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Echinodermata: Blastozoa: Diploporita): Paleobiology and Systematics

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Echinodermata: Blastozoa: Diploporita): Paleobiology and Systematics Th e North American Holocystites Fauna (Echinodermata: Blastozoa: Diploporita): Paleobiology and Systematics T. J. Frest, H. L. Strimple, and C. R. C. Paul Bulletins of American Paleontology Number 380, March 2011 BULLETINS OF AMERICAN PALEONTOLOGY Established 1895 Paula M. Mikkelsen Warren D. Allmon Editor-in-Chief Director Editorial Board Jason S. Anderson, University of Calgary Kenneth Angielczyk, Field Museum of Natural History Carlton Brett, University of Cincinnati Ann F. Budd, University of Iowa Peter Dodson, University of Pennsylvania Daniel Fisher, University of Michigan Dana H. Geary, University of Wisconsin-Madison Peter J. Harries, University of South Florida John Pojeta, United States Geological Survey Carrie E. Schweitzer, Kent State University Geerat J. Vermeij, University of California at Davis Emily H. Vokes, Tulane University (Emerita) William Zinsmeister, Purdue University Bulletins of American Paleontology is published semiannually by Paleontological Re- search Institution. For a list of titles and to order back issues online, go to http://museum- oftheearth.org. Numbers 1-94 (1895-1941) of Bulletins of American Paleontology are available from Periodicals Service Company, Germantown, New York, http://www.pe- riodicals.com. Subscriptions to Bulletins of American Paleontology are available in hard-copy and electronic formats to individuals or institutions; non-US addressees pay an additional fee for postage and handling. For current rates, additional information or to place an order, see http://museumoftheearth.org or contact: Publications Office Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York 14850-1313 Tel. (607) 273-6623, ext. 20 Fax (607) 273-6620 [email protected] This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Th e North American Holocystites Fauna (Echinodermata: Blastozoa: Diploporita): Paleobiology and Systematics T. J. Frest, H. L. Strimple, and C. R. C. Paul Bulletins of American Paleontology Number 380, March 2011 ISSN 0007-5779 ISBN 978-0-87710-493-3 Library of Congress Catalog Card Number 2011920734 © 2011, Paleontological Research Institution, 1259 Trumansburg Road, Ithaca, New York 14850, U. S. A. THE NORTH AMERICAN HOLOCYSTITES FAUNA (ECHINODERMATA, BLASTOZOA: DIPLOPORITA): PALEOBIOLOGY AND SYSTEMATICS T. J. Frest Deixis Consultants, 2517 NE 65th Street, Seattle, Washington 98115, USA1 H. L. Strimple Department of Geology, University of Iowa, Iowa City, Iowa 52242, USA2 and C. R. C. Paul Department of Earth Sciences, University of Bristol, Wills Memorial Building, Queens Road, Bristol, BS8 1RJ, UK; email [email protected] abstract The Holocystites fauna of central North America includes most known Silurian Diploporita (Echinodermata: Blastozoa). This distinctive diploporite association, widespread in the Wenlockian of Wisconsin, Illinois, Indiana, Kentucky, Ohio, and Tennessee, consists of eight genera in the Aristocystitidae, Sphaeronitidae, and Holocystitidae. Species of Holocystites Hall, 1864, and Triamara Tillman, 1967, are particularly characteristic. The fauna is best known from the Osgood Member, Salamonie Dolomite (late Llandoverian-early Wenlockian) of southeastern Indiana. Expanded quarrying operations near Napoleon, Ripley County, Indiana, add materially to knowledge of the Holocystites fauna. Thousands of specimens were recovered, including some in life position. Information from this and other localities allows formulation of a paleoecological model for the Holocystites fauna, which is tested against previously known distributional information. Most Silurian diploporites were low-level feeders with relatively inefficient subvective systems as compared with co-occurring camerate crinoids. In the Osgood, they required firm attachment sites in comparatively quiet, offshore, dominantly soft-bottomed environments with a moderate rate of continuous terrigenous sedimentation, conditions limited in southeastern Indiana to the Ripley Island positive area. Two major adaptive strategies are recognized: one relatively eurytopic group comprising species with free adults with elongate thecae, narrow bases, and aboral, stem-like processes, and a more stenotopic group of globular, large-based, completely sessile (attached) species. New taxa include a species of Holocystites Hall, 1864 (H. clavus n. sp.), a new genus (Paulicystis n. gen.) related to Trematocystis Jaekel, 1899, but with uniquely large ambulacral facets, and a new Pentacystis-like genus (Osgoodicystis n. gen.). The fauna also has an advanced sphaeronitid (Finitiporus n. gen.), the only Silurian sphaeronitid yet known. Both the Sphaeronitidae and Holocystitidae are revised. Subfamilies are established in both (Sphaeronitidae: Sphaeronitinae and Herpetocystinae; Holocystitidae: Holocystitinae, Pentacystinae, and Trematocystinae), based largely on peristome morphology. Holocystites is divided into three new subgenera (Holocystites, Megacystites n. sgen., and Sepulticystis n. sgen.) on the basis of pore morphology. Evolutionary trends are noted in the Holocystitidae toward reduction in number of thecal and peristomial plates, enlargement of the subvective system, elevation of the theca. Humatipore morphology becomes more specialized and efficient, but average size decreased. Detailed specific and generic phylogenies are constructed, using both traditional and quantitative phenetic methods. All produced similar results. Osgood diploporite biostratigraphy is revised and a zonal scheme presented. Osgood diploporites are strongly endemic. 1Deceased 10 April 2008. Because Harrell Strimple was unable to INTRODUCTION contribute to the present manuscript beyond the first draft (Frest, The late Llandoverian-early Wenlockian Osgood (Osgood 1983), any remaining errors of fact or judgement are solely the Member, Salamonie Dolomite of Pinsak & Shaver, 1964) of responsibility of the senior author. Indiana, Kentucky, and Tennessee has long been a focus of 2Deceased 21 August 1983. paleontological interest because of its extraordinary diploporite 3Corresponding author, whose contribution (since September 2009) fauna. It served as the exemplar nonpareil of the Holocystites fau- has been almost entirely confined to proofing, indexing, and editorial na (Paul, 1971), reported from six horizons in the Wenlockian matters necessary to correct minor typographical errors introduced of eastern North America. The Holocystites fauna includes all when the original typescript was scanned using OCR software. As a known Silurian Diploporita (Echinodermata, Blastozoa), with result, some of the sections, especially those dealing with phylogeny, will seem dated now. New taxa introduced here are to be referred to the exception of the more cosmopolitan Gomphocystitidae, as "Frest & Strimple in Frest et al., 2011." common to Europe and North America (Bockelie, 1979). 2 Bulletins of American Paleontology, No. 380 Major impetus for the present work was provided by the Member and are only known from the American and European discovery of an important new Osgood diploporite locality in echinoderm Providences (Text-fig. 2). The Holocystites fauna is Ripley County, Indiana. Expanded quarrying operations at the endemic to the Eastern North American Province. New Point Stone Company's Napoleon quarry [SW 1/4 sec. 21 and NW 1/4 sec. 28, T9N, R11E, Osgood 7½' quadran- PREVIOUS STUDIES gle; this is the same as Paul's (1971: 164) Locality 12; Text-fig. The initial taxonomic phase of work on the Holocystites fau- 1], due to the discovery of an unusually thick Brassfield and na dates to Hall (1861, 1864, 1868) and Winchell & Marcy largely limestone Osgood section, led to the stripping off and (1865), who described internal molds from the Racine Dolomite dumping as waste of large quantities of very fossiliferous upper and Manistique Formation and equivalents of Wisconsin and Osgood shales. This material contained thousands of diplopo- Illinois. The much richer Osgood diploporite fauna was a later rite thecae and other echinoderms, as well as a rich normal ma- discovery, with early taxonomy largely the work of S. A. Miller rine invertebrate fauna. Prior to the Napoleon discovery, only (see Miller, 1878, 1879, 1880, 1882, 1889, 1891, 1892a-b, ca. 200 diploporites had been recovered in over a century of 1894; Miller & Faber, 1892; Miller & Gurley, 1894, 1895). collecting. Extensive collecting by several Cincinnati, Ohio, One more species was described by Foerste (1917) from Ohio. area collectors, notably W. Bruce Gibson, D. Cooper, and D. By 1920, over 50 nominal species had been established, all in Bissett salvaged much of the available material. Aside from our the genus Holocystites Hall, 1864. Much of the early work was own collections, we were able to examine some of their material, overly typological by modern standards. The total number of and all three generously donated numerous critical specimens. Osgood diploporite specimens Miller had, for example, appears Most Osgood localities lack macrofossils, and all but a to have been roughly equivalent to the number of species that handful of specimens previously found were from a doz- he established. en sites in southeastern Indiana (Paul, 1971; Frest et al., Following the first burst of taxonomic interest was a lull of 1977). Descriptions of diploporite localities are included as an several decades duration. A separate genus, Triamara Tillman, Appendix herein. Osgood fossil localities in this area generally 1967, was established for one Osgood form.
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