A Crinoid Lagerstatte from Maysville, Kentucky: Paleoautecology And
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A crinoid Lagerstätte from Maysville, Kentucky: paleoautecology and taphonomy A thesis submitted to the Graduate School of the University of Cincinnati in partial fulfillment of the requirements for the degree of Master of Science in the Department of Geology McMicken College of Arts and Sciences by Mason Jane Milam B.A. Hunter College City University of New York 1993 Committee Chair: David L. Meyer, Ph.D. Abstract A Lagerstätte of Glyptocrinus decadactylus collected from the Upper Fairview Formation at Maysville, Kentucky, USA, yields new insights into the paleoautecology of camerate crinoids of the Late Ordovician. The Lagerstätte represents an autochthonous community comprised of a single mudstone interval representing an obrution deposit and containing over 400 individual glyptocrinids, including over 250 calyces. The crinoids had colonized a rise of relatively high energy within the deep subtidal zone where they were partially sheltered by plant matter to which they were attached, as suggested by columns terminating distally in coiled holdfasts of relatively consistent diameter. Turbidity flows originating in storm events led to the suspension of fine sediments by a lofting plume that intruded into and buried the crinoids in their habitat. The crinoid and plant stand on the rise may have created additional friction that slowed the sediment plume and induced deposition, signifying burial facilitated by a feedback process. Some specimens had been dead for a period of time prior to burial, suggesting more than one earlier killing event, likely related to the storms that ultimately caused the obrution. Other specimens were apparently killed by the obrution event and subjected to little or no scavenging in the resulting anoxic conditions, leading to excellent preservation. Most specimens are in trauma position, although more than 6% were preserved in spread-fan feeding posture, aboral surface up. The latter position is indicative of very rapid burial, and is often seen in stalked crinoids capable of neither crawling nor autotomizing their stems to escape. The high density of the crinoid community was accommodated by tiering, in which, in addition to the plant matter, the crinoids employed the columns of earlier-settled specimens as attachment sites; differing column lengths positioned the crinoids at various i levels within the community. Further strategy for enduring crowded conditions may have been the maintenance of small body size. Taphonomic evidence and the crinoids' own adaptive strategies suggest a community that had quickly and opportunistically colonized a zone with ample attachment possibilities, high enough current velocities for sufficient feeding, yet sheltered enough to accommodate the crinoids' weak attachment capacity in all but the strongest storms. ii iii Acknowledgements From the beginning, Dave Meyer has gone above and beyond in championing my efforts to further my education this late in the game. His kindness and support have been invaluable. Brenda Hunda provided friendship, advice, and access to fossil material. Carlton Brett’s invertebrate paleontology classes were consistently exciting and inspiring. Dan Cooper discovered and gently excavated the slab. Steve Felton knew where he’d found it and took us back there, while Bob Bergstein provided financial assistance; both of these fine Dry Dredgers were full of good advice and encouragement. Roger Cuffey graciously prepped and identified the bryozoans. Ben Dattilo very generously instructed in thin-sectioning, sanding, and polishing of mudstones, as well as providing the XRF scans, which Barry Maynard kindly helped to interpret. Patrick was very patient with my frustrations when it came to certain computer applications, and for that, as well as every kind of support imaginable, thank you. iv Table of Contents Title Page Abstract i Acknowledgements iv Table of Contents v List of Figures vii List of Tables x Introduction 1 Stratigraphic Setting 4 Maysville West 7 The Lagerstätte 10 Materials and Methods 14 Taphonomy: Introduction 30 Obrution and Autochthony 32 Paleoenvironment 35 Crinoid Remains: Introduction 38 Measurements: Materials and Methods 39 Results and Discussion 39 Preservation: Materials and Methods 47 Results and Discussion 53 Implications of Early Decay 65 Orientation: Materials and Methods 68 Results and Discussion 71 Incidental Elements 82 Geochemistry 88 Mechanism for Obrution 94 Sediment Capture 100 v Paleoautecology 102 Morphology of Glyptocrinus decadactylus 104 Implications for Paleobathymetry 104 Implications for Preservation Position 107 Implications for Attachment 110 The Question of Autotomy 111 The Community 114 The Holdfast Forest 118 Community Strategy 123 Comparison with Other Lagerstätten 129 Conclusions 134 References 136 Appendices 145 vi List of Figures Figure 1. A) General location of study area. B) Excellent illustration of Glyptocrinus decadactylus by John Agnew. C) Stratigraphic column of Maysville West with arrow indicating approximate site of crinoid slab. Column modified from T. Schramm, 2011 Masters. Figure 2. The principal sedimentary environments of the Cincinnatian. Illustration after Steve Holland in A Sea Without Fish, Indiana University Press, 2009. Figure 3. A) The Maysville West roadcut is the NE-SW trending gray streak above the center of this Google Earth photograph. GPS of cut: 38° 40.407’ North, 83° 47.830’ West. B) Northwestern side of the Maysville West roadcut. The Bellevue Limestone is at top and the Fairview Formation is roughly the next three benches down. The light gray limestone bed indicated by arrow is a seismite horizon interpreted as part of the seafloor heavily disturbed by a paleoearthquake. The crinoid bed was 2-3 meters above this horizon. C) Site of Lagerstätte indicated by arrow. Photo by D.L. Meyer. Photographs A and B and information provided by James St. John ("A" via Google Terrametrics). Figure 4. Two views of the reassembled slab. Most pieces fit together perfectly. Note the frequent overlap of individual slab pieces: measurements for depth were taken for the deposit as a single unit, not as individual pieces. Figure 5. Each individual section of the slab was assigned a letter from A to T, with the largest, first-prepared piece designated "M" or M-slab. As M-slab was originally five pieces, these are depicted as M-1 through on M-5. Not every piece of the slab was prepared. Figure 6. Schematic artwork from photograph of slab piece. Non-crinoidal elements appear yellow in artwork. A) Slab piece A. B) Slab piece B. C) Slab piece C. D) Slab piece D, which was originally in two pieces, later rejoined. E) Slab piece E. F) Slab piece F. There are no non-crinoidal elements. G) Slab piece G. There are no non-crinoidal elements. H) Slab piece H. I) Slab piece I, which was still being prepped at time of photo. The brown spot at left is the lower surface of the slab as it appeared prior to preparation. J) Slab piece J. K) Slab piece K. Photograph currently unavailable. L) Slab piece M. M) Slab piece T. Photograph currently unavailable. Figure 7. Histogram comparing calyx heights of G. decadactylus from both Maysville slab and local Cincinnatian. Figure 8. Scatterplot comparing the stem lengths and calyx heights of the Maysville slab G. decadactylus. Figure 9. Idealized examples of taphonomic grades. vii Figure 10. Examples of calyx orientation. A) Crinoid in oblique orientation directly above another specimen in lateral orientation. B) Examples of crinoids in oral-aboral position, the “starburst.” C) Crinoids in lateral position, the “modified shaving brush." Figure 11. Schematic diagram showing hypothesized stalked crinoids in: A) parabolic filter fan (feeding) position, B) trauma position of crinoid with non-muscular arm articulations (the “modified” shaving brush), C) trauma position of crinoid with well-muscled arm articulations (the shaving brush). Modified after Baumiller, et al., in Echinoderm Paleontology, 2008. Figure 12. Examples of preservation effects. A) Well-preserved specimen missing stem at distal end. B, C) Specimens showing marks of scavenging on their calyces. D,E) Specimens in which the arms are lost distal to physical control. F) Heavily decayed specimens. Figure 13. Grid system from which orientation was plotted for slab pieces M, A, B, C, D, H, I, J, and K as a single unit. Figure 14. Detail of slab. Figure 15. Orientation from slab pieces M, A, B, C, D, H, I, J, and K as a single unit. Total count: 397 elements. A) Laterally-oriented calyces in opening direction. B) Columns. Those without a coil are arbitrarily directed. C) Combined A and B. Figure 16. Orientation of calyces and stems for three defined sections, as highlighted on page right. Number of specimens in each dataset listed on rose diagram, lower right.: A) Section 1. B) Section 2. (C) Section 3. Figure 17. Orientation of fossil material for four defined areas, as highlighted on page right. Number of specimens in each dataset listed on rose diagram, lower right. Note that areas overlap. A) North. B) South. C) East. D) West. Figure 18. These crowns are relatively consistent for orientation, though not direction. Figure 19. Two crinoids, "A" in starburst, "B" in modified shaving brush, preserved at the same level on the slab. Figure 20. Other elements from the slab. Note that as the slab is prepared from underneath, these are the undersides of the specimens as preserved. A) Moult remains of Isotelus. B) Cyclonema bilix lata. C) Dalmanella (Onniella). D) Hebertella. E) Rafinesquina with encrusted inarticulate Orbiculoidea. F) Cyrtolites minor. Figure 21. Trepostome bryozoans from the slab. Atactoporella schucherti, Batostoma implicatum (or possibly Batostoma prosseri ), Batostomella gracilis, Parvohallopora ramosa, and Parvohallopora subplana. viii Figure 22. Other elements from the slab. A) Ambonychia robusta. B) Rhodophyta. C) Thin section revealing Skolithos-type burrow in vicinity of crinoid remains. Figure 23. XRF scan revealing sulfur, calcium, iron, and phosphorous as major elements of the slab. Bottom center is a crinoid fossil. Figure 24. Extracted spectra from XRF, with enlargement for manganese (Mn). Note Mn concentration at upper surface.