Silesian (Mid-Carboniferous) deltaic sedi- Philosophical Transactions of the Royal Society,
ments of the central Pennine Basin, England. In: London, Series B, 336, 113-134.
CURRAN, H. A. (ed.) Biogenic Structures: Their LOCKLEY, M. G., HUNT, A. P. & MEYER, C. A. 1994.
Use in Interpreting Depositional Environments. Vertebrate tracks and the ichnofacies concept:
Society of Economic Paleontologists & Mineralo- implications for palaeoecology and palichno-
gists, Special Publications, Tulsa, Oklahoma, 35, stratigraphy. In: DONOVAN, S. K. (ed.) The Palaeo-
99-149. biology of Trace Fossils. Wiley, Chichester, 241-
FLINT, S., STEWART, D. J. & VAN RIESSEN, E. D. 1989. 268.
Reservoir geology of the Sirikit oilfield, Thailand: LOPEZ GAMUNDI, O., ALVAREZ, L. et al. 1989. Cuencas
lacustrine deltaic sedimentation in a Tertiary Intermontanas. In: CHEBLI, G. & SPALLETTI, L. A.
intermontane basin. In: WHATELEY, M. K. & (eds) Cuencas Sedimentarias Argentinas. Serie
PICKERING, K. T. (eds,) Deltas: Sites and Traps Correlacion Geologica, 6, 123—167.
for Fossil Fuels. Geological Society, London, MANGANO, M. G., BUATOIS, L. A., Wu, X., SUN, J. &
Special Publications, 41, 223-237. ZHANG, G. 1994. Sedimentary fades, depositional
FREY, R. W. & GOLDRING, R. 1992. Marine event beds processes and climatic controls in a Triassic lake,
and recolonization surfaces as revealed by trace Tanzhuang Formation, western Henan Province,
fossil analysis. Geological Magazine, 129, 325-335. China. Journal of Paleolimnology, 11, 41-65.
GINGRAS, M. K., MACEACHERN, J. A. & PEMBERTON, MCDONALD, N. G. & LETOURNEAU, P. M. 1988.
S. G. 1998. A comparative analysis of the ichnol- Paleoenvironmental reconstruction of a fluvial-
ogy of wave and river-dominated allomembers of deltaic-lacustrine sequence, Lower Jurassic Port-
the Upper Cretaceous Dunvegan Formation. land Formation, Suffield, Connecticut. United
Bulletin of Canadian Petroleum Geology, 46, States Geological Survey Bulletin, 1767, 24-30.
51-73. MclLROY, D. (2004). Ichnology and facies model of a
ISBELL, J. L., MILLER, M. F., BABCOCK, L. E. & tide-dominated delta: Jurassic upper Ror and He
HASIOTIS, S. T. 2001. Ice-marginal environment Formations of Kristin Field, Halten Terrace, Off-
and ecosystem prior to initial advance of the late shore Mid-Norway. In: MclLROY, D. (ed.) The
Palaeozoic ice sheet in the Mount Butters area of Application of Ichnology to Palaeoenvironmental
the central Transantarctic Mountains, Antarctica. and Stratigraphic Analysis. Geological Society,
Sedimentology, 48, 953-970. London, Special Publications, 228, 237-272.
JOHNSON, S. D., FLINT, S., HINDS, D. & WICKENS, H. MELCHOR, R. N. 2001. Icnologia y sedimentologia de
DE V. 2001. Anatomy, geometry and sequence una sucesion lacustre influenciada por tormentas:
stratigraphy of basin floor to slope turbidite Formation Los Rastros (Triasico), Talampaya,
ICHNOLOGY OF LACUSTRINE DELTAS 353

La Rioja. 4th Reunion Argentina de Icnologia and PAZOS, P. J. 2002. Paleoenvironmental framework of
2nd Reunion de Icnologia del Mercosur, S. M. the glacial-postglacial transition (Late Paleozoic)
Tucuman, Abstracts, 56. in the Paganzo-Calingasta Basin (southern South
MELCHOR, R. N. 2002. Formation Ischichuca: su America) and the Great Karoo-Kalahari Basin
distinction de las Formaciones Chanares y Los (southern Africa): ichnological implications.
Rastros (Triasico, Norte de la cuenca Ischigual- Gondwana Research, 5, 619-640.
asto-Villa Union), Argentina. In: CABALERI, N., POLLARD, J. E. 1988. Trace fossils in coal-bearing
CINGOLANI, C. A., LINARES, E., LOPEZ DE LUCHI, sequences. Journal of the Geological Society,
M. G., OSTERA, H. A. & PANARELLO, H. O. (eds) London, 145, 339-350.
15th Congreso Geologico Argentino, Buenos Aires, POLLARD, J. E., STEEL, R. J. & UNDERSRUD, E. 1982.
Actas, 1, 690-693. Facies sequences and trace fossils in lacustrine/
MELCHOR, R. N., BELLOSI, E. S. & GENISE, J. F. 2003. fan delta deposits, Hornelen Basin (M. Devonian),
Invertebrate and vertebrate trace fossils from Western Norway. Sedimentary Geology, 32, 63-
a Triassic lacustrine delta: the Los Rastros 87.
Formation, Ischigualasto Provincial Park, San PREZZI, C., VIZAN, H. & RAPALINI, A. 2001. Marco
Juan, Argentina. In: BUATOIS, L. A. & MANGANO, paleogeografico. In: ARTABE, A. E., MOREL, E. M.
M. G. (eds) Icnologia: Hacia una convergencia & ZAMUNER, A. B. (eds) El Sistema Triasico en la
entre geologia y biologia. Asociacion Paleonto- Argentina. Fundacion Museo de La Plata, La
logica Argentina, Publication Especial, 9 17-33. Plata, Argentina, 255-267.
METZ, R. 1996. Newark Basin Ichnology: the Late RETALLACK, G. J. 1983. Late Eocene and Oligocene
Triassic Perkasie Member of the Passaic For- paleosols from Badlands National Park, South
mation, Sanatoga, Pennsylvania. Northeastern Dakota. Geological Society of America, Special
Geology & Environmental Sciences, 18, 118-129. Papers, 193, 1-82.
MIDTGAARD, H. H. 1996. Inner-shelf to lower-shore- RETALLACK, G. J. 1990. Soils of the Past. Unwin
face hummocky sandstone bodies with evidence Hyman, Boston.
for geostrophic influenced combined flow, Lower RODRIGUEZ-ARANDA, J. P. & CALVO, J. P. 1998. Trace
Cretaceous, West Greenland. Journal of Sedimen- fossils and rhizoliths as a tool for sedimentological
tary Research, 66, 343-353. and palaeoenvironmental analysis of ancient
MILANA, J. P. 1998. Anatomia de parasecuencias en un continental evaporite successions. Palaeogeo-
lago de rift y su relation con la generation de graphy, Palaeoclimatology, Palaeoecology, 140,
hidrocarburos, cuenca triasica de Ischigualasto, 383-399.
San Juan. Revista de la Asociacion Geologica SEEGERS-SZABLEWSKI, G. & ISBELL, J. L. 1997. Strati-
Argentina, 53, 365-387. graphy and depositional environments of Permian
MILANA, J. P. & ALCOBER, O. 1994. Modelo tectosedi- postglacial rocks exposed between the Byrd and
mentario de la cuenca Triasica de Ischigualasto Nimrod Glaciers. Antarctic Journal of the United
(San Juan, Argentina). Revista de la Asociacion States, 32, 15-17.
Geologica Argentina, 49, 217-235. STANISTREET, I. G., LE BLANC SMITH, G. & CADLE, A. B.
MILLER, M. F. & SMAIL, S. E. 1996. Permian and Tri- 1980. Trace fossils as sedimentological and
assic biogenic structures, Shackleton Glacier and palaeoenvironmental indices in the Ecca Group
Mount Weaver areas, Transantarctic Mountains. (Lower Permian) of the Transvaal. Transactions
Antarctic Journal of the United States, 31, 5-7. Geological Society of South Africa, 83, 333-344.
MILLER, M. F. & SMAIL, S. E. 1997. A semiquantitative STIPANICIC, P. N. & BONAPARTE, J. F. 1979. Cuenca
field method for evaluating bioturbation on bed- triasica de Ischigualasto-Villa Union (provincias
ding planes. Palaios, 12, 391-396. de San Juan y La Rioja). In: TURNER, J. C. M.
MILLER, M. F., COLLINSON, J. W. & FRISCH, R. A. (ed.) 2ndSimposio de Geologia Regional Argentina.
1991. Depositional setting and history of a Academia Nacional de Ciencias, Cordoba, 1, 523-
Permian post-glacial black-shale: Mackellar For- 575.
mation, Central Transantarctic Mountains. In: TANKARD, A. J., ULIANA, M. A. et al. 1995. Tectonic
ULBRICH, H. & ROCHA CAMPOS, A. C. (eds) Gond- controls of basin evolution in southwestern
wana Seven Proceedings. Institute de Geociencias, Gondwana during the Phanerozoic. In: TANKARD,
Sao Paulo, 201-215. A. J. SUAREZ SORUCO, R. & WELSINK, H. J. (eds)
MILLER, M. F., MCDOWELL, T., SMAIL, S. E., SHYR, Y. Petroleum Basins of South America. American
& KEMP, N. R. 2002. Hardly used habitats: dearth Association of Petroleum Geologists, Memoirs,
and distribution of burrowing in Paleozoic and Tulsa, Oklahoma, 62, 5-52.
Mesozoic stream and lake deposits. Geology, 30, TREWIN, N. H., MACDONALD, D. I. M. & THOMAS,
527-530. C. G. C. 2002. Stratigraphy and sedimentology
MOSLOW, T. F. & PEMBERTON, S. G. 1988. An of the Permian of the Falkland Islands: litho-
integrated approach to the sedimentological ana- stratigraphic and palaeonvironmental links with
lysis of some Lower Cretaceous shoreface and South Africa. Journal of the Geological Society,
delta front sandstone sequences. In: JAMES, D. P. London, 159, 5-19.
& LECKIE, D. A. (eds) Sequences, Stratigraphy, TYE, R. S. & COLEMAN, J. M. 1989. Depositional pro-
Sedimentology: Surface and Subsurface. Canadian cesses and stratigraphy of fluvially dominated
Society of Petroleum Geologists, Memoirs, lacustrine deltas: Mississippi delta plain. Journal
Calgary, Alberta, 15, 373-386. of Sedimentary Petrology, 59, 973-996.
354 R. N. MELCHOR

ULIANA, M. A. & BIDDLE, K. T. 1988. Mesozoic- of Sedimentologists, Special Publications, Oxford,


Cenozoic paleogeographic and geodynamic evolu- 2, 225-239.
tion of Southern South America. Revista Brasileira VISSER, J. N. J. 1993. Sea-level changes in a back-arc-
Geociencias, 18, 172-190. foreland transition: the late Carboniferous-Per-
ULIANA, M. A., BIDDLE, K. T. & CERDAN, J. 1989. mian Karoo Basin of South Africa. Sedimentary
Mesozoic extension and the formation of Argen- Geology, 83, 115-131.
tine sedimentary basins. In: TANKARD, A. J. & WALTER, H. & SUHR, P. 1998. Lebensspuren aus
BALKWILL, H. R. (eds) Extensional Tectonics and kaltzeitlichen Bandersedimenten des Quartars.
Stratigraphy of North Atlantic Margins. American Abhandlungen des Staatlichen Museums fur Miner-
Association of Petroleum Geologists, Memoirs, alogie und Geologie zu Dresden, 43/44, 311-328.
Tulsa, Oklahoma, 46, 599-614. ZAVATTIERI, A. M. & MELCHOR, R. N. 1999. Estudio
VAN DIJK, D. E., HOBDAY, D. K. & TANKARD, A. J. palinologico preliminar de la Fm. Ischichuca
1978. Permo-Triassic lacustrine deposits in the (Triasico), en su localidad tipo (Quebrada de
Eastern Karoo Basin, Natal, South Africa. In: Ischichuca Chica), provincia de La Rioja,
MATTER, A. & TUCKER, M. E. (eds) Modern and Argentina. Asociacion Paleontologica Argentina
Ancient Lake Sediments. International Association Publicacion Especial, 6, 33-38.
An approach to the description and interpretation of
ichnofabrics in palaeosols

JORGE F. GENISE1, E. S. BELLOSI2 & M. G. GONZALEZ2


1
CONICET, Museo Paleontologico Egidio Feruglio, Av. Fontana 140, 9100 (Trelew),
Chubut, Argentina (e-mail: jgenise@mef.org.ar)
CONICET, Laboratorio de Icnologia, Museo Argentina de Ciencias Naturales,
Av. Angel Gallardo 470, (1405) Buenos Aires, Argentina

Abstract: Studies on ichnofabrics have focused mainly on marine environments. Attempts to


apply the ichnofabric methodology and theoretical framework to continental deposits bearing
palaeosols are few and poorly developed. Methodologies analysed in this contribution include
the applicability of current ichnofabric indexes and diagrams, the assessment of the destruction
of the original bedding by ichnofabrics and by other soil characters separately, and the relation-
ships between different stages of palaeosol and ichnofabric development. Many soil features
may be formed without the intervention of bioturbation, or may be the result of interactions
of physical, chemical and biological processes, in which traces of organisms may have only a
subsidiary role. Ichnofabrics can be well developed in palaeosols devoid of other soil characters
and, conversely, palaeosols showing a well-developed soil structure can bear almost no trace
fossils. This fact adds a third component to classical methods that normally consider only
original bedding and ichnofabrics. Theoretical analysis includes the possibility of recording
composite ichnofabrics in palaeosols, and the value of individual ichnotaxa as possible
indicators of subaerial conditions and environmental changes. The ecological preferences
and requirements of trace-makers provide the key to understanding composite ichnofabrics;
however, only complex traces can be certainly attributed to particular modern taxa. Insect
nests, pupal chambers and earthworm burrows are the most reliable indicators of subaerial
exposure and, in many cases, very particular environmental conditions.

Ichnofabric analysis has become an increasingly analyse the utility of different aspects of ichno-
important tool in ichnological analysis, provid- fabric in continental deposits bearing palaeosols.
ing a methodology for documenting and compar- In order to apply the ichnofabric method to
ing the extent of bioturbation and relative palaeosols the effects of bioturbation must first
chronology of infaunal tiering (Ausich & Bottjer be distinguished from physical and chemical
1982; Ekdale & Bromley 1983, 1991; Bromley & pedogenic processes. Other aspects required for
Ekdale 1986; Droser & Bottjer 1986; Bottjer & complete palaeosol description are the interrela-
Droser 1991, 1994; Taylor & Goldring 1993; tionships between different stages of palaeosol
Taylor et al. 2003). Most studies of ichnofabrics and ichnofabric development. Another particu-
have been focused on marine environments lar point to be considered herein in detail is the
(Taylor & Goldring 1993; Bottjer & Droser value of the individual components of palaeosol
1994). Few, poorly developed, attempts have ichnofabrics as possible indicators of subaerial
been made to apply ichnofabric methodology conditions and changes in environment, and the
and its concepts to continental deposits bearing possibility of recording composite ichnofabrics
palaeosols. The few references involve the utiliza- (sensu Bromley & Ekdale 1986) in palaeosols.
tion of the ichnofabric index (Droser & Bottjer
1986) as a comparative scale to evaluate the
degree of development of soil structure (Retal- Methodology
lack 1990), tiering in palaeosols (Hasiotis et al.
1993; Hasiotis & Dubiel 1994; Gonzalez et al. Review of existing methodology in
1998), the proposal of informal names for ichnofabric analysis
ichnofabrics produced by particular groups of
arthropods (Hasiotis 1997), and the use of the There are different methods for measuring the
term terrestrial ichnofabrics to describe trace extent of bioturbation and describing ichno-
fossil assemblages in palaeosols (Miller & fabrics, which are reviewed elsewhere (Taylor &
Mason 2000). Goldring 1993; Bottjer & Droser 1994; Miller &
The aim of this contribution is to present Smail 1997; Netto 2000). Two basic philosophies
examples of palaeosol ichnofabrics in order to are recognized: those that use a semi-quantitative
From: MclLROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 355-382. 0305-8719/04/S 15.00 © The Geological Society
of London.
356 J. F. GENISE ET AL.

approach (e.g. Reineck 1963; Droser & Bottjer Methodology for palaeosol ichnofabric
1986; Miller & Small 1997) and those that are analysis
mostly descriptive (e.g. Ekdale & Bromley
1991; Wetzel & Uchman 1998). The case studies presented herein involve com-
Semi-quantitative methods recognize different plex ichnofabrics, small-scale, high-resolution
categories of ichnofabric (ichnofabric indices or observations and detailed palaeoenvironmental
bioturbation indices) based on the degree of analysis. The inherent complexity of soil fabrics
disruption of the original bedding by biotur- is such that conventional methodologies were
bation, which are represented in schematic dia- deemed inadequate, and some modifications to
grams developed for different sedimentary previous descriptive methods are introduced for
settings (Droser & Bottjer 1986; Bottjer & their description and as an aid to their analysis.
Droser 1994; Miller & Smail 1997). Most of Additionally, the bioturbation indices proposed
them have been developed for studying ichno- by Taylor & Goldring (1993) have been used to
fabrics in vertical sections, whereas that of complete the descriptions, as well as a semi-
Miller & Smail (1997) was elaborated for quantitative approach to quantification of bio-
bedding planes. In turn, descriptive diagrams turbation of each tier used for composite
(tiering diagrams, TD) were proposed to illus- ichnofabrics (as suggested by Ekdale & Bromley
trate relationships of ichnofabrics with palaeoen- 1991). Seven grades of BI, based on Reineck
vironmental factors, rate of sedimentation, and (1963), were recognized by Taylor & Goldring
biological activity (e.g. Ekdale & Bromley 1991; (1993), which were adapted herein for descrip-
Pollard et al 1993; Wetzel & Uchman 1998 and tion of palaeosols according to burrow density
references therein) at particular environments and the amount of burrow overlap. These BI
or localities. A comprehensive approach to com- are: 0, no bioturbation; 1, sparse bioturbation,
bine in a single analysis the evaluation of the few discrete traces; 2, low bioturbation, low
extent of bioturbation (bioturbation index, BI), trace density; 3, moderate bioturbation, traces
as in semi-quantitative methods, with a visual discrete, overlap rare; 4, high bioturbation,
representation of ichnofabrics (ichnofabric con- high trace density with overlap common; 5,
stituent diagram, ICD) was proposed by Taylor intense bioturbation, later burrows discrete;
& Goldring (1993). The ICD considers the ichno- and 6, complete bioturbation, repeated over-
taxa present, ichnodiversity, density and order of printing. The previous authors also utilized the
emplacement. sharpness of the primary sedimentary fabric to
However, no single method has been widely define the grades (Taylor & Goldring 1993).
accepted. Critical analysis of the applicability However, in palaeosols the original bedding
of different methods has been made elsewhere can also be heavily disrupted by other soil
(e.g. Frey & Wheatcroft 1989; Ekdale & Brom- processes that are non-biological, or only par-
ley 1991; Pemberton et al 1992; Miller & Smail tially related to bioturbation: thus this disruption
1997). Some authors stated that ichnofabric of the sedimentary fabric is not exclusively
indices may be useful for describing simple related to bioturbation as in the classical
ichnofabrics, whereas descriptive methods are marine examples.
more suitable for complex ones (e.g. Ekdale In both the semi-quantitative and descriptive
& Bromley 1991). Others concluded that methods only two variables are considered: the
semi-quantitative methods are more appropri- original bedding and the ichnofabric. In palaeo-
ate for large-scale studies, involving hundreds sols, soil structures other than ichnofabrics
of box cores or thousands of metres of strati- should be also considered as destructive agents
graphic section, whereas the descriptive of the original bedding. Different soil features
method has been used with core material or that can be recognized in palaeosols, such as
outcrops of limited size (e.g. Miller & Smail horizons, peds, cutans, glaebules, crystals, and
1997). In addition, Taylor & Goldring (1993) root and animal traces, modify the original
claimed that descriptive methods provide the sedimentary fabric of the deposits in which soils
possibility of assessing and comparing a more develop (Teruggi 1971; Yaalon 1971; Andreis
complete set of data, including also the 1981; Bown & Kraus 1987; Retallack 1990;
degree of bioturbation contemplated in the Nettleton et al, 2000). Root and animal traces
semi-quantitative method. In practice, however, are the direct products of the activity of organ-
the methodology is so time consuming that its isms, and as such they are considered trace
use is restricted, in most cases, to describing fossils and deserve an ichnotaxonomical treat-
key stratigraphic surfaces or summarizing ment, whereas other soil characters are the
recurring ichnofabrics within a stratigraphic result of interactions of purely physical-chemical
unit (e.g. Mcllroy 2004b). processes, or mixed physical-chemical-biological
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 357

processes, in which traces of organisms may only whereas the last two stages (4, 5) are defined
be involved to different extent (e.g. Brewer 1976; mostly by the increasing thickness of A and B
Fitzpatrick 1984; Retallack 1990). Regarding horizons. In terms of BI, a highly developed
Ekdale & Bromley's (1983) definition of ichno- palaeosol will comprise volumetrically high
fabrics, it would be possible to consider the percentages of pedofabric and, accordingly, low
whole soil structure as an ichnofabric because it to moderate percentages of primary sedimentary
may be a direct or indirect result of bioturbation. fabric and ichnofabrics. Hence the single BI
However, soil features such as peds, cutans, value does not necessarily indicate the percentage
glaebules and crystals may be formed as well, of disruption of the original bedding (by animals
without the necessary intervention of biotur- and plants), and conversely the single percentage
bation (e.g. Buol et al 1990; Retallack 1990). of disruption is not a true reflection of the degree
For instance, Blokhuis et al. (1990) stated that of bioturbation. In palaeosols, the proportions of
Vertisols are typical cases of ped formation by all three processes must be properly described in
physical processes of shear failure, along inclined order to analyse meaningfully the fabric of the
surfaces, and cracking. Repeated wetting and deposits in which they occur. A comparison of
drying of a dispersed cracking clay soil results five study cases is presented herein utilizing a
in the fragments parting into fine wedge-shaped pedofabric/ichnofabric ternary diagram (PITD).
aggregates, because of the stress-strain regime The PITD (Fig. 1) includes the bioturbation
of the swelling soils (Blokhuis et al. 1990). index (BI), and percentages of original bedding
Even when the traces of organisms would have (OB), ichnofabrics (IF) and pedofabrics (PF).
been involved in the origin of soil characters (e.g. BI and percentage of IF were measured following
granular and crumb peds), they might be Taylor & Goldring's (1993) adapted table based
unpreserved or so distorted as to become recog- on burrow density and overlap. The percentage
nizable, thereby precluding an ichnotaxonomical of the original bedding (OB) was measured
approach. Apart from its formal definition, the according to the preservation of primary sedi-
common usage of the ichnofabric concept mentary structures. The pedofabric (PF) was
involves fabrics completely produced by traces calculated considering the development of soil
of organisms, more commonly than fabrics in features other than the recognizable trace fossils.
which traces are only partially involved. Finally, Two types of diagram have been proposed for
the development of an ichnofabric may be inde- representing descriptive methodologies: tiering
pendent of other soil characters. Ichnofabrics diagrams (TD) (e.g. Ekdale & Bromley 1991)
can be well developed in immature palaeosols, and ichnofabric constituent diagrams (ICD) pro-
obliterating the original bedding almost com- posed by Taylor and Goldring (1993). The ICD
pletely (e.g. Genise & Bown 1994b). Conversely, represents events consecutively, starting with
palaeosols showing a well-developed ped struc- the primary fabric and then ichnotaxa in order
ture may preserve almost no trace fossils of emplacement. This order reflects the infaunal
(Melchor et al. 2001). In conclusion, the fabric succession produced by changes in the environ-
directly and completely produced by root and mental conditions (e.g. Bromley & Ekdale 1986;
animal traces is considered herein to be the ichno- Buatois et al. 1997). In turn, the original tiering
fabric, whereas the fabric produced by other soil - the vertical partitioning of the habitat in
characters, resulting from the interactions of response to environmental gradients (e.g.
physical, chemical and biological processes, is Ausich & Bottjer 1982; Bromley & Ekdale
termed herein the pedofabric. It should be noted 1986) - may be not represented if it is not the
that pedofabric is distinct from the terms 'soil result of the order of emplacement. In marine
fabric' and 'soil microfabric' (e.g. Brewer 1976; environments, where colonization of substrates
Fitzpatrick 1984; Bullock et al. 1985; Retallack may progress from the surface downwards, the
1990; Buol et al. 1990), which are applied deepest tier is also the last emplaced (Bromley
mostly to the microstructure of the fine-grained & Ekdale 1986; Ekdale & Bromley 1991; Brom-
part of the soils. The stages of palaeosol develop- ley 1994). In these cases ICDs may represent tier-
ment may be used to evaluate the degree of ing and order of emplacement simultaneously.
pedofabric development. There are two scales However, in palaeosols the first colonizers of
for estimating stages of palaeosol development, the deposit may belong to the deepest tier, as
which were proposed by Bown & Kraus (1987) shown herein. Both original tiering (Hasiotis &
and Retallack (1988) respectively. Both scales Dubiel 1994; Gonzalez et al. 1998) and infaunal
are broadly based on similar criteria. The first succession (Hasiotis et al. 1993) have been
three stages are characterized by (1) no horizon recorded from palaeosols, and so descriptive
formation; (2) A horizon and incipient B hori- diagrams such as TDs, which can illustrate
zon, and (3) A and B horizons well defined, both features of ichnofabrics simultaneously,
358 J. F. GENISE ET AL.

Fig. 1. Proposed diagram for assessing palaeosol ichnofabric, pedofabric, original bedding and grades of
bioturbation index, BI. 1, Ischigualasto Formation, well-developed Vertisols; 2, Ischigualasto Formation,
poorly developed Vertisols; 3, Laguna Palacios Formation, Entisols; 4, Laguna Palacios Formation, Alfisols;
5, Asencio Formation, Ultisols; 6, Jebel Qatrani Formation, Inceptisols; 7, Sarmiento Formation, Andisols;
8, Sarmiento Formation, Alfisols.

are of fundamental importance. In addition, per- of the seafloor or major changes in palaeo-
centages of bioturbation, which can sometimes environmental conditions produce either the
be time consuming to calculate, are instrumental upward migration of the communities or a shift
for the construction of the ICDs (Taylor & from one to another. This is reflected in the
Goldring 1993). In TDs, the thickness of the juxtaposition of tiers producing composite ich-
study section, ichnotaxa, tiering and cross- nofabrics (Bromley & Ekdale 1986; Mcllroy
cutting relationships are readily understandable 2004a). In modern soils the tiering of organisms
(e.g. Bromley & Ekdale 1986). is also a well-known phenomenon (e.g. Hasiotis
In conclusion, the study cases presented herein & Bown 1992), but scarcely described from
are represented in TDs in which the BI of palaeosols. A few examples of infaunal changes
different tiers is indicated, the pedofabric is and tiering in palaeosols have been documented
depicted along with and independently from the until now, which show the importance of soil
ichnofabric, and are completed with the com- moisture and palaeo-water table as controls on
positional diagram (Fig. 1) showing total organism distribution (Hasiotis et al. 1993;
percentages of bioturbation, pedofabric and Hasiotis & Dubiel 1994).
original bedding. There are different sedimentological, palaeo-
pedological, geochemical and ichnological indi-
cators that can contribute to the understanding
Theoretical background of fluctuations of ancient water tables (e.g.
Bown & Kraus 1983; Retallack 1990; Hasiotis
One of the most important results of the study of et al. 1993). The development of independent
ichnofabrics is the recognition of the tiered pat- ichnological criteria, which allow comparisons
tern of organisms in substrates and subsequent with data gathered from other sources, may be
modifications of it through time in relation to based on two different approaches. One method-
environmental change (Ausich & Bottjer 1982; ology, proposed by Bown & Kraus (1983), is
Bromley & Ekdale 1986). The vertical stacking based on the detailed record and comparison of
of ichnofossils, or tiering, is the response of the occurrence of trace fossils in particular
trace-makers to biological, physical and chemical palaeosol types and horizons. This approach
gradients within a substrate (Bromley & Ekdale is particularly important when dealing with
1986). In the marine realm, the vertical accretion trace fossils whose producers are unknown or
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 359

uncertain. Regrettably, papers that analyse trace has received criticism and general lack of accep-
fossils and palaeosols with such a degree of detail tance (e.g. Engel 2001; Genise 2004).
are too few to provide a broad reference database The compiled information on recorded trace
(e.g. Down & Kraus 1983; Hasiotis et al 1993; fossil assemblages from palaeosols shows that
Hasiotis & Dubiel 1994; Retallack 200 Ib; they are composed of insect nests and pupation
Genise et al. 2002). A second evaluation of chambers, earthworm, millipede and crayfish
palaeoenvironmental changes involves the burrows, root traces and other ichnofossils
recognition of the producers of ichnotaxa whose affinities are obscure (Genise et al. 2000,
recorded from palaeosols. If the trace-maker table 2; Retallack 2001b). Insect nests as a
can be identified, the ecological preferences and whole are the most common trace fossils in
requirements of that taxon can then provide the palaeosols, comprising a distinct ethological
key to understanding composite ichnofabrics. category termed calichnia (Genise & Bown
A word of caution on the attribution of trace 1994a). These authors explained their abundance,
fossils to modern groups should be introduced arguing that nests are constructed structures and
before an analysis. Trace fossils range from not merely excavated ones, and consequently
very simple to very complex morphologies. have a high preservational potential. The nesting
Along this spectrum, the more complex the activities involve the provision of nests with
trace, the more reliable will be its attribution to different kinds of organic matter to feed larvae
a particular modern taxon. The principle of by adults. Both provisions and larvae are very
special quality of behavioural homology sensitive to soil temperature and moisture,
(Wenzel 1992) states that the more complicated requiring very specific environmental conditions
the behaviour, the stronger will be the postulate (e.g. Michener 1979; Grasse 1984; Cane 1991;
of homology. Behavioural homologies are Hanski & Cambefort 1991; Genise 1999).
defined as those behaviours that are similar Adults achieve these conditions basically in two
because of their origin in a common ancestor, ways: (1) by the morphology and constructional
whereas analogies are those similarities that are materials of nests and breeding chambers, and
shown by groups that are not phylogenetically (2) by selecting specific nesting sites and soil
related (Wenzel 1992). To extrapolate the ecolo- depths to locate them. Constructed walls and lin-
gical preferences and requirements of modern ings, along with the general morphology of nests,
organisms to the producers of trace fossils, it is provide the necessary isolation from soil to main-
necessary to postulate a close phylogenetic rela- tain a suitable temperature and humidity inside
tionship for both. Only behavioural homologies, breeding chambers (e.g. Halffter & Edmonds
confirmed by macro-and micromorphological 1982; Grasse 1984). Accordingly, these morpho-
studies and/or the high complexity of traces, logical devices increase the complexity of the
are useful for postulating this phylogenetic whole structure, providing the diagnostic charac-
relationship between a modern and a fossil ters for recognition of behavioural homologies
trace-maker. In subaerial settings, insect nests that permit the attribution to particular insect
and pupation chambers, and earthworm, milli- taxa (e.g. Genise 1999). In contrast, other struc-
pede and crayfish burrows are complex enough tures excavated by different groups of insects
and/or have been well studied macro- and micro- for shelter, feeding or dwelling are in most
morphologically to satisfy this special quality cases more simple, and these analogous struc-
criterion (e.g. Bown & Kraus 1983; Hasiotis & tures cannot be attributed to any particular
Mitchell 1993; Genise et al 2000; Retallack invertebrate taxa (e.g. Ratcliffe & Fagerstrom
200Ib). Conversely, other trace fossils in palaeo- 1980).
sols remain to be studied in more detail because Hence, for the purpose of palaeoenvironmen-
their morphologies are very simple and can be tal analysis, trace fossils that can be certainly
attributed indistinctly to different groups of attributed to particular invertebrate taxa and
organisms (e.g. Ratcliffe & Fagerstrom 1980). those whose attribution is doubtful should be
In modern groups, other criteria that cannot be analysed separately. Some broad-scale relation-
used with trace fossils are also employed to ships between emplacement of trace fossils and
establish behavioural homologies (Wenzel position of an ancient water table can be easily
1992). In turn, a certain degree of congruence addressed knowing the physiological require-
with the body fossil record can be proposed as ments of producers. For instance, insects that
an additional ichnological criterion to postulate are air breathers will inhabit the soil above the
homologies. For instance, the proposal that water table, whereas crayfishes and other
Triassic bee cells (Hasiotis et al. 1995) largely crustaceans, which need free water to breathe,
predate the appearance of angiosperms and the live in burrows under the water table or con-
oldest fossil bees, known from the Cretaceous, nected to watercourses (Retallack 1990; Hasiotis
360 J. F. GENISE ET AL.

& Mitchell 1993). However, a more accurate during Ischigualasto times. Trunk and more
picture of soil moisture preferences and tiering steady braided rivers occupied the central area
can be achieved by analysing particular groups of the Ischigualasto Park, whereas in northern
of trace-makers involved within the context of and southern areas floodplains were better
the resultant ichnofabric. developed (Milana et al 1998).
Previous workers (Milana & Alcober 1994)
recognized the presence of 'root traces, bioturba-
Selected study cases tion and burrows' at many horizons. In addition,
Melchor et al (2001) stated that Ischigualasto
The few examples presented herein were selected palaeosols exhibit vertic features, tabular root
particularly (1) to test the application of present systems and drab colours. Palaeosols of the
methodology, (2) to show the independent devel- lower section are better drained, and present
opment of ichnofabrics and pedofabrics, (3) to calcic and mottled horizons, whereas those of
demonstrate tiering in ancient soils, and (4) to the upper section are more clayey and rooted
illustrate the presence of composite ichnofabrics (Martinez et al 1998). In general, these alluvial
in palaeosols. It is not the purpose of this con- palaeosols are simple and show moderate to
tribution to review the palaeoenvironmental poor maturity, in accordance with rapid sedi-
conditions or any other aspects of the sedimen- mentation deduced from the presence of fresh
tary sequences included in the study cases. In feldspars (Milana & Alcober 1994). Most
the case of the Jebel Qatrani Formation from mudrocks exhibit greyish to greenish hues,
Egypt, only one of us (JFG) conducted ichno- whereas some beds are reddish, indicating
logical research at that locality: hence the better-drained conditions. Fossil amphibians,
sedimentological and palaeoenvironmental set- primitive dinosaurs and therapsids are abundant,
ting is taken exclusively from the literature and occur together with diverse plant remains
(Bown 1982; Bown & Kraus 1988). However, belonging to the Dicroidium Flora (Kokogian
the importance of this case study for the pre- et al. 1999; Zamuner et al. 2001). Vegetation
viously mentioned purposes of this contribution along the margins of fluvial channels was
necessitates its inclusion herein. dominated by subtropical (seasonal) evergreen
and sclerophyllous forests, with low diversity,
whereas on floodplains a herbaceous-arbustive
Palaeosols from the Ischigualasto Formation community developed (Artabe et al. 2001).
(Argentina) The two studied palaeosols are intercalated
in a small-scale (10m thick in average) sequence
The Ischigualasto Formation (lower Late Tri- in the upper section of the formation (Fig 2).
assic) is a continental unit of the Ischigualasto- Several of these sequences are intercalated
Villa Union Basin, which crops out in San Juan between main channel and typical floodplain
Province, western Argentina (Stipanicic 2001). deposits, and are associated with carbonaceous
It is composed mainly of grey claystones, green- mudstones deposited in lacustrine/backswamp
ish tuffaceous siltstones and bentonites, carbon- environments and fine- to medium-grained
aceous mudstones and few thin coal beds sandstones that accumulated in levees and as
intercalated with fine- to coarse-grained, cross- splay lobes, or in subordinated channels. Such
bedded sandstones (Stipanicic & Bonaparte sequences are comparable to avulsion belt
1979; Bellosi et al 200la). At the surveyed deposits (Kraus & Gwinn 1997; Farrell 2001),
localities in the Ischigualasto Provincial Park, which generally support high sedimentation
the fluvial facies are organized in fining-upward rates (Kraus 1996; Kraus & Asian 1998).
cycles formed by channelized sandstone bodies One example is a 60 cm thick, olive grey (5Y5/
with trough cross-bedding and downstream 1), smectitic clay-rich palaeosol (Figs 2, 4a) in
and lateral accretion surfaces, grading to thick which no relict of the original bedding is recog-
floodplain deposits (Fig. 2). General alluvial nized. Pedofabric is uniform and characterized
architecture and the anatomy of sandstone by small, angular blocky peds (wedge-shaped)
bodies indicate an avulsion-controlled fluvial and closely spaced slickensides. The palaeosol
system, having perennial, moderate to high shows no horizonation, and it is eroded at top
sinuosity, channels, mixed load and frequent by a medium-grained sandstone with convolute
splays (Bellosi et al. 2001a; Spalletti 2001). bedding accumulated in a minor avulsion
Floodplains were predominantly poorly drained, channel. Micromorphologically it has a well-
favouring the development of palustrine condi- defined blocky microstructure (sensu Bullock
tions locally (Milana & Alcober 1994). Land- et al. 1985) and abundant connected streaks
scape and basin configuration varied slightly of bright clay, which are aligned around ped
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 361

Fig. 2. (a) Location of stacked vertisols (asterisks) in a coarsening-upward avulsion sequence (CU Seq) of the
upper section of the Ischigualasto Formation. Arrows show position of Vertisols with in situ Equisetales stems.
The CU sequence is formed by suspension clay deposits (SC), splay sand lobes (SL) with ripple lamination and
minor sandstone channels (Ch) with deformed trough cross-bedding. Tiering diagrams from well-developed
Vertisols (b) and poorly developed ones (c) in this Formation.
362 J. F. GENISE ET AL.

surfaces. Truncated stems of equisetales in life- structures abruptly covered by a primary white
position filled by green sandstone represent the tuff, suggesting non-permanent streams. Most
only bioturbation (BI1) (Genise et al 2001; of the pyroclastic deposits (60% in thickness)
Fig. 4a). This palaeosol probably underwent are pedogenized, commonly occurring as stacked
advanced pedogenic homogenization. Thus it is non-calcic palaeosols (Sciutto 1981; Bellosi et al.
interpreted as a very well-developed Vertisol, as 2002a). Detailed sedimentary, ichnologic and
the original bedding is totally obliterated by the pedogenic features were recently published by
pedofabric. Genise (2000) and Genise et al. (2002). Well-
Another example is a less-developed Vertisol, developed and more frequent palaeosols occur
where the original bedding is clearly recognized in the middle and upper members (Sciutto
(Figs 2, 4b). The upper 20cm are composed of 1981). The Laguna Palacios Formation is
a very coarse siltstone, light olive grey (5Y 6/1), considered to be a volcaniclastic-influenced,
having relict ripple lamination and less- loess-palaeosol sequence, formed in a temperate
developed short slickensides and wedge-shaped subhumid/-seasonal climate and relatively dry
peds. In the lower 20cm wedge-shaped clay conditions (Bellosi & Sciutto 2002; Bellosi et al.
peds and slickensides are also present, but a 2002a; Genise et al. 2002). Fluctuations of the
fine lamination is still preserved. Bioturbation water table, suggested by scarce hydromorphic
is higher than in the case described previously features such as Fe-Mn nodules (Genise et al.
(BI2). Ichnofabric is composed of equisetales 2002), are also consistent with a fossil stump in
stems and Skolithos linearis in the upper siltstone growth position of the Middle Member, which
(Genise et al. 2001; Fig. 4b). The preservation of would have required rapid burial in a water-
parallel and ripple cross-lamination suggests a logged habitat (Driese et al. 1997; Retallack
poorly developed Vertisol. 200 la).
The depositional setting of the Laguna
Palacios Formation can be compared to the
Palaeosols from the Laguna Palacios Pleistocene Pampean loess plains, with few
Formation (Argentina) rivers and ponds, and savannah or prairie soils
with herbaceous or low vegetation (Teruggi
The Laguna Palacios Formation is an orange to 1957; Iriondo 1999; Sayago et al. 2001). Most
reddish brown Upper Cretaceous (Santonian- sediments were supplied as distal volcanic ash
Maastrichtian) tuffaceous succession, which falls and subordinately by fluvial streams
constitutes the upper part of the Chubut Group (Genise et al. 2002). The development of
(Feruglio 1949; Sciutto 1981). It records terres- this loess-palaeosol sequence was regulated by
trial sedimentation at the northwest periphery the balance supply versus pedogenesis, which
of the San Jorge Basin and north Deseado ultimately could have been governed by
Massif, in central Patagonia, Argentina (Bellosi fluctuations in the climate regime (Bellosi et al.
& Sciutto 2002). Palaeogeographic reconstruc- 2002a).
tion and thickness variation indicate that it The first case of palaeosol ichnofabrics shown
accumulated on basin margin topographic herein occurs in a vitric Entisol from the Upper
highs (Sciutto 1981). The Laguna Palacios For- Member (Genise et al. 2002, Fig. 3B, 4B; Figs
mation is divided into lower, middle and upper 3, 4c). The parent material is very fine (clay size
members (Sciutto 1981). The only recorded grade) vitric volcanic ash. Glass shards are
body fossils from this formation are a few entire and fresh or slightly weathered. This iso-
stems and trunks (Sciutto 1981). Two main lated bed, 60cm thick, moderately bioturbated,
facies type are recognized where the unit reaches and very pale orange (10YR8/2), is a distinct
its maximum thickness (Bellosi & Sciutto 2002). light-coloured pyroclastic deposit interbedded
Pyroclastic facies predominate, including mas- in a suite of brownish palaeosols. The original
sive sheet strata of vitrie-crystal tuffs, very fine bedding of this air-fall ash is massive or crudely
tuffs with accretionary lapilli, and bioturbated stratified, and pedofabric is wholly absent.
tuffaceous mudstones and claystones (Sciutto Micromorphologically no soil feature is present,
1981; Genise et al 2002). The remaining facies and no crystalline mineral was identified in XRD
are fining-upward cross-bedded pyroclastic sand- analysis. The ichnofabric shows two tiers (Fig.
stones and scarce intra-formational conglomer- 3). The upper tier is dominated by Taenidium
ates, which are subordinate, albeit significant, barretti and a few root traces, resulting in a
because they are mostly restricted to incised moderate bioturbation (BI3). The lower tier
channels. Several of these channel bodies, in the includes fossil bee nests, Cellicalichnus chubuten-
middle and upper members, comprise a thin sis, and also a few root traces, resulting in low
basal siliciclastic sandstone with (fluvial) current intensities of bioturbation (BI 2) (Genise et al.
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 363

Fig. 3. (a) Section of the upper member of the Laguna Palacios Formation containing two different palaeosol
types. These facies are included in a 300m thick pyroclastic loess-palaeosol succession, with intercalated fluvial
channel deposits, (b) Entisol (star) developed in very fine vitric tuff (volcanic dust), (c) Intensely bioturbated
Alfisol (asterisk) developed in tuffaceous sandstones.
364 J. F. GENISE ET AL.

Fig. 4. (a) Truncated stem of equisetales in growth position (left of hammer), in a well-developed Vertisol.
Ischigualasto Formation, (b) Abundant Skolithos in a poorly developed Vertisol, which preserves ripple
lamination. Ischigualasto Formation. Scale bar: 1 cm. (c) Eroded surface of a vitric Entisol showing fossil bee
nests (Cellicalichnus chubutensis) tunnels in relief. Laguna Palacios Formation. Scale bar: 10cm. (d) Detail of
Cellicalichnus in plan view in the same Entisol. Laguna Palacios Formation. Coin 2 cm. (e) Structured and
moderately bioturbated Alfisol; the pedofabric is composed mainly of blocky peds. Laguna Palacios
Formation, (f) Coleopteran pupal chambers (Rebuffoichnus casamiquelai) from the palaeosol horizon shown in
(e). Laguna Palacios Formation. Lens cap: 5cm.

2002). The nests, which originally would have trace-makers and roots of plants destroyed this
reached the surface of the soil, could not be part of the nests.
traced upwards into the upper tier, suggesting The second case is also included in the Upper
that the later activity of the shallow Taenidium Member, separated by 3m from the previous
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 365

one (Fig. 3). The ichnofabric is present in a well- Bown (1996) interpreted the clasts of Ford's con-
developed Alfisol, included in a sequence of glomerates as palaeosol peds. Finally, Gonzalez
composite tuffaceous palaeosols (Genise et al. et al. (1998) proposed the existence of two differ-
2002, figs 3C, F, 4B). It is the most common ent horizons characterized by different types of
palaeosol type in the Laguna Palacios Forma- ped (prismatic and nodular) and ichnological
tion. The original bedding is only very scarcely content. The abundant ichnological assemblage,
preserved in the upper horizon. The palaeosol composed of 12 ichnogenera, was described by
shows a yellowish to light brown colour (5 YR Roselli (1938, 1987) and redescribed by several
6/4), a well-developed, upper, elluvial, horizon, subsequent authors (Genise & Bown 1996;
having platy peds, and a lower illuvial one, Genise & Hazeldine 1998a, 1998b; Genise &
having angular to subangular blocky peds (Fig. Laza 1998; Genise & Verde 2000).
4e). The original deposit is a light-coloured, This kind of red ferruginous palaeosol, as
horizontal-bedded turfite (vitric medium tuffac- exemplified by the Asencio Formation, is usually
eous sandstone), preserved as a C horizon in produced by a complex of iterative destruction-
some cases. Bioturbation is high in the upper reconstruction stages related to hydromorpho-
horizons (BI4) and low to absent in the C hori- logical processes (Tardy 1992). The original Bt
zon. Recognizable trace fossils are coleopteran horizons of mature Ultisols (Gonzalez 1999) sub-
pupation chambers, attributable to Rebuffoich- sequently supported lateritization and indura-
nus casamiquelai (Fig. 4f), Taenidium barretti, tion (crusts) (Caorsi & Gorli 1958; Ford 1988c),
Skolithos linearis, Beaconites coronus (sensu becoming ferricretes or cuirasses (sensu Nahon
Keighley & Pickerill 1994) and thin root traces. 1976; Fig. 7a, b). Lateral changes in the structure
Rebuffoichnus casamiquelai is not cross-cut by from nodular to vacuolar and massive is normal
the other traces, which suggests that it was the in cuirasses because of iron transfer, leaching or
last emplaced trace. Its occurrence is patchy, in dissolution of kaolinite and quartz grains, forma-
contrast to the other ichnogenera that display a tion of voids, and ferruginization (Nahon 1986).
more homogeneous pattern of lateral distribu- The beds with nodular structure are interpreted
tion in the deposit. herein as being produced by dismantling of
such iron duricusts (Tardy 1992; Tardy &
Roquin 1992; Temgoua 2001; Figs 7a, c). Most
Palaeosols from the Asencio Formation of them are residual deposits showing no
(Uruguay) evidence of transport.
Intense and widespread bioturbation (e.g.
The Asencio Formation is a late Cretaceous- termite activity) is considered to be responsible
Palaeogene red-bed unit cropping out in western for surficial hematite duricrust dismantling
Uruguay (Bossi 1966) and partly in eastern (Eschenbrenner 1986; Grasse 1986). However,
Argentina (Genise & Zelich 2001). It is thin, 5- no undoubtedly termite-made nest structure was
15m in average thickness, and composed of identified. Irregular geometries and transitional
three to six stacked palaeosols, developed on boundaries between the ferricrete-ferricrete lag
sandstones of probable fluvial origin (Genise & are the response of gradual dismantling processes,
Bown 1996; Figs 5a, 7a). Natural outcrops are which may occur at the surface, inside, or at the
small, but the formation is well exposed in bottom of ferricretes (Temgoua 2001). Most
many stone quarries. No body fossils were ferricretes develop under seasonally tropical
recorded from this formation despite its contain- climates: mean annual temperature 30 °C and
ing one of the most diverse, well-preserved and rainfall 1300-1700 mm per year, with several
abundant assemblages of bee and coleopteran dry months (Tardy & Roquin 1992). As they
nests in palaeosols (Genise & Bown 1996). Origi- usually develop along large periods of time (105
nal bedding is nearly absent with the exception of to 106 years) it is probable that the thin Asencio
a few channelized bodies at the base of the unit sequence represents several million years.
(Pazos et al. 1998; Goso Aquilar & Guerequiz Changes in pedoenvironment produce small-
2001) and poorly preserved in trough cross- scale variations in mineralogy and structure.
beds in Dumestre quarry. The outstanding pedo- For instance, red ferralitic soils (with haematite
genized character of this unit was first recognized and kaolinite-haematite micronodules) are
by Ford (1988a, 1988 b), who also proposed that generally located in well-drained upland areas
fossil insect nests were restricted to 'conglomer- (Schwertmann 1988).
ates' [sic] sourced from coeval Oxisols developed In the selected example, two interfingered
in humid tropical conditions. Regarding the ich- horizons are identified, one nodular and poorly
nological content and sedimentological charac- consolidated (dismantling horizon) and the
teristics of the Asencio Formation, Genise & other well indurated, showing columnar
366 J. F. GENISE ET AL.

Fig. 5. (a) Asencio Formation. The originally structured Ultisols contain abundant insect nests, supported
laterization, ferricrete development and later dismantling (intraformational conglomerates), (b) Tiering diagram
shows the different fabrics in the dismantling horizon and ferruginized crust, (c) Stratigraphic interval in the
middle section of Jebel Qatrani Formation showing high-sinuosity fluvial facies and the position of the studied
Inceptisol (star) (taken from Bown 1982). (d) The Inceptisol developed in fine-grained sandstones that preserve
the original bedding originated in the upper part of a channel point-bar, with abundant rhizoliths and fossil
termite nests (Fleaglellius pagodus).

structure (duricrust) in which the original bed- and reddish, and their mean size varies from 3
ding is completely disrupted (Figs 5a, 7a). The to 10cm. The mineral composition and micro-
dismantling horizons are massive, having morphology of the nodules are similar to those
variable quantities of clay matrix (Ford 1988a). of the duricrust, where hematite-kaolinite
Their individual thickness is l-3m, and their aggregates prevail. The smallest nodules are the
geometry is lenticular. The base is, with rare more rounded ones. Clay matrix proportion
exceptions, clearly non-erosive. Lateral bound- also varies from 10% to 60%. Matrix percentage
aries of the columnar horizons are always and roundness increase as nodule size decreases.
transitional and irregular (Gonzalez 1999). In The predominance of Monesichnus, Uruguay and
some exposures, dismantling horizons surround Coprinisphaera in the nodular horizons, either in
remains of duricrusts. Nodules are subrounded their original positions or rotated, was pointed
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 367

out by Gonzalez et al (1998; Fig. 7c). The bio- 1988). Fossil leaves, fruits and woods indicate a
turbation is moderate (BI3). wet tropical to subtropical (probably monsoo-
The duricrusts are 0.5-2.0 m thick, 200-300 m nal) climate. Mangrove swamps prevailed in
long, undulatory, and wedge shaped (Ford coastline areas, whereas in the forested interior
1988b). Internal structure is defined by coarse a large and varied vertebrate fauna proliferated,
columnar to angular blocky peds (Gonzalez especially diverse mammals including Old
et al. 1998), with reddish to orange clay cutans, World primates (Bown 1982; Bown et al. 1982).
pedotubules and glaebules (Ford 1988a). Col- Apart from body fossils, ichnofossils of mam-
umns are often inclined at 10-30°. Laterally mals (rodents), invertebrates (insects, annelids,
and subordinately the structure becomes massive crabs and crayfishes) and plants (rhizoliths) are
or vacuolar, with empty channels (3—6 mm wide) also very common in the Jebel Qatrani Forma-
and macro voids (alveoles). Some horizons in the tion. Most traces are exceptionally preserved in
lower section of the unit include abundant iron pedogenized fluvial-channel sandstones, because
nodules or brecciated carbonate patches, and particular geochemical conditions prevailed
others show mottled and bleached (yellowish) during early diagenesis (Bown 1982). In some
levels (Ford 1988b). Micromorphologically localities these trace fossils formed intricate,
these horizons are characterized by a clayey- dense masses of chambers and tubes (Bown &
haematitic ground mass and thick laminated Kraus 1988).
ferroargillans coating voids and grains (Gonza- The selected example comes from the middle
lez 1999). Mineral composition is dominated by part of the Jebel Qatrani Formation, where a
weathered and fractured quartz and hematite yellowish brown, poorly developed Inceptisol
cement. Kaolinite is abundant and decreases occurs in a fine sandstone deposit, corresponding
upwards in the profile, whereas montmorillonite to the upper part of a point bar of a meandering
increases upwards (Ford 1988a). These horizons river (Genise & Bown 1994b; Fig. 5b). Relict
are dominated by Palmiraichnus and Teisseirei trough cross-bedding is the only remaining pri-
located in situ and rizoliths up to 30cm long mary physical sedimentary structure. Inceptisols
(Gonzalez et al. 1998; Fig. 7b). Bioturbation is are the most common palaeosol group in the
also moderate (BI 3). Jebel Qatrani Formation, especially in the
coarser channel deposits (Bown & Kraus 1988).
Their poor development reflects iterative deposi-
Palaeosolsfrom the Jebel Qatrani Formation tional-erosional conditions in the active meander
(Egypt) belt and minor pauses in the sedimentation
(Bown & Kraus 1988). No horizons or soil struc-
The Eocene-Oligocene Jebel Qatrani Formation ture were recognized. The ichnofabric is uniform
of northern Egypt is a sequence of fine-grained to along the palaeosol. Bioturbation intensity is
gravelly sandstones, mudstones, scarce carbo- high (BI4), with frequent trace fossil overlap.
naceous shales and freshwater limestones, most Rhizoliths and termite nests are particularly
of them showing evidence of damp pedogenesis abundant. Oblate chambers and narrow galleries
(Bown 1982; Bown & Kraus 1988). This unit compose the fossil termitaria (Fleaglellius pago-
records the sedimentation of high-sinuosity dus), which occur in the upper part of densely
rivers and overbank floods on a Palaeogene rooted palaeosols, where it is difficult to distin-
lowland coastal plain (Bown & Kraus 1988). guish between galleries and small rhizoliths
Fluvial sediment accumulation was controlled (Genise & Bown 1994b; Fig. 7d). The nest
by sea-level fluctuations in a stable tectonic system, comprising polychambered structures
setting. Variegation of the rocks in tabular forming high towers interconnected by galleries,
bands is the most striking feature of palaeosol occupies a large volume of the palaeosol.
formation.
Root and insect traces, cutans, clay-lined vugs
and red duricrusts are additional evidence of Palaeosols from the Sarmiento Formation
widespread pedogenesis. The rhizolith assem- (Argentina)
blage contains roots and stems of fossil bushy
and reedy plants and trees. Curiously, vegetal The Sarmiento Formation is an Eocene-lower
and animal bioturbation are not present in asso- Miocene pyroclastic succession known mostly
ciated argillic horizons (Bown & Kraus 1988). because of its abundant and varied fossil verte-
The recognized alluvial palaeosol types include brates. The mammal assemblages (marsupials,
wet Entisols, Inceptisols, mature gley Spodosols xenarthrans, astrapotherians, notoungulates,
and very mature Ultisols, some of them with primates and rodents) are considered the strati-
fragipan horizons (Bown 1982; Bown & Kraus graphic standards for the South America land
368 J. F. GENISE ET AL.

mammal ages (Ameghino 1906; Simpson 1940; from Kay et al. 1999; fig. 1). The palaeosol
Cifelli 1985). It is broadly exposed in central occurs in a fine (coarse silt grain size) massive
and north Patagonia (Argentina), covering tuff, 2.4m thick, highly porous, and light grey
more than 200 000 km2 and showing a relatively (N 8). Bed lithology and thickness are laterally
uniform lithology, characterized by chonites very uniform on a kilometre scale (mantle bed-
(mud and clay-size tuffs), fine tuffs, bentonites, ding). The bases of beds are sharp, horizontal
and intraformational conglomerates (Mazzoni erosional surfaces, overlain by a thin upward-
1979, 1985). Likewise, the presence of palaeosols fining intra-formational breccia. Two horizons
bearing trace fossils is another significant and may be recognized in this palaeosol (Figs 6,
well-known feature (Frenguelli 1938; Andreis 7f). The upper one is 0.40m thick, indurated,
1972; Andreis et al 1975; Spalleti & Mazzoni very light grey (5YR7/1), and shows intense bio-
1977, 1979). At the type locality, Gran Barranca, turbation and scattered specimens of Coprini-
south of Chubut province, where the unit is sphaera (BI5). The intense bioturbation is
divided in three members, there are numerous composed of a boxwork of sinuous intercon-
well-exposed palaeosols in the middle and nected burrows, 1-2 mm wide and 20mm long,
upper members. The dominant parent material having smooth walls and irregular chambers.
for palaeosols is fine volcanic ash composed of Micromorphologically this horizon is character-
rhyolitic-dacitic glass shards and subordinate ized by abundant glass shards and a vesicular
plagioclase (andesine). Opal phytoliths are also structure, with an undifferentiated b-fabric,
frequent or abundant in some levels, reaching abundant voids and channels, some of them
up to 7% of the sample (Mazzoni 1979). Pyro- coated by thin argilans, organic nodules and a
clastic deposits resulted from distal ash falls, few braided strands. Some features of this
sourced from plinian-type eruptions more than boxwork resemble modern termite nests. Copri-
500km to the NNW (Mazzoni 1985). Most of nisphaera balls are densely perforated inside
the Sarmiento Formation tuffaceous beds are this boxwork, but in the remaining palaeosol
interpreted as an aeolian loessite (Spalletti & they are undamaged. The lighter-coloured
Mazzoni 1979). lower horizon is 2.0m thick with a transitional
At Gran Barranca, the middle section (Puesto upper contact, showing coarse columnar struc-
Almendra Member) is composed of primary ture. Preservation of the original bedding is
vitric tuffs, bentonites, cross-bedded volcanic difficult to assess because of the massive charac-
sandstones, intraformational conglomerates ter (ash fall) of the primary sediment. Bioturba-
(generally palaeosol fragments) and a few lenti- tion is sparse (BI1), and composed of very thin,
cular basaltic flows (Spalletti & Mazzoni 1979; long root traces. Owing to a lack of clay (which
Fig. 6). A late Eocene-early Miocene age has precluded the formation of an argillic horizon),
been established for this member (Kay et al. the abundance and good preservation of vitric
1999). Pyroclastic deposits were partly recycled shards (volcanic ash) and the thick profile, this
by fluvial and aeolian processes, and supported palaeosol is classified as a weak to moderately
pedogenesis. Most of the tuffaceous palaeosols developed Andisol. This example is comparable
are bioturbated and possess a significant argillic to the Luquem palaeosol of Retallack et al.
horizon, Fe and Mn nodules, a moderate to (2000), a type of inmature Andisol, which pre-
well-developed b-fabric and significant argilans serves opportunistic vegetation of low specific
and ferromangans (Bellosi et al. 200Ib). Domi- diversity that grew on volcanic ash substrates.
nant trace fossils are dung-beetle nests (Coprini- The second case is included in a series of
sphaerd) (Spalletti & Mazzoni 1979; Laza 1986; stacked, reddish-orange indurated palaeosols
Escribano & Delgado 1996; Genise et al. 2000; (stake MZ-17 in Kay et al. 1999; fig. 1) associated
Bellosi et al. 200Ib), but there are also frequent to the upper erosive unconformity of this
root traces, other coleopteran traces (Teisseirei), member (Kay et al. 2001; Bellosi et al. 2002b;
bee cells (Celliforma), Beaconites cor onus, and Figs 6, 7e). Palaeosols differ in horizon thickness
large horizontal burrows (Bellosi et al. 200Ib). and boundary, textural features, ped type and
Scarce calcareous palaeosols (calcretes) contain abundance of iron nodules (pedofabric), and
almost exclusively Celliforma in association especially in the trace fossil content (ichno-
with charopid land gastropods (Bellosi et al. fabric). In the selected palaeosol the original
2002c). deposit preserved in the lower 80 cm, is massive,
Two palaeosols of Puesto Almendra Member greyish orange (10YR7/4), with an intraforma-
were selected to show tiering and pedofabric/ich- tional conglomerate. A rough stratification
nofabric relationships (Figs 6, 7e, f). The first describes the poor preservation of the original
example is in the middle section of the west end bedding. The surface horizon is 30-40 cm thick
profile of the Gran Barranca (stake MZ-10 and argillic. The pedofabric is composed of
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 369

Fig. 6. (a) Stratigraphic position of the selected palaeosols from the Puesto Almendra Member of Sarmiento
Formation. Fades association records a discontinuous sedimentation (erosional surfaces and abundant
palaeosols), with distal volcanic ash falls, local basalt flows and fluvial reworking by a low-sinuosity system,
(b) The lower example (asterisk) is an Andisol preserving the C horizon, (c) The upper example (star) is an
Alfisol located near the top of a series of stacked palaeosols related to an unconformity.

subangular blocky peds (subordinate crumb Micromorphology is characterized by abundant


peds), with ferruginized crusts at the top (Fig. fresh glass shards, a well-developed spongy
7e). Fed size varies from coarse blocky (2-5 cm) structure, laminated clay, and Fe-cutans, along
to medium crumb (2-5 mm) (Soil Survey Staff with Fe nodules. Some interconnected pores
1975), and the shape is slightly equidimensional. generate a blocky microstructure. The contact
Peds are commonly not interlocked, being between both horizons is undulatory and rela-
separated by clay materials. The crusts are dark tively sharp. This palaeosol is interpreted as a
orange (10 YR 5/6), 2-3 cm thick and folded. moderately developed Alfisol because of the
370 J. F. GENISE ET AL.

Fig. 7. (a) Laterized profile of Ultisols from the Asencio Formation. Massive to coarse columnar duricrust
(arrow) covered by a nodular dismantling horizon, (b) Duricrust showing blocky peds, rhizoliths and
Palmiraichnus castellanosi (arrow). Asencio Formation. Scale in cm. (c) Dismantling horizon showing
nodular structure and the fossil bee nest Uruguay auroranormae (arrow). Scale in cm. (d) Inceptisol of the
Jebel Qatrani Formation showing tiered chambers of the fossil termite nests (Fleaglellius pagodus) and
abundant vertical rhizoliths. Lens cap: 5 cm. (e) Alfisol showing subangular blocky peds and folded
ferruginized crusts showing a dung-beetle brood mass (Coprinisphaera) (arrow). Sarmiento Formation,
(f) Andisol showing an upper horizon with granular peds and intense bioturbation possibly produced by
termites and Coprinisphaera (arrows). The lower horizon show a coarse columnar structure and low
bioturbation. Sarmiento Formation.
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 371

coarse well-defined peds, thick argillans and terms of destruction of the original bedding by
undifferentiated b-fabric in the ground mass. the ped-forming processes.
The ichnofabric is arranged in two tiers (Fig. 6). A still clearer case of ichnofabric destruction
In the upper one, Celliforma is the only discrete by other soil processes occurs in the Ultisols of
trace, resulting in a low bioturbation (BI2). the Asencio Formation. These palaeosols show
The lower tier includes Coprinisphaera, Teisseirei well-developed pedofabrics and ichnofabrics,
barattinia, pan-shaped traces, and large hori- both responsible for the obliteration of the
zontal burrows 1.8cm in diameter (Fig. 7e). original bedding, which is absent in almost all
Bioturbation is moderate (BIS), discrete traces outcrops. In addition, the lateritization process
are frequent, and cross-cutting is rare. dismantled the duricrusts, partially destroying
the original ichnofabric. The only preserved
trace fossils in these dismantling horizons are
Discussion insect nests with thick constructed walls. Many
nests are disturbed relative to their original
Ichnofabrics and pedofabrics positions, as evidenced by random orientation
of their nest entrances, as most bees and dung-
The different cases described herein show how beetles are very constant in the orientation of
pedofabrics and ichnofabrics may independently their cells and brood masses (e.g. Halffter &
exhibit variable degrees of development, how Matthews 1966; Stephen et al. 1969). The ferri-
bioturbation and other soil processes may dis- cretization process also modified the original
rupt the original bedding by themselves or in pedofabric of Ultisols in which insects nested
combination, and how soil processes (e.g. homo- and bioturbation developed. At the other
genization, lateritization) may also disrupt extreme, the study case from the Jebel Qatrani
ichnofabrics. In palaeosols of the Ischigualasto Formation shows Inceptisols almost deprived
Formation ichnofabric is almost absent - just a of pedofabric, where intense bioturbation results
few stems of equisetales and Skolithos linearis in a complex intense ichnofabric of BI4.
in the less-developed palaeosols. Even so, the Entisols from the Laguna Palacios Formation
degree of ped development in these Vertisols is and palaeosols from the Sarmiento Formation
such that the original bedding is visible only in show intermediate cases in which pedofabric is
one of the study cases. The absence of a well- poorly or undeveloped, whereas ichnofabric
developed ichnofabric in those palaeosols may shows different degrees of development. Alfisols
be attributed to different causes: of the Laguna Palacios Formation show well-
developed ichnofabrics and pedofabrics, whose
• the absence of constructed, more preservable,
relative degree of development varies laterally
structures, such as those made by bees, dung-
along the exposed palaeosol on a 100m scale.
beetles and termites before the Cretaceous
Most of the Laguna Palacios and Sarmiento
(Genise & Bown 1994a); palaeosols differ from the remaining alluvial
• the frequent waterlogging and reducing condi-
examples in that they developed in pyroclastic
tions found in these soils (Retallack 1990);
aeolian systems. The contrasting mechanisms of
• ichnofabric destruction by intense pedogenic
sediment supply and hiatuses controlled the
homogeneization in Vertisols (Melchor et al.
soil-forming processes and soil stratigraphy in a
2001).
variety of different ways.
The last hypothesis suggests not only that non- The independent evaluation of ichnofabrics
biological soil processes may disrupt the original and pedofabrics in these cases favours a more
bedding by itself, but also that they may destroy complete analysis of the evolution of these
the ichnofabric of the deposits, thus emphasizing deposits. Different characters of the pedofabric
the importance of studying both pedofabric can be used to estimate the relative duration of
and ichnofabric in palaeosols. According to formation of soils (Retallack 1994). For instance,
independent sedimentological data, these alluvial the clayey Bt horizon of the Alfisol from Laguna
Vertisols are immature, as maturity was defined Palacios Formation suggests 104 to 103 years of
as a function of time by Bown & Kraus (1987). formation because of the well-developed clay
In addition, the scales of palaeosol development skins, ground mass, and homogeneous sepic
are based mostly on particular types of horizon microfabric (Retallack 1994). During this
(Bown & Kraus 1987; Retallack 1988) that are period of subaerial exposure water tables fluctu-
not present in palaeovertisols (Nettleton et al. ated seasonally, as revealed by hydromorphic
2000). Consequently, and in the absence of features and illuviation of clays (Genise et al.
important bioturbation, the stage of develop- 2002). The evidence of subaerial exposure pro-
ment of these palaeosols is evaluated herein in vided by coleopteran pupation chambers in
372 J. F. GENISE ET AL.

these palaeosols falls in a very different timescale. (e.g. Michener 1979; Roubik & Michener 1980;
Insect activity involved in the construction of Cane 1991; Visscher et al. 1994). The water-
Rebuffoichnus probably indicates the subaerial proofed breeding cells of an aggregation of Epi-
exposure of the deposits during only one or two charis zonata (Anthophorinae) remain beneath
seasons. Pedofabrics of the Ischigualasto Forma- the water table during the wet season (Roubik
tion appear to have been developed in less time & Michener 1980). An aggregation of Calliopsis
than that of Laguna Palacios because of the pugionis (Andreninae) that nested in a site that
rapid sedimentation rate in the first formation, was flooded after a heavy, unusual winter rain
as discussed previously. was recorded by Visscher et al. (1994). Flooding
The nesting activity of termites that produced was not fatal, although bees emerged later in the
the ichnofabric dominated by Fleaglellius pago- season, affecting the pollen collection and repro-
dus in the Jebel Qatrani Formation suggests ductive success. Accordingly, most bee nests are
several years of colony growth by apposition of located in well-drained soils (Linsley 1958;
chambers and simultaneous foundation of new Batra 1984), and the cell wall is lined with
colonies (Genise & Bown 1994b). This fact, water-repellent lipids to maintain the moisture
along with the abundance of rhizoliths, indicates conditions (Cane 1991). The ichnogenera Copri-
a longer period of subaerial exposure and lower nisphaera, Fontanai and Monesichnus, present in
water tables than that suggested by the nesting the Asencio and Sarmiento ichnofabrics, are
of solitary insects in the Laguna Palacios, considered to be brood-masses of dung-beetles
Asencio and Sarmiento Formations. Nests of (Sauer 1955; Roselli 1987; Genise 1993; Genise
solitary insects may be constructed, provisioned & Laza 1998). Poorly drained and occasionally
and closed in only one day, whereas those of flooded soils are unfavourable for many digging
social insects may be maintained for years, and species of dung-beetle, whereas the period and
their micromorphology can even reveal their duration of flooding determine the success or
age (e.g. Jonkman 1980). Regardless of this, failure of reproduction (Lumaret 1983). One spe-
high concentrations of nests of solitary insects, cies nesting in soils that were flooded during
along with well-developed pedofabrics, such as winter showed a high mortality of eggs and
from the Asencio Formation, may indicate a larvae (Kirk 1983). In addition, Hanski and
very long period of subaerial exposure and low Cambefort (1991) stated that waterlogged soils
water tables, interrupted only by heavy seasonal are generally poor for all dung-beetles owing to
rains, attested to by the presence of abundant alteration of the droppings that these insects
dismantled horizons. The Egyptian example utilize to provision their nests. In conclusion, in
demonstrates the independence of ichnofabric nests of bees and dung-beetles, excessive water
and pedofabric development, as the intense sub- content in the soil produces problems related to
aerial bioturbation produced by termites and larval mortality, decay of provisions, oxygen dif-
roots is not accompanied by the development fusion and alteration of droppings. Thus the ich-
of other soil characters. notaxa mentioned previously may be regarded as
indicators of well drained to sporadically flooded
palaeoenvironments and low water tables at the
Bee and coleopteran trace fossils time of their emplacement in the soils.
Apart from calichnia, other recognizable
The ichnofabrics of Laguna Palacios, Asencio coleopteran structures, fossil pupation cells are
and Sarmiento show fossil bee cells and/or common in palaeosols (Roselli 1938, 1987; Retal-
dung-beetle brood masses as their dominant lack 1984; Johnston et al. 1996; Genise 1999;
components. Remains of nests or cells attribu- Genise et al. 2002). They are represented in the
table to bees are included in the ichnofamily present study by the ichnogenera Teisseirei in
Celliformidae, comprising the ichnogenera the Asencio and Sarmiento ichnofabrics and by
Palmiraichnus, Celliforma, Corimbatichnus, Rebuffoichnus in the Laguna Palacios locality.
Uruguay, Ellipsoideichnus, Rosellichnus and Such structures are chambers constructed by
Cellicalichnus (Genise 2000). In soil bee nests, larvae of different groups of coleopterans to
excessive moisture causes liquefaction or decay contain and protect pupae before emergence as
of provisions by fungal attack, which may also adult (Retallack 1984; Johnston et al. 1996). It
destroy the larvae (Michener 1979; Rozen in. is difficult to attribute these chambers to par-
litt. in Michener 1979). In addition, in water- ticular coleopteran taxa with known ecological
logged soils, larvae are exposed to poor oxygen preferences (Genise et al. 2002), but at least as
diffusion (Visscher et al 1994). There are a the trace-makers are air breathers it is possible
few exceptional records of bees nesting in to ascertain that they were constructed under
periodically or sporadically submerged sites subaerial conditions above the water table.
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 373

Ant and termite trace fossils by fossil termite nests such as those of the Jebel
Qatrani Formation would indicate higher moist-
The ichnogenus Fleaglellius, which is dominant ure conditions and more frequently flooded soils
in the ichnofabric of the study case from the than ichnofabrics dominated by bee and dung-
Jebel Qatrani Formation, is considered to be a beetle traces. Fossil plants and mammals of the
termite nest (Genise & Bown 1994b). Another Jebel Qatrani Formation also indicate a wet
possible termite nest presented herein is that climate (Bown 1982; Bown et al 1982; Bown &
recorded from the second case study of the Kraus 1988).
Sarmiento Formation. Fossil termite nests are In conclusion, ichnofabrics dominated by
one of the most common trace fossils in palaeo- calichnia and pupation chambers indicate sub-
sols (Bown & Laza 1990; Genise & Bown aerial conditions. The construction of nests and
1994b; Genise 1997), sometimes comprising other underground activities take place during
complex termitic ichnofabrics (Genise & Bown periods of subaerial exposure of deposits. It is
1994b). Termites as a whole prefer high atmos- important to note that insect nests can be trans-
pheric and soil moisture (e.g. Collins 1969; ported, and that they have been recorded as
Grasse 1986), and consequently their nests may clasts in conglomerates (e.g. Andreis 1981;
be found in periodically waterlogged soils (e.g. Bown & Ratcliffe 1988). Therefore not only the
Lee & Wood 1971; Grasse 1984). However, as identification of nests, but also their position in
with bee or dung-beetle nests, waterlogging situ, are needed before the subaerial exposure
produces similar problems of gas exchange (e.g. of deposits can be determined.
Schmidt 1960; Roy-Noel 1972; San Jose et al
1989) or fungal or microbial infections (e.g.
Grasse 1984). Thus species that inhabit soils Earthworm trace fossils
that are seasonally waterlogged show particular
behaviours or nest features, such as those that Fossil earthworm burrows indicate a moist soil
retreat into epigeous mounds during the wet environment. Edaphichnium lumbricatum Bown
season (Lee & Wood 1971; Matthews 1977). and Kraus 1983 is considered to be an earth-
Other species provide their nests with special worm burrow because of its morphology, the
chimneys (Roy-Noel 1972), perforation systems, presence of faecal pellets, and the concentration
and air or sand envelopes (Schmidt 1960) to of calcium carbonate in the burrows and pellets.
maintain the microenvironmental conditions One of the most important requirements of
inside nests, which are very specific in terms of earthworms is adequate soil moisture, because
moisture and concentrations of C>2 and CC>2 respiration depends on the diffusion of gases
(Grasse 1984). A similar case is that of fossil through the moistened body wall (Lee 1985).
ant nests, also common in ichnofabrics of Consequently, earthworms inhabit soils in
Tertiary palaeosols in North and South America which water is confined to films on the surface
(Laza 1982; Bown et al 1997). In contrast to of soil aggregates or is held in pore spaces by
termites, which are restricted mostly to the capillary forces, and the relative humidity of
stable microenvironment of their nests, most air-filled spaces is 100% or slightly less (Lee
ants construct less elaborate structures but have 1985). In waterlogged soils or those where free
the ability to move their eggs and larvae from water is no longer present, earthworms cannot
place to place in response to environmental survive (Lee 1985). In seasonal tropical climates
changes (e.g. Wheeler 1910; Holldobler & earthworms construct spherical aestivation
Wilson 1990). Colonies move frequently - flood- chambers to spend the dry season at deeper
ing is one of the factors that can trigger these layers in the soil profile (Jimenez et al. 2000).
movements - although there are records of ants This type of trace, of great environmental
surviving several hours or even days submerged value, has recently been found in palaeosols
by floodwater (Holldobler & Wilson 1990). (Verde et al. 2002). In accordance with the
There are few records of ants that nest in low moist environments preferred by earthworms,
lands and also construct mounds to which they E. lumbricatum was found in gleyed palaeosols
retreat during the wet season to avoid high of the Willwood Formation (Bown & Kraus
water tables (e.g. Bruch 1916; Bonetto et al. 1983). It should be noted that a complete ichno-
1961), like termites. In summary, ants and - taxonomical review of pellet-filled burrows from
particularly - termites nest in well-drained to different environments (e.g. Richter & Richter
seasonally flooded soils. In the latter case nests 1939; Pickerill 1989 and references therein) is
are provided with particular devices, such as still lacking, and would be critical to correctly
epigeous mounds, chimneys, or special walls. In distinguish earthworm from other similar
terms of soil moisture, ichnofabrics dominated burrows.
374 J. F. GENISE ET AL.

Trace fossils of uncertain affinities Ichnotaxa such as Skolithos, Scoyenia, Taeni-


dium, Beaconites, Macanopsis, Cylindricum and
In contrast, another group of palaeosol ichno- other named and unnamed trace fossils have
fossils is that composed of those ichnotaxa that been cited from palaeosols (e.g. Genise et al.
(1) cannot be attributed unequivocally to a parti- 2000; Hasiotis 2000; Retallack 200Ib) as well as
cular group of producers, and (2) are recorded from marine and lacustrine environments (e.g.
from different continental and marine deposits. Alpert 1974; Hantzschel 1975; Bown & Kraus
As Retallack (1990) pointed out, these tubular 1983; Frey et al. 1984; Keighley & Pickerill
trace fossils, which are common in modern 1994; Buatois & Mangano 1995). With the excep-
soils, are not diagnostic of them because they tion of Scoyenia beerboweri (Retallack 200Ib), it
are also found in lacustrine and marine environ- is impossible to distinguish between specimens in
ments (e.g. Ratcliffe & Fagerstrom 1980). Ichno- subaerial environments from those occurring in
fabrics of Ischigualasto, Laguna Palacios, and subaqueous ones. The attribution of most of
Sarmiento show this type of trace fossil. In these trace fossils to particular groups of insects,
contrast, they are absent from those showing or even invertebrates, is thus, in such cases, at
the most developed subaerial assemblages, best speculative. Furthermore, to extract
Asencio and Jebel Qatrani. It is possible that, palaeoecological and other inferences from
in the future, micromorphological studies may them (e.g. Hasiotis & Dubiel 1994; Hasiotis
aid in the attribution of these trace fossils to 2000) without a previous accurate analysis of
modern taxa, but currently they are of uncertain their affinities is of dubious utility. The Triassic
affinities. The identification of ichnotaxa belong- meniscate burrows attributed to soil bugs by
ing to air breathers is critical for assessing the Hasiotis & Dubiel (1994) are similar to those
subaerial exposure of the deposits, particularly recorded from subaqueous environments
for those showing poorly developed palaeosols. (Keighley & Pickerill 1994). In addition, in the
Palaeosols and/or subaerial trace fossils may original contribution by Willis & Roth (1962)
occur in fluvial (e.g. Bown & Kraus 1983), cited as source by Hasiotis & Dubiel (1994),
lacustrine (e.g. Edwards et al. 1998) and marine it is stated that no tunnels or channels are pre-
(e.g. Curran 1994) deposits during periods of sent, a fact already mentioned by RatclifTe and
subaerial exposure. In theory, if palaeosol devel- Fagerstrom (1980). Soil bugs have their mouth-
opment is very weak, a trace fossil suite related to parts adapted for sucking fluids from roots or
the original subaqueous conditions of these other plant materials (e.g. Carver et al. 1991).
deposits could be preserved along with trace Observations of the feeding habits of these
fossils produced under subaerial conditions insects are particularly important in the inter-
(e.g. Curran 1994; Buatois et al. 1998; Retallack pretation of Triassic meniscate burrows. Their
200 Ib). It has been stressed that the presence menisci are ungraded, and stained with alter-
of root traces is a post-depositional process, nating zones of oxidized and unoxidized iron
independent from the origin of deposits, that compounds (Hasiotis & Dubiel 1994). This
indicates subaerial exposure and change of pattern is commonly interpreted as the result of
environment (Bockelie 1994; Curran 1994). the original alternation of organic-rich (faecal)
Similarly, a subaqueous assemblage could be material with organic-poor (sediment) material
developed if a subaerially exposed deposit were (D'Alessandro & Bromley 1987; Hasiotis et al.
later to be submerged (e.g. Driese & Foreman 1993; Keighley & Pickerill 1994). The presence
1991; Curran 1994), and subaerial and sub- of such alternating menisci containing faecal
aqueous trace fossils may occur together in material in burrows made by insects, which do
deposits subsequently submerged and exposed not ingest sediment, is unlikely (e.g. Frey et al.
to air (Frey et al. 1984 and references therein). 1984), and even more in soil bugs that feed on
Recently, Retallack (200Ib) analysed the possi- root fluids. This type of meniscate burrow in
bility of recording fossil burrows representing Willwood palaeosols was attributed to earth-
aquatic environments predating soil formation worms by Bown & Kraus (1983).
or resulting from later inundations by lake or The only possibility of gathering some
lagoonal waters. Different criteria, such as environmental data from this group of trace fos-
density of burrows in relation to other soil sils of uncertain affinities is when its presence is
characters, cross-cutting relationships with analysed in combination with other palaeosol
carbonate nodules, burrow collapse and fillings and sedimentological data and compared with
and associated ichnofauna, were used to demon- those from other sequences (Bown & Kraus
strate that Scoyenia beerboweri was produced 1983; Retallack 1985, 2001b). The 'adhesive
during the period of soil formation (Retallack meniscate burrows' from palaeosols of the
1985, 2001b). Willwood Formation described by Bown &
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 375

Kraus (1983) were later interpreted as indicators mental conditions. The few possible clues are
of increased soil moisture and fluctuating water that:
table, owing to their presence in the lower half
they are mostly recorded from subaqueous
of the palaeosol profile (Hasiotis et al. 1993). In
environments;
addition, Hasiotis et al. (1993) recorded the pre- in some cases their lateral distribution is
sence of Scaphichnium hamatum, a dung-beetle
homogeneous, suggesting that it is not con-
nest, in the upper half of the same profile in a
trolled by soil environmental factors; and
better-drained and drier environment. Similarly,
several studies suggest that some of them occur
in the Gleysols from the Triassic Chinle Forma-
in moister conditions than insect nests.
tion the adhesive meniscate burrows are abundant
in the deepest tier just above the water table, Considering that soils may be sporadically,
characterized as having high, stable moisture con- seasonally or permanently waterlogged (e.g.
tent (Hasiotis & Dubiel 1994). In these two cases Retallack 1990), a possibility is that these ichno-
Taenidium is recorded from the deepest tiers of genera, in contrast to those attributed to
the vadose zone of gleyed palaeosols. The possible insect nests and earthworms, may be recording
producers of meniscate burrows were analysed in periods of high moisture or even waterlogging
detail by Frey et al. (1984), who pointed out the of soils.
difficulties in establishing their affinities and
palaeoenvironmental significance. In conclusion,
these authors stated that some meniscate burrows Composite ichnofabrics in the study cases
(e.g. Scoyenia gracilis and Beaconites coronus)
occur preferentially in moist to wet substrates in In the Ischigualasto Formation ichnofabrics,
shallow aquatic deposits periodically exposed to Skolithos are found at the top of poorly devel-
air or in subaerial deposits periodically submerged oped palaeosols in association with remains of
(Frey et al. 1984). These scarce data suggest a the original bedding (e.g. ripple cross-lamina-
clear distinction between the drier and well- tion), suggesting that these trace fossils were
drained conditions preferred by insect to nest or probably produced by subaquatic organisms. In
pupate and the moister and less-drained condi- contrast, in Entisols of the Laguna Palacios
tions selected by the producers of these meniscate Formation, the upper tier of Taenidium can be
trace fossils. In contrast, Scoyenia beerboweri demonstrated to obliterate the entrance of Celli-
was described from well-drained palaeosols in a calichnus, suggesting that bees opportunistically
tropical, seasonally dry, semi-arid palaeoclimate colonized the deepest layers of the soil first, and
(Retallack 200 Ib). that Taenidium is a later component of this
Apart from vertical tiering, soil organisms ichnofabric. This composite ichnofabric would
show patchy lateral distributions controlled by thus reflect a shift of palaeoenvironmental
soil texture, soil carbon content, vegetation, conditions from drier conditions (early deepest-
and population dynamics (Ettema & Wardle tier Cellicalichnus ichnocoenosis) to moister
2002). Accordingly, insect fossil nests and conditions (later shallow-tier Taenidium ichno-
pupation chambers commonly show a laterally coenosis), probably because of a raised water
heterogeneous distribution in palaeosols, as in table. Curiously, Savrda et al. (2000) found a
the case of Rebuffoichnus described herein from similar, but extreme, form of composite ichno-
one example of the Laguna Palacios Formation. fabric in the Cretaceous Tuscaloosa Formation,
In contrast, meniscate trace fossils in the same having recent Cellicalichnus-like nests cross-
ichnofabric, such as Taenidium and Beaconites, cutting Taenidium-dommatQd fabrics.
have an extended and homogeneous lateral dis- In Alfisols of the Laguna Palacios Formation,
tribution, suggesting that these trace fossils ichnofabrics are dominated by Taenidium,
were not controlled by the lateral heterogeneity Beaconites and Skolithos, which show a homoge-
of soils. Horizontal patterns of variability of neous horizontal and vertical distribution in the
ichnofabrics have been less documented than palaeosol, whereas, Rebuffoichnus - a clear
vertical ones, but more homogeneous lateral indicator of subaerial conditions - is laterally
distributions are known from subaqueous trace restricted in the same palaeosol, in which they
fossils, so much that concentration of burrows cross-cut the previously mentioned group of
(e.g. Skolithos, Ophiomorpha, Zoophycos) along traces. This composite ichnofabric thus indicates
bedding planes has been used as marker beds in changes in the soil conditions from moist or
stratigraphy (e.g. Ekdale et al. 1984). even waterlogged palaeonvironments (Taenidium,
In conclusion, this group of ichnogenera of Beaconites, Skolithos) to subaerial exposure
simple morphology is by now not clearly (Rebuffoichnus). Accordingly, illuviation of clays
indicative of a particular set of palaeoenviron- and hygromorphic features of this palaeosol
376 J. F. GENISE ET AL.

indicate seasonality and a mobile water table based on two different approaches. One
(Genise et al 2002). approach comprises the detailed record and
On the other hand, tiers composed of different comparison of the occurrence of trace fossils in
subaerial ichnotaxa (1) sharing common environ- particular palaeosol types and horizons, which
mental preferences, such as bee cells and dung- is particularly important when dealing with the
beetle brood masses, and (2) showing little, at more simple trace fossils, whose producers are
random, or no cross-cutting are interpreted unknown or uncertain. A second approach, par-
herein as the original tiering of soils. In the case ticularly important when dealing with complex
study of the Asencio Formation, ichnofabric trace fossils such as insect nests, is based on the
reveals a tiered structure, albeit somewhat blurred identification of trace-makers and the evaluation
by the laterization processes. In the dismantling of ecological preferences and requirements of
horizon the upper tier is composed mostly of their recent representatives.
Uruguay, Monesichnus and Coprinisphaera, Different groups of trace fossils in the study
whereas in the crusts Teisseirei and Palmiraichnus cases show a spectrum from the drier conditions
prevail. Similarly, in the Sarmiento Formation preferred by bees and dung-beetles to the moister
Celliforma constitutes the upper tier, whereas in ones of ants, termites and trace fossils of uncertain
the lower one Coprinisphaera, Teisseirei and two affinities. The subaerial environment is clearly
other undetermined trace fossils are dominant. indicated only by trace fossils certainly attributa-
ble to air breathers (insect nests and pupation
cells, earthworm and millipede burrows). In con-
Conclusions trast, trace fossils of uncertain affinities may be
subaerial or subaqueous in origin. A possibility
Many soil features that disrupt the original bed- is that they may be recording periods of high
ding of the deposits may be formed without the moisture, waterlogging of soils, or even a sub-
intervention of bioturbation, or may be the aqueous environment before or after the soil-
result of its interactions with physical and chemi- forming process.
cal processes. For ichnofabric analysis these soil The duration of subaerial exposure will be
features are considered to constitute the pedofab- directly related to ichnofabric development by
ric of the deposit, which is distinguished from the the subaerial suite and, as such, it will be a key
ichnofabric (the fabric directly and completely factor, along with the development of the
produced by plant and animal traces). The use- pedofabric, in understanding the dynamics of
fulness of this distinction is supported by the post-depositional soil processes.
analysis of study cases that show that ichno-
fabrics can be intense in palaeosols devoid of The authors thank D. Mcllroy, G. Retallack and J. de
Gibert for the critical review of the manuscript. This
other soil characters and, conversely, palaeosols research was partially supported by grants from the
showing a well-developed pedofabric can bear National Scientific Research Council of Argentina
almost no trace fossils. In addition, bioturbation (CONICET-PIP 717/98), the National Agency of
and other soil processes may disrupt the original Scientific and Technical Promotion of Argentina
bedding by themselves or in combination, and (FONCYT-PICT 6156/99), and the National Science
soil processes (e.g. homogenization, lateritiza- Foundation of the USA (EAR 00-87636).
tion) may also disrupt ichnofabrics. The pedo-
fabric is an additional component to classical
ichnofabric analysis, which normally considers References
only original bedding and ichnofabrics. This ALPERT, S. P. 1974. Systematic review of the genus
complexity of palaeosol fabrics requires some Skolithos. Journal of Paleontology, 48, 661-669.
modifications to previous methodologies to AMEGHINO, F. 1906. Les formations sedimentaires du
better describe and interpret palaeosol ichnofab- Cretace superieur et du Tertiaire de Patagonie
rics. The analysis comprises: avec un parallele entre leurs faunes mammalogi-
ques et celles de 1'ancien continent. Anales del
• tiering diagrams, in which the bioturbation Museo Nacional de Historia Natural de Buenos
index of different tiers is indicated; Aires, 15, 1-568.
• the pedofabric, depicted along with and ANDREIS, R. 1972. Paleosuelos de la Formacion Mus-
independently from the ichnofabric; and ters (Eoceno Medio), Laguna del Mate, Provincia
• a pedofabric/ichnofabric ternary diagram de Chubut, R. Argentina. Revista de la Sociedad
showing percentages of bioturbation, pedo- Argentina de Mineralogia, Petrografia, Sedimento-
fabric and original bedding. logia, 3, 91-97.
ANDREIS, R. 1981. Identification e importancia geolo-
The record of palaeoenvironmental changes gica de los paleosuelos. Universidade Federal do
in composite ichnofabrics of palaeosols may be Rio Grande do Sul, Porto Alegre, Livro-texto 2.
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 377

ANDREIS, R., MAZZONI, M. M. & SPALLETTI, L. A. BOTTJER, D. & DROSER, M. L. 1991. Ichnofabric and
1975. Estudio estratigrafico y paleoambiental de basin analysis. Palaios, 6, 199-205.
las sedimentitas terciarias entre Pico Salamanca y BOTTJER, D. & DROSER, M. L. 1994. The history of
Bahia Bustamante, provincia de Chubut, Repub- Phanerozoic bioturbation. In: DONOVAN, S. K.
lica Argentina. Revista de la Asociacion Geologica (ed.) The Paleobiology of Trace Fossils. Wiley,
Argentina, 30, 85-103. New York, 155-176.
ARTABE, A. E., MOREL, E. M. & SPALLETTI, L. A. 2001. BOWN, T. M. 1982. Ichnofossils and rizoliths of the
Paleoecologia de las floras triasicas argentinas. In: nearshore fluvial Jebel Qatrani Formation (Oligo-
ARTABE, A. E., MOREL, E. M. & ZAMUNER, A. cene), Fayum Province, Egypt. Palaeogeography,
(eds) El Sistema Tridsico en la Argentina. Funda- Palaeoclimatology, Palaeoecology, 40, 255-309.
cion Museo de La Plata 'Francisco P. Moreno', BOWN, T. M. & KRAUS, M. J. 1983. Ichnofossils of the
La Plata, 199-225. alluvial Willwood Formation (Lower Eocene),
AUSICH, W. I. & BOTTJER, D. J. 1982. Tiering in suspen- Bighorn Basin, Northwest Wyoming, USA.
sion-feeding communities on soft substrata Palaeogeography, Palaeoclimatology, Palaeoecol-
throughout the Phanerozoic. Science, 216,173-174. ogy, 43, 95-128.
BATRA, S. T. W. 1984. Solitary bees. Scientific Ameri- BOWN, T. M. & KRAUS, M. J. 1987. Integration of
can, 250, 120-127. channel and floodplain suites in aggrading fluvial
BELLOSI, E. S. & SCIUTTO, J. C. 2002. Laguna Palacios systems. 1. Developmental sequence and lateral
Formation (San Jorge Basin, Argentina): an relations of lower Eocene alluvial palaeosols,
Upper Cretaceous loess-palaeosol sequence from Willwood Formation, Bighorn Basin, Wyoming.
Central Patagonia. Resumenes de la IX Reunion Journal of Sedimentary Petrology, 57, 587-601.
Argentina de Sedimentologia, p. 15. BOWN, T. M. & KRAUS, M. J. 1988. Geology and
BELLOSI, E. S., JALFIN, G. A., Bossi, G. E., BOGGETTI, paleoenvironment of the Oligocene Jebel Qatrani
D., CHEBLI, P. & MURUAGA, C. 2001a. Facies y Formation and adjacent rocks, Fayum Depres-
sedimentacion. In: ARTABE, A. E., MOREL, E. M. sion, Egypt. US Geological Survey Professional
& ZAMUNER, A. (eds) El Sistema Tridsico en la Papers, 1452.
Argentina. Fundacion Museo de La Plata 'Fran- BOWN, T. M. & LAZA, J. H. 1990. A miocene fossil
cisco P. Moreno', La Plata, 103-129. termite nest from southern Argentina and its
BELLOSI, E. S., LAZA, J. & GONZALEZ, M. 2001b. Icno- paleoclimatological implications. Ichnos, 1, 73-79.
faunas en paleosuelos de la Formacion Sarmiento BOWN, T. M. & RATCLIFFE, B. C. 1988. The origin of
(Eoceno-Mioceno), Patagonia central. Resumenes Chubutolithes Ihering, ichnofossils from the
de la IV Reunion Argentina de Icnologia y II Eocene and Oligocene of Chubut Province, Argen-
Reunion de Icnologia del Mercosur, Tucuman, p. 31. tina. Journal of Paleontology, 62, 163-167.
BELLOSI, E. S., GONZALEZ, M. & GENISE, J. F. 2002a. BOWN, T. M., KRAUS, M. J., WING, S. L., FLEAGLE,
Paleosuelos y sedimentacion cretacica de la J. G., TIFFNEY, B. H., SIMONS, E. L. & VONDRA,
Cuenca San Jorge (Grupo Chubut) en la sierra C. F. 1982. The Fayum primate forest revisited.
de San Bernardo, Patagonia central. Adas del The Journal of Human Evolution, 11, 603-632.
15° Congreso Geologico Argentino, El Calafate, BOWN, T. M., HASIOTIS, S. T., GENISE, J. F., MALDO-
CD-ROM. NADO, F. & BROUWERS, E. M. 1997. Trace fossils
BELLOSI, E. S., GONZALEZ, M., KAY, R. H. & MADDEN, of Hymenoptera and other insects and paleonvir-
R. F. 2002b. El valle inciso colhuehuapense. onments of the Claron Formation (Paleocene
Resumenes de la IX Reunion Argentina de Sedimen- and Eocene), Southwestern Utah. Bulletin of the
tologia, Corboba, p. 49. US Geological Survey, 2153, 42-58.
BELLOSI, E. S., MIQUEL, S. E., KAY, R. H. & MADDEN, BREWER, R. 1976. Fabric and Mineral Analysis of Soils.
R. F. 2002c. Un paleosuelo mustersense con Wiley, New York.
microgastropodos terrestres (Charopidae) de la BROMLEY, R. G. 1990. Trace Fossils. Unwin Hyman,
Formacion Sarmiento, Eoceno de Patagonia cen- London.
tral: significado paleoclimatico. Ameghiniana, 39, BROMLEY, R. G. 1994. The palaeoecology of bioero-
465^77. sion. In: DONOVAN, S. K. (ed.) The Palaeobiology
BLOKHUIS, W. A., KOOISTRA, M. J. & WILDING, L. P. of Trace Fossils. Wiley, New York, 134—154.
1990. Micromorphology of cracking clayey soils BROMLEY, R. G. & EKDALE, A. A. 1986. Composite
(Vertisols). In: DOUGLAS, L. A. (ed.) Soil Micro- ichnofabrics and tiering of burrows. Geological
morphology: A Basic and Applied Science. Devel- Magazine, 123, 59-65.
opments in Soil Science, Elsevier, Amsterdam, BRUCH, C. 1916. Contribution al estudio de las hormi-
19, 123-148. gas de la provincia de San Luis. Revista del Museo
BOCKELIE, J. F. 1994. Plant roots in core. In: DONOVAN, de La Plata, 23, 291-357.
S. K. (ed.) The Paleobiology of Trace Fossils. BUATOIS, L. A. & MANGANO, M. G. 1995. The paleon-
Wiley, New York, 175-199. vironmental and paleoecological significance of
BONETTO, A. A., MANZI, R. & PIGNALBERI, C. 1961. the lacustrine Mermia ichnofacies: an archetypical
Los 'tacurues' de Camponotus punctulatus subaqueous nonmarine trace fossil assemblage.
(Mayr). Notas ecologicas. Physis, 22, 217-224. Ichnos, 4, 151-161.
Bossi, J. 1966. Geologia del Uruguay. Universidad de la BUATOIS, L. A., JALFIN, G. & ACENOLAZA, F. G. 1997.
Republica, Departamento de Publicaciones, Permian nonmarine invertebrate trace fossils
Coleccion Ciencias, 2. from southern Patagonia, Argentina: ichnologic
378 J. F. GENISE ET AL.

signatures of substrate consolidation and coloni- EKDALE, A. A., BROMLEY, R. G. & PEMBERTON, S. G.
zation sequences. Journal of Paleontology, 71, 1984. Ichnology: the use of trace fossils in sedimen-
324-336. tology and stratigraphy. Society of Economic
BUATOIS, L. A., MANGANO, M. G., MAPLES, C. G. & Paleontologists and Mineralogists, Tulsa, Okla-
LANIER, W. P. 1998. Ichnology of an Upper homa, p. 316.
Carboniferous fluvio-estuarine paleovalley: the EKDALE, A. A. & BROMLEY, R. G. 1991. Analysis of
Tonganoxie Sandstone, Buildex Quarry, Eastern composite ichnofabrics: an example in uppermost
Kansas, USA. Journal of Paleontology, 72, 152- Cretaceous chalk of Denmark. Palaios, 6, 232-
180. 249.
BULLOCK, P., FEDOROFF, N., JONGERIUS, A., TURSINA, ENGEL, M. S. 2001. A monograph of the Baltic bees and
T. & BABEL, U. 1985. Handbook of Soil Thin evolution of the Apoidea (Hymenoptera). Bulletin
Section Description. Waine Research Publications, of the American Museum of Natural History, 259,
Albrighton. 1-192.
BUOL, S. W., HOLE, F. D. & MCCRACKEN, R. J. 1990. ESCHENBRENNER, V. 1986. Contribution des termites a
Genesis y clasificacion de suelos. Editorial Trillas, la microagregation des sols tropicaux. Pedologie,
Mexico. 22, 397-^08.
CANE, J. H. 1991. Soils of ground-nesting bees (Hyme- ESCRIBANO, V. & DELGADO, A. G. 1996. Aportes al con-
noptera: Apoidea): texture, moisture, cell depth ocimiento de nidos fosiles de Sacrabaeidae
and climate. Journal of the Kansas Entomological (Coleoptera) del Terciario (Eoceno temprano)
Society, 64, 406-413. del Chubut. Naturalia Patagonica, Ciencias de la
CAORSI, J. H. & GONI, J. C. 1958. Geologia Uruguaya. Tierra, 4, 17-27.
Boletin del Instituto de Geologia del Uruguay, 37, ETTEMA, C. H. & WARDLE, D. A. 2002. Spatial soil
1-73. ecology. Trends in Ecology and Evolution, 17,
CARVER, M., GROSS, G. F. & WOODWARD, T. E. 1991. 177-183.
Hemiptera. In: CSIRO, DIVISION OF ENTOMOLOGY FARRELL, K. M. 2001. Geomorphology, facies architec-
(ed.) The Insects of Australia L Melbourne Univer- ture, and high-resolution, non-marine sequence
sity Press, Carlton, 429-509. stratigraphy in avulsion deposits, Cumberland
CIFELLI, R. L. 1985. Biostratigraphy of the Casa- Marshes, Saskatchewan. Sedimentary Geology,
mayoran, Early Eocene, of Patagonia. American 139, 93-150.
Museum Novitates, 2820, 1-26. FERUGLIO, E. 1949. Descripcion Geologica de la Patago-
COLLINS, M. S. 1969. Water relations in termites. In: nia, Tomo I. Direction General de Yacimientos
KRISHNA, K. & WEESNER, F. M. (eds). Biology of Petroliferos Fiscales, Buenos Aires.
Termites, Vol. I. Academic Press, New York, FITZPATRICK, E. A. 1984. Micromorphology of Soils.
433^58. Chapman & Hall, London.
CURRAN, A. H. 1994. The paleobiology of ichnocoe- FORD, I. 1988a. Conglomerados con nidos de insectos
noses in Quaternary, bahamian-style carbonate fosiles: Formacion Palmitas (provisorio) - Ter-
environments: the modern to fossil transition. In: ciario Inferior (tentative). Adas del 6to Panel de
DONOVAN, S. K. (ed.) The Paleobiology of Trace Geologia del Literal y Ira Reunion de Geologia
Fossils. Wiley, New York, 83-104. del Uruguay, Salto, 47^9.
D'ALESSANDRO, A. & BROMLEY, R. 1987. Meniscate FORD, I. 1988b. Paleoclima y paleogeografia del Creta-
trace fossils and the Muensteria-Taenidium prob- cico Superior-Terciario Inferior en el Uruguay: un
lem. Palaeontology, 30, 743-763. nuevo modelo interpretative. Adas del 6to Panel
DRIESE, S. G. & FOREMAN, J. L. 1991. Traces and de Geologia del Litoral y Ira Reunion de Geologia
related chemical changes in a Late Ordovician del Uruguay, Salto, 50-53.
paleosol, Glossifungites ichnofacies, southern FORD, I. 1988c. Asociacion caolinita-montmorillonita
Appalachians, USA. Ichnos, 1, 207-219. en perfiles tipo de la Formacion Asencio (Ks).
DRIESE, S., MORA, C. & ELICK, J. 1997. Morphology Adas del 6to Panel de Geologia del Litoral y Ira
and taphonomy of root and stump casts of the Reunion de Geologia del Uruguay, Salto, 42-46.
earliest trees (Middle and Late Devonian), FRENGUELLI, J. 1938. Nidi fossili di Scarabeidi e
Pennsylvania and New York, USA. Palaios, 12, Vespidi. Bolletino Societta Geologia Italiana, 57,
524^537. 77-96.
DROSER, M. L. & BOTTJER, D. J. 1986. A semiquantita- FREY, R. W. & WHEATCROFT, R. A. 1989. Organism-
tive field classification of ichnofabrics. Journal of substrate relations and their impact on sedimen-
Sedimentary Petrology, 56, 558-559. tary petrology. Journal of Geological Education,
EDWARDS, N., JARZEMBOWSKI, E. A., PAIN, T. & 37, 261-279.
DALEY, B. 1988. Cocoon-like trace fossils from FREY, R. W., PEMBERTON, S. G. & FAGERSTROM, J. A.
the lacustrine-palustrine Bembridge Limestone 1984. Morphological, ethological and environ-
Formation (Late Eocene), Southern England. mental significance of the ichnogenera Scoyenia
Proceedings of the Geologists' Association, 109, and Ancorichnus. Journal of Paleontology, 58,
25-32. 511-528.
EKDALE, A. A. & BROMLEY, R. G. 1983. Trace fossils GENISE, J. F. 1993. Trazas fosiles de insectos en paleo-
and ichnofabric in the Kj01by Gaard Marl, suelos. In: MELCHOR, R. N. (ed.) Nuevas tendencias
Upper Cretaceous, Denmark. Bulletin of the en el estudio de trazas fosiles. Facultad de Ciencias
Geological Society of Denmark, 31, 107-119. Exactas y Naturales (UNLPam) La Pampa, 49-59.
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 379

GENISE, J. F. 1997. A fossil termite nest from the GONZALEZ, M. G. 1999. Los paleosuelos de la Forma-
Marplatan stage-age (late Pliocene) of Buenos cion Laguna Palacios (Cretacico Superior) de
Aires province. Argentina, as paleoclimatic indica- Patagonia y la Formacion Asencio (Cretacico
tor. Palaeogeography, Palaeoclimatology, Palaeo- Superior-Terciario Inferior) de Uruguay. Boletim
ecology, 136, 139-144. do 5° Simposio sobre o Cretdceo do Brasil, 65-70.
GENISE, J. F. 1999. Paleoicnologia de Insectos. Revista de GONZALEZ, M. G., TOFALO, O. R. & PAZOS, P. 1998.
la Sociedad Entomologica Argentina, 58, 104-116. Icnologia y paleosuelos del Miembro del Palacio
GENISE, J. F. 2000. The ichnofamily Celliformidae for de la Formacion Asencio (Cretacico Superior-
Celliforma and allied ichnogenera. Ichnos, 1, Terciario Inferior) del Uruguay. Adas del II
267-284. Congreso Uruguay o de Geologia, 38^42.
GENISE, J. F. 2004. Ichnotaxonomy and ichnostrati- Goso AGUILAR, C. & GUEREQUIZ, R. 2001. Hipotesis
graphy of chambered trace fossils in palaeosols acerca del origen de las columnas en las Grutas
attributed to coleopterans, ants and termites. In: del Palacio, Fm. Mercedes-Asencio (Ks), Depto
MclLROY, D. (ed.) 2004. The Application of de Flores (Uruguay). Actas del XI Congreso
Ichnology to Palaeoenvironmental and Strati- Latinoamericano de Geologia y III Congreso
graphic Analysis. Geological Society, London, Uruguayo de Geologia, Montevideo, Contribution
Special Publications, 228, 419^53. 236, 1-13.
GENISE, J. F. & BOWN, T. M. 1994a. New Miocene GRASSE, P. 1984. Termitologia, Tome II. Masson, Paris.
scarabeid and hymenopterous nests and Early GRASSE, P. 1986. Termitologia, Tome III. Masson, Paris.
Miocene (Santacrucian) paleoenvironments, HALFFTER, G. & MATHEWS, G. 1966. The natural
Patagonian Argentina. Ichnos, 3, 107-117. history of dung beetles of the subfamily Scarabaei-
GENISE, J. F. & BOWN, T. M. 1994b. New trace fossils nae. Folia Entomologica Mexicana, 12-14, 1-312.
of termites (Insecta: Isoptera) from the Late HALFFTER, G. & EDMONDS, W. D. 1982. The nesting
Eocene-Early Miocene of Egypt, and the recon- behaviour of dung beetles: an ecological and
struction of ancient isopteran social behaviour. evolutive approach. Publicaciones del Instituto de
Ichnos, 3, 155-183. Ecologia de Mexico, 10, 1-176.
GENISE, J. F. & BOWN, T. M. 1996. Uruguay Roselli and HANSKI, I. & CAMBEFORT, Y. 1991. Resource partition-
Rosellichnus n. ichnogen. two ichnogenera for ing. In: HANSKI, I. & CAMBEFORT, Y. (eds) Dung
clusters of fossil bee cells. Ichnos, 4, 199-217. Beetle Ecology. Princeton University Press,
GENISE, J. F. & HAZELDINE, P. L. 1998a. The ichno- Princeton, 331-349.
genus Palmiraichnus Roselli for fossil bee cells. HANTZSCHEL, W. 1975. Trace fossils and problematica.
Ichnos, 6, 151-166. In: TEICHERT, C. (ed.) Treatise on Invertebrate
GENISE, J. F. & HAZELDINE, P. L. 1998b. 3D-recon- Paleontology (2nd edn, Part W). Geological
struction of insect trace fossils: Ellipsoideichnus Society of America, Boulder, CO/Kansas Univer-
meyeri Roselli. Ichnos, 5, 167-175. sity Press, Lawrence, KS.
GENISE, J. F. & LAZA, J. H. 1998. Monesichnus ameghi- HASIOTIS, S. T. 1997. Ant, termite, bee and wasp, and
noi Roselli: a complex insect trace fossil produced crayfish ichnofabrics: how to differentiate their ich-
by two distinct trace makers. Ichnos, 5, 213-223. nofabric signatures. Abstracts of the Fourth Inter-
GENISE, J. F. & VERDE, M. 2000. Corimbatichnus national Workshop on Ichnofabrics, San Salvador, 1.
fernandezi: a cluster of fossil bee cells from the HASIOTIS, S. T. 2000. The invertebrate invasion
Late Cretaceous-Early Tertiary of Uruguay. and evolution of Mesozoic soil ecosystems: the
Ichnos,!, 115-125. ichnofossil record of ecological innovations. In:
GENISE, J. F. & ZELICH, M. R. 2001. Trazas fosiles de GASTALDO, R. A. & DIMICHELE, W. A. (eds)
insectos de la Formacion Puerto Unzue (Cretacico Phanerozoic Terrestrial Ecosystems. The Paleonto-
Superior-Paleogeno) de Entre Rios. Resumenes de logical Society Papers, 6, 141-169.
la IV Reunion Argentina de Icnologia y Segunda HASIOTIS, S. T. & BOWN, T. M. 1992. Invertebrate trace
Reunion de Icnologia del Mercosur, Tucuman, 44. fossils: the backbone of continental ichnology. In:
GENISE, J. F., MANGANO, M. G., BUATOIS, L. A., LAZA, MAPLES, C. G. & WEST, R. R. (eds) Trace Fossils.
J. H. & VERDE, M. 2000. Insect trace fossil associa- Short Courses in Paleontology, 5, 15-33.
tions in palaeosols: the Coprinisphaera ichnofacies. HASIOTIS, S. T. & DUBIEL, R. F. 1994. Ichnofossil
Palaios, 15, 49-64. tiering in Triassic alluvial palaeosols: implications
GENISE, J. F., CONTRERAS, V. H. & CILLA, G. 2001. for pangean continental rocks and paleoclimate.
Trazas de Equisetales en paleosuelos de la Forma- Memoirs of the Canadian Society of Petroleum
cion Ischigualasto (Triasico) de San Juan. Resu- Geologists, 17,311-317.
menes de la IV Reunion Argentina de Icnologia y HASIOTIS, S. T. & MITCHELL, C. E. 1993. A comparison
Segunda Reunion de Icnologia del Mercosur, Tucu- of crayfish burrow morphologies: Triassic and
man, 46. Holocene paleo- and neoichnological evidence,
GENISE, J. F., SCIUTTO, J. C, LAZA, J. H., GONZALEZ, and the identification of their burrowing signa-
M. G. & BELLOSI, E. 2002. Fossil bee nests, coleop- tures. Ichnos,2, 291-314.
teran pupation chambers and tuffaceous palaeo- HASIOTIS, S. T., ASLAN, A. & BOWN, T. M. 1993.
sols from the Late Cretaceous Laguna Palacios Origin, architecture, and paleoecology of the
Formation, Central Patagonia (Argentina). Early Eocene continental ichnofossil Scaphich-
Palaeogeography, Palaeoclimatology, Palaeoecol- nium hamatum, integration of ichnology and
ogy, 111, 215-235. paleopedology. Ichnos, 3, 1-9.
380 J. F. GENISE ET AL.

HASIOTIS, S. T., DUBIEL, R. F. & DEMKO, T. M. 1995. LAZA, J. H. 1986. Icnofosiles de paleosuelos del
Triassic hymenopterous nests: insect eusociality Cenozoico mamalifero de Argentina. I Paleogeno.
predates angiosperm plants. Abstracts of Papers Boletin de la Asociacion Paleontologica Argentina,
at the Meeting of the Rocky Mountain Section of 15, 19.
the Geological Society of America, 27, 13. LEE, K. E. 1985. Earthworms: Their Ecology and
HOLLDOBLER, B. & WILSON, E. O. 1990. The Ants. Relationships with Soils and Land Use. Academic
Harvard University Press, Cambridge, MA. Press, London.
IRIONDO, M. 1999. The origin of silt particles in the LEE, K. E. & WOOD, T. 1971. Termites and Soils.
loess question. Quaternary International, 62, 3-9. Academic Press, London.
JIMENEZ, J. J., BROWN, G. G., DECAENS, T., FEIJOO, A. LINSLEY, E. G. 1958. The ecology of solitary bees.
& LAVELLE, P. 2000. Differences in the timing of Hilgardia, 21, 543-599.
diapause and patterns of aestivation in tropical LUMARET, J. P. 1983. Structure des peuplements de
earthworms. Pedobiologia, 44, 677-694. coprophages Scarabaeidae en region mediter-
JOHNSTON, P. A., EBERTH, D. A. & ANDERSON, P. K. raneenne fran9aise: relations entre les conditions
1996. Alleged vertebrate eggs from Upper Cretac- ecologiques et quelques parametres biologiques
eous redbeds, Gobi Desert, are fossil insect des especes. Bulletin de la Societe Entomologique
(Coleoptera) pupation chambers: Fictovichnus de France, 88, 481-495.
new ichnogenus. Canadian Journal of Earth MARTINEZ, R. N., ALCOBER, O. A. & MILANA J. P.
Sciences, 33, 511-525. 1998. Variation vertical del ambiente sedimentario
JONKMAN, J. C. 1980. The external and internal struc- de la Formacion Ischigualasto (Triasico superior,
ture and growth of nests of the leaf-cutting ant Carniano), San Juan, Argentina. Acta Geologica
Atta vollenweideri¥orQ\, 1893 (Hym.:Formicidae). Lilloana, 18, 166-167.
Part II. Zeitschrift fur angewandte Entomologie, MATHEWS. A. G. 1977. Studies on Termites from the
89, 217-246. Mato Grosso State, Brazil. Academia Brasileira
KAY, R. F., MADDEN, R. H., VUCETICH, M. G., CAR- de Ciencias, Rio de Janeiro.
LINI, A. A., MAZZONI, M. M., RE, G. H., HEIZLER, MAZZONI, M. M. 1979. Contribution al conocimiento
M. & SANDEMAN, H. 1999. Revised geochronology petrografico de la Formacion Sarmiento. Revista
of the Casamayoran South America Land de la Asociacion Argentina de Mineralogia, Petro-
Mammal Age: climatic and biotic implications. logia y Sedimentologia, 10, 33-54.
Proceedings of the National Academy of Sciences, MAZZONI, M. M. 1985. La Formacion Sarmiento y el
96, 13235-13240. vulcanismo paleogeno. Revista de la Asociacion
KAY, R. F., MADDEN, R. H., BELLOSI, E., CARLINI Geologica Argentina, 40, 60-68.
A. A., HEIZLER, M., RE, G., VILAS, F. & VUCETICH MclLROY, D. 2004a. A review of some ichnological
M. G. 2001. Puesto Almendra - Colhue-Huapi concepts, methodologies, applications and fron-
contact at Gran Barranca. Ameghiniana, 38, 35R. tiers. In: MclLROY, D. (ed.) 2004. The Application
KEIGHLEY, D. G. & PICKERILL, R. K. 1994. The of Ichnology to Palaeoenvironmental and Strati-
ichnogenus Beaconites and its distinction from graphic Analysis. Geological Society, London,
Ancorichnus and Taenidium. Palaeontology, 37, Special Publications, 228, 3-27.
305-337. MclLROY, D. 2004b. Ichnofabrics and sedimentary facies
KIRK, A. A. 1983. The biology of Bubas bison (L.) of a tide-dominated delta: Jurassic He Formation
(Coleoptera: Scarabaeidae) in southern France of Kristin Field, Haltenbanken, offshore mid-
and its potential for recycling dung in Australia. Norway. In: MclLROY, D. (ed.) 2004. The Applica-
Bulletin of Entomological Research, 73, 129-136. tion of Ichnology to Palaeoenvironmental and Strati-
KOKOGIAN, D., SPALLETTI, L. ET AL. 1999. Los deposi- graphic Analysis. Geological Society, London,
tos continentales triasicos. In: CAMINOS, R. (ed.) Special Publications, 228, 237-272.
Geologia Argentina. Anales del Institute de Geolo- MELCHOR, R. N., GENISE, J. F. & VERDE, M. 2001.
gia y Recursos Minerales, 29, 377-398. Invertebrate trace fossils from Triassic continental
KRAUS, M. J. 1996. Avulsion deposits in lower Eocene sequences of San Juan province, Argentina. Publi-
alluvial rocks, Bighorn Basin, Wyoming, USA. cation Especial de la Asociacion Paleontologica
Journal of Sedimentary Research, 66, 354-366. Argentina, 7, 127-131.
KRAUS, M. J. & ASLAN, A. 1998. Palaeosol sequences in MICHENER, C. D. 1979. Biogeography of the bees. Annals
floodplain environments: a hierarchical approach. of the Missouri Botanical Garden, 66, 277-347.
In: THIRY, M. (ed.) Palaeoweathering, Palaeo- MILANA, J. P. & ALCOBER, O. A. 1994. Modelo tectosedi-
surfaces and Related Continental Deposits. Inter- mentario de la cuenca Triasica de Ischigualasto (San
national Association of Sedimentology, Special Juan, Argentina). Revista de la Asociacion Geologica
Publications, Blackwell, Oxford, 27, 303-321. Argentina, 49, 217-235.
KRAUS, M. J. & GWINN, B. 1997. Facies and facies MILANA, J. P., ALCOBER, O. A. & MARTINEZ, R. N.
architecture of Paleogene floodplain deposits, 1998. La mega-arquitectura depositacional de la
Willwood Formation, Bighorn Basin, Wyoming, Formacion Ischigualasto como control paleo-
USA. Sedimentary Geology, 114, 33-53. ecologico de las faunas de paleovertebrados.
LAZA, J. H. 1982. Signos de actividad atribuibles a Atta Acta Geologica Lilloana, 18, 169-170.
(Myrmicidae) en el Mioceno de la Provincia de La MILLER, M. F. & SMAIL, S. E. 1997. A semiquantitative
Pampa, Republica Argentina. Signification paleo- field method for evaluating bioturbation on
zoogeografica. Ameghiniana, 19, 109-124. bedding planes. Palaios, 12, 391-396.
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 381

MILLER, W. R. & MASON, T. R. 2000. Stellavelum RETALLACK, G. J. 1994. The Environmental Factor
arborensis igen. sp. nov., Stellavelum uncinum Approach to the Interpretation of Palaeosols.
igen., isp. nov. and Termitichnus namibiensis isp. Special Publications of the Soil Science Society
nov.; new ichnofossils from Cenozoic deposits of of America, 33, 31-63.
Namaqualand, South Africa. Ichnos, 7, 195-215. RETALLACK, G. J. 200la. Soils of the Past (2nd edn).
NAHON, D. 1976. Cuirasses ferrugineuses et encroute- Blackwell, Oxford.
ments calcaires au Senegal Occidental et en RETALLACK, G. J. 200 Ib. Scoyenia burrows from
Mauritanie, systemes evolutifs: geochimie, struc- Ordovician palaeosols of the Juniata Formation
tures, relais et coexistence. Memoirs de Sciences in Pennsylvania. Palaeontology, 44, 209-235.
Geologiques, 44, 1-232. RETALLACK, G. J., BESTLAND, E. & FREMD, T. 2000.
NAHON, D. 1986. Evolution of iron crusts in tropical Eocene and Oligocene Paleosols of Central
landscapes. In: COLMAN, S. M. & DETHEER, D. P. Oregon. Special Papers of the Geological Society
(eds) Rates of Chemical Weathering of Rocks and of America, 344, 1-192.
Minerals. Academic Press, London, 169-191. RICHTER, R. & RICHTER, E. 1939. Marken und Spuren
NETTLETON, W. D., OLSON, C. G. & WYSOCKI, D. A. aus alien Zeiten. III. Eine Lebens-Spur (Synco-
2000. Palaeosol classification: problems and prulus pharmaceus}, gemeinsam den rheinischen
solutions. Catena, 41, 61-92. und bohmischen Ordovicicum. Senckenbergiana,
NETTO, R. G. 2000. Icnofabricas: conceptos y aplica- 21, 152-168.
bilidad. Not as del Cur so de Actualizacion sobre ROSELLI, F. L. 1938. Apuntes de geologia y paleontolo-
Icnologia: aplicaciones en la geologia sedimentaria gia uruguaya. Sobre insectos del Cretacico del
y la industria petrolera. II Congreso Latinoameri- Uruguay o descubrimiento de admirables instintos
cano de Sedimentologia y VII Reunion Argentina constructivos de esa epoca. Boletin de la Sociedad
de Sedimentologia, Mar del Plata, 13-25. Amigos de las Ciencias Naturales 'Kraglievich-
PAZOS, P. J., TOFALO, O. R. & GONZALEZ, M. 1998. La Fontana', 1, 72-102.
paleosuperficie Yapeyu: significado estratigrafico ROSELLI, F. L. 1987. Paleoicnologia: nidos de insectos
y paleoambiental en la evolution del Cretacico fosiles de la cubertura Mesozoica del Uruguay.
Superior del Uruguay. Actas del II Congreso Publicaciones del Museo Municipal de Nueva
Uruguayo de Geologia, Montevideo, 59-63. Palmira, 1, 1-56.
PEMBERTON, S. G., FREY, R. W., RANGER, M. J. & ROUBIK, D. W. & MICHENER, C. D. 1980. The seasonal
MAC£ACHERN, J. 1992. The conceptual framework cycle and nests of Epicharis zonata, a bee whose
of ichnology. In: PEMBERTON, S. G. (ed.) Applica- cells are below the wet-season water table (Hyme-
tions of Ichnology to Petroleum Exploration. noptera, Anthophoridae). Biotropica, 12, 56-60.
SEPM Core Workshops, Calgary, 17, 1-32. ROY-NOEL, J. 1972. Recherches sur 1'ethologie des
PICKERILL, R. K. 1989. Compaginatichnus: a new isopteres de la presqu'ile du Cap-Vert (Senegal).
ichnogenus from Ordovician flysch of eastern Bulletin Biologique de France et Belgique, 106,
Canada. Journal of Paleontology, 63, 913-919. 193-283.
POLLARD, J. E., GOLDRING, R. J. & BUCK, S. G. 1993. SAN JOSE, J. J., MONIES, R., STANSLY, P. A. & BENTLEY,
Ichnofabrics containing Ophiomorpha: their B. L. 1989. Environmental factors related to the
significance in shallow-water facies interpretation. occurrence of mound-building nasute termites in
Journal of the Geological Society, 150, 149-164. Trachypogon savannas of the Orinoco Llanos.
RATCLIFFE, B. C. & FAGERSTROM, J. A. 1980. Inverte- Biotropica, 21, 353-358.
brate lebensspuren of Holocene floodplains: SAUER, W. 1955. Coprinisphaera ecuadoriensis, un fosil
their morphology, origin and paleoecological singular del Pleistocene. Boletin del Instituto de
significance. Journal of Paleontology, 54, 614- Ciencias Naturales del Ecuador, 1, 123-132.
630. SAVRDA, C. E., BLANTON-HOOKS, A. D. et al. 2000.
REINECK, H. E. 1963. Sedimentgefuge im Bereich der Taenidium and associated ichnofossils in fluvial
siidlichen Nordsee. Abhandlungen der senckenber- deposits, Cretaceous Tuscaloosa Formation,
gische naturfoschende Gesellschaft, 505, 1—138. Eastern Alabama, Southeastern USA. Ichnos, 1,
RETALLACK, G. J. 1984. Trace fossils of burrowing 227-242.
beetles and bees in an Oligocene palaeosol, Bad- SAYAGO, J., COLLANTES, M., KARLSON, A. & SANABRIA,
lands National Park, South Dakota. Journal of J. 2001. Genesis and distribution of the Late
Paleontology, 58, 571-592. Pleistocene and Holocene loess of Argentina: a
RETALLACK, G. J. 1985. Fossil soils as grounds for regional approximation. Quaternary International,
interpreting the advent of large plants and animals 76/77, 247-257.
on land. Philosophical Transactions of the Royal SCHMIDT, R. S. 1960. Functions of Apicotermes nests.
Society of London, Series B, 309, 105-142. Insectes Sociaux, 1, 357-368.
RETALLACK, G. J. 1988. Field recognition of palaeo- SCHWERTMANN, U. 1988. Occurrence and formation of
sols. In: REINHARDT, J. & SIGLEO, W. R. (eds) iron oxides in various pedoenvironments. In:
Palaeosols and Weathering through Geologic STUCKI, J. W., GOODMAN, B. A. & SCHWERTMANN,
Time: Principles and Applications. Special U. (eds) Iron and Soils and Clay minerals. Reidel,
Papers of the Geological Society of America, Dordrecht, 267-306.
216, 1-20. SCIUTTO, J. C. 1981. Geologia del Codo del Rio
RETALLACK, G. J. 1990. Soils of the Past. Unwin Senguerr, Chubut, Argentina. Actas del VIII
Hyman, Boston. Congreso Geologico Argentino, III, 203-219.
382 J. F. GENISE ET AL.

SIMPSON, G. G. 1940. Review of the mammal bearing Journal of the Geological Society of London, 150,
Tertiary of South America. Proceedings of the 141-148.
American Philosophical Society, 83, 649-710. TAYLOR, A. M., GOLDRING, R. & GOWLAND, S. 2003.
SOIL SURVEY STAFF. 1975. Soil Taxonomy: A Basic Analysis and application of ichnofabrics. Earth
System of Soil Classification for Making and Inter- Science Reviews, 60, 227-259.
preting Soil Surveys. United States Department of TEMGOUA, E. 2001. Les accumulations ferrugineuses
Agriculture, National Resources Conservation actuelles de bas de versants en zone forestiere
Service, Agriculture Handbook, Washington, DC. humide du Sud-Cameroun: evolutions petrologi-
SPALLETTI, L. A. 2001. Evolution de cuencas sedi- ques des fades et des elements traces en relation
mentarias. In: ARTABE, A. E., MOREL, E. M. & avec le cuirassesment. Memoires de Geologie de la
ZAMUNER, A. (eds) El Sistema Tridsico en la Argen- Universite de Lausanne, 38, 1-134.
tina. Fundacion Museo de La Plata 'Francisco P. TERUGGI, M. E. 1957. The nature and origin of Argen-
Moreno', La Plata, 81-101. tine loess. Journal of Sedimentary Petrology, 27,
SPALLETTI, L. A. & MAZZONI, M. M. 1977. Sedimento- 322-332.
logia del Grupo Sarmiento en un perfil ubicado al TERUGGI, M. E. 1971. Criterios para el reconocimiento
sudeste del Lago Colhue Huapi, provincia de y estudio de los paleosuelos. Revista de la Asocia-
Chubut. Obra del Centenario del Museo de La cion Geologica Argentina, 26, 485-490.
Plata, Geologia, 4, 261-285. VERDE, M., GENISE, J. F., UBILLA, M. & JIMENEZ, J. J.
SPALLETTI, L. A. & MAZZONI, M. M. 1979. Estratigra- 2002. Camaras de estivation de lombrices en
fia de la Formation Sarmiento en la barranca sur sedimentos del Pleistocene tardio de Uruguay
del lago Colhue-Huapi, provincia del Chubut. (Formacion Sopas), implicancias icnologicas y
Revista de la Asociacion Geologica Argentina, 34, paleoambientales. Adas de las II Jornadas
271-281. Uruguayas del Cenozoico, Montevideo, 87.
STEPHEN, W. P., BOHART, G. E. & TORCHIO, P. F. 1969. VISSCHER, P. K., VETTER, R. S. & ORTH, R. 1994.
The Biology and External Morphology of Bees, with Benthic bees? Emergence phenology of Calliopsis
a Synopsis of the Genera of Northwestern America. pugionis (Hymenoptera: Andrenidae) at a season-
Agricultural Experimental Station, Oregon State ally flooded site. Annals of the Entomological
University, Corvallis, OR. Society of America, 87, 941-945.
STIPANICIC, P. N. 2001. Antecedentes geologicos y WENZEL, J. W. 1992. Behavioural homology and phylo-
paleontologicos. In: ARTABE, A. E., MOREL, E. M. geny. Annual Review of Ecology and Systematics,
& ZAMUNER, A. (eds) El Sistema Tridsico en la 23, 361-381.
Argentina. Fundacion Museo de La Plata WETZEL, A. & UCHMAN, A. 1998. Deep-sea benthic
'Francisco P. Moreno', La Plata, 1-21. food content recorded by ichnofabrics: a concep-
STIPANICIC, P. N. & BONAPARTE, J. F. 1979. Cuenca tual model based on observations from Paleogene
Triasica de Ischigualasto-Villa Union (provincias flysch, Carpatians, Poland. Palaios, 13, 533—546.
de San Juan y La Rioja). In: LEANZA, A. F. (ed.) WHEELER, W. M. 1910. Ants. Columbia University
Geologia Regional Argentina. Academia Nacional Press, New York.
de Ciencias, Buenos Aires, 507-536. WILLIS, E. R, & ROTH, L. M. 1962. Soil and moisture
TARDY, Y. 1992. Diversity and terminology of lateritic relations of Scaptocoris divergens Troeschner
profiles. In: MARTINI, I. P. & CHESWORTH, W. (eds) (Hemiptera: Cydnidae). Annals of the Entomologi-
Weathering, Soils andPalaeosols. Developments in cal Society of America, 55, 21-32.
Earth Surface Processes, Elsevier, Amsterdam, 2, YAALON, D. H. 1971. Paleopedology: Origin, Nature
379-405. and Dating of Palaeosols. Israel University Press,
TARDY, Y. & ROQUIN, C. 1992. Geochemistry and Jerusalem.
evolution of lateritic landscapes. In: MARTINI, ZAMUNER, A. B., ZAVATTIERI, A. M., ARTABE, A. E. &
I. P. & CHESWORTH, W. (eds) Weathering, Soils MOREL, E. M. 2001. Paleobotanica. In: ARTABE,
and Palaeosols. Developments in Earth Surface A. E., MOREL, E. M. & ZAMUNER, A. (eds) El
Processes, Elsevier, Amsterdam, 2, 407-443. Sistema Tridsico en la Argentina. Fundacion
TAYLOR, A. M. & GOLDRING, R. 1993. Description Museo de La Plata 'Francisco P. Moreno', La
and analysis of bioturbation and ichnofabric. Plata, 143-184.
Development of early Palaeozoic ichnofabrics:
evidence from shallow marine siliciclastics

MARY L. DROSER 1 , S0REN JENSEN1 & JAMES G. GEHLING2


1
Department of Earth Sciences University of California, Riverside, CA 92521, USA
(e-mail: Mary.Droser@ucr.edu)
2
South Australian Museum, Division of Natural Sciences, North Terrace,
Adelaide 5000, South Australia, Australia

Abstract: Earliest Cambrian fine-grained sediments appear to have been firm close to the
sediment-water interface. As a result there was a high preservational potential of shallow
tiers. There is limited evidence for a mixed layer at this time; rather, most of the preserved
trace fossils were open burrows. Later in the Cambrian, depth of sediment mixing increased
but firmground conditions are still found relatively close to the sediment-water interface.
The development of the mixed layer and properties of Cambrian muddy sediments have
numerous stratigraphic and ichnological consequences. These include secular trends in the
preservation of event beds and shallow-tier trace fossils including Rusophycus and Cruziana.

Trace fossils and ichnofabric have long played a interface at which an animal(s) lives (Ausich &
role in stratigraphic and sedimentological ana- Bottjer 1982). Thus, in modern marine environ-
lyses (e.g. Bromley 1996). The advent of sediment ments, invertebrate animals live at a variety of
mixing has important implications for a number depths within the sediment, and thus occupy a
of sediment properties. Sediment mixing by number of different tiers. Of course, animals
animals greatly influences oxygenation of the may live in, but not mix, the sediment. Sus-
sediments, the geochemistry of the sediments pension-feeding animals that build a domicile
and the distribution of organic material, and is open vertical burrow, such as a Skolithos-type
important for nutrient cycling (Aller 1983; Mcll- burrow, do not actively mix the sediment.
roy & Logan 1999). The onset of bioturbation Many researchers have assumed that the
might have played a part in shifts in the carbon development of the sediment mixing and thus
and oxygen isotope record, nutrient cycling, the mixed layer occurred at or near the Neopro-
and the distribution of organic material (Brasier terozoic-Cambrian transition, but recent work
1990; Brasier & Mcllroy 1998; Mcllroy & Logan has indicated a more complicated history. Earliest
1999). Another aspect of sediment mixing is the Cambrian strata show very little evidence of a
destruction of primary bedding and changes in mixed layer. Indeed, data suggest that the lack
the rheological properties of the sediment. The of a well-developed mixed layer resulted in fine-
preservation potential of physical sedimentary grained sediments that were firm close to or at
structures was highest before the advent of the sediment-water interface at this time because
infaunal activity in the latest Proterozoic (e.g. of a lack of bioturbation or mixing and a resultant
Eriksson et al. 1998). An additional aspect of well-developed mixed layer (Droser et al. 2002a,
the rise of bioturbation is the increased destruc- 2002b). We have previously suggested that firm
tion of thin storm event beds. Sepkoski et al. muddy sediments that form at a shallow depth
(1991) proposed a secular trend in event bed (e.g. are not exposed at the surface because of
preservation as a result of increased infaunal erosion of overlying soft or soupy sediments)
activity: that is, as depth and extent of bioturba- can be directly related to the shallow sediment
tion increase, event beds are less well preserved. mixing.
The upper portion of the sediment that is In this paper we review the evidence for firm
actively being burrowed and thoroughly mixed substrates in Lower Cambrian strata and present
is known as the sediment mixed layer. Applied relevant data from Lower Cambrian through
originally to deep-sea sediments (Berger et al. Cambro-Ordovician strata to evaluate the
1979), this term is now commonly used to refer nature and timing of the development of the
to the actively burrowed zone from a variety mixed layer. We have examined several aspects
of marine environments (e.g. Bromley 1996; of the Cambrian ichnological record including:
Savrda & Bottjer 1989). Related, but certainly
not synonymous, is the concept of tiering. trace fossil preservation;
Tiering is the depth below the sediment-water preserved depth of bioturbation;

From: MC!LROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 383-396. 0305-8719/04/S15.00 © The Geological Society
of London.
384 M. L. DROSER ET AL.

nature of ichnofabric (all aspects of the trace sandstone, siltstone and mudstone. In this
fossil record include features such as mottled setting, even in earliest Cambrian strata, trace
bedding resulting from sediment mixing fossils are conspicuous. Indeed trace fossils are
where discrete trace fossils cannot be identi- an important part of lowermost Cambrian
fied); and stratigraphy and define the base of the Cambrian
nature of the substrate, which has been recog- system (Crimes 1987; Narbonne et al 1987;
nized as a factor in trace fossil preservation. Brasier et aL 1994). Of the units that we exam-
ined, the Lower Cambrian formations exhibit
This paper is based on field and laboratory a number of shared ichnological and sedimento-
examination of a large number of Cambrian logical characteristics. These characteristics are
siliciclastic units, trace fossils and sedimentary ubiquitous in Cambrian strata of the earliest
structures. The bulk of the record through this Cambrian (Nemakit-Daldynian) (Fig. 1).
interval is siliciclastic, and trace fossils are best
observed in relatively thin-bedded siliciclastic
sediments. These include units of the Cambro-
Ordovician Bell Island Group (~120m) and Preserved depth of bioturbation
Lower Ordovician Wabanna Group (20m), In modern settings, burrows, tracks and trails
Newfoundland; the Lower Cambrian Mickwitzia produced near the surface have a very low
sandstone (10m), Sweden; the Cambro-Ordovi- chance of preservation because of physical pro-
cian Bynguano Formation (30m), New South cesses resulting in erosion, and because the
Wales, Australia; the Lower Cambrian Lontova potential trace fossils are destroyed by those
(10m) and Liikati (15m) formations, Estonia; animals that subsequently burrow deeply into
the Lower Cambrian Wood Canyon (100m), the sediment. Lowermost Cambrian sediments
Pioche (20m) and Harkless (5m) formations, preserve a range of trace fossils that are inter-
western USA; the Lower Cambrian Arumbera preted as representing shallow tiers: that is, the
Sandstone (200m) and the Cambro-Ordovician burrows did not extend more than a few
Pacoota Sandstone (200m), Northern Terri- centimetres at most below the sediment-water
tories, Australia. interface (Droser et aL 2002a, 2002b).
We examined parts of these units that These shallow-tiered burrows include various
represent deposition on the shelf below fair- 'treptichnids' (including Treptichnus pedum).
weather wavebase and above maximum storm They consisted of additions of curved elements;
wavebase, as determined by independent the burrows themselves were open and the tops
sedimentological criteria such as hummocky extended to the sediment-water surface (Fig. 1).
cross-stratification, presence of event beds and The geometry and style of preservation of these
stratigraphic context. The measured portions of trace fossils suggest that they formed less than
these units (indicated in parentheses above) are a few centimetres below the sediment-water sur-
characterized by heterolithic bedding ranging face (Droser et al. 2002b). Other burrows include
from the centimetre to decimetre scale. Detailed Gyrolithes, a corkscrew-type trace fossil that has
logs were made of all sections, and selected a preserved depth of 1-2 cm. The diameter of this
intervals were described at the centimetre scale. burrow is of the order of millimetres, and it is
The sedimentology, ichnofabrics and trace interpreted to have been an open-burrow
fossil taxa were described in relation to the system. Gyrolithes is abundant in the Chapel
sedimentary context (e.g. preservation). Rock Island Formation of Newfoundland and in the
samples were taken at decimetre intervals where Lower Cambrian units of Baltica (Fedonkin
possible, cut into slabs and polished. Selected 1983; Jensen & Mens 1999)
slabs were X-rayed.

Quality of preservation
Characteristics of the ichnological record of
lowermost Cambrian strata Even though 'treptichnid' burrows were con-
structed close to the sediment-water interface
Below we summarize results from previous work. they have sharp walls without actively reinforced
The greatest diversity of Cambrian trace fossils margins, and in certain cases delicate surface
have been reported from settings representing ornamentation is preserved. Compaction of
deposition below fair-weather wavebase and the burrows is also relatively minor. Several
above storm wavebase (e.g. shelf) that have a other shallow-tier trace fossils show excellent
heterolithic bedding characterized by moderately preservation of detail, including Gyrolithes and
thin, generally centimetre-scale, interbedded Rusophycus. This quality of preservation is
EARLY PALAEOZOIC ICHNOFABRICS 385

Fig. 1. Schematic representation of the ichnofabric and sediment response to storm-related erosion positioned
at approximate stratigraphic level. Sand is indicated by stippled pattern, mud by white. The mixed layer is
depicted by dense patterning. Increased depth of mixing leads to increased erodability and partial removal of
the shallower tier.

ubiquitous in the Lower Cambrian units rather resistant to erosion allowing the trapping
examined. of sand in burrows rather than the destruction
of the burrows (Fig. 1).

Styles of preservation
The nature of ichnofabric
In most shallow marine settings, particularly in
the Palaeozoic, well-preserved burrows on the Some animals that burrow do not leave well-
base of sandstone beds are created by animals defined discrete trace fossils. Instead, the record
that burrow through the sand to the interface produced is one of some degree of homogeniza-
with the underlying finer-grained sediment (Sei- tion, where primary sedimentary structures are
lacher 1970, 1985). In Lower Cambrian strata a not preserved. The final texture has a mottled
fundamentally different style of preservation appearance. This is direct evidence of a mixed
appears to be common, for shallow tiers in parti- layer. For example, in a modern setting under
cular. The burrow may be cast by a source bed to normal marine conditions in environments not
which it remains attached, or it may be cast by characterized by rapid deposition such as pro-
sand that bypassed the seafloor, and thus the delta settings, where there is alternating deposi-
cast is subsequently attached to the base of a tion of sand and mud, these two lithologies
different bed (adhered preservation), or may could be completely mixed, depending on the
even be preserved as a sand-filled burrow com- rates of deposition. In sedimentary rocks of the
pletely in silt (floating preservation) (Droser earliest Cambrian, rare isolated homogenized
et al. 2002b). This is a common type of preserva- beds occur less than 1 cm in thickness. Otherwise,
tion of treptichnids, Gyrolithes and Palaeophy- there is no evidence of sediment mixing. Sand
cus/Planolites-type burrows, and is the most and mud beds remain remarkably discrete, with
common style of trace fossil preservation in all sharp boundaries.
the Lower Cambrian units examined. This type
of preservation does occur in the Phanerozoic,
and reflects penecontemporaneous erosion (e.g. Properties of earliest Cambrian muddy
Hallam 1975) sediments
This type of preservation requires that
burrows are open and possibly empty. The The features described above, such as the preser-
preservation of shallow-tier trace fossils in this vation of sharp burrow margins with delicate
manner suggests that the muddy sediment was scratch marks and the low degree of compaction,
386 M. L. DROSER ET AL.

characterize firmground conditions (Goldring Ichnological record of the late Early


1995). A firmground indicates stiff but uncemen- Cambrian to earliest Ordovician
ted sediment. In modern settings, firmgrounds
are exposed at the surface after erosion of There was an increase in the diversity of trace
upper layers; firm conditions at depth generally fossils throughout the Early Cambrian, and the
result from advanced dewatering and compac- majority of marine ichnogenera had appeared
tion (Bromley 1996). In the Early Cambrian, by the Atdabanian (Crimes 1992). Of particular
compaction would not be a requisite process, note is the increase in range of size, which is
but, rather, silty mud would be deposited and significant as, other factors being equal, the
in the absence of bioturbators would tend to rate of sediment mixing is proportionally linked
dewater more rapidly (Droser et al. 2002a). to animal cross-section (Piper & Marshall 1969;
These firmgrounds are not comparable to firm Thayer 1983; Mcllroy & Logan 1999). Trilobites
carbonate sediments that exhibit early lithifica- appear in the Atdabanian (though predated by
tion. That is, when eroded, these sediments pre- trilobite trace fossils), and were among the
sumably could have quickly disaggregated into larger animals that disturbed sediment.
individual grains rather than erode as mud All of the units examined share a number of
chips, though flat pebble conglomerates and significant ichnological characteristics in spite
chips resulting from organic binding can occur. of the fact that they represent a variety of shallow
In recent years there has been a growing body subtidal terrigenous clastic environments. In
of evidence that terminal Proterozoic sediment comparison with the earliest Cambrian strata:
surfaces were bound by microbial mats to a far
greater extent than would be typical of most of Shallow-tier burrows (less than 5 cm in depth)
the Phanerozoic (e.g. Seilacher & Pfluger 1994; such as Treptichnus pedum and Gyrolithes are
Gehling 1999). The siltstone-sandstone succes- not generally common in Upper Cambrian-
sion of the earliest Cambrian units that we exam- Lower Ordovician strata. However, both of
ined shows no typical mat-related structures such these trace fossils can be common in some of
as peetee structures, elephant skin texture or the Atdabanian units, such as the Mickwitzia
pyrite-rich horizons. Wrinkle marks that have Sandstone and the Pacoota Sandstone. Other
been interpreted as mat-related structures (cf. common Cambrian burrows, such as Phy-
Hagadorn & Bottjer 1997) do occur on a few codes, Teichichnus, Rusophycus and Cruziana,
surfaces in several, but not all of the units. are interpreted to be relatively shallow in
Thus there is no compelling evidence that mats depth by most workers, but would have been
were an important component of these fine- emplaced deeper than T. pedum and Gyro-
grained sediments (but see Goldring & Jensen lithes, perhaps shallower than between 5 and
1996). 10 cm (Figs 2-5). With the exception of vertical
That Cambrian and Lower Ordovician sedi- burrows such as Skolithos and Arenicolites
ments were firm near the surface is also suggested (and perhaps some Teichichnus) emplaced in
by the presence of particular sedimentary struc- sand, this second, deeper tier represents the
tures that must have been formed close to the bulk of the common Cambrian trace fossil
sediment-water interface. This includes Kullin- record. This tier is unlikely to be preserved in
gia-type scratch circles, which form when a Recent normal marine shelfal settings (e.g.
tethered organism is rotated by currents (Jensen Bromley 1996). Indeed, the presence of this
et al. 2002). The circles are formed in silts or tier is commonly used to suggest that condi-
fine sands, and are cast by overlying coarser tions were anoxic or otherwise, not normal
material. Their preservation requires that sedi- marine (Gibert & Ekdale 1999).
ment just beneath the sediment-water interface Preservation asfloatingand adhering burrows
is firm enough to imprint delicate concentric (Droser et al. 2002b) remains common
structures and also to withstand the erosion of through Upper Cambrian-Lower Ordovician
currents in subtidal shallow-marine settings. strata (Fig. 4b), and also occurs in younger
Scratch circles are most common in lowermost strata (Simpson 1957; R. Goldring, personal
Cambrian terrigenous clastic strata including communication 2003). Open mud burrows
the Chapel Island, the Uratanna, and the Torne- cast by sand are also common in these strata.
trask formations, as well as the Mickwitzia sand- Quality of preservation (e.g. scratch marks and
stone of the units that we have examined, and minor compaction) is very high in all of these
they are reported from the Khmelnitsk Forma- units. A conspicuous element of this interval
tion of the Ukraine (see Jensen et al. 2002) and is the trace fossils Rusophycus and Cruziana,
Arumbera Sandstone (D. Mcllroy, personal which are common in all the formations exam-
communication, 2003). ined in spite of the fact that they represent a
EARLY PALAEOZOIC ICHNOFABRICS 387

Fig. 2. Ichnology and ichnofabric of the Lower Ordovician of Grebs Nest Point Formation, Grebes Nest
Point, Bell Island, (a) Lower surface of bed with diverse trace fossils preserved by sand in mud. Prominent
trace fossils include Cruziana and several Trichophycus. Diameter of coin is about 2.5cm. (b) Abundant
sand-filled Trichophycus in mud and sharp erosively based event bed. Scale bar is 20 mm.

range of depositional environments. Scratch commonly pipe sediment from one to the
marks are routinely exquisitely preserved. other (Figs 3, 4). This is in contrast to shelfal
Nearly all these units have at least thin mud sediments of the Mesozoic that, when viewed
beds that are thoroughly mixed as viewed in in core, are thoroughly mixed (e.g. Bockelie
cut slab and/or X-ray. This represents direct 1991, figs 2, 3).
evidence of a mixed layer. However, at least
at the scale of centimetres, mud and sand Compared with the earliest Cambrian it is more
beds remain discrete, although burrows common to find burrows that penetrate sands.
388 M. L. DROSER ET AL.

Fig. 3. Polished vertical section of heterolithic bedding in the Grebes Nest Point Formation (Lower
Ordovician), Grebes Nest Point, Bell Island. Note prominent vertical spreite to the left of centre and narrow
vertical burrows (Phycodes wabanensis) in lower portions cutting through sandy beds. Top event bed shows
minor bioturbation. Scale bar is 20mm.

Tops of sands are nevertheless generally intact, of a firm substrate below a shallow mixed layer,
and storm-generated subtidal heterolithic bed- in contrast to the Lowermost Cambrian, where
ding generally is well preserved (Droser & Li evidence for a mixed layer is scant (Fig. 1). Our
2001, and see below). Where sedimentation rate hypothesis is illustrated in Fig. 1. A shallow
was low, the tops of sands may be reworked. mixed layer would be present in mud exposed
For example, the Mickwitzia Sandstone and at the surface. During storm events, we envisage
Liikati Formation have beds that are disturbed some erosion of the mixed layer followed by
by Diplo crater ion and Rhizocorallium (e.g. Opik deposition of sand. Below we discuss specific
1929; Jensen 1997). examples of ichnofabric.
All these units can and do have several An example of heterolithic ichnofabric: the
tiers, which result in intervals of complex ichno- Lower Ordovician Grebes Nest Point Formation.
fabric, with trace fossils exhibiting cross-cutting The Grebes Nest Point Formation of the
relationships. Wabana Group (Newfoundland) provides a
particularly instructive example of Cambro-
Ordovician ichnofabric. In a detailed study of
Properties of late Early Cambrian-earliest the ichnology of the Cambro-Ordovician Bell
Ordovician sediments Island and Wabana groups, Fillion & Pickerill
(1990) recognized that the muds were cohesive
Evidence for a near-surface firmground, such as in a variety of intertidal and shallow subtidal
the occurrence of the shallow-tiered trace fossils facies, based on sharp burrow outlines and
Treptichnus pedum and Gyrolithes and Kullin- preservation of fine surface sculpture. Our data
gia-type scratch circles, is rare above the Ruso- support these conclusions.
phycus avalonensis zone. All the characteristics Bases of beds often show a high diversity of
discussed above are consistent with the presence traces fossils (Fig. 2a). In addition to Rusophycus
EARLY PALAEOZOIC ICHNOFABRICS 389

Fig. 4. Polished vertical sections from the Grebes Nest Point Formation (Lower Ordovician), Grebes Nest
Point, Newfoundland. Scale bars are 10mm. (a) Sand cut by Phycodes wabanensis. (b) Rusophycus polonicus
in adhering preservation, (c) Sharply defined Cruziana isp. Note darker sediment in fill of burrow compared
with adjacent sand, suggesting an intrastratal formation for this particular specimen, (d) Lower portion of slab
has thin sands partly reworked by Planolites montanus; truncated and overlain by sandstone with small-scale
cross-lamination.

and Cruziana (discussed further below), Tricho- burrow margins (see also Pillion & Pickerill
phycus, Teichichnus and Phycodes are common. 1990). Phycodes burrows cross-cut thin mud
The different ichnospecies of Phycodes and Tei- and sand beds (Figs 3, 4a).
chichnus are probably shallow tiers (Pillion & Trichophycus occurs commonly in some inter-
Pickerill 1990). Phycodes wabanensis is generally vals. Preservation ranges from floating to
preserved with relatively distinctly defined adhered to cast. Burrows may also be 'washed
390 M. L. DROSER ET AL.

Fig. 5. Field photographs of the Lower Cambrian Arumbera sandstone, Northern Territories. Scale bar is
20 mm. (a) Abundant sand-filled burrows in mud. Member 3 at Daily Gorge, (b) Diverse preservation of
sandfilled burrows in mud. Member 3 at Shannon Bore.

out' where the type of preservation is dependent Trichophycus can be cross-cut within the mud-
upon the depth of erosion of overlying material. stone by Teichichnus, providing further evidence
Burrow margins are sharply defined, as are that Trichophycus was not deeply emplaced.
scratch marks. Trichophycus burrows preserved Teichichnus is a common and conspicuous
within the mud (attached to the overlying sand- Cambrian trace fossil (see Bland & Goldring
stone or not) act as tubular tempestites (Fig. 1995; Droser & Li 2001), and can also occur as
2b) (cf. Wanless et al 1988): that is, open bur- a tubular tempestite (Jensen 1997). Relatively
rows that are infilled with sediment that would sharply defined margins with bioglyphs suggest
otherwise bypass this palaeoenvironment. These that the sediment was moderately cohesive at
EARLY PALAEOZOIC ICHNOFABRICS 391

the time of burrow construction (Pillion & tion. Although individual burrows do not result in
Pickerill 1990). significant particle mixing, they would have
In the Grebes Nest Point as well as the Lower allowed the oxygenation of sediment to depths
Cambrian Mickwitzia sandstone and Liikati of several tens of centimetres. Especially under
Formation, mudstones may be bioturbated. conditions of slow sedimentation the fabric may
However, bedding down to the centimetre scale be dominated by these burrows (cf. Mcllroy
is generally preserved (Fig. 3). Thus thin tempes- 2004, fig 12).
tites are recognizable. Interestingly, in spite of Also within this high-energy setting may be
the fact that these mudstones apparently were found preservation of shallow tiers formed in
bioturbated, they still become cohesive at a mud. Examples include Rusophycus latus in the
relatively shallow depth, as indicated by the Pacoota Sandstone, Northern Territories, Aus-
prevalence of preserved shallow-tier burrows. tralia, and the Bynguano Formation of New
South Wales (Droser et al 1994). The preserved
details indicate that the mud must have been
The relevance of Skolithos piperock: the cohesive at the time of trace fossil emplacement.
Cambro-Ordovician Pacoota and Bynguano Both the Pacoota Sandstone and the Bynguano
formations Formation include successions of sandstones
with Skolithos burrows piping down through
By the latest Tommotian-Atdabanian an ichno- abundant, well-preserved Rusophycus occurring
fabric dominated by vertical trace fossils such as on the base of beds (Fig. 6). In some cases,
Skolithos and Diplocraterion (e.g. Westergard where the Skolithos do not penetrate the base
1931; Droser 1991; Mcllroy 2004) first appeared of the bed, it is possible to see original primary
in the stratigraphic record. These burrows bedding such as cross-stratification in the beds
extended to depths of decimetres and commonly casting the Rusophycus. Amalgamation and ero-
create a dense fabric often referred to as 'pipe sion surfaces are common. Mudstone is rarely
rock'. These are typical of high-energy sand-domi- preserved. Despite ubiquitous amalgamation,
nated shallow marine environments. There has there are numerous levels of Rusophycus (Fig.
been some debate as to the role of this fabric in 7), which indicate firmground conditions within
the context of the evolution of the early infauna the depth of trilobite burrowing. Primary sedi-
(e.g. Bottjer & Ausich 1985; Miller & Byers mentary structures appear to continue to the
1984). As pointed out by Thayer (1983) among base of the Rusophycus, and this is most consis-
others, these burrows most likely represent dwell- tent with a casting scenario of preservation
ing burrows and do not generate intense bioturba- (Fig. 8, mode k; see also Baldwin 1977).

Fig. 6. Field photograph (oblique lower view) of beds in the Pacoota sandstone, Ellery Creek, Northern
Territories. Note several basal surfaces covered with Rusophycus latus, and sides of sandstone beds with
Skolithos. Scale bar is 50 mm.
392 M. L. DROSER ET AL.

Fig. 7. Three types of preservation of sandstone and ichnocoenoses of the Bynguano Formation, New South
Wales (modified from Droser et al. 1994). (a) Amalgamated sandstones. No predepositional (e.g. Rusophycus)
preserved, (b) Amalgamated sandstones with some trace fossils preserved as casts on the base of sandstone
beds, (c) Sandstones interbedded with mudstones. Burrows are commonly cast on the base of sandstone beds
and appear to reflect open firmground burrows.

Fig. 8. Possible pathways for the formation of Rusophyus. (a) Muddy sediment with a shallow mixed layer,
beneath which is firmer sediment, (b, c) Intrastratal burrowing along the interface of sand and mud creates a
situation where the burrow is immediately cast. Subsequent erosion may lead to removal of sand except for the
burrow cast (f), followed by deposition of sand (i) or mud (h), resulting in styles of preservation that appear to
be particularly common in the Lower Palaeozoic, (d) Erosion of mud without deposition of sand leads to
exposed firm muds. Preservation of open mud burrows generally is not considered likely. The relatively stiff
nature of Lower Palaeozoic muds close to the sediment-water interface may, however, have made this rather
common, (k) A fill showing features of physical deposition is to be expected, (e) It can also be envisaged that
burrowing took place within mud and that firmer sediment was subsequently exposed and cast (g, k).
EARLY PALAEOZOIC ICHNOFABRICS 393

Preservation of Rusophycus and Cruziana may occur in the Cambrian, all depending on
the physical conditions.
Trilobite traces provide useful indicators of sedi- There are a number of implications of these
ment properties in part because they are rela- findings. Below we briefly list potential implica-
tively large and three-dimensional. It is thus tions of these data, ranging in order from the
relatively easy to determine the relationship of obvious to the conjectural.
the burrows to the encasing sediments in the
field. The quality of preservation of the scratch
marks can also be instructive. As discussed Increase in the depth and extent of the mixed
below, these also may provide insights into the
properties of sediments at this time. layer
It is generally assumed that the arthropod-type There is an increase in trace fossil size (Mcllroy &
trace fossils Rusophycus, and in particular Cruzi- Logan 1999) and diversity (Crimes 1992) from
ana, did not form at great depths below the sedi- the earliest Cambrian to the Early Ordovician.
ment-water interface and, in the case of traces In the earliest Cambrian the very shallowest tiers
such as Cruziana semiplicata, probably no more were commonly preserved. Shallow-tier burrows
than a few centimetres below the sediment- emplaced a few centimetres or even millimetres
water interface. Delicate preservation of leg were preserved through the Early Ordovician.
imprints suggests that the sediment encountered Data from Neoproterozoic-Ordovician strata
was relatively cohesive. Whether such features indicate that there was virtually no mixed layer
could be preserved as casts of originally open sur- in the Neoproterozoic and a minimal mixed
face furrows or must have originally formed layer in the earliest Cambrian. Atdabanian-
along a sand-mud interface has been a matter earliest Ordovician strata appear to have had a
of some contention (see Seilacher 1970; Crimes shallow mixed layer of the order of 5-1 Ocm.
1975; Baldwin 1977; Goldring 1985). Regardless Throughout this latter interval, bioturbation
of the precise mechanism of preservation it is remained of moderate depth in shelfal settings.
likely that a cover of less cohesive mud was This supports suggestions of a late origination of
washed out (cf. Miller & Rehmer 1982). The extensive mixing (e.g. Larson & Rhoads 1983).
conditions for the preservation of this type of
trace fossil may therefore have been particularly
favourable. The processes leading to the preser-
vation of Rusophycus and Cruziana are illu- Decreased preservation potential of event
strated in Fig. 8. All of these scenarios require beds
that sediments be firm at relatively shallow
depths, either so that trilobites burrow down to The results presented here further substantiate
the firmground through a mixed layer (Fig. 8e) the suggestion of Sepkoski et al. (1991) that
or so that it is shallow enough that erosion of there is a secular trend in the preservation of
the overlying mixed layer is commonplace (Fig. event beds. Results from this study suggest
8b, c, d). This suggests that the mixed layer was that, in particular, thin event beds will be best
relatively shallow, following the argument of preserved in the Neoproterozoic and earliest
Droser et al. (2002a, 2002b). Cambrian, but that Cambrian and earliest Ordo-
vician event beds also have high preservation
potential. This will, again, depend on the deposi-
Discussion tional environment and rates of sedimentation.

The characteristics described above are based on


examination of the 11 units listed in addition to Increased depth of firmground conditions
the Neoproterozoic-earliest Cambrian units from near surface in the Early Cambrian
studied for previous work (Droser et al. 2002b).
We present these characteristics as a model for As a consequence of the shallow mixed layer,
understanding early Palaeozoic substrate condi- muddy sediments were firm relatively close to
tions. We view these as generalizations to which the sediment-water interface. Evidence from low-
there will probably be exceptions. In particular, ermost Cambrian strata indicates the presence of
trace fossil preservation and substrate conditions firmground conditions at or near the sediment
will depend on the depositional environment and surface. All of the formations examined for this
rates of sedimentation. Indeed, many of these paper have extensive evidence of firmground
features may be present in any strata of Phaner- conditions, of the order of 5-1 Ocm below the
ozoic age, and Phanerozoic-style preservation sediment-water interface. Future work will
394 M. L. DROSER ET AL.

determine the subsequent nature of the develop- 1970). Trilobites do decline in diversity through
ment of the mixed layer and thus the deepening this interval (Adrain et al. 1998). However, a
of firmground conditions within the sediment. deepening of the mixed layer through the Palaeo-
Related to this, Bottjer et al. (2000) have sug- zoic would result in the deterioration in the
gested that many Early Cambrian metazoans preservation potential of this type of trace fossil.
were well suited for matground conditions. Pre- Clearly, a number of other factors must also be
sumably, they would have also been ideally considered and tested, but changes in the preser-
suited for firmground conditions. vation potential of these trace fossils may be
important.
Decreased trace fossil preservation potential We acknowledge partial funding for the fieldwork from
of shallow-tier burrows National Science Foundation (grant EAR-9219731 to
MLD) and the National Geographic Society. This
Cohesive sediment close to the sediment-water paper benefited from conversations with N. Hughes,
interface may explain the rich Lower Palaeozoic P. Myrow, M. Kennedy and G. Narbonne. V.
Droser, R. Droser, A. Dzaugis and M. Dzaugis
record of well-preserved marine trace fossils. The provided field assistance. D. A. Droser facilitated field-
common occurrence of trace fossils preserved as work. This paper also benefited greatly from reviews
floating or adhering trace fossils (Fig. 8) is prob- by R. Goldring and J. Hagadorn. We are also grateful
ably a result of firm muddy sediments. Casting of to D. Mcllroy for organizing the symposium and
open or washed-out burrows may have been par- providing a very helpful review.
ticularly likely at this time. The greatest potential
for the preservation of marine trace fossils is in
Precambrian and earliest Cambrian sediments.
In earliest Cambrian strata, very shallow-tiered References
millimetre-scale Gyrolithes and Treptichnus ADRAIN, J. M., FORTEY, R. A. & WESTROP, S. R. 1998.
pedum occur in abundance. Although they are Post-Cambrian trilobite diversity and evolution-
present in younger strata and sometimes in ary faunas. Science, 280, 1922-1925.
some abundance, they are never as ubiquitous. ALLER, R. C. 1983. The effects of macrobenthos on
The presence of graphoglyptid trace fossils such chemical properties of marine sediments and over-
as Palaeodictyon in Cambrian shallow-water lying water. In: McCALL, P. L. & TEVESZ, J. S.
sediments could be a further indication of (eds) Animal-Sediment Relations: the Biogenic
preservation of shallow-tiered trace fossils in Alteration of Sediments. Plenum Press, New
York, 53-102.
this setting (Crimes & Fedonkin 1994). AUSICH, W. I. & BOTTJER, D. J. 1982. Tiering in sus-
pension-feeding communities on soft substrata
throughout the Phanerozoic. Science, 216, 173—
Nature of Lower Palaeozoic trace fossil 174
record BALDWIN, C. T. 1977. Internal structure of trilobite
trace fossils indicative of an open surface furrow
Most of the discrete trace fossils preserved in origin. Palaeogeography, Palaeoclimatology,
these heterolithic settings, with the exception of Palaeoecology, 21, 273-284.
those piping into the sand, are firmground trace BERGER, W. H., EKDALE, A. A. & BRYANT, P. P. 1979.
fossils. These consist largely of open burrows in Selective preservation of burrows in deep-sea
carbonates. Marine Geology, 32, 205-230.
mud. The evidence for very earliest Cambrian BLAND, B. H. & GOLDRING, R. 1995. Teichichnus Seila-
mobile deposit feeders in muds is low (Droser cher 1955 and other trace fossils (Cambrian?) from
et al. 2002b). In the earliest Cambrian virtually the Charnian of Central England. Neues Jahrbuch
all of the trace fossils were emplaced in firm- fur Geologie und Palaontologie, Abhandlungen,
grounds. As the mixed layer further developed, 195, 5-23.
and thus firmground conditions receded into BOCKELIE, F. J. 1991. Ichnofabric mapping and
the sediment, softground trace fossils would interpretation of Jurassic reservoir rocks of the
have become more common and ultimately Norwegian North Sea. Palaios, 6, 206-215.
would have dominated the ichnofauna. BOTTJER, D. J. & AUSICH, W. I. 1985. Abundant and
diverse early Paleozoic infauna indicated by
the stratigraphic record. Comment. Geology, 13,
83-84.
Decreased preservation of Rusophycus and BOTTJER, D. J., HAGADORN, J. W. & DORNBOS, S. Q.
Cruziana 2000. The Cambrian substrate revolution. GSA
Today, 10, 1-7.
Rusophycus and Cruziana decline in occurrence BRASIER, M. 1990. Nutrients in the early Cambrian.
and abundance through the Palaeozoic (Seilacher Nature, 347, 521-522.
EARLY PALAEOZOIC ICHNOFABRICS 395

BRASIER, M. & MC!LROY, D. 1998. Neonereites uniser- GOLDRING, R. 1985. The formation of the trace fossil
ialis from c. 600 Ma year old rocks in western Cruziana. Geological Magazine, 122, 65—72.
Scotland and the emergence of animals. Journal GOLDRING, R. 1995. Organisms and the substrate:
of the Geological Society, London, 155, 5-12. response and effect. In: BOSCENCE, D. W. J. &
BRASIER, M., COWIE, J. & TAYLOR, M. 1994. Decision on ALLISON, P. A. (eds) Marine Palaeoenvironmental
the Precambrian-Cambrian stratotype. Episodes, Analysis from Fossils. Geological Society,
17, 3-8. London, Special Publications, 83, 151-180.
BROMLEY, R. G. 1996. Trace Fossils: Biology, Taphon- GOLDRING, R. & JENSEN, S. 1996. Trace fossils and bio-
omy and Applications. Chapman & Hall, London. fabrics at the Precambrian-Cambrian boundary
CRIMES, T. P. 1975. The production and preservation of interval in western Mongolia. Geological Maga-
trilobite resting and furrowing traces. Lethaia, 8, zine, 133,403-415.
35-48. HAGADORN, J. W. & BOTTJER, D. J. 1997, Wrinkle
CRIMES, T. P. 1987. Trace fossils and correlation of structures: microbially mediated sedimentary
late Precambrian and early Cambrian strata. structures common in subtidal siliciclastic settings
Geological Magazine, 124, 97-119. at the Proterozoic-Phanerozoic transition. Geol-
CRIMES, T. P. 1992. Changes in the trace fossil biota ogy, 25, 1047-1050.
across the Proterozoic-Phanerozoic boundary. HALLAM, A. 1975. Preservation of trace fossils. In:
Journal of the Geological Society, London, 149, FREY, R. W. (ed.) The Study of Trace Fossils.
637-646. Springer, New York, 289-334.
CRIMES, T. P.& FEDONKIN, M. A. 1994. Evolution and JENSEN, S. 1997. Trace fossils from the Lower Cambrian
dispersal of deepsea traces. Palaios, 9, 74-83. Mickwitzia sandstone, south-central Sweden.
DROSER, M. L. 1991. Ichnofabric of the Paleozoic Sko- Fossils and Strata, 42, 1-110.
lithos ichnofacies and the nature and distribution JENSEN, S. & MENS, K. 1999. A Lower Cambrian
of Skolithos piperock. Palaios, 6, 316-325. shallow-water occurrence of the branching deep-
DROSER, M. L. & XING Li 2001. The Cambrian water type trace fossil Dendrorhaphe from the
radiation and the diversification of sedimentary Lontova Formation, eastern Latvia. Palaontolo-
fabrics. In: ZHURAVLEV, A. Yu. & RIDING, R. gische Zeitschrift, 73, 187-193.
(eds) The Ecology of the Cambrian Radiation. JENSEN, S., GEHLING, J. G., DROSER, M. L. & GRANT,
Columbia University Press, New York, 137-169. S. W. F. 2002. A scratch circle origin for the medu-
DROSER, M. L., HUGHES, N. L. & JELL, P. A. 1994. soid fossil Kullingia. Lethaia, 35, 291-299.
Infaunal communities and tiering in Early Palaeo- LARSON, D. W. & RHOADS, D. C. 1983. The evolution
zoic nearshore clastic environments: trace-fossil of infaunal communities and sedimentary fabrics.
evidence from Cambro-Ordovician of New South In: TEVESZ, J. S. & McCALL, P. L. (eds) Biotic
Wales. Lethaia, 27, 273-283. Interactions in Recent and Fossil Benthic Commu-
DROSER, M. L., JENSEN, S. & GEHLING, J. G. 2002a. nities. Plenum Press, New York, 627-648.
Trace fossils and substrates of the terminal Proter- MclLROY, D. 2004. Some ichnological concepts,
ozoic—Cambrian transition: implications for the methodologies, applications and frontiers. In:
record of early bilaterians and sediment mixing. MclLROY, D. (ed.) The Application of Ichnology
Proceedings of the National Academy of Sciences to Palaeoenvironmental and Stratigraphic Analysis.
of the United States of America, 99, 12572-12576. Geological Society, London, Special Publications,
DROSER, M. L., JENSEN, S., GEHLING, J. G., MYROW, P. 228, 3-27.
& NARBONNE, G. M. 2002b. Lowermost Cambrian MclLROY, D. & LOGAN, G. A. 1999. The impact of
Ichnofabrics from the Chapel Island Formation, bioturbation on infaunal ecology and evolution
Newfoundland: implications for Cambrian sub- during the Proterozoic-Cambrian transition.
strates. Palaios, 17, 3-15. Palaios, 14, 58-72.
ERIKSSON, P. G., CONDIE, K. C. et al. 1998. Pre- MILLER, M. F. & BYERS, C. W. 1984. Abundant and
cambrian (pre-vegetational) clastic sedimentation diverse early Paleozoic infauna indicated by the
systems. Sedimentary Geology, 120, 5—53. Stratigraphic record. Geology, 12, 40-^43.
FEDONKIN, M. A. 1983. Besskeletnaya fauna podols- MILLER, M. F. & REHMER, J. 1982. Using biogenic struc-
kogo pridnestrovya. In: VELIKANOV, V. A., tures to interpret lithologic boundaries: an example
ASEEVA, M. A. & FEDONKIN, M. A. (eds) Vend from the Lower Devonian of New York. Journal of
Ukrainy. Naukova Dumka, Kiev, 129-139 Sedimentary Petrology, 52, 887-895.
PILLION, D. & PICKERILL, R. K. 1990. Ichnology of NARBONNE, G. M., MYROW, P., LANDING, E. & ANDER-
the Upper Cambrian? to Lower Ordovician Bell SON, M. A. 1987. A candidate stratotype for the
Island and Wabana groups of eastern Newfound- Precambrian-Cambrian boundary, Fortune Head,
land, Canada. Palaeontographica Canadiana, 1, Burin Peninsula, southeastern Newfoundland.
1-119. Canadian Journal of Earth Sciences, 24, 1277-1293.
GEHLING, J. G. 1999. Microbial mats in terminal OPIK, A. 1929. Eine Corophioides-f&ima aus dem est-
Proterozoic siliciclastics: Ediacaran death masks. nischen unterkambrischen 'fossilleeren' Sandstein.
Palaios, 14, 40-57. Ada et Commentationes Universitatis Tartuensis
DE GIBERT, J. M. & EKDALE, A. A. 1999. Trace fossil A, 15:2, 30^2.
assemblages reflecting stressed environments in PIPER, D. J. W. & MARSHALL, N. F. 1969. Bioturbation
the Middle Jurassic Carmel Seaway of central of Holocene sediments of La Jolla deep sea fan.
Utah. Journal of Paleontology, 73, 711-720. Journal of Sedimentary Petrology, 39, 601-606.
396 M. L. DROSER ET AL.

SAVRDA, C. E. & BOTTJER, D. J. 1989. Anatomy and the biological overprint. In: EINSELE, G., RICKEN,
implications of bioturbated beds in Black Shale W. & SEILACHER, A. (eds) Cycles and Events in
sequences: examples from the Jurassic Posido- Stratigraphy. Springer, Berlin, 298-312.
nienschiefer (Southern Germany). Palaios, 4, SIMPSON, S. 1957. On the trace-fossil Chondrites. Quar-
330-342. terly Journal of the Geological Society of London,
SEILACHER, A. 1970. Cruziana stratigraphy of 'non- 112, 475-500.
fossiliferous' Palaeozoic sandstone. In: CRIMES, THAYER, C. W. 1983. Sediment-mediated biological
T. P. & HARPER, J. C. (eds) Trace Fossils. Seel disturbance and the evolution of marine benthos.
House Press, Liverpool, 447-476. In: TEVESZ, M. J. S. & McCALL, P. L. (eds)
SEILACHER, A. 1985. Trilobite palaeobiology and sub- Biotic Interactions in Recent and Fossil Benthic
strate relationship. Transactions of the Royal Communities. Plenum, New York, 479-625.
Society of Edinburgh, 76, 231-237. WANLESS, H. R., TEDESCO, L. R. & TYRRELL, K. M.
SEILACHER, A. & PFLUGER, F. 1994, From biomats to 1988. Production of subtidal tubular and surficial
agricultural revolution: In: KRUMBEIN, W. E., tabular tempestites by Hurricane Kate, Caicos
PATERSON, D. M. & STAL, L. J. (eds) Biostabiliza- Platform. Journal of Sedimentary Petrology, 58,
tion of Sediments. Bibliotheks und Informations- 73-75.
system der Carl von Ossietzky Universitat, WESTERGARD, A. H. 1931. Diplocraterion, Monocrater-
Oldenburg, 97-105. ion and Scolithus from the Lower Cambrian of
SEPKOSKI, J. J., BAMBACH, R. K. & DROSER, M. L. 1991. Sweden. Sveriges Geologiska Under sokning,
Secular changes in Phanerozoic event bedding and C372, 1-25.
Trace fossils in the aftermath of mass extinction events

RICHARD J. TWITCHETT1 & COLIN G. BARRAS2 3


1
School of Earth, Ocean and Environmental Sciences, University of Plymouth,
Drake Circus, Plymouth PL4 8AA, UK
(e-mail: rtwitchett@plymouth.ac.uk)
2Department of Palaeontology, The Natural History Museum, Cromwell Road,
London SW7 5BD, UK
^Department of Earth Sciences, School of Geography, Earth and Environmental Sciences,
University of Birmingham, Birmingham B15 2TT, UK

Abstract: Ichnology has great potential to advance our understanding of mass extinction
events and yet is currently an underutilized resource in such studies. Here we review
published ichnological studies for the Ordovician-Silurian, Permian-Triassic and
Cretaceous-Tertiary extinction-recovery intervals. In addition, new information regarding
the Triassic-Jurassic ichnological record from England, Austria and the western USA is
presented. Trace fossils provide important information on the ecological response of the
benthic community at such times. In the immediate post-extinction aftermath, the ichno-
diversity, burrow size, depth of bioturbation, and ichnofabric index of the sediments are
all much reduced. There is an increase in all these parameters through the post-extinction
recovery period. In some cases, the stepwise reappearance of certain distinctive ichnotaxa
(e.g. Diplocraterion, Rhizocorallium and Thalassinoides) may be of some stratigraphic use.
Evidence from Permian-Triassic studies indicates that recovery took longer at low (tropical)
palaeolatitudes than mid-high palaeolatitudes. Trace fossils also provide important
information on palaeoenvironmental change through the extinction-recovery interval. The
application of ichnology to mass extinction studies is in its infancy, but should prove a
valuable tool in future research.

Understanding mass extinction events is crucial Application of ichnology to mass extinction


to understanding the evolutionary history of studies
life on Earth. One tool that is currently under-
utilized in such studies, but which has much to Palaeoenvironmental analysis
offer, is ichnology. Our objective is to demon-
strate that studying trace fossils can enhance The use of trace fossils for palaeoenvironmental
our understanding of mass extinction episodes analysis is commonplace, and can be applied to
and, in particular, of the post-extinction recovery all parts of the geological column, including
period. We provide a review of previous studies extinction-recovery intervals. Ichnofacies analy-
in this area, including new data from the sis, based on Seilacher's (1967) concepts, allows
Permian-Triassic and Triassic-Jurassic events, environmental parameters such as substrate
and hope to stimulate others to use the trace consistency, bathymetry and hydrodynamic
fossil record in future analyses of extinction- energy to be inferred from the occurrence of
recovery episodes. particular suites of trace fossils (see Bromley
The focus of attention is directed towards 1990 for a recent discussion).
the marine trace fossil record. However, terres- The amount of bioturbation present in a
trial ichnofaunas also have much unexplored particular sedimentary unit also has palaeo-
potential. One recent example is Olsen et al.'s environmental significance. Sediments that are
(2002) study of the effects of possible extra- thoroughly bioturbated were obviously depos-
terrestrial impact events on the evolution of ited under conditions that were amenable to
the Triassic-Jurassic terrestrial vertebrate both benthic colonization (adequate oxygen,
faunas of North America, which relied entirely food supply etc.) and trace fossil preservation.
on trackway evidence left by dinosaurs and Sediments lacking bioturbation were deposited
other tetrapods. To date, this represents the under conditions that prevented colonization
only extinction-related investigation that has (e.g. anoxia, high sedimentation rates) and/or
primarily involved terrestrial trace fossil were unsuitable for preserving trace fossils.
evidence. The observed ichnofabric is related to the
From: MC!LROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 397^18. 0305-8719/04/S15.00 © The Geological Society
of London.
398 R. J. TWITCHETT & C. G. BARRAS

interplay between the levels of colonization and maker is not usually possible, and ichnotaxa
preservation. The amount of bioturbation in are only of limited use in assessing benthic
vertical section can be estimated semi-quan- diversity. However, certain trace fossils can be
titatively by using Droser & Bottjer's (1986) attributed to certain classes of organism. For
ichnofabric index (ii); for bedding plane example, Thalassinoides and Ophiomorpha are
exposures, Miller & Smail's (1997) scheme can attributed to anomuran crustaceans, based on
be applied. comparisons with modern burrows (e.g. Bromley
One area of palaeoenvironmental analysis 1990). Resting traces (cubichnia) are often
that is of particular importance to extinction particularly easy to assign as the shape of the
studies is the reconstruction of benthic oxygen trace fossil reflects the morphology of the trace-
levels. In their recent review of Phanerozoic maker (e.g. Lockeia traces are attributed to
extinction events, Hallam & Wignall (1997) con- bivalves, Asteriacites lumbricalis to ophiuroids
cluded that most were associated with episodes etc.). Such assignments allow the presence or
of oceanic anoxia. Ichnology is particularly absence of certain groups to be documented,
useful in the assessment of ancient benthic which may enable comparison with modern
oxygen levels. As oxygen levels decline, the benthic communities. For example, crustaceans
amount and depth of bioturbation decreases, as are usually the last group to reappear after
do ichnodiversity and burrow diameter (e.g. modern, low-latitude hypoxic events (e.g.
Rhoads & Morse 1971; Savrda & Bottjer 1986). Harper et al. 1991). Similarly, in the low palaeo-
Dysaerobic sediments tend to be poorly bio- latitude sediments of northern Italy, Thalassi-
turbated by a single ichnotaxon, such as Chon- noides is one of the last ichnotaxa to reappear
drites, which typically occupies the deepest tier after the Permian-Triassic anoxic event (it is
in normal, aerobic environments (Bromley & absent until the Anisian).
Ekdale 1984; Savrda & Bottjer 1987). Sediments Another important ecological aspect of the
deposited under continuously anoxic conditions trace-making organisms that can be assessed
lack bioturbation. through study of their trace fossils is that of
size. Body size is a key element in animal evolu-
tion (Jablonski 1996). It is a fundamental charac-
Palaeoecological studies ter of living organisms, with implications for
many aspects of an animal's biology, behaviour
Trace fossils are records of the activities of and ecology. A growing number of studies have
organisms, and therefore also provide biological shown that the aftermaths of mass extinction
and ecological information. In the marine realm, events are characterized by fossil animals of
most ichnotaxa are the products of the actions of unusually small size. Termed the 'Lilliput effect'
soft-bodied, or lightly mineralized, invertebrates, by Urbanek (1993), this phenomenon affects a
which are rarely (if ever) fossilized. Yet, in range of animal groups and all extinction
modern marine ecosystems, and presumably in events studied to date (e.g. Girard & Renaud
ancient ones too, soft-bodied organisms com- 1996; Twitchett 2001). Burrow diameter is a
prise the vast bulk of the benthic biota (Allison good proxy for size of the trace-making organ-
& Briggs 1991, p. 26). Usually our only glimpse isms, and certainly a burrowing animal cannot
of these taxa in the fossil record is through the have a diameter larger than its burrow, so the Lil-
record of Konservat-Lagerstatten (e.g. the Bur- liput effect may be expected in post-extinction
gess Shale). However, no known Konservat- ichnogenera too. Indeed, results from northern
Lagerstatte spans an extinction episode, and no Italy show that there is an order of magnitude
extinction episode has Konservat-Lagerstatten decrease in burrow diameter across the Per-
in the immediate pre-extinction and post- mian-Triassic extinction event (Twitchett 1999).
extinction intervals. For example, in order to In shelly taxa, the Lilliput effect is a temporary
attempt an assessment of how the Permian phenomenon, affecting the immediate post-
extinction events affected soft-bodied marginal extinction aftermath, and may be related to a
marine taxa, Briggs and Gall (1990) were decrease in food supply (Twitchett 2001; Price-
forced to compare the Anisian Ores a Voltzia Lloyd & Twitchett 2002) or some other tempor-
lagerstatte with several Carboniferous ones. ary environmental disturbance such as low
Thus trace fossils are the only record of the oxygen levels, temperature change or salinity
direct response of the soft-bodied benthic inver- fluctuation (Hallam 1965). It is followed by a
tebrate community to mass extinction events. subsequent increase in body size in the later
Do changes in ichnodiversity reflect real stages of post-extinction recovery, presumably
changes in benthic diversity? Trying to match a as conditions improve. Likewise, there is an
particular trace fossil to a particular trace- increase in burrow size through the later recovery
ICHNOSTRATIGRAPHY & MASS EXTINCTION 399

intervals of the Early Triassic (Twitchett 1999)


and Early Jurassic (discussed more fully below).
No burrow size data have been published for
any other extinction episode.
Trace fossils are primarily records of beha-
viour, and hence provide information on the
ecological response of the benthic community
to extinction events. For example, they can be
used to identify opportunistic behaviour (e.g.
Ekdale 1985; Vossler & Pemberton 1988),
which may be one of the life strategies likely to Fig. 1. Maximum levels of tiering of suspension-
characterize the surviving communities of the feeding communities in marine environments during
immediate post-extinction interval (Harries et al. the Phanerozoic. Vertical axis shows distance from
1996). Changes in feeding strategy are also sediment-water interface in centimetres. PC,
likely to occur during extinction intervals, for Precambrian; Mz, Mesozoic; Cz, Cenozoic. Arrows
example in response to food chain collapse. mark positions of the five major mass extinction
During the end-Cretaceous event, plankton events of the Phanerozoic. Redrawn from Ausich &
productivity collapse appears to have led to the Bottjer (2001).
preferential extinction of filter feeders and the
preferential survival of detritivores (Sheehan zoic extinction events have yet to be investigated,
et al. 1996). Under such conditions, it might be and doubtless the picture will change further as
predicted that trace fossil assemblages in the our knowledge improves.
immediate aftermath of such an event would be
dominated by the traces of deposit feeders,
and that the dwelling burrows of infaunal sus- Ichnos tra tigraphy
pension feeders would not return until suitable
levels of primary production were re-established. In certain situations trace fossils have a strati-
Finally, trace fossils are critical in studies of graphic use, and have proved to be a valuable
the tiering of infaunal communities. Different tool in correlation, especially during the Pre-
taxa are adapted to live at different depths (i.e. cambrian-Cambrian transition (e.g. Crimes
within different tiers) below the seafloor, and 1992) and the lower Palaeozoic (e.g. Seilacher
the term 'tiering' describes the vertical distri- et al. 2002). Crimes (1987) gave a comprehensive
bution of the infaunal community that results. account of the distribution of ichnotaxa through
In their major study of Phanerozoic tiering of a number of Precambrian-Cambrian boundary
suspension-feeding communities in shelf sea sections, showing that there is a stepwise
settings (below wavebase), Bottjer & Ausich appearance of ichnogenera through the earliest
(1986) showed that during the early Palaeozoic Palaeozoic, and that the order of appearance is
only the shallowest tiers (down to 12cm below similar worldwide. The stepwise appearance is
the seafloor) were occupied. During the Carboni- to be expected as different trace-making groups
ferous, organisms began to exploit the deeper originate and/or novel behavioural and burrow-
tiers (down to 1 m below the seafloor), and ing strategies evolve. The first appearance
infaunal tiering remained at these depths through datum of the ichnotaxon Treptichnus (formerly
to the present. Phycodes) pedum actually defines the base of
The results of this initial study suggested that the Cambrian (Landing 1994). The value of
the level of infaunal tiering was unaffected by these earliest Cambrian traces is that they are
mass extinction intervals, implying greater present in rocks lacking abundant shelly remains,
stability of infaunal communities, compared and hence are the only potentially useful strati-
with epifaunal ones, in the face of environmental graphic tools available.
change (Bottjer & Ausich 1986). However, recent Similar conditions may also apply in the
research (e.g. Twitchett 1999) has shown that the aftermath of mass extinction events. There is
depth of infaunal tiering can be severely reduced certainly a stepwise reappearance of ichnotaxa
during extinction episodes. This has been after some of the major extinction episodes,
demonstrated for the Permian-Triassic extinc- which is unrelated to facies change and is com-
tion event, and is reflected in Ausich & Bottjer's parable in different localities. One example is in
(2001) Phanerozoic tiering diagram (Fig. 1). the aftermath of the end-Permian event (Twitch-
However, with the exception of the Triassic- ett 1997), which is discussed further below. In
Jurassic event described below, the infaunal addition, post-extinction strata often contain a
tiering histories of the remaining major Phanero- depauperate and poorly preserved shelly fauna,
400 R. J. TWITCHETT & C. G. BARRAS

which can make traditional biostratigraphy Palaeophycus, Planolites and Protopaleodictyon —


difficult. In the immediate extinction aftermath whereas the sediments of the lowest Silurian Allt
there may be a 'dead zone' (Harries & Kauffman Goch Formation preserve just four: Chondrites,
1990), where body fossil taxa are completely Helminthopsis, Palaeophycus and Planolites.
absent, resulting in significant gaps in the Post-extinction ichnodiversity is 50% of the pre-
ranges of biostratigraphically useful groups extinction levels, and, according to McCann
such as ammonoids or conodonts (e.g. Looy (1990), the ichnotaxa that remain are the most
et al 2001; Twitchett et al 2001). In such cases common members of the pre-event ichnofauna.
ichnostratigraphy may be one solution to the Time is required for ichnodiversity to recover:
problems of correlation. the missing ichnotaxa do not reappear until the
Aberystwyth Grits Formation (Upper Llandov-
ery).
Ordovician-Silurian (O-S) event Are these ichnodiversity changes the result of
real diversity changes in the benthic infauna, or
The Late Ordovician (Ashgill) interval witnessed are they simply due to changing sea-level?
extinction of some 24% of marine families Although the drop in ichnodiversity across the
(Benton 1995) and is unique among Phanerozoic O-S boundary also corresponds with a transgres-
extinction events in being intimately linked with sion, it is difficult to determine actual depths
a short (less than 1 Ma) episode of glaciation from the sediments (McCann 1990). The trace
(e.g. Brenchley et al 1994). The patterns of fossil record is interpreted as being, at least
extinction and recovery are diachronous in a partly, the result of real biotic change in trace-
number of marine groups, and were clearly maker communities. Ecological changes, such
driven by changes in climate, oceanographic as organism size and depth of burrowing, may
state and sea-level (e.g. Armstrong 1996; also have occurred during the O-S interval, but
Finney et al 1999). Thus far, MeCann's (1990) have yet to be studied.
study of the deep-sea ichnofaunal record of the McCann (1990) also showed that the ichno-
Llandeilo to Upper Llandovery formations of diversity of deep-sea environments has changed
the Welsh Basin is the only detailed study of over the Phanerozoic. There are broad similari-
trace fossils through the Ordovician-Silurian ties with curves of marine family diversity
mass extinction event and recovery (Fig. 2). based on shelly fossils (e.g. Sepkoski 1984;
The data of McCann (1990) indicate an overall Benton 1995): an initial Cambrian-Ordovician
increase in ichnodiversity from the Llandeilo increase, low diversity through the latest Palaeo-
through into the Llandovery. This is probably, zoic and early Mesozoic, and a dramatic increase
in part, a local expression of the global increase through the later Mesozoic and Tertiary.
in the number of taxa colonizing deep-sea envir- Decreases in ichnogeneric diversity of deep-sea
onments through the lower Palaeozoic (Orr environments occur across the O-S, P-Tr and
2001) and in part reflects changes in trace fossil K-T mass extinction intervals, as well as through
preservation through the different formations the Carboniferous-Permian interval (McCann
(McCann 1990). However, superimposed on 1990).
this overall increase in ichnodiversity is a short-
term decrease associated with the O-S boundary.
The decrease in ichnodiversity from the Late Devonian event
uppermost Ordovician Llangranog Formation to
the basal Silurian Gaerglwyd Formation can be The Late Devonian event is not considered
simply explained by the facies change from further in this present work because there are
sandstone-dominated to mudstone-dominated no detailed studies of the trace fossil record of
facies (Fig. 2). Indeed, this facies control on this event, although the presence of bioturbation
ichnodiversity is recorded throughout the succes- is sometimes mentioned in sedimentological
sion (McCann 1990). However, when only the studies (e.g. Chen & Tucker 2003).
sandstone-dominated formations are compared
(in order to reduce the problems of facies control)
a decrease in ichnodiversity is still recorded across Permian-Triassic (P-Tr) event
the O-S boundary, with the earliest Silurian Allt
Goch and Grogal Formations 'having lower abun- The P-Tr mass extinction event was the most
dances than might be predicted' (McCann 1990). severe of the Phanerozoic, with some 48% of
The uppermost Ordovician Llangranog For- marine families, and possibly in excess of 95%
mation contains eight ichnotaxa - Chondrites, of species, becoming extinct (e.g. Erwin 1993;
Circulichnus, Cochlichnus, Gordia, Helminthopsis, Hallam & Wignall 1997). Study of the P-Tr
ICHNOSTRATIGRAPHY & MASS EXTINCTION 401

Fig. 2. Stratigraphy, ichnotaxa ranges and sea-level curve through the Ordovician-Silurian succession of the
Welsh Basin. Arrow indicates direction of sea-level rise. Solid line indicates presence in formation; dashed line
indicates absence. From data in McCann (1990).

event has undergone something of a renaissance Pre-event ichnofauna


since the late 1980s and through the 1990s
(Benton 2003). A significant part of this recent Typically, uppermost Permian (Changhsingian)
effort has included detailed ichnological ana- sediments are well-bioturbated, with a diverse
lyses. One of the triggers that led to this renewal trace fossil assemblage that may include Diplo-
of interest was Hallam's (1989) suggestion that craterion, Palaeophycus, Planolites, Rhizocoral-
the P-Tr event was caused by an episode of lium, Skolithos, Thalassinoides and Zoophycos
oceanic anoxia. A number of field studies soon (Table 1). This diverse assemblage disappears in
followed (e.g. Wignall & Hallam 1992) and, the very latest Changhsingian or earliest Triassic
because of their utility in assessing benthic (Griesbachian) with the onset of benthic oxygen
oxygen levels, trace fossils often featured in restriction, which is diachronous on a global
these investigations. scale (Wignall £ Twitchett 2002a).
402 R. J. TWITCHETT & C. G. BARRAS

Table 1. Latest Permian (Changhsingian) ichnofaunas documented to date

Locality Formation Ichnotaxa Reference

Northern Italy Bellerophon Fm. Diplo crater ion, Planolites, Rhizocorallium, Twitchett (1999)
Skolithos, Zoophycos
Spitsbergen Kapp Starostin Fm. Chondrites, Diplo crater ion, Planolites, Wignall et al. (1998)
'Zoophycos
East Greenland Schuchert Dal Fm. Palaeophycus, Planolites, Rhizocorallium Twitchett et al. (2001)
Sichuan, China Dalong Fm. Thalassinoides , Zoophycos Wignall etal (1995)
Northern Oman Maqam Fm. Chondrites, Palaeophycus, Rhizocorallium, pers. observ.
Skolithos, Thalassinoides

Disappearance of the pre-event ichnofauna the only ichnotaxon present in the immediate
usually occurs over several decimetres or metres post-extinction environments of the earliest
of strata. In southern Austria, ichnofabric index Griesbachian is Planolites, which represents the
declines from ii5-6 to iil-2 over some 15m of fodinichnia of deposit-feeding vermiform organ-
strata (Twitchett & Wignall 1996). In East Green- isms (Twitchett & Wignall 1996; Twitchett 1999).
land similar changes occur over just 50cm of In contrast, the pre-extinction trace fossil assem-
strata, which probably represent only a few tens blage was a diverse mixture of domichnia and
of thousands of years (Twitchett et al. 2001). Dis- fodinichnia (Fig. 3; Table 1), indicating the
appearance of the pre-event ichnofauna is inter- presence of both suspension-feeding and
preted as reflecting the collapse of the latest deposit-feeding animals, including crustaceans.
Permian benthic ecosystem, which, in East Green- The temporary disappearance of suspension-
land at least, occurs at the same time as collapse offeeding organisms could indicate a collapse in
the terrestrial plant communities (Looy et al. primary production and inadequate food
2001; Twitchett et al. 2001). As well as a reduction supply (cf. Sheehan et al. 1996).
in the amount of bioturbation and ichnodiversity, However, the disappearance of suspension-
an accompanying decrease in burrow diameter is feeding domichnia could also be related to low
also recorded (e.g. Wignall et al. 1995; Wignall oxygen conditions. In a study of the infauna
& Hallam 1996; Twitchett 1999). of the Cretaceous Greenhorn Formation, Sage-
man & Bina (1997) showed that domichnia
were confined to well-oxygenated intervals, and
Immediate post-extinction aftermath as oxygen levels decreased, domichnia dis-
appeared until only the fodinichnia of deposit
Earliest Griesbachian ichnofaunas are typically of feeders were left. Small size could also be the
low diversity and small size (1-5 mm), with shal- result of either oxygen stress or inadequate
low penetrating (< 1 cm) bioturbation confined food supply (or both). Modern low-energy, oli-
to a few discrete horizons separated by metres of gotrophic environments are characterized by a
laminated, unbioturbated sediments. This pattern small, sparse, deposit-feeding infauna (Jumars
indicates that environmental stress (of some sort) & Wheatcroft 1989). A size decrease is also
prevented benthic colonization; only when condi- observed in the shelly macrofauna through
tions improve slightly can a pioneering benthic the Permian-Triassic interval, which has been
community colonize the substrate. This is typical related to food shortage (Price-Lloyd & Twitch-
of dysoxic environments, although Erwin (1993, ett 2002). Reduction in the depth of burrowing
p. 246) has argued that the absence of bioturba- is probably more likely to be caused by low
tion at this time is the result of the extinction of oxygen conditions, although food supply may
the benthos, rather than low oxygen conditions, also have an effect (Jumars & Wheatcroft
and that benthic conditions were otherwise 1989). Other evidence of low productivity
normal. However, this argument does not explain levels can be found, such as a negative shift
the observed changes in burrow size and depth of in <S13C and the low organic carbon content of
burrowing, and, in addition, there is plenty of the sediments (e.g. Twitchett 2001). Thus the
independent geochemical evidence for low immediate post-extinction trace fossil record
oxygen conditions at this time (Wignall & Twitch- may be the result of more than one type
ett 1996, 2002a). of environmental stress: inadequate food
Are there any other possible causes of environ- supply and low oxygen conditions are both
mental stress at this time? In northern Italy, contenders.
ICHNOSTRATIGRAPHY & MASS EXTINCTION 403

Early Trlassie recovery ichnofauna of small size (Twitchett 1999). Ichno-


diversity remains fairly high, however, and the
Despite the increasing number of studies that reddish micaceous fine sandstones and siltstones
have documented ichnofaunal change through were clearly deposited in a shallow, well-
the P-Tr boundary, there are still relatively few oxygenated environment. Monotaxic assem-
studies of trace fossil response and recovery blages of Asteriacites lumbricalis (produced by
through the later Early to Middle Triassic. ophiuroids) are common within this unit. The
Only the shallow water, low palaeolatitude, Campil Member was rapidly deposited, and
mixed carbonate-siliciclastic Werfen Formation may have been affected by low salinity levels
(of northern Italy) has been investigated in any (Twitchett 1999), which could have affected
detail (Twitchett & Wignall 1996; Twitchett both the size of the benthos and the amount of
1999). burrowing activity that occurred.
The domichnia of suspension feeders (e.g. The Spathian sediments of the Dolomites
Skolithos, Arenicolites) reappear in the lower region were deposited in a range of environments,
Siusi Member (late Griesbachian) of the from peritidal settings to offshore shelf. Bioturba-
Werfen Formation (Fig. 3). These sediments tion is usually high (ii3-6), and burrow diameters
were still deposited under some degree of benthic are larger than at any other time in the Early
oxygen restriction (Wignall & Twitchett 1996), Triassic. Spathian sediments record the first
but the P-Tr anoxic event was already beginning reappearance of Rhizocorallium, which com-
to weaken in this region (Wignall & Twitchett monly exceeds 10mm in diameter. Of the
2002a). Arenicolites and Skolithos penetrate only common late Permian ichnotaxa, only Thalassi-
some 1-2 cm into the substrate, and burrow dia- noides fails to appear by the Spathian. However,
meters are small (1-2 mm). They represent the the overlying Middle Triassic units (separated by
activities of a post-event pioneering community, an unconformity from the underlying Werfen
penetrating the upper surfaces of thin, distal Formation) are often well bioturbated by large
storm beds. Thalassinoides.
There is an improvement in benthic oxygena-
tion in the lower-middle Siusi Member (Wignall
& Twitchett 1996) and a subsequent increase Early Triassic ichnostratigraphy
in ichnodiversity, burrow size and the amount
and depth of bioturbation. Increase in burrow Can the stepwise reappearance of ichnotaxa
size at this time is mirrored by a similar through the Early Triassic recovery be used to
increase in the size of the shelly macrofauna correlate strata? Certainly, the pattern of re-
(Price-Lloyd & Twitchett 2002). Ichnotaxa appearance of different ichnotaxa, as well as
present include Cochlichnus, Catenichnus, Lock- the increase in burrow size and amount of
eia and Palaeophycus. In the shallower storm- bioturbation through the Werfen Formation of
deposited, micaceous fine sandstones of the the Dolomites, can be used to correlate with
upper Siusi Member (lower Dienerian in age) less fossiliferous deposits in adjacent regions
depth of bioturbation increases again with the (Twitchett 1997). The Servino Formation of
reappearance of Diplo crater ion (Twitchett Lombardy is one example.
1999). The Dipio crater ion burrows at this level The Servino Formation is a relatively thin,
are usually 3-4 mm in diameter, and may reach lateral equivalent of the Werfen Formation,
10cm in depth. which contains very few macrofossil remains
The overlying Gastropod Oolite Member, (Cassinis 1968). It is lithologically variable
which is Dienerian in age, records a further across Lombardy, and correlation with the
increase in burrow size (Twitchett 1999), ichno- Werfen Formation is often problematic (e.g.
diversity (Twitchett & Wignall 1996), and the Assereto et al. 1973; Posenato et al. 1996). In
amount of bioturbation. Ichnofabric indices addition, outcrop of the Servino Formation is
(sensu Droser & Bottjer 1986) may reach ii5 often patchy on the grass-covered hillsides,
for some beds, but are typically ii3. The sedi- which makes detailed mapping difficult. The
ments record deposition in a mixed carbonate- discovery of conodonts (Twitchett 1997, 2000)
siliciclastic environment above storm wavebase may aid correlation in the future, but a systema-
and below fair-weather wavebase. Both pre- tic study of this microfauna has yet to be under-
event and post-event ichnofaunas are present, taken. Meanwhile, trace fossils remain the best
and benthic conditions were clearly improving. means of correlation in the field.
The overlying Smithian age Campil Member, Two key ichnostratigraphic levels can be
however, records a temporary return to poorly identified. The first is the appearance of Diplo-
bioturbated sediments (iil-2) containing an craterion, which occurs in the thin storm beds
404 R. J. TWITCHETT & C. G. BARRAS

Fig. 3. Generalized stratigraphy of the Permian-Triassic strata of the Dolomites, northern Italy, showing
distribution of ichnogenera and sea-level curve. Ichnogeneric ranges from Twitchett & Wignall (1996) and
Twitchett (1997, 1999). Sea-level curve and stratigraphy modified from Broglio Loriga et al (1986). AH,
Andraz Horizon; GOM, gastropod oolite member; sp, supratidal; in, intertidal; sb, subtidal; 1, oolites; 2, mud
cracks; 3, wave ripples.

of micaceous fine sandstone, interbedded with correlated with those of the upper Siusi Member
laminated siltstones, in the lower pelitic unit of the Werfen Formation, suggesting a lower Die-
(Twitchett 1997; Fig. 4). Despite the common nerian age for this part of the Servino Formation.
occurrence of Diplocraterion, burrow density This supports previous lithological correlation, as
is never very great, and so there is minimal the overlying carbonate unit is usually correlated
disturbance to the primary sedimentary structures with the Gastropod Oolite Member (Assereto
(ii2). These Diplocraterion assemblages can be etal. 1973; Fig. 4).
ICHNOSTRATIGRAPHY & MASS EXTINCTION 405

Fig. 4. Generalized stratigraphy of the Servino Formation of eastern Lombardy (after Assereto & Rizzini
1975) showing distribution of trace fossils (data from Twitchett 1997, 2000, and recent field observations) and
correlation with the Werfen Formation.

The second important marker is the appear- also have stratigraphic value (Twitchett 1999).
ance of Rhizocorallium. The upper silt stone- Certainly, there is some correspondence between
dominated units of the Servino Formation the trace fossil record of the Lower Triassic
contain common Rhizocorallium (Fig. 4). Ichno- deposits of the western USA and that of northern
fabric index is high (ii3-5), and burrow diameters Italy. Both regions record deposition in similar
are larger than in the underlying parts of the environments at similar (tropical) palaeolati-
Servino Formation. In the Werfen Formation, tudes, and record similar post-extinction patterns
Rhizocorallium first appears in the Spathian of ecological recovery in the shelly fauna (cf.
(Val Badia through to San Lucano Members). Schubert & Bottjer 1995; Twitchett 1999). Both
This age designation agrees well with other regions record increasing burrow size, depth of
evidence, such as the rare occurrences of the bioturbation, amount of bioturbation and ichno-
foraminiferan Meandrospira pusilla within the diversity through the Early Triassic. The highest
upper parts of the Servino Formation, and levels of bioturbation occur in the Spathian
the fact that the underlying Middle Pelitic Unit deposits of the western USA (Schubert & Bottjer
is lithologically similar to the Campil Member 1995), although even here burrow depths do not
(Fig. 4; Twitchett 1997). exceed 10cm.
Globally, the first appearances of Diplocrater- Preliminary assessment by one of us (RJT)
ion and Rhizocorallium in the Early Triassic may indicates that the Griesbachian Dinwoody
406 R. J. TWITCHETT & C. G. BARRAS

Formation contains just simple horizontal fodi- Middle Triassic records are known, although
nichnia such as Planolites. U-shaped burrows Zoophycos is common in many later Mesozoic
with spreiten first occur in the Sinbad Limestone and Tertiary shelf sea sediments, and was
Member of the San Rafael Swell (southern extremely abundant in Upper Permian sediments
Utah), which is upper Nammalian (i.e. probably (e.g. Wignall et al. 1998). One possible explana-
Smithian) in age (Schubert & Bottjer 1995). tion is that the Permian Zoophycos trace-
These small-diameter burrows are steeply maker(s) became extinct during the P-Tr crisis,
inclined to the bedding, but are never vertical, and it then took many millions of years for the
and penetrate just a few centimetres into the sub- behaviour to be 'reinvented' by other organisms
strate. They could be classified as either Diplocra- (an Elvis ichnotaxon?). Alternatively, the habitat
terion or Rhizocorallium. The first undoubted preferences of the Zoophycos trace-maker(s) may
Rhizocorallium occur within the Spathian have changed, and they may have become
Virgin Limestone Member of the Moenkopi restricted to habitats that are (so far) unrecorded
Formation. In southern Nevada, the Virgin in the earliest Mesozoic sedimentary record.
Limestone contains a diverse ichnofauna, which Support is provided by Bottjer et al. (1988),
also includes Arenicolites, Asteriacites, Cochlich- who demonstrated that, since the Palaeozoic,
nus, Planolites and Skolithos. Finally, it appears Zoophycos has been restricted to deeper-water
that Thalassinoides, which is common in similar environments.
facies in the Permian (Schubert & Bottjer
1995), is completely absent from the Lower
Triassic sediments of the western USA, but Summary
does occur in the Middle Triassic (D. Bottjer
personal communication). This is identical to The P-Tr trace fossil record has been well studied
the post-extinction distribution of Thalassinoides at a number of localities worldwide, representing
in northern Italy (Twitchett 1999). a range of depositional environments. Ichno-
diversity, burrow size, depth and amount of
bioturbation are all severely reduced across the
Faster recovery at higher latitudes? P-Tr boundary. Only the small (millimetre-
sized), simple fodinichnia of deposit-feeding ani-
At low palaeolatitudes, Thalassinoides, attri- mals are present, sporadically, in the immediate
buted to the activities of crustaceans, is one of post-extinction aftermath, and penetrate just a
the last ichnotaxa to reappear (in the Middle few centimetres below the sediment surface.
Triassic, some 10 million years after the P-Tr Patterns of infaunal recovery are less well under-
extinction event). After recent, small-scale stood than details of the biotic crisis, but several
anoxic events, crustaceans are typically last to interesting ecological patterns are beginning
become re-established (e.g. Harper et al. 1991). to emerge. There is an increase in burrow
This suggests that patterns of benthic ecological size, ichnodiversity and depth of bioturbation
response may be broadly similar at vastly differ- through the Early Triassic, but the timing of
ent temporal scales, and may indicate that the recovery varies between regions. Lower-latitude
reappearance of Thalassinoides can be used to regions apparently took longer to recover than
mark the return to 'normal' marine conditions. higher latitudes (at least in the northern hemi-
However, the record at higher northern sphere). The stepwise reappearance of certain
palaeolatitudes is somewhat different. In East ichnotaxa (Diplo crater ion, Rhizocorallium and
Greenland, small, shallow-tier Thalassinoides Thalassinoides) may have some stratigraphic
are present in the upper part of the Wordie use for correlation between regions at similar
Creek Formation (personal observation), which palaeolatitude.
is dated as latest Griesbachian-early Dienerian
in age (Wignall & Twitchett 2002b). In Spitsber-
gen, Thalassinoides occurs in the Dienerian-age, Triassic-Jurassic (Tr-J) event
shallow shoreface deposits of the Vardebukta
Formation, soon after disappearance of oceanic The Tr-J extinction event was identified by
anoxia (Wignall et al. 1998). These data indicate Newell (1967) as one of the five major Phanero-
that, following the end-Permian event, crusta- zoic extinction episodes, with an estimated 80%
ceans reappeared sooner in mid-high palaeo- of species becoming extinct (e.g. Palfy et al.
latitudes than in the tropics. 2000), although our understanding of this
One common element of the pre-event ichno- event remains poor, as few complete marine
fauna that apparently fails to reappear, even by boundary sections are known. Some authors
the Middle Triassic, is Zoophycos. No Early or have argued that levels of extinction may have
ICHNOSTRATIGRAPHY & MASS EXTINCTION 407

Fig. 5. Sedimentary log of the Triassic-Jurassic of Pinhay Bay, southern England, showing the distribution of
trace fossils and ichnofabric indices (ii). Horizontal ichnofabric calculated from methods of Miller & Smail
(1997). Vertical ichnofabric follows Droser & Bottjer's (1986) scheme.

been overestimated, as disappearances usually Bay (Somerset) and Pinhay Bay (Dorset).
coincide with major facies change (Cuny 1995; Despite differences in the thickness of the
Hallam 2002). Blue Lias at these two sites, the trace fossil
The tracks of Late Triassic and Early Jurassic records of both show remarkable similarity, and
tetrapods have been used to study the faunal so only Pinhay Bay is discussed in detail here
response of terrestrial vertebrates through the (Fig. 5).
Tr-J interval (Olsen et al. 2002). This represents To the west of Pinhay Bay (SY 295895), the
the only dedicated study of the terrestrial trace top of the Rhaetian Lilstock Formation is locally
fossil record through any of the major extinction heavily bioturbated by 6-1 Omm diameter Diplo-
events. Here we present the first analysis of the craterion, which penetrate up to 13 cm. The lack
marine trace fossil record through the Tr-J of a more diverse ichnofauna in the Lilstock
boundary, from our investigations in southern Formation is probably due to deposition under
England, Austria, and Nevada. unusual salinity conditions (Hallam & El
Shaarawy 1982). Diplocraterion disappears at
the Formation boundary, and does not reappear
England until the angulata Zone of the Hettangian, where
burrow diameter is significantly smaller than in
Sections through the Uppermost Triassic Lilstock the Rhaetian (Fig. 5).
Formation and the Lower Jurassic Blue Lias The lowest pre-planorbis beds of the Blue Lias
Formation have been examined at St Audries Formation are laminated and unbioturbated,
408 R. J. TWITCHETT & C. G. BARRAS

Fig. 6. Changes in the depth of bioturbation through the Triassic-Jurassic of Pinhay Bay. Open circles
represent mean burrow depth; closed circles, maximum burrow depth. P values indicate the statistical
significance (Kolmogorov-Smirnov test) between Lilstock Formation and planorbis Zone datasets, and between
planorbis Zone and bucklandi Zone datasets.

which may indicate an episode of anoxia. Areni- between the prQ-planorbis and planorbis beds
colites and small Thalassinoides appear in the (Fig. 6). Both the ichnodiversity and the degree
upper prQ-planorbis beds at Pinhay Bay, and of bioturbation continue to increase through
there is a coincident increase in vertical ichnofab- the overlying liasicus Zone with the (re)appear-
ric index (sensu Droser & Bottjer 1986). Burrow ance of Chondrites (Fig. 5).
depth, measured from Arenicolites, never exceeds Maximum ichnodiversity is reached in the
5 cm. Relatively small burrow size, shallow depth angulata Zone, which contains Arenicolites, Chon-
of bioturbation and low ichnofabric index are drites, Diplocraterion, Palaeophycus, Planolites,
consistent with an oxygen-restricted (dysaerobic) Rhizocorallium and Thalassinoides. Ichnodiversity
environment (e.g. Rhoads & Morse 1971; remains at this level through the Sinemurian
Savrda & Bottjer 1986). A recent study (Wignall (bucklandi and semicostatum Zones). Maximum
2001) revealed the presence of abundant pyrite depth of burrowing also increases in the angulata
framboids in the lowest pre-planorbis beds, Zone to 17cm (Fig. 6), as a consequence of the
while spectral gamma ray analysis showed an reappearance of Diplocraterion.
exceptionally low Th/U ratio. These observa- However, the diameters of these Diplocraterion
tions, too, are indicative of anoxic deposition. burrows are still significantly smaller than those
However, Arenicolites and Thalassinoides are of the Rhaetian examples, and pre-Jurassic
not typical dysaerobic ichnotaxa, and the small size is not reached in Diplocraterion until the
size of the infauna may also be due to other Sinemurian (bucklandi Zone). Most ichnotaxa,
factors, such as a decrease in food supply, and except Rhizocorallium and Chondrites, which
is typical of immediate post-extinction intervals show no size increase, are small when they first
(cf. Urbanek 1993). Ichnofabric indices are vari- (re)appear, and increase in size through the
able, and may reflect changes in sedimentation Hettangian and into the basal zones of the
rate or population density of the infauna. Sinemurian (Fig. 7). This increase in trace-
Palaeophycus, Planolites, Rhizocorallium and maker body size may be a response to improving
Thalassinoides are recorded from the overlying environmental conditions such as food supply
planorbis Zone. There is an associated increase or oxygen levels, and mirrors the size increase
in ichnofabric index (Fig. 5). However, there of the marine shelly macrofauna during the
appears to be little change in depth of burrowing same interval (e.g. Hallam 1975).
ICHNOSTRATIGRAPHY & MASS EXTINCTION 409

Fig. 7. Size changes of selected ichnogenera through the Triassic-Jurassic of Pinhay Bay. Open circles
represent mean burrow diameter; closed circles, maximum burrow diameter. P values indicate the statistical
significance between the datasets shown by the arrows at the end of each line (Kolmogorov-Smirnov test)

Central Austria ammonite fauna and, unlike the Rhaetian of


England, were deposited under normal marine
Two sections were studied by one of us (CGB), in conditions similar to those of the overlying
the Salzkammergut region of Central Austria. Hettangian. Ichnofaunal changes across the
The Rhaetian sediments of Austria contain an Tr-J boundary of Austria are thus more likely
410 R. J. TWITCHETT & C. G. BARRAS

Fig. 8. Composite sedimentary log of the Triassic-Jurassic sections of Central Austria exposed at GaiBau and
Kendelbach Gorge (see text for details). Key as in Fig. 6. Vertical ichnofabric index (ii) follows Droser and
Bottjer's (1986) scheme.

to reflect the effects of the end-Triassic mass identifiable trace fossils, and only limited
extinction, rather than facies change. evidence of bioturbation, could be found in the
The Rhaetian Kossen Formation, exposed in a Hettangian sediments.
road cutting near GaiBau, is well bioturbated Limestone samples from the Kossen Forma-
(ii3—4) and contains Diplocraterion, Planolites, tion and overlying Kendelbach Formation at
Rhizocorallium, Skolithos and Zoophycos (Fig. Kendelbach Gorge are completely homogeneous
8). The Rhaetian ichnodiversity of Austria is in vertical section. It is, however, unclear whether
therefore higher than in England, where the this homogeneity reflects the uniform nature of
sediments were deposited under unusual salinity original deposition, or is a result of total bio-
conditions (Hallam & El Shaarawy 1982). turbation, or diagenesis.
The Kossen Formation was examined again The apparent absence of trace fossils from the
approximately 60m up-section (Zankl 1971), in upper Kossen Formation may suggest a deterior-
nearby Kendelbach Gorge. Deposition of the ation of environmental conditions approaching
Kossen Formation continues to the Tr-J bound- the Tr-J boundary in Central Austria. The failure
ary, and at Kendelbach Gorge there is exposure of the ichnofauna to reappear at any point within
of the boundary itself. However, although the the overlying Hettangian sediments at Kendel-
upper Kossen Formation limestones are similar bach Gorge implies that any disturbance to the
in appearance and lithology to those at GaiBau, marine ecosystem continued to affect the trace-
they contain no readily observed trace fossils making taxa well into the Early Jurassic.
(Fig. 8). At Kendelbach Gorge, the Hettangian In contrast, the shelly fossil record suggests
Kendelbach Formation contains shallow that recovery at Kendelbach Gorge was rela-
marine limestones and is overlain by the tively rapid, and occurred even in the latest
deeper-water Sinemurian Adnet Formation. No Rhaetian (Hallam 1990). Seven metres below
ICHNOSTRATIGRAPHY & MASS EXTINCTION 411

Fig. 9. Sedimentary log of the Triassic-Jurassic section in New York Canyon, Nevada. Key as in Fig. 5.
Asterisked letters show stratigraphic positions of photographs shown in Fig. 10. Vertical ichnofabric index (ii)
follows Droser & Bottjer's (1986) scheme.

the Tr-J boundary, within the Kossen Forma- ary of Nevada appear to have been minimal.
tion, there are shale horizons containing an Trace fossil changes are thus not complicated by
impoverished bivalve fauna (Hallam 1990). salinity considerations. There is, however, a
This correlates with the decrease in bioturbation facies change in the upper Gabbs Formation
and ichnodiversity. However, prior to the (Muller Canyon Member), with a reduction in
first appearance of the ammonite Psiloceras the number of resistant limestone horizons with
planorbis in the Hettangian, Hallam documents bedding plane exposure (Fig. 9).
an increase in faunal diversity, and the A relatively diverse ichnofauna is present in the
appearance of forams, ostracods, crinoids and Mount Hyatt Member of the Gabbs Formation
brachiopods. (Fig. 9), consisting of Arenicolites, Planolites,
Rhizocorallium, Skolithos and Thalassinoides
(Fig. lOe). Ichnofabric index is low, however
Nevada (maximum ii2; Fig. lOf). The majority of these
ichnotaxa are absent from the overlying Muller
In the New York Canyon area, Gabbs Valley Canyon Member, which straddles the Tr-J bound-
Ranges, Nevada, marine limestones and siltstones ary (Taylor et al. 1983). Indistinct burrows, here
from the Upper Triassic (Rhaetian) Gabbs referred to Planolites, occur rarely (Fig. lOd).
Formation and the Lower Jurassic (Hettangian) The Muller Canyon Member contains relatively
Sunrise Formation were examined. An ammonite few limestone beds and little horizontal bedding
fauna is present throughout the section, and so, as plane exposure. In the underlying and overlying
in Austria, salinity changes across the Tr-J bound- units, trace fossils appear to be confined to the
412 R. J. TWITCHETT & C. G. BARRAS

Fig. 10. Polished blocks and field photographs indicating evidence for the presence or absence of
ichnofauna: (a) Rhizocorallium burrows from the Ferguson Hill Member; (b) polished block showing mottled
fabric cut by horizontal burrows from the Ferguson Hill Member; (c) polished block showing traces of original
bedding from the Ferguson Hill Member; (d) polished block showing mottled fabric cut by large burrows from
the Muller Canyon Member; (e) Thalassinoides burrow from Mount Hyatt Member (see Fig. 9 for stratigraphic
positions); (f) polished block showing well-preserved bedding cut by rare burrows in the Mount Hyatt
Member.
ICHNOSTRATIGRAPHY & MASS EXTINCTION 413

limestones and are most easily observed on hori- Summary


zontal surfaces. Thus facies change is the most
likely explanation for the low ichnodiversity of These data have shown that the nature of the
the Muller Canyon Member. trace fossil record changes through the Tr-J
An earlier study (Hallam & Wignall 2000) boundary of each of these three regions (Fig.
reported intense bioturbation from the central 11). Some of these changes, such as those at
part of the Muller Canyon Member, with a thor- New York Canyon, can be attributed to facies
oughly mottled fabric of indistinct traces cut by change, but others, such as those through the
abundant Helminthoida. However, other than Blue Lias Formation of England, appear to
rare Planolites burrows, no other ichnotaxa were occur with no significant change in sedimentary
found from the Muller Canyon Member in the facies. These results show that normal marine
current study, and we were unable to confirm environments of Rhaetian age contain a diverse
this observation. Furthermore, although cut and ichnofauna, which may include Arenicolites,
polished rock samples from this interval often Diplo crater ion, Planolites, Rhizocorallium, Sko-
showed a burrow-mottled fabric, the rocks them- lithos, Thalassinoides and Zoophycos. There is
selves were relatively fissile, suggesting that some some evidence that ichnodiversity, burrow size,
of the original bedding remains. depth of bioturbation and ichnofabric index are
Arenicolites, Planolites, Rhizocorallium, Sko- much reduced in the uppermost Rhaetian (e.g.
lithos and Thalassinoides all reappear in the pre-planorbis beds of England), presumably as
Ferguson Hill Member of the Sunrise Formation a result of environmental deterioration, which
(Fig. lOa), which is dated as upper Hettangian to may be attributed to dysoxia. There is an
lower Sinemurian (Taylor et al. 1983). Chondrites increase in ichnodiversity, ichnofabric index,
and Diplo crater ion are also present in this burrow size and depth of burrowing through
unit. The amount of bioturbation recorded in the Hettangian as benthic oxygen levels improve,
individual beds varies. Some beds have a mottled although rates of recovery vary. Ichnodiversity
fabric, cut by prominent burrows (e.g. Fig. lOb), returned to Rhaetian levels by the angulata
whereas others show the remains of original Zone of the Hettangian (at least in England
bedding (e.g. Fig. lOc). There is little difference and Nevada; Fig. 11). Ichnotaxa recorded at
in burrow diameter between the ichnotaxa of this time include Arenicolites, Chondrites, Diplo-
the Gabbs and those of the Sunrise Formations. craterion, Palaeophycus, Planolites, Rhizocoral-
This suggests that palaeoenvironments of the lium and Thalassinoides. Burrow size of some
Ferguson Hill Member and Mount Hyatt ichnotaxa (e.g. Diplocraterion, Fig. 7) continues
Member were similar. to increase through into the Sinemurian, which

Fig. 11. Trends in ichnodiversity through the Triassic-Jurassic intervals of England, Central Austria and
Nevada, USA. Gap in the Austrian record is the result of diagenesis (see text for details).
414 R. J. TWITCHETT & C. G. BARRAS

parallels the Early Jurassic rise in body size of lar to the patterns observed through the P-Tr and
macroinvertebrates such as ammonites and Tr-J intervals.
bivalves (Hallam 1975; Dommergues et al. Trace fossils have also been used in novel
2002). Changes in the depth of burrowing palaeoenvironmental analyses of the so-called
(Fig. 6) indicate that in the latest Rhaetian and 'tsunami beds' of the southern USA (Savrda
earliest Hettangian (planorbis and Hastens Zones) 1993) and Mexico (Ekdale & Stinnesbeck
only the shallowest tiers are occupied (cf. Fig. 1). 1998). Following the Alvarez et al (1980)
The stepwise reappearance of some ichnotaxa, impact hypothesis, and especially with the identi-
such as Chondrites and Diplocraterion, may have fication of the crater itself beneath Chicxulub
ichnostratigraphic value. Certainly, in the UK, (Hildebrand et al. 1991), evidence was sought
Diplo crater ion appears to be absent from Jurassic for the giant, impact-generated, tsunamis that
strata until the angulata Zone beds of the Blue were hypothesized to have devastated the coastal
Lias (Fig. 5). If the angulata Zone is considered plains surrounding the present-day Gulf of
to be the level at which 'normal' infaunal com- Mexico. Several spherule-bearing sandstones,
munities have reappeared, then post-extinction up to a few metres thick and of K-T age, were
recovery is clearly far more rapid than, for soon identified as tsunami deposits, based on
example, after the P-Tr event. limited sedimentological evidence such as their
coarse grain size (in otherwise fine-grained sedi-
ments) and apparently rapid deposition (e.g.
Cretaceous-Tertiary (K-T) event Bourgeois et al 1988).
Although the interpretation of these units as
Trace fossils have, thus far, contributed relatively tsunami deposits fitted perfectly with the expec-
little to our understanding of the benthic tations of supporters of the impact model, such
response to the K-T event. The only study of 'hypothesis-driven interpretations' (Stinnesbeck
the marine trace fossil record through the extinc- et al 1994) were not universally accepted.
tion-recovery interval is Ekdale and Bromley's Bohor (1996) suggested that they were turbidite
(1984) account of the ichnology of the K-T deposits, which may have been triggered by tsu-
boundary chalks of Denmark. They found that namis or seismic events related to the impact.
in deeper basinal areas a diverse ichnofauna Others argued against any seismic or tsunami
containing Zoophycos, Chondrites and Thalassi- involvement and instead interpreted them as
noides continues to the top of the Maastrichtian, the normal transgressive infilling of incised
where there is a sudden change to the Thalassi- valleys cut during an episode of sea-level fall
noides-dominatQd ichnofauna of the lower (e.g. Savrda 1993; Stinnesbeck et al 1993). One
Danian limestones. The disappearance of Zoo- key prediction of this latter hypothesis is that
phycos, and its absence from the lower Danian deposition was not catastrophic, but took place
sediments, is attributed to shallowing across over an extended period of time.
the boundary and a change to more soupy The trace fossil studies of Savrda (1993), who
substrates (Ekdale & Bromley 1984). Certainly, investigated the Clayton Sands of Alabama,
there is no evidence for a change in oxygen and Ekdale & Stinnesbeck (1998), who analysed
levels, as the sediments are well bioturbated the sandstones of the Mendez Formation, north-
throughout. However, there may be a preserva- eastern Mexico, have proven to be crucial in the
tional issue here as the lower Danian chalks are correct interpretation of these deposits. In both
highly compacted, making burrow identification cases, the 'tsunami beds' could be divided into
problematic (Ekdale & Bromley 1984). three subunits, each with a different suite of
In shallower parts of the basin, Zoophycos trace fossils. For example, the lower unit of the
declines in abundance in the Upper Maastrichtian Clayton Sands contains rare Ophiomorpha,
chalks, which is again attributed to sea-level fall emplaced prior to deposition of the overlying
(Ekdale & Bromley 1984), so that the uppermost, middle unit, which contains Thalassinoides. The
grey Maastrichtian chalks are dominated by Tha- upper unit contains Thalassinoides and a
lassinoides. These are overlain by the Lower number of horizons with small diameter (3-
Danian Fish Clay, which records an episode of 5 mm), shallow penetrating (less than 1 cm)
oxygen restriction evinced by laminated, black, Planolites. The multiple horizons of trace fossil
carbonaceous sediments. Ichnofabric index, ich- emplacement indicate that deposition of the
nodiversity and burrow size all apparently decline Clayton Sands took place over an extended
in the lower part of the Fish Clay, although quan- period of time, with breaks in deposition allow-
titative data are lacking, and increase again in the ing the burrowing infauna to colonize the sub-
transition to the Danian chalks above. These strate (Savrda 1993). The results of Ekdale &
changes in the trace fossil record are clearly simi- Stinnesbeck (1998) were similar, and so the
ICHNOSTRATIGRAPHY & MASS EXTINCTION 415

tsunami origin for these units could be rejected. ALVAREZ, L. W., ALVAREZ, W., ASARO, F. & MICHEL,
The studies of Savrda (1993) and Ekdale & H. V. 1980. Extraterrestrial causes of the
Stinnesbeck (1998) show the immense value of Cretaceous-Tertiary extinction. Science, 208,
ichnology in palaeoenvironmental analysis, and 1095-1108.
ARMSTRONG, H. A. 1996. Biotic recovery after mass
how it is crucial to correct interpretation of
extinction: the role of climate and ocean state in
sedimentary units, especially when testing the postglacial (Late Ordovician-Early Silurian)
hypotheses of catastrophic deposition. Despite recovery of the conodonts. In: HART, M. B. (ed.)
the conclusions of these two studies, so-called Biotic Recoveries from Mass Extinction Events.
tsunami deposits continue to be identified in the Geological Society, London, Special Publications,
Gulf of Mexico region (e.g. Tada et al. 2002) 102, 105-117.
without any reference to the trace fossil assem- ASSERETO, R. L. & RIZZINI, A. 1975. Reworked ferroan
blages they contain. dolomite grains in the Triassic 'oolite a gastero-
podi' of the Camoniche Alps (Italy) as indicators
of early diagenesis. Neues Jahrbuch fur Geologie
Summary und Palaontologie Abhandlungen, 148, 215-232.
ASSERETO, R. L., BOSELLINI, A., FANTINI SESTINI, N. &
The trace fossil records of four of the five major SWEET, W. C. 1973. The Permian-Triassic bound-
Phanerozoic mass extinction events have been ary in the Southern Alps (Italy). In: LOGAN, A. &
HILLS, L. V. (eds) The Permian and Triassic
analysed, but coverage is variable. To date, the Systems and Their Mutual Boundary. Memoirs of
Permian-Triassic trace fossil record has been the Canadian Society of Geology and Petrology,
the most intensively studied of these and has Calgary, Alberta, 2, 176-199.
been important in palaeoenvironmental analysis, AUSICH, W. I. & BOTTJER, D. J. 2001. Sessile inverte-
and crucial in providing insights into the ecologi- brates. In: BRIGGS, D. E. G. & CROWTHER, P. R.
cal response of the marine benthos. Trace fossils (eds) Palaeobiology H. Blackwell Science, Oxford,
are the only records we have of the response of 384^386.
the soft-bodied burrowing infauna to major BENTON, M. J. 1995. Diversification and extinction in
biotic events of the past. The post-extinction the history of life. Science, 268, 52-58.
BENTON, M. J. 2003. When life nearly died: the greatest
stepwise reappearance of ichnotaxa may have
mass extinction of all time. Thames and Hudson,
some ichnostratigraphic use, but only within London, New York.
similar environmental settings at similar palaeo- BOHOR, B. F. 1996. A sediment gravity-flow hypothesis
latitudes. Trace fossils have great potential for for siliciclastic units at the K/T boundary, north-
increasing our understanding of these important eastern Mexico. In: RYDER, G., FASTOVSKY, D. &
events in the evolution of life on Earth: we have GARTNER, S. (eds) The Cretaceous-Tertiary Event
only just begun to scratch the surface. and Other Catastrophes in Earth History. Geo-
logical Society of American Special Papers,
CGB's fieldwork in Austria was funded by a Sylvester Boulder, Colorado, 307, 183-195.
Bradley Award (Palaeontological Association). BOURGEOIS J., HANSEN, T. A., WIBERG, P. L. & KAUFF-
Fieldwork in England was funded by the John Ray MAN, E. F. 1988. A tsunami deposit at the
Trust, and fieldwork in Nevada was funded by the Cretaceous-Tertiary boundary in Texas. Science,
Geological Society of London. R. Scott and W. 241, 567-570.
Barras are thanked for assistance in the field. C. Paul BOTTJER, D. J. & AUSICH, W. I. 1986. Phanerozoic
is thanked for taking us through the coastal sections development of tiering in soft substrata suspension
west of Pinhay Bay. RJT thanks D. Sciunnach for shar- feeding communities. Paleobiology, 12, 400-^420.
ing his knowledge of the Lower Triassic sediments of BOTTJER, D. J., DROSER, M. L. & JABLONSKI, D. 1988.
eastern Lombardy, and D. Bottjer, M. Fraiser and S. Palaeoenvironmental trends in the history of
Pruss for similar discussions on the trace fossil record trace fossils. Nature, 333, 252-255.
of the western USA. Thanks to staff in the Department BRENCHLEY, P. J., MARSHALL, J. D. et al. 1994. Bathy-
of Earth Sciences (University of Bristol, UK) and metric and isotopic evidence for a short lived Late
Department of Earth and Planetary Science (UC Ordovician glaciation in a greenhouse period.
Berkeley) for use of facilities. Thorough reviews by Geology, 22, 295-298.
P. Wignall and M. Garton greatly improved the quality BRIGGS, D. E. G. & GALL, J.-C. 1990. The continuum in
of this publication. soft-bodied biotas from transitional environments:
a quantitative comparison of Triassic and Carbo-
niferous Konservat-Lagerstatten. Paleobiology,
References 16, 204-218.
BROGLIO LORIGA, C., NERI, C. & POSENATO, R. 1986.
ALLISON, P. A. & BRIGGS, D. E. G. 1991. Taphonomy The Lower Triassic of the Dolomites and Cadore.
of nonmineralised tissues. In: Allison, P. A. & In Permian and Permian—Triassic Boundary in
Briggs, D. E. G. (eds) Taphonomy: Releasing the the South-Alpine Segment of Western Tethys.
Data Locked in the Fossil Record. Plenum Press, IGCP Project 203, Excursion Guidebook, Brescia,
New York, 25-70. 29-34.
416 R. J. TWITCHETT & C. G. BARRAS

BROMLEY, R. G. 1990. Trace Fossils: Biology and Taph- marine invertebrates. Philosophical Transactions
onomy. Unwin, London. of the Royal Society, Series B, 325, 437-^55.
BROMLEY, R. G. & EKDALE, A. A. 1984. Chondrites: a HALLAM, A. 1990. Correlation of the Triassic-Jurassic
trace fossil indicator of anoxia in sediments. boundary in England and Austria. Journal of the
Science, 224, 872-874. Geological Society, 147, 421^24.
CASSINIS, G. 1968. Studio stratigrafico del 'Servino' di HALLAM, A. 2002. How catastrophic was the end-
Passo Valdi (Trias inferiore dell' alta Val Caffaro). Triassic mass extinction? Lethaia, 35, 147-157.
Atti dell' Instituto Geologico della Universita di HALLAM, A. & EL SHAARAWY, Z. 1982. Salinity
Pavia, 19, 15-39 [in Italian]. reduction of the end-Triassic sea from the Alpine
CHEN, D. & TUCKER, M. E. 2003. The Frasnian- region into northwestern Europe. Lethaia, 15,
Famennian mass extinction: insights from 169-178.
high resolution sequence stratigraphy and HALLAM, A. & WIGNALL, P. B. 1997. Mass Extinctions
cyclostratigraphy in South China. Palaeogeo- and Their Aftermath. Oxford University Press,
graphy, Palaeoclimatology, Palaeoecology, 193, Oxford.
87-111. HALLAM, A. & WIGNALL, P. B. 2000. Facies changes
CRIMES, T. P. 1987. Trace fossils and correlation of late across the Triassic-Jurassic boundary in Nevada,
Precambrian and early Cambrian strata. Geologi- USA. Journal of the Geological Society of
cal Magazine, 124, 97-119. London, 157, 49-54.
CRIMES, T. P. 1992. Changes in the trace fossil biota across HARPER, D. E. JR, McKiNNEY, L. D., NANCE, J. M. &
the Proterozoic-Phanerozoic boundary. Journal of SALZER, R. R. 1991. Recovery responses of two
the Geological Society, London, 149, 637-646. benthic assemblages following an acute hypoxic
CUNY, G. 1995. French vertebrate faunas and the event on the Texas continental shelf, northwestern
Triassic-Jurassic boundary. Palaeogeography, Gulf of Mexico. In: TYSON, R. V. & PEARSON, T. H.
Palaeoclimatology, Palaeoecology, 119, 343-358. (eds) Modern and Ancient Continental Shelf
DOMMERGUES, J. L., MONTUIRE, S. & NfilGE, P. 2002. Anoxia. Geological Society, London, Special
Size patterns through time: the case of the Early Publications, 58, 49-64.
Jurassic ammonite radiation. Paleobiology, 28, HARRIES, P. J. & KAUFFMAN, E. G. 1990. Patterns of
423-^34. survival and recovery following the Cenomanian-
DROSER, M. L. & BOTTJER, D. J. 1986. A semiquantita- Turonian (Late Cretaceous) mass extinction in the
tive field classification of ichnofabric. Journal of Western Interior Basin, United States. In: KAUFF-
Sedimentary Petrology, 56, 558-559. MAN, E. G. & WALLISER, O. H. (eds) Extinction
EKDALE, A. A. 1985. Paleoecology of the marine Events in Earth History. Lecture Notes in Earth
endobenthos. Palaeogeography, Palaeoclimatol- History, 30, Springer, Berlin, 277-298.
ogy, Palaeoecology, 50, 63-81. HARRIES, P. J., KAUFFMAN, E. G. & HANSEN, T. A.
EKDALE, A. A. & BROMLEY, R. G. 1984. Sedimentology 1996. Models for biotic survival following mass
and ichnology of the Cretaceous-Tertiary bound- extinction. In: HART, M. B. (ed.) Biotic Recovery
ary in Denmark: implications for the causes of from Mass Extinction Events. Geological Society,
the terminal Cretaceous extinction. Journal of London, Special Publications, 102, 41-60.
Sedimentary Petrology, 54, 681-703. HlLDEBRAND, A. R., PENFIELD, G. T., KRING, D. A.,
EKDALE, A. A. & STINNESBECK, W. 1998. Trace fossils PILKINGTON, M. & CAMARGO, Z. A. 1991. Chicxu-
in Cretaceous-Tertiary (KT) boundary beds in lub crater: a possible Cretaceous/Tertiary bound-
northeastern Mexico: implications for sedimen- ary impact crater in the Yucatan Peninsula.
tation during the KT boundary event. Palaios, Geology, 19, 867-871.
13, 593-602. JABLONSKI, D. 1996. Body size and macroevolution. In:
ERWIN, D. H. 1993. The Great Paleozoic Crisis: life and JABLONSKI, D., ERWIN, D. H. & LIPPS, J. H. (eds)
Death in the Permian. Columbia University Press, Evolutionary Paleobiology. Chicago University
New York. Press, Chicago, 256-289.
FINNEY, S. C., BERRY, W. B. N. ET AL. 1999. Late JUMARS, P. A. & WHEATCROFT, R. A. W. 1989.
Ordovician mass extinction: a new perspective Responses of benthos to changing food quality
from stratigraphic sections in central Nevada. and quantity, with a focus of deposit feeding and
Geology, 27, 215-218. bioturbation. In: BERGER, W. H., SMETACEK, V. S.
GIRARD, C. & RENAUD, S. 1996. Size variation in & WEFER, G. (eds) Productivity of the Ocean: Past
conodonts in response to the Upper Kellwasser and Present. Wiley, Chichester, 235-253.
crisis (Upper Devonian of the Montagne Noire, LANDING, E. 1994. Precambrian-Cambrian boundary
France). Comptes Rendus de I'Academie des global stratotype ratified and a new perspective
Sciences, Serie Ha, 323, 435^42. of Cambrian time. Geology, 22, 179-182.
HALLAM, A. 1965. Environmental causes of stunting in LOOY, C. V., TWITCHETT, R. J., DILCHER, D. L., KONIJ-
living and fossil marine benthonic invertebrates. NENBURG-VAN CITTERT, J. H. A. & VISSCHER, H.
Palaeontology, 8, 132-155. 2001. Life in the end-Permian dead zone. Proceed-
HALLAM, A. 1975. Evolutionary size increase and long- ings of the National Academy of Sciences, USA, 98,
evity in Jurassic bivalves and ammonites. Nature, 7879-7883.
258, 493^96. McCANN, T. 1990. Distribution of Ordovician-Silurian
HALLAM, A. 1989.The case for sea-level change as a ichnofossil assemblages in Wales: implications for
dominant causal factor in mass extinctions of Phanerozoic ichnofaunas. Lethaia, 23, 243-255.
ICHNOSTRATIGRAPHY & MASS EXTINCTION 417

MILLER, M. F. & SMAIL, S. E. 1997. A semiquantitative Cretaceous-Tertiary event and Other Catastrophes
field method for evaluating bioturbation on bed- in Earth History. Geological Society of America
ding planes. Palaios, 12, 391-396. Special Papers, Boulder, Colorado, 307, 183-195.
NEWELL, N. D. 1967. Revolutions in the History of Life. STINNESBECK, W., BARBARIN, J. M. ET AL. 1993. Deposi-
Geological Society of America Special Papers, tion of channel deposits near the Cretaceous-
Boulder, Colorado, 89, 63-91. Tertiary boundary in northeastern Mexico:
OLSEN, P. E., KENT, D. V. ET AL. 2002. Ascent of catastrophic or 'normal' sedimentary deposits.
dinosaurs linked to the Iridium anomaly at the Geology, 21, 797-800.
Triassic-Jurassic boundary. Science, 296, 1305- STINNESBECK, W., KELLER, G., ADATTE, T. & MACLEOD,
1307. N. 1994. Deposition of channel deposits near the
ORR, P. J. 2001. Colonization of the deep-marine Cretaceous-Tertiary boundary in northeastern
environment during the early Phanerozoic: the Mexico: catastrophic or 'normal' sedimentary
ichnofaunal record. Geological Journal, 36, 265- deposits. Reply. Geology, 22, 955-956.
278. TADA, R., NAKANO, Y. ET AL. 2002. Complex tsunami
PALFY, J., MORTENSEN, J. K., CARTER, E. S., SMITH, P. waves suggested by the Cretaceous-Tertiary
L., FRIEDMAN, R. M. & TIPPER, H. W. 2000. boundary deposit at the Moncada section, western
Timing the end-Triassic mass extinction: first on Cuba. In: KOEBERL, C. & MACLEOD, K. G. (eds)
land, then in the sea? Geology, 28, 39^2. Catastrophic events and mass extinctions: impacts
POSENATO, R., SCIUNNACH, D. & GARZANTI, E. 1996. and beyond. Gelogical Society of America, Special
First report of Claraia (Bivalvia) in the Servino Papers, Boulder, Colorado, 356, 109-123.
Formation (Lower Triassic) of the western TAYLOR, D. G., SMITH, P. L., LAWS, R. A. & GUEX, J.
Orobic Alps, Italy. Rivista Italiana di Paleontolo- 1983. The stratigraphy and biofacies trends of
gia e Stratigrafia, 102, 201-210. the Lower Mesozoic Gabbs and Sunrise forma-
PRICE-LLOYD, N. & TWITCHETT, R. J. 2002. The Lilli- tions, west-central Nevada. Canadian Journal of
put effect in the aftermath of the end-Permian Earth Science, 20, 1598-1608.
mass extinction event. GSA Annual Meeting TWITCHETT, R. J. 1997. Palaeoenvironments of the
Program with Abstracts, 34, 355. Lower Triassic of the Dolomites, northern Italy.
RHOADS, D. C. & MORSE, J. W. 1971. Evolutionary and PhD thesis, University of Leeds.
ecologic significance of oxygen-deficient marine TWITCHETT, R. J. 1999. Palaeoenvironments and faunal
basins. Lethaia, 4, 413-428. recovery after the end-Permian mass extinction.
SAGEMAN, B. B. & BINA, C. R. 1997. Diversity and Palaeogeography, Palaeo climatology, Palaeoecol-
species abundance patterns in Late Cenomanian ogy, 154, 27-37.
black shale biofacies, Western Interior, US. TWITCHETT, R. J. 2000. A high resolution biostratigra-
Palaios, 12, 449^66. phy for the Lower Triassic of northern Italy.
SAVRDA, C. E. 1993. Ichnosedimentological evidence Palaeontology Newsletter, 43, 19-22.
for a non-catastrophic origin of Cretaceous- TWITCHETT, R. J. 2001. Incompleteness of the Permian-
Tertiary boundary sands in Alabama. Geology, Triassic fossil record: a consequence of productivity
21, 1075-1078. decline? Geological Journal, 36, 341-353.
SAVRDA, C. E. & BOTTJER, D. J. 1986. Trace fossil TWITCHETT, R. J. & WIGNALL, P. B. 1996. Trace fossils
model for reconstruction of paleo-oxygenation in and the aftermath of the Permo-Triassic mass
bottom waters. Geology, 14, 3-6. extinction: evidence from northern Italy. Palaeo-
SAVRDA, C. E. & BOTTJER, D. J. 1987. The exaerobic geography, Palaeoclimatology, Palaeoecology,
zone, a new oxygen-deficient marine biofacies. 124, 137-151.
Nature, 327, 54-56. TWITCHETT, R. J., LOOY, C. V., MORANTE, R.,
SCHUBERT, J. K. & BOTTJER, D. J. 1995. Aftermath of VISSCHER, H. & WIGNALL, P. B. 2001. Rapid and
the Permian-Triassic mass extinction event: synchronous collapse of marine and terrestrial
paleoecology of Lower Triassic carbonates in the ecosystems during the end-Permian mass extinc-
western USA. Palaeogeography, Palaeoclimatol- tion event. Geology, 29, 351-354.
ogy, Palaeoecology, 116, 1-39. URBANEK, A. 1993. Biotic crises in the history of Upper
SEILACHER, A. 1967. Bathymetry of trace fossils. Silurian graptoloids: a palaeobiological model.
Marine Geology, 5, 413-428. Historical Biology, 7, 29-50.
SEILACHER, A., LUNING, S., MARTIN, M. A., KLITZSCH, VOSSLER, S. M. & PEMBERTON, S. G. 1988. Skolithos in
E., KHOJA, A. & CRAIG, J. 2002. Ichnostrati- the Upper Cretaceous Cardium Formation: an
graphic correlation of Lower Palaeozoic elastics ichnofossil example of opportunistic ecology.
in the Kufra Basin (SE Libya). Lethaia, 35, 257- Lethaia, 21, 351-362.
262. WIGNALL, P. B. 2001. Sedimentology of the Triassic-
SEPKOSKI, J. J. JR 1984. A kinetic model of Phanerozoic Jurassic boundary beds in Pinhay Bay (Devon,
taxonomic diversity. III. Post-Paleozoic fami- SW England). Proceedings of the Geologists Asso-
lies and mass extinctions. Paleobiology, 10, 246- ciation, 112, 349-360.
267. WIGNALL, P. B. & HALLAM, A. 1992. Anoxia as a cause
SHEEHAN, P. M., COOROUGH, P. J. & FASTOVSKY, D. E. of the Permian-Triassic mass extinction: facies
1996. Biotic selectivity during the K/T and Late evidence from northern Italy and the western
Ordovician extinction events. In: RYDER, G., United States. Palaeogeography, Palaeoclimatol-
FASTOVSKY, D. & GARTNER, S. (eds) The ogy, Palaeoecology, 93, 21—46.
418 R. J. TWITCHETT & C. G. BARRAS

WIGNALL, P. B. & HALLAM, A. 1996. Fades change and WIGNALL, P. B., HALLAM, A., LAI, X. & YANG, F. 1995.
the end-Permian mass extinction event in S.E. Palaeoenvironmental changes across the Permian/
Sichuan, China. Palaios, 11, 587-596. Triassic boundary at Shangsi (N. Sichuan, China).
WIGNALL, P. B. & TWITCHETT, R. J. 1996. Oceanic Historical Biology, 10, 175-189.
anoxia and the end-Permian mass extinction. WIGNALL, P. B., MORANTE, R. & NEWTON, R. 1998.
Science, 272, 1155-1158. The Permo-Triassic transition in Spitsbergen:
WIGNALL, P. B. & TWITCHETT, R. J. 2002a. Extent, <513C0rg. chemostratigraphy, Fe and S geochemis-
duration and nature of the Permian-Triassic try, fades, fauna and trace fossils. Geological
superanoxic event. In: KOEBERL, C. & MACLEOD, Magazine, 135, 47-62.
K. G. (eds) Catastrophic Events and Mass Extinc- ZANKL, H. 1971. Upper Triassic carbonate facies in the
tions: Impacts and Beyond. Geological Society of Northern Limestone Alps. In: MULLER, G. (ed.)
America Special Papers, Boulder, Colorado, 356, Sedimentology of parts of Central Europe. Guide-
395^13. book to Excursions held during the VIII Inter-
WIGNALL, P. B. & TWITCHETT, R. J. 2002b. Facies national Sedimentological Congress 1971 in
analysis of the Latest Permian-Earliest Triassic Heidelberg, Germany. Kramer, Frankfurt am
of Jameson Land, East Greenland. Journal of the Main, 147-186.
Geological Society of London, 159, 691-703.
Ichnotaxonomy and ichnostratigraphy of chambered trace fossils in
palaeosols attributed to coleopterans, ants and termites
JORGE F. GENISE

CONICET, Museo Paleontologico Egidio Feruglio, Av. Fontana 140, 9100 (Trelew),
Chubut, Argentina (e-mail: jgenise@mef.org.ar)

Abstract: Most recorded trace fossils in palaeosols are burrows and chambers attributed to
bees, ants, termites and coleopterans. Ichnogenera attributed to bees are grouped in the ichno-
family Celliformidae, whereas those attributed to ants, termites and coleopterans are included
herein in the new ichnofamilies Pallichnidae, Krausichnidae and Coprinisphaeridae respec-
tively. Shape, type of wall, fillings and associated burrows of chambers are the main morpho-
logical ichnotaxobases used for this classification; they are weighed with regard to the
behaviour and architecture of the supposed trace-makers. Coprinisphaeridae are spherical,
pear-shaped or ovoid structures, having active or passive fillings and constructed walls. The
ichnogenera included are: Fontanai, Coprinisphaera, Eatonichnus, Monesichnus, Teisseirei and
Rebuffoichnus, attributed to coleopterans. The similar Pallichnidae show lined or structureless
walls, and include Pallichnus, Fictovichnus and Scaphichnium, also attributed to coleopterans.
The Krausichnidae constitutes trace fossils composed of chambers of different shapes inter-
connected by burrow systems of inconsistent diameter or isolated chambers associated with
burrow systems of different diameters. The Krausichnidae include Attaichnus, Parowanichnus,
Krausichnus, Archeoentomichnus, Tacuruichnus, Vondrichnus, Fleaglellius, Termitichnus and
Syntermesichnus, attributed to ants and termites. The stratigraphic ranges of insect ichnotaxa
in palaeosols are reviewed and compared with the body fossil record of potential trace-
makers, revealing that in most cases insect trace and body fossils show similar ranges. As
stated by earlier authors, the Cretaceous was a critical period during which the oldest body
fossils of dung-beetles, bees, termites and ants are recorded, whereas the trace fossils of these
groups are recorded from this period or shortly after, during Cenozoic times.

The grouping of ichnogenera in ichnofamilies is an architecture, the more difficult it is to find


uncommon but commendable trend in some analogous structures outside the insect realm,
branches of ichnology, where highly significant Probably the most similar trace fossils are those
ichnotaxobases can be used to make coherent produced by another group of arthropods:
groupings of ichnotaxa (e.g. Genise 2000; Bertling crustaceans. Tunnels associated with chambers
et al. 2003; Rindsberg & Martin 2003). Roselli are known from crayfishes, whose trace fossils
(1939), a pioneer in insect palaeoichnology, are included in the ichnogenus Camborygma
first suggested a higher taxonomy for the Hasiotis & Mitchell 1993. However, Cambor-
hymenopteran and coleopteran trace fossils that ygma shows a distinctive bioglyph composed of
he described. In his contribution he grouped scrape and scratch marks, knobby and hum-
the supposed hymenopteran nests in the mocky surfaces, pleopod and body impressions,
family 'Nidus Himenopterogenosidae* and those all of which distinguish it from similar ichno-
of coleopterans in 'Nidus coleopterogenosidae' genera attributed to social insect nests. In
(Roselli 1939). Recently, Genise (2000) created addition, interconnected tunnels of very different
the first ichnofamily for insect trace fossils in diameters are absent, as in other known modern
palaeosols, Celliformidae, to include Celliforma crustacean constructions (Bromley 1990). The
and allied ichnogenera. This contribution repre- ichnogenera Ophiomorpha and Thalassinoides,
sents an attempt to classify many of the remaining also attributed to crustaceans, commonly show
chambered trace fossils in palaeosols, attributed to burrow systems devoid of chambers (Bromley
Coleoptera, Hymenoptera and Isoptera (Genise 1990), but Verde & Martinez (2004) described
1999), and to provide a general picture of the chambers having tiny tunnels radiating vertically
stratigraphic ranges of insect ichnotaxa in palaeo- from the upper part of the chambers, in connec-
sols. It also represents an attempt to identify the tion with both ichnogenera. Spongeliomorpha
major behavioural features of these groups as shows burrow systems in association with cham-
reflected in the morphology of their trace fossils. bers in one ichnospecies: S. sicula D'Alessandro
Few trace fossils constructed by organisms & Bromley 1995. The presence of large vertical
other than insects can be compared morphologi- shafts, small chambers below the maze of tunnels
cally with insect traces. The more complex the and the typical criss-cross pattern of grooves in
From: MC!LROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 419^53. 0305-8719/04/$15.00 © The Geological Society
of London.
420 J. F. GENISE

Spongeliomorpha (D'Alessandro & Bromley 1969; Halffter & Edmonds 1982; Grasse 1984;
1995) and the upper radiating tunnels in the Holldobler & Wilson 1990). These excavated
chambers associated with Ophiomorpha and Tha- chambers are used without further modifications
lassinoides separate these trace fossils from those for nesting or pupation, and in other cases they
of social insects. are used to house more complex constructions
The morphology of insect ichnogenera devoid for nesting or pupation. The knowledge of
of tunnel systems has few similarities with other insect nest architecture was developed mostly
known ichnological structures. The specimen through entomological studies, with each group
illustrated by Hantzschel (1975) of Amphorichnus of insects having its own nomenclature for exca-
papillatus Myannil 1966 superficially resembles vated chambers and constructed structures of
Fictovichnus (Johnston et al. 1996); however, different functions. Some terms used in the ento-
the former is a filling of an amphora-like mological literature are pupation cell (e.g.
hollow ending in a distinct apical protuberance Scholtz 1988; Skelley 1991), brood cell (Stephen
(Pemberton et al. 1988; Edwards et al. 1998). et al. 1969; Batra 1984), brood ball chamber
Another case is that of the cocoons of earth- (Halffter & Matthews 1966; Halffter & Edmonds
worms and leeches described by Manum et al. 1982), fungus garden chamber (Holldobler &
(1991) attributed to the genera Burejospermum, Wilson 1990) and royal cell (Grasse 1984),
Dictyothylakos and Pilothylakos. The former among others. The functions of these chambers
two were previously considered as a seed and a are diverse, but usually they are related to nesting
palynomorph respectively. These cocoons are activities or pupation - in sum, to the successful
more likely secretions of organisms (Manum development of larvae. Fossil nests and pupation
et al. 1991) than structures resulting from their cells of Coleoptera, Hymenoptera and Isoptera
activity, and as such they are ruled out as trace respectively are the most common insect trace
fossils (Bertling et al. 2003). Also, the structure fossils in palaeosols (Genise et al. 2000), a fact
of these cocoons differs from that of insects that was related to the high potential of preserva-
(Manum et al. 1991), whose cocoons are true tion of these constructed or lined structures
trace fossils. The ichnofossil Lithoplaision (Genise & Bown 1994a).
ocalae Diblin et al. 1991 may superficially resem- Females of most solitary bees and wasps nest-
ble an insect chamber, but its conical shape and ing in soils, as well as dung-beetles, prepare
marine invertebrate remains in the wall are chambers or structures constructed inside them,
important differences from insect chambers. in which they provision food (pollen, nectar,
Continental ichnology, and particularly insect prey, carrion or vertebrate excrement), lay an
palaeoichnology, is an exciting topic that is egg, and close the entrance immediately after.
developing quickly in a changing scene, in A single larva feeds on these provisions and com-
which new discoveries occur daily. Thus this con- pletes its development without the assistance of
tribution is written with the conviction that the adults in most cases (Evans 1963; Halffter &
classification and stratigraphy of trace fossils Matthews 1966; Batra 1984). In other groups of
proposed herein will probably need to be solitary insects having recorded trace fossils in
updated in the near future. However, a first palaeosols, such as weevils (e.g. Lea 1925; John-
impression is presented herein, with the under- ston et al. 1996; Genise et al. 2002b), the larvae
standing that it will help to order the somewhat are not restricted to chambers prepared by
chaotic ichnotaxonomy of insect trace fossils, adults. Their development takes place in the
providing a new standpoint from which to soil, in which they move freely, feeding on vege-
observe and analyse insect behaviour as reflected table matter until their pupation, whereupon
by trace fossils. they prepare a cell that protects them during
this critical period before emergence as adults
(e.g. Loiacono & Marvaldi 1994). In turn,
Theoretical background social insects such as ants, termites and some
bees provision and inhabit the underground
Even the most complex insect trace fossils in nests in which they lay eggs, and rear larvae
palaeosols can be morphologically divided into that are confined to the interior of chambers
two components: burrows (tunnels, shafts (Michener 1974; Grasse 1984; Holldobler &
and galleries) and chambers. The latter term Wilson 1990).
has no specific definition in the ichnological These main behavioural differences, and a
literature and glossaries (e.g. Ekdale et al. 1984; large number of more specific ones, gave rise to
Bromley 1990). However, it is commonly used the great morphological diversity of insect nest-
to name distinct enlargements of burrows in the ing and pupation structures in soils and palaeo-
entomological literature (e.g. Stephen et al. sols. Thus the available ichnotaxobases by
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 421

which to classify ichnogenera in ichnofamilies depressed outline in cross-section (Fig. 3c). Sca-
depend on the features of chambers and asso- phichnium Bown & Kraus 1983 is unique because
ciated structures, especially shape, wall, fillings it has a peculiar hamate or lunate shape.
and burrows. Accordingly, the former ichnofam- Fossil termite and ant nests show a more or
ily proposed earlier for insect trace fossils in less continuous spectrum, from flat chambers in
palaeosols, Celliformidae Genise 2000, was Krausichnus Genise & Bown 1994b, Fleaglellius
based on various characters of cells, the usual Genise & Bown 1994b and Archeoentomichnus
name that hymenopterists give to the brood Hasiotis & Dubiel 1995a, to spherical chambers
chambers made by wasps and bees (Evans in Attaichnus Laza 1982, Termitichnus Bown
1963; Stephen et al. 1969; Michener 1974). 1982 and Vondrichnus Genise & Bown 1994b.
In most cases, other complementary characters
are needed to separate these ichnotaxa. However,
Ichnotaxobases some morphological discontinuities can be
recognized in the spectrum of shapes. Krausich-
Four ichnotaxobases, the most common charac- nus and the unnamed termite nests from the Plio-
ters used as the basis of ichnotaxa, were listed cene and Pleistocene of Africa (Coaton 1981;
and analysed by Bromley (1990): general form, Schuster et al. 2000) display low, flat, tabular
type of wall structure, type of branching, and chambers, whose floor and roof are parallel.
nature of the fill. Of these ichnotaxobases, Vertical pillars commonly accompany these
branching is a less important character for classi- chambers, probably to reinforce the whole struc-
fying insect fossil nests, than the cross morphol- ture (Fig. 5b). These tabular chambers are clearly
ogy of the entire burrow system is considered distinguishable from other flattened, but more
herein as an effective ichnotaxobase for insect oval, high chambers (e.g. Fleaglellius) in which
traces. the roof is more arched with respect to the floor
(Fig. 6b). Tabular chambers are apparently also
present in Archeoentomichnus (Hasiotis &
Shape Dubiel 1995a) and in Termitichnus namibiensis
Miller & Mason 2000. However, in the latter
Excavated and constructed chambers show a the tabular chambers seem to be more likely
morphological continuum that ranges from flat the result of a tiered arrangement of meniscate
and tabular shapes to spherical ones. Fortu- burrows, probably made by another organism.
nately, discontinuities exist within this spectrum The complex architecture of social insects may
and also some complementary characters that result in secondary chamber systems within a
favour the separation of the range as a whole primary chamber system. This is true, for
into discrete units such as ichnogenera. The instance, in Krausichnus trompitus, where the tab-
shape of the chambers is an important character ular chambers are grouped in spindle-shaped
for termite and ant nests, but it is even more cri- structures that are, in turn, connected with
tical for separating trace fossils attributed to bees other spindles (Genise & Bown 1994b). Similarly,
and beetles, because in the latter cases the asso- in Tacuruichnus, a single chambered trace fossil
ciated burrows are rare. Celliformidae (attribu- supports a boxwork with chambers in the thick
ted to bees) can be recognized by the presence peripheral wall (Genise 1997) (Fig. 5c).
of cells having rounded backs and flat or conical
tops showing a spiral design that was constructed
from the outside by the adult bee (Fig. 1). In con- Types of wall
trast, closed pupation cells attributed to beetles
have both extremes rounded and smoothed This ichnotaxobase is the most difficult to ana-
because the larvae themselves construct them lyse because of the different common usages of
from the inside. Among trace fossils attributed the term 'wall' (e.g. Retallack 200Ib) and the
to beetles, there is a clear morphological discon- complexity that walls can reach in insect traces.
tinuity between spherical or pear-shaped traces, Commonly speaking, the term 'wall' is applied
namely Coprinisphaera Sauer 1955, Fontanai indistinctly to two different structures: two-
Roselli 1939 and Pallichnus Retallack 1984, and dimensional surfaces (e.g. 'burrow boundary' of
ovoid ones: Monesichnus Roselli 1987, Fictovich- Bromley 1990, 1996) and discrete three-dimen-
nus Johnston et al. 1996, Teisseirei Roselli 1939, sional constructions. This fact introduces some
Rebuffoichnus Roselli 1987 and Eatonichnus confusion, and is discussed below. In an
Bown et al. 1997. Among the ovoid forms, it is excavated chamber the wall is the boundary
impossible to distinguish different shapes, with between the cavity and the soil - for instance
the single exception of Teisseirei, which shows a the brood ball chamber wall in dung-beetle
422 J. F. GENISE

Fig. 1. Schematic drawings to demonstrate common features in nesting structures, showing different types of
wall built by some dung-beetles, bees and termites. In all cases the insects first excavate a chamber in soil and
then construct within it one or more brood balls (dung-beetles), a cell (bees) or a complete nest (termites),
respectively. Constructed walls may show an inner lining (bees) or may bear a system of galleries and chambers
with lined walls (termites). In dung-beetles and termites, constructed structures are separated from the
excavated chamber by a space, whereas in bees the constructed wall is built against the chamber. Drawings are
based on data from Coprini (Scarabaeidae) (Halffter & Edmonds 1982), Emphorini (Apidae) (Hazeldine 1997;
Genise & Poire 2000) and Nasutitermitinae (Termitidae) (Grasse 1958; Grasse 1984).

nests (e.g. Halffter & Edmonds 1982) (Fig. 1). constructed bee cell, the wall is commonly a
In these nests one or more brood masses, each three-dimensional structure removable from the
one having its own constructed wall, may be soil. This constructed wall is in turn contained
located inside this excavated brood ball chamber within an excavated chamber with its own two-
(Halffter & Edmonds 1982) (Fig. 1). In a dimensional wall (e.g. Hazeldine 1997) (Fig. 1).
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 423

Furthermore, in some termite nests the external (1990, 1996) considered those lined walls covered
wall of the central part of the nest is a construc- by 'dust films' to be collected passively in mucus-
tion that contains a boxwork that has lined walls lined burrows. Insects actively produce similar
separated by a space (paraecie) from an exca- linings with fine material by two main methods:
vated cavity (e.g. Grasse 1958) (Fig. 1). In sum, the addition of transported material, or fluidiza-
each constructed wall in a soil produces at least tion. Among those that transport material from
three surfaces: the inner and outer surfaces of elsewhere, many species of termite usually line
the constructed structure and the surface of the the internal surfaces of constructed or excavated
bearing cavity, each surface of which may in chambers with faecal and/or regurgitated
turn be considered to be a wall. Different types material (e.g. Noirot 1970; Grasse 1984). This
of wall may be present and related in various lining is well preserved in the ichnofossil Krau-
ways within a single nest and potentially the sichnus trompitus Genise & Bown 1994b (Fig.
same trace fossil. Confusingly, in nests of differ- 5b). Some halictine bees are known to line cells
ent insect taxa the wall may bear different with clay material transported from the tunnel
names, particularly in termite nests, in which (Sakagami & Michener 1962). The clay lining
the specific nomenclature is more developed of Rosellichnus arabicus is probably derived in
(e.g. Grasse 1984). this manner (Genise & Bown 1996). Bees com-
The complexity of chamber walls in insect monly line cells with water-repellent lipids
traces is related to the necessity of maintaining (Cane 1991) after smoothing the chamber wall
very specific environmental conditions inside (Batra 1984). This smoothing behaviour triggers
nests to protect larvae and provisions. To achieve a fluidization process in the soil adjacent to the
these conditions insects commonly line or con- chamber that results in a distinct lining of fine
struct walls by adding different types of organic material sourced from the same surrounding
matter to the soil material (Michener 1979; soil (Genise & Poire 2000). The bee's movements
Grasse 1984; Cane 1991; Hanski & Cambefort against the water-saturated soil pellets of the wall
1991; Genise 1999). Genise & Bown (1994a) pro- during its construction increase the pore pres-
posed that insect fossil nests are the most sure, which in turn produces the escape of
common trace fossils in palaeosols because of water, drawing the fine material towards the
the original incorporation of organic matter in inner surface of the wall (Genise & Poire 2000).
the wall structure, which enhances the probabil- Fossil bee cells included in Celliforma Brown
ity of being consolidated by diagenetic processes. 1934, Uruguay Roselli 1939, Ellipsoideichnus
Bromley (1990, 1996), when describing the Roselli 1987; Palmiraichnus Roselli 1987 and
most common taxobases for invertebrate ichno- Cellicalichnus Genise 2000 show this type of
taxonomy, recognized seven categories of wall lining. Similar layers of clay material are found
structure: no structure (no lining), dust film, con- in the walls of the coleopteran pupation cham-
structional walling (constructional lining), zoned bers Pallichnus Retallack 1984 and Fictovichnus
fill, wall compaction, diagenetic haloes, and wall Johnston et al. 1996, who proposed that they
ornament. At least five of these are represented in were built by coleopteran larvae. Lined walls
insect architecture and have been recorded in are easily recognizable because they show a
different insect trace fossils in palaeosols. In clear discontinuity and a smooth internal sur-
two ichnogenera, both attributed to ants - face, whereas externally they intergrade gradu-
namely Attaichnus Laza 1982 and Parowanichnus ally with the host sediment (Retallack 1984;
Bown et al. 1997 - the chamber wall shows no Johnston et al. 1996). Accordingly, trace fossils
particular structure, a fact that probably reflects having lined walls are usually preserved as
the absence of linings in the walls of ant nests detached casts of smooth surfaces, or they are
(Bown et al 1997). preserved in situ in rocks, from where they can
However, most recorded insect trace fossils in be distinguished because of the different texture
palaeosols have lined or constructed walls. To and colour of the lining.
line a wall is to cover a surface (e.g. excavated Of all the categories of wall distinguished by
chamber wall) with a 'plaster' or coating (e.g. Bromley (1990, 1996), the constructed ones are
secretions, clay, faecal material), whereas to con- the most common in insect fossil nests. Usually,
struct a wall is to add 'bricks and mortar' (e.g. their producers first excavate a chamber and
soil pellets or coarse grains and fine material) then build a wall through successive addition of
to produce a discrete, three-dimensional struc- soil pellets or sand grains (e.g. Halffter & Mat-
ture (Fig. 1). A further complication in insect thews 1966; Stuart 1969; Noirot 1970; Lee &
architecture is that insects may also actively line Wood 1971; Hazeldine 1997). There are several
the inner surface of constructed walls with fine important differences between lined and con-
material (Noirot 1970; Hazeldine 1997). Bromley structed walls that arise from the particular
424 J. F. GENISE

behaviours involved. Usually, linings are thin in are generally preserved as ovoid internal casts,
comparison with constructed walls; however, whose complex surface sculpture can be inter-
intermediate thickness may produce some confu- preted as scratch marks (cast in positive relief)
sion that would preclude a clear distinction. Lin- from depressions in the chamber wall. Similar
ings are usually made from fine soil or plant casts preserving the original sculpture of the
material, secretions, or excretions that are chamber wall are known in Teisseirei barattinia
applied as a coating on an excavated or con- (Melchor et al. 2002, fig. 12 I). The constructed
structed surface (e.g. Sakagami & Michener wall of both ichnospecies of Eatonichnus shows
1962; Stephen et al 1969; Stuart 1969; Noirot a helical pattern that is present on both wall sur-
1970; Lee & Wood 1971; Cane 1991). In contrast, faces, although more clearly in the outer surface.
constructed walls are made from unsorted soil E. utahensis also exhibits a distinct superimposed
material, within which coarse grains or dry bioglyph (Bown et al. 1997) (Fig. 2d). External
faecal pellets are added one at a time and bioglyphs are more pronounced when a space is
bound by fine soil or faecal material (e.g. Halffter present between the constructed wall and the
& Matthews 1966; Stuart 1969; Noirot 1970; excavated chamber. The external ornamentation
Hazeldine 1997; Cosarinsky 2001). In lined of bee (Apidae, Emphorini) cells shows flattened
walls the insect interacts with only one (inner) pellets because the constructed wall is built
surface of the chambers, whereas in constructed against the cavity boundary (e.g. Hazeldine
walls both inner and outer surfaces result from 1997; Genise & Poire 2000) (Fig. 1). In contrast,
the behaviour of the insect. Thus the outer as the wall of Apicotermitinae (Termitidae) nests -
well as the inner surface may show a bioglyph, which are separated by a space from the bearing
as in Eatonichnus (Fig. 2d). Constructed walls excavated chambers - show a very pronounced,
are discrete and resistant structures that can be complex sculpture (e.g. Grasse 1984).
removed entirely from soils and, when preserved,
from the host rock. Accordingly, many trace
fossils bearing them are preserved not only as Fillings
casts or in situ in palaeosols, but also, and more
frequently, as complete structures, removed Filling material and structure is another impor-
from the rock matrix, showing internal and tant taxobase in invertebrate ichnotaxonomy.
external bioglyphs. In addition, insect trace fos- Passive fill enters a burrow gravitationally,
sils having constructed walls can be transported whereas active fillings are emplaced by the
and re-deposited (e.g. Andreis 1972, 1981) in trace-maker (Bromley 1990, 1996). Both kinds
contrast to most trace fossils, which are generally of fill occur in insect trace fossils. As described
in situ (e.g. Ekdale et al. 1984). These walls are previously, adult insects provision their nests
characteristic of Palmiraichnus Roselli 1987, with different kinds of organic matter to rear
Rosellichnus Genise & Bown 1996, Uruguay their larvae. Some dung-beetles are known to
Roselli 1939, Coprinisphaera Sauer 1955, Fonta- arrange the provision of their brood masses in
nai Roselli 1939, Monesichnus Roselli 1987, Teis- a meniscate pattern (Halffter & Matthews 1966;
seirei Roselli 1939, Rebuffoichnus Roselli 1987, Halffter & Edmonds 1982) (Fig. 3a). Such menis-
Eatonichnus Bown et al. 1997, Termitichnus cate fills are seen in Coprinisphaera, Monesichnus,
Bown 1982, Fleaglellius Genise & Bown 1994b, Eatonichnus and Scaphichnium (Bown & Kraus
Vondrichnus Genise & Bown 1994b, Tacuruich- 1983; Hasiotis et al. 1993; Bown et al. 1997;
nus Genise 1997 and Archeoentomichnus Hasiotis Genise & Laza, 1998; Duringer et al. 2000a,
& Dubiel 1995a. 2000b) (Figs 2d, 3a). These authors attributed
Bromley (1990, 1996) mentioned diagenetic their material to dung-beetles based on the
haloes as a particular category of wall lacking meniscate structure of the fills. The absence of
ichnotaxonomical value. Hasiotis et al. (1993) an active fill in Rebujfoichnus, Pallichnus, Ficto-
interpreted the external coating of Scaphichnium vichnus and Teisseirei is concomitantly taken to
hamatum as diagenetic cement, but no particular suggest that these ichnotaxa represent coleop-
wall was described for this ichnofossil (Bown & teran pupation cells (Retallack 1984; Johnston
Kraus 1983; Hasiotis et al. 1993). The last cate- et al. 1996; Genise et al. 2002a; Melchor et al.
gory mentioned by Bromley (1990, 1996) - wall 2002). Despite the fact that bees, ants and
ornamentation - seems to be more probably a termites also provision their nests actively (e.g.
wall feature than a type of wall in itself. Orna- Stephen et al. 1969; Grasse 1984; Holldobler &
mentation may, potentially, be present in several Wilson 1990), their trace fossils show no recog-
different types of wall. Edwards et al. (1998) nizable active fill (e.g. Laza 1982; Genise &
described unnamed trace fossils from the Bern- Bown 1994b; Hasiotis & Dubiel 1995a; Bown
bridge Limestone Formation (England), which et al. 1997; Genise 2000).
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 425

Burrow system Brown 1934; Uruguay Roselli 1939; Palmiraich-


nus Roselli 1987; Ellipsoideichnus Roselli 1987;
Burrows associated with fossil bee cells are Rosellichnus Genise & Bown 1996; Corimbatich-
uncommon (e.g. Cellicalichnus and Ellipsoideich- nus Genise & Verde 2000; and Cellicalichnus
nus) (Genise 2000), and the same is true for Genise 2000. These ichnogenera can be recog-
coleopteran trace fossils. The coleopteran trace nized by the presence of cells having rounded
fossil Pallichnus is the single exception: it shows bases and flat tops, which commonly bear a
an associated burrow system, albeit only obser- spiral closure. Cells may be isolated or clustered,
vable microscopically (Retallack 1984). The and may be connected by tunnels of similar dia-
poor preservation of burrows in fossil nests of meters.
solitary insects may be a result of the absence The second group comprises the new ichno-
of constructed or lined burrow walls, in contrast family Coprinisphaeridae introduced herein,
to those of brood and pupation cells (Genise & and includes unnamed trace fossils and spherical,
Bown 1994a). pear-shaped or ovoid ichnogenera with active or
The burrows within nests of solitary insects passive fill and constructed walls. Ichnogenera
have a constant diameter that corresponds to included are: Fontanai Roselli 1939; Teisseirei
the body diameter of their constructor, as in Roselli 1939; Coprinisphaera Sauer 1955; Mone-
other invertebrate trace fossils (Ekdale et al sichnus Roselli 1987; Rebuffoichnus Roselli
1984). The situation is different in ant and ter- 1987; and Eatonichnus Bown et al. 1997. These
mite nests, whose burrow systems result from ichnogenera are interpreted as the brood
the cooperative work of many. Burrows are com- masses of dung-beetles or pupation chambers
monly very complex, and burrow diameter is of various coleopteran taxa.
variable. In many cases, larger burrows (first The third group, also representing trace fossils
order) are connected with medium-sized ones attributed to pupation chambers and brood
(second order) and smaller ones (third order), masses of beetles, consists of ichnofossils lacking
the latter representing individual passages that a constructed wall. Such ichnotaxa are included
match the size of the workers (Grasse 1984; in the new ichnofamily Pallichnidae, and include:
Sands 1987; Genise & Bown 1994b) (Fig. 6c, Scaphichnium Bown & Kraus 1983; Pallichnus
d). Even when individual chambers are similar Retallack 1984; and Fictovichnus Johnston et al.
to those constructed by coleopterans, the asso- 1996. Pallichnus shows two features that distin-
ciated burrow systems clearly distinguish the guish it from somewhat similar coprinisphaerid
constructions of social from solitary insects. ichnogenera. The Pallichnidae lack a discrete
The burrow systems may: constructed wall. This fact suggested to Retal-
arise from the base of isolated chambers (e.g. lack (1984) that these trace fossils would prob-
Tacuruichnus)\ ably represent pupation cells of Geotrupinae or
be present in the wall or in the interior of Scarabaeinae rather than brood masses. Also,
chambers (e.g. Tacuruichnus, Termitichnus the chambers are connected to lateral tunnels
qatranii); or that radiate from a vertical shaft, another charac-
connect different chambers (e.g. Krausichnus, ter that is absent in Coprinisphaeridae. Fictovich-
Termitichnus, Vondrichnus, Fleaglellius, Arche- nus resembles Pallichnus in having a lined wall
oentomichnus) (Genise & Bown 1994b; Hasio- defined on its inner surface by a clay layer (Retal-
tis & Dubiel 1995a; Genise 1997). lack 1984; Johnston et al. 1996). Fictovichnus has
also been attributed to pupation chambers of
This ichnotaxobase distinguishes ant and termite beetles (Johnston et al. 1996). Scaphichnium
nests from others and enables them to be classi- lacks a lined or constructed wall, and its charac-
fied together in a suprageneric group. teristic hamate or lunate shape with a bulbous
termination is very different from both Pallichnus
and Fictovichnus. Its inclusion in Pallichnidae is
Ichnofamilies of palaeosol trace fossils tentative. The meniscate filling of Scaphichnium
attributed to insects probably reflects provisioning organic matter
inside the excavated chambers for rearing
Insect trace fossils in palaeosols can be grouped larvae, as seen in modern examples of the Geo-
into four morphological groups or ichnofamilies trupinae (Hasiotis et al. 1993).
based on the ichnotaxobases described above. Some internal casts of Coprinisphaeridae may
The first ichnogeneric group comprises the ich- be indistinguishable from Pallichnidae. Such is
nofamily Celliformidae, erected by Genise the case for internal casts of Rebuffoichnus
(2000), and includes the following ichnogenera, resembling Fictovichnus (Fig. 3e), as well as inter-
all of which are attributed to bees: Celliforma nal casts of Coprinisphaera, which may resemble
426 J. F. GENISE

Key to separate ichnofamilies of chambered trace fossils in palaeosols


1 Trace fossils composed of cells having rounded bases and flat tops that commonly
bear a spiral closure Celliformidae Genise
Trace fossils showing another combination of characters 2
2 Trace fossils composed of: (a) chambers of different shapes interconnected by
a boxwork showing burrows of different diameters; or (b) isolated chambers
bearing an internal boxwork; and/or (c) chambers from which a burrow
system radiates. Intersecting grooves, scratch and scrape markings absent Krausichnidae ifam. nov.
Trace fossils showing another combination of characters 3
3 Trace fossils composed of isolated or clustered, spherical, pear-shaped
or ovoid chambers, surrounded by a discrete, constructed wall Coprinisphaeridae ifam. nov.
Trace fossils composed of spherical, ovoid, hamate, or lunate chambers
lacking a constructed wall Pallichnidae ifam. nov.

Pallichnus. However, internal casts are com- the interconnecting tunnels were constructed by
monly associated with complete specimens in different trace-makers (Duringer et al. 2000b).
outcrop, leaving no doubt as to the identity of Later, the same authors recognized termites as
the trace fossils (e.g. Genise et al. 2002a). possible constructors of these tunnel systems
Ovoid trace fossils included in the Coprini- (Duringer et al. in press). The traces are thus a
sphaeridae and Pallichnidae may commonly composite trace fossil (sensu Pickerill 1994) in
show a broken end, or may be perforated or which the termites exploit food reserves intended
flattened following the emergence of the adult for larvae of Coleoptera.
insect. In these cases, specimens may take a
celliformid-like shape (e.g. Edwards et al.
1998). However, the flattened end lacks the Systematic ichnology
spiral design as of well-preserved Celliformidae.
The fourth group, the new ichnofamily Krau- Ichnofamily Coprinisphaeridae ifam. nov.
sichnidae, consists of trace fossils composed
either of chambers of different shapes intercon- Type ichnogenus. Coprinisphaera Sauer 1955.
nected by burrow systems of different diameters, Diagnosis. Trace fossils consist of spherical,
or of isolated chambers associated with burrow subspherical, pear-shaped, ovoid, or sub-ovoid
systems of different diameters. Ichnogenera chambers, generally isolated, rarely clustered.
include: Attaichnus Laza 1982; Termitichnus Chambers are surrounded by a discrete con-
Bown 1982; Syntermesichnus Bown & Laza structed wall, which may show a circular or
1990; Krausichnus Genise & Bown 1994b; Flea- ovoid hole. Some ichnogenera show empty or
glellius Genise & Bown 1994b; Vondrichnus passively filled chambers, whereas in others
Genise & Bown 1994b; Archeoentomichnus active infill is the norm.
Hasiotis & Dubiel 1995a; Parowanichnus Bown
et al. 1997; and Tacuruichnus Genise 1997. Ichnogenus Fontanai Roselli 1939 (Fig. 2c)
These ichnogenera are attributed to the work of v*1939 Fontanai Roselli p. 79, figs 23, 24, 31(8).
social insects, namely ants and termites. The p. 1975 'Nidos fosiles petreos de Coleopteros'
presence of tunnels of different diameters in the Francis p. 553.
same structure is the distinctive feature for recog- 1976 Fontanaichnus Roselli p. 167 (junior
nition of traces constructed by the cooperative synonym),
work of specialized types of individual (of differ- p. 1981 'Nidos fosiles de Coleopteros'
ing sizes) within a society (Genise 1997). Sprechmann, Bossi and Da Silva p. 266.
A recently described but unnamed trace fossil 1982 Fontanaichnus Roselli; Martinez p. 58.
from the Pleistocene of Chad (Duringer et al. v!987 Fontanaichnus Roselli; Roselli p. 34, pi. II,
2000a, 2000b) deserves further comment. The fig. 6; pi. Ill, fig. 5.
trace fossil represents several specimens of Copri- p. 1988 'Nidos de insectos fosiles (coleopteros)'
nisphaera connected by a tunnel system. As such, Ford p. 47.
it could be seen as a link between Coprinisphaer- 1990a Fontanaichnus Roselli; Retallack, p. 219,
idae and Krausichnidae that would almost pre- figs 201 A,B.
clude their separation. Such structures are 1993 Fontanai Roselli; Genise p. 53.
unknown among dung-beetle traces, a fact that 1994 Fontanaichnus Roselli; Donovan p. 209, figs
originally suggested to Duringer et al. (2000a, 8.5 A, B.
2000b) that specimens of Coprinisphaera and 1994a Fontanai Roselli; Genise & Bown p. 112.
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 427

Key to ichnogenera within Coprinisphaeridae


1 Spherical, subspherical or pear-shaped chambers 2
Ovoid or sub-ovoid chambers 3
2 Spherical chambers showing a raised rim surrounding the emergence hole Fontanai Roselli
Spherical, subspherical or pear-shaped chambers lacking a raised rim to the
aperture Coprinisphaera Sauer
3 Ovoid structures, external wall showing a helical design Eatonichnus Bown et al.
Ovoid structures showing smooth or rugose outer wall 4
4 Chambers showing meniscateMonesichnus Roselli
Empty or passively filled chambers 5
5 Chambers having a depressed outline in cross-section; inner surface may show
scratch marks; some specimens show a short tunnel at the entrance or
antechamber Teisseirei Roselli
Chambers having rounded outline in cross-section Rebuffoichnus Roselli

1996 Fontanai Roselli; Johnston, Eberth & v!941 'Nidos de escarabeidos' Frenguelli p. 87.
Anderson p. 522. 1950 'Bolas de Cangagua' Bruet p. 280, pi. I, figs
1998 Fontanai Roselli; Genise & Laza p. 213. 2,3.
1998 Fontanaichnus Roselli; Buatois, Mangano, *1955 Coprinisphaera Sauer p. 123, figs 1-5.
Genise & Taylor p. 226. 1955 Cangabola Lengerken p. 937, figs 6-8.
2000 Fontanai Roselli; Genise, Mangano, 1956 Coprinisphaera Sauer; Sauer p. 550, figs 1—4.
Buatois, Laza & Verde p. 54. 1959 'Nidos de Scarabaeidae' Halffter p. 174.
2000 Fontanai Roselli; Krell p. 891. 1959 Coprinisphaera Sauer; Sauer p. 119.
2002 Fontanaichnus Roselli; Buatois, Mangano 1962 Coprinisphaera Sauer; Hantzschel W189.
& Acenolaza p. 189. 1966 'Fossil scarab balls' Halffter & Matthews
Type and only known ichnospecies. Fontanai p. 154.
kraglievichi Roselli 1939. 1966 'Nidos fosiles de escarabeidos' Camacho
Diagnosis. Spherical chambers having a thick p. 490, pi. XVI figs n, o.
constructed wall and an emergence hole sur- 1972 'Nidos de escarabajos' Andreis p. 91.
rounded by a raised rim or neck. p. 1975 'Nidos fosiles petreos de Coleopteros'
Remarks. This ichnogenus is pending the Francis p. 553.
ichnotaxonomic revision along with the closely 1975 Coprinisphaera Sauer; Hantzschel p. W52.
related Coprinisphaera (Laza, personal commu- 1976 Devincenzichnus Roselli p. 167 (junior
nication). Possible trace-makers are dung- synonym).
beetles (Scarabaeinae). The presence of a neck 1977 'Paleonidos de escarabeidos' Spalletti &
is the only feature that allows the separation Mazzoni p. 267.
of Fontanai from Coprinisphaera. The neck 1981 'Nidos de escarabeidos' Pascual & Bondesio
may be interpreted as the remains of a separate p. 125.
egg chamber that some dung-beetles construct 1981 'Nidos de escarabajos' Andreis p. 34.
over the provision chamber in their brood p. 1981 'Nidos fosiles de Coleopteros'
masses (Halffter & Matthews 1966). However, Sprechmann, Bossi & Da Silva p. 266.
if the neck is interpreted as the remains of a 1982 'Nidos de escarabeidos' Alonso, Gonzalez &
former egg chamber, it would be possible to Pelayes p. 2.
trace a continuous morphological series between 1982 Coprinisphaera Sauer; Martinez p. 48.
both ichnogenera. 1982 Devincenzichnus Roselli; Martinez p. 48.
1986a 'Icnofosiles de Scarabaeinae' Laza p. 13.
Ichnogenus Coprinisphaera Sauer 1955 1986b 'Nidos de Scarabaeinae' Laza p. 19.
(Fig. 2a, b) 1987 'Heliocopris dung ball' Sands p. 423.
v!938a 'Bolas de escarabeidos' Frenguelli p. 348. v!987 Devizenzichnus Roselli, Roselli p. 21, pi. I,
v!938b 'Pallottole di Scarabeidi' Frenguelli p. 77, fig. 1.
fig. 5; pi. VII, figs 1-8. *v!987 Martinezichnus Roselli p. 22, pi. I, figs 2,
v!939 Devincenzia Roselli p. 81, figs 26, 27, 28 2a (new synonym).
and 31 (5-6) (non Kraglievich 1932). *v!987 Madinaichnus Roselli p. 23, pi. I, fig. 3
v!939a 'Nidos fosiles de Escarabeidos' Frenguelli (new synonym).
p. 270. *v!987 Microicoichnus Roselli p. 49, pi. I, fig. 8
v!939b 'Nidos fosiles de Escarabeidos' Frenguelli (new synonym).
p. 379, figs 4^-9, pis I-II. p. 1988 'Nidos de coleopteros' Ford p. 47.
v!940 'Nidos fosiles de Escarabeidos' Frenguelli v!990 'Nidos de escarabeidos' Laza & Reguero
p. 70. p. 245.
428 J. F. GENISE

Fig. 2. Coprinisphaeridae. (a) Coprinisphaera ecuadoriensis Sauer 1955: thick constructed wall, empty chamber,
and emergence hole. Eocene-Miocene Sarmiento Formation, Argentina; MACN-LI1802. Bar: 1 cm. (MACN-
LI, Museo Argentine de Ciencias Naturales, Laboratorio de Icnologia.) (b) Holotypes of Devincenzichnus
murguiai Roselli 1939 (left) (MFLR 479), Martinezichnus francisi Roselli 1987 (centre) (MFLR607), and
Madinaichnus larranagai Roselli 1987 (right) (MFLR 641). Note the almost continuous range of ball and hole
sizes. Late Cretaceous-Early Tertiary Asencio Formation, Uruguay. Bar: 1 cm. (MFLR: Museo Francisco
Lucas Roselli, Nueva Palmira, Uruguay.) (c) Fontanai kraglievichi Roselli 1939: raised rim around the
emergence hole. Late Cretaceous-early Tertiary Asencio Formation, Uruguay; MACN-LI1786. Bar: 1 cm.
(d) Eatonichnus utahensis (Guilliland and La Rocque, 1952). Holotype: external bioglyph and internal chamber
with meniscate filling. Palaeocene Colter Formation, USA; OM20201. Bar: 1 cm. (Photograph courtesy of
T. Bown.) (OM: Ohio State University Orton Geological Museum.)

1990a Coprinisphaera Sauer; Retallack p. 219, 1990b 'Dung beetle trace fossils' Retallack
fig. 201 G-I. p. 436.
1990a Devincenzichnus Roselli; Retallack p. 219, 1991 Coprinisphaera Sauer; Retallack p. 182.
fig. 201 C, D. 1993 Coprinisphaera Sauer; Genise p. 50.
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 429

1993 Devincenzichnus Roselli; Genise p. 50. 2000b 'Boules de bousiers fossiles' Duringer,
1994 Coprinisphaera Sauer; Donovan p. 209, fig. Brunet, Cambefort, Beauvilain, Mackaye,
8.5 G-I. Vignaud & Schuster p. 259, figs 1-10.
1994 Devincenzichnus Roselli; Donovan p. 209, 2000 'Boules-nids fossiles de bousiers' Schuster,
fig. 8.5 C, D. Duringer, Nel, Brunet, Vignaud & Mackaye
1994a Coprinisphaera Sauer; Genise & Bown, p. 17.
p. 109, figs 3, 4. 200la Coprinisphaera Sauer; Retallack p. 142.
1995 Coprinisphaera Sauer; Genise & Cladera 200la Devincenzichnus Roselli; Retallack p. 142.
p. 78, figs IE, Fig. IF, left and right. 200la Martinezichnus Roselli; Retallack p. 142.
1995 Martinezichnus Roselli; Genise & Cladera 200la Madinaichnus Roselli; Retallack p. 142.
p. 78, figs IE, Fig. IF, centre. 2002 Martinezichnus Roselli; Buatois, Mangano
1995 'Nidos de escarabajos' Fontaine; Ballesteros & Acenolaza p. 22.
& Powell p. 12. 2002 Microicoichnus Roselli; Buatois, Mangano
1996 'Nidos de escarabajos' Iriondo & Krohling & Acenolaza p. 189.
p. 43. 2002 Madinaichnus Roselli; Buatois, Mangano &
1996 Devincenzichnus Roselli; Veroslavsky & Acenolaza p. 189.
Martinez p. 41, pi. I, fig. 5. 2002 Devincenzichnus Roselli; Buatois, Mangano
1996 Devincenzichnus Roselli; Johnston, Eberth & Acenolaza p. 189.
& Anderson p. 522. 2002 Coprinisphaera Sauer. Buatois, Mangano &
1996 Coprinisphaera Sauer; Johnston, Eberth & Acenolaza p. 189.
Anderson p. 522. Type ichnospecies. Coprinisphaera ecuadoriensis
1998 Devincenzichnus Roselli; Buatois, Sauer 1955.
Mangano, Genise & Taylor p. 226. Diagnosis. Spherical, subspherical and pear-
1998 Martinezichnus Roselli; Buatois, Mangano, shaped chambers having a constructed wall.
Genise & Taylor p. 226. Some specimens may show a hole (possibly an
1998 Madinaichnus Roselli; Buatois, Mangano, emergence hole). Internal cavities empty or
Genise & Taylor p. 226. containing a meniscate or passive fill.
1998 Microicoichnus Roselli; Buatois, Mangano, Included ichnospecies. Coprinisphaera murgiai
Genise & Taylor p. 226. Roselli 1939; C. francisi Roselli 1987; C.
1998 Coprinisphaera Sauer; Buatois, Mangano, larranagai Roselli 1987; C. lafurcadai Roselli
Genise & Taylor p. 227. 1987; C. ecuadoriensis Sauer 1955; C. frenguellii
1998 Microicoichnus Roselli; Genise & Laza Genise & Bown 1994a.
p. 213. Remarks. Coprinisphaera is one of the most
1998 Madinaichnus Roselli; Genise & Laza p. 213. common trace fossils in the Tertiary palaeosols
1998 Martinezichnus Roselli; Genise & Laza of South America, and it was appropriately one
p. 213. of the first recorded insect trace fossils (Fren-
1998 Devincenzichnus Roselli; Genise & Laza guelli 1938a; Roselli 1939). It has subsequently
p. 213. been described from different localities and ages
1998 Coprinisphaera Sauer; Genise & Laza p. 220. of South and Central America, Asia, Antarctica
1999 Coprinisphaera Sauer; Genise p. 110. and Africa (Genise et al 2000; Krell 2000 and
1999 Martinezichnus Roselli; Genise p. 110. references therein). The mention of 'nidos de
1999 Madinaichnus Roselli; Genise p. 110. escarabajos o escarabeidos' [nests of dung-beetles
1999 Microicoichnus Roselli; Genise p. 110. or scarabs] is common in the Argentinean
2000 Coprinisphaera Sauer; Genise, Mangano, sedimentological and palaeontological literature
Buatois, Laza & Verde p. 54. (e.g. Andreis 1972, 1981; Spalletti & Mazzoni
2000 Martinezichnus Roselli; Genise, Mangano, 1977; Pascual & Bondesio 1981; Alonso et al.
Buatois, Laza & Verde p. 54. 1982; Laza 1986a, 1986b; Laza & Reguero
2000 Madinaichnus Roselli; Genise, Mangano, 1990; Fontaine et al. 1995; Iriondo & Krohling
Buatois, Laza & Verde p. 54. 1996). In these cases, it is clear that the authors
2000 Microicoichnus Roselli; Genise, Mangano, recognized the typical spherical or pear-shaped
Buatois, Laza & Verde p. 54. forms described by Frenguelli (1938b, 1939b),
2000 Coprinisphaera Sauer; Krell p. 890. and as such these occurrences are attributed
2000 Microicoichnus Roselli; Krell p. 891. herein to Coprinisphaera ispp. The trace fossils
2000 Madinaichnus Roselli; Krell p. 891. described by Frenguelli (1938b, 1939b) are the
2000 Martinezichnus Roselli; Krell p. 891. same as those recorded by HahTter (1959), Cama-
2000a 'Fossil dung-beetle brood ball' Duringer, cho (1966) and Halffter & Matthews (1966). In
Brunet, Cambefort, Likius, Mackaye, turn, Uruguayan sedimentologists (e.g. Francis
Schuster & Vignaud p. 277, figs 3-6. 1975; Sprechmann et al. 1981; Ford 1988)
430 J. F. GENISE

mentioned coleopteran nests described by Roselli group. Possible trace-makers are dung-beetles
(1939) which may be assigned indeterminately to (Scarabaeinae) (Bown et al. 1997). The affinity
Coprinisphaera or Fontanai. with dung-beetle brood masses is suggested by
Roselli (1987) created Martinezichnus, Madi- its shape and meniscate fillings, similar to the ich-
naichnus and Microicoichnus based on the nogenus Monesichnus. In addition, some modern
different sizes of the chambers and the sizes of dung-beetles are known to excavate helical tun-
emergence holes, but these ichnotaxobases nels (Bown et al. 1997). However, the producer
show significant overlap, precluding any clear- of this trace fossil is unknown because there are
cut distinction between the different ichnogenera no modern analogues for this structure.
(Fig. 2b). They are formally considered herein as
synonyms following Genise (1999). The probable Ichnogenus Monesichnus Roselli 1987 (Fig. 3a)
trace-makers are dung-beetles (Scarabaeinae). v*1987 Monesichnus Roselli p. 39, pi. I, fig. 7.
v!994 Monesichnus Roselli; Laza, Genise &
Ichnogenus Eatonichnus Bown et al. 1997 Bown p. 397.
(Fig. 2d) 1997 Monesichnus Roselli; Bown, Hasiotis,
v*1997 Eatonichnus Bown, Hasiotis, Genise, Genise, Maldonado & Browers p. 52.
Maldonado & Browers p. 52, figs 8C-F, 9. 1998 Monesichnus Roselli; Buatois, Mangano,
1998 Eatonichnus Bown, Hasiotis, Genise, Genise & Taylor p. 226.
Maldonado and Browers; Genise & Laza v!998 Monesichnus Roselli; Genise and Laza
p. 214. p. 213, figs 3-5.
1999 Eatonichnus Bown, Hasiotis, Genise, 1999 Monesichnus Roselli; Genise p. 110.
Maldonado & Browers; Genise p. 111. 2000 Monesichnus Roselli; Genise, Mangano,
2000 Eatonichnus Bown, Hasiotis, Genise, Buatois, Laza & Verde p. 54.
Maldonado & Browers; Genise, Mangano, 2000 Monesichnus Roselli; Krell p. 891.
Buatois, Laza & Verde p. 54. 2001 Monesichnus Roselli; Retallack p. 142.
2000 Eatonichnus Bown, Hasiotis, Genise, 2002 Monesichnus Roselli; Buatois, Mangano &
Maldonado & Browers; Krell p. 891. Acenolaza p. 189.
v2001 Eatonichnus Bown, Hasiotis, Genise, Type and only known ichnospecies. Monesichnus
Maldonado & Browers; Genise, Cladera & ameghinoi Roselli 1987.
Tancoff p. 45. Diagnosis. Discrete structures, fusiform to
Type ichnospecies. Eatonichnus utahensis ovate, composed of a constructed, unpatterned
(Gilliland & La Rocque 1952). wall (sometimes showing a longitudinal
Diagnosis. Trace fossils composed of closely groove), and an internal cavity that in some
appressed whorls, tightly spiralled around a cen- cases is empty and in others exhibits a meniscate
tral cylindrical cavity and converging terminally, fill (Genise & Laza 1998).
thereby forming a closed, spindle-shaped helix. Remarks. Monesichnus is morphologically simi-
Helices may be either sinistral or dextral, with lar to Eatonichnus, but its unpatterned wall clearly
whorls inclined away from the transverse axis. distinguishes it from the latter. Its meniscate fill,
Whorl diameter constant along the helix and interpreted as the provisions of a dung-beetle
invariably greater than the diameter of the cen- brood mass (Fig. 3a), distinguishes it from other
tral cavity. Central cavity fill closely packed, ichnogenera such as Teisseirei andebuffoichnus,
with or without meniscae (Bown et al. 1997). which are interpreted as pupation chambers.
Included ichnospecies. E. utahensis (Gilliland Some broken specimens of Monesichnus may
& La Rocque 1952); E. claronensis Bown et al. also be empty, their fills probably lost by weather-
1997. ing. Even in these cases, the rounded cross-section
Remarks. The helical design of the outer wall of the chamber distinguishes it from Teisseirei,
is unique in Coprinisphaeridae, but its ovoid and the absence of a rounded emergence hole pre-
shape and meniscate fillings relate it to the unpat- cludes its assignment to Rebuffoichnus.These
terned Monesichnus. The distinction between the structures are similar to brood masses of the
two named ichnospecies of Eatonichnus is based representatives of the modern dung-beetle
on their very dissimilar sizes, whorl inclination, genera Dichotomius, Gromphas and Oruscatus
cavity fill and external ornamentation. A third, (Scarabaeinae) (Genise & Laza 1998).
unnamed, ichnospecies has also been described
based on its larger size and external bioglyph Ichnogenus Teisseirei Roselli 1939 (Fig. 3b, c)
(Bown et al. 1997). Later, Genise et al. (2001) v*1939 Teisseirei Roselli p. 82, figs. 29, 30, 31(7).
recorded Argentinean specimens of E. claronen- 1976 Tesseirichnus Roselli p. 167 (junior
sis that show intermediate sizes, suggesting that synonym).
size may be a misleading ichnotaxobase for this 1982 Tesseirichnus Roselli; Martinez p. 61.
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 431

Fig. 3. Coprinisphaeridae. (a) Monesichnus ameghinoi Roselli 1987 (left): wall and meniscate filling. Late
Cretaceous-Early Tertiary Asencio Formation, Uruguay; MACN-LI233. A sectioned sample of a brood mass
of the dung-beetle Oruscatus davus (Scarabaeinae) (right) showing the same type of wall and fillings. Bar: 1 cm.
(b) Teisseirei barattinia Roselli 1939: antechamber and bioglyph in the chamber wall; included in a piece of
rock matrix. Late Cretaceous-Early Tertiary Asencio Formation, Uruguay; MACN-LI876. Bar: 1 cm.
(c) T. barattinia Roselli 1939: cross-section showing the constructed wall and the depressed outline of the
chamber. Eocene-Miocene Sarmiento Formation, Argentina; MPEF-IC 253. Bar: 1 cm. (MPEF-IC: Museo
Paleontologico Egidio Feruglio-Icnologia). (d) Rebuffoichnus casamiquelai Roselli 1987. Late Cretaceous
Laguna Palacios Formation, Argentina; MACN-LI1202. Bar: 1 cm. (e) R. casamiquelai: internal cast with
remains of the outer wall. Late Cretaceous Laguna Palacios Formation, Argentina; MACN-LI1181. Bar: 1 cm.

v!987 TeisserichnusRoselli p. 24, pi. I, fig. 5 1996 Teisseirei Roselli; Veroslavsky and
(lapsus). Martinez p. 41, pi. I, fig. 4.
v!987 Isociesichnus Roselli p. 38, pi. II, fig. 5 1998 Teisseirei Roselli; Buatois, Mangano,
(new synonym). Genise & Taylor p. 226.
1990a Teisseirichnus Roselli; Retallack p. 219. 1998 Isociesichnus Roselli; Buatois, Mangano,
1993 Teisseirei Roselli; Genise p. 53. Genise & Taylor p. 226.
432 J. F. GENISE

1998 Teisseirei Roselli; Genise & Laza p. 213. v!999 Rebuffoichnus Roselli; Genise, Sciutto,
1998 Isociesichnus Roselli; Genise & Laza p. 213. Laza, Gonzalez & Bellosi p. 29.
2000 Teisseirei Roselli; Genise, Mangano, 2000 Rebuffoichnus Roselli; Krell p. 892.
Buatois, Laza & Verde p. 54. 2000 Rebuffoichnus Roselli; Genise, Mangano,
v2001 Teisseirei Roselli; Genise & Zelich p. 44. Buatois, Laza & Verde p. 54.
2001 Teisseirei Roselli; Retallack p. 142. 200la Rebuffoichnus Roselli; Retallack p. 142.
v2002 Teisseirei Roselli; Melchor, Genise & v2002 Rebuffoichnus Roselli; Genise, Sciutto,
Miquel p. 35, fig. 12 A-E, I. Laza, Gonzalez & Bellosi p. 230, figs 5D, 7.
Type and only known ichnospecies. Teisseirei 2002 Rebuffoichnus Roselli; Buatois, Mangano &
barattinia Roselli 1939. Acenolaza p. 189.
Diagnosis. Depressed chambers slightly v2002 RebuffoichnuRoselli: Genise, Laza,
arched downwards and having constructed Fernandez & Frogoni p. 160.
walls. Some specimens may show a small, Type and only known ichnospecies.
rounded antechamber. Inner surface of the wall Rebuffoichnus casamiquelai Roselli 1987.
displays, in well-preserved specimens, a distinct Diagnosis. Sub-ovoid to subcylindrical struc-
lining bearing small elliptical scratches oriented tures composed of a wall, whose exterior aspect
mostly longitudinally (modified from Melchor is rugose and lumpy, whereas the interior is
et al 2002). smooth or showing a faint bioglyph. The internal
Remarks. The internal bioglyph (Fig. 3b) and chamber is ovoid and has a circular cross-sec-
the depressed cross-section (Fig. 3c) distinguish tion. The wall may be perforated by a rounded
this ichnogenus from other Coprinisphaeridae. hole (Genise et al. 2002b).
These trace fossils commonly show three preser- Remarks. Rebuffoichnusdiffers from other
vational morphologies: (1) empty or passively more or less ovoid insect trace fossils, such as
filled chambers found in situ in palaeosols; (2) Monesichnus, in lacking any active fill and in
isolated, detached clasts composed of the the presence of a rounded hole. Trace fossils
chamber fillings; or (3) empty or passively filled from the Quaternary of Australia described by
chambers surrounded by a thick wall. Specimens Lea (1925) and Read (1974) were assigned to
found in situ in palaeosols of the Asencio Forma- an unnamed ichnospecies of Fictovichnus by
tion suggested that these chambers were merely Johnston et al. (1996). However, the type ichno-
excavated structures (Genise 1999). However, species of Fictovichnus lacks a constructed wall as
the recent finding of hundreds of specimens bear- seen in the material of Lea (1925) and Read
ing constructed walls in the Tertiary of Patagonia (1974). These specimens have a general aspect
and re-examination of previous discoveries sug- and similar features to those of Rebuffoichnus
gests that Teisseirei barattinia has a constructed casamiquelai (Genise et al. 2002b). R. casamique-
wall. The holotype (and only documented lai is attributed to pupation chambers of
specimen) of Isociesichnus diplocamara Roselli Curculionidae, Scarabaeidae, or Tenebrionidae
1987 is actually a structure composed of two (Johnston et al. 1996). However, species of the
attached specimens of Teisseirei barattinia. Curculionidae are the most likely constructors
Thus it is regarded herein as a junior synonym because of the circular body outline, which
of Teisseirei. Despite the excellent preservation would accord with the circular holes in the
of many specimens that display an internal bio- trace fossils (Genise et al. 2002b). This assump-
glyph showing the scratches of the constructors, tion is also confirmed by the discovery of the con-
the origin of Teisseirei is unknown. Its shape structor within a Rebuffoichnufrom Australia
and the absence of active fill indicate the lack (Lea 1925).
of original provisions, and suggest that it is prob-
ably a coleopteran pupation chamber.
Ichnofamily Pallichnidae ifam. nov.
Ichnogenus Rebuffoichnus Roselli 1987 (Fig. 3d, e)
1925 'Calcareous insect puparia' Lea p. 35, pi. I, Type ichnogenus. Pallichnus Retallack 1984.
figs1-20. Diagnosis. Group of ichnogenera comprising
v* 1987 Rebuffoichnus Roselli p. 24, pi. I, fig. 4; pi. subspherical, ovoid, hamate or lunate chambers
Ill, fig. 4. lacking a constructed wall. Chambers are sur-
p. 1996 Fictovichnus Johnston, Eberth and rounded by linings or diagenetic haloes. Fillings
Anderson p. 516, figs 2, 3C-D. may be passive or meniscate.
1998 RebuffoichnusRoselli; Genise and Laza Remarks. Scaphichnium, the first named ichno-
p. 213. genus of the ichnofamily (Bown & Kraus 1983),
1998 Rebuffoichnus Roselli; Buatois, Mangano, shows peculiar characters that make its inclusion
Genise & Taylor p. 226. in the Pallichnidae tentative. Pallichnus Retallack
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 433

Key to identify the ichnogenera of Pallichnidae


1 Hamate to lunate with bulbous termination, unlined walls, meniscate filling Scaphichnium Bown and Kraus
Spherical or ovoid shape, lined walls, empty chambers 2
2 Spherical shape Pallichnus Retallack
Ovoid shape Fictovichnus Johnston et al.

1984 is herein designated as the type ichnogenus. 71994 'Egglike concretions' '? lizard eggs'
The presence of associated tunnels as found in Mikhailov, Sabath & Kurzanov p. 106, figs
Pallichnus is an unusual trait for this ichnofamily 7.16D-F, 7.20.
and is therefore not considered as diagnostic. ?1994a 'Pseudoeggs' Hirsch p. 281, fig. 11.3A.
Apart from the known ichnogenera, many ?1994b 'Pseudoeggs' Hirsch p. 145, fig. 10-6F.
unnamed ovoid-shaped trace fossils that are 71996 Celliforma sp. Veroslavsky & Martinez
devoid of constructed walls may be included in p. 41, pi. I, fig. 2.
this ichnofamily, including cocoons (e.g. Bown v!996 'Nodulos ovoidales' Sciutto & Martinez
et al. 1997), pseudoeggs (e.g. Hirsch 1994a), p. 74.
misidentified eggs (e.g. Johnston et al. 1996) and * 1996 Fictovichnus Johnston, Eberth & Anderson
pupation chambers (e.g. Edwards et al. 1998). p. 521, figs l,3A-Band4.
v!997 'Wasp traces' Bown, Hasiotis, Genise,
Ichnogenus Scaphichnium Bown & Kraus 1983 Maldonado & Browers p. 48, figs 6A, C-E,
(Fig. 4a) 8A-B.
*1983 Scaphichnium Bown & Kraus p. 106, figs v!998 'Cocoon-like trace fossils' Edwards,
4C, 5E-G, 6C, E, 9B, D. Jarzembowski, Pain & Daley p. 25, figs 3-6.
1993 Scaphichnium Bown & Kraus; Hasiotis, 1999 Fictovichnus Johnston, Eberth & Anderson;
Asian & Bown p. 2, figs 2, 4. Genise, Sciutto, Laza, Gonzalez & Bellosi p. 29.
1998 Scaphichnium Bown & Kraus; Genise & 2000 Fictovichnus Johnston, Eberth & Anderson;
Lazap. 214. Genise, Mangano, Buatois, Laza & Verde p. 54.
1998 Scaphichnium Bown & Kraus; Buatois, 2002 Fictovichnus Johnston, Eberth & Anderson;
Mangano, Genise & Taylor p. 227. Genise, Sciutto, Laza, Gonzalez & Bellosi
1999 Scaphichnium Bown & Kraus; Genise p. 110. p. 230.
2000 Scaphichnium Bown & Kraus; Genise, v2002 'Cocoons' Melchor, Genise & Miquel
Mangano, Buatois, Laza & Verde p. 54. p. 24, fig. 12G-H.
2000 Scaphichnium Bown & Kraus; Krell p. 892. Type ichnospecies. Fictovichnus gobiensis
Type and only known ichnospecies. Johnston et al. 1996.
Scaphichnium hamatum Bown & Kraus 1983. Diagnosis. Ellipsoid chambers enveloped by
Diagnosis. Discrete, hook-shaped to lunate, thin clay-rich zone, the inner surface of which
meniscate endostratal burrow fills, oriented is clearly defined and smooth. The outer surface
with long axis vertical to concave upward and of the clay-rich zone is gradational with the sur-
with rounded, bulbous, lower (distal) termina- rounding matrix. The long axis of the chamber is
tions (Bown & Kraus 1983). parallel to bedding, with a terminal exit hole that
Remarks. Scaphichnium hamatum is an unusual may be subterminal or medial (on the upper
insect trace fossil in many aspects. Its particular surface of the chamber relative to bedding).
shape, meniscate fill and lack of a lined or con- Where the ichnogenus occurs in calcretized
structed wall confer to this structure a very dis- soils, the chamber is commonly surrounded by
tinctive architecture unknown in other trace variably developed calcite halo. Trace fossils
fossils. Consequently, its inclusion in Pallichni- occur as hollow structures with a wall formed
dae is tentative. These trace fossils are attributed of a clay-rich zone and calcite halo, or as egg-
to brood masses of Scarabaeinae beetles, pos- shaped internal moulds, depending on local
sibly Geotrupinae (Hasiotis et al. 1993). calcretization and induration of host sediment
(Johnston et al. 1996).
Ichnogenus Fictovichnus Johnston et al. 1996 Included ichnospecies. F. gobiensis Johnston et al.
(Fig. 4c) 1996 (=parvus Johnston et al. 1996 syn. nov.).
71982 'Ovoid vesicles' Freytet & Plaziat p. 65, pi. Remarks. Both named ichnospecies of Ficto-
49, figs G, H. vichnus - F. gobiensis and F. parvus - were diag-
71987 'Cocoons' Ritchie p. 435, pi. 11.14, figs 9, nosed on the basis of differing size, probably
10. reflecting the fact that they were made by differ-
71994 'Cocoons' Thackray p. 796. ent species of insect. However, the potential
434 J. F. GENISE

Fig. 4. Schematic drawings of Pallichnidae and Krausichnidae described by Bown & Kraus (1983), Retallack
(1984), Johnston et al (1996), Bown et al. (1997), and Hasiotis & Dubiel (1995a). (a) Scaphichnium hamatum
Bown & Kraus 1983: Lower Eocene Willwood Formation, USA. (b) Pallichnus dakotensis Retallack 1984:
Oligocene Brule Formation, USA. (c) Fictovichnus gobiensis Johnston et al. 1996: Late Cretaceous Djadokhta
Formation, Mongolia, (d) Parowanichnus formicoides Bown et al. 1997: Palaeocene-Eocene Claron Formation,
USA. (e) Idealized reconstruction of Archeoentomichnus metapolypholeos Hasiotis & Dubiel 1995a: Late Triassic
Chinle Formation, USA.

existence of trace fossils of intermediate size showing an ovoid shape (e.g. Freytet & Plaziat
would confuse the ichnospecific taxonomy: thus 1982; Ritchie 1987; Thackray 1994; Veroslavsky
the two ichnospecies are considered herein to & Martinez 1996; Sciutto & Martinez 1996;
be synonymous. A third, unnamed ichnospecies Bown et al. 1997; Melchor et al. 2002). They are
from the Quaternary of Australia, included in attributed herein tentatively to Fictovichnus
Fictovichnus by Johnston et al. (1996), is trans- ispp. In addition, Johnston et al. (1996 and
ferred to Rebuffoichnus owing to the presence references therein) mentioned a long list of doubt-
of a constructed wall in the Australian material ful vertebrate fossil eggs or pseudoeggs that are
(Genise et al. 2002a). also probably attributable to Fictovichnus (e.g.
Many ovoid casts were described or mentioned Hirsch 1994a, 1994b; Mikhailov et al. 1994).
as 'cocoons', 'ovoid structures' and/or illustrated However, most of these cocoons or pseudoeggs
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 435

lack any evidence of the emergence hole, an (Retallack 1984). These trace fossils are inter-
important diagnostic character to definitively preted as pupation cells of scarabaeid beetles,
associate these dubiofossils with this ichnogenus. particularly Geotrupinae and Scarabaeinae
In contrast, the cocoon-like trace fossils (Retallack 1984).
described by Edwards et al. (1998) show evidence
of an emergence hole, a complex bioglyph and, in
some cases, an associated burrow, features Ichnofamily Krausichnidae ifam. nov.
important enough to suggest a new ichnospecies
of Fictovichnus. This ichnogenus is attributed to Type ichnogenus. Krausichnus Genise & Bown
pupation chambers of Coleoptera, probably 1994b.
Curculionidae, Tenebrionidae or Scarabaeidae Diagnosis. Group of ichnogenera showing
(Johnston et al. 1996). chambers associated with burrow systems, com-
posed in most cases of burrows of very different
Ichnogenus Pallichnus Retallack 1984 (Fig. 4b) diameters. Burrows are devoid of scratch marks
*1984 Pallichnus Retallack p. 580, figs 7, 9, 10. and/or intersecting grooves. Chambers have no
1990b Pallichnus Retallack; Retallack p. 221, radiating tunnels from their upper parts and are
figs 204, 206. commonly linked to a burrow system that nor-
1993 Pallichnus Retallack; Genise p. 50. mally interconnects them with other chambers.
1994b Pallichnus Retallack; Genise & Bown Chambers may be empty, passively or actively
109. filled, and/or they may contain secondary systems
1996 Pallichnus Retallack; Johnston, Eberth & of burrows of different diameters and smaller
Anderson p. 522. chambers on or within their inner walls.
1998 Pallichnus Retallack; Buatois, Mangano, Remarks. The first described and most repre-
Genise & Taylor p. 227. sentative ichnogenera of this ichnofamily,
1999 Pallichnus Retallack; Genise p. 110. Attaichnus Laza 1982 and Termitichnus Bown
2000 Pallichnus Retallack; Retallack, Bestland & 1982, bear names relating to ants and termites
Fremd p. 178. respectively. However, names such as Termitich-
2000 Pallichnus Retallack; Genise, Mangano, nidae or Attaichnidae should be avoided because
Buatois, Laza & Verde p. 54. Krausichnidae comprises trace fossils that can be
2000 Pallichnus Retallack; Krell p. 891. attributed indistinctly to ants or termites, and
2000 Pallichnus Retallack; Genise & Poire p. 7. also because trace fossil names should not
2002 Pallichnus Retallack; Melchor, Genise & make reference to possible producers (e.g. Brom-
Miquel p. 29. ley 1990). In contrast, Krausichnus shows one of
Type and only known ichnospecies. Pallichnus the most complex morphologies of the group,
dakotensis Retallack 1984. and its name does not refer to the presumed con-
Diagnosis. Nearly spherical chambers, defined structor. Among the ichnotaxa included within
by thin wall of dark clay and organic matter; the Krausichnidae, Termitichnus, Vondrichnus
inner boundary of wall sharp and smooth, such and Fleaglellius constitute a well-defined mor-
that the internal cast is easily separated from phological group that may deserve the creation
rock matrix. The outer boundary of the wall of an ichnofamily distinct from the Krausichni-
grades outward into the surrounding matrix. dae once they are better known.
One side of the chamber is disrupted to form In addition to named ichnogenera, there is an
large, irregularly circular, exit cavity, usually extensive pedologic literature in relating to the
about half the diameter of the main chamber. origin of laterites, dealing with unnamed trace
Each nearly spherical chamber is arranged at fossils attributed to termites, mostly from the
end of short branches from the vertical burrow, Pliocene and Pleistocene of tropical areas (e.g.
so that the exit cavity faces into the branch Machado 1983; Grasse 1986; Schaefer 2001). It
burrow; both vertical and branch burrows of is almost impossible to deal with the ichnotaxon-
slightly lesser diameter than the nearly spherical omy of this material because in most cases the
chamber (Retallack 1984). supposed nests were only studied micromorpho-
Remarks. The presence of a lined wall clearly logically and show no clear boundaries. Meaning
distinguishes this trace fossil from those included that structures were only studied micromor-
in Coprinisphaeridae. The preservation of phologically (not macromorphologically) macro-
tunnels is unusual among those ichnogenera morphological studies are lacking. Conversely,
attributed to the work of solitary insects, this kind of study is needed in already named
grouped in Celliformidae, Coprinisphaeridae trace fossils, and could usefully be conducted
and Pallichnidae. Accordingly, tunnel remains in order to evaluate the degree of reliability of
in Pallichnus are detected only in thin sections the attribution of these ichnogenera to termites
436 J. F. GENISE

Key to the ichnogenera of Krausichnidae


1 Chamber walls neither constructed nor lined 2
Chamber walls constructed or lined 3
2 Spherical chambers having only one main vertical burrow connected to the
lower part or, rarely, another one at the top of the chamber. Secondary
burrows connecting the main ones and chambers at different points.
The distribution of chambers is similar throughout the structure Attaichnus Laza.
Oblate to subspherical chambers interconnected by burrows of similar
diameter arising from top, bottom and sides of chambers. Chambers and
burrows more numerous in the central part of the nest Parowanichnus Bown et al.
3 Structures composed of tiered, tabular and flat chambers supported by pillars
and ramps 4
Structures composed of subspherical, ovoid or oblate chambers 5
4 Tiered, tabular and flat chambers composing spindle-shaped to columnar
compound chambers lacking an external, discrete, wall. Compound chambers
interconnected by a burrow system and/or isolated burrows Krausichnus Genise & Bown
Tiered, tabular and flat chambers composing a columnar, central structure
surrounded by a discrete wall. A burrow system connects this structure to
peripheral oblate chambers and possibly other columns Archeoentomichnus Hasiotis & Dubiel
5 Structures composed of a single, large, cup-shaped chamber surrounded by a
thick constructed wall bearing a boxwork of secondary burrows and chambers
and a peripheral system of radiating burrows with differing diameters Tacuruichnus Genise
Structures composed of a system of chambers having similar sizes 6
6 Chambers connected by primary burrows of similar sizes 7
Chambers connected by primary and secondary burrows 8
7 At most three apposed, obovate chambers. Commonly having a rind of
anastomosing burrows at top and sides of chambers. Diffuse arrangement
of chambers Vondrichnus Genise & Bown
Two or more apposed chambers commonly forming towers. Bases and tops
of chambers convex upwards without a rind of anastomosing burrows Fleaglellius Genise & Bown
8 Spherical to subspherical chambers. Burrows, simple or compound and well
differentiated from chambers, having much smaller diameters Termitichnus Bown
Elongate to oblate chambers. Burrows consistently simple, in some cases
comparable in size to chambers Syntermesichnus Bown & Laza

or ants. For instance, whereas Krausichnus, posed constructors (e.g. Bown & Laza 1990;
Termitichnus and Attaichnus leave few doubts Laza 1995, 1997; Hasiotis & Dubiel 1995a).
about their termite and ant origin respectively,
Tacuruichnus is known from a single specimen Ichnogenus Attaichnus Laza 1982 (Fig. 5a)
and requires further confirmation of its structure v*1982 Attaichnus Laza p. 112, pis II, III.
using more material. Vondrichnus, Fleaglellius 1993 Attaichnus Laza; Genise p. 53.
and Parowanichnus are comparatively simple 1997 Attaichnus Laza; Bown, Hasiotis, Genise,
structures lacking the distinctive boxwork Maldonado & Browers p. 45.
(sensu Ekdale et al. 1984) composed of tunnels 1998 Attaichnus Laza; Buatois, Mangano,
of different diameters. Finally, Syntermesichnus Genise & Taylor p. 227.
and Archeoentomichnus were described from 1999 Attaichnus Laza; Genise p. 111.
fragmentary material, from which the whole 2000 Attaichnus Laza; Genise, Buatois,
structure and ichnogeneric diagnoses were Mangano, Laza 7 Verde p. 55.
inferred. Micromorphological confirmation of 2001a Attaichnus Laza; Retallack p. 143.
the biological affinities of these trace fossils is cri- Type and only known ichnospecies. Attaichnus
tical, moreover, as the Triassic Archeoentomich- kuenzelii Laza 1982.
nus predates the oldest (Cretaceous) termite Diagnosis. System of spherical chambers
body fossils by about 150 million years (Hasiotis interconnected by primary and secondary bur-
& Dubiel 1995a) (Table 1). rows. Primary burrows connected to the cham-
The taxonomy of this ichnofamily is complex bers vertically to the lower part, forming inside
because the trace fossils were described using dif- a folded rim in some cases. A second primary
ferent criteria to select taxobases and particularly burrow can be connected at the opposite side of
because the diagnoses were strongly influenced the chamber. The secondary burrows inter-
by the architecture of modern nests of the sup- connect chambers with primary ones. The
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 437

Fig. 5. Krausichnidae. (a) Attaichnus kuenzelii Laza 1982: chamber and burrow casts. Late Miocene Cerro
Azul Formation, Argentina (MACN-LI1787, 1788, 1791, 1792). Bar: 1 cm. (b) Krausichnus trompitus Genise &
Bown 1994b, close-up of the holotype: flat chambers with parallel roofs and floors, vertical pillars (centre), and
dark linings and fillings. Late Eocene-Oligocene Jebel Qatrani Formation, Egypt. Bar: 1 cm. (c) Tacuruichnus
farinai Genise 1997: holotype. Close to the knife it is possible to see the external wall bearing a system of
burrows and chambers; on the left, large burrows radiating from the cup-shaped nest. Late Pliocene Barranca
de Los Lobos Formation, Argentina. Bar: 10cm.

chamber system occupies a conical area up to 7 m of the other representatives of Krausichnidae.


in diameter and 3 m in height, in which the cham- The large (140-170 mm) spherical chambers con-
bers are regularly distributed (modified from nected by one or at most two primary burrows at
Laza 1982). the base and at the top are unmistakable traits.
Remarks. This trace fossil is known only from This trace fossil is preserved as detached casts
the type locality. However, its morphological of chambers and burrows having no indication
features are clearly distinguishable from those of the presence of a lined or constructed wall
438 J. F. GENISE

(Fig. 5a). Laza (1982) attributed this trace fossil V71998 Krausichnus Genise & Bown; Genise,
to ants of the genus Atta because of the gross Pazos, Gonzalez, Tofalo & Verde p. 12.
morphology of the structure, the size and shape 1999 Krausichnus Genise & Bown; Genise
of chambers and burrows, the folded rim of p. 112.
primary burrows entering to the chambers, and 2000 Krausichnus Genise & Bown; Genise,
the secondary burrows. Buatois, Mangano, Laza & Verde p. 55.
2000 Krausichnus Genise & Bown; Miller &
Ichnogenus Parowanichnus Bown et al., 1997 Mason p. 210.
(Fig. 4d) 72000 'Termitieresfossiles' Schuster, Duringer,
*1997 Parowanichnus Bown, Hasiotis, Genise, Nel, Brunet, Vignaud & Mackaye p. 15,
Maldonado & Brouwers p. 45, figs 4, 5. figs 3-5.
1999 Parowanichnus Bown, Hasiotis, Genise, 2002 Krausichnus Genise & Bown; Pazos, Tofalo
Maldonado & Brouwers; Genise p. 111. & Sanchez-Bettuci p. 34.
2000 Parowanichnus Bown, Hasiotis, Genise, 2002 Krausichnus Genise & Bown; Buatois,
Maldonado & Brouwers; Genise, Buatois, Mangano & Acenolaza p. 189.
Mangano, Laza & Verde p. 55. Type ichnospecies. Krausichnus trompitus Genise
Type and only known ichnospecies. & Bown 1994b.
Parowanichnus formicoides Bown et al. 1997. Diagnosis. Tiered arrangement of tabular, flat
Diagnosis. System composed of oblate to chambers with roofs and floors flat and parallel.
subspherical chambers interconnected by Chambers exhibit vertical pillars, partition walls
burrows of similar diameter. Burrows connected and conspicuous linings. Successive chambers
at tops, sides and bottoms of chambers. The are connected by tiny passages. Arrangement of
gallery network is crudely trellate in plan, tiered chambers may take various forms, such
forming a grid-like lattice with descending shaft as spindles or columns, resulting in compound
galleries and lateral tunnel galleries set more or chambers without surrounding walls. These
less perpendicular to one another. Lined or compound chambers may be interconnected by
constructed walls are lacking. Burrows, and the isolated or interconnecting burrows (modified
chambers they connect, radiate away from the from Genise & Bown 1994b).
centre of the structure and gradually decline in Included ichnospecies. K. trompitus Genise &
number. The chamber system occupies an area Bown 1994b; K. altus Genise & Bown 1994b.
up to approximately 1 m in height and 3.3m in Remarks. The original ichnogeneric diagnosis
width (modified from Bown et al. 1997). included details of the shape and arrangement
Remarks. Parowanichnus differs from Attaich- of compound chambers, but it is now clear that
nus in having: (1) a much smaller total nest such taxobases are more useful at ichnospecific
volume; (2) much smaller and less densely level. A third, unnamed ichnospecies (Coaton
packed chambers; (3) chambers oblate to hemi- 1981; Schuster et al. 2000) demonstrates that
spherical rather than globular; (4) galleries of the morphological features that compose a recur-
one basic size (smaller than in Attaichnus); (5) rent architecture of ichnogeneric rank include
galleries providing access equally to top, sides tabular and tiered chambers showing pillars
and bottom of chambers; and (6) chambers and and forming compound chambers without exter-
burrows more densely grouped in the central nal walls. The shape of the compound chambers
part of the structure (Bown et al. 1997). As with and their interconnections are useful to distin-
Attaichnus, chambers and burrows lack linings guish ichnospecies. The whole structure may
or constructed walls. This trace fossil is attributed consist of a single compound chamber or a
to ants because its structure is composed of system of related compound chambers. Another
burrows connected with chambers, and it lacks ichnogenus that shows tiered tabular chambers
lined or constructed walls (Bown et al. 1997). is Archeoentomichnus, in which these structures
are connected with simple elongate oblate
Ichnogenus Krausichnus Genise & Bown, 1994 chambers.
(Fig. 5b) In K. trompitus and K. altus, plus the Chadian
71981 'Fossilized nests of Hodotermitidae' material (Schuster et al. 2000), the combination
Coatonp. 79, fig. 1. of chambers and burrows indicates a social
1993 'Chevron-shaped chambers' Bown & insect. The conspicuous linings and the con-
Genise p. A58. structed pillars and vertical walls specifically
v*1994b Krausichnus Genise & Bown p. 169, suggest termites (Genise & Bown 1994b;
figs6H-I, 7, 8D, 9-11, 12A. Schuster et al. 2000). Accordingly, Coaton
1998 Krausichnus Genise & Bown; Buatois, (1981) attributed his unnamed fossil nests to
Mangano, Genise & Taylor p. 227. the Hodotermitidae.
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 439

Ichnogenus Archeoentomichnus Hasiotis & Diagnosis. Cup-shaped structure composed of


Dubiel 1995a (Fig. 4e) a wall bearing a net of anostomosing burrows
1993 Archeoentomoichnos Hasiotis & Dubiel surrounding a chamber. Exteriorly the wall is
p. 177 (nomen nudum). connected to a gallery system composed of bur-
1994 Archeoentomichnus Hasiotis & Dubiel p. 8 rows of different diameters (Genise 1997).
(nomen nudum). Remarks. Tacuruichnus looks like a large
*1995a Archeoentomichnus Hasiotis & Dubiel; specimen of Termitichnus qatranii; however, in
Hasiotis & Dubiel p. 121, figs 3-5. the latter ichnogenus the chambers are spherical
1995b Archeoentomoichnus Hasiotis & Dubiel; and closed, because they would have been
Hasiotis & Dubiel p. 86, fig. 1.6 (lapsus). located completely below ground (Genise &
1998 Archeoentomichnus Hasiotis & Dubiel; Bown 1994b; Genise 1997). Tacuruichnus is the
Buatois, Mangano, Genise & Taylor p. 225. only krausichnid represented by a structure com-
1999 Archeoentomichnus Hasiotis & Dubiel; posed of a single large chamber. However, the
Genise p. 112. characteristic burrow and chamber systems of
2000 Archeoentomichnus Hasiotis & Dubiel; the ichnofamily are present in the chamber's
Genise, Buatois, Mangano, Laza & Verde external wall and periphery (Fig. 5c).
2000 p. 54. Its architecture closely resembles the hypo-
2000 Archeoentomichnus Hasiotis & Dubiel; geous part of the nest of Cornitermes cumulans
Hasiotis p. 154. (Nasutitermitinae) (Genise 1997). The presence
200la Archaeoentomonichnus Retallack p. 143 of a single chamber, interpreted as a nest, is
(lapsus). also compatible with nests of C. cumulans. It
Type and only known ichnospecies. would be important to find more specimens to
Archeoentomichnus metapolypholeos Hasiotis determine the complete morphology and the
& Dubiel 1995a. taxonomic affinity of this trace fossil.
Diagnosis. Peripheral part of the structure Ichnogenus Vondrichnus Genise & Bown, 1994
composed of large and small anostomosing gal- (Fig. 6a)
leries and elongate and oblate chambers. Central
part columnar and partially subterranean, *1994b Vondrichnus Genise & Bown p. 165,
internally with stacked floor levels and slender, figs 5J; 6A, B.
steeply inclined, connecting ramps. Galleries 1998 Vondrichnus Genise & Bown; Buatois,
connect the central part to elongate oblate Mangano, Genise & Taylor p. 227.
chambers and possibly to other columnar parts 1999 Vondrichnus Genise & Bown; Genise
(modified from Hasiotis & Dubiel 1995a). p. 112.
Remarks. The original diagnosis is slightly 2000 Vondrichnus Genise & Bown; Genise,
modified to avoid interpretative terms such as Mangano, Buatois, Laza & Verde p. 54.
periecie or endoecie that correspond specifically 2002 Vondrichnus Genise and Bown; Buatois,
to termite architecture. This ichnogenus is Mangano & Acenolaza p. 189.
described from fragmentary material. The diag- Type and only known ichnospecies. Vondrichnus
nosis is based on an idealized reconstruction, obovatus Genise & Bown 1994b.
rather than the holotype's morphology as Diagnosis. Diffuse, polychambered, excavated
illustrated by Hasiotis & Dubiel (1995a). The subterranean systems. Obovate chambers occur
reconstructed morphology is similar to that of in dense swarms of near 300 in cross-section.
Krausichnus, but in Archeoentomichnus the Burrows simple, branched or unbranched, exit-
tiered chambers are connected with elongate, ing from one or more points on periphery of
oblate, simple chambers, which are lacking in chamber and comprising a dense mass of anasto-
Krausichnus. Moreover, the tabular chambers mosing burrows that may connect chambers.
are thicker, and the central part of the nest less Sediment in the centre of the chambers is alveolar
organized in general aspect than in Krausichnus. and commonly arranged in concentric bands.
Chambers expanded by apposition of 1-3
chambers against one another (Genise and
Ichnogenus Tacuruichnus Genise 1997 Bown 1994b).
(Fig. 5c) Remarks. Vondrichnus differs from Termitich-
v*1997 Tacuruichnus Genise p. 140, figs 2, 3. nus in: (1) consistently smaller mean chamber
1999 Tacuruichnus Genise; Genise p. 112. size; (2) obovate form of chamber; (3) tighter
2000 Tacuruichnus Genise; Genise, Buatois, packing of associated chambers; (4) absence of
Mangano, Laza & Verde p. 55. isolated chambers; (5) larger number of galleries
Type and only known ichnospecies. Tacuruich- in each cluster; (6) lack of gallery ornamentation;
nus farinai Genise 1997. (7) greater density of galleries between chambers;
440 J. F. GENISE

Fig. 6. Krausichnidae. (a) Vondrichnus obovatus Genise & Bown 1994b. Late Eocene-Oligocene Jebel Qatrani
Formation, Egypt. Bar: 1 cm. (b) Fleaglellius pagodus Genise & Bown 1994b: three-chambered tower. Late
Eocene-Oligocene Jebel Qatrani Formation, Egypt. Bar: 1 cm. (c) Syntermesichnus fontanae Bown & Laza
1990: interconnected chambers lined with a lighter fine material; cross-section of an individual thin passage
(on the left). Miocene Pinturas Formation, Argentina; MACN-LI81. Bar: 1 cm. (d) Termitichnus simplicidens
Genise & Bown 1994b: main burrows (particularly visible on the right) radiate from the base of the chamber.
Late Eocene-Oligocene Jebel Qatrani Formation, Egypt. Bar: 10cm.

and (8) absence of compound galleries. Vondrich- authors, such as size, packing of chambers and
nus differs from Fleaglellius in: (1) never having number of burrows, are undoubtedly very
more than two apposed chambers; (2) rarely useful for dividing the morphological complex
having only vertically apposed chambers; (3) of Termitichnus-Fleaglellius-Vondrichnus; how-
not having convex-upward chambers; (4) ever, they are not very satisfactory as ichnotaxo-
commonly possessing a rind of anastomosed bases in a more general context. Vondrichnus and
peripheral galleries at the tops and sides of Fleaglellius show a primary burrow system
chambers; and (5) occurring as diffuse but inter- devoid of secondary tunnels that distinguish
connected groups of chambers (Genise & Bown them from Termitichnus. Vondrichnus can be
1994b). Some of the characters stated by the clearly separated from Fleaglellius because in
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 441

the latter the chambers are convex upwards and 71993 Termitichnus Bown; Smith, Mason &
form 'towers' by vertical apposition (Genise & Ward p. 592, fig. 16.
Bown 1994b) (Fig. 6a, b). Possible trace-makers V1994b Termitichnus Bown; Genise & Bown
are termites, possibly Macrotermitinae (Genise p. 160, figs 4, 5, 8A-C.
& Bown 1994b). The very simple architecture 1997 Termitichnus Bown; Genise p. 140.
of this trace fossil necessitates further exami- 1998 Termitichnus Bown; Buatois, Mangano,
nation of micromorphological characters to Genise & Taylor p. 227.
confirm its affinities. 1998 Termitichnus Bown; Smith & Mason p. 555,
figs 12, 13, 14A.
Ichnogenus Fleaglellius Genise & Bown, 1994 1999 Termitichnus Bown; Genise p. 112.
(Fig. 6b) 2000 Termitichnus Bown; Genise, Mangano,
1994b Fleaglellius Genise & Bown p. 167, figs Buatois, Laza & Verde p. 54.
6C-G, 8E-F. 2000 Termitichnus Bown; Miller & Mason p. 208,
1998 Fleaglellius Genise & Bown; Buatois, figs 12-16.
Mangano, Genise & Taylor p. 227. 2001a Termitichnus Bown; Retallack p. 143.
1999 Fleaglellius Genise & Bown; Genise p. 112. 2002 Termitichnus Bown; Buatois, Mangano &
2000 Fleaglellius Genise & Bown; Genise, Acenolaza p. 189.
Mangano, Buatois, Laza & Verde p. 54. Type ichnospecies. Termitichnus qatranii Bown,
2002 Fleaglellius Genise & Bown; Buatois, 1982.
Mangano & Acenolaza p. 189. Diagnosis. Diffuse, polychambered excavated
Type and only known ichnospecies. Fleaglellius subterranean systems with spherical to sub-
pagodus Genise & Bown 1994b. spherical chambers connected to one another
Diagnosis. Diffuse, polychambered, excavated by a network of simple and/or compound gal-
subterranean system. Chambers oblate, with leries of different diameters. Primary burrows
bases and tops of chambers almost invariably arise from the base of chambers. Chambers
convex-upward and generally apposed. Succes- may be isolated, as expanded globular clusters
sive chambers are added vertically or near verti- of chambers or in associated constellate aggrega-
cally, with the top of the lower completely tions of tens to hundreds of chambers (modified
overlapped by the base of the upper chamber, from Genise & Bown 1994b).
such that completed structures make up towers Included ichnospecies. T. qatranii Bown 1982;
of 2-35 chambers. Towers are widely dispersed T. simplicidens Genise & Bown 1994b; T. nami-
in groupings of up to 40 specimens and are con- biensis Miller & Mason 2000.
nected by dense masses of simple galleries. Gal- Remarks. The diagnosis has been modified to
leries, both branched and unbranched, connect exclude characters such as size and fill of cham-
different towers at all levels (Genise & Bown bers, which are more useful for ichnospecific
1994b). diagnosis. Also excluded are those characters
Remarks. This ichnogenus differs from its referred to 'nest expansion', which is a concept
closest morphological counterpart, Vondrichnus, linked to the supposed constructors rather than
in that Fleaglellius'. (1) always has more than one to documented morphology. The basal position
and up to 35 apposed chambers, always added of the primary burrows is an important character
vertically; (2) invariably has individual chambers added to the diagnosis, because it is constant and
that are noticeably convex-upwards; (3) lacks a very representative in both ichnospecies. Differ-
peripheral rind of anastomosed galleries at tops ences from Fleaglellius and Vondrichnus were
and/or sides of chambers; and (4) lacks a diffuse commented on in previous sections. Attaichnus
distribution of individual chambers (Genise & has consistently spherical chambers always, and
Bown 1994b). Possible trace-makers are termites a single primary burrow arises from the base of
of unknown affinities (Genise & Bown 1994b). chambers, and in some cases other from the
The particular kind of enlargement of nest by top. The architecture of Syntermesichnus is
vertical apposition of chambers is unknown in clearly different from that of Termitichnus',
subterranean termites, but resembles that of however, in terms of ichnotaxobases it is difficult
certain subaerial nests. This ichnogenus requires to translate these differences avoiding measure-
micromorphological analysis of its affinities. ments and other taxobases of dubious merit. In
addition the commonest type of Termitichnus
Ichnogenus Termitichnus Bown, 1982 (Fig. 6d) comprises spherical or subspherical chambers
v*1982 Termitichnus Bown p. 259, figs 2-7. from which one to five primary burrows arise,
71984 Termitichnus Bown; Tandon & Naug some of which may connect another distant
p. 285, figs 7B, 9B-E. chamber (Fig. 6d). In contrast, Syntermesichnus
1993 Termitichnus Bown; Genise p. 54. takes on the aspect of a boxwork of burrows
442 J. F. GENISE

and chambers of rather similar diameters the chambers, suggest that the inclusion of this
(Fig. 6c). trace fossil in Termitichnus would require
Not all of the trace fossils attributed to Termi- further work. On the other hand, although the
tichnus after its original description (Bown 1982) absence of external wall makes some specimens
and redescription (Genise & Bown 1994b) belong comparable to Krausichnus, these specimens
to that ichnogenus. Smith et al. (1993) reported are incomplete and eroded (Miller & Mason
supposed Termitichnus from the late Pleistocene 2000). Possible trace-makers include termites
of Namibia, and proposed a Termitichnus ichno- belonging to Macrotermitinae (Genise & Bown
facies, but their figured specimen does not closely 1994b).
resemble Termitichnus. Probable Termitichnus
have subsequently been illustrated from the Ichnogenus Syntermesichnus Bown & Laza 1990
Tertiary of Namibia by Smith & Mason (1998): (Fig. 6c)
these are discussed below with unnamed Krau- v*1990 Syntermesichnus Bown & Laza p. 74, figs
sichnidae. 2-4.
An additional ichnospecies of Termitichnus 1993 Syntermesichnus Bown & Laza; Genise
from early Pleistocene deposits of South Africa, p. 54.
T. namibiensis, is composed of isolated or inter- 1995 Syntermesichnus Bown & Laza; Genise and
connected chambers, surrounded by a thick Cladera p. 80, fig. 2C-D.
wall and a network of simple tunnels (Miller & 1995 Syntermesichnus Bown & Laza;
Mason 2000). Chambers of T. namibiensis are Constantino p. 460.
filled with meniscate, tiered, galleries. A thick 71997 Syntermesichnus Bown & Laza; Smith &
wall and peripheral tunnel system surround iso- Kitchingp. 41, figs 16, 17.
lated or interconnected subspherical chambers 1997 Syntermesichnus Bown & Laza; Bown,
typical of Termitichnus. The attribution to this Hasiotis, Genise, Maldonado & Browers p. 46.
ichnogenus to fossil termite nests is also sup- 1998 Syntermesichnus Bown & Laza; Buatois,
ported by the tiered arrangement of shelves, as Mangano, Genise & Taylor p. 227.
in other fossil and modern termite nests (e.g. 1999 Syntermesichnus Bown & Laza; Genise
Sands 1987). However, some characters raise p. 112.
doubts about the ichnotaxonomical placement 2000 Syntermesichnus Bown & Laza; Genise,
of this trace fossil. Chambers apparently lack Mangano, Buatois, Laza & Verde p. 54.
the characteristic primary tunnels of Termitich- 2002 Syntermesichnus Bown & Laza; Buatois,
nus, and the interpretation of the internal struc- Mangano & Acenolaza p. 189.
ture is confusing. It is not clear how to Type and only known ichnospecies.
interpret a gallery that extends horizontally and Syntermesichnus fontanae Bown & Laza 1990.
also shows a meniscate filling, which commonly Diagnosis. Peripheral part of the structure,
results from the backfilling of burrows of similar tabular, with large and small anastomosing
diameter from that of the trace-maker (e.g. burrows and elongate, oblate chambers. Large
D'Alessandro & Bromley 1987; Keighley & burrows arising from chambers, but small bur-
Pickerill 1994). Also, the turning points at the rows branch from larger ones. Systems of small
end of galleries are unusual, in that the thin passages at one level extend the structure in the
layers between two successive shelves commonly horizontal plane and permit communication to
become thicker towards the sides before joining other levels. The same style of branching is
the external wall (e.g. Sands 1987). These thick- present at all levels. Walls of burrows and
enings help to reinforce the whole structure. In chambers lined with compacted sediment (finer
T. namibiensis the thickenings are disconnected than host matrix) (modified from Bown & Laza
from the outer wall at the turning points. In 1990).
addition, T. namibiensis also lacks the ramps, Remarks. Modifications of the original diag-
pillars and openings that are common in Krau- nosis were made to avoid any reference to
sichnus, as well as the thin-layered ovoids and modern termite nests. Syntermesichnus shows
the hive of modern termite nests (e.g. Sands one of the simplest morphologies of Krausichni-
1987; Genise & Bown 1994b). In sum, the inter- dae and, consequently, one of the most difficult
nal structure of the chambers of T. namibiensis to define and separate from the other Krausich-
(Miller & Mason 2000, fig. 12) plus the above- nidae. The whole structure lacks definite limits
mentioned characters leave some doubts about and, in the field, resembles a diffuse boxwork of
the trace-makers of this internal structure, tunnels and chambers of more or less similar dia-
which may be different from those of the meter occupying entire beds (Genise & Bown,
chambers. These characters, plus the absence of unpublished data). The abundance of similar
primary burrows arising from the base of Syntermesichnus-\ikG traces in pyroclastic
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 443

deposits from the Cretaceous and Tertiary of structure. The fifth type is represented by three
southern South America (Genise, unpublished small ovoids showing pits, protrusions, ramps
data) gives this trace fossil a particular impor- and openings respectively. These resemble simi-
tance. The chamber and burrow systems, lined lar structures from the Tertiary of Namibia
with fine material, suggested to Bown & Laza attributed to Termitichnus by Smith and Mason
(1990) that Syntermesichnus was a fossil termite (1998), and also unnamed trace fossils from the
nest. However, the lack of a definite architecture early Miocene of Ethiopia (Bown & Genise
(e.g. nest limits) and its pervasive presence in 1993). They superficially resemble Termitichnus,
beds preclude a definite attribution without but lack the associated burrow system, which is
micromorphological analysis of the lined walls. diagnostic. The sixth type is similar to the first
Trace fossils comparable to Syntermesichnus type but also has chambers and pores arranged
are described from the Jurassic Elliot Formation in anastomosing columnar and alveolar struc-
of South Africa (Smith & Kitching 1997), tures. No other trace fossils are comparable to
although this attribution needs further analysis. the sixth type. The last type, from the upper
Possible trace-makers are termites of the genus part of the Laetoli beds, consists of thin-layered
Syntermes (Bown & Laza 1990). However, as ovoids that show a layered internal structure
discussed above, this attribution requires micro- having ramps and connecting openings sur-
morphological study and the examination of rounded by a sculptured wall. This type is very
further material to look for well-delimited similar to unnamed trace fossils described by
nests. In addition, as Constantino (1995) noted, Coaton (1981) and Schuster et al. (2000) from
the descriptions of Syntermes nests in the litera- the Pleistocene of South Africa and the Pliocene
ture are inadequate, so that the affinities of of Chad respectively, and is attributable to a
Syntermesichnus should be considered doubtful. distinct ichnospecies of Krausichnus. However,
the ovoids from Laetoli have a constructed
Unnamed trace fossils attributable to wall, an unusual trait for Krausichnus that is
Krausichnidae lacking in the other African material. Miller &
Various trace fossils from the Pliocene Laetoli Mason (2000) considered these structures attri-
Formation of Tanzania, all attributable to butable to their ichnospecies Termitichnus nami-
Krausichnidae, were described by Sands (1987). biensis.
He recognized seven basic types. Sands (1987) related most of these types of
The first type is composed of systems of trace fossil to different parts of Macrotermitinae
anastomosing burrows of different diameters, nests at different stages of development and/or
which may show vertical shafts associated with constructed in different types of soil, with two
flattened chambers. These systems are com- exceptions: the vertical shaft with flattened
parable to the structures attributed by Smith chambers, which more probably relates to ant
et al. (1993) to Termitichnus. However, these nests of Camponotus, and the thin-layered
structures are not clearly compatible with the ovoids, which are attributable to neither the
ichnogeneric diagnosis. The second type with Macrotermitinae nor the Hodotermitidae
thick-layered ovoids (= chambers) shows par- (Sands 1987).
ticular features, such as the presence of a In summary, the Laetoli material (Sands 1987)
surrounding cavity, that are unknown from represents a spectrum of samples of krausichnids
other fossil termite nests. These ovoids lack an that share common features with other represen-
external wall and associated burrow systems. tatives of the ichnofamily from Africa (e.g.
They somewhat resemble very roughly ichno- Coaton 1981; Bown & Genise 1993; Smith et al.
species of Krausichnus in having a layered struc- 1993; Genise & Bown 1994b; Smith & Mason
ture, but the layers are unusually thick. A third 1998; Schuster et al. 2000). This similarity
type of trace fossil with shafted chambers is not reflects the origin of these trace fossils in
comparable to any other Krausichnidae. This common lineages of African termites. Ichnotaxo-
type is composed of a bell-shaped chamber nomically, the Laetoli material presents some
from which vertical shafts arise, which are problems. Considering the architecture of
capped by smaller shafts, chambers and pores. Macrotermitinae nests, each type described is
The fourth type, composed of a single ovoid, is more likely a part of a more complex structure
similar to the second type, but has an external than a complete structure in itself. This fact reiter-
wall and ramps and passages from one floor to ates a common trend in evolution of behaviour in
the next. The fourth type can be attributed to insects, in which complex architectures and
neither Krausichnus nor Termitichnus because behaviours are the result of the addition of
Krausichnus does not have an outer wall and simple behaviours and architectures respectively.
Termitichnus does not have a comparable layered One possibility is to include the types one and six
444 J. F. GENISE

(tunnel systems) in Syntermesichnus, interpreting with the fragmentary nature of the material,
this ichnogenus as diffuse boxworks of tunnels makes any ichnotaxonomical consideration
and chambers. It would also be possible to very difficult. There are still other unnamed
include the thick- and thin-layered ovoids in trace fossils, cited as termite or ant nests (e.g.
Krausichnus or Termitichnus, but important Tandon & Naug 1984; Iriondo & Krohling
features distinguish these ovoids from the 1996; Tauber 1996; Andreis & Cladera 1998)
known ichnospecies. Another standpoint might that are only mentioned and will require analysis
be to include each Laetolian type in a new ichno- and proper description.
taxon, even if each is part of a more complex A sound description of this unnamed material,
structure; this procedure would be supported by in the light new understanding of krausichnid
the fact that some of these types are recurrent as morphology, will aid incorporation of this
individual trace fossils in other localities and material into the emerging ichnotaxonomic
ages in Africa (e.g. Coaton 1981; Bown & framework.
Genise 1993; Smith & Mason 1998; Schuster
et al. 2000).
Other unnamed Krausichnidae were described Ichnostratigraphy
by Laza (1995, 1997) from Pliocene and Pleisto-
cene deposits of Argentina. They include seven There are 25 described ichnogenera attributed
types of trace fossil: two attributed to termites, to insect trace fossils in palaeosols, 19 of which
and five to ants. One type of termite nest is are reviewed herein, and the remaining six in a
currently being redescribed by Laza (personal previous contribution (Genise 2000) (Table 1).
communication) and two ant nests described in Almost all of them show comparable strati-
1995 were redescribed by Laza (1997). Hasiotis graphic ranges that, in turn, accord with our
& Demko (1996) also described a supposed ant present knowledge of the evolutionary history
nest from the Jurassic Morrison Formation. of their trace-makers: bees, beetles, ants and
The importance of all this material for our termites (e.g. Crowson 1981; Kuschel 1983;
knowledge of the diversity of Krausichnidae is Krishna 1990; Jarzembowski & Ross 1996;
unquestionable. However, in all cases the diag- Grimaldi et al. 1997; Grimaldi 1999; Engel
noses and descriptions are influenced by the sup- 2000; Grimaldi & Agosti 2000; Krell 2000;
posed modern analogue, which, in combination Schaefer 2001; Nel et al in press).

Table 1. Stratigraphic ranges and abundance of insect ichnotaxa in palaeosols

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25

Pleistocene 4 1 1 3 1 i
Pliocene 4 1 1 1 1 2
Miocene 3 1 1 1 1 1 2 4 2 1
Oligocene 2 1 1 1 1 1 1 3 1
Eocene 1 5 2 1 3 1 4 1 1 1 1 1 1 1 3 1 1 1
Palaeocene 1 1 2
Upper Cretaceous 1 3
Middle Cretaceous
Lower Cretaceous
Jurassic & 1
Triassic i
Permian
Carboniferous
Devonian
Silurian
Ordovician
Cambrian

Coprinisphaeridae (1-6), Pallichnidae (7-9), Krausichnidae (10-18) and Celliformidae (19-25). 1, Fontanai;
2, Coprinisphaera; 3, Eatonichnus; 4, Monesichnus; 5, Teisseirei; 6, Rebuffoichnus; 7, Pallichnus; 8, Fictovichnus;
9, Scaphichnium; 10, Attaichnus; 11, Parowanichnus; 12, Krausichnus; 13, Archeoentomichnus; 14, Tacuruichnus;
15, Vondrichnus; 16, Fleaglellius; 17, Termitichnus; 18, Syntermesichnus; 19, Palmiraichnus; 20, Celliforma;
21, Corimbatichnus; 22, Uruguay; 23, Rosellichnus; 24, Ellipsoideichnus; 25, Cellicalichnus. Numbers in cells
represent formations in which the ichnotaxon is recorded. Circles indicate the oldest body fossil of the potential
trace-maker.
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 445

The Coprinisphaeridae Pliocene Chad Formation, Chad (Duringer et al.


2000a, 2000b), the Pleistocene Ensenada
Among the Coprinisphaeridae, Fontanai has Formation, Argentina (Frenguelli 1938a), the
been recorded only from the Late Cretaceous- Pleistocene Tezanos Pinto Formation, Argentina
Early Tertiary Asencio Formation of Uruguay (Iriondo & Krohling 1996), the Pleistocene Tafi
(Roselli 1939). These redbed deposits could not del Valle Formation, Argentina (Fontaine et al.
be dated precisely until now because of the com- 1995), and from an unnamed Pleistocene forma-
plete absence of datable rocks or fossil remains tion, from Ecuador (Sauer 1955). Eatonichnus
other than the fossil nests (Genise et al. 2004). has been recorded from the Palaeocene Colter
Genise et al. (2002a) suggested a possible Formation, USA (Gilliland & La Rocque
Eocene age for this unit, owing to the presence 1952), the late Palaeocene-Eocene Claron
of an important unconformity with the under- Formation, USA (Bown et al. 1997), and the
lying Cretaceous Yapeyu Formation (Pazos early Palaeocene Penas Coloradas Formation,
et al. 2002), comparison of the ichnofauna with Argentina (Genise et al. 2001). Monesichnus is
other Palaeogene deposits in South America, another ichnogenus that has been recorded
and the abundance and diversity of dung-beetle exclusively from the Late Cretaceous-Early
nests. This abundance and diversity could reflect Tertiary Asencio Formation of Uruguay (Roselli
the increased availability of herbivore faeces 1987).
corresponding to the Eocene diversification and The previously mentioned ichnogenera of
abundance of South American herbivores. In Coprinisphaeridae are attributed to Scarabaei-
Table 1 all the records from this formation are nae (Frenguelli 1938; Laza 1986b; Bown et al.
considered to be Eocene. Because this formation 1997; Genise & Laza 1998; Genise 1999; Krell
is one of the richest in fossil insect nests - it 2000), whose oldest body fossils come from the
includes 10 of the 25 described ichnogenera - it Late Cretaceous strata (Krell 2000). In this case
produces a peak of diversity for this age that the body fossils shortly predate the trace fossils
may be an artefact, not simply because the age of the group, as there is no ichnological record
of this formation is unknown, but also because for the Cretaceous. Moreover, the presence of
intensive research on other formations and Coprinisphaera is taken as indicative of Cenozoic
continental deposits has just begun. Never- deposits in South America (Genise et al. 2000).
theless, it cannot be overlooked that, during the HalfTter & Edmonds (1982) and Cambefort
early Eocene, a climatic optimum occurred (1991) suggested that dung-beetles might have
(e.g. Zachos et al. 2001), which would have diversified by the end of the Cretaceous, helped
favoured the abundance and diversification of by the radiation of the herbivorous dinosaurs
some groups of insects (e.g. Wilf & Labandeira and/or the increase of mammal excrement. The
1999). only Cretaceous trace fossils attributable to
Coprinisphaera and Celliformaare the most dung-beetles (non-Scarabaeinae) are those
widely recorded ichnogenera in palaeosols. described by Chin & Gill (1996) from dinosaur
Coprinisphaera has been recorded from the coprolites.
aforementioned Asencio Formation in Uruguay The other two ichnogenera of Coprinisphaeri-
(Roselli 1939, 1987; Genise et al. 2004), the dae deserve particular attention. Teisseirei has
early Eocene Casamayor Formation, Argentina been recorded from the Late Cretaceous-Early
(Frenguelli 1938b; Laza 1986a), the late Eocene Tertiary Asencio Formation, Uruguay (Roselli
Musters Formation, Argentina (Andreis 1972; 1939), the Eocene Gran Salitral Formation,
Laza 1986a), the late Eocene La Meseta Forma- Argentina (Melchor et al. 2002), and the Eocene-
tion, Antarctica (Laza & Reguero 1990), the Miocene Sarmiento Formation, Argentina
Eocene-Miocene Sarmiento Formation (Laza (Bellosi et al. 2001). Despite the well-preserved
1986a; Bellosi et al. 2001), the Oligocene macro- and micromorphological characters of
Deseado Formation, Argentina (Frenguelli this trace fossil, as well as its abundance in Tertiary
1938b; Laza 1986a), the early Miocene Pinturas deposits of South America, the trace-maker
Formation, Argentina (Genise & Bown 1994a), cannot yet be identified more precisely than as
the late Miocene Collon-Cura Formation, probably being a coleopteran. Hence it is impossi-
Argentina (Frenguelli 1939; Laza 1986b), the ble to compare the stratigraphic range of Teisseirei
late Miocene Paso de las Carretas Formation, with that of the body fossil record of any particular
Argentina (Pascual & Bondesio 1981), the Plio- coleopteran. Rebuffoichnus has been recorded
cene Monte Hermoso Formation, Argentina from the Late Cretaceous Laguna Palacios
(Laza 1986b), the Pliocene Piquete Formation, Formation, Argentina (Genise et al. 2002b), the
Argentina (Alonso et al. 1982), the Pliocene Late Cretaceous-Early Tertiary Asencio
Laetoli Formation, Kenya (Sands 1987), the Formation, Uruguay (Roselli 1987), and the
446 J. F. GENISE

Pleistocene of Australia (Lea 1925; Johnston et aL The Krausichnidae


1996). This ichnogenus, probably attributable to
the work of weevils (Genise et al. 2002b), is one Representatives of Krausichnidae show, in most
of only two that crosses the K-T boundary, being cases, similar characteristics: body fossils predat-
recorded from the Late Cretaceous to the Pleisto- ing trace fossils and scarcity of data, with some
cene. As such, it is one of the oldest trace fossils exceptions. Attaichnus and Parowanichnus, the
that can be definitely attributed to insects two ichnogenera attributed to ants, have been
(Genise et al 2002b). The body fossil record of recorded exclusively from the Miocene Epecuen
their probable constructors, the curculionids, Formation, Argentina (Laza 1982) and from
extends back to Upper Jurassic-Lower Cretaceous the Eocene Claron Formation, USA (Bown
deposits (Crowson 1981; Kuschel 1983; Jarzem- et al. 1997) respectively. The oldest body fossils
bowski & Ross 1996). As a whole, the body fossil of ants come from the Cretaceous of the USA
record of possible trace-makers predates the and France (Grimaldi et al. 1997; Grimaldi &
Coprinisphaeridae, even though the thick con- Agosti 2000; Nel et al. in press).
structed walls confer to these trace fossils a high Krausichnus has been recorded from the Late
preservation potential. Cretaceous-Early Tertiary Asencio Formation,
Uruguay (Genise et al. 1998), the late Eocene
Qasr el Sagha Formation and late Eocene-Oligo-
The Pallichnidae cene Jebel Qatrani Formation, Egypt (Genise &
Bown 1994b), the late Miocene Baynunah For-
The Pallichnidae pose a different problem from mation, Abu Dhabi Emirate (Bown & Genise
that of the Coprinisphaeridae. Scaphichnium 1993), the Pliocene of Chad (Schuster et al.
has been recorded exclusively from the Eocene 2000), and the Pleistocene of South Africa
Willwood Formation, USA (Bown & Kraus (Coaton 1981). Tacuruichnus has been recorded
1983), whereas its possible producers, the Geo- exclusively from the late Pliocene Barranca de
trupinae (Hasiotis et aL 1993), are recorded los Lobos Formation, Argentina (Genise 1997),
since the late Oligocene (Krell 2000). Pallichnus whereas Vondrichnus and Fleaglellius from the
has been recorded only from the Oligocene late Eocene-Oligocene Jebel Qatrani Formation,
Brule Formation in the USA (Retallack 1984), Egypt (Genise & Bown 1994b). Termitichnus
whereas its possible producers, the Geotrupinae shows a broader distribution, being recorded
or Scarabaeinae (Retallack 1984), are recorded from the late Eocene-Oligocene Jebel Qatrani
from the late Oligocene and Late Cretaceous Formation, and the Late Eocene Qasr el Sagha
respectively (Krell 2000). Fictovichnus, in a Formation, Egypt (Bown 1982), the early Mio-
broad sense, may include very simple trace fossils cene Kashab Formation, Egypt (Genise &
showing a morphology that has been recorded Bown 1994b), the Plio-Pleistocene Boulder For-
from the Late Jurassic Morrison Formation, mation, India (Tandon & Naug 1984), the late
USA (Hirsch 1994b), the Late Cretaceous Pleistocene Homeb Silts, Namibia (Smith et al.
Barun Goyot Formation, Mongolia (Mikhailov 1993), the late Pleistocene Sossus Sand, Namibia
et al. 1994), the Late Cretaceous Djadokhta (Smith & Mason 1998), and the Pleistocene of
Formation, Mongolia (Johnston et al. 1996), South Africa (Miller & Mason 2000). Finally,
the Late Cretaceous Bajo Barreal Formation, Syntermesichnus has been recorded from the
Argentina (Sciutto & Martinez 1996), the Palaeo- Miocene Pinturas Formation, Argentina (Bown
cene of Uruguay (Veroslavsky & Martinez 1996), & Laza 1990), and there is a doubtful record
the Eocene Claron Formation, USA (Bown et al. from the Early Jurassic Elliot Formation,
1997), the Eocene of France (Freytet & Plaziat South Africa (Smith & Kitching 1997). With
1982; Hirsch 1994a), the Eocene Gran Salitral the exception of this last record, all other occur-
Formation, Argentina (Melchor et al. 2002), rences are predated by the oldest fossil termites
the late Eocene Bembridge Limestone Forma- from the Lower Cretaceous (Krishna 1990).
tion, England (Edwards et al. 1998), the Miocene The case of Archeoentomichnus from the
Higewi Formation, Kenya (Thackray 1994), and Triassic Chinle Formation, USA (Hasiotis &
the Pliocene of Tanzania (Ritchie 1987). It is Dubiel 1995a) deserves particular attention
impossible to determine a unique trace-maker because it is the only datum that does not fit
for Fictovichnus, whose makers may have the general picture shown in Table 1. These
included the Tenebrionidae, Curculionidae, authors attributed this trace fossil to termites
Scarabaeidae (Johnston et al. 1996), or even despite the fact that it would predate the oldest
other families of Coleoptera. This precludes body fossils by 150 million years (Hasiotis &
any precise comparison with the body fossil Dubiel 1995a). Many invertebrate trace fossils
record of the coleopteran families. are more preservable than their producers:
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 447

accordingly, fossil bee nests predate the oldest can unequivocally be attributed to insects are
bees (Elliott & Nations 1998; Genise 2000). few, and are recorded from only three Late
However, the few million years involved in this Cretaceous formations in the USA, Mongolia
difference between the oldest bee trace fossils and Argentina (Johnston et al. 1996; Elliott &
and the oldest fossils of the probable construc- Nations 1998; Genise et al. 2002b). The record
tors is an expected one. It is consistent with our of insect fossil nests from rocks of this age
previous knowledge of the evolutionary history accords with the body fossil record of their prob-
of bees and their relationship with the coevolving able constructors and/or reflects their supposed
plant groups (Grimaldi 1999; Engel 2001). In the evolutionary history. A scenario proposed by
case of Archeoentomichnus, the palaeoentomolo- Genise & Bown (1994a) stated that the diversity
gical significance that would result from the of insect fossil nests in palaeosols increased
discover of a Triassic termite nest predating the significantly after the Cretaceous and not earlier.
oldest termites by 150 million years would This hypothesis is based on the fact that most
require sound proof, such as micromorphologi- common trace fossils in palaeosols are construc-
cal studies, to be accepted as such. At present, tions belonging to termites, bees and dung-
pending further work, this occurrence is not beetles, groups that arose during that period
accepted. based on body fossil evidence. This hypothesis
The whole interpretation of Mesozoic evolu- was later corroborated by the discovery of Late
tion of palaeosol ichnofaunas attributed to Cretaceous bee nests and coleopteran pupal
insects by Hasiotis (2000) is mostly unsupported, chambers (Johnston et al. 1996; Elliot & Nations
in that the attribution of these early purported 1998; Genise 2000; Genise et al. 2002b). The
trace fossils to insects is not well documented Genise and Bown hypothesis is also supported
or well founded. The conclusions are based by the increase in abundance of records of
largely on poorly supported interpretations of insect fossil nests in Tertiary palaeosols, in
Triassic and Jurassic trace fossils from the contrast to the general absence of records from
Chinle and Morrison Formations. These inter- pre-Cretaceous deposits.
pretations are at odds with the strong evidence Although Labandeira & Sepkoski (1993)
that indicate that ants, termites, bees and dung- stated that the appearance and expansion of
beetles arose and diversified during the Cretac- angiosperms had no influence on insect familial
eous (Krishna 1990; Labandeira 1998; Grimaldi diversification, their quantitative analysis did
1999; Krell 2000; Engel 2001; Schaefer 2001). not evaluate the ecological importance of the
This conclusion is also supported by the ichno- families involved. The origin of termites, ants,
logical record of insect nests in palaeosols bees and dung-beetles during the Cretaceous
(Table 1). Accordingly, Labandeira (1998), was probably related to the origin and diversifi-
Grimaldi (1999), Genise (2000) and Engel cation of angiosperms and to the broad-scale
(2001) objected to such interpretations of ecological changes that resulted in the K-T
Chinle and Morrison trace fossils because of mass extinction event. These insect trace-
inadequate documentation. makers would have played a major role in this
Triassic and Jurassic trace fossils are unknown event as the new soil colonizers of the emergent
from other studied deposits of similar age, which ecosystems. The record of latosols, from the Cre-
adds significance to the Chinle and Morrison taceous onwards in the stratigraphic column, has
trace fossils, as increasing the necessity for been strongly related to the radiation of termites
sound descriptions and interpretations of the and angiosperms (Schaefer 2001). In contrast to
fossils. Other Triassic and Jurassic palaeosols other groups that became extinct or less domi-
studied are of low ichnodiversity, composed nant following the K-T event, insect ichnofossils
mainly of simple trace fossils such as Skolithos, in palaeosols show that their producers were part
Macanopsis, Taenidium and Edaphichnium, of the new flourishing ecosystems.
none of which can be certainly attributed to This evolutionary scenario is reflected in the
insects (Retallack 1976, 1980; Smith & Kitching stratigraphic range and abundance of insect
1997; Melchor et al 2001; Genise et al 2004). trace fossils in palaeosols reflected in Table 1,
The morphology of trace fossils in palaeosols which shows the absence of Celliformidae,
ranges from very simple to very complex. The Coprinisphaeridae, Pallichnidae and Krausichni-
attribution of traces of very simple morphology dae in pre-Cretaceous rocks. It also shows that
to modern taxa is a misleading procedure the oldest record of two of these ichnofamilies
because they can commonly be attributed to sev- came from Cretaceous rocks, which contain a
eral different groups of organisms (Ratcliffe & lower diversity and abundance of nests than
Fagerstrom 1980; Retallack 1990b; Genise et al Tertiary ones. Even so, Cretaceous nests are
2004). Trace fossils in Mesozoic palaeosols that very important ones, in relation to the origin of
448 J. F. GENISE

important groups of insects and their building BOWN, T. M. HASIOTIS, S. T., GENISE, J. F., MALDO-
behaviour. Tertiary rocks have the most diverse, NADO, F. & BROUWERS, E. M. 1997. Trace fossils
abundant and well-preserved assemblages of of Hymenoptera and other insects and paleonvir-
insect fossil nests, in accordance with the diversi- onments of the Claron Formation (Paleocene
fication of insects and their building behaviour. and Eocene), Southwestern Utah. Bulletin of the
United States Geological Survey, 2153, 42-58.
BERTLING, M., BRADDY, S. ETAL. 2003. Draft proposal
The initial manuscript benefited from comments by A. to emend the Code with respect to trace fossils:
Uchman and A. Rindsberg. I thank also D. Mcllroy, request for comments. Bulletin of Zoological
M. Verde and A. Rindsberg for improving the final Nomenclature, 60, 141-142.
version. This research was partially supported by a BROMLEY, R. G. 1990. Trace Fossils. Unwin Hyman,
grant from the National Agency of Scientific and London.
Technical Promotion of Argentina (FONCYT-PICT BROMLEY, R. G. 1996. Trace Fossils: Biology, Taphon-
6156/99) to the author. omy and Applications. Chapman & Hall, London.
BROWN, R. W. 1934. Celliforma spirifer, the fossil larval
chambers of mining bees. Journal of the Washing-
References ton Academy of Sciences, 24, 532-539.
BRUET, E. 1950. Le loess de la Republique de 1'Equa-
ALONSO, R. N., GONZALEZ, C. E. & PELAYES, H. A. teur et ses nids fossiles d'insectes. Revue Francaise
1982. Hallazgo de roedores y nidos de escara- de Entomologie, 17, 280-283.
beidos en el Terciario Superior de la Sierra de BUATOIS, L. A., MANGANO, M. G. & ACENOLAZA, F. G.
Vaqueros, Salta, Republica Argentina. Revista 2002. Trazas Fosiles. Museo Paleontologico Egidio
del Instituto de Ciencias Geologicas, 5, 1-3. Feruglio, Trelew. Edition Especial MEF No. 2.
ANDREIS, R. 1972. Paleosuelos de la Formacion Mus- BUATOIS, L. A., MANGANO, M. G., GENISE, J. F. &
ters (Eoceno Medio), Laguna del Mate, Provincia TAYLOR, T. N. 1998. The ichnologic record of
de Chubut, Republica Argentina. Revista de la the continental invertebrate invasion: evolutionary
Sociedad Argentina de Mineralogia, Petrografia y trends in environmental expansion, ecospace
Sedimentologia, 3, 91-97. utilization, and behavioural complexity. Palaios,
ANDREIS, R. 1981. Identification e importancia geolo- 13, 217-240.
gica de los paleosuelos. Editora de Universidade CAMACHO, H. 1966. Invertebrados fosiles. Eudeba,
Federal do Rio Grande do Sul, Brasil. Buenos Aires.
ANDREIS, R. & CLADERA, G. 1998. Sistemas fluviales CAMBEFORT, Y. 1991. Biogeography and evolution. In:
entrelazados neocretacicos en la Patagonia HANSKI, I. & CAMBEFORT, Y. (eds) Dung Beetle
septentrional, Argentina: Facies, ciclicidad y Ecology. Princeton University Press, Princeton,
paleocorrientes. Resumenes de la Septima Reunion 51-67.
Argentina de Sedimentologia, 99-101. CANE, J. H. 1991. Soils of ground-nesting bees (Hyme-
BATRA, S. T. W. 1984. Solitary bees. Scientific Ameri- noptera: Apoidea): texture, moisture, cell depth
can, 250, 120-127. and climate. Journal of the Kansas Entomological
BELLOSI, E. S., LAZA, J. H. & GONZALEZ, M. G. 2001. Society, 64, 406-413.
Icnofaunas en paleosuelos de la Formacion Sar- CHIN, K. & GILL, B. D. 1996. Dinosaurs, dung-beetles,
miento (Eoceno-Mioceno), Patagonia Central. and conifers: participants in a Cretaceous food
Resumenes de la IVReunion Argentina de Icnologia web. Palaios, 11, 280-285.
y Segunda Reunion de Icnologia del Mercosur, COATON, W. G. H. 1981. Fossilised nests of Hodoter-
Tucuman, 31. mitidae (Isoptera) from the Clanwilliam district,
BOWN, T. M. 1982. Ichnofossils and rizoliths of Cape Province. Journal of the Entomological
the nearshore fluvial Jebel Qatrani Formation Society of South Africa, 44, 79-81.
(Oligocene), Fayum Province, Egypt. Palaeogeo- CONSTANTINO, R. 1995. Revision of the Neotropical
graphy, Palaeoclimatology, Palaeoecology, 40, genus Syntermes Holmgren (Isoptera: Termiti-
255-309. dae). The University of Kansas Science Bulletin,
BOWN, T. M. & GENISE, J. F. 1993. Fossil nests and 55,455-518.
gallery systems of termites (Isoptera) and ants COSARINSKY, M. 2001. Micromorfologia del nido de
(Formicidae) from the early Miocene of Southern Cornitermes cumulans (Kollar) (Isoptera, Termiti-
Ethiopia and the late Miocene of Abu Dhabi dae). Resumenes de la Cuarta Reunion Argentina
Emirate, UAE. Abstracts with Programs of the de Icnologia y Segunda Reunion de Icnologia del
Annual Meeting of the Geological Society of Mercosur, Tucuman, 36.
America, 25, 58. CROWSON, R. A. 1981. The Biology of the Coleoptera.
BOWN, T. M. & KRAUS, M. J. 1983. Ichnofossils of Academic Press, New York.
the alluvial Willwood Formation (Lower Eocene), D'ALESSANDRO, A. & BROMLEY, R. 1987. Meniscate
Bighorn Basin, Northwest Wyoming, USA. trace fossils and the Muensteria-Taenidium prob-
Palaeogeography, Palaeoclimatology, Palaeo- lem. Palaeontology, 30, 743-763.
ecology, 43, 95-128. D'ALESSANDRO, A. & BROMLEY, R. 1995. A new
BOWN, T. M. & LAZA, J. H. 1990. A Miocene fossil ichnospecies of Spongeliomorpha from the
termite nest from southern Argentina and its Pleistocene of Sicily. Journal of Paleontology, 69,
paleoclimatological implications. Ichnos, 1, 73-79. 393-398.
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 449

DIBLIN, M. C, RANDAZZO, A. F. & JONES, D. S. 1991. FRANCIS, J. C. 1975. Esquema bioestratigrafico regio-
Lithoplaision ocalae: a new trace fossil from the nal de la Republica Oriental del Uruguay. Adas
Ocala Limestone (Eocene), Florida. Ichnos, 1, del Primer Congreso Argentino de Paleontologia y
255-260. Bioestratigrafia, 2, 539-568.
DONOVAN, S. K. 1994. Insects and other arthropods as FRENGUELLI, J. 1938a. Bolas de escarabeidos y nidos de
trace-makers in non-marine environments and vespidos fosiles. Physis, 12, 348-352.
palaeoenvironments. In: DONOVAN, S. K. (ed.) FRENGUELLI, J. 1938b. Nidi fossili di Scarabeidi e Ves-
The Paleobiology of Trace Fossils. Wiley, New pidi. Bolletino Societta Geologia Italiana, 57, 77-96.
York, 200-220. FRENGUELLI, J. 1939a. Sobre nidos fosiles del Neuquen
DURINGER, P., BRUNET, M., CAMBEFORT, Y., BEAUVI- y Rio Negro. Revista de la Sociedad Entomologica
LAIN, A., MACKAYE, H. T., VIGNAUD, P. & Argentina, 10, 270.
SCHUSTER, M. 2000a. Des boules de bousiers FRENGUELLI, J. 1939b. Nidos fosiles de insectos en el
fossiles et leurs terriers dans les sites a Australo- Terciario del Neuquen y Rio Negro. Notas del
pitheques du Pliocene tchadien. Bulletin de la Museo de La Plata (Paleontologia}, 4, 379-402.
Societe Geologique du France, 171, 259-269. FRENGUELLI, J. 1940. Viaje a la zona central andina de
DURINGER, P., BRUNEI, M., CAMBEFORT, Y., LIKIUS, la Patagonia septentrional. Revista del Museo de
A., MACKAYE, H. T., SCHUSTER, M. & VIGNAUD, La Plata (Seccion Oficial), 1940, 53-76.
P. 2000b. First discovery of fossil dung-beetle FRENGUELLI, J. 1941. Viaje a los territorios patagonicos
brood balls and nests in the Chadian Pliocene del Neuquen y del Chubut. Revista del Museo de
Australopithecine levels. Lethaia, 33, 277-284. La Plata (Seccion Oficial), 1941, 80-91.
DURINGER, P., SCHUSTER, M., CAMBEFORT, Y., NEL, A., FREYTET, P. & PLAZIAT, J. C. 1982. Continental
BRUNET, M., VIGNAUD, P. & MACKAYE, H. T. (in carbonate sedimentation and pedogenesis - late
press). Predation of dung-beetles brood-balls by Cretaceous and early Tertiary of Southern
termites in Chadian Pliocene. A possible explana- France. Contributions to Sedimentology, 12.
tion for unusual dung-beetles gallery networks. GENISE, J. F. 1993. Trazas fosiles de insectos en paleo-
Implications for dung-beetles brood-balls preser- suelos. In: Melchor, R. N. (ed.) Nuevas tendencias
vation. Palaeogeography, Palaeoclimatology, en el estudio de trazas fosiles. Facultad de Ciencias
Palaeoecology. Exactas y Naturales (UNLPam), Santa Rosa, La
EDWARDS, N., JARZEMBOWSKI, E. A., PAIN, T. & DALEY, Pampa, 49-59.
B. 1998. Cocoon-like trace fossils from the lacus- GENISE, J. F. 1997. A fossil termite nest from the
trine-palustrine Bembridge Limestone Formation Marplatan stage-age (late Pliocene) of Buenos
(Late Eocene), Southern England. Proceedings of Aires province, Argentina, as paleoclimatic
the Geologists' Association, 109, 25-32. indicator. Palaeogeography, Palaeoclimatology,
EKDALE, A. A., BROMLEY, R. G. & PEMBERTON, S. G. Palaeoecology, 136, 139-144.
1984. Ichnology: The Use of Trace Fossils in GENISE, J. F. 1999. Paleoicnologia de Insectos. Revista
Sedimentology and Stratigraphy. Society of Eco- de la Sociedad Entomologica Argentina, 58, 104-
nomic Paleontologists and Mineralogists, Tulsa, 116.
Oklahoma. GENISE, J. F. 2000. The ichnofamily Celliformidae for
ELLIOTT, D. K. & NATIONS, J. D. 1998. Bee burrows in Celliforma and allied ichnogenera. Ichnos, 1,
the Late Cretaceous (Late Cenomanian) Dakota 267-284.
Formation, Northeastern Arizona. Ichnos, 5, GENISE, J. F. & BOWN, T. M. 1994a. New Miocene
243-253. scarabeid and hymenopterous nests and Early
ENGEL, M. S. 2000. A new interpretation of the oldest Miocene (santacrucian) paleoenvironments, Pata-
fossil bee (Hymenoptera: Apidae). American gonian Argentina. Ichnos, 3, 107-117.
Museum Novitates, 3296, 1—11. GENISE, J. F. & BOWN, T. M. 1994b. New trace fossils
ENGEL, M. S. 2001. A Monograph of the Baltic Bees and of termites (Insecta: Isoptera) from the Late
Evolution of the Apoidea (Hymenoptera). Bulletin Eocene-Early Miocene of Egypt, and the recon-
of the American Museum of Natural History, 259, struction of ancient isopteran social behaviour.
1-192. Ichnos, 3, 155-183.
EVANS, H. E. 1963. Predatory wasps. Scientific Ameri- GENISE, J. F. & BOWN, T. M. 1996. Uruguay Roselli and
can, 208, 145-154. Rosellichnus n. ichnogen: two ichnogenera for
FONTAINE, J., BALLESTEROS, J. M. & POWELL, J. E. 1995. cluster of fossil bee cells. Ichnos, 4, 199-217.
Artefactos del comportamiento de escarabajos GENISE, J. F. & CLADERA, G. 1995. Application of
(Coleoptera, Scarabaeidae) como evidencias computerized tomography for studying insect
paleoclimaticas y paleoambientales en el Cuater- traces. Ichnos, 4, 77-81.
nario (Pleistocene Superior) del valle de Tafi, GENISE, J. F. & LAZA, J. H. 1998. Monesichnus ameghi-
provincia de Tucuman, Argentina. Resumenes de noi Roselli: a complex insect trace fossil produced
la Quinta Reunion Nacionalde la Asociacion Argen- by two distinct trace makers. Ichnos, 5, 213-223.
tina de Ciencias del Comportamiento, Tucuman, 12. GENISE, J. F. & POIRE, D. G. 2000. Fluidization in insect
FORD, I. 1988. Conglomerados con nidos de insectos constructions in soils. Ichnos, 1, 127-134.
fosiles: Formation Palmitas (provisorio) - Ter- GENISE, J. F. & VERDE, M. 2000. Corimbatichnus
ciario Inferior (tentative). Adas del Sexto Panel fernandezi: a cluster of fossil bee cells from the
de Geologia del Litoral y Primera Reunion de Late Cretaceous-Early Tertiary of Uruguay.
Geologia del Uruguay, Salto, Uruguay, 47-49. Ichnos,!, 115-125.
450 J. F. GENISE

GENISE, J. F. & ZELICH, M. R. 2001. Trazas fosiles de Proceedings of the National Academy of Sciences,
insectos de la Formation Puerto Unzue (Cretacico USA, 97, 13678-13683.
Superior-Paleogeno) de Entre Rios. Resumenes de GRIMALDI, D., AGOSTI, D. & CARPENTER, J. M. 1997.
la Cuarta Reunion Argentina de Icnologia y New and rediscovered primitive ants (Hymen-
Segunda Reunion de Icnologia del Mercosur, optera: Formicidae) in Cretaceous amber from
Tucuman, 44. New Jersey, and their phylogenetic relationships.
GENISE, J. F., PAZOS, P. J., GONZALEZ, M. G., TOFALO, American Museum Novitates, 3208.
R. O. & VERDE, M. 1998. Hallazgo de termiteros y HALFFTER, G. 1959. Etologia y paleontologia de S-
tubos meniscados en la Formation Asencio carabaeinae. Ciencia, 19, 165-178.
(Cretacico Superior-Terciario Inferior) R.O. del HALFFTER, G. & EDMONDS, W. D. 1982. The Nesting
Uruguay. Resumenes de la Tercera Reunion Argen- Behaviour of dung-beetles. An ecological and Evolu-
tina de Icnologia y Primera Reunion de Icnologia tive Approach. Publicaciones del Institute de
del Mercosur, Mar del Plata, 12-13. Ecologia de Mexico, Mexico D.F., 10, 1-176.
GENISE, J. F., SCIUTTO, J. C, LAZA, J. H., GONZALEZ, HALFFTER, G. & MATHEWS, G. 1966. The Natural
M. G. & BELLOSI, E. 1999. Fossil bee nests and History of Dung-Beetles of the Subfamily Scara-
coleopteran pupation chambers from the Laguna baeinae. Folia Entomologica Mexicana, Mexico
Palacios Formation (Late Campanian-Maastrich- D.F., 12-14, 1-312.
tian) from Central Patagonia. Abstracts of the HANSKI, I. & CAMBEFORT, Y. 1991. Resource partition-
Seventh International Symposium on Mesozoic ing. In: HANSKI, I. & CAMBEFORT, Y. (eds) Dung
Terrestrial Ecosystems, Buenos Aires, 28-29. Beetle Ecology. Princeton University Press, Prince-
GENISE, J. F., MANGANO, M. G., BUATOIS, L. A., LAZA, ton, 331-349.
J. H. & VERDE, M. 2000. Insect trace fossil associa- HANTSZCHEL, W. 1975. Trace fossils and problematica.
tions in palaeosols: the Coprinisphaera ichnofacies. In: TEICHERT, C. (ed.) Treatise on Invertebrate
Palaios, 15, 49-64. Paleontology (2nd edn), Part W (Supplement 1).
GENISE, J. F., CLADERA, G. & TANCOFF, S. 2001. La pre- Geological Society of America & Kansas Univer-
sencia de Eatonichnus claronensis en el Paleoceno sity Press, 269.
del Chubut (Argentina). Resumenes de la Cuarta HASIOTIS, S. T. 2000. The invertebrate invasion and
Reunion Argentina de Icnologia y Segunda Reunion evolution of Mesozoic soil ecosystems: the
de Icnologia del Mercosur, Tucuman, 45. ichnofossil record of ecological innovations. In:
GENISE, J. F., LAZA, J. H., FERNANDEZ, W. & FROGONI, GASTALDO, R. A. & Di MICHELE, W. A. (eds)
J. 2002a. Camaras pupales fosiles de coleopteros: Phanerozoic Terrestrial Ecosystems. Papers of the
el icnogenero Rebuffoichnus Roselli. Revista del Paleontological Society, Boulder, Colorado and
Museo Argentina de Ciencias Naturales, n.s,, 4, Lawrence, Kansas, 6, 141-169.
159-165. HASIOTIS, S. T. & DEMKO, M. 1996. Terrestrial and
GENISE, J. F., SCIUTTO, J. C., LAZA, J. H., GONZALEZ, freshwater trace fossils, Upper Jurassic Morrison
M. G. & BELLOSI, E. 2002b. Fossil bee nests, Formation, Colorado Plateau. In: MORALES, M.
coleopteran pupation chambers and tuffaceous (ed.) The Continental Jurassic. Bulletin of the
palaeosols from the Late Cretaceous Laguna Pala- Museum of Northern Arizona, 60, 355-370.
cios Formation, Central Patagonia (Argentina). HASIOTIS, S. T. & DUBIEL, R. F. 1993. Continental trace
Palaeogeography, Palaeoclimatology, Palaeoecol- fossils of the Upper Triassic Chinle Formation,
ogy, 111, 215-235. Petrified Forest National Park, Arizona. In:
GENISE, J. F., BELLOSI, E. S. & GONZALEZ, M. A. 2004. LUCAS, S. G. & MORALES, M. (eds) The Nonmarine
An approach to the description and interpretation Triassic. Bulletin of the New Mexico Museum of
of ichnofabrics in palaeosols. In: MC!LROY, D. Natural History and Science, 3, 175-178.
(ed.) 2004. The Application of Ichnology to HASIOTIS, S. T. & DUBIEL, R. F. 1994. Termite-nest
Palaeoenvironmental and Stratigraphic Analysis. ichnofossils in crevasse splay deposits, lower part
Geological Society, London, Special Publications, of the Petrified Forest Member, Upper Triassic
228, 355-382. Chinle Formation, Petrified Forest National
GILLILAND, W. N. & LA ROCQUE, A. 1952. A new Xeno- Park, Arizona. Research Abstracts of the Petrified
helixl from the Paleocene of Utah. Journal of Forest National Park, 3, 8-9.
Paleontology, 26, 501-504. HASIOTIS, S. T. & DUBIEL, R. F. 1995a. Termite (Insecta:
GRASSE, P. 1958. Sur le nid et la biologic de Cornitermes Isoptera) nest ichnofossils from the Upper Triassic
cumulans (Kollar), termites bresilien. Insectes Chinle Formation, Petrified Forest National Park,
Sociaux, 5, 187-199. Arizona. Ichnos, 4, 119-130.
GRASSE, P. 1984. Termitologia, Tome 2. Masson Ed., HASIOTIS, S. T. & DUBIEL, R. F. 1995b. Continental
Paris. trace fossils, Petrified Forest National Park: tools
GRASSE, P. 1986. Termitologia, Tome 3. Masson Ed., for paleohydrologic and paleoecosystem recon-
Paris. structions. Technical Report of the National Park
GRIMALDI, D. 1999. The co-radiations of pollinating Service, 16, 82-88.
insects and angiosperms in the Cretaceous. Annals HASIOTIS, S. T. & MITCHELL, C. E. 1993. A comparison
of the Missouri Botanical Garden, 86, 373-406. of crayfish burrow morphologies: Triassic and
GRIMALDI, D. & AGOSTI, D. 2000. A formicinae in Holocene paleo- and neoichnological evidence,
New Jersey Cretaceous amber (Hymenoptera: and the identification of their burrowing signa-
Formicidae) and early evolution of the ants. tures. Ichnos, 2, 291-314.
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 451

HASIOTIS, S. T., ASLAN, A. & BOWN, T. M. 1993. LAZA, J. H. 1986a. Icnofosiles de paleosuelos del Ceno-
Origin, architecture, and paleoecology of the zoico mamalifero de Argentina. I Paleogeno. Bole-
Early Eocene continental ichnofossil Scaphich- tin de la Asociacion Paleontologica Argentina, 15,19.
nium hamatum, integration of ichnology and LAZA, J. H. 1986b. Icnofosiles de paleosuelos del
paleopedology. Ichnos, 3, 1-9. Cenozoico mamalifero de Argentina. II Neogeno.
HAZELDINE, P. L. 1997. Comportamiento de nidifica- Boletin de la Asociacion Paleontologica Argentina,
cion de cuatro especies de Ptilothrix Smith 15, 13.
(Apidae, Emphorini). Physis, 54, 27-41. LAZA, J. H. 1995. Signos de actividad de insectos. In:
HIRSCH, K. F. 1994a. The fossil record of vertebrate ALBERDI, T. M., LEONE, G. & TONNI, E. P. (eds).
eggs. In: DONOVAN, S. K. (ed.) The Paleobiology Evolucion biologica y climdtica de la region
of Trace Fossils. Wiley, New York, 269-294. pampeana durante los ultimos cinco millones de
HIRSCH, K. F. 1994b. Upper Jurassic eggshells from the anos. Consejo de Investigaciones Cientificas de
Western Interior of North America. In: CARPEN- Espana, Madrid, 347-361.
TER, K., HIRSCH, K. F. & HORNER, J. R. (eds) LAZA, J. H. 1997. Signos de actividad atribuibles a dos
Dinosaur eggs and babies. Cambridge University especies de Acromyrmex (Myrmicinae, Formici-
Press, Cambridge, 137-150. dae, Hymenoptera) del Pleistoceno en la provincia
HOLLDOBLER, B. & WILSON, E. O. 1990. The Ants. de Buenos Aires, Republica Argentina. Signifi-
Harvard University Press, Cambridge, MA. cado paleoambiental. Revista de la Universidad
IRIONDO, M. & KROHLING, D. M. 1996. Los sedimentos Guarulhos, Geociencias, 2, 56-62.
eolicos del noroeste de la llanura pampeana LAZA, J. H. & REGUERO, M. A. 1990. Extension
(Cuaternario superior). Adas del Decimotercer faunistica de la antigua region neotropical en la
Congreso Geologico Argentino y Tercer Congreso Peninsula Antartica durante el Eoceno. Ameghini-
de Hidrocarburos, 1, 27-48. ana, 26, 245.
JARZEMBOWSKI, E. A. & Ross, A. J. 1996. Insect origi- LAZA, J. H., GENISE, J. F. & BOWN, T. M. 1994. Arqui-
nation and extinction in the Phanerozoic. In: tectura y origen de Monesichnus ameghinoi Roselli,
HART, M. B. (ed.) Biotic Recovery from Mass revelada por tomografia computada. Ameghini-
Extinction Events. Geological Society, London, ana, 31, 397.
Special Publications, 102, 65-78. LEA, A. M. 1925. Notes on some calcareous insect
JOHNSTON, P. A., EBERTH, D. A. & ANDERSON, P. K. puparia. Records of the South Australian Museum,
1996. Alleged vertebrate eggs from Upper Cretac- 3, 35-36.
eous redbeds, Gobi Desert, are fossil insect LEE, K. & WOOD, T. 1971. Termites and Soils.
(Coleoptera) pupation chambers: Fictovichnus Academic Press. London.
new ichnogenus. Canadian Journal of Earth LENGERKEN, H. v. 1955. Die Brutbirnen von Delto-
Sciences, 33, 511-525. chilum (Coleopt. Lamellicornia). Wissenschaftliche
KEIGHLEY, D. G. & PICKERILL, R. K. 1994. The Zeitschrift der Martin-Luther-Universitat Halle-
ichnogenus Beaconites and its distinction from Wittenberg, 4, 933-940.
Ancorichnus and Taenidium. Palaeontology, 37, LOIACONO, M. S. & MARVALDI, A. E. 1994. Biologia y
305-337. danos ocasionados. In: LANTERI, A. A. (ed.) Bases
KRAGLIEVICH, L. 1932. Una gigantesca ave fosil del para el control integrado de los gorgojos de la alfalfa.
Uruguay, Devincenzia gallinai. Anales del Museo Ediciones De La Campana, La Plata, 49-56.
Nacional de Montevideo (serie 2), 3, 323-353. MACHADO, A. DE B. 1983. The contribution of termites
KRELL, F. 2000. The fossil record of Scarabaeoidea of to the formation of laterites. Proceedings of the
the Mesozoic and Tertiary (Coleoptera: Poly- Second Seminar on Lateritisation Process, Sao
phaga). Invertebrate Taxonomy, 14, 871-905. Paulo, 261-270.
KRISHNA, K. 1990. Isoptera. In: GRIMALDI, D. A. (ed.) MANUM, S. B., BOSE, M. N. & SAWYER, R. T. 1991.
Insects from the Santana Formation, Lower Cretac- Clitellate cocoons in freshwater deposits since the
eous, Brazil. Bulletin of the American Museum of Triassic. Zoologica Scripta, 20, 347-366.
Natural History, 195, 76-81. MARTINEZ, S. 1982. Catalogo sistematico de los
KUSCHEL, G. 1983. Past and present of the relict family insectos fosiles de America del Sur. Revista de la
Nemonychidae (Coleoptera, Curculionoidea). Facultad Humanidades y Ciencias (serie Ciencias
GeoJournal, 7.6, 499-504. de la Tierra), 1, 29-84.
LABANDEIRA, C. C. 1998. The role of insects in the late MELCHOR, R. N., GENISE, J. F. & VERDE, M. 2001.
Jurassic to middle Cretaceous ecosystems. In: Invertebrate trace fossils from Triassic continental
LUCAS, S. G., KIRKLAND, J. I. & ESTEP, J. W. sequences of San Juan province, Argentina. Publi-
(eds) Lower and Middle Cretaceous Terrerstrial cacion Especial de la Asociacion Paleontologica
Ecosystems. New Mexico Museum of Natural Argentina, 1, 127-132.
History and Science Bulletin, 14, 105-124. MELCHOR, R. N., GENISE, J. F. & MIQUEL, S. 2002.
LABANDEIRA, C. C. & SEPKOSKI, J. J. 1993. Insect Ichnology, sedimentology and paleontology of
diversity in the fossil record. Science, 261, 310- Eocene calcareous palaeosols from a palustrine
315. sequence, South West La Pampa, Central Argen-
LAZA, J. H. 1982. Signos de actividad atribuibles a tina. Palaios, 17, 16-35.
Atta (Myrmicidae) en el Mioceno de la Provincia MICHENER, C. D. 1974. The Social Behavior of Bees.
de La Pampa, Republica Argentina. Signification The Belknap Press of Harvard University, Cam-
paleozoogeografica. Ameghiniana, 19, 109-124. bridge, MA.
452 J. F. GENISE

MICHENER, C. D. 1979. Biogeography of the bees. RETALLACK, G. J. 1990a. The work of dung-beetles and
Annals of the Missouri Botanical Garden, 66, its fossil record. In: BOUCOT, A. J. (ed.) Evolution-
277-347. ary Paleobiology of Behaviour and Coevolution.
MIKHAILOV, K., SABATH, K. & KURZANOV, S. 1994. Elsevier, Amsterdam, 214—226.
Eggs and nests from the Cretaceous of Mongolia. RETALLACK, G. J. 1990b. Soils of the Past. Unwin
In: CARPENTER, K., HIRSCH, K. F. & HORNER, J. R. Hyman, Boston.
(eds) Dinosaur Eggs and Babies. Cambridge RETALLACK, G. J. 1991. Miocene Palaeosols and Ape
University Press, Cambridge, 88-115. Habitats of Pakistan and Kenya. Oxford Univer-
MILLER, W. R. & MASON, T. R. 2000. Stellavelum sity Press, Oxford.
arborensis igen., isp. nov., Stellavelum uncinum RETALLACK, G. J. 2001a. Soils of the Past (2nd edn).
igen., isp. nov. and Termitichnus namibiensis isp. Blackwell. Oxford.
nov.; new ichnofossils from Cenozoic deposits RETALLACK, G. J. 200 Ib. Scoyenia burrows from
of Namaqualand, South Africa. Ichnos, 7, 195- Ordovician palaeosols of the Juniata Formation
215. in Pennsylvania. Palaeontology, 44, 209-235.
MYANNIL, R. M. 1966. Concerning vertical burrows RETALLACK, G. J., BESTLAND, E. A. & FREMD, T. 2000.
formed in Ordovician limestones of the Baltic Eocene and Oligocene Palaeosols of Central
region (in Russian). In: GEKKER, P. F. (ed.) Orga- Oregon. Special Papers of the Geological Society
nizm i sreda v geologicheskom proshlom. Nauka, of America, 344, 1-192.
Moscow and Lenningrad, 200-207. RINDSBERG, A. K. & MARTIN, A. J. 2003. Arthrophycus
NEL, A., PERRAULT, G., PERRICHOT, V. & NERAUDEAU, in the Silurian of Alabama (USA) and the problem
D. (in press). The oldest ant in the Lower Cretac- of compound trace fossils. Palaeogeography,
eous amber of Charente-Maritimes (SW France) Palaeoclimatology, Palaeoecology, 192, 187-219.
(Insecta: Hymenoptera: Formicidae). Ada Geolo- RITCHIE, J. M. 1987. Trace fossils of burrowing
gica Hispanica. Hymenoptera from Laetoli. In: LEAKEY, D. M. &
NOIROT, C. 1970. The nests of termites. In: KRISHNA, K. HARRIS, J. M. (eds) Laetoli, a Pliocene Site in
& WEESNER, F. M. (eds) Biology of Termites, Northern Tanzania. Oxford University Press,
Volume 2. Academic Press, New York, 73-126. Oxford, 433^38.
PASCUAL, R. & BONDESIO, P. 1981. Sedimentitas ceno- ROSELLI, F. L. 1939. Apuntes de geologia y paleon-
zoicas. Relatorio del Octavo Congreso Geologico tologia uruguaya. Sobre insectos del Cretaceo del
Argentine, 117-153. Uruguay o descubrimientos de admirables
PAZOS, P. J., TOFALO, O. R. & SANCHEZ-BETTUCCI, L. instintos constructivos de esa epoca. Boletin de
2002. El Neocretacico-Terciario de la cuenca la Sociedad Amigos de las Ciencias Naturales
Chacoparanaense en el Uruguay: aspectos estrati- 'Kraglievich-Fontana', I, 72-102.
graficos y paleoambientales. Resumenes de la ROSELLI, F. L. 1976. Contribucion al estudio de la geo-
Novena Reunion Argentina de Sedimentologia, paleontologia de los departamentos de Colonia y
34. Soriano Uruguay. IMCO (ed.), Montevideo.
PEMBERTON, S. G., FREY, R. W. & BROMLEY, R. G. ROSELLI, F. L. 1987. Paleoicnologia: nidos de insectos
1988. The ichnotaxonomy of Conostichus and fosiles de la cubertura Mesozoica del Uruguay.
other plug-shaped ichnofossils. Canadian Journal Publicaciones del Museo Municipal de Nueva
of Earth Sciences, 25, 866-892. Palmira, 1, 1-56.
PICKERILL, R. K. 1994. Nomenclature and taxonomy of SAKAGAMI, S. F. & MICHENER, C. D. 1962. The Nest
invertebrate trace fossils. In: DONOVAN, S. K. (ed.). Architecture of the Sweat Bees (Halictinae). Uni-
The Palaeobiology of Trace Fossils. Wiley, New versity of Kansas Press, Lawrence, Kansas, 135.
York, 3-42. SANDS, W. A. 1987. Ichnocoenoses of probable termite
RATCLIFFE, B. C. & FAGERSTROM, J. A. 1980. Inverte- origin from Laetoli. In: LEAKEY, D. M. & HARRIS,
brate lebensspuren of Holocene floodplains: their J. M. (eds) Laetoli, a Pliocene Site in Northern Tan-
morphology, origin and paleoecological signifi- zania. Oxford University Press, Oxford, 409^4-33.
cance. Journal of Paleontology, 54, 614-630. SAUER, W. 1955. Coprinisphaera ecuador• ensis, un fosil
READ, J. F. 1974. Calcrete deposits and Quaternary singular del Pleistocene. Boletin del Instituto de
sediments, Edel Province, Shark Bay, Western Ciencias Naturales del Ecuador, 1, 123-132.
Australia. Memoirs of the American Association SAUER, W. 1956. Coprinisphaera ecuadoriensis (Bola de
of Petroleum Geologists, 22, 250-282. Cangahua) y las esferas elaboradas actualmente
RETALLACK, G. J. 1976. Triassic palaeosols in the por escarabajos de la familia Scarabaeidae. Boletin
Upper Narrabeen Group of New South Wales. de Informaciones Cientificas Nacionales, 8, 550-
Part I: features of the palaeosols. Journal of the 555.
Geological Society of Australia, 23, 383-399. SAUER, W. 1959. Merkwiirdige Kugeln in Tuffen
RETALLACK, G. J. 1980. Middle Triassic megafossil Ecuadors und ihre Deutung. Natur und Volk, 89,
plants and trace fossils from Tank Gully, Canter- 118-124.
bury, New Zealand. Journal of the Royal Society SCHAEFER, C. E. 2001. Brazilian latosols and their B
of New Zealand, 10, 31-63. horizon as long-term biotic constructs. Australian
RETALLACK, G. J. 1984. Trace fossils of burrowing Journal of Soil Research, 39, 909-926.
beetles and bees in an Oligocene palaeosol, SCHOLTZ, C. H. 1988. Biology of Sparrmannia flava
Badlands National Park, South Dakota. Journal Arrow (Coleoptera: Scarabaeidae: Melolonthi-
of Paleontology, 58, 571-592. nae). The Coleopterists Bulletin, 42, 57-62.
ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 453

SCHUSTER, M., DURINGER, P., NEL, A., BRUNEI, M., STEPHEN, W. P., BOHART, G. E. & TORCHIO, P. F. 1969.
VIGNAUD, P. & MACKAYE, H. T. 2000. Decouverte The Biology and External Morphology of Bees, with
de termitieres fossiles dans les sites a vertebras du a Synopsis of the Genera of Northwestern America.
Pliocene tchadien: description, identification et Agricultural Experimental Station, Oregon State
implications paleoecologiques. Comptes Rendus University, Corvallis, Oregon.
Academic des Sciences de Paris (Sciences de la STUART, A. M. 1969. Social behavior and communica-
Terre et des Planetes), 331, 15-20. tion. In: KRISHNA, K. & WEESNER, F. M. (eds)
SCIUTTO, J. C. & MARTINEZ, R. D. 1996. El Grupo Biology of Termites, Volume 1. Academic Press,
Chubut en el anticlinal Sierra Nevada, Chubut, New York, 193-232.
Argentina. Actas del Decimotercer Congreso Geo- TANDON, S. K. & NAUG, B. 1984. Facies-trace fossil
logico Argentina y Tercer Congreso de Exploracion relationship in a Plio-Pleistocene fluvial sequence:
de Hidrocarburos, 1, 67-75. The Upper Siwalk Subgroup, Punjab Sub-Hima-
SKELLEY, P. E. 1991. Observations on the biology of laya, India. Palaeogeography, Palaeodimatology,
Stephanucha thoracica Casey (Coleoptera: Scara- Palaeoecology, 47, 277-299.
baeidae: Cetoniinae). The Coleopterists Bulletin, TAUBER, A. A. 1996. Los representantes del genero
45, 176-188. Protypotherium (Mam. Notoungulata, Intera-
SMITH, R. M. H. & KITCHING, J. 1997. Sedimentology theridae) del Mioceno Temprano del sudoeste de
and vertebrate taphonomy of the Tritylodon la provincia de Santa Cruz. Misceldnea de la
Acme Zone: a reworked palaeosol in the Lower Academia Nacional de Ciencias de Cordoba, 95,
Jurassic Elliot Formation, Karoo Supergroup, 3-29.
South Africa. Palaeogeography, Palaeoclimatol- THACKRAY, G. D. 1994. Fossil nest of sweat bees
ogy, Palaeoecology, 131, 29—50. (Halictinae) from a Miocene palaeosol, Rusinga
SMITH, R. M. H. & MASON, T. R. 1998. Sedimentary Island, western Kenya. Journal of Paleontology,
environments and trace fossils of Tertiary oasis 68, 795-800.
deposits in the Central Namib Desert, Namibia. VERDE, M. & MARTINEZ, S. 2004. A new ichnogenus for
Palaios, 13, 547-559. crustacean trace fossils from the late Miocene
SMITH, R. M. H., MASON, T. R. & WARD, J. D. 1993. Camacho Formation of Uruguay. Palaeontology,
Flash-flood sediments and ichnofacies of the 47, 39^9.
Late Pleistocene Homeb Silts, Kuiseb River, VEROSLAVSKY, G. & MARTINEZ, S. 1996. Registros
Namibia. Sedimentary Geology, 85, 579-599. no depositacionales del Paleoceno-Eoceno del
SPALLETTI, L. A. & MAZZONI, M. M. 1977. Sedimento- Uruguay: nuevo enfoque para viejos problemas.
logia del Grupo Sarmiento en un perfil ubicado al Revista de la Universidade Guarulhos (Serie Geo-
sudeste del Lago Colhue Huapi, provincia de ciencias), 1, 32—41.
Chubut. Obra del Centenario del Museo de La WILF, P. & LABANDEIRA, C. C. 1999. Response of
Plata (Geologia), 4, 261-285. plant-insect associations to Paleocene-Eocene
SPRECHMANN, P., Bossi, J. & DA SILVA, J. 1981. Cuen- warming. Science, 284, 2153-2156.
cas del Jurasico y Cretacico del Uruguay. In: ZACHOS, J., PAGANI, M., SLOAN, L., THOMAS, E. &
VOLKHEIMER, W. & MusACCHio, E. (eds) Cuencas BILLUPS, K. 2001. Trends, rhythms, and aberra-
Sedimentarias del Jurasico Cretacico de America tions in global climate 65 Ma to present. Science,
del Sur, Volumen 1, 239-270. 292, 686-693.
This page intentionally left blank
A stratigraphy of marine bioerosion
RICHARD G. BROMLEY

Geological Institute, University of Copenhagen, 0ster Voldgade 10,


DK-1350 Copenhagen K, Denmark (e-mail: rullard@geo.geol.ku.dk)

Abstract: About 65 ichnogenera and a number of bioerosional trace fossils that are unnamed
are catalogued with respect to their stratigraphic ranges. In most cases, corresponding
stratigraphic studies of the trace-makers are not possible because (1) the rank of taxonomic
ascription is too high to be meaningful and (2) not all members of a high taxon are bio-
eroders. For example, radulation traces of chitons are known from Jurassic to Recent,
whereas chitons have a body fossil record back to the Early Palaeozoic. Similarly, whereas
the round drill-hole Oichnus paraboloides is known from Cambrian to Recent, the only
identified makers of this trace fossil, naticid gastropods, range from Cretaceous to Recent.
The stratigraphic ranges of bioerosion ichnotaxa emphasize the two marine revolutions of
the Phanerozoic: there is marked increase in diversification during the Ordovician-Devonian
interval and since the Triassic.

Most groups of trace fossils are generally consid- today. Many ichnotaxa are badly in need of revi-
ered of little use as stratigraphic indicators, sion; many forms in the 'grey area' that surrounds
because they provide more information on the ichnology have been described but not named
behaviour of the trace-maker than on its bio- (e.g. damage to prey skeletons during attempts
logical identity, and because most ichnotaxa are at predation); some apparent trace fossils, such
stratigraphically long-ranging. Among the excep- as embedment structures (e.g. Tremichnus), may
tions to this are the hard substrate trace fossils and not strictly be trace fossils at all; and some trace
their bioeroding progenitors; but even within this fossils are very poorly known or understood and
group, many trace fossils are long-ranging and therefore provide minimal stratigraphic or bio-
remain biologically anonymous. logical information.
Some attempts at treating bioerosional ichno- In her unpublished thesis, Plewes (1996) made
taxa stratigraphically have been made before. good progress in the revision of several ichno-
Kobluk et al (1978) covered the Early Palaeo- taxa, having had singular success in borrowing
zoic; Wilson & Palmer (1990, 1992) and Palmer type material from museums. Some of her
(1982, fig. 5) the Cambrian to Cretaceous; taxonomic discoveries are followed in the
Ekdale et al (1984, fig. 10.12) had a single nomenclature used in this paper.
figure; and Bromley (1994) included a general-
ized statement. Taylor & Wilson (2003) cited
the stratigraphic range of the more important Attribution to a trace-maker
macrobioerosion ichnotaxa in their table 2.
Palmer (1982) expressed pessimistic feelings as Attempts at attribution of hard-substrate trace
he embarked on a synthesis of hardground com- fossils to the organism that produced them
munities through time. Nevertheless, despite a involve all degrees of precision. A few can be
widespread lack of consensus on the nomencla- ascribed to individual biological species (e.g.
ture of hard-substrate trace fossils, and a highly several endolithic algae and Cyanobacteria).
variable degree of accuracy in the identification Indeed, considering the far greater potential for
of the trace-makers, an updating of the strati- preservation as fossils that these microborings
graphy of bioerosional trace fossils is attempted have over their trace-makers, it is surprising
here. these trace fossils were not included in The
In the present study, all modes of bioerosion are Fossil Record 2 (Benton 1993).
treated, i.e. internal bioerosion (boring, duro- In the case of most endolithic Bryozoa, the
phagy) and external bioerosion (scratching, etch- boring moulds the external shape of the animal
ing) at all scales (macro, meso and micro). On precisely. This has led bryozoologists to treat
the other hand, no attempt is made at complete- the borings as body fossils and, therefore, the
ness of treatment. Revision of the bioerosional names as biological taxa (Pohowsky 1974).
ichnotaxa continues, but still leaves much to be Although Taylor (1993) considered this an
desired. Kobluk et al (1978, p.163) said 'the 'unfortunate dual nomenclature', as long as the
taxonomy of macroborings is sadly confused same name is used by bryozoologists as ichnolo-
and crude,' and much the same might be said gists there need be little misunderstanding. This

From: MC!LROY, D. (ed.) 2004. The Application of Ichnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 455-479. 0305-8719/04/S15.00 © The Geological Society
of London.
Fig. 1. Stratigraphic ranges of bioerosive ichnotaxa and some other bioerosive structures. The chart has been coloured so as to emphasize the possible four 'ages of
bioerosive activity'.
A STRATIGRAPHY OF MARINE BIOEROSION 457

perfection of body moulding by the boring is also microborings of endolithic Cyanobacteria


seen in the acrothoracican barnacles, microbial having been found in the Proterozoic (Campbell
borers and phoronids (Plewes 1994). Tavernier 1982; Golubic et al. 1985, 1999; Knoll et al. 1986,
et al. (1992) have commented in depth on this 1989). The earliest is dated at 1700 Ma (Zhang &
situation in respect of microbial borings. Golubic 1987, Fig. 1) or more probably 1500 Ma
At the other end of the scale are the borings (Golubic et al. 1999). These Proterozoic borings
of sponges and worms, where the accuracy of (some containing bodily preserved borers) are
trace-maker identification in some cases leaves identical to living cyanobacterial borings today.
even the phylum in doubt. Nevertheless, some Although earlier studies were mostly made
attributions are wildly exaggerated. Rigby et al. using thin sections, for example by Hessland
(1993), apparently considering 'Clionidae' and (1949) in Ordovician limestones, real advances
'boring sponge' to be synonymous (as did de in biotaxonomy and ichnotaxonomy were made
Laubenfels 1955 in the Treatise on Invertebrate only when three-dimensional casts of the micro-
Paleontology), extended the range of Clionidae borings could be made in resin (Golubic et al.
from Early Cambrian to Recent on the basis of 1975; Golubic 1990).
trace fossils. It seems unlikely, however, that The morphology of the borings of micro-
the family arose before Jurassic and possibly organisms in many cases is species-specific and
much later. allows precise identifications with living trace-
Hard substrate trace fossils are classically making species. However, some forms are
divided into macroborings versus microborings, extinct, having no living counterparts. Ichnotaxa
for the study of which a hand lens or a scanning based on boring morphology are now used, so
electron microscope is used, respectively. How- that all forms are treated alike as trace fossils.
ever, as Taylor et al. (1999) pointed out, there Indeed, it is good to see that a description of a
is an intermediate category (mesoborings) for new species of endolithic alga, Hyella vacans,
which the optical microscope is a necessary includes the morphology of its boring (Gektidis
tool. There are no firm boundaries between & Golubic 1996); it is Recent and so, as unfossi-
these size classes, and they are not used here. In lized material may not form the basis of new
the following, ichnogenus is the basic rank for trace fossil taxa, the boring itself is not named.
treatment. Ichnogenera are grouped either
under major trace-maker group (bivalve, Cavernula Radtke 1991
sponge, etc.) and trace type (boring, surficial The ichnogenus Cavernula is represented so far
etching trace, etc.), or, in trace fossil types by one species. Its morphology is similar to the
where the trace-makers are unknown or variable, codiolum stage of the green alga Gomontia poly-
in morphological groups (e.g. 'small round holes rhiza. Cavernula pediculata has been described
in shells'). Ichnofamilies have been characterized from the Palaeogene (Radtke 1991), the Cretac-
by some workers for a few groups but not all, so eous (Hofmann 1996), the Jurassic (Glaub
these are not used here. 1994) and the Triassic (Schmidt 1992).
Stratigraphic ranges of the ichnogenera and
various other trace fossils are arranged in Eurygonum Schmidt 1992
Figure 1. The only species of this ichnogenus represents a
boring similar to that of the living cyanobacter-
ium Mastigocoleus testarum. It has been reported
Algal, cyanobacterial and fungal borings from the Upper Jurassic (Glaub 1994), the
Triassic (Schmidt 1992; Balog 1996) and the
The microborings produced by endolithic photo- Permian (Balog 1996).
autotrophic microorganisms (algae and Cyano-
bacteria) are ubiquitous in carbonate substrates Fasciculus Radtke 1991
of the illuminated seafloor today, and the chemo- This ichnogenus represents the borings of several
heterotrophs (mainly fungi) occur in virtually all species of the living cyanobacterium Hyella.
marine environments. Fungal borings are However, one ichnospecies, F. grandis Radtke,
common over a broad bathymetric spectrum. has been compared with the endolithic rhizoid
Algal, cyanobacterial and fungal borings have of the green dasycladacean macro-alga Aceta-
therefore received much attention as indicators bularia crenulata (Radtke et al. 1997). Fasciculus
of palaeobathymetrical conditions (e.g. Glaub is known from the Palaeogene (Radtke 1991), the
et al. 2001,2002). Cretaceous (Hofmann 1996), the Upper Jurassic
The fossil record of these microorganisms has (Glaub 1994), the Triassic (Schmidt 1992; Balog
received close study over the last few decades and 1996), the Permian (Balog 1996) and the Silurian
their trace fossils are known to be the oldest, (Bundschuh 2000).
458 R. G. BROMLEY

Hyellomorpha Vogel, Golubic & Brett 1987 (Glaub 1994; Hofmann 1996), Jurassic (Glaub
The shape of this trace fossil is also considered to 1994), Triassic (Schmidt 1992; Balog 1996), Per-
closely resemble that of the boring of the young mian (Balog 1996) and Silurian (Bundschuh
thallus of the living cyanobacterium Hyella. 2000).
This form is described from the Late Cretaceous
(Maastrichtian) by Schnick (1992) and the Polyactina Radtke 1991
Devonian by Vogel et al (1987). This ichnogenus represents the borings of fungi
of, among others, the genus Conchyliastrum. It
Orthogonum Radtke 1991 is known from the Middle Eocene to Upper Oli-
Named for its somewhat right-angular branching gocene (Radtke 1991), the Cretaceous (Hofmann
pattern, this ichnogenus comprises three rather 1996), the Middle Jurassic (Glaub 1994), the
contrasting ichnospecies. O. tubulare Radtke is Triassic (Schmidt 1992; Balog 1996), the Permian
considered to be the work of an unknown (Balog 1996) and the Silurian (Bundschuh 2000).
heterotrophic endolith. O. spinosum Radtke
also appears to be the boring of a heterotrophic Reticulina Radtke 1991
microorganism, and O. fusiferum Radtke is This microboring has been compared with that
compared with the heterotrophic fungus Ostra- of the living green alga, Ostreobium quekettii. It
coblabe implexa. Schmidt (1992) added an ichno- occurs in the Lower Eocene and Oligocene
species considered comparable to a red alga, and (Radtke 1991), the Cretaceous (Hofmann
Glaub (1994) has added three further ichno- 1996), the Jurassic (Glaub 1994), the Triassic
species, one compared with a modern hetero- (Schmidt 1992; Balog 1996) and the Silurian
troph of unknown taxonomic affiliation, (Bundschuh 2000).
whereas the other two ichnospecies have no
modern counterparts so far. Rhopalia Radtke 1991
Considering this mixture of different trace- This ichnogenus equates with borings of the
makers, it is unlikely that the stratigraphic green-algal genera Eugomontia and Phaeophila,
range of the ichnogenus will be meaningful; but and occurs in the Eocene and Oligocene
this is a general problem in ichnology. Ortho- (Radtke 1991), the Cretaceous (Hofmann
gonum has been reported from the Palaeogene 1996), the Upper Jurassic (Glaub 1994) and the
(Radtke 1991), the Cretaceous (Glaub 1994; Triassic (Schmidt 1992).
Hofmann 1996), the Upper Jurassic (Glaub
1994), the Triassic (Schmidt 1992) and the Silur- Saccomorpha Radtke 1991
ian (Bundschuh 2000). Schmidt (1992) consid- Representing the borings of the living fungal
ered a microboring in Ordovician ostracods genera Dodgella, Lithopythium and Phytho-
described by Olempska (1986) to be conspecific phthora, this ichnogenus occurs in the Eocene
with his O. tripartitum. and Oligocene (Radtke 1991), the Cretaceous
(Glaub 1994; Hofmann 1996), Middle and
Palaeoconchocelis Campbell, Kazmierczak & Upper Jurassic (Glaub 1994) and the Triassic
Golubic 1979 (Schmidt 1992).
This name covers the bodily remains of the endo-
lithic phase of the bangiacean red alga Porphyra Scolecia Radtke 1991
nereocystis. Borings of this type are common This ichnogenus contains ichnospecies that com-
today and have been described from the Palaeo- pare with both green algae and Cyanobacteria. It
gene (Radtke 1991), the Upper Jurassic to Lower has been described from the Eocene and Oligo-
Cretaceous (Glaub 1994), the Triassic (Schmidt cene (Radtke 1991), the Cretaceous (Hofmann
1992) and the Silurian (Bundschuh 2000). The 1996), the Jurassic (Glaub 1994), the Triassic
type material was Silurian in age (Campbell (Schmidt 1992; Balog 1996), the Permian
et al. 1979). It is so well preserved (deriving (Balog 1996) and the Silurian (Bundschuh 2000).
from a well core) that organic remains of the
boring rhodophyte are preserved. The name
applies to the alga, and in fact the boring remains Small resetted borings
unnamed. However, in the other references here,
the name has been applied to the boring. This group includes the Dendrina-\ikQ rosette
trace fossils, particularly common in the Cretac-
Planobola Schmidt 1992 eous and known otherwise from the Devonian.
The four ichnospecies are compared with The group has no modern counterparts (Hof-
modern borings of fungi and cyanobacteria. mann 1996). There is probably room for some
This ichnogenus is known from the Cretaceous synonymy among the following forms, and the
A STRATIGRAPHY OF MARINE BIOEROSION 459

group needs revising and rationalizing. T. J. other trace-makers. The structural plan is gener-
Palmer (personal communication 2003) consid- ally an anastomosing network of canals that in
ers it likely that all these forms are borings of most cases swell to form rounded chambers.
Foraminifera. Commonly the chambers dominate the boring
and obscure the design of the network. Numer-
Dendrina Quenstedt 1848 ous apertures connect the chambers and canals
The type ichnospecies, D. belemniticola Magde- to the substrate surface. The number of cham-
frau 1937, was compared to Hyellomorpha by bers ranges from a single cavity to hundreds.
Schnick (1992), who considered it likely to be The wall of the boring has a special sculpture
the work of an alga. Hofmann (1996), who intro- created by the removal of chips of substrate by
duced several new ichnospecies, found these the borer. The preservation potential of the
never to occur together with light-dependent boring in carbonate substrates is far superior to
borings (with one exception), suggesting that that of the opaline spicules of the skeleton of
the organism responsible for Dendrina ispp. was the sponge. Details of spiculation are necessary,
not a photoautotroph. Bertling & Insalaco both of megascleres and microscleres, however,
(1998) attributed an ichnospecies of Dendrina for the identification of the sponge, and so the
to boring foraminifera. This species-rich ichno- taxonomy of the borers is usually unknown.
genus has been reported from the Cretaceous Nevertheless, recent studies by Calcinai et al.
and Jurassic. (2003) indicate that the chip pattern on the
walls may show itself to be diagnostic to genus
Dendrorete Tavernier, Campbell & Golubic 1992 level.
The only ichnospecies, D. balani, resembles Clio- The characteristic wall sculpture allows even
nolithes Clarke, but lacks certain details of the the smallest sponge borings, possibly represent-
branching structure of that ichnogenus (Plewes ing borings by the earliest post-larval growth
1996). D. balani occurs in Pliocene barnacle stage, little larger than a millimetre, to be recog-
skeleton. nized (Bundschuh et al. 1989; Glaub 1994).
Nododendrina Vogel, Golubic & Brett 1987 Sponge borings become abundant in the
This and the following two ichnogenera were Mesozoic and remain so to today. Most of
considered junior synonyms of Clionolithes by these borings are referred to ichnogenus Entobia.
Plewes (1996), which she considered valid. However, a number of papers describing sponge
Certainly, Nododendrina nodosa Vogel, Golubic borings of Palaeozoic age usually do not refer
& Brett 1987 looks very like C. radicans Clarke these to an ichnotaxon. Mikulas (1994b), never-
1908, but this decision cannot be taken in this theless, has described a Devonian boring that
paper. Both Clionolithes, which was considered closely resembles Entobia. These Palaeozoic
a sponge boring by Clarke (1908) and Plewes sponge borings constitute an intriguing, but
(1996), and Nododendrina are based on Devonian little-known group of trace fossils.
material.
Entobia Bronn 1837
Platydendrina Vogel, Golubic & Brett 1987 Ichnogenus Uniglobites Pleydell & Jones 1988 is
Hofmann (1996) described a new ichnospecies considered a junior synonym of Entobia, having
from the Upper Cretaceous. The original but a single chamber. Several sponges today
description was of Devonian material (Vogel fuse their expanding chambers together with
et al 1987). growth, some ending with only a single chamber
(Bromley & D'Alessandro 1989).
Ramodendrina Vogel, Golubic & Brett 1987 Sponges of different families produce similar
Vogel et al. (1987) described Devonian material. borings. Sponges of the family Clionidae are
the dominant endolithic sponges today. How-
Globodendrina Plewes, Palmer & Haynes 1993 ever, species of other living groups of sponges,
The authors considered G. monile Plewes, Palmer e.g. the genus Aka [Siphonodictyon] of the
& Haynes 1993 to have been made by a foramini- family Adociidae, also produce borings that
feran. The monotypic ichnogenus is of Late may be included in Entobia (Riitzler 1971; Brom-
Jurassic age. ley & D'Alessandro 1989; Reitner & Keupp
1991).
The stratigraphic range of Entobia is
Sponge borings undoubted from Jurassic to today. Jurassic
entobians have been reported by many authors
The borings of sponges possess several character- (e.g. Fursich et al. 1994; Bertling 1999; Perry &
istics that distinguish them from the work of Bertling 2000).
460 R. G. BROMLEY

Reports of Triassic sponge borings are usually Worm borings


associated with question marks (Szulc 1990).
Resin casting, however, has revealed tiny borings Caulostrepsis Clarke 1908
showing sponge-chip wall ornament in the U-shaped borings that have a vane connecting
Triassic (Schmidt 1992). The Middle to Late the limbs of the U-boring. Polychaete annelids
Devonian entobian documented by Mikulas produce Caulostrepsis today. Spionid poly-
(1994b) is very convincing. Further convincing chaetes of the genus Polydora are the best-
Ordovician sponge borings have been reported documented producers, and this has led to the
by Kobluk (198la), less convincingly by Pickerill belief that all Caulostrepsis are produced by
& Harland (1984) and Lindstrom (1979). In Spionidae (e.g. Hantzschel 1975). However,
Lower Cambrian archaeocyath reefs, however, other polychaetes also produce these borings
Kobluk (1981b) described cavities showing today, such as the small eunicid Lysidice ninetta
scalloped walls, and associated sponge chips (Bromley 1978). Boring polychaetes are almost
and spicules. never preserved as body fossils, but see Cameron
It thus seems clear that boring sponges were (1969).
active, but rarely documented, throughout the The ichnogenus Caulostrepsis ranges from
Palaeozoic. Devonian (Clarke 1908) to Recent, and is abun-
dant in the Jurassic (e.g. Fiirsich et al. 1994) and
Cretaceous (e.g. Voigt 1971). Bodily preserved
Topsentopsis de Laubenfels 1955 spionid worms have been described from the
De Laubenfels erected this (ichno)-genus to Cambrian (Glaessner 1976), but very few taxa
replace the homonym Topsentia Clarke 1921 of that large family are borers.
(non Berg 1899, fide de Laubenfels 1955). He
himself regarded its sponge affinities as doubtful. Conchotrema Teichert 1945
The specimen consists of a cavity from which a This name is in common use. However, the reali-
few straight canals radiate. Plewes (1996) re- zation that Conchotrema is identical to and thus a
examined Clarke's holotype and concluded that junior synonym of Talpina (Voigt 1972; Plewes
the central structure was a random hole and 1996) renders it of uncertain value, and it will
not a trace fossil. She concluded that Topsentop- not be considered in this paper.
sis is a nomen dubium.
It is interesting that Mikulas (1992) has docu- Cunctichnus Fiirsich, Palmer & Goodyear 1994
mented Early Cretaceous structures having a This is a millimetre-wide boring having very
rather similar appearance to Topsentopsis. They short and stubby branches. The terminations of
are, however, much more regularly developed the branches have a stunted and slightly broa-
and more convincing as trace fossils; he named dened appearance. It has only been reported
them Entobia Solaris. from the Jurassic (Fiirsich et al. 1994; Bertling
& Insalaco 1998).
Clionolithes Clarke 1908 Helicotaphrichnus Kern, Grimmer & Lister 1974
The validity and coverage of this ichnogenus has
been discussed by many authors, and opinions The spionid worm Polydora commensalis lives
commensally together with hermit crabs, boring
are very varied (Clarke 1921; Solle 1938; Teichert
a Trypanites-likG tube up the columella of the
1945; Vogel et al. 1987). Plewes (1996) examined
shell. The trace fossil has been found in Recent,
Clarke's type material and concluded that the
ichnogenus is valid; surface sculpture of similar, Pleistocene and Pliocene material of California
and Baja California (Kern et al. 1974; Walker
better preserved material indicated that Cliono-
1989; Feige & Fiirsich 1991), in the Miocene of
lithes is likely to be the boring of a sponge.
Europe (Kern 1979; Baluk & Radwanski 1979a,
The trace fossil seems to occur chiefly in the
1979b), and in the Eocene of Mississippi
Devonian. (Walker 1992).

Cicatricula Palmer & Palmer 1977 Lapispecus Voigt 1970


Found in a Middle Ordovician hardground, Ichnogenus Lapispecus is a winding, cylindrical
Cicatricula retiformis Palmer & Palmer rather boring. It differs from Trypanites in having a
resembles Clionolithes and may also be placed short, lateral vane extending along it locally,
among the probable sponge borings. It differs especially on the inside of curves (Voigt 1970,
in covering a larger area and apparently lacking 1971; Bromley 1972). A single Early Pleistocene
a roof, being an open, etched network on the occurrence has been reported (Bromley &
substrate surface. D'Alessandro 1987).
A STRATIGRAPHY OF MARINE BIOEROSION 461

Maeandropolydora Voigt 1965 The small spiral bioclaustration structures


Voigt (1965) attributed Maeandropolydora, based Helicosalpinx Oekentorp 1969 (Devonian) and
on two ichnospecies, to spionid polychaetes Torquaysalpinx Plusquellec 1968 (Devonian),
because the morphology includes pouch-like considered commensals with tabulate corals
developments closely similar to Caulostrepsis and stromatoporoids (Stel 1976), somewhat
individuals. These pouches are connected by resemble Spirichnus. However, they are not
winding cylindrical tubes. branched and are probably not in any way
The ichnogenus has been identified in Recent related to that ichnogenus.
material (Feige & Ftirsich 1991), and three
further ichnospecies have been defined on the Talpina von Hagenov 1840
basis of Pleistocene material (Bromley & D'Ales- The work of Voigt (1972, 1975, 1978) demon-
sandro 1987). The trace fossil is common in the strated that Talpina networks are the borings of
Cretaceous (Voigt 1965; Wilson 1986) and Juras- phoronid worm pseudocolonies. He also demon-
sic (Hoffmann & Krobicki 1989; Bertling & Insa- strated (Voigt 1972) that Conchotrema Teichert
laco 1998), and possibly occurs in the Triassic 1945 is a junior synonym of Talpina, as was con-
(Schmidt 1992). Palaeozoic reports are less con- firmed by Plewes (1996).
vincing; Mikulas (1994a) identified Maeandropo- Talpina is abundant today and is well known
lydora of Devonian age and Bundschuh (2000) from the Cretaceous (Voigt 1972) and Jurassic
used the name for small fragments in the Silurian. (Fursich et al. 1994). The ichnogenus ranges
back to the Late Devonian (Thomas 1911;
Palaeosabella Clarke 1921 Rodriguez & Gutschick 1970).
Examination of type material led Plewes (1996)
to conclude that Palaeosabella Clarke is a valid Trypanites Magdefrau 1932
and useful name. This provides a name for The revalidation of the ichnogenus Palaeosabella
long, tubular borings that, in contrast to Trypa- has rendered many records of Trypanites
nites, expand distally as an acute cone. Palaeosa- untrustworthy (Plewes 1996; Palmer et al.
bella is thus a senior synonym for Specus 1997). These two forms are similar, Trypanites
Stephenson, a name that has been little used. being cylindrical whereas Palaeosabella slightly
This causes a problem in the present case expands distally. Thus only where the trace
because it means that some trace fossils named fossil has been described fully or illustrated in
Trypanites may in fact be Palaeosabella. Only full profile can a record be accepted. The state-
in very careful studies (e.g. Pemberton et al. ment that Trypanites is the oldest macroboring
1980) can this be detected, using three-dimen- and is abundant in the Early Cambrian may
sionally exposed material. still be true; James et al. (1977, fig. 3B) illustrated
Palmer et al. (1997) suggest that Trypanites is what does appear to be true Trypanites, but this
dominantly a post-Palaeozoic trace fossil, needs to be confirmed.
whereas Palaeosabella is chiefly distributed in Relatively fat and short Trypanites, about 1—
the Palaeozoic. For the moment, the following 5 mm in diameter, are produced today by sipun-
evidence is available. culan worms (Rice 1969; Bromley 1978), whereas
Some sipunculans make Palaeosabella-likQ thin and slender ones are bored by polychaetes
borings today (Rice 1969). The boring of a com- (Bromley 1978). Trypanites is common through-
mensal worm within the bivalve Corbula is clearly out the Cenozoic and Mesozoic (e.g. Bromley &
Palaeosabella, having a Miocene to Recent range D'Alessandro 1987; Cole & Palmer 1999).
(J. K. Nielsen 1999). The partly bioclaustrated Pickerill (1976) described (under the name
tubes in oysters that Voigt (1965) called Ostreo- Vermiforichnus Cameron 1969) Late Ordovician
blabe may be referred to Palaeosabella', they are borings in brachiopod shells that are probably
of Late Cretaceous age. Stephenson (1952) Trypanites. The oldest undoubted Trypanites
based Specus on Late Cretaceous material. On may be middle Ordovician (Kobluk & Nemcsok
the other hand, Cameron (1969) and Pemberton 1982). These authors recorded scolecodont
et al. (1980) described Palaeosabella from Devo- remains in the borings, which was considered to
nian material (using the names Vermiforichnus indicate a polychaete trace-maker.
and Trypanites, respectively). The record would
appear to be incomplete.
Bivalve mollusc and barnacle etchings
Spirichnus Fursich, Palmer & Goodyear 1994
These branched and axially spiralled tubes Centrichnus Bromley & Martinell 1991
appear to be known only from the Late Jurassic Very shallow etching scars on carbonate sub-
(Fursich et al. 1994; Bertling & Insalaco 1998). strates are produced by the basal surface of
462 R. G. BROMLEY

balanid cirripeds (Boekschoten 1967; Miller & (Pojeta & Palmer 1976). This is the earliest
Brown 1979). The rather deeper scars produced direct evidence of mytilid bioerosion. The
by verrucid cirripeds and the byssal plug of grooves, locally abundant, have since been
anomiid bivalves have been named C. concentri- found empty, i.e. lacking the bivalve, and have
cus and C. eccentricus respectively. Bromley received the name Petroxestes pera Wilson &
(1999) published a Pleistocene C. eccentricus Palmer 1988. Tapanila & Copper (2002) reported
that lay in situ directly beneath the byssal notch Petroxestes from the Silurian.
of its anomiid trace-maker. Recently, a single specimen of a groove in a
C. eccentricus occurs from Campanian to Miocene oyster shell has been identified as Pet-
Recent, and C. concentricus from Miocene to roxestes pera (Pickerill et al. 2001). Until further
Recent (Radwanski 1977; Bromley & Martinell convincing material is described, however, such
1991). The earliest anomiids from Middle and drastic extension of the range of this ichnospecies
Late Jurassic do not appear to produce Centrich- must remain uncertain.
nus (Fursich & Palmer 1982; Todd & Palmer
2002a). Teredolites Leymerie 1842
Clavate borings in wood (lignic) substrates have
apparently all been attributed to bivalve trace-
Bivalve borings makers (Kelly & Bromley 1984). These belong
to two families: the Teredinidae (shipworms)
Gastrochaenolites Leymerie 1842 from Early Cretaceous to today, and the Phola-
This ichnogenus includes clavate borings in lithic didae from Middle Jurassic to today (Kelly
substrates. These are mostly borings of endo- 1988; Evans 1999). At present, two ichnospecies
lithic bivalves, but other trace-makers may be are recognized. T. clavatus Leymerie is produced
involved. It should be noted that Recent corallio- by species ofMartesia and Xylophaga today, and
philid (=magilid) gastropods bore chambers in fossil trace-makers include Martesia, Xylophaga
coral that might be included in Gastrochaenolites, and Opertochasma, whereas T. longissimus
although fossil examples of these do not appear Kelly & Bromley is produced by the teredine
to have been described (Wenz 1939; Gohar & shipworms.
Soliman 1963; Soliman 1969; Bromley 1970a, Both ichnospecies become abundant by the
fig. 4d; Massin 1982, 1983, 1988, 1990, 1992; end of the Cretaceous. For example, Mikulas
Zibrowius & Arnaud 1995). Nevertheless, some et al. (1995) documented crowded individuals
of these gastropod borings differ from Gastro- of T. clavatus of Cenomanian age, and Huggett
chaenolites in having an extra canal diverging et al. (2000) recorded a diversity of six teredinid
from near the aperture (Massin 1987). Some species producing T. longissimus in an Eocene
sipunculan worms may also produce Gastrochae- community.
nolites (Rice 1969; Ekdale et al. 1984).
Gastrochaenolites becomes abundant and
diverse in the Jurassic (e.g. Kelly & Bromley Gastropod borings and surficial etchings
1984; Oschmann 1989). The oldest Gastrochaeno-
lites containing its trace-maker (Lithophaga sp.) It was noted above that some gastropods bore
is Triassic (Kleemann 1994). Another Gastro- cavities in coral substrates today that might be
chaenolites producer, Gastrochaena, has its assigned to ichnogenus Gastrochaenolites. One
bodily first appearance also in the Triassic (J. G. group of boring gastropods, however, produces
Carter in Wilson & Palmer 1990). borings that are characteristic enough to be
Early Pennsylvanian Gastrochaenolites have assigned an ichnogeric name: the vermetid gas-
been described from rocky shore environments tropods (Lamy 1930; Bromley 1970a; Savazzi
(Webb 1993, 1994; Wilson & Palmer 1998). 1996).
Although in this case the borer is not preserved,
the morphology of these (G. anauchen) corre- Renichnus Mayoral 1987b
sponds exactly to that of bivalve borings. The Mayoral (1987b) assigned the name R. arcuatus
oldest Gastrochaenolites, also lacking its trace- to etchings produced by vermetids that spiralled
maker, is G. oelandicus of Early Ordovician age at an angle to the substrate surface. As a result,
(Ekdale & Bromley 2001; Ekdale et al. 2002). the substrate was etched only where the spiral
came into contact with the surface. This pro-
Petroxestes Wilson & Palmer 1988 duced a series of kidney-shaped depressions -
The mytilid bivalve Coralliodomus was found hence the name. However, many vermetids
fitted well within short, deep grooves on the spiral in the plane of the interface, in which
under side of Late Ordovician stromatoporoids case the whole spiral is etched into the substrate.
A STRATIGRAPHY OF MARINE BIOEROSION 463

As all intermediates are found, the one name is Lowenstam 1971) and is therefore considerably
probably adequate for all vermetid etchings harder than calcite. Similarly, chiton radulae
(Radwanski 1977). contain denticles hardened with magnetite
The depth to which the gastropod etches itself (Lowenstam 1962). The radulae of these molluscs
into the substrate varies, and these trace fossils are therefore adapted to surficially eroding
are intermediate between borings and surface carbonate substrate and allowing the animals
scars. to exploit the endolithic microbes as a food
Renichnus is known from Pliocene to today source.
(Mayoral 1987b; Taddei Ruggiero 1999). The fine sculpture that this activity imparts to
the substrate has a relief in the region of 100 jam
and therefore has a fair fossilization potential.
Gastropod etched and rasped pits and scars The sculpture has been named Radulichnus inopi-
natus Voigt. This name covers both gastropod
Many different groups of gastropods etch or and chiton radulation, which can be distin-
abrade (or both) home depressions in their sub- guished on the basis of morphology. The
strate. Most celebrated are the homing scars of holotype is a gastropod radulation sculpture.
intertidal patelliform gastropods (limpets) (e.g.
Orton 1914; Bromley & Hanken 1981; Lindberg Radulichnus Voigt 1977
& Dwyer 1982). Some modern limpets make pits The trace fossil is abundant and widespread
on the surface of other living molluscs (Vermeij today, both of chiton origin (e.g. Jiick &
1998). The great holes supposedly punched in Boekschoten 1980) and gastropod (e.g. Bromley
giant ammonites by Late Cretaceous playful or & Hanken 1981). Reports of fossil examples are
hungry mosasaurs (KaufTman & Kesling 1960; not common, undoubtedly because of their
Kauffman 1990), or due simply to implosion small scale, but the study of acmaeid Radulichnus
damage (Keupp 1991), have been reinterpreted on Cretaceous ammonites by Akpan et al. (1982)
as the homing pits of limpets (Kase et al. 1994, is outstanding. Radulichnus was also found in
1998; Seilacher 1998). association with the limpets living on ammonites
Scars produced on host shells beneath station- described by Kase et al (1998).
ary parasitic capulid gastropods are also well It is not known when Radulichnus was first
known today (Sharman 1956; Boss 1965; Brom- produced. Chitons (Runnegar et al. 1979) or
ley 1981), and are reported from the Pleistocene related forms (Stinchcomb & Darrough 1995)
(Matsukuma 1978). occur in the Late Cambrian. Radulation trace
Less well known are the scars produced by hip- fossils are known from the Cambrian, but only
ponicid gastropods on the shells of other living in matgrounds, not in hard substrates (Seilacher
gastropods. Vermeij (1998) described and illu- 1995; Bottjer 2000; Dornbos & Bottjer 2000;
strated Recent material and found correspond- Dornbos et al. in press).
ing pits in Early Pliocene and Late Miocene
material. The presence of hipponicid body fossils
in the same beds increased confidence in attribut- Bryozoans
ing the pits, but Noda (1991) actually found a
Pliocene hipponicid in place over its etched scar. Owing to the 'dual nomenclature' for bryozoans,
Pit etching has been very rarely mentioned in as mentioned above, the degree of 'splitting' as
association with the genus Crepidula (slipper lim- biotaxa is excessive in an ichnological context.
pets). However, Walker (1992) has found that Therefore the biofamily level has been chosen,
Crepidula adunca, which attaches to the external where this is known, and the following is based
surface of gastropod shells while the host snail is largely on the work of Pohowsky (1978) and
living and after hermit crabs have taken over, Taylor (1993).
etches a characteristic scar in the shell. Compar- Our present knowledge of stratigraphical
able scars have been found on Eocene gastropod ranges suggests that first appearances of endo-
shells. lithic bryozoan taxa occur rather smoothly
None of these scars have received ichnotaxo- through time. Most of these appear to belong
nomic treatment. to the Ctenostomata. In the following, the taxa
are discussed in stratigraphical order of first
occurrence, youngest first.
Gastropod and chiton radulation Mayoral (1987a) designated the ichnogenus
Stellichnus for the boring produced by the
The radula of most patellid and acmaeid gastro- Early Pliocene ctenostome bryozoan Paravinella
pods contains opal and goethite (Runham 1961; (Mayoral & Reguant 1995).
464 R. G. BROMLEY

The Immergentiidae and Terebriporidae both earliest occurrence of Leptichnus is Maastrich-


appear in the Late Cretaceous and continue to tian, Late Cretaceous.
Recent. Immergentia cruciata (Magdefrau 1937)
is the oldest known immergentiid, Santonian,
Late Cretaceous (Taylor 1993). Vogel et al Brachiopod etching
(1987) transferred /. devonica Richards (1974)
to Orbignyopora Pohowsky 1978. While the morphology of some attachment scars
Taylor (1993) gave the range for the Terebri- of pediculate brachiopods may vary (Bromley &
poridae as Late Cretaceous (Maastrichtian) to Surlyk 1973), the large majority conform to a
Recent. Mayoral (1988), considering Terebripora similar plan and have been covered by a single
a biogenus, has published the ichnogenus Pina- ichnospecies, Podichnus centrifugalis Bromley &
ceocladichnus as an ichnological equivalent. Surlyk, despite the wide taxonomic variety of
Mayoral et al. (1994) have claimed Ordovician the trace-making brachiopods. Where the etch-
orthids to contain Pinaceocladichnus, but this ing trace was emplaced in a living brachiopod,
form may be more closely related to the Ropalo- the shell can exhibit considerable reaction in
nariidae (Todd 2000). the form of wound tissue (Taddei Ruggiero
Foraripora Voigt & Soule 1973 comprises a 1999).
single ichnospecies, F. pesavis (somewhat resem-
bling a minute bird trackway). Its age is Late
Podichnus Bromley & Surlyk 1973
Cretaceous (Voigt 1979).
The Spathiporidae range from Jurassic to The trace fossil is usually a clearly visible meso-
Recent (Taylor 1993). Mayoral (1988), consider- boring etched shallowly into the substrate. How-
ever, several occurrences have been recorded in
ing Spathipora to be a biogenus, has published
epoxy casting of substrates, e.g. in the Triassic
the ichnogenus Pennatichnus as an ichnological
(Schmidt 1992; Glaub & Schmidt 1994), the
equivalent.
Devonian (Vogel et al. 1987) and the Silurian
The Penetrantiidae range from Triassic
(Bundschuh 2000). Alexander (1994) has also
(Rhaetic) to Recent (Taylor 1993).
Pohowsky (1978) declared Iramena danica recorded Podichnus from the Carboniferous.
The range is Silurian to Recent.
Boekschoten 1970 (Danian) a junior synonym
of Penetrantia. Fiirsich et al. (1994) recorded
Iramena isp., of Late Jurassic age.
The Cookobryozoonidae are known from two Crustacea: acrothoracican borings
localities only, of Triassic age, and the Voigtelli-
dae range from the Permian to Maastrichtian The borings of acrothoracican cirripeds are small
(Taylor 1993). cavities shaped somewhat like a sock, which in
Fischerella is restricted to the Late Mississip- many ichnospecies has a partial calcitic lining
pian and Casterella to the Late and Middle (Seilacher 1969; Tomlinson 1969). Biologists
Devonian (Pohowsky 1978). treat the fossil borings as body fossil moulds, in
The family Orbygnyoporidae ranges from the same way as the description of Bryozoa.
Silurian to Triassic (Taylor 1993), although Ichnologists, however, use the first 'genus' avail-
Cuffey et al. (1981) claimed Orbignyopora isp. able, Rogerella, as the ichnogenus (Bromley &
indet. from the Maastrichtian. D'Alessandro 1987; Plewes 1996).
The Ropalonariidae comprise one certain
(ichno)species, Ropalonaria venosa Ulrich 1879 Rogerella Saint-Seine 1951
from the Late Ordovician, the oldest known Acrothoracican borings are abundant today and
bryozoan boring. Two other (ichno)species, of back through the Mesozoic. Webb (1994) found
Late Jurassic and Early Cretaceous age respec- them in rocky shore environments of the Early
tively, possibly belong to Ropalonaria Mississippian. Rodriguez & Guttschick (1970)
(Pohowsky 1978). The latter two (ichno)species described them from Late Devonian material.
were accepted by Taylor (1993), who thereby The oldest examples may be those commensal
gave a range for the Ropalonariidae as Ash- with the gastropod Platyceras, itself parasitic
gillian to Aptian. on crinoids of Middle Devonian age (Baird
Leptichnus Taylor, Wilson & Bromley 1999 is a et al. 1990). The unabraded aperture of this
surficial etching trace of cheilostome bryozoans. form is much wider than is usual for Rogerella.
At least nine Recent species of cheilostome pro- Bundschuh (2000) reported Silurian acrothor-
duce it today. There are two ichnospecies. acican borings, but her illustrations are not
Palmer & Plewes (1993) published a fine illustra- very convincing. The range, then, is Devonian,
tion under the nomen nudum Bothrioichnus. The maybe Silurian, to Recent.
A STRATIGRAPHY OF MARINE BIOEROSION 465

Crustacea: ascothoracican borings comparatively weak and rare (Vermeij et al.


1982). Sublethal injury and repair was reported
Small, round holes, usually millimetric in size, in Carboniferous ammonoids by Bond & Saun-
are produced by endoparasitic ascothoracican ders (1989). Devonian brachiopods (Leighton
cirripeds in irregular echinoids. They tend to be 2001) and gastropods (Lindstrom & Peel 2003)
placed near the genital pores of the apical were predated by unidentified durophages, at
system, but are larger than these (Brattstrom which time another increase in predation level
1936, 1937, 1947). In some cases the holes lie at has been detected (Signor & Brett 1984). Elliot
other locations on the aboral surface of the & Brew (1988) found the beak of a nautiloid
echinoid (Madsen & Wolff 1965). lodged within the valve of a Devonian brachio-
The published fossil record of ascothoracican pod, and Peel (1984) described damage to
borings is negligible. This must be due in part Silurian gastropods that compared to that
to the great similarity of the borings to drill- inflicted by Sepia today, suggesting cephalopod
holes of predatory gastropods. Madsen & Wolff predation.
(1965) attributed two holes in the test of a Late There are also reports of Early Palaeozoic
Cretaceous holasteroid echinoid to ascothor- durophagy. Indeed, according to Peel et al.
acican parasites. Two round holes that Kier (1996) and Ebbestad & Peel (1997), Silurian
(1981) described in an Early Cretaceous spatan- and Ordovician gastropods show 'heavy
goid echinoid may be of ascothoracican origin, damage' by predators. Alexander (1986) consid-
although that author interpreted them as the ered nautiloids as likely producers of predator
work of parasitic gastropods. damage on Ordovician brachiopods. Cambrian
Voigt (1959, 1967) attributed cyst-like cavities, evidence of durophagy is dominated by bitten
having a slit-like entrance, in Danian octocorals trilobites (Babcock 1993; Pratt 1998).
to the parasitism of ascothoracicans, naming
the (unpreserved) animals Endosacculus moltkiae
and Endosacculus? najdini. These may be bio- Crustacean smashing traces
claustration structures.
Crustaceans produce still other forms of trace
fossils in hard substrates that are particularly dif-
Shell chipping and peeling ficult to classify. One of these, the durophagous
activities of Stomatopoda or mantis shrimps,
Chipping and peeling of shells is a common has received an ichnogenus, Belichnus Pether
source of trace fossils. It is a form of durophagy, 1995. These are holes produced in gastropod
the damage of hard skeletal material by preda- and bivalve prey skeletons by blows struck by
tion. Durophagy can be extremely difficult to the club-shaped chelipods of smasher stomato-
distinguish from physical or diagenetic damage pods. Such damage is difficult to distinguish
(Elliott & Bounds 1987; Lescinsky & Benninger from other damage unless there is evidence of
1994). impact. The inwardly bent manner of chipping
Today, shell chipping of bivalves and shell of the shell provides this evidence.
peeling of prey gastropods is dominantly the
work of crustaceans (Cadee 1968; Shoup 1968; Belichnus Pether 1995
Bishop 1975; Vermeij 1982, 1983b; Shigemiya Stomatopods are active durophagous carnivores
2003), but fish and sharks (see below) and even today (Caldwell & Dingle 1975, 1976). Pether
some asteroids (Mauzey et al. 1968) are known (1995) based the ichnotaxonomy on Holocene
to be durophagous. material, but Geary et al. (1991) interpreted
Among brachyuran crabs the habit is Plio-Pleistocene shell damage as stomatopod
especially abundant and, in one study, peeling activity, and Bamk & Radwanski (1996) have
frequency in a given area of seafloor was corre- extended this to the Miocene.
lated loosely with the number of resident species
of the shameface crab, Calappa (Vermeij et al.
1980; Vermeij 1982). However, shell peeling is Borings in coral and rock by Decapoda
also practised by many other genera of crabs.
The stratigraphy of shell chipping and peeling Many crustaceans bore various forms of cham-
is fairly straightforward. Durophagous shell bers in coral (e.g. Andre & Lamy 1933; Zibro-
breakage is abundant in the Neogene (Rad- wius 1984) and rock. The thalassinideans are
wanski 1977; Martinell 1989). In fact this perhaps best known: for example, Upogebia
abundance begins in the Jurassic, and Pre- operculata bores branched passages in coral
Jurassic shell breakers have been considered (Kleemann 1984; Scott et al. 1988; Williams &
466 R. G. BROMLEY

Ngoc-Ho 1990; Fonseca & Cortes 1998) and Monteillet et al. 1982; Nebelsick 1999). Corre-
Alpheus saxidomus produces sizable chambers spondingly, many cases of bite traces in the
within basalt (Holthuis 1980; Fischer & Meyer fossil record have been ascribed to fish and
1985). Such structures do not seem to have sharks. Fish grazing on coral reefs was consid-
been described from the fossil record. ered by Hubbard et al. (1990) to be chiefly impor-
tant from the base of the Palaeogene onward.
Ichnological nomenclature has not been
Echinoid borings attempted for this variable group of trace fossils.
Reports of trace fossils produced by such
Several species of regular echinoid bore bowl- activity, however, are particularly numerous in
shaped, hemispherical or deeper pits in rock sur- Cretaceous prey skeletons (e.g. KaufTman 1972;
faces in shallow water (Markel & Maier 1966, Thies 1985; Giessler 1991; Dortangs 1998; Neu-
1967; Warme 1975). The boring process is mann 2000). Older occurrences were reported
mechanical, so the activity is not restricted to by Holder (1973) on Early Jurassic belemnites
carbonate substrates: for example, Allouc et aL and Devonian ammonites, and by Hansen &
(1996) described Recent ones in dolerite, and Mapes (1990) on Carboniferous and Devonian
Abel (1935) in granite. nautiloids.

Circolites Mikulas 1992


Fossil structures interpreted as echinoid borings Small round holes in shells
have been reported many times from the Pliocene
(Martinell & Domenech 1986; Agusti et al 1990; A large variety of trace-makers make and have
Watkins 1990; Gibert et aL 1998), and Miocene made small round holes in shells and tests for a
examples were published by Radwanski (1965, variety of reasons. Some holes penetrate right
1969, 1970). Cenomanian material was described through the shell, others end blind because the
by Gtazek et al. (1971), Lewy & Avni (1988, in drilling process was interrupted, and still others
association with body fossils of the echinoid end blindly through intent. The purpose of
Goniopygus menardi) and Mikulas (1992). Juras- drilling the holes includes predation (penetrative
sic echinoid pits were reported by Fursich (1979). when complete), parasitism (either penetrative or
The range is from Jurassic to Recent. non-penetrative), or mere attachment to the sub-
strate (chiefly non-penetrative). This variability
has led to the designation of three ichnogenera:
Echinoid bite traces Oichnus Bromley 1981 for penetrative holes,
and Sedilichnus Miiller 1977 and Tremichnus
Bite traces produced by plucking and grazing Brett 1985 for non-penetrative pits. However,
regular echinoids are abundant in the present because ichnotaxa are based on morphology,
marine environment (e.g. Krumbach 1914; and because the two morphological forms are
Krumbein & Van der Pers 1974; Bak 1994). not clearly different, there has recently been dis-
They are less common in the fossil record, cussion as to whether Tremichnus should be con-
undoubtedly because of their shallow topogra- sidered a junior synonym of Oichnus (Pickerill &
phy and their poor preservation potential. Donovan 1998; Nielsen & Nielsen 2001, 2002;
Todd & Palmer 2002b; Donovan & Jagt 2002;
Gnathichnus Bromley 1975 Donovan & Pickerill 2002.) This suggested syno-
There are surprisingly few reports of Gnathichnus nymy is followed here, even though Sedilichnus
of Neogene and Palaeogene age (e.g. Martinell may be the senior synonym.
1982; Barrier & D'Alessandro 1985). In contrast, Oichnus is produced by a wide range of trace-
descriptions of Cretaceous examples are numerous making organisms. As the trace fossils contain
(e.g. Bromley 1970b, 1975,1994; Voigt 1972,1975, few fingerprints to indicate the taxonomic posi-
1996; Breton et al. 1992). Jurassic occurrences tion of the trace-maker, confident identification
have been documented by Palmer (1982) and of the borer is not common. Small (millimetric)
Riegraf (1973), and from the Triassic by Michalik round drill-holes are dominantly produced
(1977, 1980) and Fursich & Wendt (1977). today by predatory gastropods of the Naticidae
and Muricacea, both arising in the Cretaceous
(Taylor et al. 1983; Tracey et al. 1993). The
Bite traces attributed to fish and sharks holes produced by these two groups are con-
sidered to be morphologically distinguishable
There are many reports of bite traces caused by (Carriker 1981), as Oichnus paraboloides and O.
fish predation and grazing in the Recent (e.g. simplex respectively (although Herbert & Dietl
A STRATIGRAPHY OF MARINE BIOEROSION 467

2002 found some Recent muricids produce bor- et al. 1998; Harper & Wharton 2000), and have
ings indistinguishable from those of naticids). been found in parasitized crinoid columns
Thus a good guess at the trace-maker may be (Feldman & Brett 1998).
made for material of Cretaceous or younger Oichnus appears to be unimportant from the
age (e.g. Carriker 1998). Nevertheless, it cannot Carboniferous to Triassic. However, Fiirsich &
be ignored that octopods can make drill-holes Jablonski (1984) found borings resembling O.
closely similar to those of muricaceans, and paraboloides of Triassic age, and Ausich & Gur-
that both ichnospecies occur throughout the rola (1979) found both O. simplex and O. parabo-
Palaeozoic. loides in Carboniferous brachiopods. Baumiller
Recently, the identity of some presumed early (1990), Baumiller et al. (1999) and Hoffmeister
naticid gastropods has been revised, transferring et al. (2003) reported drill-holes in Carboniferous
them to the non-boring family Ampullospiridae crinoids and brachiopods.
(Kase & Ishikawa 2003). These authors claim The Devonian period has provided evidence of
that the Naticidae first appeared in the Late much shell-drilling (Kowalewski et al. 1998).
Cretaceous Campanian, and that the four species Devonian blastoids contain holes attributed to
of gastropod that accompany the supposed platyceratid gastropods (Baumiller 1993, 1996;
earliest naticid borings in the English Middle Baumiller & Macurda 1995), and Devonian bra-
Albian (Mid-Cretaceous) Blackdown Greensand chiopods display drill-holes resembling both O.
are not naticids but ampullospirids. Concerning paraboloides (Smith et al. 1985) and O. simplex
the nature of the trace-maker, the Albian (and (Leighton 2001). Much-parasitized Silurian
other O. paraboloides dated earlier than the crinoids were described by Brett (1978).
Campanian) 'remain mysterious' (Kase & The oldest trace fossils referable to Oichnus
Ishikawa 2003, p. 406). occur in the Cambrian (Matthews & Missarz-
The life-long borings made by ectoparasitic hevsky 1975; Conway Morris & Bengtson 1994)
capulid gastropods usually have diagnostic and Late Precambrian (Bengtson & Zhao
features that allow their attribution (Orr 1962; 1992), the latter in the tube of the earliest
Matsukuma 1978; Bromley 1981). known animal to produce a mineralized skeleton!
In the Palaeozoic, the parasitic gastropods of
the family Platyceratidae have been found in
place over their drill-hole in Mississippian Minute round holes in shells: borings by and
crinoids (Baumiller 1990) and Devonian brachio- in Foraminifera
pods (Baumiller et al. 1999) and blastoids
(Baumiller 1996). The boreholes are close in Increasing interest is being shown in the minute
shape to O. simplex. borings in microfossils, especially foraminifera.
Some of the borings generated by predatory Some of these have been produced by foramini-
octopuses are morphologically characteristic as fera. This group of trace fossils represents a
O. ovalis (Bromley 1993), but not all. Neither size-class of its own (K. S. S. Nielsen 1999; Niel-
do all living species of octopus bore holes. The sen & Nielsen 2001). Several new ichnospecies of
earliest identified octopus boring is Pliocene in Oichnus have been based on these minute trace
age (Robba & Ostinelli 1975), or possibly Cretac- fossils, ranging from about 10 to 60 um in dia-
eous (Harper 2000), whereas rare, octopus-like meter, and some new ichnogenera are being
body fossils date back to the Carboniferous established, e.g. Dipatulichnus Nielsen & Nielsen
(Kluessendorf & Doyle 2000). Boreholes have a 2001 and Stellatichnus Nielsen & Nielsen 2001
much better preservation potential than octopus (see also Nielsen et al. 2003).
bodies (Engeser 1990), and this may indicate that The minute, globular pit Planobola macrogota
the hole-drilling habit is relatively new in the Schmidt 1992 is comparable with Oichnus in
octopods. Foraminifera, being 20-100 um in diameter
Small round holes in echinoids, produced by (e.g. Bundschuh 2000). It occurs in trilobite,
endoparasitic ascothoracican barnacles, are brachiopod and coral skeleton, and has not
treated under crustacean borings. been reported in foraminiferans.
The parasitic foraminiferan Planorbulinopsis
Oichnus Bromley 1981 parasita creates rounded, flat-floored pits about
Small round holes in shells are abundant today 0.5 mm wide in its host, the foraminiferan Alveo-
and since the Late Cretaceous, when the murica- linella quoii (Banner 1971). The pits were not
cean and naticid gastropods began boring (e.g. named, the material being Recent.
Vermeij et al. 1980; Kowalewski et al. 1998; Late Cretaceous orbitoid Foraminifera have
Kase & Ishikawa 2003). They are also common been reported that contain relatively large
in Jurassic bivalves and brachiopods (Harper (nearly 1 mm wide) borings (Baumfalk et al.
468 R. G. BROMLEY

1982). These borings have a looped or horseshoe The small bioclaustration structures, Ordo-
shape to fit the limitations of the substrate, but vician to Devonian, including Helicosalpinx
have side branches and crossovers. They were Oekentorp 1969, Torquaysalpinx Plusquellec
not named, and were interpreted as the work of 1968, Chaetosalpinx Sokolov 1948, Catellocauda
an anomalinid foraminiferan, Talpinella cunicu- Palmer & Wilson 1988, Hicetes Clarke 1908 and
laria, specimens of which were usually present others, are considered commensal with bryozo-
within the borings. ans, tabulate corals and stromatoporoids (Stel
In fact, Foraminifera themselves have been 1976; Tapanila 2002, in press). As yet, little is
shown to produce a variety of borings (Matteucci known about them.
1980; Cedhagen 1994; Freiwald & Schonfeld
1996; Venec-Peyre 1996). Hallock & Talge
(1994) and Hallock et al (1998) described the Discussion
predatory borings of Floresina amphiphaga in
other foraminifera. The present survey of ranges of bioerosional ich-
Several reports of fossil borings attributed to nogenera is more comprehensive than previous
foraminifera include some resetted borings such ones (e.g. Kobluk et al. 1978; Palmer 1982), but
as Globodendrina monile, Late Jurassic (Plewes it does not reveal many surprises. As seen in
et al. 1993; Bertling & Insalaco 1998). The Fig. 1, there are two periods of diversification,
Jurassic foraminiferan Troglotella incrustans both of which have been recognized before.
bores in the juvenile growth stage (Schmid & The 'Mesozoic marine revolution' is well
Leinfelder 1996). Venec-Peyre (1996) identified marked by the abundance and diversity of
20 species known to bore from Jurassic to bioerosional trace fossils (Bertling 1999; Perry
Recent. & Bertling 2000). Vermeij (1977, 1983a) saw a
substantial increase in durophagous and drilling
predation in the middle Mesozoic, and corre-
Fossil borings and bioclaustration of sponding changes in gastropod shell architecture.
uncertain origin Brett (1988) drew attention to the much greater
abundance of deeply boring bivalves and sponges
Grahn (1981) described minute round holes in the Cretaceous than earlier. Palmer (1986) sug-
made in Ordovician chitinozoans, supposedly gested that an increase in predation might have
by bacteria and fungi. triggered this trend. According to Reitner &
Keupp (1991), the sponge genus Aka appeared
Phrixichnus Bromley & Asgaard 1993 in the Triassic and became endolithic in the
This strange boring somewhat resembles Gastro- Jurassic, producing giant borings. By the Cretac-
chaenolites, but has a growth-line ornament and eous, Aka tended to occupy deeper, tranquil
a somewhat quadrate cross-section. P. phrix is waters, whereas the newly appearing endolithic
the only ichnospecies. It occurs today (Bromley clionid sponges generally colonized the shallow,
& Asgaard unpublished observations), yet the energetic waters. It is hoped that new work on
trace-maker remains unknown. It was first pub- ichnogenus Entobia will support these trends.
lished in Pliocene material (Bromley & Asgaard The apparent dearth of Oichnus in the Jurassic
1993), and has been reported from the Miocene reported by Kowalewski et al. (1998) does not
rocky shore environment (Domenech et al. seem to be real (Harper et al. 1998; Harper &
2001). Wharton 2000).
A Mid-Palaeozoic precursor of the Mesozoic
Planovolites Mikulas 1992 revolution was identified by Signor & Brett
This trace fossil comprises depressions about (1984) and Brett (1988) as the diversity of hard-
50 cm wide and 1-1.5 cm deep, and is considered ground communities increased markedly within
to be an algal-grazing area, perhaps by echinoids the Ordovician and Devonian, and this is also
or molluscs. P. homolensis (Early Cretaceous) is clearly represented by the record of bioerosive
the only ichnospecies (Mikulas 1992). trace fossils. Gastrochaenolites and Petroxestes
make their first appearance at this time, as do
Diorygma Biernat 1961 Talpina, acrothoracican barnacles and Caulos-
This is a bioclaustration structure caused by an trepsis (e.g. Wilson & Palmer 1998). There is
endoparasitic or commensal worm in Devonian also a rapid appearance of durophagous preda-
atrypid brachiopods. The worm was considered tors in the Devonian: placoderms, other fish
to be a phoronid by MacKinnon & Biernat and some arthropods (Signor & Brett 1984).
(1970). There is no sign of bioerosion, and this Before the first, mid-Palaeozoic revolution, the
may not be a true trace fossil. bioerosion process was not impressive. There
A STRATIGRAPHY OF MARINE BIOEROSION 469

were boring worms producing Trypanites and I. Glaub is thanked for having provided considerable
Palaeosabella; endolithic sponges were present assistance with the literature on microbioerosion
but uncommon; drilling durophagy existed, during the preparation of this paper.
apparently on a small scale. All these forms
were present in the Early Cambrian. In the Late
Proterozoic we have an isolated but remarkable References
occurrence of Oichnus as well as several records
of Cyanobacteria. The earliest bioerosion ABEL, O. 1935. Vorzeitliche Lebensspuren. Gustav
demonstrated is an endolithic cyanobacterium Fischer, Jena.
from the Early Proterozoic dated at 1700 Ma. Aousxi, J., DOMENECH, R., JULIA, R. & MARTINELL, J.
A further slight increase in bioerosional ichno- 1990. Evolution of the Neogene Basin of Emporda
diversity is suggested for the Neogene. It is not (NE Spain). Paleontologia i Evolutio, Memoria
clear whether this is a genuine mini-revolution, Especial, 2, 251-267.
or merely a function of taphonomy. Generally AKPAN, E. B., FARROW, G. E. & MORRIS, N. 1982.
Limpet grazing on Cretaceous algal-bored ammo-
speaking, these latest sediments have suffered nites. Palaeontology, 25, 361-367.
less diagenesis than the older, which may have ALEXANDER, R. R. 1986. Resistance to and repair of
led to an increase in the preservation potential shell breakage induced by durophages in Late
of trace fossils. Ordovician brachiopods. Journal of Paleontology,
60, 273-285.
ALEXANDER, R. R. 1994. Distribution of pedicle boring
Conclusions traces and the life habit of Late Paleozoic lei-
orhynchid brachiopods from dysoxic habitats.
The approximately 65 ichnogenera catalogued in Lethaia, 27, 227-234.
ALLOUC, J., LE CAMPION-ALSUMARD, T. & LEUNG
this study plus several types of unnamed trace TACK, D. 1996. La bioerosion des substrats
fossil have very variable quality of information magmatiques en milieu littoral: 1'exemple de la
to impart. However, those that appear to have a presqu'ile du Cap Vert (Senegal occidental). Geo-
meaningful stratigraphic representation combine bios, 29, 485-502.
to emphasize the diversity of trends of the ANDRE, M. & LAMY, E. 1933. Crustaces xylophages et
Phanerozoic (Fig. 1). The Cambrian radiation lithophages. Bulletin de I'lnstitut oceanographique
did not express itself among the bioerosional de Monaco, 626, 1-23.
trace fossils. However, the Mid-Palaeozoic and AUSICH, W. I. & GURROLA, R. A. 1979. Two boring
Mesozoic marine revolutions played a strong organisms in a Lower Mississippian community
role. of southern Indiana. Journal of Paleontology, 53,
335-344.
Most of the behavioural-biological groups of BABCOCK, L. E. 1993. Trilobite malformations and the
bioerosion that have been outlined in this paper fossil record of behavioral asymmetry. Journal of
have survived to the present day. The present Paleontology, 67, 217-229.
bioerosion community may therefore be classi- BAIRD, G. C, BRETT, C. E. & TOMLINSON, J. T. 1990.
fied as comprising three chronological categories. Host-specific acrothoracid barnacles on Middle
The Early Palaeozoic forms include Trypanites, Devonian platyceratid gastropods. Historical
possibly Palaeosabella, Oichnus, sponge and Biology, 4, 221-244.
cyanobacterial borings, and damage by preda- BAK, R. P. M. 1994. Sea urchin bioerosion on coral
tory attack. The Late and Mid-Palaeozoic reefs: place in the carbonate budget and relevant
variables. Coral Reefs, 13, 99-103.
forms include the endolithic algae, acrothor- BALOG, S.-J. 1996. Boring thallophytes in some
acican barnacles and mytilid boring bivalves, Permian and Triassic reefs: bathymetry and
ctenostome bryozoans, much durophagy, and bioerosion. In: REITNER, J., NEUWEILER, F. &
Podichnus. The Mesozoic-Cenozoic bioerosion GUNKEL, F. (eds) Global and Regional Controls
community is very diverse, both biologically on Biogenic Sedimentation. I. Reef Evolution.
and ichnologically, and is dominated by endo- Research Reports, Gottinger Arbeiten zur
lithic sponges, a variety of boring and etching Geologic und Palaontologie, Sonderband 2, 305-
bivalves, rasping gastropods and chitons, biting 309.
echinoids and a host of minor groups. BALUK, W. & RADWANSKI, A. 1979a. Additional data
There appears to be yet another increase in bio- on the organic communities and facies develop-
ment of the Korytnica Basin (Middle Miocene;
erosive biodiversity in the Neogene. This effect Holy Cross Mountains, central Poland). Ada Geo-
may have been assisted somewhat by taphonomy: logica Polonica, 29, 225-238.
relative to the older sediments, the younger sedi- BALUK, W. & RADWANSKI, A. 1979b. Boring cteno-
ments have generally been affected only mildly stomate bryozoans from the Korytnica Clays
by diagenetic processes that might decrease the (Middle Miocene; Holy Cross Mountains, central
preservation potential of trace fossils. Poland). Acta Geologica Polonica, 29, 243-252.
470 R. G. BROMLEY

BALUK, W. & RADWANSKI, A. 1996. Stomatopod pre- Boss, K. J. 1965. Note on Lima (Acesta) angolensis.
dation upon gastropods from the Korytnica Nautilus, 79, 54-58.
Basin, and from other classical Miocene localities BOTTJER, D. J. 2000. Radular grazing traces in the
in Europe. Ada Geologica Polonica, 46, 279—304. Lower Cambrian of China: implications for the
BANNER, F. T. 1971. A new genus of the Planorbu- Cambrian substrate revolution. Annual Meeting
linidae an endoparasite of another foraminifer. Geological Society of America, Reno, Nevada.
Revista Espanola de Micropaleontologia, 3, 113— Abstract 50471.
128. BRATTSTROM, H. 1936. Ulophysema oresundense n. gen.
BARRIER, P. & D'ALESSANDRO, A. 1985. Structures bio- et sp., eine neue Art der Ordnung Cirripedia
geniques et physiques dans les Sables de Pavigli- Ascothoracica. Arkiv for Zoologi, 28A (23), 1-10.
ana, Reggio Calabria (Italic). Rivista Italiana di BRATTSTROM, H. 1937. On the genus Ulophysema
Paleontologia e Stratigrqfia, 91, 379^408. Brattstrom with description of a new species
BAUMFALK, Y. A., FORTUIN, A. R. & MOK, R. P. 1982. from East Greenland. Meddelelser om Gronland,
Talpinella cunicularia n. gen., n. sp., a possible for- 118 (7), 1-24.
aminiferal parasite of Late Cretaceous Orbitoides. BRATTSTROM, H. 1947. On the ecology of the ascothor-
Journal of Foraminiferal Research, 12, 185-196. acid Ulophysema oresundense Brattstrom. Studies
BAUMILLER, T. K. 1990. Non-predatory drilling of on Ulophysema oresundense 1. Kungliga Fysiogra-
Mississippian crinoids by platyceratid gastropods. fiska Sdllskapets Handlingar, 58, 1-77.
Palaeontology, 33, 743-748. BRETON, G., NERAUDEAU, D. & CUENCA-BOULAT, C.
BAUMILLER, T. K. 1993. Boreholes in Devonian 1992. Gnathichnus stellarum ichnosp. nov., trace
blastoids and their implications for boring by de broutage d'un echinide du Campanien des
platyceratids. Lethaia, 26, 41^47. Charentes (France). Revue de Paleobiologie, 11,
BAUMILLER, T. K. 1996. Boreholes in the Middle Devo- 219-229.
nian blastoid Heteroschisma and their implications BRETT, C. E. 1978. Host-specific pit-forming epizoans
for gastropod drilling. Palaeogeography, Palaeo- on Silurian crinoids. Lethaia, 11, 217-232.
climatohgy, Palaeoecology, 123, 343-351. BRETT, C. E. 1985. Tremichnus: a new ichnogenus of
BAUMILLER, T. K. & MACURDA, D. B. 1995. Borings in circular-parabolic pits in fossil echinoderms. Jour-
Devonian and Mississippian blastoids (Echino- nal of Paleontology, 59, 625-635.
dermata). Journal of Paleontology, 69, 1084-1089. BRETT, C. E. 1988. Paleoecology and evolution of
BAUMILLER, T. K., LEIGHTON, L. R. & THOMPSON, D. L. marine hard substrate communities: an overview.
1999. Boreholes in Mississippian spiriferide bra- Palaios, 3, 374^378.
chiopods and their implications for Paleozoic BROMLEY, R. G. 1970a. Borings as trace fossils and
gastropod drilling. Palaeogeography, Palaeodima- Entobia cretacea Portlock, as an example. In:
tology, Palaeoecology, 147, 283-289. CRIMES, T. P. & HARPER, J. C. (eds) Trace Fossils.
BENGTSON, S. & ZHAO, Y. 1992. Predatorial borings in Geological Journal Special Issues, 3, 49-90.
Late Precambrian mineralised exoskeletons. BROMLEY, R. G. 1970b. Predation and symbiosis in
Science, 257, 367-369. some Upper Cretaceous clionid sponges. Medde-
BENTON, M. J. 1993. The Fossil Record 2. Chapman & lelser fr a Dansk Geologisk Forening, 19, 398^05.
Hall, London. BROMLEY, R. G. 1972. On some ichnotaxa in hard
BERTLING, M. 1999. Late Jurassic reef bioerosion: the substrates, with a redefinition of Trypanites
dawning of a new era. Bulletin of the Geological Magdefrau. Palaontologische Zeitschrift, 46, 93-
Society of Denmark, 45, 173-176. 98.
BERTLING, M. & INSALACO, E. 1998. Late Jurassic BROMLEY, R. G. 1975. Comparative analysis of fossil
coral/microbial reefs from the northern Paris and Recent echinoid bioerosion. Palaeontology,
Basin: facies, palaeoecology and palaeobiogeogra- 18, 725-739.
phy. Palaeogeography, Palaeoclimatology, BROMLEY, R. G. 1978. Bioerosion of Bermuda reefs.
Palaeoecology, 139, 139-175. Palaeogeography, Palaeoclimatology, Palaeoecol-
BEERNAT, G. 1961. Diorygma atrypophilia n. gen. n. sp.: ogy, 23, 169-197.
a parasitic organism of Atrypa zonata Schnur. BROMLEY, R. G. 1981. Concepts in ichnotaxonomy illu-
Acta Palaeontologica Polonica, 6, 17-28. strated by small round holes in shells. Acta Geolo-
BISHOP, G. A. 1975. Traces of predation. In: FREY, gica Hispdnica, 16, 55-64.
R. W. (ed.) The Study of Trace Fossils. Springer, BROMLEY, R. G. 1993. Predation habits of octopus
New York, 261-281. past and present and a new ichnospecies, Oichnus
BOEKSCHOTEN, G. J. 1967. Palaeoecology of some ovalis. Bulletin of the Geological Society of
Mollusca from the Tielrode Sands (Pleistocene, Denmark, 40, 167-173.
Belgium). Palaeogeography, Palaeoclimatology, BROMLEY, R. G. 1994. The palaeoecology of bioero-
Palaeoecology, 3, 311-362. sion. In: DONOVAN, S. K. (ed.) The Palaeobiology
BOEKSCHOTEN, G. J. 1970. On bryozoan borings from of Trace Fossils. Wiley, Chichester, 134^154.
the Danian at Fakse, Denmark. In: CRIMES, T. P. BROMLEY, R. G. 1999. Anomiid (bivalve) bioerosion on
& HARPER, J. C. (eds) Trace Fossils. Geological Pleistocene pectinid (bivalve) shells, Rhodes,
Journal Special Issues, 3, 43-48. Greece. Geologic en Mijnbouv, 78, 175-177.
BOND, P. N. & SANDERS, W. B. 1989. Sublethal injury BROMLEY, R. G. & ASGAARD, U. 1993. Endolithic com-
and shell repair in Upper Mississippian ammo- munity replacement on a Pliocene rocky coast.
noids. Paleobiology, 15, 414-428. Ichnos, 2, 93-116.
A STRATIGRAPHY OF MARINE BIOEROSION 471

BROMLEY, R. G. & D'ALESSANDRO, A. 1987. Bioerosion CEDHAGEN, T. 1994. Taxonomy and biology ofHyrrok-
of the Plio-Pleistocene transgression of southern kin sarcophaga gen. et sp. n., a parasitic foramini-
Italy. Rivista Italiana di Paleontologia e stratigra- feran (Rosalinidae). Sarsia, 79, 65-82.
fia, 93, 379-442. CLARKE, J. M. 1908. The beginnings of dependent life.
BROMLEY, R. G. & D'ALESSANDRO, A. 1989. Ichno- Bulletin of the New York State Museum, 121, 146-
logical study of shallow marine endolithic sponges 196.
from the Italian coast. Rivista Italiana di Paleonto- CLARKE, J. M. 1921. Organic Dependence and Disease:
logia e Stratigrqfia, 95, 279-314. Their Origin and Significance. Yale University
BROMLEY, R. G. & HANKEN, N.-M. 1981. Shallow Press, New Haven.
marine bioerosion at Vard0, arctic Norway. COLE, A. R. & PALMER, T. J. 1999. Middle Jurassic
Bulletin of the Geological Society of Denmark, 29, worm borings, and a new giant ichnospecies of
103-109. Trypanites from the Bajocian/Dinantian uncon-
BROMLEY, R. G. & MARTINELL, J. 1991. Centrichnus, formity, southern England. Proceedings of the
new ichnogenus for centrically patterned attach- Geologists' Association, 110, 203-209.
ment scars on skeletal substrates. Bulletin of the CONWAY MORRIS, S. & BENGTSON, S. 1994. Cambrian
Geological Society of Denmark, 38, 243-252. predators: possible evidence from boreholes.
BROMLEY, R. G. & SURLYK, F. 1973. Borings produced Journal of Paleontology, 68, 1—23.
by brachiopod pedicles, fossil and Recent. Lethaia, CUFFEY, R. J., FELDMANN, R. M. & POHLABLE, K. E.
6, 349-365. 1981. New Bryozoa from the Fox Hills Sandstone
BRONN, H. G. 1837. Lethaea geognostica oder Abbildun- (Upper Cretaceous, Maestrichtian) of North
gen und Beschreibungen der fur die Gebirgsforma- Dakota. Journal of Paleontology, 55, 401-409.
tionen bezeichnendsten Versteinerungen 1, (Atlas) DE LAUBENFELS, M. W. 1955. Porifera. In: MOORE, R. C.
1-672; 2, (Text) 673-1350. E. Schweizerbart, (ed.) Treatise on Invertebrate Paleontology, E.
Stuttgart. Geological Society of America and University of
BUNDSCHUH, M. 2000. Silurische Mikrobohrspuren - Kansas Press, Lawrence, KS, 21-112.
ihre Beschreibung und Verteilung in verschiedenen DOMENECH, R., GlBERT, J. M. DE & MARTINELL, J.
Faziesrdumen (Schweden, Litauen, Groflbritannien 2001. Ichnological features of a marine transgres-
und USA). Doctoral thesis, Johann Wolfgang sion: Middle Miocene rocky shores of Tarragona,
Goethe University, Frankfurt am Main. Spain. Geobios, 34, 99-107.
BUNDSCHUH, M., GLAUB, I., HOFMANN, K., RADTKE, DONOVAN, S. K. & JAGT, J. W. M. 2002. Oichnus Brom-
G. & VOGEL, K. 1989. Bohrorganismen helfen, fos- ley borings in the irregular echinoid Hemipneustes
sile Meeresbecken zu rekonstruieren. Forschung Agassiz from the Type Maastrichtian (Upper
Frankfurt, 3, 56-63. Cretaceous, The Netherlands and Belgium).
CADEE, G. C. 1968. Molluscan biocoenoses and thana- Ichnos, 9, 67-74.
tocoenoses in the Ria de Arosa, Galicia, Spain. DONOVAN, S. K. & PICKERILL, R. K. 2002. Pattern
Zoologische Verhandelingen, 95, 1—121. versus process or informative versus uninforma-
CALCINAI, B., ARILLO, A., CERRANO, C. & BAVES- tive ichnotaxonomy: reply to Todd and Palmer.
TRELLO, G. 2003. Taxonomy-related differences Ichnos, 9, 85-87.
in the excavating micro-patterns of boring DORNBOS, S. Q. & BOTTJER, D. J. 2000. Radular grazing
sponges. Journal of the Marine Biological Associa- traces from the Lower Cambrian of China: impli-
tion of the United Kingdom, 83, 37-39. cations for the Cambrian substrate revolution.
CALDWELL, R. L. & DINGLE, H. 1975. Ecology and Geological Society of America Annual Meeting,
evolution of agonistic behaviour in stomatopods. Abstracts with Programs, 32, 301.
Naturwissenschaften, 62, 214—222. DORNBOS, S. Q., BOTTJER, D. J. & CHEN, J. Y. in press.
CALDWELL, R. L. & DINGLE, H. 1976. Stomatopods. Evidence for seafloor microbial mats and asso-
Scientific American, 234, 81-89. ciated metazoan lifestyles in Lower Cambrian
CAMERON, B. 1969. New name for Palaeosabella prisca phosphorites of southwest China. Lethaia.
(McCoy), a Devonian worm-boring, and its DORTANGS, R. W. 1998. Sporenfossielen. In: JAGT, J.
preserved probable borer. Journal of Paleontology, W. M., LELOUX, J. & DHONDT, A. V. (eds) Fossie-
43, 189-192. len van de St Pietersberg. Grondboor en Hamer,
CAMPBELL, S. E. 1982. Precambrian endoliths discov- 53, 150-151.
ered. Nature, 299, 429^31. EBBESTAD, J. O. R. & PEEL, J. S. 1997. Attempted
CAMPBELL, S. E., KAZMIERCZAK, J. & GOLUBIC, S. 1979. predation and shell repair in Middle and Upper
Palaeoconchocelis starmachii gen. n., sp. n., an Ordovician gastropods from Sweden. Journal of
endolithic rhodophyte (Bangiaceae) from the Paleontology, 71, 1007-1019.
Silurian of Poland. Ada Palaeontologica Polonica, EKDALE, A. A. & BROMLEY, R. G. 2001. Bioerosional
24, 405-108. innovation for living in carbonate hardgrounds
CARRIKER, M. R. 1981. Shell penetration and feeding in the Early Ordovician of Sweden. Lethaia, 34,
by naticacean and muricacean predatory gastro- 1-12.
pods: a synthesis. Malacologia, 20, 403-422. EKDALE, A. A., BROMLEY, R. G. & PEMBERTON, S. G.
CARRIKER, M. R. 1998. Predatory gastropod traces: a 1984. Ichnology: The Use of Trace Fossils in Sedi-
comparison of verified shallow-water and pre- mentology and Stratigraphy. Society of Economic
sumed deep-sea boreholes. American Malacologi- Paleontologists and Mineralogists, Short Courses,
cal Bulletin, 14, 121-131. Tulsa, Oklahoma, 15, 1-317.
472 R. G. BROMLEY

EKDALE, A. A., BENNER, J. S., BROMLEY, R. G. & nov. from Lee Stocking Island, Bahamas. Nova
GIBERT, J. M. DE. 2002. Bioerosion of Lower Hedwigia, Beiheft, 112, 93-100.
Ordovician hardgrounds in southern Scandinavia GIBERT, J. M. DE, MARTINELL, J. & DOMENECH, R.
and western North America. Acta Geologica 1998. Entobia ichnofacies in fossil rocky shores,
Hispdnica, 37, 9-13. Lower Pliocene, northwestern Mediterranean.
ELLIOT, D. K. & BOUNDS, S. D. 1987. Causes of damage Palaios, 13, 476-487.
to brachiopods from the Middle Pennsylvanian GIESSLER, M. 1991. Echinocorys mit BiBspuren.
Naco Formation, central Arizona. Lethaia, 20, Aufschlufi, 42, 117-120.
327-335. GLAESSNER, M. F. 1976. Early Phanerozoic annelid
ELLIOT, D. K. & BREW, D. C. 1988. Cephalopod preda- worms and their geological and biological signifi-
tion on a Desmoinesian brachiopod from the cance. Journal of the Geological Society, London,
Naco formation, central Arizona. Journal of 132, 259-275.
Paleontology, 62, 145-147. GLAUB, I. 1994. Mikrobohrspuren in ausgewahlten
ENGESER, T. 1990. Phylogeny of the fossil coleoid Ablagerungsraumen des europaischen Jura und
Cephalopoda (Mollusca). Berliner geowissenschaf- der Unterkreide (Klassifikation und Palokologie).
tliche Abhandlungen (A), 124, 123-191. Courier Forschungsinstitut Senckenberg, 174, 1—
EVANS, S. 1999. Wood-boring bivalves and boring lin- 324.
ings. Bulletin of the Geological Society of Denmark, GLAUB, I. & SCHMIDT, H. 1994. Traces of endolithic
45, 130-134. microboring organisms in Triassic and Jurassic
FEIGE, A. & FURSICH, F. T. 1991. Taphonomy of bioherms. Kaupia, Darmstadter Beit rage zur Nat-
the Recent molluscs of Bahia la Choya (Gulf of urgeschichte, 4, 103-112.
California, Sonora, Mexico). Zitteliana, 18, 89- GLAUB, I., VOGEL, K. & GEKTIDIS, M. 2001. The role of
133. modern and fossil cyanobacterial borings in
FELDMAN, H. R. & BRETT, C. E. 1998. Epi- and endo- bioerosion and bathymetry. Ichnos, 8, 185-195.
biontic organisms on Late Jurassic crinoid col- GLAUB, L, GEKTIDIS, M. & VOGEL, K. 2002. Micro-
umns from the Negev Desert, Israel: implications borings from different North Atlantic shelf areas:
for co-evolution . Lethaia, 31, 57-71. variability of the euphotic zone extension and
FISCHER, R. & MEYER, W. 1985. Observations on rock implications for paleodepth reconstructions.
boring by Alpheus saxidomus (Crustacea: Alphei- Courier Forschungsinstitut Senckenberg, 237, 25-
dae). Marine Biology, 89, 213-219. 37.
FONSECA, A. C. & CORTES, J. 1998. Coral borers of the GLAZEK, J., MARCINOWSKI, R. & WIERZBOWSKY, A.
Eastern Pacific: Aspidosiphon (A.) elegans (Sipun- 1971. Lower Cenomanian trace fossils and trans-
cula: Aspidosiphonidae) and Pomatogebia rugosa gressive deposits in the Cracow Upland. Acta
(Crustacea: Upogebiidae). Pacific Science, 52, Geologica Polonica, 21, 433-448.
170-175. GOHAR, H. A. F. & SOLIMAN, G. N. 1963. On the
FREIWALD, A. & SCHONFELD, J. 1996. Substrate pitting biology of three coralliophilids boring in living
and boring pattern of Hyrrokkin sarcophaga corals. Publications of the Marine Biological
Cedberg, 1994 (Foraminifera) in a modern deep- Station, Al-Ghardaqa, Egypt, 12, 99-126.
water coral reef mound. Marine Micropaleontol- GOLUBIC, S. 1990. Shell-boring fungi and algae. In:
ogy, 28, 199-207. BOUCOT, A. J. (ed.) Evolutionary Paleobiology of
FURSICH, F. T. 1979. Genesis, environments, and ecol- Behavior and Coevolution. Elsevier, Amsterdam,
ogy of Jurassic hardgrounds. Neues Jahrbuch fur 347-353.
Geologie und Paldontologie, Abhandlungen, 158, GOLUBIC, S., PERKINS, R. D. & LUKAS, K. J. 1975.
1-63. Boring microorganisms and microborings in
FURSICH, F. T. & JABLONSKI, D. 1984. Late Triassic carbonate substrates. In: FREY, R. W. (ed.) The
naticid drillholes: carnivorous gastropods gain a Study of Trace Fossils. Springer, New York, 229-
major adaptation but fail to radiate. Science, 259.
224, 78-80. GOLUBIC, S., YUN, Z. & CAMPBELL, S. E. 1985. Early
FURSICH, F. T. & PALMER, T. J. 1982. The first true evolution of morphological complexity in
anomiid bivalve? Palaeontology, 25, 897-903. prokaryotes (Cyanophyta, Cyanobacteria). In:
FURSICH, F. T. & WENDT, J. 1977. Biostratigraphy and MLIKOVSKY, J. & NOVAK, V. J. A. (eds) Evolution
palaeoecology of the Cassian Formation (Triassic) and Morphogenesis. Academia, Praha, 355-368.
of the southern Alps. Palaeogeography, Palaeo- GOLUBIC, S., LE CAMPION-ALSUMARD, T. & CAMPBELL,
climatology, Palaeoecology, 22, 257—323. S. E. 1999. Diversity of marine Cyanobacteria. In:
FURSICH, F. T., PALMER, T. J. & GOODYEAR, K. L. 1994. CHARPY, L. & LARKUM, A. W. D. (eds) Marine
Growth and disintegration of bivalve-dominated Cyanobacteria. Bulletin de 1'Institut Oceanogra-
patch reefs in the Upper Jurassic of southern phique, Monaco, Special Issues, 19, 53-76.
England. Palaeontology, 37, 131-171. GRAHN, Y. 1981. Parasitism on Ordovician Chitinozoa.
GEARY, D. H., ALLMON, W. D. & REAKA-KUDLA, M. L. Lethaia, 14, 135-142.
1991. Stomatopod predation on fossil gastropods HANTZSCHEL, W. 1975. Trace Fossils and Problema-
from the Plio-Pleistocene of Florida. Journal of tica. In: TEICHERT, C. (ed.) Treatise on Invertebrate
Paleontology, 65, 355-360. Paleontology W (Supplement 1, 2nd edn). Geo-
GEKTIDIS, M. & GOLUBIC, S. 1996. A new endolithic logical Society of America and University of
cyanophyte/cyanobacterium: Hyella vacans sp. Kansas Press, Lawrence, KS, 1-269.
A STRATIGRAPHY OF MARINE BIOEROSION 473

HAGENOV, F. VON 1840. Monographic der Riigenschen HUGGETT, J. M., GALE, A. S. & EVANS, S. 2000. Carbo-
Kreideversteinerungen, 2. Abth. Radiarien und nate concretions from the London Clay (Ypresian,
Annulaten, nebst Nachtragen zur ersten Abthei- Eocene) of southern England and the exceptional
lung. Neues Jahrbuch fur Mineralogie, Geognosie, preservation of wood-boring communities.
Geologic und Petrefaktenkunde, 1840, 631-671. Journal of the Geological Society, London, 157,
HALLOCK, P. & TALGE, H. K. 1994. A predatory fora- 187-200.
minifer, Floresina amphiphaga, n. sp., from the JAMES, N. P., KOBLUK, D. R. & PEMBERTON, S. G. 1977.
Florida Keys. Journal of Foraminiferal Research, The oldest macroborers: Lower Cambrian of
24,210-213. Labrador. Science, 197, 980-983.
HALLOCK, P., TALGE, H. K., WILLIAMS, D. E. & JiicH, P. J. W. & BOEKSCHOTEN, G. J. 1980. Trace
HARNEY, J. N. 1998. Borings in Amphistegina fossils and grazing traces produced by Littorina
(Foraminiferida): evidence of predation by Flore- and Lepidochitona, Dutch Wadden Sea. Geologie
sina amphiphaga (Foraminiferida). Historical en Mijnbouw, 59, 33-42.
Biology, 13, 73-76. KASE, T. & ISHIKAWA, M. 2003. Mystery of naticid
HANSEN, M. C. & MAPES, R. H. 1990. A predator-prey predation history solved: evidence from a 'living
relationship between sharks and cephalopods in fossil' species. Geology, 31, 403-406.
the late Paleozoic. In: BOUCOT, A. J. (ed.) Evolu- KASE, T., SHIGETA, Y. & FUTAKAMI, M. 1994. Limpet
tionary Paleobiology of Behavior and Coevolution. home depressions in Cretaceous ammonites.
Elsevier, Amsterdam, 189-192. Lethaia, 27, 49-58.
HARPER, E. M. 2000. Hunting the killer octopus. KASE, T., JOHNSTON, P. A., SEILACHER, A. & BOYCE, J.
Palaeontology Newsletters, 45, Abstracts 2000, 15. 1998. Alleged mosasaur bite marks on Late
HARPER, E. M. & WHARTON, D. S. 2000. Boring preda- Cretaceous ammonites are limpet (patellogastro-
tion and Mesozoic articulate brachiopods. Palaeo- pod) home scars. Geology, 26, 947-950.
geography, Palaeoclimatology, Palaeoecology, 158, KAUFFMAN, E. G. 1972. Ptychodus predation upon a
15-24. Cretaceous Inoceramus. Palaeontology, 15, 439-
HARPER, E. M., FORSYTHE, G. T. W. & PALMER, T. 444.
1998. Taphonomy and the Mesozoic marine revo- KAUFFMAN, E. G. 1990. Mosasaur predation on ammo-
lution: preservation state masks the importance of nites during the Cretaceous: an evolutionary
boring predators. Palaios, 13, 352-360. history. In: BOUCOT, A. J. (ed.) Evolutionary Paleo-
HERBERT, G. S. & DIETL, G. P. 2002. Tests of the biology of Behavior and Coevolution. Elsevier,
escalation hypothesis: the role of multiple preda- Amsterdam, 184-189.
tors. Geological Society of America Abstracts KAUFFMAN, E. G. & KESLING, R. V. 1960. An Upper
with Programs, 34, 538-539. Cretaceous ammonite bitten by a mosasaur.
HESSLAND, I. 1949. Investigations of the Lower Ordo- University of Michegan Museum, Paleontological
vician of the Siljan District, Sweden. 2, Lower Contributions, 15, 193-248.
Ordovician penetrative and enveloping algae KELLY, S. R. A. 1988. Turnus? davidsoni (de Loriol), the
from the Siljan District. Bulletin of the Geological earliest British pholadid wood-boring bivalve,
Institute, Uppsala, 33, 409^28. from the Late Jurassic of Oxfordshire. Proceedings
HOLDER, H. 1973. Miscellanea cephalopodica. Geologie of the Geologists' Association, 99, 43-47.
und Palaontologie, Munster, 29, 39-76. KELLY, S. R. A & BROMLEY, R. G. 1984. Ichnological
HOFFMANN, M. & KROBICKI, M. 1989. Oyster buildup nomenclature of clavate borings. Palaeontology,
within the disaerobic-facies mudstones (Middle 27, 793-807.
Jurassic, central Poland): example of benthic KERN, J. P. 1979. The ichnofossil Helicotaphrichnus
island colonization. Annales Societatis Geolo- commensalis in the Korytnica basin (Middle Mio-
gorum Poloniae, 59, 299-330. cene; Holy Cross Mountains, Central Poland).
HOFFMEISTER, A. P., KOWALEWSKI, M., BAMBACH, R. K. Acta Geologica Polonica, 29, 239-242.
& BAUMILLER, T. K. 2003. Intense drilling in the KERN, J. P., GRIMMER, J. C. & LISTER, K. H. 1974. A
Carboniferous brachiopod Cardiarina cordata new fossil spionid tube, Pliocene and Pleistocene
Cooper. Lethaia, 36, 107-118. of California and Baja California. Journal of
HOFMANN, K. 1996. Die mikro-endolithischen Spuren- Paleontology, 48, 978-982.
fossilien der borealen Oberkreide Nordwest- KEUPP, H. 1991. BiBmarken oder postmortale Implo-
Europas und ihre Faziesbeziehungen. Geologisches sionsstrukturen? Fossilien, 3/1991, 275-280.
Jahrbuch, A, 136, 1-151. KIER, P. M. 1981. A bored Cretaceous echinoid.
HOLTHUIS, L. B. 1980. Alpheus saxidomus new species, Journal of Paleontology, 55, 656-659.
a rock boring snapping shrimp from the Pacific KLEEMANN, K. 1984. Lebensspuren von Upogebia
coast of Costa Rica, with notes on Alpheus simus operculata (Crustacea, Decapoda) in karibischen
Guerin-Meneville, 1856. Zoologische Mededelin- Steinkorallen (Mareporaria, Anthozoa). Beitrage
gen. Leiden, 55 (4), 47-58. zur Palaontologie von Osterreich, 11, 35-57.
HUBBARD, D. K., MILLER, A. I. & SCATURO, D. 1990. KLEEMANN, K. 1994. Associations of corals and boring
Production and cycling of calcium carbonate in a bivalves since the Late Cretaceous. Fades, 31,
shelf-edge reef system (St Croix, US Virgin 131-140.
Islands): applications to the nature of reef systems KLUESSENDORF, J. & DOYLE, P. 2000. Pohlsepia mazo-
in the fossil record. Journal of Sedimentary Petrol- nensis, an early 'octopus' from the Carboniferous
ogy, 60, 335-360. of Illinois, USA. Palaeontology, 43, 919-926.
474 R. G. BROMLEY

KNOLL, A. H., GOLUBIC, S., GREEN, J. & SWETT, K. LOWENSTAM, H. A. 1962. Magnetite in denticle capping
1986. Organically preserved microbial endoliths in Recent chitons (Polyplacophora). Bulletin of the
from the late Proterozoic of East Greenland. Geological Society of America, 73, 435^438.
Nature, 321, 856-857. LOWENSTAM, H. A. 1971. Opal precipitation by marine
KNOLL, A. H., SWETT, K. & BURKHARDT, E. 1989. gastropods (Mollusca). Science, 111, 487-490.
Paleoenvironmental distribution of microfossils MACKINNON, D. I. & BIERNAT, G. 1970. The probable
and stromatolites in the Upper Proterozoic Back- affinities of the trace fossil Diorygma atrypophilia.
lundtoppen Formation, Spitsbergen. Journal of Lethaia, 3, 163-172.
Paleontology, 63, 129-145. MADSEN, F. J. & WOLFF, T. 1965. Evidence of the
KOBLUK, D. R. 198la. Lower Cambrian cavity-dwell- occurrence of Ascothoracica (parasitic cirripeds)
ing endolithic (boring) sponges. Canadian Journal in Upper Cretaceous. Meddelelser fra Dansk
of Earth Sciences, 18, 972-980. Geologisk Forening, 15, 556-558.
KOBLUK, D. R. 1981b. Middle Ordovician (Chazy MAGDEFRAU, K. 1932. iiber einige Bohrgange aus dem
Group) cavity-dwelling boring sponges. Canadian Unteren Muschelkalk von Jena. Palaontologische
Journal of Earth Sciences, 18, 1101-1108. Zeitschrift, 14, 150-160.
KOBLUK, D. R. & NEMCSOK, S. 1982. The macroboring MAGDEFRAU, K. 1937. Lebens Spuren fossiler 'Bohr'-
ichnofossil Trypanites in the Middle Ordovician Organismen. Beitrdge zur naturkundlichen For-
bryozoan Prasopora: population behaviour and schung in Sudwestdeutschland, 2, 54—67.
reaction to environmental influences. Canadian MARKEL, K. & MAIER, R. 1966. Uber die Beweglichkeit
Journal of Earth Sciences, 19, 679-688. von Seeigeln. Naturwissenschaft, 53 (20), 535.
KOBLUK, D. R., JAMES, N. P. & PEMBERTON, S. G. 1978. MARKEL, K. & MAIER, R. 1967. Beobachtungen an
Initial diversification of macroboring ichnofossils lochbewohnenden Seeigeln. Natur und Museum,
and exploitation of the macroboring niche in the 97, 233-243.
lower Paleozoic. Paleobiology, 4, 163-170. MARTINELL, J. 1982. Echinoid bioerosion from the
KOWALEWSKI, M., DULAI, A. & FuRSiCH, F. T. 1998. A Pliocene of NE Spain. Geobios, 15, 249-253.
fossil record full of holes: the Phanerozoic history MARTINELL, J. 1989. Interaction organismos/sustrato
of drilling predation. Geology, 26, 1091-1094. duro: la bioerosion y sus implicaciones. In:
KRUMBACH, T. 1914. Mitteilungen liber die Nahrung AGUIRRE, E. (ed.) Paleontologia. Coleccion
felsbewohnender Seeigel der nordlichen Adria. 'Nuevas Tendencias', Consejo Superior de Investi-
Notizen iiber die Fauna der Adria bei Rovigno. gaciones Cientificas, Madrid, 205-222.
Zoologischer Anzeiger, 44, 440^51. MARTINELL, J. & DOMENECH, R. 1986. Actividad bio-
KRUMBEIN, W. E. & VAN DER PERS, J. N. C. 1974. erosiva en el Plioceno marino del Emporda (Cata-
Diving investigations on biodeterioration by sea- lunya). Paleontologia i Evolucio, 20, 247-251.
urchins in the rocky sublittoral of Helgoland. Hel- MASSIN, C. 1982. Contribution to the knowledge of two
golander wissenschaftiche Meeresuntersuchungen, boring gastropods with an annotated list of the
26, 1-17. genera Magilus Montfort, 1810 and Leptoconchus
LAMY, E. 1930.Quelques mots sur la lithophagie chez les Riippell, 1835. Bulletin de I'lnstitut Royal des Sci-
gasteropodes. Journal de Conchyliogie, 74, 1-34. ences Naturelles de Belgique, Biologie, 53 (17), 1-28.
LEIGHTON, L. R. 2001. New example of Devonian pre- MASSIN, C. 1983 Note on the genus Leptoconchus Riip-
datory boreholes and the influence of brachiopod pell, 1835 (Mollusca, Gastropoda, Coralliophili-
spines on predator success. Palaeogeography, dae) with the description of two new species,
Palaeoclimatology, Palaeoecology, 165, 53-69. Leptoconchus vangoethemi sp.n. and Leptoconchus
LESCINSKY, H. L. & BENNINGER, L. 1994. Pseudo- cyphastreae sp.n., from Papua New Guinea. Bulle-
borings and predator traces: artefacts of pressure- tin de I'lnstitut Royal des Sciences Naturelles de
dissolution in fossiliferous shales. Palaios, 9, 599- Belgique, Biologie, 55, 1-16.
604. MASSIN, C. 1987. Reliquiaecava, a new genus of Coral-
LEWY, Z. & AVNI, Y. 1988. Omission surfaces in the liophilidae (Mollusca, Gastropoda). Bulletin de
Judea Group, Makhtesh Ramon Region, southern I'lnstitut Royal des Sciences Naturelles de Belgique,
Israel, and their paleogeographic significance. Biologie, 57, 79-90.
Israel Journal of Earth Science, 37, 105-113. MASSIN, C. 1988. Boring Coralliophilidae (Mollusca,
LEYMERIE, A. 1842. Suite du memoire sur le terrain Gastropoda): coral host relationship. Proceedings
cretace du Departement de 1'Aube. Memoires de of the 6th International Coral Reef Symposium,
la Societe Geologique de France, 5, 1-34. Townsville, Australia, 3, 177-184.
LINDBERG, D. R. & DWYER, K. R. 1982. The topogra- MASSIN, C. 1990. Biologie et ecologie de Leptoconchus
phy, formation and role of the home depression of peronii (Lamarck, 1818) (Gastropoda, Corallio-
Collisella scabra (Gould) (Gastropoda: Acmaei- philidae) recolte en Papouasie Nouvelle-Guinee,
dae). The Veliger, 25, 229-234. avec une redescription de 1'espece. Bulletin de
LINDSTROM, A. & PEEL, J. 2003. Shell repair and mode I'lnstitut Royal des Sciences Naturelles de Belgique,
of life of Praenatica gregaria (Gastropoda) from Biologie, 60, 23-33.
the Devonian of Bohemia (Czech Republic). MASSIN, C. 1992. Ecology of some Leptoconchus spp.
Palaeontology, 46, 623-633. (Gastropoda, Coralliophilidae) infesting Fungii-
LINDSTROM, M. 1979. Probable sponge borings in dae (Anthozoa, Madreporaria). In: GIUSTI, F. &
Lower Ordovician limestone of Sweden. Geology, MANGANELLI, G. (eds) Abstracts llth International
7, 152-155. Malacological Congress, Siena, 455.
A STRATIGRAPHY OF MARINE BIOEROSION 475

MATSUKUMA, A. 1978. Fossil boreholes made by shell- MIKULAS, R., PEK, I. & ZIMAK, J. 1995. Teredolites
boring predators or commensals. I. Boreholes of clavatus from the Cenomanian near Maletin
capulid gastropods. Venus, Japanese Journal of (Bohemian Cretaceous Basin), Moravia, Czech
Malacology, 37, 29^5. Republic. Vestnik Ceskeho Geologickeho Ustavu,
MATTEUCCI, R. 1980. Osservazioni sul foraminifero 70(2), 51-58.
endolitico Cymbaloporella tabellaeformis (Brady) MILLER, W. & BROWN, N. A. 1979. The attachment
nell'atollo di Male (north Male), Isole Maldive. scars of fossil balanids. Journal of Paleontology,
Geologica Romana, 19, 267-274. 53, 208-210.
MATTHEWS, S. C. & MISSARZHEVSKY, V. V. 1975. Small MONTEILLET, J., MARCHAND, B. & PLAZIAT, J.-C. 1982.
shelly fossils of late Precambrian and early Cam- Les 'cannelures horizontales', traces de broutage
brian age: a review of recent work. Journal of the fossilisables realisees en elevage sur divers sub-
Geological Society of London, 131, 289-304. strats par le poisson ouest-africain Tilapia heude-
MAUZEY, K. P., BIRKELAND, C. K. & DAYTON, P. K. lotti. Bulletin de la Societe Geologiques de France,
1968. Feeding behavior of asteroids and escape 24, 149-152.
responses of their prey in the Puget Sound MULLER, A. H. 1977. Zur Ichnologie der subherzynen
region. Ecology, 49, 603-619. Oberkreide (Campan). Zeitschrift fur geologische
MAYORAL, E. 1987a. Stellichnusnov. ichnogen., huellas Wissenschaften, Berlin, 5, 881-897.
de incrustacion atribuidas a Paravinella nov. gen. NEBELSICK, J. 1999. Taphonomic comparison between
(Bryozoa, Ctenostomata) de la formation Arenas Recent and fossil sand dollars. Palaeogeography,
de Huelva (Plioceno inferior) en la Cuenca del Palaeoclimatology, Palaeoecology, 149, 349-358.
Bajo Guadalquivir (Espana). Revista Espanola de NEUMANN, C. 2000. Evidence of predation on Cretac-
Paleontologia, 2, 33-40. eous sea stars from north-west Germany. Lethaia,
MAYORAL, E. 1987b. Action bioerosiva de Mollusca 33, 65-70.
(Gastropoda, Bivalvia) en el Plioceno inferior de NIELSEN, J. K. 1999. Commensal association ofCorbula
la Cuenca del Bajo Guadalquivir. Revista Espa- gibba (Bivalvia) and a subconical boring. Bulletin
nola de Paleontologia, 2, 49-58. of the Geological Society of Denmark, 45, 135-138.
MAYORAL, E. 1988. Pennatichnus nov. ichnogen.; Pina- NIELSEN, J. K. & NIELSEN, K. S. S. 2002. Pattern versus
ceocladichnus nov. ichnogen. e Iramena. Huellas de process or informative versus uninformative ich-
bioerosion debidas a bryozoa perforantes (Cteno- notaxonomy: reply to Todd and Palmer. Ichnos,
stomata, Plioceno inferior) en la cuenca del Bajo 9, 83-84.
Guadalquivir. Revista Espanola de Paleontologia, NIELSEN, K. S. S. 1999. Forminiferivory revisited: a pre-
3, 13-22. liminary investigation of holes in foraminifera.
MAYORAL, E. & REGUANT, S. 1995. Palaeoecology and Bulletin of the Geological Society of Denmark, 45,
taphonomy of bivalves, mainly Glycymeris insu- 139-142.
brica (Brocchi), and bryozoans from the Huelva NIELSEN, K. S. S. & NIELSEN, J. K. 2001. Bioerosion in
Sands Fm. (Lower Pliocene, SW Spain). Revista Pliocene to Late Holocene tests of benthic and
Espanola de Paleontologia, no. Homenaje al Dr. planktonic foraminiferans, with a revision of the
Guillermo Colom, 31-47. ichnogenera Oichnus and Tremichnus. Ichnos, 8,
MAYORAL, E., GUTIERREZ MARCO, J. C. & MARTINELL, 99-116.
J. 1994. Primas evidencias de briozoos perforantes NIELSEN, K. S. S., NIELSEN, J. K. & BROMLEY, R. G.
(Ctenostomata) en braquiopodos ordovicios de los 2003. Palaeoecological and ichnological signifi-
Montes de Toledo (Zona Centroiberica Meridio- cance of microborings in Quaternary foraminifera.
nal, Espana). Revista Espanola de Paleontologia, Palaeontologia Electronica, 6(2), 13.
9, 185-194. NODA, H. 1991. Fossil homing scar of gastropod
MICHAL!K, J. 1977. Systematics and ecology ofZeilleria Hipponix (Malluvium) lissus from the Pliocene
Bayle and other brachiopods in the uppermost Shinzato Formation in Okinawa Prefecture,
Triassic of the west Carpathians. Geologick southwestern Japan. Annual Report, Institute of
zbornik, Geologica Carpathica, 28, 323-346. Geoscience, University of Tsukuba, 17, 43^47.
MICHALIK, J. 1980. A palaeoenvironmental and OEKENTORP, K. 1969. Kommensalismus bei Favositi-
palaeoecological analysis of the West Carpathian den. Munstersche Forschungen zur Geologic und
part of the northernTethyan nearshore region in Palaontologie, 12, 165-217.
the latest Triassic times. Rivista Italiana di Paleon- OLEMPSKA, E. 1986. Endolithic microorganisms in
tologia e Stratigrafia, 85, 1047-1064. Ordovician ostracod valves. Ada Palaeontologica
MIKULAS, R. 1992. Early Cretaceous borings from Polonica, 31, 229-236.
Stramberk (Czechoslovakia). Casopispro Minera- ORR, V. 1962. The drilling habit of Capulus danieli
logii a Geologii, Roc, 37, 297-312. (Crosse) (Mollusca: Gastropoda). Veliger, 5, 63-67.
MIKULAS, R. 1994a. Ichnofosilie vznikle v pevnych sub- ORTON, J. H. 1914. On the breeding habits of Echinus
stratech (vrtby) ze spodniho Devonu Barrandienu. miliaris, with a note on the feeding habits of
Zprdvy o Geologickych Vyzkumech v Roce, 1993, Patella vulgata. Journal of the Marine Biological
65-66. Association, United Kingdom, 10, 254-257.
MIKULAS, R. 1994b. Sponge borings in stromatopor- OSCHMANN, W. 1989. Growth and environmental
oids and tabulate corals from the Devonian of hazards of the Upper Jurassic colonial coral Acti-
Moravia (Czech Republic). Vestnik Ceskeho Geo- nastrea matheyi (Koby) from Portugal. Palaonto-
logickeho Ustavu, 69 (1), 69-74. logische Zeitschrift, 63, 193-205.
476 R. G. BROMLEY

PALMER, T. J. 1982. Cambrian to Cretaceous changes in PLEWES, C. R. 1994. Jurassic boring phoronids: non-
hardground communities. Lethaia, 15, 309-323. boring insights into fossil record of some soft
PALMER, T. J. 1986. Adaptive radiations in Mesozoic bodied worms. Palaeontology Newsletter, 24, 24.
groups that inhabit hard substrates. Proceedings PLEWES, C. R. 1996. Ichnotaxonomic studies of Jurassic
Fourth North American Paleontological Conven- endoliths. PhD thesis, Institute of Earth Studies,
tion, Boulder, Colorado, p. A34. University of Wales, Aberystwyth.
PALMER, T. J. & PALMER, C. D. 1977. Faunal distri- PLEWES, C. R., PALMER, T. J. & HAYNES, J. R. 1993. A
bution and colonization strategy in a Middle boring foraminiferan from the Upper Jurassic of
Ordovician hardground community. Lethaia, 10, England and northern France. Journal of Micro-
179-199. palaeontology, 12, 83-89.
PALMER, T. & PLEWES, C. 1993. Borings and bioerosion PLEYDELL, S. M. & JONES, B. 1988. Borings of various
in fossils. Geology Today, 1993, 138. fauna elements in the Oligocene-Miocene Bluff
PALMER, T. J. & WILSON, M. A. 1988. Parasitism of Formation of Grand Cayman, British West
Ordovician bryozoans and the origin of pseudo- Indies. Journal of Paleontology, 62, 348-367.
borings. Palaeontology, 31, 939-949. PLUSQUELLEC, Y. 1968. Commensaux des Tabules et
PALMER, T. J., PLEWES, C. R. & COLE, A. 1997. The Stromatoporoides du Devonien Armoricain.
simple and long-ranging worm-boring Trypanites: Annales de la Societe Geologique du Nord, 88,47-56.
not so simple and long-ranging after all. Geological POHOWSKY, R. A. 1974. Notes on the study and nomen-
Society of America, Abstracts with Programs, 29, clature of boring Bryozoa. Journal of Paleontol-
A.107. ogy, 48, 556-564.
PEEL, J. S. 1984. Attempted predation and shell repair POHOWSKY, R. A. 1978. The boring ctenostomate
on Euomphalopterus (Gastropoda) from the Silur- Bryozoa: taxonomy and paleobiology based on
ian of Gotland. Bulletin of the Geological Society cavities in calcareous substrata. Bulletins of
of Denmark, 32, 163-168. American Paleontology, 73, 1-192.
PEEL, J. S., EBBESTAD, J. O. R. & LINDSTROM, A. 1996. POJETA, J. & PALMER, T. J. 1976. The origin of rock
Shell repair and failed predation in Lower boring in mytilacean pelecypods. Alcheringa, 1,
Palaeozoic gastropods from Sweden. 6th North 167-179.
American Paleontological Convention, Washington, PRATT, B. R. 1998. Probable predation on Upper Cam-
DC. Abstracts for papers. Paleontological Society brian trilobites and its relevance for the extinction
Special Publications, Knoxville, Tennessee, 8, of soft-bodied Burgess Shale-type animals.
305. Lethaia, 31, 73-88.
PEMBERTON, S. G., KOBLUK, D. R., YEO, R. K. & RISK, QUENSTEDT, F. A. 1848. Petrefaktenkunde Deutsch-
M. J. 1980. The boring Trypanites at the Silurian- lands. 1. Abtheilung, Band 1: Cephalopoden. 580
Devonian disconformity in southern Ontario. pp. L. F. Fues, Tubingen.
Journal of Paleontology, 54, 1258-1266. RADTKE, G. 1991. Die mikroendolithischen Spurenfos-
PERRY, C. T. & BERTLING, M. 2000. Spatial and tem- silien im Alt-Tertiar West-Europas und ihre palo-
poral patterns of macroboring within Mesozoic kologische Bedeutung. Courier Forschungsinstitut
and Cenozoic coral reef systems. In: INSALACO, Senckenberg, 138, 1-185.
E., SKELTON, P. W. & PALMER, T. J. (eds) RADTKE, G., GEKTIDIS, M., GOLUBIC, S., HOFMANN, K.,
Carbonate Platform Systems; Components and KIENE, W. E. & LE CAMPION-ALSUMARD, T. 1997.
Interactions. Geological Society, London, Special The identity of an endolithic alga: Ostreobium
Publications, 178, 33-50. brabantium Weber-van Bosse is recognized as
PETHER, J. 1995. Belichnus new ichnogenus, a ballistic carbonate-penetrating rhyzoids of Acetabularia
trace on mollusc shells from the Holocene of the (Chlorophyta, Dasycladales). Courier Forschungs-
Benguela region, South Africa. Journal of Paleon- institut Senckenberg, 201, 341-347.
tology, 69, 171-181. RADWANSKI, A. 1965. Additional notes on Miocene
PICKERILL, R. K. 1976. Vermiforichnus borings from littoral structures of southern Poland. Bulletin de
the Ordovician of coastal Wales. Geological I'Academic Polonese des Sciences Serie des Sciences
Magazine, 113, 159-164. Geologiques et Geographique, 13, 167-173.
PICKERILL, R. K. & DONOVAN, S. K. 1998. Ichnology RADWANSKI, A. 1969. Transgresja dolnego tortonu na
of the Pliocene Bowden shell bed, southeast poludniowych stokach Gor Swi^tokrzyskich
Jamaica. In: DONOVAN, S. K. (ed.) The Pliocene (strefa zatok i ich przedpola). Acta Geologica Polo-
Bowden Shell Bed, Southeast Jamaica. Contribu- nica, 19, 1-143.
tions to Tertiary and Quaternary Geology, 35, RADWANSKI, A. 1970. Dependence of rock-borers and
161-175. burrowers on the environmental conditions
PICKERILL, R. K. & HARLAND, T. L. 1984. Middle within the Tortonian littoral zone of southern
Ordovician microborings of probable sponge Poland. In: CRIMES, T. P. & HARPER, J. C. (eds)
origin from eastern Canada and southern Trace Fossils. Geological Journal Special Issues,
Norway. Journal of Paleontology, 58, 885-891. 3, 371-390.
PICKERILL, R. K., DONOVAN, S. K. & PORTELL, R. W. RADWANSKI, A. 1977. Present-day types of trace in the
2001. The bioerosional ichnofossil Petroxestes Neogene sequence: their problems of nomencla-
pera Wilson and Palmer from the Middle Miocene ture and preservation. In: CRIMES, T. P. &
of Carriacou, Lesser Antilles. Caribbean Journal of HARPER, J. C. (eds) Trace Fossils 2. Geological
Science, 37, 130-131. Journal Special Issues, 9, 227-264.
A STRATIGRAPHY OF MARINE BIOEROSION 477

REITNER, J. & KEUPP, H. 1991. The fossil record of the SEILACHER, A. 1995. Fossile Kunst: Albumblatter der
haplosclerid excavating sponge Aka de Lauben- Erdgeschichte (1st edn). Goldschneckverlag,
fels. In: REITNER, J. & KEUPP, H. (eds) Fossil and Tubingen.
Recent Sponges. Springer, Berlin, 102-120. SEILACHER, A. 1998. Mosasaurs, limpets or diagenesis:
RICE, M. E. 1969. Possible boring structures of sipun- how Placenticeras shells got punctured. Mitteilun-
culids. American Zoologist, 9, 803-812. gen des Museums fur Naturkunde Berlin, geowis-
RICHARDS, R. P. 1974. A Devonian Immergentia senschaftliche Reihe, 1, 93-102.
(Ectoprocta Ctenostomata) from Ohio. Journal SHARMAN, M. 1956. Note on Capulus ungaricus (L.).
of Paleontology, 48, 940-946. Journal of the Marine Biological Association,
RIEGRAF, W. 1973. BiBspuren auf jurassischen Belemni- United Kingdom, 35, 445^50.
tenrostren. Neues Jahrbuch fur Geologic und SHIGEMIYA, Y. 2003. Does the handedness of the pebble
Palaontologie, Monatshefte, 1973, 494-500. crab Eriphia smithii influence its attack success on
RIGBY, J. K., BUDD, G. E., WOOD, R. A. & DEBRENNE, two dextral snail species? Journal of Zoology,
F. 1993. Porifera. In: BENTON, M. J. (ed.) The London, 260, 259-265.
Fossil Record 2. Chapman & Hall, London, 71-99. SHOUP, J. B. 1968. Shell opening by crabs of the genus
ROBBA, E. & OSTINELLI, F. 1975. Testimonianze di Calappa. Science, 160, 887-888.
predazione sui molluschi pliocenici di Albenga. SIGNOR, P. W. & BRETT, C. E. 1984. The mid-Paleozoic
Rivista Italiana di Paleontologia e Stratigrqfia, precursor to the Mesozoic marine revolution.
81, 309-372. Paleobiology, 10, 229-245.
RODRIGUEZ, J. & GUTSCHICK, R. C. 1970. Late SMITH, S. A., THAYER, C. W. & BRETT, C. E. 1985. Pre-
Devonian-Early Mississippian ichnofossils from dation in the Paleozoic: gastropod-like drillholes in
western Montana and northern Utah. In: Devonian brachiopods. Science, 230, 1033-1035.
CRIMES, T. P. & HARPER, J. C. (eds) Trace Fossils. SOKOLOV, B. S. 1948. Kommensalism u Favositid.
Geological Journal Special Issues, 3, 407^438. hvestija Akademyi Nauk Sovietzkovo Soyza Soye-
RUTZLER, K. 1971. Bredin-Archbold-Smithsonian dynonnih Republik, ser. Biol, 1, 101-110.
biological survey of Dominica: burrowing SOLIMAN, G. N. 1969. Ecological aspects of some coral-
sponges, genus Siphonodictyon Bergquist, from boring gastropods and bivalves of the northwes-
the Caribbean. Smithsonian Contributions to tern Red Sea. American Zoologist, 9, 887-894.
Zoology, 77, 1-37. SOLLE, G. 1938. Die ersten Bohr-Spongien im euro-
RUNHAM, N. W. 1961. The histochemistry of the radula paischen Devon und einige andere Spuren. Senck-
of Patella vulgata. Quarterly Journal of the Micro- enbergiana, 20, 154-178.
scopical Society, 102, 371-380. STEL, J. H. 1976. The Paleozoic hard substrate trace
RUNNEGAR, B., POJETA, J., TAYLOR, M. E. & COLLINS, fossils Helicosalpinx, Chaetosalpinx and Torquay-
D. 1979. New species of the Cambrian and Ordo- salpinx. Neues Jahrbuch fur Geologie und Palaonto-
vician chitons Matthevia and Chelodes from Wis- logie, Monatshefte, 1976, 726-744.
consin and Queensland: evidence for the early STEPHENSON, L. W. 1952. Larger invertebrate fossils of
history of polyplacophoran molluscs. Journal of the Woodbine Formation (Cenomanian) of Texas.
Paleontology, 53, 1374-1394. United States Geological Survey, Professional
SAINT-SEINE, R. DE 1951. Un cirripede acrothoracique Papers, 242, 1-226.
du Cretace: Rogerella lecointrei nov. gen., nov. STINCHCOMB, B. L. & DARROUGH, G. 1995. Some mol-
sp. Comptes rendus hebdomadaires des Seances. luscan Problematica from the Upper Cambrian -
Academie des Sciences, Paris, 233, 1051-1053. Lower Ordovician of the Ozark Uplift. Journal
SAVAZZI, E. 1996. Adaptations of vermetid and sili- of Paleontology, 69, 52-65.
quariid gastropods. Palaeontology, 39, 157-177. SZULC, J. 1990. Diagenesis. In: SZULC, J. et al. (eds)
SCHMID, D. U. & LEINFELDER, R. R. 1996. The Jurassic International Workshop — Field Seminar: the
Lithocodium aggregatum - Troglotella incrustans Muschelkalk - Sedimentary Environments, Fades
foraminiferal consortium. Palaeontology, 39,21-52. and Diagenesis, Cracow-Opole 9—12 May 1990.
SCHMIDT, H. 1992. Mikrobohrspuren ausgewahlter International Association of Sedimentologists/
Faziesbereiche der tethyalen und germanischen Institute of Geological Sciences, Jagiellonian Uni-
Trias (Beschreibung, Vergleich und bathymetrische versity, 26-35
Interpretation). Frankfurter geowissenschaftliche TADDEI RUGGIERO, E. 1999. Bioerosive processes affect-
Arbeiten A, Geologie-Paldontologie, 12. ing a population of brachiopods (Upper Pliocene,
SCHNICK, H. 1992. Zum Vorkommen der Bohrspur Apulia). Bulletin of the Geological Society of
Hyellomorpha microdendritica Vogel, Golubic & Denmark, 45, 169-172.
Brett im oberen Obermaastricht Mittelpolens. TAPANILA, L. 2002. A new endosymbiont in Late
Zeitschrift fur geologische Wissenschaften, 20, Ordovician tabulate corals from Anticosti Island,
109-124. eastern Canada. Ichnos, 9, 109-116.
SCOTT, P. J. B., REISWIG, H. M. & MARCOTTE, B. M. TAPANILA, L. in press. Paleoecology and diversity of
1988. Ecology, functional morphology, behaviour, endosymbionts in Paleozoic marine invertebrates:
and feeding in coral- and sponge-boring species of trace fossil evidence. Lethaia.
Upogebia (Crustacea: Decapoda: Thalassinidea). TAPANILA, L. & COPPER, P. 2002. Endolithic trace
Canadian Journal of Zoology, 66, 483^495. fossils in Ordovician-Silurian corals and stroma-
SEILACHER, A. 1969. Paleoecology of boring barnacles. toporoids, Anticosti Island, eastern Canada.
American Zoologist, 9, 705-719. Ada Geologica Hispdnica, 37, 15-20.
478 R. G. BROMLEY

TAVERNIER, A., CAMPBELL, S. E. & GOLUBIC, S. 1992. A VERMEIJ, G. J. 1983b. Traces and trends of predation,
complex marine shallow-water boring trace: Den- with special reference to bivalved animals.
drorete balani n. ichnogen. et ichnospec. Lethaia, Palaeontology, 26, 455-465.
25, 303-310. VERMEIJ, G. J. 1998. Sabia on shells: a specialized
TAYLOR, J. D., CLEEVELY, R. J. & MORRIS, N. J. 1983. Pacific-type commensalism in the Caribbean Neo-
Predatory gastropods and their activities in the gene. Journal of Paleontology, 72, 465-472.
Blackdown Greensand (Albian) of England. VERMEIJ, G. J., ZIPSER, E. & DUDLEY, E. C. 1980.
Palaeontology, 26, 521-553. Predation in time and space: peeling and drilling
TAYLOR, P. D. 1993. Bryozoa. In: BENTON, M. J. (ed.) in terebrid gastropods. Paleobiology, 6, 352-364.
The Fossil Record 2. Chapman & Hall, London, VERMEIJ, G. J., ZIPSER, E. & ZARDINI, R. 1982. Break-
465-489. age-induced shell repair in some gastropods from
TAYLOR, P. D. & WILSON, M. A. 2003. Palaeoecology the Upper Triassic of Italy. Journal of Paleontol-
and evolution of marine hard substrate commu- ogy, 56, 233-235.
nities. Earth-Science Reviews, 62, 1-103. VOGEL, K., GOLUBIC, S. & BRETT, C. E. 1987. Endolith
TAYLOR, P. D., WILSON, M. A. & BROMLEY, R. G. 1999. associations and their relation to facies distribu-
A new ichnogenus for etchings made by cheilo- tion in the Middle Devonian of New York State,
stome bryozoans into calcareous substrates. USA. Lethaia, 20, 263-290.
Palaeontology, 42, 595-604. VOIGT, E. 1959. Endosacculus moltkiae n. g. n. sp., ein
TEICHERT, C. 1945. Parasitic worms in Permian vermutlicher fossiler Ascothoracide (Entomostr.)
brachiopod and pelecypod shells in Western Aus- als Cystenbildner bei der Oktokoralle Moltkia
tralia. American Journal of Science, 243, 197-209. minuta. Paldontologische Zeitschrift, 33, 211-223.
THIES, D. 1985. BiBspuren an Seeigel-Gehausen der VOIGT, E. 1965. Uber parasitische Polychaeten in
Gattung Echinocorys Leske, 1778 aus dem Maas- Kreide-Austern sowie einige andere in Muschel-
trichtium von Hemmoor (NW-Deutschland). schalen bohrende Wurmer. Paldontologische
Mitteilungen des Geologisch-Paldontologischen Zeitschrift, 39, 193-211.
Institutes der Universitdt Hamburg, 59, 71-82. VOIGT, E. 1967. Ein vermutlicher Ascothoracide (Endo-
THOMAS, A. O. 1911. A fossil burrowing sponge from sacculus (?) najdini n. sp.) als Bewohner einer
the Iowa Devonian. Bulletin of the Laboratory of kretazischen Isis aus der UdSSR. Paldontologische
Natural History, State University of Iowa, 6, 165- Zeitschrift, 41, 86-90.
166. VOIGT, E. 1970. Endolithische Wurm-Tunnelbauten
TODD, J. A. 2000. The central role of ctenostomes in (Lapispecus cuniculus n. g. n. sp. und Dodecaceria
bryozoan phylogeny. Proceedings of the llth [?] sp.) in Brandungsgerollen der oberen Kreide
International Bryozoology Association Conference, im nordlichen Harzvorlande. Geologische
Republic of Panama, Smithsonian Tropical Rundschau, 60, 355-380.
Research Institute, Balboa, 104-135. VOIGT, E. 1971. Fremdskulpturen an Steinkernen von
TODD, J. A. & PALMER, T. J. 2002a. The Jurassic bivalve Polychaeten-Bohrgangen aus der Maastrichter
genus Placunopsis: new evidence on anatomy and Tuffkreide. Paldontologische Zeitschrift, 45, 144—
affinities. Palaeontology, 45, 487-510. 153.
TODD, J. A. & PALMER, J. T. 2002b. Pattern versus pro- VOIGT, E. 1972. Uber Talpina ramosa v. Hagenow 1840,
cess or informative versus uninformative ichno- ein wahrscheinlich zu den Phoroniden gehoriger
taxonomy: comments on Nielsen and Nielsen, Bohrorganismus aus der Oberen Kreide, nebst
2001./ctoww, 9, 81-82. Bemerkungen zu den iibrigen bisher beschriebenen
TOMLINSON, J. T. 1969. Shell-burrowing barnacles. kretazischen 'Talpina'-Aften. Nachrichten der
American Zoologist, 9, 837-840. Akademie der Wissenschaften in Gottingen 2.
TRACEY, S., TODD, J. A. & ERWIN, D. H. 1993. Mathematisch-physikalische Klasse, 1972, 93—126.
Mollusca: Gastropoda. In: BENTON, M. J. (ed.) VOIGT, E. 1975. Tunnelbaue rezenter und fossiler
The Fossil Record 2. Chapman & Hall, London, Phoronidea. Paldontologische Zeitschrift, 49,
131-167. 135-167.
ULRICH, E. O. 1879. Descriptions of new genera and VOIGT, E. 1977. On grazing traces produced by the
species of fossils from the Lower Silurian about radula of fossil and Recent gastropods and
Cincinnati. Journal of the Cincinnati Society of chitons. In: CRIMES, T. P. & HARPER, J. C. (eds)
Natural History, 2, 8-30. Trace Fossils 2. Geological Journal Special
VENEC-PEYRE, M.-T. 1996. Bioeroding foraminifera: a Issues, 9, 335-346.
review. Marine Micropaleontology, 28, 19-30. VOIGT, E. 1978. Phoronidenbaue (Talpina ramose v.
VERMEIJ, G. J. 1977. The Mesozoic marine revolution: Hagenow) aus der maastrichter Tuffkreide.
evidence from snails, predators and grazers. Paleo- Natuurhistorisch Genootschap in Limburg, 28, 3-6.
biology, 3, 245-258. VOIGT, E. 1979. Wann haben sich die Feuersteine der
VERMEIJ, G. J. 1982. Gastropod shell form, breakage, oberen Kreide gebildet? Nachrichten der Akademie
and repair in relation to predation by the crab der Wissenschaften in Gottingen, 2. Mathematisch-
Calappa. Malacologia, 23, 1-12. Physikalische Klasse, 1979, 75-127.
VERMEIJ, G. J. 1983a. Shell-breaking predation through VOIGT, E. 1996. Submarine Aragonit-Losung am
time. In: TEVESZ, M. J. S. & McCALL, P. L. (eds) Boden des Schreibkreide-Meeres. Mitteilungen
Bio tic Interactions in Recent and Fossil Benthie des geologisch-paldontologischen Institutes der Uni-
Communities. Plenum Press, New York, 649-669. versitat Hamburg, 77, 577-601.
A STRATIGRAPHY OF MARINE BIOEROSION 479

VOIGT, E. & SOULE, J. D. 1973. Cretaceous burrowing WILSON, M. A. 1986. Coelobites and spatial refuges in a
bryozoans. Journal of Paleontology, 47, 21-33. Lower Cretaceous cobble-dwelling hardground
WALKER, S. E. 1989. Hermit crabs as taphonomic fauna. Palaeontology, 29, 691-703.
agents. Palaios, 4, 439^52. WILSON, M. A. & PALMER, T. J. 1988. Nomenclature of
WALKER, S. E. 1992. Criteria for recognizing a bivalve boring from the Upper Ordovician of the
marine hermit crabs in the fossil record using Midwestern United States. Journal of Paleontol-
gastropod shells. Journal of Paleontology, 66, ogy, 62, 306-308.
535-558. WILSON, M. A. & PALMER, T. J. 1990. A review of evo-
WARME, J. E. 1975. Borings as trace fossils, and the lutionary trends in carbonate hardground commu-
processes of marine bioerosion. In: FREY, R. W. nities. Paleontological Society Special Publication,
(ed.) The Study of Trace Fossils. Springer, New 5, 137-152.
York, 181-227. WILSON, M. A. & PALMER, T. J. 1992. Hardgrounds
WATKINS, R. 1990. Paleoecology of a Pliocene rocky and hardground faunas. University of Wales,
shoreline, Salton Trough region, California. Aberystwyth, Institute of Earth Studies Publica-
Palaios, 5, 167-175. tions^, 1-131.
WEBB, G. E. 1993. A Lower Pennsylvanian encrusting WILSON, M. A. & PALMER, T. J. 1998. The earliest Gas-
tabulate coral from a rocky shore environment trochaenolites (Early Pennsylvanian, Arkansas,
developed on the Mississippi-Pennsylvanian USA): an upper Paleozoic bivalve boring? Journal
unconformity surface in northwestern Arkansas. of Paleontology, 72, 769-772.
Journal of Paleontology, 67, 1064^1068. ZHANG, Y. & GOLUBIC, S. 1987. Endolithic microfossils
WEBB, G. E. 1994. Paleokarst, paleosol, and rocky- (Cyanophyta) from early Proterozoic stromato-
shore deposits at the Mississippian-Pennsylvanian lites, Hebei, China. Acta Micropalaeontologica
unconformity, northwest Arkansas. Geological Sinensis, 4, 1—12.
Society of America Bulletin, 106, 634-648. ZIBROWIUS, H. 1984. Deep-water scleractinian corals
WENZ, W. 1939. Gastropoda. In: SCHINDEWOLF, O. H. from the south-western Indian Ocean with crypts
(ed.) Handbuch der Paldontologie. Borntraeger, excavated by crabs, presumably Hapalocarcini-
Berlin-Zehlendorf, 6, 481-720. dae. Crustaceana, 43, 113-120.
WILLIAMS, A. B. & Nooc-Ho, N. 1990. Pomatogebia, a ZIBROWIUS, H. & ARNAUD, P. M. 1995. New records of
new genus of thalassinidean shrimps from Western molluscs (Leptoconchus, Lithophaga, Fungiacava)
Hemisphere tropics (Crustacea: Upogebiidae). that bore Indo-Pacific reef corals and their inter-
Proceedings of the Biological Society of Washing- actions with their hosts. Bulletin de la Musee Natio-
ton, 103, 614-616. nale d'Histoire Naturelle, Paris, 16A, 231-244.
This page intentionally left blank
Index

Note: Page numbers in bold refer to tables; those in italics refer to figures (while some are included within page
ranges).

Alberta, Canada, Viking Formation 40, 41, 42, 44, laminated sandstone 283-5, 287-8
45, 49, 50, 53, 55-7, 56 measurement 355-8
algal borings 455-6 muddy sandstone 188-91, 283, 285-7, 286
Alum Shale Member, North Yorkshire 146, 147, 148 mudstone 281
amalgamated sequence boundaries 51-7 preserved depth 383
animal behaviour 399 sandstones 193-4, 193, 283-*
ant trace fossils 373, 421, 423 siltstone 182-8, 279-83
see also Krausichnidae succession 16-17
Arab-D interval, Saudi Arabia 31 timing 383
Archeoentomichnus 434, 439, 446-7 bioturbation indices (BI) 356, 357
archosaurs see dinosaur tracks bite traces 466
Arenicolites 153, 164 bivalve trace fossils 10, 461-62
Denison Trough 285, 286, 287 Blue Lias, Lyme Regis, Dorset 151^
He Formation 241 body size 398-99
Argentina borings
palaeosol ichnofabrics 360-5, 367-71, 375 see also microborings
Triassic lacustrine deltas 335-54 acrothoracicans 464
Arizona, Bright Angel Shale 213-36 algal, cyanobacterial and fungal 457—8
arthropod trackways 7-8, 313, 318, 321, 322, 326, 346 ascothoracicans 465
Asencio Formation, Uruguay 365-7, 366, 370 attribution 455-7
Asteriacites 164, 165, 166, 167, 168 bivalves 461
Asterosoma 241, 249-50, 249, 250, 267, 303 Decapoda 465-6
Attaichnus 436, 437 echinoids 466
Australia foraminifera 467-8
Cambro-Ordovician formations 390, 391, 392 gastropods 462-3
Pebbley Beach Formation 179-212 in shells 466-7
shoreface/deltaic facies 273-310 small rosetted 458-9
Austria, Triassic-Jurassic extinction 409-11, 410 sponges 459-60
avulsion belt deposits 360, 361 worms 460-1
brachiopod etchings 464
Bandera Shale Formation, Kansas 162 brackish-water deposits 179-81, 202, 237
barnacle etchings 461-62 see also tidal flats
bathymetric reconstruction 4, 72-4 Bright Angel Shale, Arizona 213-36
bayhead delta 55-7, 268 ecosystem reconstruction 233—4
Beaconites 317, 321, 365 facies 220
bee trace fossils 362, 363, 364, 371, 372, 422, 423 ichnoguilds 223-7
see also Celliformidae lithofacies 217-23
beetles see coleopteran trace fossils palynofacies 227-30, 228-30
behavioural homologies 359 stratigraphy 215
Belichnus 465 bryozoans 455, 463—4
benthic fauna 146-7 Burniston dinosaur tracks 113-19, 113, 116, 118
benthic oxygen levels 398 burrows
Bergaueria 164, 241 chambered trace fossils 425, 435, 436-8
BI see bioturbation indices dimensions 15
Bichordites 83^, 84, 85, 87 echinoids and crustaceans 80, 82
bioerosional trace fossils 455-79 escape 10, 242, 243, 245-6, 246
see also borings; etchings Glossifungites ichnofacies 35
biogenic marks 224 homogenization 11
biomechanics of dinosaurs 95 horizontal 319
bioturbation insects 317,420
see also ichnofabrics interstratal 222
claystone and siltstone 281-83 iron-rich sandstones 225
depth and extent 10-11, 393, 397-8, 408 meniscate 375
intensity 13-14, 183, 296 Phycosiphon 245
482 INDEX

burrows (cont.) cocoons 420, 435


Savrda and Bottjer's model 142-3 coleopteran trace fossils 364, 365, 371, 372, 421, 422
shallow-tier 386, 394 see also Coprinisphaeridae; Pallichnidae
thalassinideans 79 colonization styles 16, 17-18
U-shaped 225 communities see ichnocoenoses; trace fossil
vertical 313, 314,315, 317 assemblages
Bynguano Formation, Australia 391, 392 composite ichnofabrics 375-6
Conchotrema 460
Callichurus major 79, 83 Conichnus 289
Calliopsis 372 continental environments
Cambrian see also deltaic facies; fluvial facies
base 399 fluvial ichnofaunas 215-16, 262, 315-19
Bright Angel Shale, Arizona 213-36 ichnofacies model 312-15, 572
earliest muddy sediments 385-6 lacustrine deltas 335-54
ichnological record 386—93 lacustrine ichnofaunas 319-23
radiation event 7 river-dominated facies 255, 344, 345, 347
canyons, submarine 37, 38 sequence stratigraphy 323-7
Carboniferous tidal environments 157-78 Coprinisphaera 312, 368, 369, 425, 427-30, 428, 445
Cardium Formation, Alberta 49, 51 Coprinisphaeridae 425, 426-32, 427, 428, 445-6
Caulostrep sis 460 Cornichnus 197
Cavernula 65, 66-7, 66, 457 correlation tool 18-19
cavities see microborings Cretaceous
Cellicalichnus 362, 363, 364, 423 graphoglyptids 134
Celliforma 423, 445 mass extinction event 414-15
Celliformidae 372, 419, 421, 425 palaeosols 362-7, 445-6, 447
Centrichnus 461-62 cross-bedded sandstones 287, 295
chambered trace fossils 419-53 see also hummocky cross-stratification
see also insect nests crustaceans
burrows 425 distribution 81-3
construction 420-1 ichnology 79-81
fillings 424 trace fossils 419, 464-5
ichnofamilies 425-44 Cruziana 222, 345, 346, 386, 387
ichnotaxobases 421-25 preservation 393, 394
identification 426, 427, 433, 436 tidal environments 164, 165, 166
shape 421 Cruziana ichnofacies 7, 41, 42, 45, 49, 284
wall types 421-24 Bright Angel Shale 233
channel environments 261-8, 315-17, 316 diverse proximal expression 301
channelized sandstone—mudstone 194—5, 196 Pebbley Beach Formation 184, 186, 188, 189, 190,
chiton radulation 463 191-3, 197, 208
Chondrites tide-influenced shorelines 161—2
deep-sea facies 130 Cunctichnus 460
Denison Trough 282, 288, 304 Curvolithus 164, 165, 166
ichnofabric 245, 251, 266, 267 cyanobacterial borings 457-8
ichnoguild 144-5 cyclic sediments 199-200
He Formation 241 Cylindricum 317
oxygen levels 144, 149-50, 150, 153, 154
Chondrites-Skolithos ichnofabric 242, 242, 266 Decapoda, borings 465—6
Cicatricula 460 deep-sea trace fossils 125-39, 130
Circulichnis 170 diversity 125-33, 135
Cladichnus 131 time range 726, 127-8
claystones 281,295-7, 360 deltaic facies 237
climatic control of distribution 77-92 analysis 277-81
Clionolithes 460 coarsening-upward sequence 297, 298, 302
closed lake ichnofaunas 320-1 delta front 252-7, 342-3, 342, 345-6, 346
coarse-grained facies 274 delta plain 343, 343, 348
coarsening upward sequence 297, 298, 302 Denison Trough, Australia 273-81, 290-1, 303
coastal facies 51-2, 194-9, 208, 299 distributary mouth-bar 253-5, 255, 257, 258
Cochlichnus 345, 346, 349 prodeltaic deposits 304, 301
INDEX 483

tide-dominated 237-72, 258 Echinoid Form 66, 69, 72, 74


Tonto Group 214 echinoids 81-3, 466
Triassic rift lakes, Argentina 335-54 ecology see palaeoecology
Dendrina 459 ecosystem reconstruction 233
Dendroidichnites 170 Ekdale and Mason model of oxygenation 144, 150
Dendrorete 459 Ellipsoideichnus 423
Denison Trough Permian sequences, Australia England
273-310,275,276 Blue Lias, Dorset 151-4
deltaic facies 273-81, 290-301, 303 Pinhay Bay, Dorset 407-8, 407-8
facies analysis 277-81, 282 Scalby Formation, Yorkshire 98, 113-19
offshore and shoreface deposits 281-90, 306 Toarcian, North Yorkshire 145-51
deoxygenation events 142 Triassic-Jurassic extinction 407-9
Desmograpton 131 Entisol 362, 363
Devonian mass extinction event 400 Entobia 459-60
Dietyodora 128, 129 Eocene 136, 367-71
Dimorphichnus 224 epichnial grooves 224
dinosaur tracks 93-123, 322 Equisitales 361, 362
anatomy and gait 112 erosion
experimental work 93-5, 96-100, 107-12 regressive surfaces 34-7, 38-41, 39, 42, 43
laboratory simulations 108-12 transgressive surfaces 39, 43-52, 203, 205-7
Diorygma 468 escape burrow ichnofabric 242, 243, 245-6, 266
Diplichnites 170, 171, 345 escape traces 315, 316
Diplocraterion 13, 17, 41 estuarine facies
Denison Trough 284, 286, 285-9 basin fill 202
ichnofabric 242-3, 242, 266 Bright Angel Shale 213-36
He Formation 241 channel fill 202, 203
oxygen levels 153 differentiation from offshore marine 179-212
Pebbley Beach Formation 188, 191, 193, 204, 205 He Formation 252-7, 252, 253
tidal environments 164, 165, 167 palaeoenvironments 265
Triassic 403-4, 407 Pebbley Beach Formation 194-9, 208
wide time range 77 valley fills 53-5
Diplopodichnus 170 etchings 461-3, 464
discontinuities 29-62, 204 euphotic zone 72
see also omission suites Eurygonum 457
marine flooding surfaces 43, 45, 46, 51-7, 184 experiments on dinosaur tracks 93-5, 96-100,
regressive surfaces of erosion 34-7, 38^4 107-12
sequence boundaries 34-8, 51-7, 202, 205, 207
transgressive surfaces of erosion 39, 43-52, 203, facies 21
205-7 see also deltaic facies; estuarine facies; ichnofacies;
distributary mouth-bar facies 253-5, 257, 258 lithofacies
diversity 14-15, 125-33, 135 Bright Angel Shale, Arizona 220
see also mass extinction events aftermath characterization 21
bioerosional fossils 468-9 Denison Trough, Australia 277-81, 252
changes 78 lacustrine deltas 340-5, 341, 344
salinity levels and assemblages 772 palaeocurrents, He Formation, Norway 257
Triassic-Jurassic extinction 408, 413 southern Sydney Basin 182-99
Dolomites, Italy, ichnostratigraphy 403-6 tidal channels 262
Douglas Group, Kansas 162 tide-dominated deltas, He Formation 254, 256
downhole imaging 21 failure of substrate 104-5, 106-7, 707, 116-19
drowned river valley 237 fair-weather conditions 187
dung-beetle nests 371, 372, 422 Fasciculus 65, 66, 67-8, 72-3, 74, 457
duricrusts 365, 367 feeding traces 314
dyads 227, 225, 229 ferricretes 365
dysoxia-trace fossils relationship 141-56 Fictovichnus 423, 425, 433-5, 434, 446
fillings
earthworm trace fossils 373 chambered trace fossils 424
Eatonichnus 424, 428, 430 estuarine facies 53-5, 202, 203
Echinocardium 80, 83 fine-grained sands 109-11
484 INDEX

firmgrounds Helminthopsis 170, 171, 283, 286, 304


early Cambrian 386, 393-4 Helminthorhaphe 130
Glossifungites ichnofacies 33-4 heterotrophs 72, 73
ichnofacies distribution 6 hiatus see omission suites
lacustrine environments 324, 325, 327 horizontal burrows 319
omission suites 46, 47, 49 hummocky cross-stratification (HCS) 187, 190, 191,
fish trace fossils 314, 322, 346, 466 193, 197, 255, 287
flaser bedding 195, 797 Hyellomorpha 457
flashcards 11 hypichnial casts 224
Fleaglellius 366, 372, 373, 441-2, 440, 446
flooding see marine flooding surfaces ichnocoenoses 18, 30-1
floodplain ichnocoenoses 315, 319 floodplain 315, 319
fluvial ichnofaunas oxygen-related 141-4
channels 215-16, 262, 315-17, 316 ichnofabrics
intertidal 169-73, 170, 171 see also bioturbation; palaeosol ichnofabrics
overbank374, 315-11,318 composite 375-6
flysch trace fossil assemblages 129 constituent diagram 12
Fontanai 426-7, 428, 445 He Formation, Norway 239, 241-51
footprints marine/non-marine intercalation 20
see also track morphology; trackways methods of analysis 11-18, 355-8
dinosaurs 8, 113-19 palaeoenvironments 14
indenter theory equipment 96-7, 98 scales of development 12
foraminifera borings 465-6 stacking patterns 18-19, 264, 268-9
forced-regressive shorefaces 37^3, 39 tidal channels 263
Freitag Formation, Australia 299, 300 ichnofacies
freshwater ichnofaunas see also Cruziana ichnofacies; Glossifungites
Bright Angel Shale 213-36 ichnofacies; Skolithos ichnofacies
fluvial 215-16, 262, 315-19 concept 4-7, 18
lacustrine 319-23 continental environments model 312-15, 312
lacustrine deltas 38-47 Corinisphaera 312
sequence stratigraphy 326-7 distribution 30
frontiers 19-22 fluvio-lacustrine succession 311-33
fugichnia 191, 193, 197 incised valley complexes 54
fungal borings 457-8 Mermia 312, 314-15, 316, 319, 320, 322, 324, 335,
348
gamma-ray spectrometry 145 Nereites 4
Gastrochaenolites 462 palaeoenvironments 5, 6
gastropod trace fossils 462-3, 467 Pebbley Beach Formation 200, 201
genetic stratigraphy 29 Scoyenia 4, 312-14, 318, 320, 322, 324, 326, 327
Globodendrina 66, 69,11, 459 substrate types 30-4
Glockerichnus 131 Teredolites 32-3
Glossifungites ichnofacies 4, 31, 33-4, 35, 49, 56 Trypanites 32
assemblages 36, 38 vertebrates 119
omission suites 41, 42, 46, 47 Zoophycos 4, 41, 42, 45, 144-5
Pebbley Beach Formation 205, 207 ichnofamilies see insect ichnofamilies
trace fossil associations 35, 41, 42 ichnofaunas
Gordia 170, 171, 346 fluvial 215-16, 262, 315-19
Grand Canyon, Bright Angel Shale 213-36 fluvio-estuarine intertidal 169-73, 170, 171
graphoglyptids 129-31 intertidal 162-3
grazing trails 314, 318 lacustrine 319-23
Grebs Nest Point Formation 387, 388-91 latest Permian 401-2, 402
Gyrochorte 168, 241, 242, 243, 266 open-marine intertidal 164-7, 185, 202
Gyrolithes 241, 386, 388 restricted-bay intertidal 167-9, 167
Rhaetian 405, 409-13
hardgrounds 30, 32 salinity levels and diversity 772
HCS see hummocky cross-stratification trace fossil assemblages 336-7
Helicotaphrichnus 460 ichnogenera
Helminthoidichnites 170, 346 estuarine facies 194-9
INDEX 485

Lyme Regis, Dorset 152-4, 752 mud 257-8, 260


marine fades 182-94 open-marine 164—7, 164, 166
range and diversity 126, 127-8 restricted-bay 167-9, 167
ichnoguilds sand 258-9, 260
Bright Angel Shale, Arizona 223-7 trace fossil distribution 775
Zoophycos-Chondrites 144-5 iron-rich sandstones 219-20, 279, 222-3, 231
ichnometry 15 Ischigualasto Formation 360-2, 364, 375
ichnostratigraphy 7-8 Ischigualasto-Villa Union basin 338-10, 339, 340
Cambrian 399-400
Coprinisphaeridae 445-6 Jebel Qatrani Formation, Egypt 367, 370
Krausichnidae 446-8 Jet Rock Member, North Yorkshire 146, 147
Pallichnidae 446 Joffre Shoreface Complex 49, 50, 55, 56
Triassic 403-6 Judy Creek Field, Alberta 41, 43, 44
ichnotaxa Jurassic
attribution 359 see also Rhaetian
bioerosional 455-79 archosaur footprints 8
climatic control 78 Blue Lias, Dorset 151-4
stratigraphic range 77, 128-33, 128, 729 He Formation, Norway 237-72
ichnotaxobases 421-25 mass extinction event 406-14
ichnotaxonomy 3^ palaeosol ichnofaunas 447
chambered trace fossils 419-53 Toarcian, North Yorkshire 145-51
composition 72
microborings 65-6 Kansas, USA, outcrops and stratigraphy 158, 759
vertebrate tracks 119-20 Kaybob Field, Alberta 41-3, 42
ICZN see International Code of Zoological key stratigraphic surfaces 19, 29-62
Nomenclature kinematics of track morphology 103^4
IHS see inclined heterolithic stratification Konservat-Lagerstatten 398
He Formation, Norway 237-72 Kossen Formation, Austria 410-11
conceptual facies model 254, 256 Kouphichnium 170
facies 255, 257, 261-8 Krausichnidae 426, 435-43, 437, 438, 441, 446-7
ichnofabrics241-51 Krausichnus 420, 423, 435-6, 438-9, 438, 443^
palaeoenvironments 252-61, 257 Kristin Field, offshore Norway 238-41, 238, 240
sedimentology 238-41, 240, 251-69
trace fossil assemblages 241 laboratory-simulated tracks 108-12
incised shoreface deposits 37-43, 44 lacustrine deltas
incised submarine canyons 37, 38 facies associations 340—45, 341, 344
incised valley complexes 52-5 marginal environments 343-5, 343, 347-9
inclined heterolithic stratification (IHS) 195, 198, 203 trace fossil assemblages 336-7, 341, 343, 345-9
indenter theory 94-5, 96-7, 98, 102-3 lacustrine ichnofaunas 319-23
infaunal communities 15-16, 81-3 Laguna Palacios Formation 362-5, 363, 364, 375
insect ichnofamilies 419-20, 425-44 lake basin types 324, 325
Celliformidae 419, 421, 425 laminated mudstone 199
Coprinisphaeridae 425, 426-32, 427, 428 laminated sandstone 187-93, 284, 285-90
Krausichnidae 426, 432, 433-42, 436, 437, 440 Lapispecus 460
Pallichnidae 425, 432-5, 433, 434 latitudinal distribution 78, 79, 81-3
stratigraphic range 444 Lias see Blue Lias; Toarcian
insect nests 8, 315, 419, 420 Lingulella2l3,233
ants 421 Lingulichnus 168
bees 362, 363, 364, 370, 371, 372 liquefaction failure 117-19
beetles 364, 365, 371, 372, 420 lithofacies, Bright Angel Shale 217-23, 220
termites 366, 370, 372, 373, 421 Lockeia 164, 165, 166, 167, 168, 241
wall types 421-24, 422 locomotion traces 313, 314, 318, 345, 349
International Code of Zoological Nomenclature loess sequence 362, 363
(ICZN) 3-4, 120 loosegrounds 30
intertidal flats Lower Aldebaran Sandstone 292, 297-9, 302
see also tidal flats lower shoreface deposits 792
fluvial ichnofaunas 169-73, 770, 777 lowstand incised shorefaces 37^43, 39, 44
model 158-62,160 Lyme Regis, Dorset, Blue Lias 151-4
486 INDEX

Macaronichnus Monesichnus 430, 431


Denison Trough 289-96, 297-9, 304, 306 Monocraterion 241
Pebbley Beach Formation 189, 191, 194 Monomorphichnus 213, 224
Maeandropolydora 461 mouth-bar facies 253-5, 253, 255, 265
Magenta Beds, Arizona 219-20, 219, 222-3, 231 mud flats 161
Mannville Group-Joli Fou Formation contact 51, 52 muddy sandstone 188-91, 281-5
marginal lacustrine environment 343-5, 343, 348-9 mudstones 281, 385-6
marine bioerosion 455-79 MZD see maximum zone of deformation
attribution to trace-maker 455-7
borings 457-61, 462, 464-6, 468 nektonic fauna 146
etchings 461-63, 464 Neogene graphoglyptids 133-6, 134-6
radulation 463 neoichnology 21, 94
round holes 466-8 Nereites 130, 164, 166, 167
stratigraphic range 456 Nereites ichnofacies 4
marine environments nests see insect nests
climatic control 77-92 Nevada, USA, Triassic-Jurassic extinction 411-13
ichnofabric 20 Nododendrina 459
ichnofacies 5-6 nomenclature 3—4, 65-6
siliclastic sediments 383-97 see also ichnotaxonomy
marine facies 253 non-deltaic shoreface deposits 281-90
estuarine deposit differentiation 179-212 non-marine ichnofabric 20
Pebbley Beach Formation 182-94, 202, 208 non-marine ichnofacies 5-6
marine flooding surfaces (MFS) 43, 45, 46, 51-7, Norway, Jurassic tide-dominated delta 237-72
184
mass extinction events aftermath 397^16 oceanic anoxic event 145
Cretaceous-Tertiary event 414-15 offshore deposits 281-90
late Devonian event 400 lacustrine facies 340-42
Ordovian—Silurian event 401 marine 179-212
Permian-Triassic event 400-406 tide-dominated 261
Triassic-Jurassic event 406-14 trace fossil assemblages 303, 304
Mauretania, microborings 63-76 Oichnus 466-7
maximum zone of deformation (MZD) 100-2 Oligocene diversity 136
medium-grained sands 111-12 omission suites 30^
meniscate trace fossils 313, 314, 317, 322, 375, 424 firmgrounds 46, 47, 49
Mermia 323 Glossifungites ichnofacies 41, 42, 46, 47
Mermia ichnofacies 312, 314-15, 316, 319, 320, 322, OPB see optical pedobarograph
324, 335, 350 open lake ichnofaunas 321-23
Mesozoic open marine ichnofaunas 164-7, 164, 166, 185, 202
bioerosion diversity 468 Ophiomorpha 79, 80-1, 80, 130
graphoglyptids 134 Cenozoic occurrences 84
trace fossil diversity 125, 126, 128, 131 ichnofabric 244-5, 244, 266
methodology of ichnology 9-19 He Formation 241
MFS see marine flooding surfaces organism distribution 81-3
microborings 63-76, 455-7 spatangoid trace fossils 83-7
bathymetric interpretation and model 72^ optical pedobarograph (OPB) 97-9, 112
morphology 66-72 ORB see oxygen-restricted biofacies
minute round holes 467-8 Ordovician
Miocene 83-4, 136 Grebs Nest Point Formation 387, 388-91
Mirandaichnium 170 ichnological record 386-93
Missouri, USA, outcrops and stratigraphy 158, 159 mass extinction event 400, 401
mixed layers see bioturbation Orthogonum 65, 66, 68-70, 69, 74, 458
models Ostrebium 72
conceptual facies 254, 256 overbank ichnofaunas 317-19, 318
continental ichnofacies 310-13, 312 oxygen-restricted biofacies (ORB) 145, 146-8, 151-2
oxygenation 141^, 150 oxygenation 8-9
tidal flats 158-62 evidence in Toarcian deposits 146-8
moisture/density relationships 105-6, 114-19 lacustrine environments 314, 322
mollusc shell microborings 63-76, 65 Lias, Dorset 151-2
INDEX 487

oxygenation (cont.) discontinuity surfaces 204


measurement 145 estuarine facies 194-9
models 141^ ichnofacies 200, 201
marine facies 182-94
Pacoota Sandstone Formation, Australia 75, 391 sequence stratigraphy 199-207, 200, 201, 206
palaeobathymetry 4, 72-4 trace fossil summary 202
palaeoclimate criteria 78 transgressive surfaces of erosion 203, 205-7
Palaeoconchelis 73, 458 pedofabric/ichnofabric ternary diagram (PITD) 357,
palaeoecology 358
analysis 398-9 pedofabrics 357, 360, 368, 371-2
Bright Angel Shale, Arizona 213-36 pellets 79
microborings 63-76 Pennsylvanian stratigraphy, USA 759
Tonto Group 214-16 permeability of substrate 106-7
palaeoenvironments Permian
analysis 1, 12, 395-6, 414-15 Denison Trough, Australia 273-309
delta front 252-7 ichnofaunas 401-2, 402
estuarine 265 mass extinction event 400-6
ichnofabrics 14 Pebbley Beach Formation, Australia 179-212
ichnofacies 5, 6 petroleum industry 19-21
He Formation, Norway 257 Petroxestes 462
tidal channel 261-8 Phoebichnus 241, 245, 246, 266
tidal flats 173, 257-61,2(50 photoautotrophic endoliths 72
wave- and tide-influenced 259, 290—2 Phrixichnus 468
Palaeogene diversity 126, 127 Phycodes 223, 388
palaeogeography, eastern Grand Canyon 216 Phycosiphon 130
Palaeophycus 227, 222, 346 burrows 245
Denison Trough 282, 284, 288-90, 293, 304 Denison Trough 282, 281-3, 288, 287, 292-6,
ichnofabric 242, 243, 248, 266, 267 297-301, 306
He Formation 241 ichnofabric 245, 246, 247, 251, 266
oxygen levels 153 He Formation 241
Pebbley Beach Formation 185, 187, 189 Pebbley Beach Formation 185, 187, 188, 191, 194
tidal environments 164, 167, 168 Pinhay Bay, Dorset, England 407-8, 407-8
Palaeosabella 461 piped zones 142, 143
palaeosol ichnofabrics 355—82 PITD see pedofabric/ichnofabric ternary diagram
analysis 355-8, 371-2 Planobola 457, 467
Argentina 360-5, 367-71, 375 Planolites 80, 153
Asencio Formation, Uruguay 365-7, 366, 370 Denison Trough 283, 287, 293
bedding and structures 356-7 ichnofabric 245-9, 266, 267
composite ichnofabrics 375—6 He Formation 241
Jebel Qatrani Formation, Egypt 367, 370 Pebbley Beach Formation 185, 187, 188, 195, 197
palaeosols Planovolites 468
chambered trace fossils 419-53 plant fossils 367
ichnotaxa range 436 Platydendrina 459
palaeotopography 162, 214, 216, 276 Plectonema 72
Palaeozoic Pleistocene assemblages 85, 86-7
arthropod trace fossils 7-8 Pliocene assemblages 84-5, 86
bioerosion 468 Podichnus 464
diversity 125, 126, 127-31 Polyactina 65, 66, 68, 458
graphoglyptids 133-4 porous plate box 96-7
ichnofabrics 383-96 preservation
Pallichnidae 425, 432-5, 433, 446 controls, track morphology 93-123
Pallichnus 423, 425, 434, 435, 446 Cruziana and Rusophycus 393, 394
Palmiraichus 422, 423 potential of lacustrine ichnofaunas 327
palynofacies 213-14, 227-30, 228-30 quality 384-5
palynomorphs 231-3 styles 385
Parowanichnus 434, 437 prodeltaic deposits 304, 305
pascichnia traces 150 progradational cycles 274-7
Pebbley Beach Formation, Australia 179-212 prograding shoreface succession 300
488 INDEX

Protovirgularia 164, 165, 166, 167, 168 sheet-like 195-9, 197


Psammichnites 164, 166, 167, 168, 197 silty 281^,295-7
Pygmy Form 65, 66, 71, 74 tabular 291-5
pyrite framboids 145, 151-2 tidal channels 17
sandy siltstones 185-7, 186
quantification of bioturbation 10-11, 355-7 Sarmiento Formation, Argentina 358, 367-71
Savrda and Bottjer model of oxygenation 141^
radiation events 7 SB see sequence boundaries
radulation 463 Scalby Formation, Yorkshire 98, 113-19
Radulichnus 463 scale of observation 10
Ramodendrina 459 Scaphichnium 425, 433, 434, 446
Ravenscar, Yorkshire, Toarcian deposits 145-51 scarabaeid beetles 435
ravinement 43-5, 49 scars 463
Rebuffoichnus 364, 365, 372, 432, 434, 445 Schaubcylindrichnus 241, 243, 244, 247, 248, 266, 267
Recent microborings 63-76 Scolecia66, 69, 70-1,72,458
red-bed succession 365-7 Scolicia 81, 82, 83, 84, 85, 130
regressive surfaces of erosion (RSE) 34-7, 38-41, 39, Scoyenia ichnofacies 4, 312-14, 313, 318, 320, 322,
42,43 324, 326, 327
Renichnus 462-3 sediment feeders 223
repartition 349 sedimentary structures 163, 164, 276
research landmarks 3-9 see also cross-bedded sandstones; soft sediment
reservoirs 21, 273 deformation
residual strength of substrate 106-7 sedimentology
restricted-bay intertidal ichnofaunas 167-9, 167 context 10
Reticulina 66, 69, 70, 458 ichnofabrics 4
Rhaetian ichnofaunas 405, 409-14 He Formation, Norway 239-41, 240, 251-69
Rhizocorallium 149, 153, 164 intertidal flats and subtidal sandbars 160
Denison Trough 284, 286, 288, 287, 289 lacustrine deltas 347
He Formation 241 Pebbley Beach Formation, Australia 179-212
Pebbley Beach Formation 185, 188, 191 Seilacherian ichnofacies 4-7, 18
Servino Formation 405, 406, 412 semi-quantitative methods 355-6
Rhopalia 458 sequence boundaries (SB) 34-7, 38, 51-7, 202, 205,
rift basins 274 207
rift lakes 335-54 sequence stratigraphy 9
ripple structures 195, 196, 243, 253 case studies 34-51
river-dominated facies 253, 344, 345, 347 fluvial ichnofacies 326-7
see also fluvial ichnofaunas lacustrine ichnofacies 324, 325-6
Rogerella 464 Pebbley Beach Formation 199-207, 200, 201, 206
Rosellichnus 423, 424 Servino Formation, Italy 403-6, 405
Rosselia shallow marine siliclastics 383-96
Denison Trough 278, 284, 288, 287, 292, 296, 300 shallow-tier burrows 386, 394
ichnofabric 249-50, 249, 267 shear surfaces 107
He Formation 241 sheet-like sandstone-mudstone 195-9, 797
Pebbley Beach Formation 185, 187, 188, 191, 194 shelf storm deposits 230-1
Rotundusichnium 130 shell borings 465, 466-7
RSE see regressive surfaces of erosion shorefaces
Rusophycus 166, 386, 388, 391, 392, 393, 394 comparison with delta front 306
facies 278, 281-90
Saccomorpha 65, 66, 67, 74, 456 forced regressive and lowstand incised 37^3, 44
salinity levels 81-2, 772 prograding 300
sandbars 160, 231 transgressively incised 39, 47-51, 50
sands 109-12, 258-9, 260 siltstones 182-8, 184, 281-4, 297
sandstones silty sandstones 281-3, 295-7
bioturbated 193-4, 193 Silurian mass extinction event 400, 401
cross-bedded 290, 295 simulated dinosaur tracks 98, 108-12, 709, 770
deltaic 290-301 Siphonichnus 241, 250-1, 250, 267
laminated 187-93, 284, 285-9 Skolithos3l5
muddy 188-91, 286, 284-7, 288 Denison Trough 287, 288, 289
INDEX 489

ichnofabric 247, 248, 267, 285, 289 Talpina 461


He Formation 241 taphonomic pathways 316, 320
palaeosol 365 taxonomy see ichnotaxonomy
Pebbley Beach Formation 194, 195, 197 Teichichnus 167, 168, 389, 390
piperock 391 Bright Angel Shale 223
Skolithos ichnofacies 4, 42, 53, 161, 188, 312, 316, Denison Trough 281, 285, 303
320, 321, 324, 327 He Formation 241
Skolithos-Cruziana ichnofacies 188, 195, 208, 284 Pebbley Beach Formation 185, 187, 189-91
small borings 458-9, 467-8 Teisseirei 366, 372, 421, 424, 430-2, 431, 445
soft sediment deformation structures 293, 293, 304 temperate association 85-7
softgrounds 6, 30, 41, 42, 324, 325, 327 Teredolites 462
soil mechanics 105 Teredolites ichnofacies 32-3
soil moisture content 105-6, 114-19 termite nests 366, 370, 372, 373, 421, 422-3
soils see palaeosols see also Krausichnidae
Solemya 166 Termitichnus 441-2, 440, 443^, 446
soupgrounds 30 Tertiary
spatangoid trace fossils 80, 81, 83-7 diversity 136
Spirichnus 461 graphoglyptids 133-6, 134-6
sponge borings 459-60 insects 8
Spongeliomorpha 80-1 mass extinction event 414—15
stacking patterns 18-19, 264, 268-9 palaeosol ichnofaunas 444-6
Stiallia 170, 171 tetrads 227, 228, 229
Stiaria 170, 171, 345 Tetrapod tracks 170
stiffgrounds 30 textures see ichnofabrics
storm deposits 187-8, 189-91, 230-1, 287-8, 385 thalassinidean crustaceans 79, 81
stratigraphic range 77 Thalassinoides 41, 80-1, 86, 87, 153
bioerosive ichnotaxa 456 ichnofabric 167, 248, 249, 250, 251
ichnotaxa 77, 128-33, 128, 729 He Formation 241
insect ichnotaxa 436 Triassic 406, 412
lacustrine delta assemblages 336-7 tidal channel environments 17, 253-6, 257, 261—8
stratigraphic surfaces 19, 29-62 tidal flats 157-78
stratigraphy see ichnostratigraphy; sequence see also subtidal environments; supratidal zone
stratigraphy facies associations 260
striated traces 314 ichnofaunas variability 163-73
sub-recent microborings 63-76 model 158-62, 160
subaerially exposed surfaces 51—2 palaeoenvironments 257-61, 260
submarine canyon incision 37, 38 sedimentary structures 163, 164
substrate-controlled ichnofacies 30-4 stratigraphic implications 173-4
subsurface tracks 99, 104-5, 109-12 successions 259
subtidal environments 158-62, 160, 259-61, 260 variability 175-8
subtropical associations 83-5 tide-dominated deltas 237-72, 259
supratidal zone 160-1 tiering 15-16, 78, 82
surface traces 225, 226 Blue Lias 153
surface track simulations 99, 108—9 diagrams 357-8, 397
surfaces oxygenation events 141^, 142
see also discontinuities palaeosols 357, 358, 362, 363, 376
key stratigraphic surfaces 19, 29-62 time range and diversity 126, 127-8
shear surfaces 107 Toarcian deposits, North Yorkshire 145-51
subaerially exposed 51-2 Tonganoxichnus 170, 171
suspension feeders 223-5 Tonto Group, palaeoecology 214-16
Sydney Basin, Australia 179-212 Topsentopsis 460
Syntermesichnus 440, 442, 446 trace fossil assemblages
see also ichnofacies
tabular sandstone 288-92 Blue Lias, Dorset 152-4
Tacuruichnus 437, 439, 446 continental 311-33
Taenidium 153, 185, 315, 362, 363, 365, 375 deep-sea ichnotaxa 128-33
ichnofabric 244-5, 244, 248-9, 249, 266, 267 delta front 345-6, 346
He Formation 241 delta plain 348
490 INDEX

trace fossil assemblages (cont.} tropical associations 83-5


distribution controls 349-50 Trypanites 461
diversity and salinity levels 772 Trypanites ichnofacies 32
dysoxia relationship 141-56 TSE see transgressive surfaces of erosion
graphoglyptid proportions 132, 133 tsunami beds 414—15
He Formation, Norway 241 tubular trace fossils 374-5
intertidal 173 tuffaceous palaeosols 362-5, 363, 368
lacustrine deltas 336-7, 341, 345-9, 346, 347 tunnels 67-8
lacustrine sequence stratigraphy 324 see also burrows; graphoglyptids
mass extinction events aftermath 397-418 turbidites 322, 325-6
Mesozoic 131
Neogene 131 U-shaped burrows 225
palaeosols 359-60 UK see England
Palaeozoic 129-31 Ultisols 366, 370
Pebbley Beach Formation 202 Undichna 170, 171, 345, 346
prodeltaic v. offshore deposits 303, 304 upwelling areas 63-76
proximal Cruziana ichnofacies 191, 192 Uruguay, Asencio Formation 365-7, 365, 370
repartition 349
Toarcian, North Yorkshire 147, 148, 149-50 valleys 52-5, 162, 237
track morphology 93-123 vertebrates
Burniston dinosaur tracks 113-19 body fossils 367
description and terminology 100—2 dinosaur tracks 93—123
experimental work 93-5, 96-100, 107-12 ichnofacies 119
indenter theory 94-5, 96-7, 102-3 track taxonomy 119-20
moisture/density relationships 105-6, 114—19 trackways 313, 318, 322
surface and subsurface features 99-100, 101 vertical burrows 313, 314, 315, 317
trackways Vertisols 360-2, 361, 364
arthropods 7-8, 313, 318, 321, 322, 326, 346 Viking Formation, western Canada 40, 41, 42, 44, 45,
vertebrates 313, 318, 322 49, 50, 53, 55-7, 56
transgressive surfaces of erosion (TSE) 39, 43-52, Vondrichnus 440, 441, 446
203, 205-7
transgressively incised shorefaces 39, 47-51, 50 walls in chambered trace fossils 421-24
Treptichnus 170, 777, 346, 386, 388, 399 wave-dominated environments 259, 291, 344,
Triassic 345-6, 347
archosaur footprints 8 wavy bedding 195, 196, 197
ichnostratigraphy 403-4 woodgrounds 30, 32-3
lacustrine deltas, Argentina 335-54 worm borings 460-1
mass extinction events 399^14 wrinkle marks 226
palaeosols 360-2, 446-7
Triassic-Jurassic mass extinction event 406—16 Yorkshire, UK
Central Austria 409-11, 410 Burniston dinosaur tracks 113-19, 773, 116, 118
England 407-8, 407-9 Toarcian 145-51, 146
Nevada, USA 411-13
Trichichnus ichnofabric 251, 267 zone fossil characteristics 7
Trichophycus 387, 389-90 Zoophycos 284, 286, 287, 306, 406
tridactyl dinosaur tracks 98, 113, 116, 118 Zoophycos ichnofacies 4, 41, 42, 45, 144-5
Tripartum Form 66, 69,11, 74 Zoophycos-Chondrites ichnoguild 144—5
trophic reconstruction 231-3

Das könnte Ihnen auch gefallen