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Carboniferous Psammichnites: Systematic re-evaluation, taphonomy and autecology

The ichnogenus Psammichnites Torell 1870 includes a wide variety of predominantly horizontal, sinuous to looped, backfilled traces, characterized by a distinctive median dorsal structure. Though commonly preserved in full relief on upper bedding surfaces, some ichnospecies of Psammichnites may be preserved in negative hyporelief. Psammichnites records the feeding activities of a subsurface animal using a siphon-like device. Several ichnogenera reflect this general behavioral pattern, including Plagiogmus Roedel 1929 and the Carboniferous ichnogenera Olivellites Fenton and Fenton 1937a and Aulichnites Fenton and Fenton 1937b. Based on analysis of specimens from the United States, Spain, and the United Kingdom, three Carboniferous ichnospecies of Psammichnites are reviewed in this paper: P. plummeri (Fenton and Fenton, 1937a), P. grumula (Romano and Meléndez 1979), and P. implexus (Rindsberg 1994). Psammichnites plummeri is the most common Carboniferous ichnospecies and is characterized by a relatively straight, continuous dorsal ridge/groove, fine transverse ridges, larger size range, and non-looping geometric pattern. It represents a grazing trace of deposit feeders. Psammichnites grumula differs from the other ichnospecies of Psammichnites by having median dorsal holes or protruding mounds. The presence of mounds or holes in P. grumula suggests a siphon that was regularly connected to the sediment-water interface. This ichnospecies is interpreted as produced by a deposit feeder using the siphon for respiration or as a device for a chemosymbiotic strategy. Psammichnites implexus is characterized by its consistently smaller size range, subtle backfill structure, and tendency to scribble. Although displaying similarities with Dictyodora scotica , P. implexus is a very shallow-tier, grazing trace. Changes in behavioral pattern, preservational style, and bedform morphology suggest a complex interplay of ecological and taphonomic controls in Carboniferous tidal-flat Psammichnites . A first distributional pattern consists of guided meandering specimens preserved in ripple troughs, probably reflecting food-searching of buried organic matter concentrated in troughs. A second is recorded by concentration of Psammichnites on ripple crests and slopes. In some cases, the course is almost straight to slightly sinuous and closely follows topographic highs, suggesting a direct control of bedform morphology on trace pattern. Occurrences of Carboniferous Psammichnites most likely represent an opportunistic strategy in marginal-marine settings. Analysis of Carboniferous Psammichnites indicates the presence of a siphon-like device in the producer and reestablishes the possibility of a molluscan tracemaker.

Ichnos: An International Journal for Plant and Ani

The insect trace fossil Tonganoxichnus from the middle Pennsylvanian of Indiana: Paleobiologic and paleoenvironmental implications

The ichnogenus Tonganoxichnus , produced by one or more monuran insect taxa, is now recorded from the Middle Pennsylvanian Mansfield Formation of Indiana. Tonganoxichnus is a resting trace that has three important implications. First, it represents a recurrent behavioral pattern in Upper Carboniferous to Lower Permian marginal marine environments of North America. Second, it provides finely resolved anatomical information for axial and appendicular body structures and behaviors that are difficult to determine from body‐fossil material alone. Third, integrated sedimentologic and ichnologic observations indicate that the Tonganoxichnus assemblage, inclusive of other ichnotaxa, is common in tidal rhythmites that were developed under freshwater conditions, probably in the innermost part of estuarine systems, close to or at the fluvioestuarine transition.

Ichnos: An International Journal for Plant and Ani

A new pterosaur tracksite from the Jurassic Summerville formation, near Ferron, Utah

Pterosaur tracks (cf. Pteraichnus ) from the Summerville Formation of the Ferron area of central Utah add to the growing record of Pteraichnus tracksites in the Late Jurassic Summerville Formation and time-equivalent, or near time-equivalent, deposits. The site is typical in revealing high pterosaur track densities, but low ichnodiversity suggesting congregations or “flocks” of many individuals. Footprint length varies from 2.0 to 7.0 cms. The ratio of well-preserved pes:manus tracks is about 1:3.4. This reflects a bias in favor of preservation of manus tracks due to the greater weight-bearing role of the front limbs, as noted in other pterosaur track assemblages. The sample also reveals a number of well-preserved trackways including one suggestive of pes-only progression that might be associated with take off or landing, and another that shows pronounced lengthening of stride indicating acceleration. One well-preserved medium-sized theropod trackway ( Therangospodus ) and other larger theropod track casts (cf. Megalosauripus ) are associated with what otherwise appears to be a nearly monospecific pterosaur track assemblage. However, traces of a fifth pes digit suggest some tracks are of rhamphorynchoid rather than pterodactyloid origin, as usually inferred for Pteraichnus . The tracks occur at several horizons in a thin stratigraphic interval of ripple marked sandstones and siltstones. Overall the assemblage is similar to others found in the same time interval in the Western Interior from central and eastern Utah through central and southern Wyoming, Colorado, northeastern Arizona, and western Oklahoma. This vast “ Pteraichnus ichnofacies,” with associated saurischian tracks, remains the only ichnological evidence of pre-Cretaceous pterosaurs in North America and sheds important light on the vertebrate ecology of the Summerville Formation and contiguous deposits.

Utah

The history of dinosaur footprint discoveries in Wyoming with emphasis on the Bighorn Basin

Dinosaur traces are well known from the western United States in the Colorado Plateau region (Utah, Colorado, New Mexico, and Arizona). Utah contains the greatest abundance of known and documented dinosaur footprints and trackways. Far less well known, however, is the occurrence and distribution of dinosaur footprint-bearing horizons in Wyoming. Scientific studies over the past 10 years have shown that three of the four Middle and Upper Jurassic formations in northern Wyoming contain dinosaur footprints. Two of the footprint-bearing horizons are located in geologic intervals that were once thought to have been deposited in offshore to nearshore marine settings and represent rare North American examples of Middle Jurassic (Bajocian and Bathonian) dinosaur remains. Some of these new Wyoming sites can be correlated to known dinosaur footprint-bearing horizons or intervals in Utah. Wyoming has a great potential for additional discoveries of new dinosaur footprint-bearing horizons, and further prospecting and study is warranted and will ultimately lead to a much better understanding of the geographic distribution and behavior of the potential footprint-makers.

Wyoming

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 accommodate 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 specimens collected from the Late Carboniferous Tonganoxie Sandstone Member (Stranger Formation) of eastern Kansas, ascribed herein to Diplopodichnus biformis and Diplichnites gouldi , which include examples of intergradations between both ichnotaxa, and provide synonymy lists for both ichnospecies.

Kansas

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 faciès 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 (faciès B). This ichnocoenosis consists of Conostichus isp., Cruziana omanica, C. semiplicata, C. cf. tortworthi, Cruziana isp. Helminthopsis abeli, Monomorphichnus bilinearis, M. multilineatus, Palaeophycus tubularis, Rusophycus carbonarias, R. latus , and R. isp. The occurrence of Cruziana semiplicata , C. omanica, C. cf. tortworthi , and Rusophycus latus supports a Late Cambrian‐Tremadoc age. Slabbing of Cruziana shows complex interactions between biologic and sedimentologic processes, and suggests a predominance of exhumed traces, washed out and recast by tractive sand deposition. Sandstone soles are densely packed with biogenic structures and exhibit distinctive clusters of Rusophycus isp. that most likely represent trilobite nesting burrows. The Cruziana ichnocoenosis records the resident fauna of a protected, lower intertidal to subtidal interbar setting. The Skolithos ichnocoenosis is represented by high to low density vertical burrows of Skolithos linearis , which extend downwards to the quartzose sandstone soles of faciès B and cross the Cruziana ichnocoenosis. The Skolithos ichnocoenosis represents colonization by suspension‐feeding organisms following a major change in environmental conditions, related to the migration of lower intertidal to subtidal sandwaves. The Planolites ichnocoenosis consists exclusively of Planolites montanus within mudstones overlying wave‐rippled sandstones (facies C). The Planolites ichnocoenosis records opportunistic colonization by inf aunal deposit feeders that mined the organic‐rich fine‐grained sediment during the waning phase of storms that scoured organic detritus from the sea bottom. The section records, from base to top, a Cruziana‐Skolithos ichnofacies zone, a Skolithos ichnofacies zone and an unbioturbated zone typified by the thick‐bedded cross‐stratified quartzose sandstone (fades A). This trend reflects progressively higher energy conditions linked to the establishment of a large sand wave complex. The presence of a mixed Cruziana‐Skolithos ichnofacies in the lower interval reflects changes in substrate and energy levels, rather than water depth. Accordingly, contrasting ichnocoenoses from interbars (Cruziana) and sandwaves (Skolithos) must be considered an example of ichnofacies controlled by local parameters instead of general bathymétrie trends. Conversely, the vertical replacement of the Cruziana ichnofacies by the Skolithos ichnofacies towards the middle interval of the section reflects the environmental changes associated with the transition between the intertidal and subtidal zones. As overall tidal energy increases from supratidal to subtidal settings, the Skolithos ichnofacies tends to occur seaward of the Cruziana ichnofacies in tide‐dominated shallow marine environments. Therefore, onshore‐offshore ichnofacies replacement in tide‐dominated shallow seas is opposite to that in wave‐dominated marine settings.

Ichnos: An International Journal for Plant and Ani

Application of morphologic burrow interpretations to discern continental burrow architects: Lungfish or crayfish?

A methodology for trace fossil identification using burrowing signatures is tested by evaluating ancient and modern lungfish and crayfish burrows and comparing them to previously undescribed burrows in a stratigraphic interval thought to contain both lungfish and crayfish burrows. Permian burrows that bear skeletal remains of the lungfish Gnathorhiza , from museum collections, were evaluated to identify unique burrow morphologies that could be used to distinguish lungfish from crayfish burrows when fossil remains are absent. The lungfish burrows were evaluated for details of the burrowing mechanism preserved in the burrow morphologies together forming burrowing signatures and were compared to new burrows in the Chinle Formation of western Colorado to test the methodology of using burrow signatures to identify unknown burrows. Permian lungfish aestivation burrows show simple, nearly vertical, unbranched architectures and relatively smooth surficial morphologies with characteristic quasi‐horizontal striae on the burrow walls and vertical striae on the bulbous terminus. Burrow lengths do not exceed 0.5 m. In contrast, modern and ancient crayfish burrows exhibit simple to highly complex architectures with highly textured surficial morphologies. Burrow lengths may reach 4 to 5 m. Burrow morphologies unlike those identified in Gnathorhiza aestivation burrows were found in four burrow groups from museum collections. Two of these groups exhibit simple architectures and horizontal striae that were greater in sinuosity and magnitude, respectively. One of these burrows contains the remains of Lysoro‐phus , but the burrow surface reveals no reliable surficial characteristics. It is not clear whether Lysorophus truly burrowed or merely occupied a pre‐existing structure. The other two groups exhibit surficial morphologies similar to those found on modern and ancient crayfish burrows and may provide evidence of freshwater crayfish in the Permian. Burrows from the Upper Triassic Chinle Formation in western Colorado exhibit simple to moderately complex architectural morphologies, ranging from predominantly vertical, unbranched, with little or no chamber development to predominantly vertical, few branches, and with minor chamber development. Surficial burrow morphologies are moderate to highly textured. The burrows have scrape marks, scratch marks, mud and lag‐liners, knobby surfaces, pleopod striae, and body impressions. Although no fossil remains of the burrowing organism were found within or associated with the Chinle burrows from western Colorado, the similarity of architectural and surficial burrow morphologies to those in the Chinle of Canyonlands, Utah and to modern crayfish burrows, clearly indicates that the Colorado burrows are the product of burrowing crayfish rather than lungfish. Evaluation of burrowing signatures preserved in the architectural and surficial burrow morphologies is a very useful tool to compare and contrast Chinle burrows from different regions on the Colorado Plateau. Documentation of crayfish burrows in the Chinle of Utah and Colorado strongly suggests that other large‐diameter Chinle burrows elsewhere on the Colorado Plateau and in stratigraphically equivalent units may also be the product of crayfish activity.

Colorado, Utah