Applications of Ichnology to Petroleum Exploration A Core Workshop Organized and Edited by S. George Pemberton SEPM Core Workshop No. 17 Calgary, June 21, 1992 Copyright (992 by SEPM (Society for Sedimentary GeoISBN #0-918985-97-8 Cover illustration by Thomas D. Saunders Additional copies of this publication may be ordered from SEPM. Send your order to SEPM Post Office Box 4756 Tulsa, Oklahoma 74159-0756 US.A, © Copyright 1992 by SEPM (Society for Sedimentary Geology) Printed in the United States of AmericaPREFACE The field of ichnology (the study of animal-sediment relationships) is undergoing rapid expansion. Increased significance is being attached to trace fossils in environmental and didge- netic interpretations of rock units and in establishing basic stratigraphic frameworks. The sub. ject, therefore, is of importance not only for ichnologists but also for invertebrate and vertebrate palcontologists, palcoecologists, sedimentologists, stratigraphers, and resource geologists Unlike most other branches of paleontology, trace fossils are also amenable for study in subsur- face core and as such, have proven to be powerful tools in subsurface facies analysis and Sequence stratigraphy. The main purpose of this workshop is: a) to introduce the basic concepts of ichnology: b) to learn how to recognise basic types of trace fossils in core; c) to place these, structures in their appropriate paleontologic, sedimentologic, and stratigraphic content: and d) to Integrate this data with other lines of evidence to aid in petroleum exploration The papers presented in this volume pertain to Cretaceous deposits in the Western Interior, especially the Westem Canadian Sedimentary Basin. [have been fortunate enough to put together a series of papers that concentrate on marginal marine to shallow marine environ. ments in siliciclastic systems. Where possible the authors have tried to place the rocks in Sequence stratigraphic framework and have highlighted how the ichnological data has strength- ened their interpretation. T hope that this group of papers will serve to illustrate how an integrat- ed approach using combined ichnologic, sedimentologic, and stratigraphic lines of evidence ean constitute a powerlul approach to subsurface sis and interpretation. On & more personal note, this volume is very special to me for two reasons. First it is dedicated to my dear friend and colleague Bob Frey who passed away January 1, 1992. Secondly because most of the authors are either past or present students who have gone through my program at the University of Alberta. One of the true pleasures of the teaching profession is being able to instruct graduate students. Since starting at the University of Alberta in 1984 1 have been fortunate enough o be associated with a wonderful group of students who have been a source of great personal pride to me. Over these past eight yeurs this group has included 5 PhD and 22 MSc students. Their collective contributions have made this volume possible. S. George Pemberton Edmonton, Alberta 1992 iiiAC) OWLEDGMENTS This core workshop and volume were made possible only with the invaluable assistance of many people. | would like to thank Ray Garber of Chevron Canada for initiating the project, Ben Robertson of Chevron Canada for logistic support, and ‘Trish Auberle, Robin Dixon and the SEPM staff for handling everything in a most professional manner. I would like to express my gratitude to Ray Shepard and the staff of the Energy Resources Conservation Board Core Research Facility for providing the venue. The graphics, photography, and typesetting were done by the skilled stafl of Exxon Production Research and { thank photographer John Blankinship, ypeserter Daryll Ritter, and illustrators Donna Bartels, Judy Becker, Larry Besong, Charles Burwell, Becky Everitt, Mack Johnson, Kevin Linke, Becky Miller, and Diane Reina for their ctiorts. Additional typing was done by Joanne Wells of Exxon and Catherine Yuill and John Bourak of the University of Alberta. T would also like to thank two of my PhD students, Mike Ranger, who for the past five years has served as virtually a co-supervisor for my students and James MacEachern. who more recently has become a valuable addition to our program, Both af these people have gone way beyond what is expected and are always willing to share their experience and knowledge This volume was completed during my tenure as a visiting scientist at Exxon Production Research in Houston, ‘Texas. special thankyou to John Van Wagoner for initiating my visit: his views on sequence stratigraphy gained from years of experience in the ficld and in industry has given me the opportunity to apply a new stratigraphic perspective to ichnology. 1 would also like to thank Bob Todd and Kirt Campion for their generous support of my work und the rest of the geologists at Exxon who have made my stay so memorable. In conclusion, I must thank my wonderful wife Teresa, my daughters Sarah and Erin, and my son Joshua for all their love and support. I couldn't do what I do without them! ivLIST OF CONTRIBUTORS DAVID J. BECHTEL Department of Geology University of Alberta Edmonton, Alberta T6G 253 BRUCE M. BEYNON Chevron Canada S SW. TRACY BRADLEY Norcen Energy Resources Ltd. Calgar Canad, T2P 2X7 HOWARD G. BREKKE Department of Geology University of Alberta Edmonton, Alberta Canada, T6G 2E3 ROBERT W. FREY (Recently De University of Ge orgiat Athens, ( 30602, UL Department of Goole University of Alberta monton, Alberta. ida, T6G 21 BILL MALE AEC Oil and Gas Company 2400, 639 - Sth Ave. S.W. Calgary, Alberta, Canada, T2P OM9 SIMON A.J. PATTISON Imperial Oil Ltd. 7 - 4th Ave. S.W. nada, T2P OH6 S. GEORGE PEMBER! Department of Geology University of Alberta Edmonton, Alberta. Canada, T6G 263 INDRANEEL RAYCHAUDHURI Department of Geology University of Alberta Edmontor e ERRY E. RELNSON ‘ological Survey of Canada 3303 - 33rd, St. NW. Alberta. Canada, T2L 2A7 JOHN C, VAN WAGONER Exxon Production Research P.O. Box 2189 Houston, Texas. T1027, USA. GRANT WACH xon Production Research P.O, Box 2189 Houston, ‘Texas 77027, U.S.A. DARYL M. WIGHTMAN Alberta Cicologi P.O, Box 8330 § monton, Alberta ‘anada, T6H SXTABLE OF CONTENTS THE CONCEPTUAL FRAMEWORK OF ICHNOLOGY by S. G. Pemberton, R. W. Frey, M. J. Ranger, and J. A. MacEachern.. THE SEDIMENTOLOGY AND ICHNOLOGY OF THE LOWER CRETACEOUS ({ALBIAN) BLUESKY FORMATION IN THE KARR AREA OF WEST-CENTRAL ALBERTA by W. H. Male. 2033 ICHNOLOGICAL ASPECTS OF CRETACEOUS SHOREFACE SUCCESSIONS AND SHOREFACE VARIABILITY IN THE WESTERN INTERIOR SEAWAY OF NORTH AMERICA by J. A. MacEachem and S. G. Pemberton ICHNOLOGY AND EVENT STRATIGRAPHY: THE USE OF TRACE FOSSILS IN RECOGNIZING TEMPESTITES by S. G. Pemberton, J. A. MacEachern and M. J. Ranger,.., BS ICHNOLOGICAL AND SEDIMENTOLOGICAL CHARACTERISTICS OF OPEN MARINE TO STORM DOMINATED RESTRICTED MARINE SETTING WITHIN THE VIKING/BOW ISLAND FORMATIONS, SOUTH-CENTRAL ALBERTA by I. Raychaudhuri and S. G. Pemberton wl D ICHNOLOGICAL CHARACTERISTICS OF BRACKISH WATER DEPOSITS by S. G. Pemberton and D, M. Wightman... 141 STRATIGRAPHIC APPLICATIONS OF THE GLOSSIFUNGITES ICHNOFACIES: DELINEATING DISCONTINUITIES IN THE ROCK RECORD. by J. A. MacEachem, I. Raychaudhuri and $. G. Pemberton... 169 ICHNOLOGICAL SIGNATURE OF A BRACKISH WATER DEPOSI AN EXAMPLE FROM THE LOWER CRETACEOUS GRAND RAPIDS FORMATION, COLD LAKE OIL SANDS AREA, ALBERTA by B. M. Beynon and S. G. Pemberton... 199 viRECOGNITION AND INTERPRETATION OF ESTUARINE MUDSTONES (CENTRAL BASIN MUDSTONES) IN THE TRIPARTITE VALLE’ FILL DEPOSITS OF THE VIKING FORMATION, CENTRAL ALBERTA. by S.A. J. Pattison... ICHNOLOGY AND SEDIMENTOLOGY OF TRANSGRESSIVE DEPOSIT! TRANSGRESSIVELY-RELATED DEPOS! AND TRANSGRESSIVE SYSTEMS TRACTS IN THE VIKING FORMATION OF ALBERTA by J. A. MacEachern, D. J. Bechtel and S. G. Pemberton ea... COMPARATIVE ICHNOLOGICAL ANALYSIS OF LATE ALBIAN ESTUARINE VALLEY-FILL AND SHELF-SHOREFACE, DEPOSITS, CRYSTAL VIKING FIELD, ALBERTA by S. G. Pemberton, G. E. Reinson and J. A, MacEachern .. see 291 DEPOSITIONAL FACIES AND TRACE FOSSILS OF A LOW WAVE ENERGY SHOREFACIES SUCCESSION, ALBIAN VIKING FORMATION, CHIGWELL FIELD, ALBERTA, CANADA by I. Raychaudhuri, H. G. Brekke, S. G. Pemberton, and J. A, MacEachern ICHNOFACIES OF A WAVE-DOMINATED SHORELINE by S. G. Pemberton, J. C. Van Wagoner, and G. D. Wach...... EXAMPLES OF ICHNOFOSSIL ASSEMBLAGES IN THE LOWER CRETACEOUS WABISKAW MEMBER AND THE CLEARWATER FORMATION OF THE MARTEN HILLS GAS FIELD, NORTH-CENTRAL, ALBERTA, CANADA by T. L. Bradley and S. G. Pemberton..... 383 THE SEDIMENTOLOGY AND ICHNOLOGY OF ESTUARINE POINT BARS IN THE McMURRAY FORMATION OF THE ATHABASCA OIL SANDS DEPOSIT, NORTHEASTERN ALBERTA, CANADA by M. J, Ranger and S. G. Pemberton. GLOSSARY. viiROBERT WAYNE FREY (1938-1992) viiiDEDICATION TO ROBERT W. FREY S. George Pemberton Department of Geology, University of Alberta Edmonton, Alberta, Canada, T6G 2E3 Robert Wayne Frey's death, January 1, 192 after a long and courageous fight with can- cer, deprives Ichnology of one of its most influential figures. Bob will long be remembered for his monumental contributions to our understanding of the basic principles of organism-substrate relationships in both modern and ancient settings. In appreciation of his fundamental work, this volume is dedicated to Bob Frey's memory. Bob Frey’s career spanned a period of dynamic change in ichnology, from a time when ichnologists were few in number to the present day when paleontologists. sedimentologists and stratigraphers are concerned with trace fossil analysis to aid in their interpretations. To this rapid scientific evolution, the contributions of Bob Frey have been numerous, varied and almost always fundamental. He can be considered without doubt the “Father of American Technology”. He had a direct hand in shaping the modern principles of ichnology and we all owe him a great debt. A more detailed version of this memorial along with a complete list of Bob Frey's publications will be published soon (Pemberton, 1992) Bob Frey was born on a farm in Cleveland County, Arkansas on December 13, 1938, He graduated from the New Edinburgh High School in June 1956 and immediately joined the United States Air Force. He served from 1956 - 1960 and was honorably discharged with a top- security clearance. Bob then attended Lower Columbia College in Longview, Washington from 1960 - 1962. It was at Longview that Bob met and married Sharon Cowles who was a tower of strength for Bob during his long illness. In 1962, Bob transferred to the University of Montana in Missoula where he graduated with a B.A. in December, 1964. At Montana Bob acquired a keen interest in paleontology and decided to do graduate work at Indiana University, where he came under the tutelage of Don Hatin. From Indiana Bob received the M.A. degree in 1967 and the Ph.D degree in 1969. His dissertation, entitled “Stratigraphy, ichnology, and paleoecology of the Fort Hays Limestone Member of the Niobrara Chalk (Upper Cretaceous) in Trego County, Kansas” stands, along with Richard Osgood’s “Trace fossils of the Cincinnati area”, as a benchmark in the development of ichnology in North America. Bob successfully applied traditional paleoecological concepts to trace fossils and used them as essential elements in the interpretations of the depositional history of the Niobrara, He soon realized that in order to comprehend fossil traces, one must first observe and understand how recent organisms behave. His first introduction to modern organism-sediment relationships occurred while he was still at Indiana, He attended a field seminar at Beaufort, North Carolina and in typical Frey-style was able to translate his work into a series of four Papers in the Journal of Paleontology. This has always been a trademark of Bob Frey, knowing instinctively how to optimize his research opportunities and then disseminate that information, Afier graduation, Bob started his long and productive association with the University of Georgia, first as a Post-Doctoral Research Associate (1968 - 1969), then Research Associate, Marine Institute, Sapelo Island (1969 - 1970), Assistant Professor (1970 - 1974), Associate Professor (1974 - 1978), finally rising to the rank of Professor in 1978, It was at Sapelo Islandthat Bob teamed up with Jim Howard, the resident sedimentologist at the Skidaway Institute of Oceanography in Savannah, Georgia, The main thrust of their joint work was an evaluation of the ichnology and geologic significance of present-day benthic organisms found in all major marine environments of Georgia: the continental shelf, inlet shoals, beaches, dunes, washover fans, relicit mud deposits, estuaries, tidal flats, point bars, channel margins, and salt marshes. Their evaluations embraced: (1) characteristic environmental distributions of animals and plants and their associated ecologic and sedimentologic parameters (e.g., wave or current energy, salinity, feeding adaptations, sub- strate coherence, and sediment textures and fabrics), (2) the role of organisms as geologic agents, including not only their behavior and habitat adap- tations but also their inter-relationships with physical processes of erosion, reworking, trans- portation, and deposition of sediments: (3) potential preservability of the organisms as fossils, together with their associated physical and biogenic sedimentary structures, facies relationships, and stratigraphic sequences; and (4) fidelity of these preserved or preservable features as potential geologic indicators of ancient organisms and their adaptations, environments, and ecologic-sedimentologic parameters repre- sented in the fossil record. Although Bob and Jim were mainly responsible for the conception, coordination, execu- tion, and completion of the research; numerous graduate students also were involved. In addi- tion, during certain phases of the research they were joined by the staff of the renowned Senckenberg Institute for Marine Geology and Biology, Wilhelmshaven, Germany including Hans Reineck, and Gunther Hertweck, and Jurgen Dorjes (recently deceased). The results of this research program are impressive, rarely has a single coastal zone received such comprehensive geologic coverage. Consequently, the work attracted international attention and is widely cited in the literature. Characteristics of the Georgia coast became a model for comparison with numerous other modern and ancient systems. After perfecting their methodology in Georgia, Bob and Jim took their show on the road - literally, with the help of a bus turned into a mobile laboratory. This led to extended soirees to the Holocene of Florida, North Carolina, California, Massachusetts and Korea. Bob Frey’s publication record is most impressive with one hundred and thirty-one entries included are: eighty-five journal articles, thirty-four contributions to symposium compendia, guidebooks, short course manuals, and chapters of books: five edited volumes; five versions of an Historical Geology laboratory manual; and two monographs, His papers were scientifically innovative, lucid, and profusely illustrated, they were also virtually error-free. In every discipline there exists a body of literature that is considered “must reading” in order to fully comprehend its significance. In Ichnology, approximately twelve papers come to mind, of these, Bob authored or co-authored five of them. He edited the first summary book devoted entirely to the study of trace fossils that stood, for many years, as the definitive reference on the subject. He was co-editor and co-founder of Ichnos the first journal devoted entirely to organism-substrate rela- tionships. Bob worked hard at not only promoting Ichnology by also at helping young aspiring sci- entists. Although almost every facet of ichnology was addressed by Bob he was equally adept dis- cussing such diverse topics as salt marsh dynamics, botany, invertebrate zoology, taphonomy, mol- luscan paleoecology, diagenesis, bryzoan taxonomy, and clastic sedimentology. The quality of Bob’s work has been recognized by a number of awards and distinctions: in 1981 he was awarded a “Medal for Excellence and Creativity in Research” from theUniversity of Georgia; in 1984 he was co-winner of the “Best Paper Award” for Volume 56, Journal of Paleontology, awarded by the Society of Economic Paleontologists and Mineralogists; in 1984 he was given the R. C. Moore “Gechawk” Medal while serving as the Merrill W. Haas Distinguished Visiting Professor at the University of Kansas; in 1990 he was bestowed with the Richard Owen distinguished alumnus award from the Department of Geology, Indiana University; and has recently been nominated for the R. C. Moore Medal awarded by the Society of Economic Paleontologists and Mineralogist. Bob was a dedicated teacher who genuinely enjoyed his time in the class room. Bob was voted “Professor of the Year” in 1980 by the graduate students in the Department of Geology, University of Georgia, Reflecting the interest and mutual respect they held for each other, He served as major professor for 20 MSc. students in a diverse array of fields ranging from bivalve systematics to organic geochemistry. He directed 8 PhD students, (Paul Basan, Joe Wadsworth, Jennifer Smith, Steve Henderson Diane Kamola, M. A. Al-Aawah, Kent Sprague, and Anthony Martin). The quality of their work was a subject Bob took quiet pride in, He took particular pleasure in seeing his students grow not only as scientists but also as individuals, believe that the measure of one’s value to a discipline, can be measured by the regard in which one is held by his peers in the scientific community at large. Bob’s services as a critical reviewer were sought by the editors of 18 international journals and the program directors of five granting agencies. He served as the Field Trip Guidebook Editor for the Geological Society of America 1980 National Meeting in Atlanta and as Associate Editor for the Journal of Sedimentary Petrology. He was a major participant in 6 short courses run by the Society of Economic Paleontologists and Mineralogists, the Geological Society of America, and the United States Geological Survey, and assisted in leading 10 major field trips for various professional societies. In 1968 he initiated and co-authored (with Jim Howard) the “Ichnology Newsletter", an informal international newsletter on animal-sediment research, this publication continues to be distributed yearly. Bob is survived by his wife Sharon, a very special person who was devoted to him, his daughter Valerie and his son Eric. He was a loving husband and father who was very proud of his family. Bob was a modest, unassuming man and the only time you heard him boast was when he spoke of the accomplishments of Sharon, Valerie, or Eric. Those who knew Bob will remember him for his personal qualities - his enthusiasm, generosity, compassion, and high prin- ciples. His colleagues will sorely miss the stimulus his ideas brought to their own work, Ichnology will continue to thrive because it is built on a solid foundation constructed and nurtured by men like Bob Frey. During the last four years, Bob waged a heroic battle against cancer while still maintaining an active research program and fulfilling his teaching obligations. In doing so, he set an example in courage and dignity that will be an inspiration to all of us who knew him. Bob Frey left behind a great legacy of knowledge in Ichnology. His good friend Jim Howard said it best in the citation for Bob's “Best Paper Award” in the Journal of Paleontology “In recognition of Robert W. Frey, a nice guy who works too hard. Many of us today and in the future are and will be the fortunate recipients of his unending creative efforts, He is dedicated to his work because he likes what he is doing and he gets excited about it. We and he can’t ask for much more than that”. T was proud to have been his colleague but prouder to have been his friend. I will miss him dearly. xiACKNOWLEDGMENTS I would like to thank the entire Frey family, especially Sharon for their hospitality during my visits to Athens. They always made me feel at home, even under tragic and trying circum- stances. My thanks to Valerie Frey who took the picture of Bob that I used in this memorial, I thought it truly captured Bob’s persona. REFERENCE Pemberton, S. G, 1992. Memorial to Robert Wayne Frey (1938-1992). Ichnos, 2.THE CO! CEPTUAL FRAMEWORK OF ICHNOLOGY . George Pemberton! Robert W. Frey?, Michael J. Ranger!, and James MacEachern! 1, Department of Geology, University of Alberta, Edmonton, Alberta, : Canada, T6G 2E3. pepartment of Geology, University of Georgia, Athens, Georgia, 30602, U.S.A. (recently deceased), ABSTRACT fossils are difficult to identify and classify phylogenetically but can be assigned latively easily to various behavioral, perservational, and environmental categories, Analysis of these aspects of trace fossils, in tum, can yield information that is invaluable in sedimentary geology ‘The most significant contributions of trace fossils have been in paleoccology, sedimen- ‘ology, and environmental reconstruction, including recognition of local and regional-temporal facies changes, patterns of bioturbation, and documentation of individual paleoecological para meters. Trace fossils are potential indicators of bathymetry, currents, food supplies, aeration, Tate of deposition, depositional history, substrate stability, and salinity. ‘They are emerging as useful tools in the developing fields of sequence stratigraphy, allostratigraphy, and event stratig- raphy. TRODUCTION Trace fossils (or ichnotossils) are biologically produced sedimentary structures that include tracks, trails, burrows, borings, fecal pellets and other traces made by organisms (Fig. 1h, Excluded are markings that do not reflect a behavioral function, Owing to their nature, trace fossils can be considered as both paleontological and sedimentological entities, thereby bridging the gap between two of the main subdivisions in sedimentary geology. In fact the contributions to sedimentary geology already are considerable (Table 1), and ‘sumerous additional applications and refinements are currently underway. These contributions can be viewed in terms of traditional fields such as paleontology, stratigraphy,and sedimentol- ogy. Indeed, ichnology - like petroleum geology - is more a blending of related disciplines than a singularly unique cntity. Recent summaries dealing with general ichnological principles can be found in Ekdale et al. (1984), Frey and Pemberton (1984, 1985), Frey and Wheatcrott (1989), Bromley (1990), and Pemberton er al. (in press). This paper is an update of several others done in conjunction with Bob Frey (Frey and Pemberton, 1984, 1985; Frey et al., 1990; Pemberton et al., in press)Table 1. Major contributions of ichnology to sedimentary geology. Relative importance (indicated by the number of +'s) refers to the relative worth of that information in con- temporary sedimentary geology. (Modified from Frey and Pemberton, 1985). Disciplines and Components 1. Paleontology A. Fossil record of sofi-bodied animals B. Patterns of activity by benthic organisms >, Diversity of fossil assemblage D. Evolution of metazoans and of behaviour 2, Stratigraphy A. Recognition of discontinuities B. Delineation of event beds C. Interpretation of transgressive/regressive cycles D. Biostratigraphy of ‘unfossiliferous’ rocks E, Correlation of marker beds F, Structural attitude of beds 3. Sedimentology ‘A. Production of sediment by boring organisms B. Consolidation of sediment by suspension feeders . Alteration of grains by sediment- ingesting animals D. Sediment reworking, 1) Destruction of initial fabries and sedimentary structures 2) Construction of new fabries and sedimentary structures 4. Depositional environments and paleoecology A, Specific adaptations and behaviour of individual genera or species of organisms B. Facies and facies successions C. Bathymetry D, Temperature E. Salinity F, Depositional history: 1) Rates of deposition 2) Amounts of sediment deposited or eroded G. Aeration of water and sediments H, Substrate coherence and stability 5. Consolidation of sediments ‘A. Initial history of lithification B. Measures of compaction Geologic Setting Modern Ancient vBIOGENIC STRUCTURES BIOEROSION OTHER EVIDENCE ‘STRUCTURES oF ACTIVITY BIOGENIC SEDIMENTARY ee “ ae Borings. gnawing Tools, spider webs, scrapings, bitings, & related traces cag cases, & similar structures BIOSTRATIFICATION ‘STRUCTURES BIOTURBATION ‘STRUCTURES, ‘Tracks, trails, burrows, raving patterns, & Jother disruptive structures} BIODEPOSITION ‘STRUCTURES Fecal pellets, fecal castings, pseudoteces, coprolites, & others stromatolites, byssal mats, biogenic graded bedding, & related phenomena Figure 1. Major relationships among biogenic structures. (Modified from Frey and Pemberton, 1985.) THE CONCEPTUAL FRAMEWORK OF ICHNOLOGY The importance of ichnology to the fields to stratigraphy, paleontology, and sedimentol- ogy stems from the following characteristics displayed by trace fossils: (1) long temporal range ~ although a disadvantage in refined biostratigraphy, it greatly facilitates paleoecological com- Parisons of rocks differing in age; (2) narrow facies range - reflects similar responses by trace- making organisms to given sets of paleoecological parameters: (3) no secondary displacement - biogenic sedimentary structures, where preserved intact, are closely related to the environment in which they were formed; (4) occurrence in otherwise unfossiliferous rocks - trace fossils are commonly enhanced by the vary diagenetic processes that typically obliterate tests and shells especially in siliciclastics; and (5) creation by non-preservable soft-bodied biota - many trace fossils are formed by the activities of organisms that generally are not preserved because they lack hard parts; such organisms, in many environments, represent the greatest biomass, Such characteristics are very useful in facies analyses, including reconstruction of indi- vidual palececological factors, sedimentary dynamics, and the documentation of local and regional temporal facies changes.Table 2. Basic concepts in the study of biogenii structures. (modified from Frey, 1971, 1973.) Differentiation of Biogenic Structures 1, Biogenic structure — in ichnology, tangible evidence of activity by an organism, fossil or recent, other than the production of body parts, Embraces the entire spectrum of substrate traces or structures that reflect a bchav- ioural function: biogenic sedimentary structures, bioerosion structures, and other miscellaneous features rep- resenting activity. Excludes moulds of body fossils that result from the passive contact between body parts and the host substrate, but not imprints made by the body parts of active organisms. A. Biogenic sedimentary structure — a biogenic structure produced by the activity of an organism upon or ‘within an unconsolidated particulate substrate: bioturbation structures, biostratification structures, and biodeposition structures. 1) Bioturbation structure — a biogenic sedimentary structure that reflects the disruption of biogeni and physical stratification features or sediment fabrics by the activity of an organism: trucks, burrows, and similar structures. 2) Biostratification structure —a biogenic sedimentary structure consisting of stratification features imparted by the activity of an organism: biogenic graded bedding, byssal mats, certain stroma- tolites, and others. 3) Biodeposition structure — a biogenic sedimentary structure that reflects the production or concen- tration of sediments by the activities of an organism: fecal pellets, pseudofeces, products of bio- erosion, and others, B. Bioerosion structure — a biogenic structure excavated mechanically or biochemically by an organism into a rigid substrate: borings, gnawings, scrapings, bitings, and related traces, Disciplines and Components 2. Ichnology — the over-all study of traces made by organisms, including their description, classification, and inter- pretation. Divisions include paleoichnology for fossils, and neoichnology for recent ones. A. Trace — in ichnology, an individually distinctive biogenic structure, especially one that is related more or less directly to the morphology of the organism that made it: tracks, trails, burrows, borings, coprolites, fecal castings, and similar features, fossil or recent. Excludes biostratification structures and other traces lacking diagnostic anatomical features. B. Lebensspur — synonymous with trace (plural = lebensspuren). C. Iehnocoenose — an association of environmentally related traces: somewhat analogous to a community. Components include the ichnofauna, or animal traces, and the ‘ehnoflora, or plant traces (such as algal borings and certain root patterns). D. Trace fossil — a fossil trace (= ichnofossil) E., Ichnofacies — the fossil record of an ichnocoenose; typically recurs through long intervals of geologic time and is more or less chatacteristic of a given sct of environmental conditions, 3. Ethology — in ichnology, the study or interpretation of the behaviour of organisms as reflected by their traces. 4, Toponomy — in ichnology, the description and classification of lebensspuren with respect to their mode of reservation and occurrence; includes stratinomic (fabrication) and taphonomtc (destruction) aspects, as well as geometry and configuration, 5. Taxonomy — in ichnology, the classification of trace fossils Prevalent taxa include the ichnogenus and ichnospecies. ceording to their systematics and nomenclature,MARTINSSON SEILACHER 1964 1970 FULL RELIEF EXICHNIA EXOGENIC SEMI RELIEF (EPIRELIEF) EPICHNIA FULL RELIEF ENDICHNIA SEMI RELIEF (HYPORELIEF) HYPICHNIA ENDOGENIC FULL RELIEF EXICHNIA CHAMBERLAIN SIMPSON 1971 1957 DIAGENETIC PRESERVATION EPIGENIC BED JUNCTION PRESERVATION CONCEALED BED JUNCTION eee! PRESERVATION INTERGENIC BURIAL PRESERVATION Figure 2. Preservational classification of trace fossils in relation to siliciclastics. Comparison of the terminologies of Simpson (1957), Seilacher (1964b), Martinsson (1970) and Chamberlin (1971). (Modified and enlarged from Bromley, 1990.)Like all disciplines, ichnology has evolved a set of terms needed for the conveyance of its concepts, The more basic ichnological terms and concepts (Fig. 1, Table 2) revolve around behavior by organisms, as opposed to their body paris. Benthic activity, together with its physical manifestation as a trace, is the single most fundamental ingredient of ichnology. Unfortunately, ichnologists are beginning to use the prefix “ichno” ad nauseum, in many inelegant and superflu ous ways. It started innocently enough with ichnofacies, ichnofossil, ichnogenera and ich- nospecies but quickly escalated to include inchnoassemblage, ichnoguild, ichnoassociation, ichn- diversity, ichnostratigraphy, ichnogram, and ichnotaxobase, to name a few. As pointed out by Frey and Pemberton (1991), perhaps ichnologists should be using more restraint, or else at some point only we practitioners will know the meanings of our terms, concepts, and expositions. CLASSIFICATION OF TRACES Unique classification schemes have been developed in order to decipher trace fossils. Historically, the more important classifications have included the phylogeny of the tracemaker and especially the descriptive, preservational, taxonomic and behavioral implication of the traces, Descriptive-genetic Classification Virtually all classifications are genetic to some extent. Even a descriptive classification in ichnology presupposes that the structures were produced biogenically. Numerous descriptive- genetic terms are useful in classifying trace fossils (Table 3). Among the more fundamental terms are burrow walls, burrow fillings, burrow linings, spreiten structures, and burrows versus burrow Preservational Classifications Classifications of the stratigraphic or spatial arrangements and modes of preservation of trace fossils are both descriptive and interpretive. These preservational concepts may be reduced to two basic facets: (1) toponomy, including modes of occurrence and the mechanical-sedimen- tological processes of alteration and preservation, and (2) physiochemical (pre- through post- diagenetic) processes of preservation and alteration. ‘Toponomic (or stratinomic) classification schemes (Table 4, Fig. 2) have been devised by Seilacher (1953, 1964 a, b), Simpson (1957), Martinsson (1965, 1970), and Chamberlain (1971). Most of these schemes generally attempt to relate the position of the trace with respect to the main casting medium which is commonly sandstone or siltstone. Diagenetic preservations have been stressed by numerous authors (e.g. Simpson, 1957; Frey, 1971; Hallam, 1975; Ekdale et al., 1984). The burrowing activities of benthic organisms can result in significant changes in porosity-permeability patterns that will have a significant effect on later diagenesis. Likewise, burrow linings that may be simple secretions of mucus or Particulate walls, agglutinated by organic compounds are preferred sites for subsequent mineral- ization. Therefore, diagenetic processes that obliterate other fossils may even enhance the preservability of trace fossils. 6MARTINSSON SEILACHER 1964 1970 FULL RELIEF EXICHNIA EXOGENIC SEMI RELIEF (EPIRELIEF) EPICHNIA FULL RELIEF ENDICHNIA SEMI RELIEF (HYPORELIEF) HYPICHNIA ENDOGENIC FULL RELIEF EXICHNIA CHAMBERLAIN SIMPSON 1971 1957 DIAGENETIC PRESERVATION EPIGENIC BED JUNCTION PRESERVATION CONCEALED BED JUNCTION eee! PRESERVATION INTERGENIC BURIAL PRESERVATION Figure 2. Preservational classification of trace fossils in relation to siliciclastics. Comparison of the terminologies of Simpson (1957), Seilacher (1964b), Martinsson (1970) and Chamberlin (1971). (Modified and enlarged from Bromley, 1990.)Behavioral Classification Perhaps the single most important ingredient of ichnology is the functional interpretation of individual traces, Fundamental behavioral (or ethological) patterns are dictated and modified not only by genetic preadaptations, but also by prevailing environmental parameters. Ekdale er al. (1984) recognized seven basic categories of behavior (Fig. 3, Table 5); rest- ing traces (cubichnia), locomotion traces (repichnia), dwelling traces (domichnia), grazing traces (pascichnia), feeding burrows (fodinichnia), farming systems (agrichnia), and escape traces (fugichnia). Ekdale (1985) added predation traces (praedichnia), and Frey et al. (1987) emphasized the importance of equilibria (/ugichnia) to all other behavioral patterns (Fig, 4). These fundamental behavioral pattems are genetically controlled, but are not phylogenet- ically restricted, The basic cthological categories have generally persisted throughout the Phanerozoic. Individual tracemakers have evolved, but basic benthic behavior has not. For example, deposit feeders are preadapted to low energy environments where deposited foodstuffs are most abundant; they do not fare well in turbulent water settings, The opposite is true for sus- pension feeders. Similarly, aquatic, invertebrate locomotion traces can be preserved only in fine grained, low energy, quiescent environments such as lagoons. This ability to discern the behav- ioral trends of benthic organisms represented in the rock record greatly facilitates environmental interpretations. i F t FONG TRACE Figure 3. Ethological classification of trace fossils, and relationships of these to body fossils. Overlap of cate- gories acknowledges the intergradations inherent in nature; depending on local circumstance, for example, the spec- {tum of escape structures might span the entire circumference of the circle. (Modified and enlarged from Frey and Pemberton, 1985.)Table 4. Toponomic classification of bioturbation structures. (Modified from Frey and Pemberton, 1985) General Terms " Topenomy te desc arcsec ebasure wh ape thet modo precoated eccutenen gas on Win aSraum of she re casing madum), ar scene terran ole mechanical ong or processes of wosernin el vcore Tale 2 Bioturbaton — reworking of sedans by an arin. Rewatking by macberhcs or misGbarhes seed enplchisurtaton Bouma tere crab) — goss tne inpated to sade by exe eb pealy css rae, cone, hpeneaeg braen ai: tees, eo ncn enc morphology. Where burons ae somoatat lst rom ard we thus mow dst neily re eee a cacr aur mote ‘+ Linoturbation — process of suecestvn ensoses of ioerosen sederaen-cementaton-tcwresic. Ctasiteation by Selacher (1955, 1954, 19640) 1. Doserive Terms ‘A. Semirtet—a roe loss presen ata tha nrisce: bounday wits an clevage vl 1) Boundary relet— a serie not volving csavage preservation: mypote' arog 2) yore? — a boundary ral oxcurg anna eae ola staum: eft may Se concave or conven ©) Epielot— a boundary reel occuring onthe ap stature! may be concave or cone 2 Cleavage rte — a serie n whch sdusac line ae sor utr predun srclMae pag ot nase amine ls wo or sity reese vrcalepttan oa given abesspus, ry eased specimen ef whch osentes tenga toontny oes 8. Fulrolet— a secur presrved wih san 2 Genetic Tome ‘A Exogene — a sail stucturs covered by some eifenng ta tht fe host ube 8. Endogene —® vce ed actvely or patel wii re host sadnen © ntergene — an oréopenc seca pocced att bed uncon Classitcation by Simpson (157) 2 ealagmon preservation — aoe sts spear in rl at bed archon of conact. ego sy ccd ath ineae betwen bs of dito ihgy 2 Gonceued be function preservation ~ indi masses of bowled seiner ops bok mn bed ol Stren Melony. a roe nee are sandstone; no obvious cernecton win an overing sale bed aia because t was raved by wosen * Dlgenati preservation — acs ois reser as pacomparona roles neue prokwances Presa ted Ging ea apes: ore fu wer sae ‘alg ye rece perosy a he ed burows eave fat of be mar, Fequety coer bd jnciona a sears a ately siaceny soe ‘Sores ard shies. ‘Burl preservaion — sed bites subsautnysxnumed by sirens wowing ava he st het matin Coherent uo ge yon he sent sce and, where preserved, wre Bure auch by ater sade, Clasticatin by Martnsson (1985, 1970) 1 Eletnon — a stucre preserved athe wpe sutaoo he man hoy he using med may cpa a ge HE 2. pn ge 2 Hypihnion~ a stuctre preserved a elon sutane ft man acy fhe eetirg meu may acpew ts arabe ogiare ee, tenia aoe 3 Endlehnion ~ a stctr preserves whine mar body othe casing radon, 4 Exlchnion — a starve preserved ousiae he main bog fhe casing mau 69, edie ppd no Unda bed Ciassifeation by Chambetain (19713) 1. Eplgene —a suc erucure cove by sane dieing tom hat lhe hot satu. 2. etergene —anerdopenic shure seduced a alie-lope neta ob junc 8. Endogone ~ a sci ad actly or assay within he host seenon “Oa ao ener NG le / Le NL igure 4. Behavioral classification of lebenssparen. With environmental fluctuation, virtually all traces are {nlergradational with the fugichnia - escape or equilibrium structures. Bromley (1990) proposed a new ethologie category, the equilibrichnia to encompass such cases, (Modified trom Frey et al_, 1987.)Table 5. Ethological classification of invertabrate trace fossils. (Adapted from Frey and Pemberton, 1985.) Definition Characteristic Morphology Examples Resting traces (Cubichnia) ‘Shallow depressions made by animals that sete onto ‘Trovgh-tke rele, recording o some extent the laeroventral —_Asteracites: ‘or aig int the substrate surface, Emphasis ison eclusion. morphology ofthe animal: ideally, structures aro isolated, ‘Lockeia ‘May include shallow, ephemeral domicile. burmay inorgrade wits crawling races or escape stucures. Ausophycus Crawling traces (Repichnia) ‘Trackways and epistatal or nrastatal trails made by Linear or sinuous overall stuctures, some branched: foot Auilchnites ‘organisms traveling from one place to another. ‘bons or conbnuous grooves, commonly annulated: complete Cruzlana Emphasis is upon locomotion. Secondary activites may form may be preserved, or may appear as cloavage reels. _pchnites be involved ‘Soolia Grazing traces (Pascichnia) Grooves, paterned pis, and furrows, many of them Unbranched, nonoverlapping, curved to tight coiled pattems Helminthoida ‘discontinuous, made by mobile deposit loaders or algal ‘or delcalely conetucod spreten dominate. patterns gonoraly — Lophectenum ‘grazers a or under the substrate subsurface. Emphasis fetlect maximum ulzaton of fod resources; complete form Nerotes IS upon feeding behaviour analogous to stp ming tay be proservod. Overall stuctue fonds tobe planar Spirophycus Feeding traces (Fodinichna) More o less temporary butrows constructed by deposit Singlo, branched, or unbranched, cyincrical to sinuous shafts Chondites feeders; he structures also may provide shelter for the '5¢ Usshaped burrows, or complex, parallel o concentric burow — Gyrophyiites ‘organisms, Emphasis is on feeding behaviour analogous repetvons (spreiten sutures); walls not commonly ined, —-Phycodos to underground mining’. Some tend tobe gradational Unlass by mucus. Oriented at varous angs wih espectto Rossel, with dweling structures. ‘bedoing: complete focm may be preserved. Dwelling structures (Domichnia) Burows, borings, or dweling tubes providing more cress ___ Simple, biureated or U-shaped structures perpendicular Dipioerateron permanent domictes, mostly or the hemisessie suspension or iclned at various angles to bedding, or branched Opniomorpha feeders or, in some cases, camivores. Emphasis is on burrow or boring systems having vertical and horizontal ‘Skoltnos habitation’ Secondary achwties may be dscemible. Components: buttow walls typically lined. Complete form Trypanies may be preserved. Escape structures (Fugit) puren of various kins modified or made anew by ically repetitive resting traces; biogenic laminae ether nested funnels animals nest oopens esubtate degredaton ot {en echelon ot as nested funnels or chev, U-in-U Usd spree ‘aggradation. Emphasis s upon eagusimont or equiibium _sproon Burrows, and other structures relecting displacement down-warped between relative substrate postion and the configuration Of animals upward or downward wih respec tothe laminae. fof contained traces. Intergradatonal wih other behavioural orginal substvate surface. Complete from mat be preserved, calogoris. ‘especialy in aggraded substrates. Farming structures (Agrchnia) Fgulary patted burrow systems in which the activites Horizontal tunnels organized in compl, regular, geometic Belorhaphe of permanent dweling and feeding are combined. Emphasis pallens such as meanders, spirals, and hexagonal meshworks. Paleodicyon Is on both habitation and feeding using farming’ or Complete forms may bo presorved ‘Spromapne “rapping strategies. Predation traces (Prascichnia) Various types of traces resulting from obvious predation ‘Simple, circular holes drited in shells by carnivorous Contichous ‘Typically hese are Boerosion structures prodiced on nard gastropods or cephalopods; be marks on ammonites made Oichnus. biological materials, such as shel bone, or coral, Gnawings {and other bite traces are included in tis group. Emphasis ‘by aquatic repties; and dsincive patterns of chipped margins of gasvopod and bivae sells that have been ‘on feeding by predation and predator-prey relationships are aitacked by camivorous crabs. Stuctures tend o be solated iscemibe THE ICHNOFACIES CONCEPT Perhaps the essence of trace fossil research involves the grouping of characteristics ich- nofossils into recurring ichnofacies. The concept, developed by Adolph Seilacher in the nine- teen-fifties and ninteen-sixties, was based originally on the fact that many of the parameters that control the distribution of tracemakers change progressively with increased water depth. Nine recurring ichnofacies have been recognized, cach named for a representative ichnogenus (Fig. 5): Scoyenia, Trypanites, Teredolites, Glossifungites, Psilonichnus, Skolithos, Cruziana, Zoophycos, and Nereites. These ichnofacies reflect adaptations of tracemaking organisms to environmental factors such as substrate consistency, food supply, hydrodynamic energy, salinity, water turbidity, sedimentation rate, temperature, and oxygen levels among others (Frey and Pemberton, 1984; Frey ef al., 1990; Bromley and Asgaard, 1991). 10seyesen |soofydooz| euejznig [snuyaruoyisa|sey6unyssor5| BOOT Say | BZ TERE SIE RTS IOTSOEE TOTES “ee lFe. [eo eee e* BB AES rd Cg SaTSS ou t | 02 18S, %e, 9u0Z fesonnang eaysyoea ‘pues ‘arensang,Woodground | Hardground | Firmground | Softground Scoyenia Freshwater Psilonichnus Teredolites Trypanites Glossitungites Skolithos High Energy Cruziana Medium Energy ¢ Marine Zoophycos Nereites Low Energy Figure 6, Relationship of ichnofacies to substrate, as opposed to softground ichnofacies which have been differenti- ated on environmental factors, hardground, firmground, and woodground ichnofacies arc differentiated from one another by substrate type and consistency. (Modified from Bromley et al., 1984.) The non-marine assemblage (Scoyenia) is general and in need of revision; the marine softground ichnofacies (Psilonichnus, Skolithos, Cruziana, Zoophycos, and Nereites) are distributed accord- ing to numerous environmental parameters; and the traces in firmground (Glossifungites), wood- ground (Teredolites) and hardground (Trypanites) ichnofacies are distributed on the basis of sub- strate type and consistency (Fig. 6). Representative occurrences of the various ichnofacies are summarized below. However, each may appear in other settings, as dictated by characteristic sets of recurrent environmental parameters. Scoyenia Ichnofacies At present, the only named ichnofacies for nonmarine environments is the Scoyenia ich- nofacies (Fig. 7). Frey e¢ al. (1984) concluded that the Scoyenia ichnofacies remains a valid concept within appropriate limits and is suggested of deposition in the shore of ephemeral lakes and the overbank of sluggish rivers. The Scoyenia assemblage is characterized by: (1) small horizontal, lined, barkfilled feeding burrows; (2) curved to tortuous feeding burrows; (3) sinuous crawling traces; (4) vertical, unlined, cylindrical to irregular shafts; and (5) tracks and trails. Prospects for the recognition of additional archetypal nonmarine ichnofacies remain encouraging. For example, Ekdale et al. (1984) and Frey and Pemberton (1987) noted that dis- tinct suites of trace fossils characterize aeolian dunes, paleosols, and lakes. The ichnofos found in these environments were recently summarized by Ekdale et al. (1984), Retallack (1984), Frey et al. (1984), Miller (1984), Pollard (1988), Gray (1988), Maples and Archer (1989), Bromley (1990), and Bromley and Asgaard (1991).1. Scoyenia 2. Ancorichnus 3. Cruziana 4. Skolithos . Footprints ‘Scoyenia Ichnofacies Figure 7. Trace fossil association considered to be indicative of the Scoyenia ichnofacies; as emended by Frey etal. (1984), (Modified from Frey and Pemberton, 1984.) Psilonichnus Ichnofacies The Psilonichnus ichnofacies (Fig. 8) represents a mixture of marine, quasi-marine, and nonmarine conditions, Typical environments include the backshore, dunes, washover fans, and supratidal flats. Frey and Pemberton (1987) noted that such environments are subject to extreme variations in energy levels, sediment types, and physical and biogenic sedimentary structures. The environments may also be strongly affected by torrential rains and storm surges. Marine Processes generally dominate during spring tides and storms, whereas aeolian processes predom- inate during neap tides and nonstorm periods. The Psilonichnus assemblage is characterized by: (1) vertical shafts ranging from small structures, some with bulbous based cells, or larger, irregularly J- & or U-shaped dwelling struc- tures; (2) invertebrate and vertebrate crawling and foraging traces; (3) vertebrate tracks and coprolites; (4) low density and diversity; (5) invertebrates, mostly predators or scavangers; and (6) vertebrates, mostly predators or herbivores. Skolithos Ichnofacies The Skolithos ichnofacies (Fig. 9) is indicative of relatively high levels of wave or cur- rent energy, and typically is developed in slightly muddy to clean, well-sorted, loose or shifting 131. Psilonichnus 2. Macanopsis 3. Vertebrate Track Not To Scale Psilonichnus Ichnotacies Figure 8. Trace fossil association characteristic of the Psilonichnus ichnofacies. (Modified form Pemberton ef al., 1990.) particulate substrates. Increasing energy levels enhance physical reworking, thus obliterating the biogenic structures and preserving physical sedimentary structures. Such conditions commonly occur on the foreshore and shoreface of beaches, bars and spits, but similar conditions some- times occur on tidal deltas and submarine fans. Due to the fundmental relationships between water agitation, sediment transport and animal distribution, most tracemakers found in these set- tings are suspension feeders. ‘The organisms typically construct deeply penetrating, more or less permanent domiciles. The Skolithos ichnofacies is characterized by: (1) predominantly vertical, cylindrical, or U-shaped burrows; (2) protrusive and retrusive spreiten in some U-burrows, which develop in response to substrate aggradation or degradation; (3) few horizontal structures; (4) few structures produced by mobile organisms; (5) low diversity, although individual forms may be abundant; (6) mostly dwelling burrows constructed by suspension feeders or passive carnivores; and (7) vertebrate traces, particularly in low energy intertidal settings. Cruziana Iehnofacies The Cruziana ichnofacies (Fig. 10) is most characteristic of subtidal, poorly-sorted and unconsolidated substrates. Conditions typically range from moderate energy levels in shallow 141. Ophiomorpha 2. Diplocraterion 3. Arenicolites 4. Skolithos Not To Scale ‘Skolithos Iehnofacies Figure 9. Trace fossil association characteristic of the Skolithos ichnofacies. (Modified from Frey and Pemberton, 1984.) waters below fair weather wave base but above storm wave base, to low energy levels in deeper, quieter waters. Sediment deposition rates range from negligible to appreciable, but not normally rapid. With reduced (but no negligible) energy levels, food supplies consist of both suspended and deposited components. Either fraction may predominate locally, or the two may be inter- mixed. Characteristic organisms include both suspension and deposit feeders, as well as mobile carnivores and scavengers. Trails of epibenthic and endobenthic foragers also may be common and reflect the abundance, diversity, and accessability of food. The Cruziana ichnofacies is characterized by: (1) mixed association of vertical, inclined, and horizontal structures; (2) the presence of structures constructed by mobile organisms; (3) generally high diversity and abundance: and (4) mostly feeding and grazing structures construct- ed by deposit feeders. Zoophycos Ichnofacies Of all the marine ichnofacies, this assemblage is the most debated and least understood The ichnogenus Zoophycos has an extremely broad paleobathymetric range, hence its designa- tion as namebearer for a supposedly depth-related ichnofacies has long been controversial (Miller, 1991). In popular bathymetric schemes, the Zoophycos ichnofacies (Fig. 5) typically is 154. Aulichnites 5. Thalassinoides 6. Chondrites 7. Teichichnus 8. Asterosoma 9. Rosselia 10. Planolites ‘Not To Scale Cruziana Ichnotacies Figure 10, Trace fossil association characteristic of the Cruziana ichnofacies. (Modified from Frey and Pemberton, 1984.) Portrayed as an intermediary between the Cruziana and Nereites ichnofacies (Fig. 11). More specifically, the original designation placed it in areas below storm wave base but free of turbidi- ty currents, within the broad depositional gradient that defines the outer shelf to slope transition (Scilacher, 1967). Frey and Seilacher (1980) considered one of the major environmental controls represent- ed by the Zoophycos ichnofacies to be lowered oxygen levels associated with abundant organic material in quiet-water settings. ‘To some extent, those conditions do occur in the broad area across the shelf-slope break, and hence the popularized bathymetric placement of the ichnofacies is warranted. However, such reducing conditions replete with a dominance of Zoophycos, per- haps even better known in shallower water, epeiric deposits (Marinisch and Finks, 1982) espe- cially in the Paleozoic (Bottjer et al. 1987; Miller, 1991). ‘The Zoophycos ichnofacies is characterized by: (1) low diversity, although individual traces may be abundant; (2) grazing and feeding structures produced by deposit feeders; and (3) horizontal to gently inclined spreiten structures. Nereites Ichnofacies The Nereites ichnofacies (Fig. 12) occurs in bathyal to abyssal quiet but oxygenated water, commonly influenced by turbidity currents. Animals exploiting such environments have two major concerns: (1) scarcity of food, relative to more abundant supplies in shallower set- tings, and (2) periodic disruption by strong down-canyon bottom currents or actual sediment- 161. Phycosiphon 2. Zoophycos 3. Spirophyton Not To Scale Zoophycos Ichnotacies Figure 11. Trace fossil association characteristic of the Zoophycos ichnofacies. (Modified from Frey and Pemberton, 1984,) gravity flows. Over geologic time, the overall community developed two component parts: pre- turbidite and post-turbidite assemblages. The pre-turbidite resident association is characteristic of quiet, normal conditions and is dominant wherever the substrate is frec of the influence of tur- bidity currents. It tends to be overwhelmed or eliminated by severe erosion or turbulence, how- ever, and is replaced by the post-turbidite association immediately following a turbidity current. Later, as conditions revert to a normal, low-energy setting, the pre-turbidite association gradual. ly reestablishes itself. Pre-turbidite animals thus comprise a stable, persistent community well adapted to quiet conditions, derived mainly from original early-Paleozoic colonizers of the deep- sea floor. In contrast, post-turbidite animals represent a more opportunistic, less stable communi ty better adapted to turbidite colonization, derived mainly from subsequent evolutionary immi- grants from shallower waters (Frey and Seilacher, 1980), The Nereites ichnofacies is characterized by: (1) high diversity but low abundance; (2) complex horizontal grazing and patterned feeding-dwelling structures (agrichnia); (3) numerous crawling-grazing traces and fecal castings; and (4) structure produced by deposit feeders, engers, or harvesters. av- Substrate-Controlled Ichnofacies The remaining three ichnofacies are specialized, substrate-controlled, and environmental ly, non diagnostic in scope. The Glossifungites ichnofacies is environmentally wide-ranging, but only develops in firm, unlithified substrates such as dewatered muds. Compaction dewatering results from burial and the substrates are made available to trace-makers if exhumed by later crosion (Pemberton and Frey, 1985), Exhumation can occur in terrestrial environments as a result of channel mean- dering or valley incision, in shallow-water environments as a result of meandering tidal chan- nels, coastal erosion, erosive shoreface retreat, or as a result of submarine channels cutting 171. Spirorhaphe 2, Urohelminthoida 3. Lorenzinia 4. Megagrapton 5. Paleodictyon 6. Nereites 7. Cosmorhaphe Not To Seale Nereites Ichnofacies Figure 12. Trace fossil association characteristic of the Nereites ichnofacies, (Modified from Prey and Pemberton, 1984.) through previously deposited sediments (Hayward, 1976; Fiirsich and Mayr, 1981; Pemberton and Frey, 1985), Such horizons commonly form at bounding discontinuities and may be critical in the evolving concept of sequence stratigraphy (Savrada, 1991; Pemberton et al., in press: The Glossifungites ichnofacies (Fig. 13) is characterized by: (1) vertical, cylindrical U-or tear-shaped pseudo-borings, sparsely to densely branching dwelling burrows, and/or mixtures of burrows and pseudo-borings; (2) protrusive spreiten in some burrows that develop mostly through animal growth (funnel-shaped Rhizocorallium and Diplocraterion {formerly Glossifungites}); (3) animals that leave the burrow to feed (c.g., crabs), as well as suspension feeders; and (4) low diversity, but commonly abundant individual structures. The Trypanites ichnofacies (Fig. 14) develops in fully lithified substrates such as hard- grounds, reefs, rocky coasts, beachrock and other omission surfaces. As such, development of this ichnofacies also corresponds to discontinuities that have major sequence stratigraphic signif- icance (Pemberton, ef al., 1980). The traces are characterized by: (1) cylindrical to vase-, tear- or U-shaped to irregular domiciles of suspension feeders or passive carnivores; (2) raspings and gnawings of algal graz- ers and similar organisms (mainly chitons, limpets, and echinoids); (3) moderately low diversity, although the borings and scrapings of individual ichnogenera may be abundant; and (4) borings oriented perpendicular to the substrate which may include large numbers of overhangs. In con- trast to the Glossifungites ichnofacies, the walls of the borings cut through hard parts of the sub- strata rather than skirting around them. 181. Echinold Grooves 2 Rogerella 3. Entobia 4. Trypanites 5. Gastrochaenoliies 6. Polychaete Boring Not To Scale Teypanites Ichnotacies Figure 14, Trace fossil association characteristic of the Pemberton, 1984.) ‘panites ichnofacies. (Modified from Frey and ") 7 EF ESS G q 4 aa 1. Teredolites Not To Ses Teredolites ichnotacies Figure 15. Trace fossil association characteristic of the Teredolites ichnofacies, (Modified from Bromley et al., 1984.) 19upon recurring ichnocoenoses (Seilacher, 1967, 1978; Frey and Pemberton, 1984, 1985). Ifa particular ichnocoenose tends to occur repeatedly within a given bathymetric setting, so much the better, but water depth per se is rarely, if ever, a governing factor. Ichnofacies, therefore, are best viewed in the context of actual deposition, One of the most fundamental tenets of modern ichnofacies analysis is that all available evidence - physical, chemical, or biological - should be integrated and utilized in interpretations. For bathymetric assessments, those collective observations, at the very least, should be placed in the context of proximality trends, whether emphasized from a sedimentological viewpoint (Nittrouer ef al., 1984; Clifton, 1988) or an ichnological one (Crimes, 1973; Wetzel, 1981: Howard and Frey, 1984). Associations between, and configurations of, biogenic and physiogenic sedimentary structures are powerful combinations in the reconstruction of environmental gradi- ents (Wightman er al., 1987; Moslow and Pemberton, 1988); they are especially useful where otherwise prevalent trends have been modified by episodic events or other environmental fluctu- ations (Aigner and Reineck, 1982; Pemberton and Frey, 1984: Easthouse and Driese, 1988; Frey, 1990; and many others). Numerous authors have noted occurrences of certain ichnofacies in settings outside the zone specified in the original paradigm, and have used these discrepancies as an argument against the validity or usefulness of the overall ichnofacies concept. For instance, if each shelf sequence involved only a “normal” beach-to-offshore trend then the classic onshore Skolithos ichnofacies would indeed give way to the offshore Cruziana ichnofacies, virtually without exception. But in many settings, the nearshore zone includes bays, lagoons, estuaries, deltas and tidal flats, and the offshore zone includes bars, shoals, submarine canyons, or ridges or other fea~ tures that might disrupt the “normal trend”. For similar reasons, the Skolithos ichnofacies may appear on proximal parts of deep-sea fans (Crimes, 197; Crimes er al.,1981) or the Zoophycos ichnofacies may appear in silled marine basins or restricted lagoons (Miller, 1991). In short, the idealized ichnofacies succession works well in “normal” situations (Frey and Pemberton, 1984); yet one should not be surprised to find nearshore assemblages in offshore sediments, and vice-versa, if these accumulated under conditions otherwise like those preferred by the trace-making organisms (Frey et al., 1990). The basic consideration rests not with such inanimate backdrops as water depth or distance from shore, or some particular tectonic or phys- iographic setting, but rather with such innate, dynamic controlling factors as substrate consisten- cy, hydraulic energy, rates of deposition, turbidity, salinity, oxygen levels, toxic substances, the quality and quantity of available food, and the ecologic or ichnologic prowess of the tracemakers themselves. Resulting ichnocoenoses are related to bathymetry only where particular combina- tions of environmental parameters are aligned with bathymetry. * QUANTIFICATION OF BIOTURBATION Because of the extreme number of potential variables involved, no one has yet devised a satisfactory classification scheme for burrow-mottled sediment, bioturbate textures, or bioturba- tion processes (Frey and Pemberton, 1985). Most sedimentologic studies have relied upon the subjective process of the visual estimation of the percentages of biogenically reworked or unre- worked sediment (see Reineck, 1963, 1967). This scheme (Table 6) is highly useful from a gen- 20eral descriptive standpoint, but sheds no light on any genetic relationships (e.g. Ekdale and Bromley, 1983) Recently, in a series of papers Drosser and Bottjer (1986, 1989, 1991) have taken this Concept and presented it in visual form (Fig, 16). This scheme is based on the amount of origi- nal physical sedimentary structures disrupted by biogenic reworking from no bioburbation (index 1) to complete homogenization (index 6). ‘The six categories defined by Drosser and Bottjer (1986) are: (1) no bioturbation recorded, all original sedimentary structures preserved; 2) discrete, isolated trace fossils, up to 10% of original bedding disturbed; (3) approximately 10 to 40% of original bedding disturbed, burrows are generally isolated, but locally overlap: (4) last vestiges of bedding discernable; approximately 40 to 60% disturbed, burrows overlap and are Not always well defined; (5) bedding is completely disturbed, but burrows are still discrete in Places and the fabric is not mixed; and (6) bedding is nearly or totally homogenized. Although it is valuable to note the degree of bioturbation, it is just a descriptor and as such, has no genetic value. One near-fatal flaw in trying to characterize the changes in rates or intensities of bioturbation on a temporal basis (as this scheme attempts) is that local environmen- tal controls on bioturbation are apt to exert too much control and impart too much local variation, As documented in detail on Korean tidal flats, the intensity of bioturbation changes abruptly on both large and small scales, both vertically and laterally in the sequence (Frey eal. 1989). PALEOENVIRONMENTAL SIGNIFICANCE OF TRACE FOSSILS The concept of functional morphology, a basic premise employed by ecologists and pale- ecologists in environmental reconstructions, is equally applicable to ichnology. In fact, trace fossils are unique in that they represent not only the morphology and ethology of the tracemak- ing organism but also the physical characteristics of the substrate; they are closely linked to the environmental conditions prevailing at the time of their construction. Frey and Scilacher (1980) re-emphasized that such variables as bathymetry (Fig. 5), temperature (Fig. 17), volumes of sed- iment deposited or croded (Fig. 18), aeration of water and sediment (Fig, 19), and substrate coherence and stability, have a profound effect on resultant trace fossil distributions and mor- Phology, and hence, can be used in the determination of original biological, ethnological, and sedimentological conditions. The application of ichnology to paleoenvironmental analysis goes far beyond the mere establishment of gross or archetypal ichnofacies, For instance, shallow-water, coastal marine environments comprise a multitude of sedimentological regimes, which are subject to large fluc- tuations in many physical and ecological parameters. In order to fully comprehend the deposi- tional history of such zones in the rock record, it is imperative to have some reliable means of differentiating subtle changes in these parameters. Detailed investigations of many of these Coastal marine zongs in Georgia have shown the value of utilizing biogenic sedimentary struc tures (in concert with physical sedimentary structures) in delineating them (Frey and Pemberton, 1987). The application of these studies in deciphering palcoenvironments has also proven invaluable. The use of trace fossils in the interpretation of freshwater deposits is becoming incteas- ingly important, Recent work by Pollard (1988), Maples and Archer (1989), and Bromley and Asgaard (1991), among others, have stressed the abundance and diversity of tracemaking organ- 21Table 6. Classification of bioturbation, based on destruction of primary sedimentary structures. (Adapted from Reineck, 1967.) Classification Unbioturbated Very slightly bioturbated Slightly bioturbated Moderately bioturbated Highly bioturbated Intensely bioturbated (vestiges of some physical structures still discernable Completely bioturbated A B c D Figure 16.Schematic diagrams of estimates on degree of bioturbation (ichnofabric index). (A) for thin bedded stra ta, (B) for thick bedded strata dominated by Skolithos. (C) for thick bedded strata dominated by Ophiomorpha. (D) for fine-grained deepwater environments, (Modified from Droser and Bottjer, 1986, 1989, 1991.) 22isms in freshwater environments and emphasized their potential importance in paleoenvironmen- tal reconstructions. Distinct differences in trace fossil types and abundance have been reported from a wide range of freshwater-terrestrial environments, in both ancient and recent settings (Ekdale et al., 1984). Recently, marginal marine environments (including tidal channels, estuaries, bays, shal- low lagoons, delta plains, etc.) have been recognized with more frequency in the rock record. Such environments characteristically display steep salinity gradients, which, when combined with corresponding changes in temperature, turbulence, exposure, and oxygen levels, result ina Physiologically stressful environment for numerous groups of organisms. The typical trace fos- sil suite in such environments reflects these stresses and is characterized by: (1) low diversity; (2) ichnotaxa which represent an impoverished marine assemblage rather than a true mixture of marine and freshwater forms; (3) a dominance of morphologically simple structures constructed by trophic generalists; (4) a mixture of elements which are common to both the Skolithos and Cruziana ichnofacies; (5) assemblages that are commonly dominated by a single ichnogenus; and (6) diminished size compared to fully marine counterparts (Wightman et al., 1987; Beynon et al., 1988). One of ichnology’s greatest strengths, the bridging of sedimentology and paleontology, in some respects, can be its greatest liability. Sedimentologists tend to use a strict uniformitarian approach to paleoenvironmental interpretation and rely heavily on modern analogues. Paleontologists, on the other hand, must temper their observations in the light of organic evolu- tion. Although trace fossils can be considered as biogenic sedimentary structures and are diffi- Cult to classify phylogenetically, they are constructed by biological entities and are thus subject- ed at least to some degree to evolutionary trends. For example, occurrences of well developed terrestrial trace fossil assemblages are much more prevalent in post-Cretaceous rocks. This development corresponds to the evolutionary explosion of the insects, brought on by the diversi- fication of the angiosperms in the Late Cretaccous. Prior to this time terrestrial substrates may not have been as extensively bioturbated due to a paucity of tracemakers. Likewise, patterned grazing traces, which characterize deep-sea environments, show a trend toward more complex organization through most of the Phanerozoic. This trend may be related to the evolution of more efficient foraging strategies (Seilacher, 1986). For these reasons, paleocnvironmental interpretations based on trace fossils must be considered not in strict uniformitarian terms, but rather, in actualistic ones. Equally important, unique quantitative environmental indicators are indeed rare in the Yeological record, and ichnology is no exception (Frey and Seilacher, 1980). However, trace fossils can supply a wealth of environmental information that cannot be obtained in any other way and which should not be ignored. Their potential usefulness is accentuated when fully inte- grated with other (chemical, physical, and biological) lines of evidence. Combined studies of Physical and biogenic sedimentary structures constitute a powerful approach to facies analysis SUMMARY ‘The conceptual framework of ichnology and its significance to sedimentary geology is based on the following: 23a Temperature (°C) 6 9 2 (16 18 Hour Figure 17, Temperature variation within an intertidal sand flat during a tidal cycle, Tomales Bay, California. Deep burrowing species experience less thermal variation than near surface organisms. ‘The substra stabilize the thermal environment. (Modified from Johnson, 1965.) acts as a buffer to Figure 18. Bioturbation.patterns in different depositional settings. (A) Slow, continuous deposition, where rate of bioturbation equals or exceeds that of deposition. (B) Rapid deposition (or deposition under anoxic conditions), where animal activity is precluded; bioturbation reaches down from a depositional break at the top of the unit, (C) Alternating rapid and slow deposition with erosion; note truncation of burrows at the erosion surface; (D) Slow, continuous deposition without erosion; the nearly invisible omission surfaces are emphasized by an omission suite of trace fossils produced by animals during phases of non-deposition; depositional phases favor a different animal community and different behavior patterns. (E) Rhythmic deposition interspersed with erosion; note truncated bur- rows. (Modified from Howard, 1978 24A. TYPE2 TYPES ‘Small Thalassinoides ‘or Large Chonadrties Planoutes /Taentdtum (Large A. ethest?) nieso MOuUTE Well- Oxygenatod Anasrobic Planoittes | Thalasstnotdes Thalassinoides | _Zeophycos Zoophycos Burrow Size Depth in Sediment Pore Water Oxygenation Figure 19. Schematic diagrams summarizing details of trace fossil associations in oxygen-depleted environments, (A) Diagram iltastrating bureow size bioturbation depth, and tering across @ hypothetical seafloor oxygenation 2 dient (modified from Savrda and Bottjer, 1987).(B) Trace fossil assemblages in the Kimmeridge Clay. distributed according to an oxygen gradient. With increasing oxygen there is a general inervase in burrow depth and destruc Hon of primary fine lamination (modified from Wignall. 1991). (C) Oxygen-related trace fossil ti established from various Mesozoic marine strata based on cross-cutting rek 2 relationships lationships. Larger burrow types are pro: gressively climinated and deeper tiers migrate upward as the level of bottom-water oxygenation decreases (modified from Bromley and Ekdale, 1984).A) Biogenic structures represent the activity of soft-bodied organisms that are not gener- ally preserved. Such organisms (including many entire phyla) are commonly the dominant com- ponent of the biomass of many environments. B) Biogenic structures are commonly enhanced by diagenesis and can be used in hori- zons where physical sedimentary structures have been masked, For example, in oil sands deposits bitumen staining obliterates most physical structures, but due to the concentration of clay minerals, it enhances the visibility of biogenic structures. ©) Biogenic structures can be associated with facies that do not contain any other fossils. In many siliciclastic regimes, diagenesis dissolves most of the shelly fauna and trace fossils rep- resent the only clue as to the original biogenic component of the unit. D) Biogenic structures can be used for the paleoecological reconstruction of depositional environments. E) Biogenic structures are sensitive to fluctuations in sedimentary dynamics. Bioturbation patterns are important in recognizing event beds and sedimentation patterns. F) Biogenic structures are sensitive indicators of substrate coherence. The Trypanites, Glossifungites, and Teredolites ichnofacies are substrate-controlled, and are emerging as impor- tant elements in the evolving concepts of sequence stratigraphy. G) Biogenic structures ordinarily cannot be transported and therefore represent the origi- nal environmental position of the trace making animal. H) Biogenic structures are sensitive to changes in certain ecological parameters that are otherwise difficult to ascertain. Trace fossil models are emerging to recognize changes in such parameters as salinity and oxygen levels. }) Subtle changes in the distribution of biogenic structures can in fact be mapped. J An integrated approach utilizing physical, chemical, biological, and ichnological lines of evidence constitutes a powerful tool for facies interpretation. ACKNOWLEDGEMENTS Funding was provided by a grant from the Natural Science and Engineering Research Council of Canada (No. A0816) to Pemberton, We arc grateful to the drafting department at Exxon Production Research for the figures and to Joanne Wells for typing the manuscript. This Paper was completed during tenure as a visiting scientist at Exxon Production Research by Pemberton who is grateful to John Van Wagoner, Kirt Campion, and Bob Todd for their gener- ous hospitality and assistence. 26REFERENCES Aigner, T. and H. E, Reineck, 1982. Proximiality trends in modern storm sands from the Helgoland Bight (North Sea) and their implications for basin analysis. Senckenbergiana Maritima, 14: 183-215. Beynon, B. M., and S. G. Pemberton, D. A. Bell and C. A. 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