plied Ichnology ort Course A S The Use of Trace Fossils in Sequence Stratigraphy, Exploration and Productioi asAPPLIED ICHNOLOGY: THE USE OF TRACE FOSSILS IN SEQUENCE STRATIGRAPHY, EXPLORATION AND PRODUCTION GEOLOGY SHORT COURSE NOTES SEPM SHORT COURSE 18 HOUSTON, 2006 S. George Pemberton James A. MacEachern* Ichnology Research Group, Department of Earth & Atmospheric Sciences, University of Alberta, Edmonton, Alberta, T6G 2E3 Department of Earth Sciences, Simon Fraser University, Burnaby, British Columbia, VSA 186 Reproduction of Notes Courtesy of ConocoPhillips‘TABLE OF CONTENTS. General Principles of Ichnology ‘The Conceptual Framework of Ichnology Classification of Traces Descriptive ~Genetic Classification Preservational Classification Behavioural Classification Phylogenetic Classification The Ichnofacies Concept ‘Archetypal Ichnofacies Non-marine Ichnofacies Marine Sott Ground Ichnofacies Substrate Controlled Ichnofacies Evaluation of Models Paleoenvvironmental Significance of Trace Fossils ‘Summary Ichnology of Marine Clastie Environment ‘The Basic Softground Ichnofacies, Psilonichnus Ichnofacies Skolithos Ichnofacies Cruziana Ichnofacies Zoophycos Ichnofacies Nereites Ichnofacies ‘An Integrated Ichnological-Sedimentological Model of Shoreface Successions and Shoreface Variability Introduction Shoreface Subenvironments, Offshore Complex Lower Middle Shoreface Complex Upper Shoreface-Foreshore Complex Shoteface Variability Summary ‘The Ichnological Expression of Storm Deposits Episodic Depositional Events Turbidites ‘Tempestites Ichnology of Storm Deposits Preservational Potential ‘Transit Time u ul 13 16 20 21 21 24 2 30 30 32 4 38 38 38 4 4s 45 49, 38 35 7 n 82 85 87 87 87 89 95Summary ‘The Application of Ichnofacies Analysis to the Evaluation of Marginal Marine Reservoirs Biological and Ichnological Implications of Brackish Water Biological Response Ichnological Implications ‘Typical Brackish Water Ichnofossil Assemblages Brackish Water Channel Assemblages Brackish Water Bay-Like Environments, Conclusions Cryptobioturbation and the Work of Sedimentologically Friendly Organisms ‘Signs of Awaited Truce on the Physical Biogenic Battleground ‘What Is and What Isn’t Cryptobioturbation? ‘Taking it to the Fooail Record Known Tracemakers of Cryptobioturbation Final Comments on Cryptobioturbation Significance of Trace Fossils to Genetic Stratigraphy ‘Substrate ~Controlled Ichnofacies Glossifungites Ichnofacies Ichnological Applications to Genetic Stratigraphy ‘Sequence Boundaries Transgressive Surfaces Amalgamated Sequence Boundaries and Flooding Surfaces ‘The Glassifungites Ichnofacies and Porosity/Permeat Conclusions Paleoenvironmental Significance of Trace Fossils Ichnological Recognition of Non-marine Environments ‘Subaerial (Terrestrial) Environments Eolianites Paleosols Subaqueous (Freshwater) Environments Alluvial Environments Crevasse Splays, Lacustrine Environments Non-marine Deposits and Relative Sea Level Changes Tchnological Recognition of Shoreface Environments Shoreface Variability and the Influence of Storms Storm Deposition and the Ichnology of Tempestites Resultant Ichnological Assemblage Relative Sea level Changes and Shoreface Settings Ichnological Recognition of Deltaic Environments 104. 105 105 105 12 116 116 118 122 130 130 130 132 135 139 140 141 148 Is 11 154 159 165 169 170 170 170 170 172 173, 173, 4 176 7 178 183 183 185 191 195Fluvially Dominated Deltas Wave-Dominated Deltas ‘Tidally Dominated Deltas Ichnology of Deltas and Delta Recognition Facies and the Identification of Subenvironments of Deltas Facies of the Subaerial Delta Plain Delta Evolution and the Role of Relative Sea level Change Ichnological Applications to Incised Valley Fill Successions General Estuarine Incised Valley Models ‘Wave-Dominated Estuaries Tide-Dominated Estuaries Stratigraphic Organization of Incised Valley Fills Ichnology and Sedimentology of Incised Valley Complexes, Brackish Water Ichnology Stratigraphic Discontinuities Facies of the Incised Valley Fill Bay Head Delta Complex Distributary Channels and Point Bars Bay Head Delta Front Central Basin Complex Estuary Mouth Complex Channel-Fill Complex ‘Comparison of Viking and Glauconite Incised Valleys Technological Applications to Deep Marine Deposits Shelf Environments Tide-Dominated Shelves ‘Strom-Dominated Shelves Deep-Water Environments Pre-Turbidite Assemblages and Graphoslyptids Low Oxygen Suites in the Deep Sea Post-Turbidite Assemblages Relative Sea Level and Submarine Fans References: Depositional Systems Referenes 199 201 202 202 207 209 213 213 215 216 217 aR, 218, 223 225 228 225 228 230 232 235, 26 239 239 239 240 241 243 248 250 251 263GENERAL PRINCIPLES OF ICHNOLOGY ‘Trace Fossils (or ichnofossils) are biologically produced sedimentary structures that include tracks. trails, burrows, borings, fecal pellets and other traces made by organisms. Excluded are markings that do not reflect a behavioral function. Owing to their nature, trace fossils can be considered as both pale- ‘ontological and sedimentological entities, thereby bridging the gap between two of the main subdivisions in sedimentary geology. Four major categories are recognized: (1) Bioturhation Structures- « biogenic structure that reflects the disruption of biogenic and physical stratification features or sediment fabries by the activity of an organism: includes tracks, trails, burrows, and similar structures; (2) Biostratification Structures- a biogenic sedimentary structure consisting of stratification imparted by the activity of an ‘organism: biogenic graded bedding, byssal mats, certain stromatolites, and similar structures; (3). Biode- positional Structures- a biogenic sedimentary structure that reflects the production or concentration of sediments by the activity of an organism: includes fecal pellets, pseudofeces, products of bioerosion, and similar structures; and (4) Bioerosion Structures- a biogenic structure excavated mechanically or bi chemically by an organism into a ridged substrate: includes borings, gnawings, scrapings, bitings, and related traces (Fig. 1). In fact the contributions to sedimentary geology already are considerable (Table 1), and numerous additional applications and 1 ts are currently underway, These contributions can be viewed in terms of traditional fields such as paleontology, stratigraphy, and sedimentology. Indeed, ichnology - like Petroleum geology - is more a blending of related disciplines than a singularly unique entity. Recent summaries dealing with general ichnological principles can be found in Pemberton et al. (1992) and Bromley (1996), ‘Table 1. Major contributions of ichnology to sedimentary geology. Relative importance (indicated by the number of +) refers to the relative worth ofthat information in contemporary sedimentary geology. (Modified from Frey and Pemberton, 1985). Geologic Setting Disciplines and Components Modern Ancient 1. Paleontology A. Fossil record of soft-bodied animals : B. Patterns of activity by benthic organisms C. Diversity of fossil assemblage D. Evolution of metazaans 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 - 1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, . - 1D. Sediment reworking: 1) Destruction of initial fabrics and sedimentary structures . ~ 2) Construction of new fabrics 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 . . Bathymetry . D. Temperature . . E. Salinity F. Depositional history 1) Rates of deposition . 2) Amounts of sediment deposited or eroded + G. Actation of water and sediments H. Substrate coherence and stability . 5. Consolidation of sediments A. Initial history of lthific B. Measures of compaction — . ‘THE. CONCEPTUAL FRAMEWORK OF ICHNOLOGY ‘The importance of ichnology to the fields to statigraphy, paleontology, andl sedimentology stems from the following characteristics displayed by trace fossils: (1) long temporal range - although a disad- vantage in refined biostratigraphy, it greatly facilitates paleoecological comparisons of rocks differing in age (Fig, 2); 2) narrow facies range - reflects similar responses by tracemaking organisms to given sets of paleoecological parameters (Fig. 3); (3) no secondary displacement - biogenic sedimentary structures, where preserved intact, are closely related to the environment in which they were formed (Fig. 4); (4) occurrence in otherwise unfossiliferous rocks - trace fossils are commonly enhanced by the very diage- netic processes that typically obliterate tests and shells, especially in siliciclastics (Fig. 5), (5) ereationBiogenic Sedimentary Structures _ | Bioturbation Structure: Dinosaur Track Bioturbation Structure: Rosselia Biostratification Structure produced by the Ghost Crab, Ocyopoda quadrara Bioerosion Structure: Sponge Boring Entobia Fecal Mass of the Lugworm, Arenicola marina Figure 1. Major relationships among biogenic structures. (Modified trom Frey and Pemberton, 1985.)Ichnological Principles Most trace fossils have long temporal ranges. Arenicolites Dsapkyenr Figure 2. Trace fossils exhibit long temporal ranges because basic behaviour has not changed since the dawn of the Phanerozoic. 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 (Table 2); such organisms, in many environments, represent the greatest biomass (Fig. 6); (6) a particular structure may be produced by the work of two ‘or more different organisms living together, or in succession, within the structure (Fig. 7); (7) the same individual or species of organism may produce different structures corresponding to different behaviour patterns (Fig. §); (8) the same individual may produce different structures corresponding to identical behaviour but in different substrates (Fig. 9). (e.g. in sand, in clay. or at sand-clay interfaces): and (9) identical structures may be produced by the activity of systematically different trace-making organisms. ‘where behaviour is similar (Fig. 10) (Ekdale et al.. 1984),Such characteristics are very useful in facies analyses, including reconstruction of individual paleoecological factors. sedimentary dynamics, and the documentation of local and regional temporal facies changes. Like all disciplines, ichnology has evolved 1 set of terms needed for the conveyance ofits concepts. The more basic ichnological terms and concepts (able 3) revolve around behavior by organisms. as opposed to their body parts. Benthic activity. together with its physical manifestation as a trace. is the single most fundamental ingredient of 4Ichnological Principles Most trace fossils are largely facies dependant. Diplocraterion __ Ophiomorpha ‘Skolichos Figure 3. Most trace fossils have a narrow facies range reflecting the organisms response to a given set of paleoecological parameters Ichnological Principles No secondary displacement - biogenic sedimentary structures, where preserved intact, are closely related to the environment in which they were formed Bored beachrock Diplocraterion emplaced at top of a coal bed. Caution you must be able to recognize that clasts can be subject to secondary ‘transportation Cretaceous, Grand Rapids bivalve Teredo Formation, Alberta | | Teredolites in log, Lower “Modern log bored by the | Figure 4. Generally trace fossils display no secondary displacement. 3Ichnological Principles Trace fossils are common in rocks that otherwise are unfossiliferous. The Lower Cretaceous Viking ‘The Upper Cretaceous Formation, Alberta Cardium Formation, Alberta Figure 5. Trace fossils are common in rocks that are otherwise unfossiliferous. Ichnological Principles Non-preservation of the causative organism. Schaubeylindrichnus Conichnus The makers of most burrows systems are comonly never preserved in the burrow. Figure 6. Trace fossils are gen any other way ally created by soft bodied biota that are generally not preserved 6‘Table 2, The approximate number of species, habitat, and fossil record of the soft hadied so-called ‘Minor Phyla” (data from Laverack and Dando, 1979 and Briggs and Crowther, 1990), Phylum Nomertini Nematoda Nematomorpha Sipuncula Echiura Pogonophora Priapulida Phoronida Cephalochordata Chaetognatha Urochordata ‘Onychophora Annelida Approximate Number of Species 800 500,000 250 320 135 100 9 Is 120 8,750 Habitat* MT ETM F, (one M) £5 <5 5 £ ££ © EZ E ™ 2 3 * F = freshwater; T= terrestrial; M= marine Fossil Record Jurassic-Recent Cambrian-Recent Eocene-Recent 2Cambrian-Recent Cretaceous-Recent Nil Nil Nil Cambrian-Recent Nil Cambrian-Recent Nil Cambs Recent Cambrian-RecentIchnological Principles Multiple architects may produce a single structure; such composite burrows are difficult to interpret Figure 7. Trace fossils may represent the work of two or more different organisms. Ichnological Principles ‘The same individual or species can produce different structures corresponding to different behavior ‘The worm, Nereis ean produce both Asterosoma and Rosselia depending on Figure 8. The same individual may produce different structures corresponding to different behaviour. 8Ichnological Principles The same individual may produce different structures corresponding with identical behavior but in different substrates ‘The thalassinid shrimp, Axius serratus will produce Ophiomorpha in sandy substrates and Thalassinoides in muddy substrates. Ophiomorpha Figure 9. The same individual may produce different structures corresponding to identical behaviour in different substrates Ichnological Principles Identical structures maybe produced by the ‘activity of systematically different tracemaking organisms where behavior is simil ‘Skolithos cau be constructed by multitude of different groups of organisms ‘including Polychaetes, Siphunelids, Phoronids and Nematodes. Figure 10. Identical structures maybe produced by different organisms that behave the 9‘Table 3. Basic concepts in the study of biogenic 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 behavioural function: biogenic sedimentary structures, bioerosion structures and other miscellaneous features representing 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 biogenic and physical stratification features or sediment fabrics by the activity of an organism: tracks, burrows, and similar structures. 2) Biostratitication structure - a biogenic sedimentary structure consisting of stratification features imparted by the activity of an organism:biogenic graded bedding, byssal mats, certain stromatolites, and others. 3) Biodepasition structure - biogenic sedimentary structure that reflects the production or concentration of sediments by the activities of an organism: fecal pellets, pseudofeces, products of biocrosion, 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. Iehnology - the over-all study of traces made by organisms, including their description, classification, and interpretation. 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 revert Excludes biostratfication structures and other traces lacking diagnostic anatomical features B. _Lebensspur - synonymous with trace (plural = lebensspuren). Ichnocoenose - an association of environmentally related traces; somewhat ‘analogous to a community. Components include the ichnofauna, or animal tuaces, and the iehnoflora, or plant aces (such as algal borings and certain 100t patterns). 10‘Trace fossil ~ a fossil trace (= ichnofossil). E, _Iehnofacies - the fossil record of an ichnocoenose; typically recurs through long intervals of geologic time and is more or less characteristic of a given set of environmental conditions, 3. Ethology ~ in ichnology, the study or mterpretation of the behaviour of organisms catty their traces. ‘Toponomy - in ichnology, the description and classification of lebensspuren with respect to their ‘mode of preservation and occurrence; includes stratinomic (fabrication) and taphonomic (destruction) aspects, as well as geometry and configuration. ‘Taxonomy ~ in ichnology, the classification of trace fossils according to their systematics and nomenclature. Prevalent taxa include the ichnogenus and ichnospecies. CLASSIFICATION OF TRACES Unique classification schemes have been developed in order to decipher trace fossils. Histori- cally, 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 ichnol- ‘ogy presupposes that the structures were produced biogenically. Numerous descriptive-genetic terms are useful in classifying trace fossils (Table 4). Among the more fundamental terins are burrow walls, burrow fillings, burrow linings, spreiten structures, and burrows versus burrow systems. ‘Table 4. Basic descriptive-genetic terms used in ichnology. (Adapted from Pemberton, er al., 1992,) ‘Tracks and ‘Trails ‘Trackway - a succession of tracks refills, ‘Track ~ an impression left in underlying sediment by an individual foot or podium, ‘Trackway ~ a succession of tracks reflecting a single excursion oF trip. Runway - a surface groove, trench, or depression used repeatedly as a pathway. ‘Trail - a continuous groove produced during locomotion by an animal having a part of its body in contact with the substrate surface, or a continuous subsurface trace made by an animal travelling from ‘one point to another. Undertrack ~ biogenic structure produced by deformation of sediment laminae directly beneath a track and usually preserved in cleavage relief. uBurrows/Borings Burrow - an excavation made within unconsolidated sediment. Burrow system - highly ramified and (or) interconnected burrows Boring - an excavation made into a hard substrate (e.g, rock, shell or wood). Shaft - a dominantly vertical burrow, or a dominantly vertical component of a burrow system having prominent vertical and horizontal parts. ‘Tunnel - a dominantly horizontal burrow, or a dominantly horizontal component of a burrow system having prominent vertical and horizontal parts (~ gallery) Cell ~ a subsurface chamber, along or at the end of a shaft or tunnel, generally ovoid or spherical in shape, and larger than the diameter of the associated shaft or tunnel. Cells along otherwise straight burrow components have been called “turn-arounds” Burrow lining - a thickened burrow wall, typically constructed by organisms as a structural reinforce- ‘ment. May consist of 1) host sediments retained essentially by mucus impregnation; 2) pelletoidal aggregates of sediment shoved into the wall; 3) detrital particles selected and cemented like masonry; ‘or 4) leathery or felted tubes consisting mostly of chitinophosphatic or other secretions by organisms. Burrow linings of types 3 and 4 are commonly called dwelling tubes. Chimney - a tube-like structure constructed above the substrate surface as a continuation or extension of the burrow aperture; generally made of mud or clay. Burrow fill - sediments filling a burrow. Sediment fill may be either active, if emplaced by animals, or passive. if done by gravity. Active fill is termed backfill whenever meniscate laminae, etc, show that the animal packed sediment behind itself as it moved through the substrate. Miscellaneous Configuration - in ichnology, the spatial relationships of lebensspuren, including the disposition of ‘component parts and their orientation with respect to bedding and (or) azimuth direction. Spreite - a blade-like to sinuous, U-shaped, or spiralled structure (plural = spreiten) consisting of sets or co-sets of closely juxtaposed, repetitious, parallel or concentric feeding or dwelling burrows or ‘grazing traces. Individual burrows or groves comprising the spreite commonly anastomose into a single trunk or stem (as in Daedalus) or are strung between peripheral “support” stems (as in Rhizocoraltiun). Retrusive spreiten are extended upward, or proximal to the initial point of entry by the animal, and protrusive spreiten are extended downward, or distal to the point of entry. Xenoglyph - omament imparted to the wall of a trace fossil by structural characteristics of the substrate; primarily a non-biogenic sculpture. Bioglyph ~ omament impasted to the wall of a wave fossil by the bunowi ‘organism; a biogenic sculpture, Rhizolith - general term for a trace fossil or body fossil of plant roots: includes thizocretions. root ‘custs, root colds, root petrifactions and root tubles Graphoglyptid ~ complex horizontal tunnel system that serves as both a permanent dwelling and farm ‘or trap for obtaining food. tor boring aetivity of anPreservational Classi cations Classifications of the stratigraphic or spatial arrangements and modes of preservation of trace fos- sils are both descriptive and interpretive. These preservational concepts may be reduced t0 two basic facets: (1) toponomy, including modes of occurrence and the mechanical-sedimentological processes of alteration and preservation, and (2) physiochemical (pre- through post-diagenetic) processes of preserva tion and alteration, “Toponomic (or stratinomic) classification schemes (Iuble 9, Fig. 11) have been devised by Sei- lacher (1953, 1964), Simpson (1957), Martinsson (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; 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 lin- ings that may be simple secretions of mucus or particulate walls, agglutinated by organic compounds are preferred sites for subsequent mineralization. Therefore, diagenetic processes that obliterate other fossils may even enhance the preservability of trace fossils EPICHNIA ENDICHNIA 1 hhypichnial ridges (hypichnial groove casts) ‘exichnial burrow casts ‘A. MARTINSSONS CLASSIFICATION B. CHAMBERLAIN CLASSIFICATION Figure 11, Preservational classification of trace fossils in relation to silicielastics. Comparison of the terminologies of Simpson (1957), Seilacher (1964b), Mastinsson (1970) and Chamberlin (1971). (Modified and enlarged from Bromley, 1990.) 13‘Table 5. Toponomic classification of bioturbation structures. (Modified from Pemberton er al., 1992) General Terms ‘Toponomy - the description and classification of lebensspuren with respect to their mode of preserva- tion and occurrence (position on or within a stratum, or relative to the casting medium), and secondly, the interpretation ofthe mechanical origin or processes of preservation of traces (see Table 2). Bioturbation - reworking of sediments by an organism, Reworking by meiobenthos or microbenthos 1s termed eryptobioturbation Rioturhate texture (= ichnofabric) — gross texture imparted to sediments by extensive bioturbation: typically consists of dense, contorted, or interpenetrating burrows or other traces, few of which are distinct morphologically. Where burrows are somewhat less crowded and are thus more distinct individually, the sediment is said to be burrow mottled. Lithoturbation ~ process of succes bioerosion. episodes of bicerosion - sedimentation - cementation Classifie mn by Seilacher (1964) Descriptive Terms A. Semirelief - a trace fossil preserved at a lithologic interface: boundary reliefs and cleavage reliefs 1) Boundary relief a semirelief not involving cleavage preservation: hyporeliefs and epireliefs a) Hyporelief — a boundary ve concave or convex, ») Epirelief —a boundary relief occurring on the top of a stratum: relief may be concave orconvex. 2) Cleavage relief — a semirelief in which subsurface laminae are deformed during production of the surficial trace: parting of these laminae (as by weathering fissility) reveals vertical repetition of a given lebbensspur, any isolated specimen of which resembles a single boundary relief, occurring on the sole of a stratum, relief may be Full relief — a structure preserved within a stratum, Genetic ‘Terms ‘A. Exogene — a surficial structure covered by sediment differing from that of the host substrate. B, _Endogene — a structure filled actively or passively within the host sediment CC. Intergene — an endogenie structure produced at u bed junction. 4Classification by Simpson (1987) Red junetion preservation - trace fossils appear in relief at a bed junction or contact. Biogenic activity ‘occurred at the interface between beds of different lithology. Concealed bed junction preservation - individual masses of burrow-filled sediment appear to be isolated within a bed of different lithology; e.g. a shale-filled burrow in sandstone; no obvious connection with an overlying shale bed exists because it was removed by erosion Diagenetic preservation — trace fossils preserved as precompactional nodules or nodule protuberances. Presumably formed during early diagenesis; pore fluids were controlled by the reduced porosity of the filled burrows relative to that of the matrix. Frequently observed at bed junctions in sequences of alternating argillaceous limestones and shales. Burial preservation - filled burrows subsequently exhumed by currents winnowing away the soft host matrix. Coherent burrow linings thus lay on the sediment surface and, where preserved, were buried Quickly by later sediment Classification by Martinsson (1970) Epichnion ~ a structure preserved at the upper surface of the main body of the casting medium: may appear as a ridge or groove, e-g., epichnial ridge. Hypichnion - a structure preserved at the lower surface of the main body of the casting medium: may appear as a ridge or groove: e.g, hypichnial groove, Endichnion - a structure preserved within the main body of the casting medium. Exichnion - a structure preserved outside the main body of the casting medium: e.g., sediment piped into underlying bed. Classification by Chamberlain (1971) Epigene - a surficial structure covered by sediment differing from that of the host stratum. Intergene - an endogenic structure produced at a ltho-logic interface or bed junction. Endogene - a structure filled actively or passively within the host sediment.Behavioral Classification Perhaps the single most important ingredient of ichnology is the functional interpretation of indi- vidual traces. Fundamental behavioral (or ethological) patterns are dictated and modified not only by genetic preadaptations, but also by prevailing environmental parameters. Ekdale et al. (1984) recognized seven basic categories of behavior (Fig. 12, Table 6); resting traces (cuhichnia), locomotion traces (repichnia), dwelling traces (damichnia), grazing traces (pascich- nia), 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 equi- libria (ugichnia) to all other behavioral patterns (Fig. 13). ‘These fundamental behavioral pattems are genetically controlled, but are not phylogenetically restricted. The basic ethological categories have generally persisted throughout the Phanerozoic. Indi- vidual 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 suspension feeders. Similarly, aquatic, inverte- brate locomotion traces can be preserved only in fine grained, low energy, quiescent environments such as lagoons. This ability to discem the behavioral trends of benthic organisms represented in the rock record ‘greatly facilitates environmental interpretations (Fig. 14). Figure 12. Ethological classification of trace fossils, and relationships of these to body fossils. Overlap of categories acknowledges the intergradation inherent in nature; depending on local circumstance, for example, the spectrum of escape structures might span the entire circumference of the circle. (Modified and enlarged from Frey and Pemberton, 1985.) 16‘soreasqns popesdie 4 ‘pousosad aq mu wos s.jduo “sams aensqnsfeUrLO rusts pades-umoy eras ur sung parson sounds ou2ioydi, (emyouuog) ssumanas Suman -ponsesaud ag Zeus wo) 21du0> Burppeq “saurtonns 3uy/94p YH opssoy, ‘onpedsos ya sofBur snowea ye poo “snonu kqss3jun euonepes 29 0; pust aug ~ Summ puncsBspun, 01 speyduig “susie2i0 aptnoxd Ans os sunisnns 4819929} ‘ys snonuss ca eoupUIé>“paxsuagun 4 “PO4UEA BUS asodop sq poionasuos swauing Ksuoduo1 89] 40 309, (euypurpo,y semen Susp snskydonds -punjd agor spusymmanasyos9K0 “poaasad 9q Kou Bayar duns, 4 snodoysue sno1seyeq Buspe9s ued st soy sydwa “20eprsqns aensqns 99 s9pun De saZeB |p 8te so s9poopasodap 2) qou Xq peur ‘snonunuo2sip twa4p Jo Aur ‘sau; pu se pausonrd “9x00 (ouuypiseg) some Buz, posqoaursg, sosouaisy sidux $Bojoudiowy onsuorerey oni (Zo61 1 19 wouDquiAg WON pardepy) “S[ss0 99% STRIGUIDAUT JO VOMPOUSSAD WIAD|ONG “S SRL a18 emaro, smyrna ydomionds wosioypooqog aaxdopong sojduxer us sBumnung “sqes> snavon Kq poyENE asosaid 29 At sts05 21qEt40) ‘iepurau sb yons waned Bqoydioyy onsuo12804 ‘Suyasous [2109 29°9009 aby uo paanpoud saumianns uwontoo1d snowgo wos vwonuysarrs as Feeding _— [Resting \ : ‘Dwelling — [Locomotion| ian Pascichnia Domichina Cubichnia Repichnia Figure 13. Rehavioral classification of lebensspuren. With environmental fluctuation, virtually all traces are intergradation: with the fugichnia - escape or equilibrium structures. Bromley (1990) proposed a new ethologic category, the equilibrichnia to encompass such cases. (Modified from Frey et al., 1987.) Ethological Classification Figure 14, Major behavioural classes. 19Phylogenetic Classification Perhaps one of the most frustrating albeit most fascinating facets of ichnology is the attempt to establish the zoological affinities of specific ichnofossils. Chamberlain (1971) pointed out that difficultics arise in any systematic classification of ichnofossils, partly because extensive comparisons of trace mor phology and ethology with that of most modem organisms has not yet been made. Furthermore, ichnofos- sils mostly reflect behaviour of animals, and only to a small extent reflect their anatomy or morphology. ‘The result is that more than one genus or specics of ichnofossil may have been constructed by a single species of animal, or conversely different species of animals may have made identical species or genera of trace fossils (Frey and Seilacher, 1980). For example, specimens of Skolithos linearis at one locality may show affinities to the phoronids (Fenton and Fenton, 1943) whercas at another locality they may show affinities to onuphid polychaetes (Curran and Frey, 197). Finally, a nestler or commensal organism in some instances may be better suited for prescrvation within a burrow than the animal that constructed the burrow (rey, 1975). ‘Therefore, each occurrence of a given ichnofossil most be treated on an indi- vidual basis; sweeping gencralizations on their zoological affinitics should be avoided. Notable excep- tions include certain types of distinctive, hard substratc borings and those rare instances where the trace~ ‘making organism is preserved within the burrow. In most cases, however, allibuting & particular ave « particular soft-bodied organism depends on a uniformitarian approach (Pickerill and Forbes, 1978) and other indirect lines of evidence (Fig. 15). Phylogenetic Classification Although very difficult sometimes it is possible to match the trace making organism to the individual trace fossil. One must approach this with caution but it can assist in paleoecological reconstructions. Lockeiais thought to represent Conichaus is thought to represent the eee, eee ez sang opiate. gate ees neem Seale eras ‘gure 15. Although difficult, itis possible to establish the zoological affinities of specific ace fossils. 2»‘THE ICHNOFACIES CONCEPT Perhaps the essence of trace fossil research involves the grouping of characteristic ichnofossils into recurring ichnofacies. ‘This concept developed by Adolf Seilacher in the nineteen-fifties and nineteen- sixties, was based originally on the Fact that many of the parameters that control the distribution of trace~ rmukers tend to change progressively with increased water depth (Fig. 16). Because of the potential geological value of this hathymetric relationship, the Seilacherian ichnofa- ies sequence soon came to be regarded almost exclusively (albeit erroneously) as a relative paleobathom- eter. Today the ichnofacies remain valuable in environmental reconstruction (Fig. 17), but paleobathym elty is only one aspect of the modem ichnotacies concept (Frey ef al., 1990) ‘The ichnofacies concept also is related closely to two other concepts: ichnocoenoses and facies sensu stricto, Simply said, an ichnocoenose is an association of contemporaneous, environmentally related traces, somewhat analogous to a community of organisms, and an ichnofacies is the preserved record of that ichnocoenose (Frey and Pemberton, 1985). The Seilacherian ichnofacies are merely distinc- tive, recurrent, archetypal associations of traces made most useful when put in the context of the original ichnocoenoses. In practice, however, different workers (Bromley, 1990) have applied the term ichnocoenose (some- times spelled ichnocoenosis) in different ways. The concept clearly applies not only to traces in modem settings but also to their ancient counterparts as interpreted from the rock record (Frey and Pemberton, 1987). Modem ichnococnoses (Dérjes and Hertweck, 1975) are studied mainly w gait uniformitarian insights into possible fossil equivalents. Second, ichnofacies and reconstructed ichnocoenoses are part of the total aspect of the rock and therefore, like lithofacies, are subject to Walther’s Law. For example, isolated bored shells or clasts do not itute the Trypanites ichnofacies. Rather, there should be some semblance of stratifica- tion, lateral continuity, and vertical succession. This relation is another strength of ichnology, however, interpretations of ichnofaunas are improved substantially when the traces are studied in the context of host rocks and their implications, in themselves co Archetypal Iehnofacies [Nine recurring ichnofacies have heen recognized, each named for a representative ichnogenus: Scoyenia, Trypanites, Teredolites, Glossifungites, Psitonichnus, Skolithos, Cruziana, Zoophy cos, and Nereites. These trace fossil associations (Fig 16) reflect adaptations of tracemaking organisms to numerous environmental factors such as substrate consistency, food supply, hydrodynamic energy, and salinity and oxygen levels (Pemberton et al, 1992)). ‘Traces in nonmarine and brackish marine set- tings are in need of further study; the marine softground ichnofacies (Psilonichnus, Skolithos, Cruciana, Zoophycos, and Nereites) are distributed according to numerous environmental parameters; traces in the firmground (Glossifungites), woodground (Teredolites), and hardground (Trypanites) ichnofacies are dis- tributed on the basis of substrate type and consistency (Fig. 18). Representative occurrences of the various ichnofacics arc summarized below. Ilowever, each may ‘appear in other settings, as dictated by characteristic sets of recurrent environmental parameters. From the a(S861 ‘uousquiag pur ous woxy paytPoHy) omponynd (9F, ‘Soasydoog (sz ‘munzua0 aydoads (og ‘aydner poishyd [woo] “stuoIpras If westeIp andouss “gy aundty, senuedsuy rye ouTULIp Krew HN 0:3: TruuvoH:AUD Jo ons “oatsnjox9 20U Ne “arineyUasaidas v UF 19s SoFDesDUYD! DUNE TuLNoAI Zu lsnuycruerisa| sayjesey |soodydooz| euerzug | soumoys samoparay SERB de hada: 2 vez sens ayensans powpipsuco ius)Seilacher's Concept of Recurring Ichnofacies TRACE FOSSILS mm BEHAVIOUR ma ENVIRONMENT Trace fossils are a manifestation of behaviour which can be modified by the environment. ECOLOGICAL CONTROLS The distribution and behaviour of benthic organisms is limited by a number of interrelated ecological controls, including: 1. Sedimentation Rate 5. Turbidity 2, Substrate Coherence 6. Light 3. Salinity 7. Temperature 4. Oxygen Level 8. Water Energy FOOD RESOURCE PARADIGM The Food Resource Paradigm is a limiting factor on the distribution of tracemaking organisms in shallow marine environments FOOD PARTICLES DEPOSITED + ervaasxx ——_- Figure 17, Major controls on the distribution of tracemaking organisms as summarized in the Seilacherian ichnofacies models.wooDGROUND| HARDGROUND | FIRMGROUND | SOFTGROUND Termitichnus | reresciat Scoyenia Mermia Teredolites. | Trypanites | Glossifungites | Psilonichnus Skolithos ten ees Cruziana eum ery Marine Zoophycos Low Eaeey Nereites Figure 18, Relationship of ichnofacies to substrate, as opposed to softground ichnofacies which have been differentiated on environmental factors, hardground, firmground, and woodground ichnofacies ave differentiated from oie another by substrate type and consistency. (Modified from Bromley et al., 1984) standpoint of ethological requirements of tracemaking organisms, for example, certain intertidal backbar- rier environments are not all that different from certain subtidal forebarrier environments and may contain Virtually identical suites of lebensspuren. Non-Marine Ichnofacies At present, nonmarine environments have been characterized by three main ichnofacies the Scoy~ enia ichnofacies, the Mermia ichnofacies, and the Termitichnus ichnofacies (Table 7). Frey er al. (1984) concluded that the Scoyenia ichnofacies remains a valid concept within appropriate limits and is sug- gested of deposition in the shore of ephemeral lakes and the overbank of sluggish rivers. The Scoyenia assemblage is characterized by: (1) small horizontal, lined, backfilled feeding burrows; (2) curved to tortuous feeding burrows; (3) sinuous crawling traces; (4) vertical, unlined, cylindrical to irregular shatts, and (5) vertebrate and arthropod tracks and trails. Buatois and Méngano, 1995 concluded that the Mermia ichnofacies typifies unconsolidated, fine-grained, permanent subaqueous substrates, and well-oxygenated, low-energy lake bottoms. The Mermia assemblage is characterized by: (1) dominance of horizontal to sub-horizontal grazing and feeding traces produced by mobile deposit feeders, (2) subordinate occurrence of locomotion traccs, (3) generally high to moderate divetsity and abundance, and (4) low abundance of specialized grazing structures (Buatois and Mangano, 1995). Smith et al. (1993) named the Termitichnus ichnofacies as a distinct terrestrial assemblage characterizing paleosols. The Termitichnus ichnofacies is characterized by: (1) vertebrate tracks and trails, (2) low diversity and abundance, (3) dominated by bee, beetle, and termite nests, (4) few feeding structure, and (5) rhizoliths, coprolites, and leaf miners Smith et al, 1993). 24“sounpuoqut jam pue ‘spuod ‘saumte9y p Anus soumonuns Aarnuowypas qeoiskyd{ “siwowuas!Aus ‘utmur uou ‘onenbe wou pur omenbe £yyny 9998109, sreipouuiaruy “wosard 9q eur sorerofon onerberwos ‘nuaBiowqns Sujuiovag -svou9 Suypojouysamyeay asseduned Ie, eH, “UYpUNge ag ew Sumanss Jo Spury UDA 16 NO} KeaHd 409 Pur SNOUT I emp pyres fosuap o1 fase 0 “sBupog podeys eo) nfissoyp uso} © noss019 ambos wr 'So-ej0uyo! euowepestioy) “aeasqns ay) jouorsou> pers -seunejouyprsorem ——-swouucui soiree yoeq snetseA 30 (orexisqns o1ix) sorsejouys, semjopesay SUss0q a9RAy, INSHIAIIEZEY quacuostau orqnuag TeNdAT, Taser “wrousngouny pu uoLaquiad WON] paXepy) suorteay| Cur yemmUUOSIAUD (9AISn|Dx9 OU Ing) LOUILDD sa} puE SUONIDOSSE ESS} ARN pafjonuos axeAKgns jedKayor? SULINIOY “6 a1GELOn the other hand, many kinds of environmental parameters do tend to change progressively with water depth and distance from shore (Fig. 16), and these gradients affect corresponding changes in we distribution of physical and biogenic sedimentary structures (Table 8). To that extent, trace fossil associa tions are indeed useful in paleobathymetry. Equally important is the long temporal duration of most kinds of trace fossils. ‘These basic benthic behavioural patterns are more nearly like stable ecologic niches than individualistic records of particular animal species (Frey and Seilacher, 1980). As long as the functional niche remains advantageous under given environmental conditions, many different animal species, over long intervals of geologic time, may be expected to exploit it: their preserved traces are strikingly similar and have equivalent significance. Hence, although we conveniently speak of the “Skolithos animal” as the architect for a particular kind of dwelling structure, numerous different animal species actually were involved. ‘The longevity of recur- rent ichnofacies thereby exceeds the longevity of recurrent biofacies by a considerable margin, and are correspondingly more useful as archetypical models not only for environmental interpretation but also for comparisons of depositional environments of widely differing ages. Furthermore, the purpose for recognizing these recurrent ichnofacies must not be overlooked. Inter- preted in terms of the original ichnocoenoses, these ichnofacies are merely archetypal facies models with ‘which the local ichnofacies may be compared. ‘The archetypes are intended to supplement, not supplant, local ichnofacies designations, some of which are quite distinctive; corresponding ichnofaunas may be restricted (Serna, 1986), intergradational (Marintsch and Finks, 1982), or diverse (Dam, 1990). ‘The idealized ichnofacies succession works well in most “normal” situations (Frey and Pemberton, 1984), including distributions according to salinity gradicnts (ef. Bromlcy and Asgaard, 1991); yet one should not be surprised to find nearshore assemblages in offshore sediments, and vice versa, if these accu- mulated under conditions otherwise like those preferred by the tracemaking organisms, for example. The basic consideration rests not with such inanimate backdrops as water depth or distance from shore, or some particular tectonic or physiographic setting, but rather with such innate, dynamic controlling factors as substrate consistency, hydraulic energy, rates of deposition, turbidity, oxygen and salinity levels, wxie substances, the quality and quantity of available food, and the ecologic or ichnologic prowess of tracemak- ers themselves (e,g., Vossler and Pemberton, 19884). Finally, the models should not be divorced from associated pattems of bioturbation. Numerous local ichnofacies, particularly those representing low-energy conditions and slow rates of deposition, are set in a complex biogenic fabric. Several generations of burrows may be discernible via their crosscut- ting relationships, showing that the same volume of sediment passed repeatedly through various styles of reworking. In environmental reconstruction, such ichnologic fabrics may be equally as important as the individual, named trace fossils (Pemberton and Frey, 1984). The residology of traces and fabrics may be complicated indeed (Frey and Wheaterott, 1989), PALEOENVIRONMENTAL SIGNIFICANCE OF TRACE FOSSILS ‘The concept of functional morphology, a basic premise employed by ecologists and paleoecolo- sists in environmental reconstruction, is equally applicable to ichnology. In fact, trace Fossils are unique in that they represent not only the morphology and ethology of the tracemaking organism but also the 2physical characteristics ofthe substrate; they are closely inked to the environmental conditions prevailing a the tine of Uheit constuction, Fiey and Seilacher (1980) re-emphasized that such variables as bathym- etry, temperature, volumes of sediment deposited or eroded, aeration of water and sediment, and substrate coherence and stability. have a profound effect on resultant trace fossil distributions and morphology. and hence, can be used in the determination of original biological, ethnological, and sedimentological condi tions The application of ichnology to paleoenvironmental analysis goes far beyond the mere establish- ment af gmss or archetypal ichnafacies. For instance, shallow-water, coastal marine environments com prise a multitude of sedimentological regimes, which are subject to large fluctuations in many physical and ecological parameters. In order to fully comprehend the depositional history of such zones in the tock record, itis imperative to have some reliable means of differentiating subtle changes in these param- ters, Detailed investigations of many of these coastal marine zones in Georgia have shown the value of utilizing biogenie sedimentary structures (in concert with physical sedimentary structures) in delineating them (Frey and Pemberton, 1987). The application of these studies in deciphering paleoenvironments has also proven invaluable. The use of trace fossils in the interpretation of freshwater deposits is becom- ing increasingly important. Recent work by Buatois and Mangano (1995), among others, have stressed the abundance and diversity of tracemaking organisms in freshwater environments and emphasized their potential importance in palcocnvironmental reconstruction. Distinet 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 a, 1984), Recently, marginal marine environments (including tidal channels, estuaries, bays, shallow lagoons, delta plains, etc.) have been recognized with more fieyuency in Ue 1k record, Such envie ronments characteristically display steep salinity gradients, which, when combined with corresponding changes in temperature, turbulence, exposure, and oxygen levels, result in a physiologically stressful envi- ronment for numerous groups of organisms. The typical trace fossil 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 mor- photogically simple structures constructed by trophic generalists; (4) a mixture of elements which are common to both the Skolithas and Cruziana ichnofacies: (5) assemblages that are commonly dominated by a single ichnogenus; and (6) diminished size compared to fully marine counterparts (Pemberton and Wightman, 1992). 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 hhand, must temper their observations in the light of organic evolution. Although trace fossils can be considered as biogenic sedimentary structures and are difficult to classify phylogenetically, they are con- structed by biological entities and are thus subjected 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 diversification of the angiosperms in the Late Cretaceous. Prior to this time terrestrial substrates may not have been as extensively bioturbated due to a paucity of tracemakers. Likewise, pat- temed grazing traces, which characterize deep-sea environments, show a trend toward more complex organization through most of the Phancrozoie. This trend may be related to the evolution of more efficient foraging strategies (Seilacher, 1986). For these reasons, paleoen vironmental interpretations based on trace 33fossils must be considered not in strict uniformitarian terms, but rather, in actualistic ones. Equally impor- tant, unique quantitative environmental indicators are indeed rare in the geological 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 1s accentuated when fully integrated 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 se the following (Figs. 19): entary geology is based on ‘A) Biogenic structures represent the activity of soft-bodied organisms that are not generally pre- served. Such organisms (including many cntire phyla) arc commonly the dominant component of the biomass of many environments. B) Biogenic structures are commonly enhanced by diagenesis and can be used in horizons 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 mineral, it enhances the visibility of biogenic structures. ©) Biogenic structures can be associated with facies that do not contain any other fossils. In many siliviclastic regimes, diagenesis dissolves most of the shelly fauna and trace fossils represent the only clue as to the original biogenic component of the unit D) Biogenic structures can be used forthe paleoecological reconstruction of depositional environ- ments, 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, Glossifung- ites, and Teredolites ichnofacies are substrate-controlled, and are emerging as important elements in the evolving concepts of sequence stratigraphy. G) Biogenic structures ordinarily cannot be transported and therefore represent the original envi- ronmental position of the trace making animal H) Biogenic structures are sensitive to changes in certain ecological parameters that are otherwise Aifficultto ascertain, Trace fossil models are emerging to recognize changes in such parameters as salinity and oxygen levels. D An integrated approach utilizing physical, chemical, biological, and ichnological lines of e' dence constitutes @ powerful tool for facies interpretation, 34The Conceptual Framework of Ichnology A. Biogenic structures represent the activity of soft-bodied organisms that are not generally perserved. Such organisms (include many entire phyla) are commonly the dominant conponent of the boimass of many environments. B. Biogenic structures are commonly enhanced by diagenesis and can be used in horizons where physical sedimentary structures have been masked. In the example below from the Cayman Islands the coral head has been dissolved by perculating ground water and the borings having a more resistant cement were not affected and are preserved. C. Biogenic structures can be associated with faciess that do not contain any other fossils. In many siliciclastic regimes, diagenesis dissolves most of the shelly fauna and trace fossils represent the only clue as to the original biogenic component of the unit, Figure 19 . Concepts in the study of trace fossils, 35The Conceptual Framework of Ichnology D. Biogenic structures can be used for the paleoecological reconstruction of depositional environments, In the MeMurray Formation IUS beds ave Interpreted asthe lateral aceretion deposits of meandering channels. The trace fostile indicate that the channel was probably esturine in origin, E. Biogenie structures are sensitive to fluctuations in sedimentary dynamics. Bioturbation patterns are important in recognizing event beds and ‘sedimentation patterns. {In clastic regimes episodic depositional events are common Including: turbiates, tempesites, selsmites, pnyto-detrital pulses, lateral delta lobe switches and inundites. F. Biogenic structures are sensitive indicators of substrate coherence. The Trypanites, Glossifungites and Teredolites ichnofacies are substrate-controlled and are emerging as important elements in the evolving concepts of sequence stratigraphy. Figure 19. (Continued). 36The Conceptual Framework of Ichnology G. Blogente structures ordinarily cannot be transported and therefore represent the original environmental position of the trace making animal, In Drumeller te transgression ls marked by the Teredolites lehnafacies that Ie Incked Into the exposed peat bed. ‘Commonly such transgressions bottom out on peats because they are diicut to erode, H. Biogenic structures are sensitive to changes in certain ecological parameters that are otherwise difficult to ascertain. Trace fossil models are emerging (o recognize changes in such parameters as salinity and oxygen levels. In the Manville Group of Southern Alberta, a ‘monospeciti Gyrolies-dominated assemblage |y present in many of the deposits. This ‘assemblage is commonly associated with the Interpretation of brackish water environments. I. An integrated approach utilizing physical, chemical, biological and ichnological lines of evidence ‘constitutes a powerful tool for facies interpretation. Figure 19, (Continuew) 37ICHNOLOGY OF MARINE CLASTIC ENVIRONMENTS Shallow-water, coastal marine environments comprise a multitude of sedimentological regimes, which are subject to large fluctuations in many physical and chemical parameters. In order to comprehend. fully the depositional history of such zones represented in the rock record, it is imperative to have some reliable means of differentiating subtle changes in these parameters. Detailed investigations of many ‘coastal marine zones have illustrated the value of using biogenic sedimentary structures in delineating such ecological parameters as oxygenation, salinity, and energy levels, For instance, Darjes and Hertweck (1975) subdivided the coastal zone into three major environments, based primarily on the position of mean, high water, mean low water, and wave base. ‘Their faunistic investigations also confirmed the importance ‘of minimum and maximum ve hase as distinet houndaries separating animal communities (Fig. 20) From a paleoenvironmental point of view, marine clastic environments have received previous in ‘depth analysis and will be discussed only briefly here. Nonmarine and marginal marine environments, ‘on the other hand, are poorly understood and we will attempt to outline their basic ichnological characteristics. ‘THE BASIC SOFTGROUND ICHNOFACIES. Softground ichnofacies tend to be differentiated from one another by variables that typically are depth related. The Zoophycos and Nereites assemblages are more characteristic of deep-water environments, whereas the Psilonichnus, Skolithos, and Cruciana ichnofacies are rep- resented in nearshore marine or coastal environments. For example, in the Cretaccous of the Wester Interior of North America the inter-deltaic marine shoreface can be zoned ichnologically (Fig. 20). This zonation is based on the food-resources paradigm, which is influenced by relative energy levels (Fig. 17). (Specific details on these softground ichnofacies are given in Table 8.) Recent summaries of the ichnology of marine shoreface environments can be found in publications by MacEachern and Pemberton (1992), Pemberton et al. (1992) and Bromley (1996). Details ‘outlining the effect of storms on the distribution of trace fossils in shoreface settings were outlined by Pem- berton and Mactiachem, 1997). Similarly, the ichnological distinction between river-dominated delta-front environments and the inter-deltaic shoreface was summarized recently by Moslow and Pemberton (1988). ‘The Zoophycos and especially the Nereites ichnofacies tend to characterise deep-water environ- ‘ments, including outer shelf, slope, and bathyal to abyssal settings (Fig. 16). For details on the ichnology of deep marine deposits see the papers. Psilonichnus Iehnofacies The Poilonichnus iclmofacies (Fig 21, Table 8) represents a mixture of marine, Minor Behaviors tube dvellers are pass Fair-eather suites are subenv Figure 20. Idealized shore face mode! of softground ichnofacies successions, based on observations of Cretaceous strata of Western Interior Seaway ot North America (modified from Pemberton and MacEachem, 1995), 39PSILONICHNUS ICHNOFACIES sae 1. Psilonichnus 2. Macanopsis + Low diversity and abundance, * Horizontal trails formed by the crawling and foraging of * Dominated by J-, Y-, or U- insects and tetrapods. shaped dwelling burrows made by ghost crabs. * The ephemeral tracks, trails and fecal pellets of insects, + Vertical shafts of insects and reptiles, birds and mammals; spiders (Macanopsis). these are predominantly predators and herbivores. * Root penetrations are common. ‘The Ghost Crab Ocyopoda quadrata, Sapelo Island, Georgia Figure 21. Schematic of the Psilonichnus ichnofacies (modified from Pemberton etal. 1992) 40Marine processes generally dominate during spring tides and storm surges, whereas maritime aeolian pro- cesses peedaminate during neap tides and nonstorm periods. Such conditions can react in alternations of ‘wind and water depositions producing complex truncated laminae dipping shallowly in diverse directions. Because of their topographic position few such substrates are available to benthic marine animals The only persistent, notable exceptions are amphibious crabs of the Family Ocypodidze, which include both scavengers and surficial deposit feeders: these animals typically excavate J Y-, or U- shaped dwell ing burrows referable to the trace fossil Psilonichnus (Fig. 21). Other biogenic structures are generated by essentially terrestrial organisms and include the vertical shafts of insects and spiders; the horizontal tunnels of other insects and tetrapods; and the ephemeral tracks, trails, and fecal pellets of insects, reptiles, birds, and mammals. The other major type of biogenic. structure that is common in the Psilonichnus ichnofacies relates to plant-root penetrations The types of plants able to exploit these substrates range from intertidal halophytes, on the distal margins of some washover fans, to maritime or terrestrial grasses, weeds, vines, shrubs, bushes, and trees on dunes. To the extent that ocypodid crab burrows may appear in the uppermost foreshore or the upper part of estuarine point bars, the Psilonichnus ichnofacies may slightly overlap the Skolithos ichnofacies: how- ever, the boundary between these two ichnofacies is normally distinct. In contrast, because ofits poten- tially large numbers of terrestrial traces, the Psilonichnus ichnofacies may be broadly intergradational not only with the Seoyenia ichnofacies but also with several other, as yet unnamed nionmarine ichmocenoses Skolithos Ichnofacies The Skolithos ichnofacies (Fig. 22, Table 8) is indicative of relatively high levels of wave ‘or current energy, and typically is developed in clean, well-sorted, loose or shifting particulate substrates. Abrupt changes in rates of deposition, erosion, and physical reworking of sediments are frequent. Such conditions commonly occur on the foreshore and shoreface of beaches, bars, and spits: associated stratification features usually consist of finer, parallel to subparallel, gently seaward dipping laminae to large- and small-scale trough crossbeds, including ripple laminae. As dictated by fundamental intertelationships among water agitation, sediment transport, and animal distribution, most tracemakers found here are suspension feeders. Substrates serve mainly as an anchoring ‘medium. The organisms typically construct deeply penetrating, more or less permanent domiciles (Fig.23). Depth of burrowing in the intertidal zone is controlled in part by tidal range and height ofthe low water table. During low tide, moist sediments at depth help buffer the organisms against desiccation and s perature shock, and also help provide respiratory water. In both intertidal and high-energy subtidal settings, deep burrowing is one means of escaping the instability of the ever-shifting substrate surface (Fig. 24) ity ortem- The Skolithos ichnofacies ordinarily grades landward into supratidal or terrestrial zones. and seaward into the Cruziana ichnofacies. The landward boundary tends to be more abrupt than the latter. With reduced energy, it also may grade into intertidal or shallow subtidal extensions of the Cruziana ichnofacies; ecotonal or mixed Skolithes - Cruziana associations are common in both recent (Howard and Frey, 1973) and ancient settings Finally, the Skolithos ichnofacies may appear in slightly to substantially deeper water deposits wherever energy levels, food supplies, and hydrographic and substrate characteristics are suitable (Crimes et al., 1981). Potential examples include submarine canyons, deep-sea fans, 4- Predominantly vertical, cylindrical, or U-shaped burrows. - Protrusive or retrusive spreiten in some U-burrows, which respond to subtrate aggradation and degradation. - Few horizontal structures. - Few structures produced by mobile organisms. SKOLITHOS 1CHNOFACIES 1. Ophiomorpha 2. Diplocraterion 3 Arenicolites 4. Skolithos - Low diversity, although individual forms may be abundant. - Mostly dwelling burrows constructed by suspension feeders or passive carnivores. - Vertebrate traces, particularly in low-energy settings. Figure 22. Schematic of the Skolithos ichnofacies (modified from Pemberton et al. 1992). 2SKOLITHOS ICHNOFACIES Characteristic Ichnogenera Skolithos Diplocraterion Horizontal Vertical Ophiomorha Ophiomorpha —— Arenicolites Figure 23. Characteristic trace fossils commonly associated with the Skolizhos ichnofacies. 3Ophiomorpha Association Diplocraterion’ parallelum ‘Storm Sand Association Skolithos Ichnofacies Pipe Rock Association Arenicolites variabilis Skolithos linearis Figure 24. The different expressions of th Skolithos ichnofacies. Paleozoic pipe-ock contains profuse numbers of Skolithos and Arenicolites; the Mesozoic and nozoic are dominated by an Opiomorpha assemblage: and tempesites display a modified opportunistic assemblage 44and bathyal slopes swept by strong contour currents. Therefore, as emphasized previously, paleobathymetric interpretations cannot be based solely on checklists of wace fossil names. evaluation of associated physical sedimentary structures, stratigraphic position, and other evidence is, essential, even in normal beach-to-offshore sequences. Cruziana Iehnofacies ‘The Cruciana ichnofacies (Fig. 25, Table 8) is most characteristic of subtidal, poorly sorted unconsolidated substrates. Conditions typically range from moderate energy levels in shallow ‘waters below fair-weather (minimum) wave base but above storm (maximum) wave base, to low energy levels in deeper, quieter waters. Sediment textures and bedding styles exhibit considerable diversity, including thinly bedded, well-sorted silts and sands, discrete mud and shell layers, interbedded muddy and clean silts and sands, and extremely poorly sorted beds derived from any of the above through intense bioturbation. Physical sedimentary structures, where not modified or destroyed through bioturba- tion, include parallel, subparallel, or ripple laminated sand, or trough cross-bedded, megarippled sand. With reduced but not negligible energy levels, food supplies consist of both suspended and depos- ited components; either fraction may predominate locally, or the two may be intermixed. Characteristic organisms therefore include suspension and deposit feeders, as well as mobile camivores and scavengers (Fig. 26). Because of lowered energy and abrupt shifts in temperature and salinity levels, burrows tend to be constructed horizontally rather than vertically, although scattered vertical or steeply inclined burrows, ‘occur. Profusions of burrows may be present at stable, low-energy sites. Trails of epibenthic and endoben- thic foragers also may be common and reflect the abundance, diversity, and accessibility of food (Fig.27).. In shallow waters, periodic scour by storm waves and renewed deposition following their cessation may incorporate storm layers within a sequence of otherwise low-energy deposits (Pem= berton and Frey, 1984). Development of hummocky cross-statification may involve the introduction of new sediment as well as the reworking of previously deposited sediment. Any of these conditions may yield burrow truncations and escape structures. Increased energy and allied parameters thus represent a temporary excursion of Skolithos -type conditions into a Cruziana -type setting. However, this over- all eclding style differs from that of the main Skalithos ichnofacies, in which stratification features, sub- strate scour, burrow truncations, and escape structures are contained entirely within sequences of high- energy deposits. Furthermore, the storm layers eventually are overprinted with Cruziana -type traces. Zoophycos chnofacies Of all recurrent marine ichnofacies, this assemblage is most debated and least understood. The ichnogenus Zoophycos has an extremely broad paleobathymetric range; hence its designation as namebearer for a supposedly depth-related ichnofacies has long been controversial (see Kotake, 1989, 1991; Olivero and Gaillard, 1996; and Uchman and Demfrcan, 1999). In popular bathymetric schemes, the Zoophycos ichnofacies (Fig. 28, Table 8) typically is portrayed as an intermediary between the Cruziana and Nevetes ichnofacies, at a position corresponding more or less to the continental slope. More specifically, the original designation placed it in lysch-mollasse areas below wave base and free of turbidites, within a broad depositional gradient (Seilacher, 1967). AAs reevaluated recently (Seilacher, 1978; Frey and Seilacher, 1980), one of the major environ- 4sES sz andy wo SHTOVAONHOI VNVIZQNYO “sunsjunse gout Aq aprur soamanays Jo a909s94¢ ~ 46CRUZIANA ICHNOFACIES Chondrites and Rhizocorallium zZ a Individual Asterosoma arms Complete Asterosuma Figure 26. Characteristic trace fossils commonly associated with the Crusiana ichnofacies.. 47CRUZIANA ICHNOFACIES Characteristic Assemblages, Upper Cretaceous Horseshoe Canyon Formation, Alberta Weta Lower Shavotnie Aerie Figure 27. Distal-proximal trends seen in the Crugiana ichnofacies from the Upper Cretaceous Horsecanyon Formation, near Drumheller, Alberta, Distinct assemblages characterize the distal offshore, the proximal offshore, the distal lower shoreface and the proximal lower shoreface. 48mental controls represented by the ichnofacies is lowered oxygen levels associated with organics in quiet-water settings, To the extent that these conditions actually obtain on shelf-edge gradients, the popularized bathymetric placement of the ichnofacies (Fig. 16) is suitable, However, such reducing conditions replete with a dominance of Zoophycos. are perhaps even better known in shallower water, epeiric deposits (Marintsch and Finks, 1982). ‘Considering the above characteristics of the ichnofacies, together with the widespread distribution of individual specimens of Zoophycos in both shallow- and deep-water deposits (Frey et al, 1990; Olivers and Gaillard, 1996; Lichman and Demircan, 1999), we speculate that the Zanphycos animal simply was broadly adapted in most ecologic respects (Fig. 29). If tolerated not only a considerable range of water depths but also numerous substrate types, variable food resources, and. different energy and oxygen levels. Its traces therefore appear in the Cruziana through Nereites ichnofacies (Crimes et al 1981). Indeed, in final analysis, its tolerance may be its most distinguishing environmental characteristic. The animal was able to complete successfully with diverse tracemakers under Cruziana- and Nereites- type conditions, but few other animals were able to compete with it under the restricted conditions outline above. Because of its singular prominence in many restricted settings, the association seems to warrant its own ichnofacies designation. Conversoly, the less restrictive the environment at a given site, the less distinctive the ichnofacies as a separate entity. In numerous places the ichnofacies is hardly dis- ccemible in the broad transition from the Skolithos or Cruziana to the Nereites ichnofacies, especially on unstable ancient slopes originally subject to turbidity flows or swept by shelf-edge or contour currents. Whatever the major environmental implications of the Zoophycos ichnofacies and its variants, the final word is not yet in. The most important factors in the distribution of the animal, in addition to its own opportunism, evidently include water depth, depth of burrowing, sediment cohesive- ness, and interstitial or bottom-water oxygen levels (Kotake, 1989, 1991; Olivero and Gaillard, 1996; and Uchman and Demirean, 1999). Stressed quiet-water environments, particularly those exhibiting anoxia, seem to be the primary common denominator, even though the animal itself was cosmopolitan. Nereites Ichnofacies In most respects, bathymetric implications of the Nereites ichnofacies (Fig. 30, Table 8) are less ‘equivocal than those of any other recurrent ichnofacies. Although numerous trace fossils otherwise typi- cal of shallow-water deposits occasionally range into deep-sea deposits. the reverse is not ordinarily true. In addition to water depth, turbidite deposition strongly influenced original environmental inter- pretations of the Nereites association. However, depth- and energy-related variables seem to be more important than turbidite deposition per se (Crimes et al., 1981, Leszczynski and Seilacher, 1991; Miller, 1993). For example, the trace assemblage persists today on distant abyssal plains essentially beyond reach of turbidity currents but is absent among well-developed, shallow-water turhidite sequences. Nevertheless, most Nervites associations studied to date occur in turbidites, probably because the stratigraphic record of deep-water deposits examined in this context mainly represent subsid- ing basins or subduction zones rather than the broad abyssal plain. Associated sediments thus may consist of virtually any lithology, except that the ratio of turbidite sand to hemipelagic or pelagic mud tends to diminish toward distal extremities of the deposit, and carbonates become 49ZOOPHYCOS ICHNOFACIES 1. Zoophycos 2. Zoophycos. 3. Spirophyton - Low diversity, though individual traces may be abundant. - Simple to moderately complex, efficiently executed grazing and feeding structures produced by deposit feeders. - Horizontal to gently inclined spreiten structures distributed in delicate sheets, ribbons, lobes, or spirals (flattened forms of Res Zoophycos or, in pelitic sediments, ao. Phycosiphon). Zoophycos, Upp Cardium Form: Cretaceous| in, Alberta Figure 28, Schematic of the Zooplyeos ichnofacies (moditied from Pemberton et ul. 1992), 50ZOOPHYCOS ICHNOFACIES Characteristic Forms of Zoophycos Figure 29, Characteristic forms of Zoophycos that are prevelent in the Zoophycos ichnofacies, 31+ High diversity but low abundance. + Complex horizontal grazing traces and patterned feeding/dwelling structures reflecting highly organized efficient behaviour. + Spreite are typically nearly planar, although Zoophycos forms are spiraled, multilobed and complex. NEREITES I1CHNOFACIES 1. Spirorhaphe 2. Urohelminthoida 3. Lorenzinia 4. Megagrapton 5. Paleodictyon 6. Nereites 7. Nereites + Numerous crawling and/or grazing traces and sinuous fecal castings (Helminthoida, Cosmorhaphe) that are mostly intrastratal. + Structures produced by deposit feeders and scavengers. * Structures associated with trapping or farming microbes within permanent open domiciles (Paleodictyon, Megagrapton). Figure 30, Schematic of the Nerites ichnofacies (modified from Pemberton et al. 1992),increasingly scarce as the calcite compensation depth is approached. Bouma sequences or modified Sequences are common locally, and physical sedimentary sinictires may include flute, groove, ounce, and load casts as well as prod marks, flame structures, and linguoid and other ripples. Animals exploiting lower bathyal to abyssal environments have two major concems: 1) scarcity of food, relative to more abundant supplies in shallower settings, and 2) periodic disruption by strong down- canyon bottom currents or actual turbidity currents. In response to the last factor. and over long spans of geologic time, the overall community ultimately developed two component parts: pre-turbidite and post- turbidite associations, as documented by their respective traces (Leszczynski and Seilacher, 1991; Miller, 1993; Tunis and Uchman, 1996). The pre-turbiite resident association is characteristic of quiet, normal conditions and is dominant wherever the substrate is free of the influence of turbidity currents. It tends to be overwhelmed of eliminated by severe erosion or turbulence, however, and is replaced by the post- turbidite association after cessation of the turbidity current. As conditions then revert to a normal, low-energy setting, the pre-turbidite association gradually reestablishes itself. Pre-turbidite animals thus comprise a stable, persistent community welll adapted to quiet conditions, detived mainly from original early-Paleozoic colonizers of the deep-sea floor. In contrast, post-turbidite animals rep- resent a more opportunistic, less stable community better adapted to turbidite colonization, derived ‘mainly from subsequent evolutionary immigrants from shallower waters (Frey and Seilacher, 1980) In addition to pre-and postturbidite associations, numerous turbidites display ichnologic gradients along depositional dips (Crimes et al, 1981). Where strong bottom currents issue from submarine canyons or course along fan channels, components of the Skolithos ichnofacies may be present. Otherwise, proximal parts of turbidites may be characterised by rosetted or radiating traces and gently meandering forms of Scolicia. Medial parts may be indicated by spiraled or tightly meandering traces, while pattemed networks typify distal parts although other traces are generally present (Fig. 31). Zoophycos is common locally in various settings, but it tends to be multilobed and in places is more complex than in the Zoophycos _ichnofacies. Finally, the Nereites ichnofacies is recognized via sedimentary coring in ancient, unconsolidated, well-bedded fine sediments. including distal turbidites and pelagic rhythmites, of the pres- cent ocean basins, However, the association per se, if present, tends not to be preserved on great expanses of abyssal plain, where sedimentation and bioturbation are more or less constant rather than episodNEREITES |CHNOFACIES Characteristic Ichnogenera oS Paleodictyon + REE Tre Deep Sea Cores: Zoophycas, Chondrites, and Planolites Nereites Helminthoida Cosmorhaphe Figure 31. Characteristic trace fossils commonly associatd with the Nereites ichnofacies, 34AN INTEGRATED ICHNOLOGICAL-SEDIMENTOLOGICAL MODEL OF SHORFFACE SUCCESSIONS AND SHOREFACE VARIABILITY INTRODUCTION ‘The shoreface consists of a seaward -sloping sediment wedge extending from the low tide mark, gener- ally to the fairweather (minimum) wave base, comesponding to approximately 10-20 m of water depth. (Fig. 32) With greater storm-domination, effective wave base is lowered to depths approaching storm weather wave base. As a result, erosively amalgamated tempestites of “lower shoreface” alfinity may actually be deposited well below fairweather wave base. The shoreface setting is dominated by wave ‘energy and, as a result of decreasing wave interaction with the substrate in a seaward direction, shows a pronounced basinward fining. The shoreface is classically divided into three suben vironments (from sea- ward to landward): the lower, the middle and the upper shoreface. The boundaries between these are not always clearly defined (Reinson, 1984). Although the large-scale regional context may vary, the specific subenvironments (Fig.33) of the shoreface are not significantly different whether they occur as part of a strandplain, barrier island or wave/storm-dominated delta (free from interference from active distributary channels) Figure 32, Modern coastal environments. A. Beach ridge, Atlantic coast; B, Coast of South Carolina showing multiple tidal inlets, 35Sapelo Island, Georgia Figure 33, Sapelo Island, Georgia showing the multitude of coastal depositional environments, 56‘The shoreface grades distally into offshore units and landward into foreshore deposits. A complete showeface progradational succession reflects offshore to foreshore environments; consequently, these adja cent environments bear inclusion in any discussion of shoreface deposits per se. There are several excellent outcrop examples of Cretaceous shoreface deposits in the Wester Inte- rior Seaway, including the Turonian Cardium Formation, the Campanian Virgelle Member of the Milk River Formation, the Campanian Appaloosa Sandstone of the Rearpaw-Horseshoe Canyon Transition, and Maastrichtian Blood Reserve Sandstone, of southern Alberta. The Late Albian-Cenomanian Bootlegger Member of the Blackleaf Formation and Campanian Eagle Sandstone of Montana, as well as the Campan- tan Star Point and the Blackhawk formations of the Book Cliffs, Utah are also examples. In addition, there exist large numbers of excellent shoreface successions preserved within the subsurface of Alberta, such as the Albian Bluesky Formation, the Albian Cadotte Member of the Peace River Formation, the Albian Viking Formation and the Turonian Cardium Formation. The integration ofthe sedimentology and ichnology within these deposits affords the opportunity to characterise the facies and facies successions (Pig. 34), and to explain the observed facies vatiabilty SHOREFACE SUBENVIRONMENTS ‘The shoreface and adjacent environments can easily be grouped into three depositional complexes: the offshore complex, the lower-middle shoreface complex, and the upper shoreface-foreshore complex. Each complex corresponds to discrete depositional parameters, both with respect to physical processes and biogenic structures. Coastal morphodynamies are subject to a wide range of variables depending on whether the coast is prograding or in transgression. For example. estuaries are generally a function of transgression while riverine-dominated deltas and strandplains are more apt to develop along prograding coastlines (Fig. 35). Offshore Complex ‘The offshore zone is regarded to lie below minimum (Fairweather) wave base and above maximum (storm-weather) wave base. The distribution of trace fossils in this zone is related to the food resources paradigm (Fig. 36) where the Cruziana ichnofacies is prevalent in zones where food particles are depos- ited. This results in increased inter-specific competition and a corresponding increase in specialized feed ing strategies. In shallower water food particles are suspended and the animals can remain stationary in burrows while the food comes to them (Fig. 37). The offshore is commonly subdivided into a lower ‘and upper subzone (e.g. Pemberton et al., 1992a). Howard and Frey (1984) subdivided the offshore into “lower”, “middle” and “upper” offshore subzones, based on outcrops within the Book Cliffs of Utah; in many respects, their “lower” offshore is more reminiscent of shelfal silty shales. Lower Offshore ‘The lower offshore, as defined here, comprises dark silty shales, commonly thoroughly homogenized by biogenic reworking. Fine to very fine sandstone beds are present, though in low abundance. Thin beds are predominantly intensely bioturbated, although locally, remnant undulatory parallel laminae are preserved. Thicker beds tend to show sharp, erosive bases, with moderately- to well-developed parallel amination. Mast burrowing i restricted ta the taps af beds. “These preserved beds are interpreted as distal tempestites. 37Shoreface Model Cretaceous Western Interior Dominant Structures Ichnological Assemblages TT F Psilonichnus atin |_Backshore t Ichnofacies High t Tide | 2 | Foresnow [P| Macaronichnas Es | 2 {Assemblage __ Law THE (EE) cee] i i $3| Shoreface ' E i Skolithos : + =| Migae [fe lehnofacies +4/- Burrowed Sst EB | Shoreface THE TER) comm [Pt Pen S,WaveRipples 2423 Jey | Shoreface | 2 | | Cruciana pair weather -Burrowed Muddy j [8% |33 ae "wave base Sst iy ie H s y Archetypal tT Upper | iF Cruziana Burrowed Sandy Mast. { otthee ft E <3 wHes | Le Sg -Wave Ripples | 5 if ee j : cs | | Distal 1 | 22] offshore |} Cruziana 41-Wave Ripples | if ' ' FE ' \ g ! i i ~ i z Storm i 3 ave base Burowed Mdst | Shett teviareHCs | Zoophycos 4Siommetndiced Tehnofacies Wave Ripples? ' Dominant Processes +» Dominant Behaviors Subordinate Processes = = Subordinate Behaviors Minor Processes Minor Behaviors Figure 34. Shoreface model showing the distribution of ichnological assemblages based on examples from the Cretaceous Western Interior. 58Increasing Tidal Power ——— Inereasing Wave Power ———> Transgressive ian Prograding Figure 35. Differences between the coastal dnamics of prograding and transgressing shorelines. (Modified after Boyd et al, 1992) Food Resources Paradigm —=zaiX*—aez———E— FOOD PARTICLES SUSPENDED SARS FOOD PARTICLES DEPOSITE! ~<—— skourHos —__» < — cruziana — a ire 36. Food Resource paradigm for the Skolithos - Cruziana ichnofacies that are characteristic of shoreface environments. 59‘sums sorjauoys [ro1dAi ur Soronyouyo! SuLIHWL ma] |YS SNstEarONNYD Jo UONNSO “LE aun ‘sOVIERIOHS aaa | xaMo1 aQVEROUS waa agons “OE‘The ichnological suite may be regarded as a “distal” or “outer” Craziana assemblage, incorporat- ing a relatively low diversity of both deposit feeding and tw a lesser degree, gruzing/foraging structures, ‘Common ichnogeneta include sina, thit-walled Schaubcytindrichnus, Planolites, Hetminshopsts, Phyco- siphon, Chondtites, Zoophycos and Thalassinoides (Fig. 38). Less common forms include small Astero- soma and rave Paluevphycus, These facies ure relatively common in the lower cycles of the Viking For- imation, in the Cardium Formation. Howard and Frey (1984) also noted the presence of Thalassinoides suevicus, Ancorichnus capronus and Ophiomorpha annulata in the Book Cliffs of Utah (their “middle offshore). Frey (1990) observed the additional presence of Rosselia socialis and Cylindrichnus concen rricus within the lower offshore deposits within the Spring Canyon Member of the Blackhawk Forma- tion, Coal Creek Canyon, Utah. A. capronus, 0. annulata, R. Socialis and C, concentricus reflect bur- rows initially established in tempestites, rather than in the fairweather deposits of the lower offshore. In Drumheller, the Upper Cretaceous Horseshoe Canyon Formation 1s dominated by a Helminthopsis rich assemblage (Fig. 39). Lower Offshore Lower offshore assemblage in the Viking Formation of Alberta is dominated by the grazing structures Helminthopsis, Phycosiphon and Zoophycos Figure 38. Characteristic trace fossils of the lower offshore zone. ‘The lower offshore is dominated by diminutive deposit feeding structures and grazing/foraging struc- tures which typically oceur in large numbers. Most of these structures show minimal vertical penetra tion of the bed. Under such circumstances, distal storm beds may show little biogenic reworking, except near their tops, even if the tempestite is relatively thin. Since these distal storm beds are deposited well below fairweather wave base, physical processes during non-storm periods are not competent 9 modily them, and hence, such beds have a high preservation potential (Dott, 1983, 1988; Whealcrofl, 1990). The fairweather deposits of the lower offshore commonly display high abundance, high diversity and fairly uniform distribution of ichnogenera conesponding (o a distal Cruziana ichnotucies 6Offshore Environments: Horseshoe Canyon Fm ower'sonerace ” ‘TE vroxmar 1 OFFSHORE | DISTAL ‘ OFFSHORE | TRANSGRESSIVE j sromerace sanw Proximal Offshore Association Figure 39. Recurring associations of trace fossils in distal and proximal offshore environments, Upper Cretaccous Horseshoc Canyon Formation, Alberta, 62Upper Offshore ‘The upper olfshore is considerably more variable than the lower offshore, attributable to the greater ‘degree of storm and wave interaction with the substrate. The interval commonly consists of intensely bio turbated silty and very fine. to fine-grained sandy shales, interbedded with thin, very fine to fine grained sandstone beds. The sandstones range from virtual biogenic homogenization to erosively based beds pos sessing remnant low angle (15°) parallel laminae and oscillation or combined flow ripple laminae, These beds are interpreted as storm-generated in origin, with local preservation of waning flow deposits, Less intensely burrowed upper offshore intervals may show more regular interbedding of shales and storm- ‘generated sandstones, probably reflecting greater storm domination, greater storm frequency, andlor high sedimentation rates. This has been referred to as the “offshore transition”. More commonly, tempestites show moderate to intense degrees of burrowing near their tops. ‘The ichnological suite may be regarded as a diverse Cruciana assemblage. Some grazing/foraging structures such as Helminthopsis, Phycosiphon and Zoophycos may also be included. The bulk of the suite consists of ichnogenera such as Planolites, Teichichnus, Schaubeylindrichnus, Palaeophycus, Thalassi noides, Cylindrichnus, Chondrites, Siphonichnus, Lockeia, small Asterosoma, small Rosselia, and rare Ophiomorpha and Rhizocorallium (Fig. 40). Rare suspension feeding structures, such as Skolithos, Are nicolites and Diplocraterion, may also be present in sandicr fairweather units, although they are also asso- ciated with tempestites. ‘Tempestites may contain fugichnia as well. This facics is well developed in the ‘Viking Formation, the Cardium Formation and within the Aberdeen Member of the Blackhawk Forma- tion in the Book Cliffs, Utah. Within the Book Cliffs, Ancorichnus capronus , the crawling structure Scolicia (Howard and Frey, 1984; Frey, 1990) and Cosmorhaphe have also been noted. Some of the sandstone beds may also contain Ophiomorpha nodosa and Skolithos linearis, probably reflecting oppor tunistic colonisation of distal storm beds. In the Upper Cretaccous Horseshoe Canyon Formation the trace uite is dominated by Teichichnus, Chondrites, and Thalassinoides (Pig . 39) ‘As in the lower offshore silty shales, burrowing in the upper offshore is typically intense, with a high Upper Offshore fossil Upper offshore assemblage in the Viking Fm. ineludes; Scolicia, Thalassinoides, Teichichnus, Phycosiphon, Helminthopsis, Planolites, and Schaubeylindrichnus Figuro 40. Characteristic trace fossils from upper offshore zones, 63diversity and uniform distribution of ichnogenera, This suite displays an overwhelming dominance of deposit feeding behaviour over grazing/foraging behaviour. The introduction of small numbers of suspei= sion feeding structures within fairweather deposits, as well as those associated with tempestites, is distin tive, The ichnological assemblage is characteristic of the Cruziana ichnofacies. Distal tempestites depos- ited in the upper offshore typically display a greater degree of biogenic reworking than the thinner storm beds of the lower offshore. This reflects the greater vertical penetration ofthe substrate by the more robust benthic organisms that inhabit the upper offshove (Disjes and Herwweck, 1973) ‘The lower and upper offshore are commonly dominated by fairweather deposition, although on strongly storm-dominated shorefaces, the offshore complex may consist of shaly zones interbedded with relatively unburrowed, thin tempestites. Fairweather deposits are characterised by abundant burrowing, manifest by high diversity and uniform distribution of ichnogenera, ‘The suites lack domination by ind Vidual forms. The ichnological assemblages for this depositional complex display an equilibrium or K-selected behaviour, characteristic of fully marine, unstressed settings (Pianka, 1970; Jumars, 1993). ‘Some of the sandstone beds may also contain Ophiomorpha nodosa and Skolithos linearis, probably reflecting opportunistic colonisation of distal storm beds. The distribution of ichnofossils record the typi cal character of ecological stresses imparted by episodic depositional events (cf, Pemberton et al., 1992b; Pemberton and MucEachem, 1997). Escape traces record the disturbance by animals entrained within, or buried by the flow, as they attempted to reach the sediment-water interface. Trace fossils ike Helminthop- sis, Phycosiphon, Planolites, and Teichichnus record the re-establishment of grazing and deposit-feeding una following the event, and typically occur near the tops of beds or within the mudstones capping the ‘coarser clastic units. The general absence of suspension-feeding structures suggests that most sandstone beds were quickly buried by silts and muds following the depositional event, precluding colonisation by “organisms that favour sandy substrates, ‘The primary physical structures of the sandstones and siltstones consistent with a distal tempestite (storm-bed) interpretation, in a setting lying seaward of fair-weather wave base (cf, Pemberton and Frey, 1984: Aigner, 1985; Saunders, 10R6; Seilacher and Aigner, 1992; Pemberton and MacEachemn, 1997). Many of the tempestite beds record the waning stages of storm action on the substrate, characterised by normal grading, oscillation ripple laminated tops, and suspension fall-out of silts and muds following storm abatement. Lower-Middle Shoreface Complex Lower Shoreface ‘The lower shoreface proper begins at the lower limit of fairweather (minimum) wave base, but where offshore processes continue to operate (Reinson, 1984). Under progressive storm-domination, facies of lower shoreface affinity are deposited basinward of fairweather wave base, because effective wave base is significantly lowered. Wave energy is the dominant physical process controlling deposition of the interval and most structures reflect storm deposition, including hummocky cross-stratification (HCS) (cf., Dott and Bourgeois, 1982; Duke, 1985), rarer swaley cross-stratfication (SCS) (¢f,, Leckie and Walker, 1982) and quasi-planar lamination (QPL) (cf, Amott, 1993a). Intervals not dominated by erosional amalgama- tion may show waning stage oscillation or combined flow ripple lamination capping these structures. Fairweather-generated oscillation ripples may be present but are relatively uncommon, The intensity of 4burrowing is highly variable, depending upon the degree of storm dominance. High intensity and high fre- queney storm events favour minimal preserved burrowing, while low intensity and infrequent storm events lead to thorough homogenization of the tempestites and accumulation of thick fairweather intervals. Biogenic features are dominated by the Cruziana ichnofacies, with considerable contributions from the Skolithos ichnofacies. Rarer grazing/foraging structures, largely confined to muddy zones (Fig. 41) are also locally present Deposit feeder and mobile carnivore structures of the Cruziana ichnofacies include robust Rosselia, Asterosoma, Teichichnus, Cylindrichnus, Schaubeylindrichnus, Schaubcylin- drichnus, Planolites, Thalassinoides, Ancorichnus, Siphonichnus, horizontal Ophiomorpha nodosa, O. irregulaire, rate O. annutata, Chondrites and Rhizocoratlium (Fig. 42). Pemberton and Frey (1984) also noted the presence of Gyrachorte, Taenidium and Phoebichnus within outcrop examples of the Cardium Formation, Seebe, Alberta. Grazing/foraging structures include Helminthopsis, Cosmorhaphe, Zoophy- cos, Scolicia and Aulichnites. Phycosiphon are present within storm-generated sandstones, but are gener ally rare, Elements of the Skolithos ichnofacies include Skolithos, Conichnus, rare Diplocraterion, rare Bergaueria, and the passive carnivore structure Palaeophycus. In addition, escape traces (fugichnia) and those of various opportunistic organisms may also be common, generally colonising the tops of storm beds. ‘The lower shoreface trace fossil (Fig. 43) assemblage shows a dominance of deposit feeding behav- Figure 41, Variation in trace fossil associations seen in the lower shoreface-offshore succession in the Upper Cretaccous Horseshore Canyon Formation, Alberta, 65.Lower Shoreface Assemblage Boxwork Asterosoma & Ophiomorpha Chondrites Figure 42, Characteristic trace fossils from lower shoreface zones. Diplocraterion Ophiomorha ‘our, with significant contributions of suspension feeding structures, suggesting that persistent wave shoal ing (i.e. above fairweather wave base) is important in displacing suspended mud offshore und suspending food particles above the bed, and shifting sand at the sediment-water interface. ‘The observed fairweather trace fossil suite corresponds to a proximal Cruziana assemblage (Fig. 16). In the Horseshoe Canyon Formation Saunders (1986) was able to recognize distinct assemblages that characterized the distal lower shoreface (Fig. 44) and the proximal lower shoreface (Fig. 45) Storm events impart considerable control on the character of the lower shoreface. Storms tend to uproot, destroy and/or bury resident (Fairweather) benthic communities (Pemberton and Frey, 1981; Froy, 1990; Pemberton ef ai., 1992c). During post-storm recovery, initial colonisation of the tompestite gener ally records opportunistic dwelling/suspension feeding structures, which do not typically reflect the origi nal resident community. Continued fairweather conditions may see a return to the original trace fossil suite (see discussion in the section on storms). Depending upon the degree of storm dominance, the pre served record of the lower shoreface can be quite variable. Some thin tempestites may escape thorough biogenic reworking (ef: Wheateroft, 1990) under condi tions of high frequency storms, moderate to low energy storm conditions and high sedimentation rates, Saunders and) Pemberton (1086) and Saunders (1989) noted that thin storm beds common in the lower shareface deposits of the Appaloosa sandstone of the Bearpaw-Horseshoe Canyon Formation transition, Alberta, permitted Rhizocarallium jenense, Teichichnus rectus and Rosselia socialis (Fig. 46) to re- equilibrate their burraws to each successive sediment-water interface. Rosselia socialis reflects this re- equilibration by vertically stacking their burrows (Fig. 47). spectacular example of this occurs in core ff the Alhian Grand Rapids Formation of Alberta, where stacked Rosselia burrows record repeated post- storm re-establishment at least seven times during the life of a single tracemaker (Fig. 47). 66Lower Shoreface Assemblage: Horseshoe Canyon Formation Figure 43. Schematic diagram of the lower shoreface trace fossil assemblage seen in the Upper Cretaceous Formation, Alberta, oStorm Dominated Distal Lower Shoreface 1. Macronichnus 2. Rosselia 3. Rhizocorallium 4. Teichichnus 5. Ophiomorpha Figure 44, Schematic diagram showing the trace fossil association in the storm-dominated distal lower shore of the Upper Cretaceous Horseshoe Canyon Formation, Alberta, Middle Shoreface ‘The middle shoreface extends over the zone of shoaling and initial breaking of waves (Reinson, 1984), and is characterised by high wave energy. Longshore bars are commonly present near the upper portion of the midale shoreface within intermediate (barred) states of shoreface morphodynamics (Wright ef al., 1979; Thom et al., 1986). Preserved deposits of longshore bars are rarely intercalated within middle shoreface intervals, since under fairweather conditions, bars migrate landward and weld to the foreshore (Davidson-Amott and Greenwood. 1976; Davis, 1978). ‘The sandstones tend to he well-sorted, well ‘winnowed, and medium. to fine-grained, with only minor shale, silt and shell layers. Preserved structures are predominantly low angle wedge-shaped sets of parallel laminae, constituting SCS and lesser LICS, or QPL. Oscillation ripple laminae, combined flow ripple laminae, and rarer trough cross-stratification are locally present. Storms have a strong influence on the middle shoretace, greater even than in the lower shoreface, and may constitute much of the depositional record of this zone. The SCS or QPL tempestite beds are typically erosionally amalgamated, becoming progressively more cannibalistic upwards (Aigner and Reineck, 1982) with increasing shallowing and associated enhancement of storm-induced scouring. As a result, the degree of bioturbation is highly variable. 68Proximal Lower Shoreface 1 Maceronichnus 2 Teichnichnus 3 Placophycus 4. Asterosoma 5. 6 Ophiomorph 7. yfindrshnus 8, Gyrochorte 9 Rentia 10 Figure 45. Schematic diagram showing the characteristic trace fossil assemblage of the proximal lower shoreace of the Upper Cretaceous Horseshoe Canyon Formation, Alberta Figure 46. Rosselia fivn lower shoreface zones. A. Lower Cretaceous Viking Formation; B and C. Lower Cretaceous Cadotte Member. 0Lower Shoreface: Rosselia Assemblage Lower Cretaceous Grand Rapids Rosselia Modern Tracemaker: Formation, Alberta Terebellid Polychaete = Double Mud Balls Cross Section In situ Specimen Upper Cretaceous Horsesoe Canyon Fin., Drumheller Figure 47. Rosselia trom lower shoretace zones. 70Ichnologically, the middle shoreface corresponds to a dominance of suspension feeding behaviours (con- stituting the Skotizhos ichnofacies; Figure 65) over predominantly deposit feeding behaviours (the Cruz- ana ichnofacics), although the latter traces continue to persist as subordinate elements (Howard, 1972, 1975). The bulk of the assemblage consists of Skolithos, Conichnus, Diplocraterion, Ophiomorphe (vom monly vertical components), Arenicolites, Dergaueria and Pulueophycus (Fig. 97). Howaud and Frey (1984) also noted Medousichnus loculatus within intervals of the Book Cliffs, Utah (inet “upper” shore face). Howard (1971a,b; 1975) and others have noted that Ophiomorpha systems tend wo shift from hori~ zontal to vertical in orientation as energy levels inctease. This shift in orientation may accompany the transition from the lower to middle shoreface, Fugichnia may be common, particularly associated with tempesites. Middle Shoreface Rascelia & Palaeophyeus Ophiomorpha Figure 48. Characteristic trace fossils from middle shoreface zones. Cruziuna ichnofacies elements include vobust Rosvelia (R. socialis Uomiinates; R. chonoides is also recognised in the Book Cliffs), Asterosuma, Schuubcylindrichnus, Planolites, Teichichnus, Cylindrich- nus, rare Rhicocorallium, and locally abundant, though generally rare Chondrites. Chondrites is largely restricted to muddy zones. Fiey (1990) noted that Cylindrichnus and Rosselia become more steeply inclined as energy levels inerewse. This enbancement in inclination may also accompany the transition fiom the lower to tiiddle shoreface, ing the trend observed for Ophiomorpha. ‘As in the lower shoreface, variability in storm intensity and frequency strongly affects the character of the middle shuteface, buth sedimentologically and ichnologically. As the storm effects become more pronounced, the diversity of deposit feeding structures declines markedly. This appears to be attribut- able to the abundance of well-winnowed sand and the general paucity of deposited food for the Cruztana ilmofucies travemakers to feed upon. Notable exceptions include Rosselia soctalis, Asterosoma and, t0 a lesser degree, Cyndrichnus, which may persist upwards well into the middle shoreface. ‘The preservation potential of biogenic structures in the middle shoreface Is intrinsically associated with the character of tempestite accumulation. In storm-dominated conditions, as tempestites become aprogressively more cannibalistic, only deeply penetrating structures, particularly Ophiomorpha nodosa and fugichnia, may he recorded. It is nat uncommon for storm-dominated middle shoreface sandstones to show little or no burrowing. Unfortunately, discerning whether their absence corresponds to high intensity storms (greater erosional amalgamation) or to high frequency storms (minimal time for re-establishment of benthic communities) is virtually impossible. Like the offshore complex. the lower to middle shoreface complex consists of sediments accumulated Minor Behaviors Minor Processes + Many tube dwellers ure passive carnivores rather than suspension feeders Fair weather suites are subenvironmental indicators, not event suites Figure 58. Modified shoreface model for high energy shorefaces. 81High-energy Low-energy Sheltered Beach Beach Sandflat € ree Ccocve a el ti nn — — i es cen High Energy Shoreface Low Energy Shoreface 1. Dominanted hy horizontal forms 1. Dominated by vertical forms 2. Dominated by feeding burrows 2. Dominated by dwelling burrows 3. Dominated by interstratal forms 3. Domonated by interfacial forms 4, Dominant ichnogenera 4, Dominant ichnogenera Planolites and Macaronichnus Skolithos and Ophiomorpha Figure $9. Comparison ofthe ichnology of high energy and low energy shoreface successions. toe-of the-beach position. Only in isolated examples does the zone ever actually transcend into overlying swash-laminated foreshore sandstones. ‘Ihis opposite pattem of recurrence has also been recognised from Cretaceous to Tertiary deposits ofthe Pacific west coast (Hunter, 1980; Clifton, 1981, 1988; Hunter et a, 1984: Clifton and Hunter, 1987; Leithold and Bourgeois, 1989; Saunders, 1989; 0) nd Masuda 1992). In the normal Scilacherian ichnofacies scheme the nearshore zone should be dominated by the dwell- ing burrows of suspension feeders that characterize the Skolithos ichnofacies (see Fig. 20). The Mac- aronichnus assemblage, however, seems to characterize higher energy shoreface environments where domiciles can not be established because of shifting substrates and filter feeding is precluded because of intense wave action (Fig, 58) These shorefaces are instead dominated by horizontal forms that are feeding burrows that form interstratally (Fig, 59). SHOREFACE VARIABILITY Except for textural variations, the upper shoreface-foreshore complex remains relatively consistent in character over a wide range of shoreface morphodynamic states. There are few facies clearly indica- tive of barred conditions, since most subenvironments show a low preservation potential (Greenwood and Mittler, 1985), and those that are preserved closely resemble the facies of barred systems, As well, the preserved record of the upper shoreface foreshore complex reflects fairweather conditions, and is charac= terised by ichnogenera of the Skolithos ichnofacies (Fig. 34). In contrast, the lower-middle shoreface 82complex reflects the greatest degree of facies variability, both sedimentologically and ichnologically. This portion of the shoreface may range from virlually unburrowed HCS, SCS or QPL sandstones t0 thor- ‘oughly bioturbated muddy sandstones, since both storm events and fairweather conditions may produce deposits. Although the offshore also accumulates sediment during both storm and fairweather conditions, the complex is generally fairweather dominated. Additionally, the trace fossil suite occurs in the Cruziana ichnofacies (Fig, 34). The lower-middle shoreface complex straddles the transition from the Cruziana ichnofacies to the Skolithas ichnofacies, enhancing its vanability. Variability in the preserved trace fossil record may locally reflect a combination of water turbidity, water salinity, fluctuatingfepisodic depositional rates, and storm energy conditions. Reduced oxygenation also has a profound effect upon the trace fossil assemblage (cf. Bromley and Ekdale, 1984; Raychaudhuri and Pemberton, 1992), but does not appear to exert a control on the sedimentary facies themselves. The close association that appears to exist between the lithofacies character and the observed trace fossil suite suggests that both occur in response to the same overriding mechanism. ‘The effects of enhanced water turbidity, as might be experienced near a river entering the coast, or settings proximal to a distributary channel on a storm-dominated delta, will typically be reflected by the virtual absence of suspension feeding structures within the sandstones. High water turbidity interferes with the efficiency of the organism's suspension feeding apparatus (Rhoads and Young, 1971). Water turbidity does not appear to affect predaceous organisms or most deposit feeders, and their traces may continue to occur in large numbers. As well, the nature of physical sedimentation need not necessarily change in response to water turbidity, if deposition occurs above fairweather wave base, since constant wave agitation of the bed keeps mud in suspension and moves it offshore. Fluctuations in water salinity have a marked effect on the entite tave fossil sssemblage, extending from he offshore, landward wowards the foreshore. Decreasing or (more rarely) increasing salinity is largely manifest by the disappearance of specialized feeding behaviours and the corresponding dominance of trophic generalists (cf: Pemberton and Wightman, 1992; Ranger and Pemberton, 1992). These effects are not restricted to the lower and ‘middle shoreface and should show a recognisable pattern across the entire depositional profile. In river- dominated deltaic setings, water turbidity and salinity fluctuations may go hand in hand, severely limiting the diversity and abundance of the trace fossil suite. Other ecological stresses include an oxygen stress, a sedimentation stress and a geotechnical stress (Fig. 60). Such a harsh biotie environment results in limited trace fossil suite. Differences between storm/wave-dominated deltas and storm/wave-dominated strandlines, on the other hand, may be exceedingly subtle. In the Lower Cretaceous Harmon Shale in Alberta, Mostow and Pemberton (1988) suggested that prodelta to distal delta front accumulations can be recognized by the low-diversty trace fossil suite that 1vas attributed to harsh ecological conditions related to increased water turbidity and fluctuating rates of deposition from suspension. They interpreted the event beds themselves to be flood-induced, although the primary sedimentary structures are more typical of storm-induced deposition than of delta front sediment gravity flows. Nonetheless, there isa close genetic association between storm events and enhanced flood discharge through rivers and distributaries reaching the coast, since storms are commonly associated with high rates of precipitation, This supports an interpretation of rapid deposition of post-storm suspended fines, observed in other deltaic settings (Allen, 1982; Massari and Parea, 1988; Leithold, 1989). Low bur- rowing intensities may also reflect this rapid, though sporadic, accumulation of fines. Leithold (1989) studied the shelf lying offshore of the Eel River, northern California, and compared it to the analogous Pleistocene Rio Dell Formation of northem California. She concluded that river flood events play an 83Deltaic Environments Represents one of the most biologically stressed depositional systems, the stresses include a salinity stress, an oxygen stress, a turbidity stress, a sedimentation stress and a geotechnical stress Figure 60. Characteristics of deltaic environments. important role in fine sediment accumulation in the offshore, even along high-energy, storm-ominated coasts. The high sedimentation rates and rapid suspension fallout of fines in the upper Harmon Member ae consistent with prodelta to distal delta front environments, suggesting that the Harmon/Cadoste transi- tion may actually represent accumulation in a rapidly prograding. strongly wavelstorm-dominated pro- delta setting, rather than in the offshore of a strandlines system sensu stricto. It is interesting that these transitional offshore sediments are remarkably similar to the distal facies in Allomember D and F. of the Dunvegan Formation of Alberta (Bhattacharya and Walker. 1991; Gingras et al., 1998, Coates and MacEachern, 1999) as well as in the basal Belly River Formation (Lerand and Oliver, 1975; Coates and MacEachern, 1999), both interpreted as wave-dominated deltaic settings. Depositional rates have an important effect on the degree of bioturbation, but unless associated with episodic events, do not appear to impose a control on the ethology of the organisms and thus, not upon the resultant traces. Variations in sedimentation rates are typically coupled with other environmental controls. For example, slow sedimentation rates coupled with well oxygenated, fully marine conditions favour thor- ‘ough biogenic reworking of the sediment, whereas the same rate of deposition, associated with anaerobic or dysaerobic, stratified and stagnant water, favour virtually unburrowed deposits. Likewise, enhanced depositional rates may merely favour decreased bioturbation, but if coupled with increased water turbid- ity and reduced salinity, as might be produced in ariver-dominated delta, may produce completely unbur- rowed deposits. saStorms, and to a lesser degree, fairweather waves are the dominant physical processes operating on the lower and middle shoreface. Since these deposits may constitute the bulk of the ancient record of this portion of the shoreface (Kumar and Sanders, 1976; Aigner and Seilacher, 1982; Reinson, 1984; Swift et dl., 1985; Elliott, 1986), it follows that much of the observed trace fossil variability may be attributable to variability in the predominance of storm activity. Fluctuations in sedimentation rate are intimately associ- ated with the episodic nature of storm deposition and this appears to control the ultimate character of the preserved lower and riddle shoretace deposits. In general. it is possible to identify three principal “types” of lower and middle shoreface deposits: those that are strongly storm-dominated (“high energy”), those that are moderately affected by storms (intermediate energy”) and those are only weakly affected by storms, or dominated by fairweather depo- sition (“low energy”). A complete intergradation appears to exist, both along depositional strike, depend- ing on the regional paleogeography, and vertically through the facies succession, related to shallowing, and increasing effectiveness of storm action an the hed. Much of the confusion that exists in defining the character of the lower and middle shoreface, as well in determining of the boundaries between the two subenvironments, appears to revolve around the variability imposed by the degree of storm-dominance and preservability of the fairweather deposits. SUMMARY ‘The integration of ichnology and sedimentology provides a strong depositional model tor foreshore to offshore transitions, whether they occur on strandplain, barrier istands or even wave/storm-dominated deltas, removed from the interference of distributary channels. The shoreface zone can be grouped into three main depositional complexes, each of which respond to a different set of physical parameters and displays discrete ichnological characteristics. Within each depositional complex, the subenvironments ‘can be reasonably differentiated through a combination of physical and biogenic features, with the excep- tion of the lower and middle shoreface ‘The offshore complex is typically dominated by Fairweather conditions and is characterised by a Cru- ziana ichnofacies. ‘The suite displays a predominance of deposit feeding structures with subordinate ¢grazing/foraging structures, and even rarer suspension feeding structures. Storm effects are variable but ineffective in removing the fairweather suite except under high degrees of storm domination. The offshore typically displays a K-selected equilibrium suite of trace fossils, with variable but non-dominating over- print by opportunistic (r-selected) behaviours related to tempestite colonisation, ‘The lower-middle shoreface complex is strongly affected by both storm and fairweather conditions, and hence, the preserved record shows a great degree of variability. Fairweather trace fossil a blages range from equilibrium suites of the proximal Cruciana ichnofacies in the lower shoreface, to the ‘Skolithos ichnofacies in the middle shoreface (Fig. 34), enhancing the observed variability in this deposi- tional complex ‘The upper shoreface-foreshore complex is also quite variable, but this is mainly in regard to sediment texture and morphodynamic configuration. Preserved facies are remarkably similar over a wide range of shoreface states and accumulate in response to tairweather conditions. Storm events are non-depositional and strongly erosive. ‘The trace fossil assemblage is generally sparse and corresponds to the Skolithos ich- nofacies (Fig, 34), Many intervals contain zones of Macaronichnus segregatis near the upper shoreface- foreshore transition. 85‘The main variations in the preserved record of a shoreface succession are reflected in the lower-middle “« complex. This large degree of variability can be attributed to variations in overall storm inten sity, storm frequency and relative water depth. Three main “types” of shoreface successions can be con- sidered, although a complete continuum exists between them. Amalgamated tempestites with minimal preserved biogenic structures constitute the lower middle shoreface complex of strongly storm-dominated systems, Moderately storm-dominated systems correspond to intermediate energy conditions. The lower- middle shoreface complex consists of stacked, erasively-hased storm heds, bioturbated at the top by ‘opportunistic traces and cross-cut by a resident trace assemblage associated with fairweather conditions. ‘The weakly storm-affected shorefaces constitute the lowest energy and most dissipative conditions. The lower-middle shoreface complex is characterised by thoroughly bioturbated sandstones, recording nearly ‘continuous fairweather accumulation, with remnant, largely obliterated, thin intercalated tempestites. shoret ‘Stronger storm-domination may correspond, in part, to shorefaces which are subjected to severe winter storm seasons, where high frequency events permit thick amalgamated tempestites (cf. Cook and Gorsline, 1972; Owens, 197; Hunter ef al, 1979). Lower eneigy forms may reflect those subjected to the much lower frequency hurricane storms (Dott, 1988). Amalgamated, strongly storm-dominated shoreface suc cessions are relatively abundant in the northern portions of the Western Canada Sedimentary Basin (e.¢. the Bluesky Formation, the Fahler members of the Spirit River Formation and the Cadotte Member of the Peace River Formation. Higher latitudes appear to favour higher frequency and intense winter storms, and this warrants consideration asthe overriding factor in the paleogcographie distribution of these shorcface types. Discrimination between lower and middle shoreface deposits may be possible on an ichnological basis; where well-developed fairweather suites are present the boundary corresponds to a transition from a predominantly Cruziana ichnofacies to a Skolithos ichnofacies. Under progressively greater storm domi nation, fairweather suites are insufficiently preserved to resolve the two subenvironments ichnologically and physical characteristics such as coarsening of sediment texture, decrease in thickness of fairweather deposits, and replacement of lower energy HCS beds by SCS beds, may have to be employed. In the case of QPL intervals, itis unclear whether such a distinction is possible. In any case, vertical and lateral transi- tions between shoreface “types” may occur, depending upon relative water depth, storm frequency, storm intensity and shoreline orientation in relation to dominant wave attack. ‘These conditions serve to vary the character of contemporaneous lower and middle shoreface deposits along depositional strike, probably making differentiation of these two subenvironments impractical. 86THE ICHNOLOGICAL EXPRESSION OF STORM DEPOSITS Trace fossils are proving to be one of the most important groups of fossils in delineating strati- graphically important boundaries related to sequence stratigraphy (Pemberton and MacEachem, 1995) and allostratigraphy (Pemberton, et al., 1992a). Likewise, event stratigraphy is another area where trace Fossils lave proven (o be x powerful tool for the recognition and interpretation of event beds (Seilacher, 1981, 1982a, b; Wheatcroft, 1990; Pemberton et al., 1992b; Frey and Goldring, 1992). Event layers are generally regarded as strata that are attributed to rapid, episodic deposition and include such diverse entities as: volcanic ash beds, or tephra deposits (ie. Pedersen and Surlyk, 1983), beds resulting from seismic shocks, or scismites (Scilacher, 1969); and episodic sedimentation events such as turbidites (Seilacher, 1962); tempestites, or storm deposits (Aigner, 1985), phytodetritus pulses (Rice €¢ al, 1986); and inundites, or flood deposits (Leithold, 1989). Trace fossils are known to be significant features of most of these deposits. EPISODIC DEPOSITIONAL EVENTS During the last two decades, geologists have begun to recognize the extent and significance of relatively rapid inttuxes of sediment (episodic depositional events) in the rock record. In fact, Dott (1983) has successfully argued that most of the sedimentary record, rather than reflecting day-to-day, steady state conditions. typically represents episodic or discontinnans depositional events. Initial re confined almost exclusively to the deep-water, outer continental shelf and slope environments, where gravity-induced turbidity currents constitute one of the more important operating mechanisms of deposi- Gon, More revently, however, sedimentologists have focused their attention upon episodic depositional events in shallow, coastal areas. Such events, generally the result of major storms or hurricanes, are well documented in recent environments and are being recognized with more frequency in the rock record h was In a seminal paper, Seilacher (1982a) succinctly pointed out that episodic sedimentation events (turbidites, tempestites, inundites, and phytodetritus pulses) have a number of common characteristics: (1) that they reflect the onset, culmination, and waning of water turbulence during the event, by distinctive erosional and depositional structures; (2) that they redistribute the organic and inorganic sediment mate- rial along a vertical (bottom to top) and horizontal (shallow to deep) gradient; and (3) that they change the ecological situation for benthic organisms by altering the consistency andlor the food content of the hattom for a biologically relevant period of time after the event. The two most prolific types of event beds are turbidites and tempestites ‘Turbidites Perhaps the best known event bed is the turbidite that results from a single, short-lived event, a turbidity current. First recognized by Kuenen and Migliorini (1950) turbidity currents are a form of density current that flows downslope on the ocean flour driven by gravity acting on the density difference between the suspended sediment and the surrounding sea water. Walker (1984b) stressed that turbidity currents can be considered commonplace in modern oceans and their resultant deposits are extensive and volumetrically important. 87‘Walker (1984b) summarized a set of descriptors that characterize the features of classical turbidites: (1) sandstones and shales that are monotonously interbedded through many tens ot hundieds of meters, the beds tend to have flat tops and bottoms with no scouring or channeling greater than a few centimeters, (2) sandstone beds have sharp abrupt bases, and tend to grade upward into finer sand, silt, and clay; (3) sandstone soles have abundant markings that reflect either physical (tool marks, scour marks) or biogenic (trails or burrows) activity; and (4) within the sandstone beds combinations of parallel lamination, ripple cross lamination, climbing ripple cross lamination, convolute lamination and graded bedding result in a recurring sequence that reflects stages in waning current flow. This idealized sequence was first described ‘by Bouma in 1962 and is known as the Bouma sequence. The Bouma sequence represents one of the most elegant facies models as it acts as a basis for hydrodynamic interpretations. The sequence can be divisible into a number of discrete zones or divisions using textural and structural features: (1) division A: the graded division lacks internal lamination and is ither massive or normally graded; (2) division B: is typified by parallel laminations with parting linea- tion; (3) division C: is distinguished by current npple cross lamination; (4) division D: is distinguished by the interlamination of clean silt and mud; and (5) division E: consists only partly of mud introduced by the current (Bouma, 1962).The ichnology of turbidites is very well known and a number of excellent recent papers describe pre- and post-urbidite ichnofossil assembiages (Wetzel, 1991; Miller, 199a,b; and Leszezynski and Seilacher, 1991), ‘Tempestites Major problems exist in the recognition and interpretation of storm events because shallow coastal zones comprise a multitude of depositional envisonments eonumonly differing from one another in only subtle ways. Successions of rock types and sedimentary structures that are of value in delineating deep- water turbidity currents, at present, have somewhat limited significance in shallow-water deposits. One particular type of bedding (hummocky cross-statfication), however, seems to be formed exclusively by storm waves (Walker, 1984a), Storm units represent individual sandstone beds that were deposited rapidly (from a few hours to several days) from single waning flow events (Fig. 61), which generally possess a strong oscil- latory flow component (Dott, 1983). They can superficially resemble turbidites but are generally distin- guished by the presence of wave-generated and/or combined flow sedimentary structures and shallow ‘marine faunas in the intercalated shale deposits (Johnston and Baldwin, 1986). Conceptual models of storm-derived units have been constructed (e.g. Dott and Bourgeois, 1982) and interpreted. These ideal- ized successions generally show the following characteristics: (1) storm erosion: a basal erosion surface ‘which may be undulatory, with sole marks, and intraclasts of pebbles, shells or mudstone, locally sideri- tised; 2) main storm deposition: main hummocky or swaley cross-stratification interval, locally with asso- ciated horizontal or parallel lamination; (3) waning storm deposition: combined flow andor wave rippled sand layer, indicating a progressive return to lower flow regime oscillatory conditions; and (4) post-siorm /fair-weather mud deposition: reflecting either the final suspension fall-out of storm-derived sediment (Ce. poststorm mud) oF the return to normal background sedimentation (i. fait-weather mud) Johnston and Baldwin, 1986). 88HUMMocKY ‘cROSS ‘STRATIFICATION SECONDS Figure 61. Temporal significance of the idealized storm unit (modified after Dott, 1983) ICHNOLOGY OF STORM DEPOSITS In order to illustrate the characteristic trace fossil patterns discerned in storm-influenced settings, reference is made to the well-preserved suite of trace fossils from the Upper Cretaceous (Turonian) Car dium Formation of Alberta, Canada (Pemberton and Frey, 1984; Vossler and Pemberton, 1988). Trace fossil assemblages inthis unit are complex, because they Contain elements of both the Skolithos and Cru ‘without storm lag. 5 2 E. Colonization by opportunistic Prarmeaay shallow water organisms D. Common escape Ihureowed top ee 4s f G.. Rack to. diverse resident Fehnofossi assemblage Figure 63, Summary of the ichmological features associated with an idealized storm succession. (A) ‘The succession sturts with a diverse, fair-weather resident ichnocoenose comprising elements of the Cruciana ichnotacies, (B) This assemblage is interrupted by the storm event, (C) The main bouy ofthe storm deposit consists of parallel to subparalle! lamination, representing hummocky c1uss-stratification, (D) The storm-generated stratification may contain escape structures, recording the wtiempt of organisms to burrow through the event bed. (E) The top of the storm bed is commonly biogenically disturbed by dwelling burrows of opportunistic organisms, (F) Following storm abatement, fait-weather depositional patterns resume and the equilibrium resident assemblage is re-established, producing a gradational burrowed top to the event bed. (G) The farr-weather assemblage continues to flourish. (Modified from Pemberton ef al., 1992b) 93Such laminated to burrowed successions (Fig. 63) are typical of many interpreted shoreface environments in strata from the Cretaceous of the Western Interior. Pemberton et al. (1992c) deseribed storm-dominated deposits from the Spring Canyon Member ofthe Blackhawk Formation in Utah, Figure {64 depicts the development of laminated to burrowed (lam-scram) beds typical of the lower shoreface showing storm (lam) and fait-weather (scram) trace fossil assemblages and sedimentary structures. In core, iti difficult to recognize hnimmocky and swaley crass stratification because of scale prob- Jems. However, characteristic trace fossil patterns can be used to assist in the interpretation of storm «deposits. A core (Pan Am B-1 Ricans 10-01-35-8W5) from the Cardium Formation displays a diverse and well preserved trace fossil suite including representatives of at least 15 ichnogenera: Asterosoma, Chon- drites, Diplocraterion, Helminthopsis, Palacophycus, Planolites, Phycosiphon, Rhizocorallium, Rosselia, Schaubeylindrichnus, Skolithos. Subphyllochorda, Teichichnus. Thalassinoides, and Zoophycos « Near the top of the core are a number of thin, discrete sandstone beds that have been interpreted as storm deposits. ‘These beds are characterized by: (1) sharp bases, with or without basal lags; (2) low angle parallel to subparalfel laminations, (3) escape stuctures, (4) the dwelling buows of upportunistic organist; and (5) gradational burrowed tops (Fig. 65). ‘The trace fossils in the sandstone beds are interpreted to represent the activities of opportunistic ‘organisms that rapidly colonize the surface of the storm sand. Characteristic Forms include Skolithos, and Ophiomorpha. ‘These sandstones are interbedded with bioturbated fine-grained deposits dominated by Anconichnus, Asterosoma, Chondrites, Helminthopsis, Planolites, Rhizocorallium, Rosselia, Subphyl: lochorda, Teichichnus, Terebellina, Thalassinoides, and Zoophycos. The suite represents the resident suite and is indicative of stable, fair-weather deposition. Proximal/distal trends can be recognized by docu- menting changes in the resident trace fossil suite (Fig. 66). TUPEPETIE TT PALL TOT Figure 66. Proximal-distal trends in the 10-1-35-8W5 core are associated with changes in the resident (or fair-weather) assemblage; going up core this assemblage changes, froma distal Cruziana assemblage to a proximal Cruziana assemblage. (Modified from Pemberton and MacEachern, 1997.) onRecently, Bromley and Asguard (1991) imroduced a new ichnofacies, the Arenicolites ichnofu- cies, to account for short-term, opportunistic occurences of trace fossils in incongruous settings, such as storm deposits, Such a designation is confusing because it can not be separated from the present Skolithos ichnofacies concept. The present array of ichnofacies is adequate to explain such occurrences, Follmi and Grimm (1990) have postulated the "Doomed Pionee:* concept to explain anomalous trace fossils exclusively associated with gravity flow deposits within otherwise unbioturbated sediments. ‘Such a scenario would indicate that event deposition in oxygen-depleted sedimentary environments might bbe accompanied by the appearance of allochthonous infauna. The concept (Fig. 67) involves: (1) the entrainment of living decapod crustaceans (or presumably other hardy infauna) in some type of turbulent sedimentation event; (2) these imported burrowers reworked substantial quantities of laminated, com- ‘monly organie-rich sediment, in an environment from which they were previously excluded; and (4) the persistence of oxygen-depleted environmental conditions limited the survival time of these transported infaunal dwellers and rendered them ‘doomed pioneers" (Grimm and Follmi, 1990). From the standpoint of event deposition this concept may well be important in understanding the securrence of other nnamalons trace fossil associations. Likewise, as pointed out by Féllmi and Grimm (1990), the occurrence of Thalassinoides in anoxic hemipelagic host sediments has previously been incor- rectly interpreted to indicate bottom-water ventilation and re-oxygenation on a broad regional and/or tem- poral scale, The recognition of doomed pioneer trace fossil associations as ephemeral ecologic phenomena that are prominent in some laminated sequences should permit a more precise understanding of paleo- oxygen levels and hasin history (Grimm and Féllmi, 1990) Anomalous trare fossil assariatinns noted in the Sirasun/Terang Field, offshore Indonesia were interpreted as a doomed pioneer suite (Fig. 68) This suite had considerable impact on the breaching of possible permeability barriers and resulted in an incteave of vertical permeability and connectivity. PRESERVATIONAL POTENTIAL ‘The increasing awareness of event bed deposition in the ancient record, or "uniformitarianist catastro- phism" (Kumar and Sanders, 1976) has couespondingly led ty much consideration of the preservability of such beds (e,g, Dott, 1983; 1988; Wheateroft, 1990). Observations of tempesttes in various depositional settings demonstrate a wide variety of preservation styles. The fundamental parameters revolve around the net sedimentation rate, the biogenic mixing rate, and the magnitude of physical reworking, although most proposed quantitative models operate under the assumption thatthe latter factor is absent. Wheatcroft (1990) demonstrated that neither sedimentation rate nor biogenic mixing rate is easily defined both of them masking important, complex affiliated processes and interactions. Few of these have been addressed {in detail, despite the effect they have on event bed preservation. In contrast to previous models, which have concentrated mainly on layer thickness of event beds and mixing zones, Wheateroft (1990) focused cn time scales, ‘The result has been a model relating transit time (i.e. the time necessary to bury the event bed beyond the reach of the burrowing infauna) and the dissipation time (Zc. the time required to biogeni- cally destroy the event bed). Clearly, in circumstances where the event bed is thicker than the zone of burrowing, the fraction that exceeds the burrowing zone will be preserved regardless of the duration of biogenic reworking, Compelling though this model is, the inherent variability ofthe physical and biologi- ‘cal responses to the depositional event greatly limits the accuracy of this quantification and thus limits its Utility to the interpretation of the ancient recoed 95The Concept of Doomed Pioneers ‘Shot Interval of Importation of by burial in aminateg burrowing organisms Laminates ost vent depostion Thalassinottes secmonts trom gated Inailentonous event ediren how nda uo}acent ta A B c ‘The Doomed Pioneers Concept: Critical Observations Observations Interpretations 1. Bioturbation only within event strata and in directly euhjacent sediments, 2, Thalassinoides and Ophiomorpha 43. Selective bioturbation of fine- grained intervals; avoidance of tuffaceous deposits. 4. Bioturbated intervals are sharply overlain by unbioturbated hemipelagic sediment. 5. Shelly benthos and tiered infauna are absent. 1, Conclude a genetic relationship between ‘event deposition and bioturbation; infor ‘causal relationship between inputs of allochthonous sediment and importation of burrowing fauna. 2. Crustacean dwelling burrows; foreign to anoxic basinal environments; derived from neritic setting. 3. Probably indicates successful colonization and exploitation of available nutrients in basinal environment. 4. Abrupt return to or persistence of ano» conditions sharply limits temporal duration of burrowing episode. 5. Abrupt burrowing episode was not the ‘onset of substantial ecological ion on the basin floor, absence survivability of infaunal immigrants and/or insufficient opportunity for subsequent larval recruitment. Figure 67. hematie diagram illustrating the doomed pioncers model to explain the ‘exclusive association of allochthonous event deposits with bioturbation; this example relates to the introduction of burrowers into oxygen-depleted environments. Critical obser- vations and interpretations of the doomed pioneers concept are noted, (Modified from Grimm and Fotmi, 1990.) 96The Ichnology of Graded Storm| Beds: Sirasun/Terang Field, Offshore Indonesia Ophiomorpha Terbelina Planolits Rhtzocorallia Helminthopste ‘Anconichees 1, Possible permeability aR barriers are breached Sra Fe to by deep penetrating Tontnttin Ophiomorpha that were Scolicia/Subphyllocorda emplaced following episodic deposition. 2. Example of doomed pioneer concept. 3. This greatly enhances the vertical permeability and connectir Graded Storm Bed Sirasun/Terang Field, offshore Indonesia Figure 68, Large Ophiomorpha in graded stormbed in the Sirasun/Terang Field, offshore Indonesia are interpreted as doomed pioneers that breach possible permeability barriers. ‘Transit Time Previous workers have calculated the transit time by dividing the thickriess of the zone of biogenic reworking by the sedimentation rate. Wheatcroft (1990) pointed out that the thickness of the event bed itself was important in the calculation of transit time, and proposed that the combined thickness of the biogenic reworking zone and event layer, less one half of the thickness of the event layer, all divided by the sedimentation rate was a more accurate estimation of the transit time In general, the zone of biogenic reworking is thinnest in the lower offshore to abyssal depths, where diminutive grazers and foragers dominate the benthic community. The zone thickens through the upper offshore, where more robust and mobile deposit feeders dominate. ‘The fair-weather Skolithos ichnofacies 7‘may extend the zone of biogenic reworking much deeper in lower shoreface and shallower settings, but is also characterized by a decrease in effective homogenization of the hed at those depths Sedimentation rate is highly problematic to resolve quantitatively, even using modem settings as analogs. ‘The most obvious problem revolves around the use of the average or fair-weather (ambient) sedimentation rates. ‘The mere presence of an event bed in the interval in question demonstrates that sedi- ‘mentation rates are inherently unsteady and unpredictable inthe setting. An offshore zone may experience ‘months of fair-weather deposition at a rate that is totally overshadowed by the accumulation of a single tempestite deposited over the course of days. Since these events are unpredictable, both with respect to their frequency and the magnitude of deposition, dey currently defy reliable quantification and inclusion into calculated sedimentation rates of most settings. Dissipation Time Dissipation time corresponds to the time required to biogenically destroy un event bed. Wheateroft (1990) demonstrated that there isa fundamental difference between destruction of an event bed character ized by a discrete mineralogy and chemistry (e.g. ash beds or iridium anomalies) and destruction of an event bed distinguishable only by its lithology and fabric (e.g. HCS beds and turbidites). ‘The former ‘anomalies may survive considerable biogenic reworking, becoming mixed over a considerable thickness of interval and yet still remaining detectable. In contrast, the latter is recognized on the basis of far more casily obliterated criteria (sedimentary structures, etc.) and is therefore more sensitive to the duration of biogenic activity. Dissipation time is comparatively more difficult to determine than is transit time, Previous discussion regarding the characteristics of the benthic community touched on two param- eters strongly affecting dissipation time. ‘The most obvious parameter is the size of the animals constitut- ing the infauna, Robust organisms can displace or mix sediment a greater distance per unit time than can diminutive ones. Ina very general sense, organism sizes tend to be larger in shallower water settings than in deep water, although each benthic community typically contains a wide range of infauma sizes, atribut- able to a large number of factors, ranging from differences in genera or species, to relative proportions of Jjuveniles and adults (Dérjes and Hertweck, 1975) More important with regard to dissipation time is the behavior engaged in by the organism. Mobile ‘deposit feeders and camivores tend to produce much greater disruption of the bed, in both a vertical and lateral sense, than are generated by grazers and foragers. The geazing/foraging behaviors may result in rather intense reworking of the sediment, but this is generally restricted close to the sediment-water inter- face (Ekdale, 1980). Continuous, slow accumulation of fair-weather deposition may promote thorough homogenization of the sediment laterally, but the tracemakers tend to show little propensity to penetrate deeply into the substrate (Howard, 1975; Dott, 1983, 1988). An exception is the Zoophycos tracemaker, ‘which may display substantial vertical penetration into the substrate (Kotake, 1989). However, the bio- genic structure rarely shows intense reworking of the event bed, which typically has litte in the way of deposited food (trey and Goldring, 1992). Lypically, the Zoophycos trace crosses through the event bed with minimal disturbance, until a resource-rich interval, generally constituting a buried fair-weather or ambient deposit. is encountered. Thorough reworking of the buried fair-weather sediment may then occur (Vossler and Pemberton, 1988). 98Dwelling structures and suspension feeding structures may show appreciable vertical penetration into an event bed, but typically displays minimal volumetric modification af the hed!s fabric. ‘The structures may deepen or extend areally over the lifetime of the tracemaker, but such structures clearly do not result in the degree of homogenization imparted by mobile infauna (Pemberton and Frey, 1984; Frey, 1990). As such, the actual thickness of the biogenic mixing zone is less important than the nature of the behavior employed by organisms within the mixing zone. Pechaps one of the most under-appreciated factors regarding the dissipation time is associated with the dynamic interplay between the benthic community and the depositional event itself. In many cases, the event bed follows « disturbance that largely displaces or destroys the resident benthic community Biogenic modification may not simply begin immediately after the event. Instead, the site may remain tunpopulated for a considerable period of time. Once recolonisation is initiated, it may take even more time for the community to achieve the density it possessed prior to the disturbance (Sousa, 1984). Shal- lower water settings, characterized by persistent and generally higher degrees of stress may show repopu- lation within 11 months of a disturbance, whereas more highly sensitive deep sea settings may take more than double that time to recover (Dauer and Simon, 1976; Grassle, 1977) Complete destruction of the benthic community may occur in two main ways. ‘The most obvious is related to the severity of the disturbance producing the event bed. This is particularly evident in shal- lower water settings, where erosional amalgamation of beds is more pronounced and locally, cannibalis- tic (MacEachern and Pemberton, 1992). Under these conditions, the entire benthic community is typi- cally washed out of the substrate and transported elsewhere. The severity of such disturbances decreases basinward, such that distal tempestites, and in particular turbidites may show well preserved fair-weather/ resident trace fossil suites on their basal surfaces (e.g. Seilacher, 1962; Crimes, 1977; Crimes et al., 1981; Pemberton and Frey, 1984: Frey and Goldring, 1992). The second major factor resulting in destruction of the benthic community is associated with the thick- ness ofthe event bed laid down. Once the organisms become buried, they attempt to burrow up through the bed to reach the new sediment water interface. If the event bed is too thick, none of the original resident community will survive to repopulate the post-storm substrate, minimizing the time of “effective” bbioturbation (Kranz, 1972). On the other hand, thinner event beds may be repopulated more quickly, since most of the benthic community may escape burial. Some biogenic modification of the event bed will ‘occur, purely as a function of the escupe structures. ‘Tempestites show a general basinward thinning and initially it may seem, therefore, that the thinner beds of the shelf and lower shoreface may be rapidly colonized. In actuality, because organisms of these basinal benthic communities tend to be diminutive in size, few animals may be able to reach the top of even a relatively thin event bed, AAs well, few tracemakers of the shelf and more basinal settings possess carapaces, and therefore are unlikely to survive entrainment in a tempestite or turbidite flow, unlike some organisms of shallower water settings. Another important factor surrounding the dissipation time is associated with the desirability of the event bed as a site for colonization and as a repository of food resources. The event bed generally varies ‘markedly from the normal ambient (fair-weather) sediment, to which the resident community is suited. Even those organisms that burrow out of the event bed may, upon reaching the sediment-water interface, find a substrate devoid of suitable food resources, and subsequently die. ‘Those which do survive, as well as opportunistic organisms whose larvae managed to find their way to the unexploited substrate are trophic 99generalists. ‘They typically do not burrow through the bed, but rather, establish domiciles and engage in suspension feeding, or carry out surface deposit feeding and scavenging strategies (Jumars, 1993). Penetrating domiciles show minimal lateral disturbance of the bed, and exceedingly dense populations are needed in order to obliterate a bed. Once suitable amounts of ambient sediment accumulate and the resident faunal community re-establishes itself, the infauna generally show litte inclination to penetrate deeply into the event bed, but rather, engage in behaviors suited to the ambient conditions (Rees e? al., 1977). This avoidance pattern appears most pronounced in basinal settings, where the contrast hetween behaviors suited to the event bed and those suited to the ambient substrate are greatest. This is most common where turbidites are deposited in abyssal settings. In upper offshore conditions, where robust and diverse tracemakers of more proxital Cruziana ichnofacies predominate, the differences between event bed and ambient deposit behaviors are less pronounced. In lower shoreface and middle shoreface settings, no preference for fair-weather burrowing, over event bed burrowing may be noted and for all intents and purposes, both suites are identical. ‘The final factor regarding dissipation time is the absolute time available for burrowing. As in the discussion of sedimentation rates, there is obviously considerable variability in burrowing time in any set- ting characterized by episodic deposition. In the case of tempestites, one may suspect minimal burrow. ing times to occur in shallower water settings, with progressively more time available basinward. ‘This primarily reflects the greater probability of storm interaction with the sea floor in shallower water, regard- less of storm magnitude. In the case of turbidites, however, variability in burrowing time is more closely related to basin paleogeography and, if present, the configurations of any submarine fan systems. In general, the above factors impose fundamental controls on the effectiveness of biogenic modifi tion of event beds, and consequently, on their preservation potential, Unfortunately, these factors appear to resist quantification, and therefore severely limit the effectiveness of existing mathematical models to explain event bed preservation in the rock record. Physical ceworking of the substrate is typically avoided in most of the models proposed, but clearly has a profound effect on the preservation of both the event bed and the fair-weather deposits. In the case of tempesttes, enhanced physical reworking is associated with shallower water settings. with higher ‘magnitude storms, and with higher frequency storms. The latter factor principally reflects minimal ‘weather accumulation on the tempestite and minimal colonization ofthe substrate, both of which serves to enhance the ability of successive storm events to rework earlier tempestites. Under proxinal condliions, successive tempestites may become highly cannibalistic (Aigner and Reineck, 1982), resulting in only ‘minor preservation of the beds. Despite the difficulties in mathematically modeling preservation potential of tempestites, there are ‘ number of proximal-distal trends evident on the basis of empirical observations. In general, shallow water settings favor the preservation of event beds, since higher numbers of storms enhance erosional ‘amalgamation and minimize the time available for burrowing, factors that are also responsible for the ‘minimal preservation of fair-weather deposits. These appear (o overcome the factors favoring biogenic ‘modification of event beds, such as larger animal sizes, greater bed penetration, suitability of the event bbed to colonization by the resident henthic community, and higher rates of recolonisation following a disturbance. Shallow water deposits are dominated by vertical to subvertical domiciles, minimizing event bbed modification, These settings typically correspond to proximal lower shoreface and middle shoreface environments, 100Erosional Amalgamation Burrow Penetration of Substrate Average Animal Size Rate of Recolonisation Rate of Amblent Sediment Deposition Frequency of Events Duration of Fairweather Conditions Event of Bed Avoidance by Infauna Degree of Biogenic Homogenisation Sensitivity of Benthic Community to Disturbance Rate of Sediment Displacement by Infauna ‘Tempesite Preservation Potential F tempestite preservation potential appe: Tempesite Preservation 11 NLA gure 69, Schematic representation of the main factors affecting tempestite preservation and their ive importance with respect to proximal and distal depositional positions. Note that in general, s lower in intermediate depositional sett servability occurs in proximal and distal settings, although for markedly different Higher pre- -asons,Distal settings also favor preservation of event beds, but for markedly different reasons. Fair-weather ‘deposits also have a high preservation potential, unlike shallow water settings. Factors favoring presei vation of tempestites are minimal erosional amalgamation, minimal bed penetration by infauna, diminu- tive size of infaunal organisms, general avoidance of event beds as a viable substrate, sensitivity of the benthic community to environmental disturbances, and slow rates of recolonisation following a distur- bance. These serve to overshadow conditions favoring event bed modification, such as long duration Of fuir-weuther conditions, slow rates of ambient sediment accumulation, and reduced potential of event bed accumulation. Such settings are typical of shelf and lower offshore settings in the Cretaceous of the Wester Interior Seaway. ‘The lowest preservation potential for tempestites appears to occur somewhere between the proximal and distal extremes, Under such conditions, erosional amalgamation is not as effective and displacement of the resident suite is not as common. Further, the reduced sensitivity of the benthic community to dis- turhances and generally higher rates of recolonisation, all favor more rapid biogenic modification of the event bed. ‘The variability in organism type and behaviors employed insures that the tempestite will be suitable to some of the benthic community as a viable medium for burrowing. The higher numbers of mobile deposit feeders and carnivores favors rapid and thorough modification of bed fabrics within the biogenic mixing zone, while the introduction of suspension feeders also promotes the development of deeply penetrating vertical structures. Such zones appear to correspond to proximal Cruciana ichnofacies suites ofthe upper offshore and distal portions of the lower shoreface. ‘The tempestite preservation potential (Fig. 69) seems to be consistent with observations of many ‘modern and ancient examples (Kumar and Sanders, 1976; Pemberton and Frey, 1984; MacEachern and Pemberton, 1992), Dott (1988) used the deposit of Hurricane Carla to outline this variability of preserv. tion potential. McGowan (reported in Dott, 1988) was unable to identify the prominent tempestite result- ing from Hurricane Carla, reported on by Hayes (1967), a mere 15 years after the event. Bioturbation had apparently obliterated much of the record. However, Nummedal (reported in Dott, 1988) managed to recognize the event bed further offshore (beyond 18-20 m of water depth), where biogenic modification of the tempestite had been less intense. Dott (1983) indicated that this alternating interplay of deposition, burrowing and scouring could produce a very subtle record of event deposition (Fig. 70). Concealed bed- junction preservation of the trace fossils, represented by truncated burrow tops and exotic burrow fills may represent the only clue to the episodic history of such amalgamated sequences. Figure 71 schematically shows the progressive colonization of a tempestite and the variable effects of latter erosional amalgama tion on the preserved record. Amalgamation of Bedding by Burrowing Organisms Erosion Erosion Deposition Burrowing Burrowing Rurrowing Deposition Rurrnwing Deposition a 1 2 3 4 3 6 Figure 70. Amalgamation of strata by organism burrowing; the record of deposition punctuated by two events is obscured by the overprint of bioturbation, (Modified from Dott, 1983.) 102qWOAg WHOIS UOEZIUOIOD AOyBOMIeI —_UOHEZ|UO|OD WHOIS 3S0d sajjsedwiay jo uoWez]UOIO9 an|ssaiBo1d‘The low preservation potential of hurricane.induced tempestites may also reflect the infrequent nature of such disturbances. In these settings, the tempestite is exposed to long periods of biogenic colonization and modification before it is buried below the reach of infauna. In contrast, many high latitude settings are characterized by cyclic variations in storm activity. Such settings are characterized by a winter storm, season, where disturbances are both frequent and of high magnitude, and a summer fair-weather season, where storms are less frequent and of lower intensity . Under such conditions, tempestites accumulate rapidly during winter seasons with litle or no time for biogenic colonization, Summer fair-weather sea- sons favor biogenic colonization and modification of the tempestites; however, this is largely restricted to Ue (op of the uppermost event bed. Any tempestites that may accumulate in response to the infrequent ‘and lower intensity storms typical of the fair-weather season may show significantly greater degrees of biogenic mottling. SUMMARY ‘Storms represent an important mode of deposition in most basins and are being recognized with great frequency in the rock record. In one of the best books written on stratigraphy Ager (1981) con- cluded " Nothing is world wide, but everything is episodic. In other words, the his- tory of any one part ofthe earth, like the life ofa soldier, consists of long periods of boredom and short periods of terror” (Ager, 1981, p. 106-107) ‘Tempestites contain a characteristic trace fossil suite that consists of a stable fair-weather assem- blage and an unstable starm assemblage. The fair-weather (or resident) assemblage is dominated by traces, of equilibrium (or K-selected) species while the storm (or pioneer) assemblage is dominated by traces of ‘opportunistic (or r-selected) species. ‘Tempestites show the following physical and ichnological characteristics: (a) a sharp base, with or without a basal lag: (b) parallel to subparallel laminations (hummocky or swaley cross-stratification):, (©) common escape structures; (4) dwelling burrows of opportunistic organisms; and (e) gradational bur- rowed tops. Proximal / distal trends are generally discerned by changes in the character of the fait-weather suite. The integration of ichnological characteristics with the physical sedimentary features is particularly important in recognizing tempestites. 104THE APPLICATION OF ICHNOFACIES ANALYSIS TO THE EVALUATION OF MARGINAL MARINE RESERVOIRS INTRODUCTION Marginal marine environments, including the intertidal zone, shallow lagoons, sounds, incised val- leys, estuaries, bays and delta platforms. characteristically display steep salinity gradients (Fig. 72) result- ing from variations in (a) amounts of freshwater input from rivers and runoff from land, (b) rainfall, (c) ‘evaporation, (d) tidal range and salinity content in adjacent open-ocean coastal waters, (€) morphology Of the coastal area, and (f) differences in wind divection and velocity (Dixjes and Howatd 1975). Such salinity fluctuations, combined with corresponding changes in such parameters as temperature, exposure, sedimentation rates, substrate consistency, turbulence, oxygen content, turbidity, efc. result in a physi- logically stressful environment for numerous organism groups. Wightman cr al. (1987) interpreted several coarsening upward sequences in the upper Mannville of «east central Alberta to represent stacked crevasse splays deposited in brackish water bays. They presented an integrated model, based on sedimentological, ichnological, geochemical, and palynological data to sup- port their interpretation. The ichnological component of this model was based on earlier work done by Pemberton in Ekdale et al. (1984) and has been further documented by Pemberton and Wightman (1992). Similar brackish water deposits are now being recognized with greater frequency in the rock record, and their significance in the interpretation and delineation of the marginal marine zone is becom ing mote apparent. The concept of incised valley complexes is one of the more important aspects to ‘emerge from sequence stratigraphy (Reinson et al. 1988; Van Wagoner ef al. 1990; Zaithin et al. 1994) land the recognition of these features is partly dependent upon distinguishing between fully marine and brackish water environments (Pemberton etal. 19923; MacEachern and Pemberton, 1994). Although this distinction is best made using an integrated approach, trace fossils can often provide the least equivocal, and most readily accessible data for the recognition of brackish water deposits (Wightman ef al. 1987, Pemberton and Wightman 1992; MacEachern and Pemberton 1994; and Pemberton and MacEachern 1995). BIOLOGICAL AND ICHNOLOGICAL IMPLICATIONS, OF BRACKISH WATER Biological Response Itis a commonly held belief that most invertebrate groups originated in marine environments, and that representatives of some groups subsequently adapted to brackish and freshwater conditions. The physiological mechanisms involved in the adaptive process center on the organisms capabilities to regulate ‘osmotic flooding (osmo-regulation) and the ionic concentration of body fluids (ionic regulation), due to the lower and fluctuating salt concentrations imposed upon them (Croghan 1983). ‘The extent to which these adaptations occur in different organism groups is quite variable and thus, their distributions in rela- tion to salinity gradients show marked differences (Fig. 72). The freshwater fauna contrasts sharply with the marine fauna; the former is impoverished and is dominated by relatively few groups, most notably branchiopods, other freshwater crustaceans, and insects (Croghan 1983). Marine faunas are composed of os‘STENOHALINE MARINE SPECIES: EURYHALINE MARINE SPECIES EURYHALINE OPPORTUNISTS ESTUARINE ENDEMICS POIKILOHALINITY FRESH. waTeR POLYHALINE MeSOMALINE JOLIGOHALINE| 30 % Ea % 1 3 ° ‘SALINITY () Figure 72, Classification of estuarine zones relating the Venice System classification to distribution classes of organisms. (Modified after Knox, 1986,) diverse groups of organisms that are generally considered to follow equilibrium population strategies. The brackish water fauna, on the other hand, is characterized by opportunistic euryhaline species and consists of numerous elements, including (a) an exceedingly minor freshwater component consisting of euryhaline species restricted to low salinity, (b) a marine component comprising curyhaline species, (c) a brackish component consisting of species that penetrate neither fresh nor fully sine waters, and (d) a migratory ‘component comprising species that spend only a portion of their life cycles in brackish water (Perkins 1974), Investigations on modem brackish water environments indicate that although their biotic compo- nent is highly variable, a number of generalizations may have significant ecological and paleoecological ramifications (see Pemberton and Wightman 1992 for details). In summary these are: A.) Brackish waters are generally reduced in species numbers (diversity) with respect to both freshwater and fully marine water (Fig. 73). Sanders (1968) developed a theory based on the stability or predictability of the environment and on its geological history to explain differences in species diversity of benthic organisms, Wolff (1983) indicated that the main elements of this, theory as applied to brackish waters are: (1) in brackish water environmental factors are usually unstable and unpredictable; (2) speciation in such an environment is less probable than in more stable environments (Stobodkin and Sanders, 1969); (3) extinction in such an environment is more probable than in more stable environments (Slobodkin and Sanders, 1969); (4) brackish water is ‘a geological ephemeral phenomena, thus increasing the likelihood of extinction of brackish-water Populations; (5) for these reasons in brackish water the number of species adapted to the environ- ‘ment is lower than in either the marine or the freshwater environment; and (6) the unstable and unpredictable nature of brackish Water environments also excludes most marine and freshwater speci 106161 fold ¥ purMOH WOH =] 1941 29492989 an BUC|e SONS poroyes 1 Sus N S N iraNumber of Species 0 Oligo- Polyhaline Euryhalir paling Metehaine —grachyatine) —(Eunalng os 5 18 30 40 ‘ Limnetic Figure 74, Classification of salinity levels and generalized relationship of species diver- sity with respect to salinity. Note that brackish faunas are of low taxonomic diversity (Modified from Pickerill and Brenchley, 1991.) B.) Whereas the number of freshwater species decrenses rapidly even with slight increases in salinity the reduction of marine species is more gradual (Fig. 74). Therefore, the brackish water fauna can be considered more an impoverished marine assemblage than a true mixture of freshwa- ter and marine elements. C.) Sediments of estuaries and other brackish water environments are effective at dampen- ing the influence of salinity fluctuations (Sanders et al., 1965; Knox, 1986). Therefore, the deep infaunal habitat serves as a refugium buffering the organism against rapid and extreme salinity variations (Fig. 75). As a result, Alexander e/ al., (1933) showed dhat burrowing organisms are able to penetrate a greater distance up an estuary than epifaunal organisms which are subject to the fluctuating salinity of the overlying water column D.) With decreasing salinity, the teduction of species in-groups forming a calcareous skel- ton is greater than in their counterparts lacking such a skeleton (Remanc and Schlicper, 1971), ‘This suggests that the biota would be dominated by soft-bodied, trace making organisms. Many losOf these forms (Table 2) are considered minor phyla by some paleontologists but they constitute a large percentage of the biomass in most environments. These groups, for the most part, are also associated more with saline environments than they are with freshwater environments (Table 2) and as such representatives can be found in brackish waters. E) Many — marine organ- isms undergo a size reduction (Fig 76) when subjected to less saline water (Milne, 1940). Salinity) —> ° 40 20 F) ‘There is a distinct shift in the bathymetric range of marine organisms when subjected to salinity reductions (Remane and Schlicper, 1971). For instance, the polychaete Nereis diversicolor @ common intertidal organism in fully marine environments is a dominant element of the subtidal biota in many Dutch estuaries (Wolff, 1983). Depth (em) G.) Many — organisms that inhabit brackish water environ- ments can he considered to. be ‘omnivores or trophic generalists. Wolff (1973) found in the Dutch estuaries that 35% of the species ‘were omnivores, whereas this per- centage was only 6 to 16% in adja- cent freshwater or fully marine areas. This occurrence of wi niches iy related w an opportunistic life history (Fig. 77). Goerke (1971 found that in estuaries the poly- chaete Nereis diversicolor can employ several feeding techniques and may act as a deposit feeder, a suspension feeder, a scavenger and as a predator depending on prevailing conditions. Ekdale (1983) suggested that the trace fossils gencrated by opportunistic species (Fig. 77) would clarac~ terize such unstable and unpredictable environments. © High-Tide Satinity Figure 75, Salinity of bottom water and imerstitial water ata low tide station in the upper part of the Avon- Heathcote Estuary, New Zealand. (Modified after Knox, 1986.) H,) Although brackish water environments are characterized by low species diversity, Rosenberg et al. (1977) found that estuaries have very high values of benthic biomass when com- pared to other aquatic habitats. Such environments do possess some characteristics that favor benthic organisms for example: (1) they tend to be sheltered from wind waves and open swells; (2) they are rich in food provided by river input (ie. Schelske and Odum, 1961), input from salt marshes (Ke. Teal, 1962) or mangroves (Odum and Heald, 1975), input from the coastal sea (i.e. De 109Jonge and Postma, 1974), high in situ primary production (Wolff, 1977) of, most often a combina- tion of these factors: and (3) because of the shallow depths involved, suspended food particles are readily available for benthic organisms through sinking as well as through downward transport by turbulent water movements (Postma, 1961; Wolff etal, 1976). 1) These factors suggest that the benthic biota of brackish water environ- ‘ments is indeed unique and in contrast to Bromley and Asgaard (1991) we feel that the influence of salinity gradients can hardly be divorced from the ichnocoenose concept. For the same reasons that distinct nnon-marine associations are now being recognized i.e. the Mermia association. Hudson et al (1995) recently stressed that Jurassic benthic molluscan assemblages in the Great Estuarine Group of Great Britain are controlled largely by salinity with different genera or groups of organ- isms characterizing a spectrum of mainly brackish water environments. As pointed ‘out by Pemberton and Wightman (1992) salinity gradients also influence the nature Of the ichnofauna. Although some behav- ioral pattems may occur on each side of, the salinity transition, distinct ichnocoe- noses are present in freshwater, brackish water and fully marine environments s & : i 95 30 25 20 15 10 5 Salinity () ‘These factors suggest thatthe benthic biota of brackish water environment Figure 76. Reduction of maximal length of some indeed unique and in contrast to Bromley ‘mollusks with diminishing salinity in a transect and Asgaard (1991) we feel that the influ- from North Sea to Baltic. (Modified fromRemane ence of salinity gradients can hardly be and Schlieper, 1971.) divorced from the ichnocoenose concept For the same reasons that distinct non- ‘marine associations are now being recognized e.g. the Mermia association, the Termitichnus ichnofacies, and the Scoyenia ichnofacies (Frey et al. 1984; Smith er al. 1993; Buatois and Mingano 1995) itis also Possible to recognize distinct, recurring brackish water trace fossil associations.. Hudson er al. (1995) recently stressed that Jurassic benthic molluscan assemblages in the Great Estuarine Group of Great Brit- ain are controlled largely by salinity variations with different genera or groups of organisms characterizing {spectrum of mainly brackish water environments. As pointed out by Pemberton and Wightman (1992), salinity gradients likewise influence the nature of the ichnofauna. Although some behavioral patterns may ‘occur on each side of the salinity transition, distinct ichnocoenoses are present in freshwater, brackish water and fully marine environments. 110arginal Marine Trace Fossil Assemblage. dominated by opportunistic species eemeaeend eens Eee K-solected Ichnotaxa = = — - ——— selected iehnotaxa K-SELECTED r-SELECTED (EQUILIBRIUM) (OPPORTUNISTIC) 1, Respond rapidly 1, Slow to colonize. to open niche. 2, Reach size 2. Lack equlibrium equilibrium, population size. 3. Density dependant 3. Density independant mortality. mortality. 4. Good competitors. 4, Relatively poor competitive ability. 5. High dispersal ability. 6. Efficient reproductive system Figure 77. Gradients in the stability and predictability of physical environmental conditions will eon- {rol population strategies among burrowing organisms. quilibrium (K-selected) trace fossils lourish in high-diversity assemblages under very stable and predictable conditions. Opportunistic (r-selected) trace fossils rise to prominence in low-diversity assemblages under extremely variable snd unpredict- able conditions. (Modified from Bkdale, 1983). mLIn summary, brackish water environments tend to be characterized by: (1) an impoverished marine Suite of benthic organisms (Fig. 78); (2) an abundance of trace-making infaunal animals rather than epi- faunal animals; (3) a prevalence of soft bodied organisms over shelled organisms; (4) animals exhibiting a distinct size reduction; (5) bathymetric displacement of some animals; (6) a higher percentage of benthic organisms which are trophic generalists and whose activities result in morphologically simple burrows within the assemblage (Fig. 79); and (7) a reduction in diversity is commonly coupled with higher indi- vidual densities. Figure 78. Marginal marine environments are characterized by forms typeally found in marine settings and can be considered an impoverished marine assemblage. Ichnotogical Implications Although an unfortunate paucity of studies deal specifically with biogenic structures in modem brackish water environments, work by Howard and Frey (1973) on the estuaries of the Georgia coast indicate that: (1) the diversity and abundance of biogenic structures increases seaward; (2) distinct bio- genic structures and bioturbate textures are also more diverse and abundant along the margins of the estu- aries, rather than in the deeper channels; (3) distinet burrows and dwelling tubes characterize most muds, whereas sandy muds or muddy sands may exhibit both distinct burrows and various types of bioturbateRhizoliths Ophiomorpha Escape structures Figure 79, Marginal marine environments are dominated by morphologically simple structures including Teichichnus, Planolites, Skolithos, Arenicolites, Ophiomorpha, Gyrolithes, rhizoliths ‘and common escape structures. 3General Traits Common to Brackish Water Ichnofacies ds jioturbuted Sar rn cfu in distinguish aorine from fluvial sand. Skolithos-Cylindrichnus Assoc. * such simple, vertical burrows are Zommon'to estuarine sands and sand-dominated heterolithic beds. * ypical forms: Skolithos, Cylindrichnus & Arenicolites General Brackish Suite * sical an impoverished marine nivel einpaina dori feeding generis +B dominan rophie macs gal iversity low, indivi tice wettest nay assem Figure 80. General traits and tendencies common to brackish water trace fossil associations la} - = Gyrolithes dominated Teichichnus dominated assemblage assemblage Figure 81, Marginal marine ichnofossil assemblages are commonly dominated hy one or two ichnogenera. Recurring assemblages include the Cylindrichnus-dominated suite, the Gyrolithes- dominated suite, and the Teichichnus-dominated suite. ustextures; and (4) coarser sediments tend to lack biogenic sedimentary structures. In addition, they also recognized that the entite suite of estuarine bivgenic structures generally consist of both vertical and hori zontal burrows or burrow systems and thus do not fall conveniently into any of Seilacher’s (1967) univer- sal ichnofacies. Instead, the assemblage tends to be composed of @ mixture of structures typical af both the Skolithos and the Cruziana ichnofacies. Burrows tha if preserved would consist of Skolitios, Mono- ‘raterion, Thalassinoides, Cylindrichnus, Gyrolithes, Teichichnus, Ophiomorpha, Planolites and Palaeo- hycus (Howard and Frey 1973) characterize this assemblage Rrackish marginal marine environments are widespread in the modern, and are being documented with greater frequency from the rock record. Examples of studies in which the ichnofossils have been used as evidence of the presence of at least some saline influence can be found in Beynon eal. (1988), Pemberton eral, (1992b), Ranger and Pemberton (1992), Pemberton and Wightman (1992) and MacEach- ‘em and Pemberton (1994), The general brackish water assemblage (Fig. 80) reflects inherently fluctu- ‘ating environmental parameters and is characterized by: (1) low diversity; (2) forms typically found in ‘marine environments; (3) simple structures constructed by tropic generalists; (4) suites that are commonly ‘dominated by a single ichnogenus; (5) vertical and horizontal ichnofossils that are common to both the ‘Skolithos and Cruziana ichnofacies; and (6) some forms that may be found in prolific numbers (Fig. 81), It should be stressed that such an assemblage is distnet from the fully marine Cruztana tehnofacies, that is characterized by: (1) high diversity; (2) low individual densities; (3) a mixed association of vertical, inclined, and horizontal structures; (4) presence of structures produced by mobile organisms; (5) domina- tion by feeding and grazing structures constructed by deposit feeders, some of which show remarkably complex behavior pattems resulting in complex trace fossils (Pemberton et al. 1992), ‘TYPICAL BRACKISH WATER ICHNOFOSSIL ASSEMBLAGES Unlike fully marine environments that are subject to stable and predictable ecological parameters, ‘marginal-marine, brackish water zones are characterized by a plethora of sedimentary environments that ae charucterized by unstable and unpredictable ecological parameters. As such, complex arrays of ichno- fossil assemblages are possible. It is beyond the scope of this paper to provide details on all of the possible variations. Here. we will deal with assemblages that are typical of brackish water channel systems and brackish water bay-like environments. Brack it Water Channel Assemblages ‘Channels that are characterized by brackish water include estuaries, tidal creeks, tidal inlets, and distributary channels, among others. They typically contain sedimentary structures, bedding styles and sedimentary sequences that can be interpreted to result from tidal processes (see Dalrymple, 1992, for details). They also contain a distinctive trace fossil suite that is commonly dominated by simple vertical burrows such as Skolithos and Cylindrichnus (Fig. 82). Such trace fossils are commonly associated with facies characterized hy inclined herterolithic stratification that have been interpreted to represent lateral accretion point bar deposits. In the Lower Cretaceous MeMurray Formation of northeastern Alberta, a ‘Skolithos-Cylindrichnus assemblage characterizes the inclined heterolithic stratification deposits that have been interpreted as estuarine point bars (Pemberton et al. 1982; Ranger and Pemberton 1992). These assemblages contrast with ones associated with fluvial channels which are for the most part either unbur- towed or contain horizontal insect perturbations that are typically ephemeral and have a very low preserva- tion potential (Frey er al, 1984), 16[Estuarine Point Bar Model icMurray Formation, Lower Cretaceous, Alberta 1a Skolithos linearis 1b, Cylindrichnus associations Vertical burrows in inclined beds Vertical burrows » inflating beds Down-dip reclining lurows in inclined beds.» 2a, Gprolithos yp. Veil barons > inf ying ede Dowd reclang ‘uerows in inclined Beds.» » Figure 82. Schematic model of the Skolithos-Cylindrichnus association from estuarine point bar deposits in the Lower Cretaceous McMurray Formation, northeastern Alberta. Both active and abandoned associations are recognized. 17Distinguishing between fluvial and estuarine channels has important ramifications in genetic stratigraphy. Estuaries are generally deeper and wider than the associated fluvial chanicls (Dorjes and Howard, 1975). ‘Thus, transgression followed by establishment of ant estuaty could result in significant erosion of the pro-cxisting fresh water deposits. Dalrymple (1992) stated that estuaries commonly form