M. Gabriela Mángano

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.

Sequence stratigraphic and sedimentologic significance of biogenic structures from a late Paleozoic marginal- to open-marine reservoir, Morrow Sandstone, subsurface of southwest Kansas, USA

Abstract Integrated ichnologic, sedimentologic, and stratigraphic studies of cores and well logs from Lower Pennsylvanian oil and gas reservoirs (lower Morrow Sandstone, southwest Kansas) allow distinction between fluvio-estuarine and open marine deposits in the Gentzler and Arroyo fields. The fluvio-estuarine facies assemblage is composed of both interfluve and valley-fill deposits, encompassing a variety of depositional environments such as fluvial channel, interfluve paleosol, bay head delta, estuary bay, restricted tidal flat, intertidal channel, and estuary mouth. Deposition in a brackish-water estuarine valley is supported by the presence of a low diversity, opportunistic, impoverished marine ichnofaunal assemblage dominated by infaunal structures, representing an example of a mixed, depauperate Cruziana and Skolithos ichnofacies. Overall distribution of ichnofossils along the estuarine valley was mainly controlled by the salinity gradient, with other parameters, such as oxygenation, substrate and energy, acting at a more local scale. The lower Morrow estuarine system displays the classical tripartite division of wave-dominated estuaries (i.e. seaward-marine sand plug, fine-grained central bay, and sandy landward zone), but tidal action is also recorded. The estuarine valley displays a northwest–southeast trend, draining to the open sea in the southeast. Recognition of valley-fill sandstones in the lower Morrow has implications for reservoir characterization. While the open marine model predicts a “layer-cake” style of facies distribution as a consequence of strandline shoreline progradation, identification of valley-fill sequences points to more compartmentalized reservoirs, due to the heterogeneity created by valley incision and subsequent infill. The open-marine facies assemblage comprises upper, middle, and lower shoreface; offshore transition; offshore; and shelf deposits. In contrast to the estuarine assemblage, open marine ichnofaunas are characterized by a high diversity of biogenic structures representing the activity of a benthic fauna developed under normal salinity conditions. Trace fossil and facies analyses allow environmental subdivision of the shoreface-offshore successions and suggest deposition in a weakly storm-affected nearshore area. An onshore–offshore replacement of the Skolithos ichnofacies by the Cruziana ichnofacies is clearly displayed. The lower Morrow fluvio-estuarine valley was incised during a drop of sea level coincident with the Mississippian–Pennsylvanian transition, but was mostly filled during a subsequent transgression. The transgressive nature of the estuarine infill is further indicated by the upward replacement of depauperate brackish-water trace fossil assemblages by the open-marine Cruziana ichnofacies. Additional stratal surfaces of allostratigraphic significance identified within the estuary include the bayline surface, the tidal ravinement surface, the wave ravinement surface, and a basinwide flooding surface recording inundation of the valley interfluves. A younger sequence boundary within the lower Morrow is also recorded in the Gentzler field at the base of a forced regression shoreface, demarcated by the firmground Glossifungites ichnofacies, indicating a rapid basinward facies migration during a sea-level drop. Trace fossil models derived from the analysis of Mesozoic and Cenozoic reservoirs are generally applicable to the study of these late Paleozoic reservoirs. Pennsylvanian brackish-water facies differ ichnologically from their post-Paleozoic counterparts, however, in that they have: (1) lower trace fossil diversity, (2) lower degree of bioturbation, (3) scarcity of crustacean burrows, (4) absence of firmground suites, and (5) absence of ichnotaxa displaying specific architectures designed to protect the tracemaker from salinity fluctuations. Morrow open-marine ichnofaunas closely resemble their post-Paleozoic equivalents.

Trace fossils and sedimentary facies from a Late Cambrian‐Early Ordovician tide‐dominated shelf (Santa Rosita Formation, northwest Argentina): Implications for ichnofacies models of shallow marine successions

The Santa Rosita Formation is one the most widely distributed lower Paleozoic units of northwest Argentina. At the Quebrada del Salto Alto section, east of Purmamarca, Jujuy Province, it is represented by four sedimentary facies: thick‐bedded planar cross‐stratified quartzose sandstones (A), thin‐bedded planar cross‐stratified quartzose sandstones and mudstones (B), wave‐rippled sandstones and bioturbated mudstones (C), and black and greenish gray shales (D). Paleocurrent data, sandstone architecture, and sedimentary structures from facies A and B indicate bipolar/bimodal paleoflows, suggesting the action of tidal currents. The succession is interpreted as that of a tide‐dominated shelf, with only secondary influence of wave processes. Trace fossils are restricted to facies B and C. The Cruziana ichnocoenosis is preserved on the soles of thin‐bedded planar cross‐stratified quartzose sandstones (facies B). This ichnocoenosis consists of Conostichus isp., Cruziana omanica, C. semiplicata, C. cf. tortworthi,...

Sedimentary facies and environmental ichnology of a ?Permian playa-lake complex in western Argentina

Abstract A moderately diverse arthropod ichnofauna occurs in ?Permian ephemeral lacustrine deposits of the Paganzo Basin that crop out at Bordo Atravesado, Cuesta de Miranda, western Argentina. Sedimentary successions are interpreted as having accumulated in a playa-lake complex. Deposits include three sedimentary facies: (A) laminated siltstone and mudstone; (B) current-rippled cross-laminated very fine grained sandstone; and (C) climbing and wave-rippled cross-laminated fine-grained sandstone deposited by sheet floods under wave influence in the playa-lake complex. Analysis of facies sequences suggests that repeated vertical facies associations result from transgressive-regressive episodes of variable time spans. The Bordo Atravesado ichnofauna includes Cruziana problematica, Diplocraterion isp., cf. Diplopodichnus biformis, Kouphichnium ? isp., Merostomichnites aicunai, Mirandaichnium famatinense, Monomorphichnus lineatus, Palaeophycus tubularis, Umfolozia sinuosa and Umfolozia cf. U. longula . The assemblage is largely dominated by arthropod trackways and represents an example of the Scoyenia ichnofacies. Trace fossils are mostly preserved as hypichnial ridges on the soles of facies C beds, being comparatively rare in facies A and B. Ichnofossil preservation was linked to rapid influx of sand via sheet floods entering into the lake. Four taphonomic variants (types 1–4) are recognized, each determined by substrate consistency and time averaging. Type 1 is recorded by the presence of low density assemblages consisting of poorly defined trackways, which suggests that arthropods crawled in soft, probably slightly subaqueous substrates. Type 2 is represented by low to moderate density suites that include sharply defined trackways commonly associated with mud cracks, suggesting that the tracemakers inhabited a firm, desiccated lacustrine substrate. Type 3 displays features of types 1 and 2 and represents palimpsestic bedding surfaces, resulting from the overprint of terrestrial ichnocoenoses over previously formed softground suites. Type 4 differs from type 2 only in that assemblages display a high density of traces, recorded by numerous superimposed trackways, which suggests a major time gap of subaerial exposure before sheet flood entrance. Therefore, type 4 surfaces are mostly interpreted as track imprinted omission surfaces.

Taxonomic reassessment of the ichnogenus beaconichnus and additional examples from the Carboniferous of Kansas, U.S.A.

The ichnogenus Beaconichnus (Gevers 1973), an arthropod trace fossil, includes very different forms that comprise five ichnospecies, namely B. darwi‐nunt (Gevers 1971), B. gouldi (Gevers 1971), B. ahtarcticum (Gevers 1971), B. giganteum Gevers and Twomey 1982, and B. wrrighti Gevers and Twomey 1982. The original diagnosis of Beaconichnus is rather vague and potentially may accomodate virtually every arthropod trackway described from the fossil record. In view of these problems, the validity of Beaconichnus is reassessed and each of its ichnospecies is reviewed. We conclude that B. darwinum is a junior synonym of Diplopodichnus biformis Brady 1947; B. antarcticum should be regarded as Palmich‐niunt antarcticum; and B. wrighti is a nomen nudum. Additionally, we agree with previous proposals in considering B. gouldi as the senior synonym of B. giganteum, and including it in Diplichnites Dawson 1873. Therefore, we suggest that the ichnogenus Beaconichnus is best disregarded. Additionally, we describe specimen...

The ichnogenus Curvolithus revisited

The ichnogenus Curvolithus Fritsch, 1908, originally described from the Ordovician of the Prague Basin, typically comprises ribbonlike or tonguelike, flattened, endostratal traces with three rounded lobes on the upper surface. However, considerable confusion persists regarding the ichnotaxonomic status and diagnostic features of its ichnospecies. The type specimens of this ichnotaxon, overlooked in most subsequent reports, are redescribed herein. Curvolithus multiplex Fritsch, 1908, the type species, is retained for specimens with a trilobate upper surface and a quadralobate lower surface, in contrast to the criteria adopted by subsequent authors. The other ichnospecies originally proposed from the type locality, C. gregarius Fritsch, 1908, actually consists of a series of grouped parallel scratch marks forming ridges and should be removed from Curvolithus. Subsequently, four ichnospecies were defined: C.? davidis Webby 1970; C. annulatus Badve and Ghare 1978; C. aequus Walter et al. 1989; and C. manitouensis Maples and Suttner 1990. Curvolithus? davidis shows the typical trilobation of Curvolithus apparently in its lower surface, but the morphology of the upper surface is uncertain. Accordingly, it does not warrant ichnospecific assessment, and is regarded as a nomen dubium. The nature of the annulations on the trilobate upper surface of C. annulatus is unclear, and this ichnospecies is also best considered as a nomen dubium. Curvolithus aequus has a bilobate lower surface and probably represents washed out specimens of Didymaulichnus. Finally, C. manitouensis comprises specimens with a smooth, trilobate upper surface and a smooth, quadralobate lower surface, and is best regarded as a junior synonym of C. multiplex. Curvolithus multiplex has been used incorrectly for Curvolithus with a trilobate upper surface and a trilobate to unilobate lower surface. The new ichnospecies, Curvolithus simplex, is proposed herein for such traces. Curvolithus is interpreted as a locomotion trace (Repichnia) of endostratal carnivores, possibly gastropods, flatworms, or nemerteans. Curvolithus is a component of the Cruziana ichnofacies in shallow-marine facies, either of normal salinity or slightly brackish, in the latter case typically associated with fan deltas.