Luis A. Buatois

Colonization of Brackish-Water Systems through Time: Evidence from the Trace-Fossil Record

Abstract Trace fossils in estuarine deposits of different ages have been compared to evaluate colonization history of brackish-water ecosystems and to calibrate trace-fossil, brackish-water models with respect to geologic time. This comparative analysis reveals that, although the colonization of marginal-marine, brackish-water environments was a long-term process that spanned most of the Phanerozoic, this process of invasion of fully marine organisms into restricted, marginal-marine habitats did not occur at a constant rate. Five major colonization phases can be distinguished. The first phase (Ediacaran–Ordovician) represents a prelude to the major invasion that occurred during the rest of the Paleozoic. While Ediacaran–Cambrian ichnofaunas seem to be restricted to the outermost zones of marginal-marine depositional systems, Ordovician assemblages show some degree of landward expansion within brackish-water ecosystems. Intensity of bioturbation and ichnodiversity levels were relatively low during this phase. The second phase (Silurian–Carboniferous) is marked by the appearance of more varied morphologic patterns and behavioral strategies, resulting in a slight increase in ichnodiversity. While previous assemblages were arthropod dominated, brackish-water Silurian–Carboniferous ichnofaunas include structures produced by bivalves, ophiuroids, and polychaetes. Ichnofaunas from the third phase (Permian–Triassic) seem to be characterized by the presence of crustacean burrows, reflecting the late Paleozoic crustacean radiation and adaptation of some groups to brackish-water conditions. The fourth phase (Jurassic–Paleogene) is typified by a remarkable increase in ichnodiversity and intensity of bioturbation of estuarine facies. Colonization occurred not only in softgrounds and firmgrounds, but also in hardgrounds and xylic substrates. The fifth phase (Neogene–Recent) records the onset of modern brackish-water benthos. Although still impoverished with respect to their fully marine counterparts, brackish-water ichnofaunas may reach moderately high diversities, particularly in middle- and outer-estuarine regions, and degree of bioturbation may be high in certain estuarine subenvironments. Comparative analysis of brackish-water ichnofaunas through geologic time provides valuable evidence to understand colonization of marginal-marine environments through the Phanerozoic, and allows for calibration of ichnologic models that may aid in the recognition of estuarine valley-fill deposits in the stratigraphic record.

The ichnologic record of the continental invertebrate invasion; evolutionary trends in environmental expansion, ecospace utilization, and behavioral complexity

The combined study of continental trace fossils and associated sedimentary facies provides valuable evidence of colonization trends and events throughout the Phanerozoic. Colonization of continental environments was linked to the exploitation of empty or under-utilized ecospace. Although the nonmarine trace fossil record probably begins during the Late Ordovician, significant invasion of nonmarine biotopes began close to the Silurian-Devonian transition with the establishment of a mobile arthropod epifauna (Diplichnites ichnoguild) in coastal marine to alluvial plain settings. Additionally, the presence of vertical burrows in Devonian high-energy fluvial deposits reflects the establishment of a stationary, deep suspension-feeding infauna of the Skolithos ichnoguild. The earliest evidence of plant-arthropod interaction occurred close to the Silurian-Devonian boundary, but widespread and varied feeding patterns are known from the Carboniferous. During the Carboniferous, permanent subaqueous lacustrine settings were colonized by a diverse, mobile detritus-feeding epifauna of the Mermia ichnoguild, which reflects a significant paleoenvironmental expansion of trace fossils. Paleozoic ichnologic evidence supports direct routes to the land from marginal marine environments, and migration to lakes from land settings. All nonmarine sedimentary environments were colonized by the Carboniferous, and subsequent patterns indicate an increase in ecospace utilization within already colonized depositional settings. During the Permian, back-filled traces of the Scoyenia ichnoguild record the establishment of a mobile, intermediate-depth, deposit-feeding infauna in alluvial and transitional alluvial-lacustrine sediments. Diversification of land plants and the establishment of ecologically diverse plant communities through time provided new niches to be exploited by arthropods. Nevertheless, most of the evolutionary feeding innovations took place relatively early, during the Late Paleozoic or early Mesozoic. A stationary deep infauna, the Camborygma ichnoguild, was developed in Triassic transitional alluvial-lacustrine deposits. Terrestrial environments hosted the rise of complex social behavioral patterns, as suggested by the probable presence of hymenopteran and isopteran nests in Triassic paleosols. An increase in diversity of trace fossils is detected in Triassic-Jurassic eolian deposits, where the ichnofauna displays more varied behavioral patterns than their Paleozoic counterparts. Also, a mobile, intermediate-depth, deposit-feeding infauna, the Vagorichnus ichnoguild, was established in deep lake environments during the Jurassic. In contrast to Paleozoic permanent subaqueous assemblages typified by surface trails, Jurassic ichnocoenoses are dominated by infaunal burrows. High density of infaunal deposit-feeding traces of the Planolites ichnoguild caused major disruption of lacustrine sedimentary fabrics during the Cretaceous. Most insect mouthpart classes, functional feeding groups, and dietary guilds were established by the end of the Cretaceous. Diversification of modern insects is recorded by the abundance and complexity of structures produced by wasps, bees, dung-beetles, and termites in Cretaceous-Tertiary paleosols. The increase in bioturbation migrated from fluvial and lake-margin settings to permanent subaqueous lacustrine environments through time.

Ichnology of an Upper Carboniferous fluvio-estuarine paleovalley: The Tonganoxie Sandstone, Buildex Quarry, Eastern Kansas, USA

Tidal rhythmites of the Tonganoxie Sandstone Member (Stranger Formation, Douglas Group) at Buildex Quarry, eastern Kansas, contain a relatively diverse ichnofauna. The assemblage includes arthropod locomotion (Dendroidichnites irregulare, Diplichnites gouldi types A and B, Diplopodichnus biformis, Kouphichnium isp., Mirandaichnium famatinense, and Stiaria intermedia), resting (Tonganoxichnus buildexensis) and feeding traces (Stiallia pilosa, Tonganoxichnus ottawensis); grazing traces (Gordia indianaensis, Helminthoidichnites tenuis, Helminthopsis hieroglyphica); feeding structures (Circulichnis montanus, Treptichnus bifurcus, Treptichnus pollardi, irregular networks), fish traces (Undichna britannica, Undichna simplicitas), tetrapod trackways, and root traces. The taxonomy of some of these ichnotaxa is briefly reviewed and emended diagnoses for Gordia indianaensis and Helminthoidichnites tenuis are proposed. Additionally, the combined name Dendroidichnites irregulare is proposed for nested chevron trackways. Traces previously regarded as produced by isopods are reinterpreted as myriapod trackways (D. gouldi type B). Trackways formerly interpreted as limulid crawling and swimming traces are assigned herein to Kouphichnium isp and Dendroidichnites irregulare, respectively. Taphonomic analysis suggests that most grazing and feeding traces were formed before the arthropod trackways and resting traces. Grazing/feeding traces were formed in a soft, probably submerged substrate. Conversely, the majority of trackways and resting traces probably were produced subaerially in a firmer, dewatered and desiccated sediment. The Buildex Quarry ichnofauna records the activity of a terrestrial and freshwater biota. The presence of this assemblage in tidal rhythmites is consistent with deposition on tidal flats in the most proximal zone of the inner estuary, between the maximum landward limit of tidal currents and the salinity limit further towards the sea.

Animal-substrate interactions in freshwater environments: applications of ichnology in facies and sequence stratigraphic analysis of fluvio-lacustrine successions

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

Trace fossils from a carboniferous turbiditic lake: Implications for the recognition of additional nonmarine ichnofacies

A high‐diversity ichnofauna from a Late Carboniferous turbiditic glacial lake at Cantera La Laja, northwest Argentina, includes Aulichnites cf. A. parkerensis, cf. Bergaueria isp., Circulichnis nton‐tanus, Cochlichnus anguineus, Gordia marina, Haplotichnus india‐naensis, Helminihoidichnites tenuis, Helminthopsis tenuis, Mermia carickensis, Orchesteropus atavus, Palaeophycus tubularis, Planolites beverleyensis, Rusophycus isp., Treptichnus pollardi, Undichnia britannica, Vndichnia insolencia, string pits, and rhomboidal traces. Fifteen forms are present in fine to very fine sandstones and siltstones deposited by underflow currents, muddy turbidity currents, and suspension sedimentation in the deepest parts of the lake. Underflow and turbidity currents provided food and oxygen to deep‐lake settings, allowing the development of a rich biota. Alternation of fine‐grained laminae favored preservation of delicate and very small biogenic structures. The ichnofauna analyzed is remarkably different from the Sco‐yen...

Decoupling of body-plan diversification and ecological structuring during the Ediacaran-Cambrian transition: evolutionary and geobiological feedbacks.

The rapid appearance of bilaterian clades at the beginning of the Phanerozoic is one of the most intriguing topics in macroevolution. However, the complex feedbacks between diversification and ecological interactions are still poorly understood. Here, we show that a systematic and comprehensive analysis of the trace-fossil record of the Ediacaran–Cambrian transition indicates that body-plan diversification and ecological structuring were decoupled. The appearance of a wide repertoire of behavioural strategies and body plans occurred by the Fortunian. However, a major shift in benthic ecological structure, recording the establishment of a suspension-feeder infauna, increased complexity of the trophic web, and coupling of benthos and plankton took place during Cambrian Stage 2. Both phases were accompanied by different styles of ecosystem engineering, but only the second one resulted in the establishment of the Phanerozoic-style ecology. In turn, the suspension-feeding infauna may have been the ecological drivers of a further diversification of deposit-feeding strategies by Cambrian Stage 3, favouring an ecological spillover scenario. Trace-fossil information strongly supports the Cambrian explosion, but allows for a short time of phylogenetic fuse during the terminal Ediacaran–Fortunian.

The paradox of nonmarine ichnofaunas in tidal rhythmites; integrating sedimentologic and ichnologic data from the Late Cretaceous of eastern Kansas, USA

The occurrence of trace fossil assemblages dominated by arthropod trackways and surface grazing trails within Carboniferous tidal rhythmites has puzzled sedimentologists and ichnologists, who interpreted them either as marine or nonmarine. The Virgilian (Stephanian) Tonganoxie Sandstone Member (Stranger Formation) at Buildex Quarry (eastern Kansas) consists, for the most part, of planar-laminated coarse grained siltstones deposited on an upper tidal flat, close to or at the fluvial-estuarine transition of a macrotidal estuarine paleovalley. Recurrent thickness fluctuations demonstrate the strong influence of tidal process and provide evidence that these deposits are tidal rhythmites, with thicker strata representing spring tides and thinner ones recording neap tides. The Buildex sequence hosts a moderately diverse ichnofauna composed of arthropod trackways (Dendroidichnites irregulare, Diplichnites gouldi, Diplopodichnus bifurcus, Kouphichnium isp., Mirandaichnium famatinense, Stiallia pilosa, Stiaria intermedia), grazing traces (Gordia indianaensis, Helminthoidichnites tenuis, Helminthopsis hieroglyphica), subsurface feeding traces (Treptichnus bifurcus, T. pollardi, irregular networks), apterygote insect resting and feeding traces (Tonganoxichnus buildexensis, T. ottawensis), fish traces (Undichna britannica, U. simplicitas), and tetrapod trackways. In contrast to trace fossil assemblages from brackish-water estuarine settings, the Buildex ichnofauna is characterized by moderate to relatively high ichnodiversity, ichnotaxa commonly present in terrestrial/freshwater environments, dominance of surface trails and absence of burrows, dominance of temporary structures produced by a mobile deposit-feeder fauna, a mixture of traces belonging to the Scoyenia and Mermia ichnofacies, moderate density of individual ichnotaxa, and absence of monospecific suites. This ichnofauna is thought to record the activity of a typical freshwater/terrestrial benthos. The presence of this mixed freshwater/terrestrial ichnofauna in tidal rhythmites is regarded as indicative of tidal flats that were developed in the most proximal zone of the inner estuary under freshwater conditions, more precisely in a zone between the maximum limit of landward tidal currents and the salinity limit further towards the sea. Although lithofacies distribution in estuarine valleys is mainly salinity-independent, the distribution of benthos is not. Accordingly, ichnologic studies have the potential to provide a high-resolution delineation of fluvio-estuarine transitions.

Trace fossils from Carboniferous floodplain deposits in western Argentina: implications for ichnofacies models of continental environments

Abstract Floodplain deposits from the Carboniferous Tupe Formation of western Argentina contain a low-diversity ichnofauna of invertebrates and plants. The assemblage consists of Archaeonassa fossulata, Didymaulichnus lyelli, Helminthoidichnites tenuis, Palaeophycus tubularis, Planolites isp., and root traces. This ichnofauna is characterized by the dominance of very simple forms, superficial or very shallow trace fossils, combination of locomotion, grazing, and dwelling structures, and production by arthropods and vermiform organisms. Primary sedimentary fabrics are disturbed only by plant trace fossils while animal trace fossils are mostly restricted to bedding surfaces, resulting in the virtual absence of bioturbation. The poorly-defined morphology of the ichnofossils, and the absence of backfilled striated trace fossils and structures indicative of subaerial exposure suggest subaqueous production. The envisaged depositional environment is a freshwater body developed in a floodplain and overfilled by overbank deposits. A review of floodplain ichnofaunas documented from the stratigraphic record indicates that two recurrent assemblages may be recognized. The first assemblage is characterized by low to rarely moderate diversity of invertebrate trace fossils, moderate to high diversity of vertebrate structures, and common presence of meniscate backfilled structures, bilobate trace fossils with scratch marks, arthropod trackways and tetrapod trackways. It is regarded as an example of the Scoyenia ichnofacies developed in desiccated floodplains. The second type of floodplain assemblage, illustrated by the example documented in this paper, is characterized by grazing trails, locomotion trails and dwelling burrows, low to rarely moderate ichnodiversity, simple trace fossils, and superficial to very shallow structures. Although formed in floodplain environments also, this type of assemblage is remarkably different from the Scoyenia ichnofacies. It is here regarded as an example of the Mermia ichnofacies. Basic features of these ichnofaunas reflect subaqueous production of the assemblage. The lower ichnodiversity of these floodplain assemblages in comparison with their equivalents from lacustrine basins is probably an expression of the less stable conditions and temporary nature of floodplain water bodies. Our study supports the view that ichnofacies are not indicators of sedimentary environments but reflect sets of environmental factors.

Invertebrate Ichnology of Continental Freshwater Environments

SUMMARY: The study of continental ichnofaunas has shown an explosive development during the last decade. At present, three continental archetypal ichnofacies are accepted: the Scoyenia, Mermia , and Coprinisphaera ichnofacies. Integration of ichnologic, sedimentologic, and paleobiologic information is very useful in facies analysis and sequence stratigraphy of continental successions.

Ichnology of Carboniferous tide-influenced environments and tidal flat variability in the North American Midcontinent

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