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Ichnology of a Pennsylvanian Equatorial Tidal Flat

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Concluding Remarks

Integrated stratigraphic, sedimentologic, and ichnologic analysis of the Waverly succession allows a detailed characterization of the sedimentary facies and associated trace fossils. Deposition occurred along a microtidal shoreline that was directly connected with the open sea. Paleogeographic information indicates an equatorial position.

Changes in diversity and abundance of trace fossils in the lower interval of the Waverly section reflect an overall shallowing-upward trend from lower-intertidal to fluvial facies. The highest ichnodiversity is recorded in the tidal-flat deposits, specifically in the sand-flat facies. A monospecific suite of myriapod trackways is present in the fluvial deposits towards the upper part of the lower interval.

The Waverly tidal-flat is characterized by heterogeneous distribution of biogenic structures. At a larger scale, intertidal zonation of trace fossils is evident in sand-, mixed-, and mud-flat areas of the tidal flat. Trace fossils are diverse and abundant in the sand flat, recording the activity of a varied benthic community composed of animals that were able to deal with the stressful conditions of the intertidal realm, probably as a result of the predictability of such an environment. On the other hand, mixed- to mud-flat deposits contain a low diversity of biogenic structures, reflecting extremely high temperatures and desiccation, as well as low-preservation potential due to lack of sandstone interbeds. At a smaller scale, spatial separation of species occurs within each zone of the tidal flat. Examples of ichnologic heterogeneity in Waverly include mounds characterized by dense aggregations of Protovirgularia bidirectionalis and small vertical burrows, and patchy distribution of Psammichnites implexus, which is commonly concentrated in ripple troughs.

The tidal-flat is a harsh environment where marine organisms are subjected to the extremes of terrestrial climate, heating, frost, and desiccation. Temperature, salinity, duration of exposure to subaerial conditions, energy, and substrate represent limiting factors in tidal-flat ecosystems. Changes in salinity and temperature, together with time of exposure, are extreme in the upper-intertidal zone and diminish toward the lower-intertidal zone. This gradient may have led to the increase in ichnodiversity in the sand-flat deposits at Waverly.

Ichnotaxonomic composition and dominance of horizontal structures of deposit feeders and grazers suggest a moderate- to low-energy coastal setting. However, sedimentologic and ichnologic evidence (e.g., deep gutter casts, truncation of vertical shafts) indicate occasional high-energy events that sculpted the tidal-flat surface. Substrate conditions also are highly variable, because tidal flats are regularly exposed and submerged by the tides, with concomitant fluctuations in pore-water content. Additionally, the physiography of the coast during low tide commonly is characterized by tide pools within an otherwise emergent area, which results in a range of substrate conditions along an isochronous surface. For example, morphologic variability of bivalve traces at Waverly was controlled in part by substrate fluidity. In the case of bivalve locomotion and escape structures, different morphotypes respond to the same style of motion, with substrate consistency playing a significant role in trace-fossil morphologic variability.

Analysis of Waverly tidal-flat deposits also reveals information on stratigraphic completeness. Amalgamated sandstones with preferential preservation of Lockeia siliquaria record a sequence of colonization events separated by erosional scouring and renewed deposition. Individual bedding planes represent time-averaged surfaces that record the work of successive communities of burrowing bivalves. Palimpsest surfaces are common in intertidal environments and provide evidence of complex depositional histories with considerable geologic time represented by units of relatively thin strata.

Analysis of crosscutting relationships, burrowing depth, wall sharpness, trophic types, and bauplan allows recognition of a tiering structure and ichnoguild model for the benthic fauna of the Waverly tidal flat. This tidal-flat community mainly occupied shallow tiers, with domiciles of bivalves and sea anemones being the deepest forms in the association.

Integrated ichnologic, sedimentologic, and sequence-stratigraphic information provides evidence of the depositional history and role of sea level during accumulation of tidal-flat sediments. The sedimentary units of the lower interval of the Waverly succession form part of four fining-upward parasequences of tidal-flat progradation, separated by successive flooding surfaces. These four parasequences form a progradational parasequence set. A major transgressive surface separates the fluvial facies at the tops of the lower parasequence set from the subtidal orthomyalinid packstones and wackestones that, together with the Spring Branch Limestone Member, represent a retrogradational parasequence set. In the Stull Shale Member, this set is represented by the orthomyalinid packstone and wackestone parasequence. Paleosols developed during regressive maximums on top of tidal-flat parasequences, and they do not indicate sequence boundaries. Some of the transgressive surfaces contain a firmground suite regarded as an example of the Glossifungites ichnofacies.

Study of other outcrops of the Stull Shale Member in eastern Kansas shed light on the environmental significance of the Waverly trace-fossil site in a broader regional context. The Waverly ichnofauna developed in a tidal flat dominated by normal-marine salinity and connected directly to the open sea. This tidal flat was located outside of a northeast-southwest-oriented embayment that opened toward the southwest. Trace-fossil assemblages from coeval localities in the northeast indicate brackish-water conditions in a restricted, marginal-marine setting.

The Waverly assemblage clearly qualifies as an example of the Cruziana ichnofacies. Presence of the Cruziana ichnofacies in intertidal environments represents an occurrence in water shallower than expected according to standard ichnofacies models. The classic onshore-offshore ichnofacies replacement model should be applied only in wave-dominated systems, because the opposite gradient is observed in tide-dominated systems. In fact, many deposits that were interpreted as having been formed in subtidal environments based on the presence of the Cruziana ichnofacies probably should be revised.

Attributes of tidal-flat ecosystems, such as heterogeneity, unrefined interactions, predictability, and high selective pressures, may have promoted evolutionary innovations. Comparative analysis of tidal-flat ichnofaunas through time supports this view. In the case of the Waverly ecosystem, the presence of Wilkingia and relatively deep bivalve burrows suggests incipient exploitation of the deep infaunal ecospace by bivalves, which occurred long before the Mesozoic revolution.

Acknowledgments

This monograph was written while Mángano and Buatois were recipients of a post-doctoral external award by the Argentinean Research Council (CONICET). Mángano and Buatois thank the Kansas Geological Survey for logistical support and access to facilities. Mángano also thanks financial support from Grants-in-Aid-of Research by Sigma Delta Epsilon, Graduate Women in Science, the Antorchas Foundation, the Paleontological Society, and the Mid-America Paleontological Society (MAP Award). We thank Tony Ekdale and Andrew Rindsberg for reviewing this monograph. Certain aspects of this research were discussed with the following colleagues, whose comments and information are gratefully acknowledged: Richard Bromley, Alan Byrnes, Tony Ekdale, Soren Jensen, Radek Mikuláš, Keith Miller, Ron Pickerill, John Pollard, Andrew Rindsberg, Adolf Seilacher, Alfred Uchman, and Lynn Watney. Mángano and Buatois especially thank Tom Stanley for his continuous support during their stay in Kansas and for sharing his experience in Kansas geology by locating outcrops of the Stull Shale Member, and Scott Beaty for his invaluable help and enlightening discussions. We also are grateful to Ricardo Olea for helping with the statistics of bivalve traces and for sharing with us the enthusiasm of developing this project. Janet Baker kindly shared unpublished information from her work on bioerosion in the Stull Shale Member orthomyalinid wackestones and packstones. Discussions with students of the Ichnology Course (taught at the University of Kansas by Mángano and Buatois) during a visit to the Waverly trace-fossil locality were particularly stimulating. Abdulrahman Alissa, Janet Baker, Alan Byrnes, and Craig Sundell are thanked for their help in the field. The late Al Kamb and Alice Hart were extremely helpful in curating the Waverly ichnofossils and locating trace-fossil specimens housed in the Museum of Invertebrate Paleontology of The University of Kansas. While revising type specimens, Mángano and Buatois were assisted by R. Prokop and V. Turek (National Museum, Prague), K. Westphal (Geology Museum, University of Wisconsin), M. Romano (Geology Department, Sheffield University), D. Lewis (Natural History Museum, London), W. J. Kennedy and E. Howlett (Oxford University Museum of Natural History), M. Dorling and R. Long (Sedgwick Museum, University of Cambridge), and Victor Padron and Alejandro Machado (Universidad Central de Caracas). Our warmest thanks to Mark Schoneweis and John Charlton, who provided the line-drawings and photographic illustrations for this work, respectively, and kindly helped us accomplish this project in time. We also thank Marla Adkins-Heljeson for editing the monograph and encouraging us to finish this project. Finally, Mángano and Buatois would like to acknowledge the following colleagues at the Kansas Geological Survey for their continuous encouragement and support: Abdulrahman Alissa, Scott Beaty, Peter Berendsen, Liz Brosius, Rex Buchanan, Alan Byrnes, Tim Carr, David Collins, Evan Franzeen, Kristie Luchtel, Sara Marcus, Dan Merriam, Ricardo Olea, Gina Ross, Tom Stanley, and Lynn Watney.

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