Bulletins of American paleontology (Bull. Am. paleontol.) Vol 336

^/4- 3u((ctms of mv VOLUME 100, HARVARD NUMBER 336 UNIVERSITY Review of selected North American mitrate stylophorans (Homalozoa: Echinodermata) by Ronald L Parsley Paleontological Research Institution 1259 Trumansburg Road New York, 14850 U.S.A Ithaca, JANUARY 15, 1991 ) PALEONTOLOGICAL RESEARCH INSTITUTION Officers Harry A Leffingwell J Thomas Dutro, Jr Henry W Theisen James C Show acre Roger J Howley Peter R Hoover Henry W Theisen President Vice-President Secretary Treasurer Assistant Treasurer Director Legal Counsel Trustees Edward Bruce M Bell (to 6/30/93) Carlton E Brett (to 6/30/92) J Thomas Dutro, Jr (to 6/30/93) Harry A Leffingwell Robert M Linsley Cathryn Newton Samuel T Pees (to (to 6/30/93) 6/30/92) (to 6/30/91) (to 6/30/92) A D B Picou, Jr (to 6/30/92) James C Showacre (to 6/30/93) James E Sorauf (to 6/30/9 1) John Steinmetz (to 6/30/9 Henry W Theisen (to 6/30/92) Raymond Van Houtte (to 6/30/91) Ventress (to 6/30/93) Willl^m Warren P S Jr (to 6/30/91) BULLETINS OF AMERICAN 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Shuji R JiRl Klose, II Langenheim, L Harry Jr A Leffingwell Egbert G Leigh, A Jr Lenhard Louie N Marincovich, R Jr Moore NlKO Sakae O'Hara Samuel T Pees Richard E Petit Browne J David Bukry Sybil B Burger Ruth F KrIz G Edward Lyle D Campbell L Carter Anneliese S Caster Kenneth E Caster John E DuPont J Thomas Dutro, Jr J Mark Erickson Richard J Erickson Lois S Fogelsanger A Eugene Fritsche Christopher L Garvie Ernest H Gilmour Merrill W Haas Anita G Harris Steven M Herrick Robert C Hoerle F D Holland, Jr Frederick H C Hotchkiss Davtd Jablonski Richard I Johnson David B Jones Peter Jung TOMOKl Kase David Garrett Kerr CnciL H Kindle B Picou, Jr Robert A Pohowsky John Pojeta, Jr John K Pope John Anthony Reso Arthur W Rocker Walter E Sage, III John B Saunders Judith Schiebout Miriam W Schriner Edward S Slagle David H Stansbery Jorge P Valdes Charles G Ventress William P S Ventress Emily H Yokes Harold E Yokes Christine C Wakeley Thomas R Waller Albert D Warren, Jr Gary D Webster Ralph H Willoughby Armour C Winslow Yictor a Zullo subscriptions, and contributions are all important sources of funding, and allow Institution to contmue its existing programs and services The P.R.I, Research the Paleontological Paleontology and publishes two series of respected paleontological monographs, Bultetms of American of Palaeontographica Americana, that give authors a relatively inexpensive outlet for the publication works from the pasignificant longer manuscripts In addition, it repnnts rare but important older collection of inverleontological literature The P.R.I, headquarters in Ithaca, New York, houses a America: an extensive collection tebrate type and figured specimens, among the five largest in North Membership dues, significant future paleof well-documented and curated fossil specimens that can form the basis for ontologic research, and a comprehensive paleontological research library contributions Paleontological Research Institution is a non-profit, non-private corporation, and memberships, or be U.S income tax deductible For more information on P.R.I, programs, subscriptions to P.R.I, publications, call or write: The may Peter R Hoover Director Paleontological Research Institution 1259 Trumansburg Road New York 14850 U.S.A 607-273-6623 Ithaca, ^u&itms of -ffyncrican tOlO0 VOLUME 100, JANUARY NUMBER 336 Review of selected North American mitrate stylophorans (Homalozoa: Echinodermata) by Ronald L Parsley Paleontological Research Institution 1259 Trumansburg Road New York, 14850 U.S.A Ithaca, 15, 1991 Library of Congress Card Number: 90-63453 Printed in the United States of America Allen Press, Inc Lawrence, KS 66044 U.S.A CONTENTS Page Abstract Introduction Overview of Recumbent Echinoderms Review of the Subphylum Homalozoa The Class Homostelea 6 7 The Class Ctenocystoidea The Class Homoiostelea The Class Stylophora Taxonomic Position 8 Morphology, Functional Morphology, and Life Habits Orientation Morphology Theca 10 10 Aulacophore Origins and Phylogeny of the Mitrata Acknowledgements 14 16 Systematic Paleontology Introduction 18 Abbreviations of Repository Institutions 18 Systematics Class Stylophora Order Mitrata Suborder Anomalocystitida Family Anomalocystitidae Subfamily Anomalocystitinae Genus A nomalocystites Anomalocystites cormitus Hall Genus Kierocyslis, n gen Kierocystis insertus, n sp Subfamily Enoplourinae Genus Enoploura Wetherby Enoploura punclata Bassler Enoploura balanoides (Meek) Enoploura cf E balanoides (Meek) Enoploura popei Caster Enoploura cf E popei Caster Family Placocystitidae Subfamily Placocystitinae Genus Kopficystis, n gen Kopikystis kirkfieldi, n sp Genus AteleocysUtes Billings Ateleocyshtes huxleyi Billings Aleleocystites cf A huxleyi Billings Suborder Peltocystitida Family Kirkocystidae Genus Anatijerocystis Chauvel Anatiferocystis papillata (Bassler) 18 19 19 19 20 22 22 23 23 30 32 33 33 34 34 35 35 36 37 37 37 38 38 39 Appendix; Collecting Localities 41 References Cited Plates 42 46 Index 54 LIST OF ILLUSTRATIONS Page Text-figure Reconstruction of Enoploura popei in distressed orientation normal feeding orientation An oblique view of the proximal aulacophore, styloid, and a portion of the Phylogenetic chart of the Anomalocystitida Carapace detail of Anomaiocystites cornutus Plastron detail oi Anomaiocystites cornutus Lateral view oi Anomaiocystites cornutus Internal plastron surface oi Anomaiocystites cornutus Carapace of Kwrocyslis insertus, n gen., n sp Carapace and plastron of Enoploura popei Articulating spines and articulating surface on !VI4/M'5 oi Enoploura popei Internal features of Enoploura popei Detail of the styloid of Enoploura popei Transverse cross-sections of the distal styloid blade oi Enoploura popei Carapace of Enoploura punctata Plastron of Enoploura punctata Partial reconstructions of internal surface of the carapace of Enoploura popei Carapace of Kopficystis kirkfieldi n gen., n sp Carapace and plastron oi Anatiferocystis papillata its 1 14 19 Reconstruction of Enoploura popei in 11 its distal Table Explanations of plate abbreviations used 16 foldout inside back cover foldout mside back cover foldout inside back cover ; 22 24 24 25 27 28 31 31 33 35 39 40 LIST 12 foldout inside back cover 20 Plastrons of A natiferocystts papillata aiJacophore of Enoploura popei OF TABLES Page in this paper foldout inside back cover REVIEW OF SELECTED NORTH AMERICAN MITRATE STYLOPHORANS (HOMALOZOA: ECHINODERMATA) by Ronald L Parsley Department of Geology Tulane University New Orleans, Louisiana 70 1 ABSTRACT This paper reviews most of the North American mitrate stylophoran genera and species that are assigned to two suborders, and the Peltocystida The Anomalocystitida is herein divided into two long-ranging famihes, the Anom- the Anomalocystitida and the Placocystitidae The Anomalocystitidae contains mostly North American species (Middle Ordovician-Early Devonian) that are characterized by the proximal carapace margin having three plates, and only very rarely a placocystitid plate Norih American species include, within the new subfamily Anomalocystitinae, H'illmanocyslis denticulatus Kolata and JoUie, 1982 (Middle Ordovician), Kierocyslis mserliis n gen and sp (Middle Ordovician), Anomalocystites cormtlus Hall, 1859 [= A dispariHs Hall, 1859], (Early Devonian); and within the subfamily Enoplourinae, Enoploura punctata Bassler, 1932 (Middle Ordovician), Enoploura halanoides (Meek, 872) [= iT cnistacea (Haeckel, 896) = £ wetherbyi Caster, 952 = iT meeki Caster, 1952] and Enoploura popei Caster 1952 (Late Ordovician) Species assignable to the Placocystitidae are found in North America, Europe, and Australia They are characterized by the proximal carapace having only two plates in contact with the aulacophore, the plate consistently excluded, and the placocystitid plate rarely missing North American forms discussed herein include, within the subfamily Placocystitinae, Ateleocystites hu.\leyi Billings, 1858 (Middle Ordovician), and Kopficystis kirkfieldi, n gen and sp (Middle Ordovician) It seems likely that the Southern Hemisphere subfamily Allanicytidiinae branched off from the placocystitids some time in the Early Silurian and persisted until the Early Devonian The North American representative of the Peltocystida, the kirkocystid Anatiferocystis spinosa Ubaghs, 1979 [= Kirkocyslis papillala (Bassler, 1932)] from the Middle Ordovician of Oklahoma, is represented in Bohemia as well This genus has a greatly reduced number of plates The disposition and growth of these plates are discussed herein The feeding strategies of mitrates are seen to be different than previously reported In mitrates, the aulacophore faced into the prevailing current, convex upward, and food was conveyed to the food groove under the dorsally-fused cover plates through the arcuate to chevron-shaped openings between the overlapping set of cover plate pairs The width of these openings was controlled by the degree of distal aulacophore convexity Previous representations of feeding figured the distal aulacophore concavely curved over the proximal aulacophore and proximal theca I see this as a closing mechanism, used under extreme conditions, that alocystitidae 1 MA resulted in jamming together of the overlapping fused cover plate pairs (the configuration commonly preserved in the fossil record) Origin of the Anomalocystitida derives from Reticulocarpos-like comutans of the theca in the comute ancestral stock have been lost Only It is M1-M4 thecal suggested that the distal marginal plate elements marginals are homologous between comutes and anomalocystitids INTRODUCTION In the years since Caster's seminal 1952 paper on Enoploura Wetherby, 1879, which included the first major review of many of the known North American mitrate echinoderms, a great deal of new material and knowledge of stylophorans has come to light Many new ideas have been presented concerning their classification and how they functioned as animals Ubaghs (1961 and subsequent publications) pointed out that the tail-like appendage in stylophorans was a feeding organ (aulacophore) and thereby reversed the orientation commonly assumed for these creatures Haude (1980) modified Ubaghs' aulacophore model in the mitrates by proposing that they pipetted nutrients off the sea floor through an opening in the distal end of the aulacophore Abundant examples of closed terminal ends of mitrate aulacophores show that view to be untenable This work supports Ubaghs, with some minor modifications in interpretation of the deployment of the aulacophore as a feeding organ Ubaghs' opinion on the location of the mouth and function of the aulacophore is by no means universally accepted Philip (1979, pp 458-460), for example, suggests that the distal opening in all homalozoans is the site of both mouth and anus, the gut being U-shaped Philip reflects the traditional view held by authors prior to Ubaghs' work of 1961 More recent adherents to this model are Kolata and Jollie (1982) and Kolata (1984) In the latter work, Kolata postulates a shallow burrowing habit with the distal (abaulacophoral) end protruding from the substrate along with the articulating distal spines Jefferies (1967, 1968a, and subsequent papers) has proposed that the Stylophora are not echinoderms at all, but are more properly placed in the chordates under Bulletin 336 subphylum Calcichordata Jefferies, 1967 His views have met with a storm of rebuttal (Jefferies, 1968b, Ubaghs, 1975, Philip, 1979, Chauvel, 1981, Kolata and JoUie, 1982, and Jollie, 1982), in which I concur Various aspects of stylophoran evolution are also under debate I support Ubaghs' view that homalozoans are polyphyletic and that their primary symmetry is nearly, but not truly bilateral Caster (1968, 1983), on the other hand, strongly suggests that homalozoans are monophyletic and that the ancestral echinodermal radical was triradiate Using cladistic methods, Paul and Smith (1984, pp 461-462) would derive the carpoid homalozoans from an Early Cambrian asymmetric solute radical, and by virtue of their methodology argue that the group is monophyletic the recumbent echinoderms are considered toMiddle and Late Ordovician It should be noted that articulate brachiopods and nestling clams, which derived their food from the same part of the water column, also enjoyed considerable diversity in the Middle and Late Ordovician I can only speculate why homalozoans and recumsity, if all gether, is in the bent blastozoans did not persist higher in the geologic column They were probably supplanted by more efficient forms or eliminated by durophagous predators (Signor and Brett, 1984), because there is little reason to believe that the physical aspects of this ecologic niche have significantly changed since the Early Paleozoic REVIEW OF THE SUBPHYLUM HOMALOZOA OVERVIEW OF RECUMBENT ECHINODERMS In this brief introductory look at the homalozoans The subphylum Homalozoa Whitehouse, 1941, an artificial is grouping of four morphologically similar and other recumbent echinoderm groups (i.e., pleurocystitids and rhipidocystians), I suggest that the over- classes that ecologically exploited the nutrient-rich lay- of these all morphologic similarity displayed by adaptation brought on by groups is due to convergence to similar bottom-living habits Generally all of these haps the smaller elements of the closely-adjacent mei- forms have a high degree of thecal streamlining and abundance and diversity in Middle and Late Ordovician, at which time recumbent bottom-living echinoderms of the subphylum Blastozoa are also at the acme of their abundance and diversity The class Homostelea Gill and Caster, 1960, is made up of a single order, the Cincta Jaekel, 1918, and is limited to the Middle Cambrian of Europe and North all bilaterality (but rarely are they truly bilateral) In most, a rim of substantial marginal plates surrounds flexible polyplated central surfaces on both inferior and superior surfaces In some, one or both somatic surfaces are also regularized into rigid surfaces Ventral or in- commonly concave, with the marminimize adhesion to the bottom In many forms, appendages such as columns, steles, and aulacophores were adapted to serve locomotor functions These recumbent forms not only ferior surfaces are er at and near the water-sediment Homalozoans range from ofauna interface, the Middle and per- Cambrian to Early Pennsylvanian and manifest their greatest ginals serving as runners or skids to Africa exploited a similar food source, but in doing so exer- Robison and Sprinkle, Middle Cambrian of North America, Europe (France and Bohemia), and North Africa, and is the only "carpoid" group without an cised some degree of vagility in moving from place to place using their elongated appendage, or, minimally, were sufficiently vagile to favorably orient themselves on the sea floor relative to nutrient-bearing currents Most of these forms are found in low-energy environments where lime muds to limy shales were deposited, although higher energy environments (i.e., in calcarenites and fine-grained sandstones) are associated with a number of genera These low-energy environments suggest that stylophorans fed at or very near the water-sediment interface, where nutrients accumulated Perhaps these organisms are best described The 1969, class Ctenocystoidea is also limited to the appendage class Homoiostelea Gill and Caster, 1960, rangfrom the Late Cambrian to the Middle Devonian of Europe and North America Because of their eco- The es logical convergence, is North America, Europe, South America, To date, no stylophorans have been reported from the several continental plates that make up Asia Stylophorans have a long feeding organ, Africa, first few mm above recumbent echinoderms appears to have been a true suspension feeder in the "upper" water column The total range of these recumbent feeders is from Middle Cambrian to Early Pennsylvanian The greatest diver- re- geographically the most widely-distributed class, be- organic concentrates and associated microbiota pres- None of the mentioned show ing found in the water-sediment interface this class to several of the pleurocystitid rhombiferans, such as Amecystis Ulrich and Kirk, 1921, and Pleurocystites Billings, 1854 (see, e.g Dehm, 1934) The class Stylophora Gill and Caster, 960, ranges from the Middle Cambrian to Early Pennsylvanian and as concentrate suspension feeders, utilizing the rich ent in slow current regimes in the members of markable similarities to some stylophoran genera and and Australia the aulacophore, that in some is strongly homeo- of homoiosteles This homeomorphy, as discussed below, is probably in part due to the convergence in locomotor function morphic with the stem or stele North American Mitrate Stylophorans: Parsley 49 Explanation of Plate Page Figure (Meek) UCM 37296 Holotype of Enoploura Crustacea Haeckel, 1896, carapace, right lateral, left lateral, and plastron views of a nearly complete theca with attached proximal aulacophore, x 2.25, Locality E 5-9 Enoploura punctata Bassler All specimens are syntypes (USNM 91856) from Locality F Carapace with articulating spines and distorted proximal aulacophore attached [Note the transverse ridge on the internal surface of the plastron exposed by erosion of the carapace plates.], x 3.0 Lateral view of a deformed proximal aulacophore, eroded styloid, and proximal segments of the distal aulacophore, x5.0 Nearly complete carapace with the attached poorly-preserved proximal aulacophore and part of the distal aulacophore, x 2.75 Carapace with the internal surface exposed; the distorted proximal aulacophore is also exposed [Part of the right articulated 1-4 Enoploura balanoides spine is 32 30 also present.], x3.5 Slightly distorted carapace with the movable spines articulated; proximal aulacophore incomplete and distorted [This specimen its first illustration by Bassler (1932, p 18, figure 9, lowermost specimen).], x 3.5 has been considerably prepared since 10-12 Kierocystis insertus, USNM 10 1 12 42166 new genus and (holotype), species from Locality G Carapace with incomplete proximal aulacophore and discontinuous distal aulacophore, x3.0 Poorly-preserved and incomplete plastron with proximal aulacophore, eroded styloid, and several segments of the distal aulacophore attached, x 3.0 Proximal area showing the proximalmost upper tetrameres of the proximal aulacophore inserted into the carapace, x5.0 22 Bulletin 336 50 Explanation of Plate Page Figure 1-5, 8, 10 Enoploura punctata Bassler 1-5 All specimens from Locality 30 USNM 91854, theca with (mostly) internal surface of the plastron exposed, proximal aulacophore, styloid, and short section of distal aulacophore attached and deformed, x4.0; 2, USNM 91854, deformed theca, carapace exposed, proximal aulacophore attached, and deformed, x3.5; 3, USNM 91854 (syntype: see Bassler, 1932, p 18, figure 9, specimen on upper right), weathered carapace with part of the internal surface of the plastron exposed, x3.5; 4, USNM 91854, eroded carapace showing part of the tranverse ridge on the internal surface of the 10 F 1, plastron, proximal aulacophore, and several segments of the distal aulacophore attached, x4.0; 5, USNM 91854, deformed proximal aulacophore, eroded styloid, and nearly complete ventral distal aulacophore, x3.0 USNM 91854, plastron and attached proximal aulacophore, x4.0 Locality F USNM 91854, distal end of plastron (lip) with denticulate platelets which are probably functional or vestigial analpyramid plates [Note adjacent proximal part of the articulated spine with stereom structure preserved.], x 7.5, Locality ' F 6, Enoploura cf E balanoides (Meek) Both specimens x 3.0, from Locality Q 46045, proximal carapace with proximal aulacophore and several segments of the distal aulacophore attached 46046, eroded plastron with proximal aulacophore, eroded styloid, and several distal aulacophore ossicles attached 33 UCM UCM 9,11 Enoploura cf E popei Caster ^ j ' ' 34 UCM 43378, eroded carapace with paired spines nearly articulated [Note spines on distal marginal plates of carapace Stereom structures are visible on most of the plates This specimen is unusually large for a mitrate.], x 2.0, Locality H [ 11 USNM 145586, carapace with attached poorly-preserved proximal aulacophore and part of the distal aulacophore, x2.5, Locality P Bulletins of American Paleontology, Volume 100 Plate Bulletins of American Paleontology, Volume 00 Plate North American Mitrate Stylophorans: Parsley 51 Explanation of Plate Page Figure 1-3, 6-9, 1 33 Enoploura popei Caster UCM 1-3 and plastron views [Theca is slightly distorted with the proximal part spine proximal aulacophore, well-defined styloid, and several proximal elements of the distal aulacophore 25993 (holotype), carapace, of the right attached.], x 2.0, Locality 6-8 left lateral, I UCM 46049, distal ends of distalmost styloid blades [Note the deep muscle pits and narrow slit for the food groove.], X 3.0, Locality K 1 UCM 46049, lateral view of a styloid that is composed of three fused ossicles, x6.0 Locality K UCM 46049, proximal end of styloid blade [The deep pit continuous with the lumen of the proximal aulacophore.], is X 3.0, Locality K 4, 32 Enoploura species USNM 40704, lateral and ventral views of a nearly complete aulacophore the feeding position.], x 3.0, Locality 10, 13 [Distal aulacophore appears to be preserved in J Enoploura punctata Bassler Both specimens are USNM 91854, x 3.0, Locality F 10 Incomplete carapace with a spine and most of the aulacophore attached 13 Eroded carapace with eroded proximal aulacophore and eroded proximal segments of distal aulacophore Enoploura cf E balanoides (Meek) 31309, proximal aulacophore with partal styloid blade and poorly-preserved distal aulacophore segments attached, UCM x3.0 Locality L 30 33 Bulletin 336 52 Explanation of Plate Page 37 Figure 1-4 Ateleocystites A huxleyi Billings MCZ half, cf 1063, incomplete and eroded carapace with an attached distorted proximal aulacophore and the proximal or so of the distal aulacophore, x4.0 Locality M MCZ 2064, proximal carapace with the proximal aulacophore and several poorly-preserved segments attached [Note the well-preserved placocystitid plate.], x4.0, Locality M USNM USNM aulacophore 93348, incomplete plastron with poorly-preserved proximal aulacophore and styloid; distal aulacophore essentially complete, x2.5 Locality distal M 401499, distally-incomplete plastron, proximal aulacophore, weathered lacophore, x2.0 Locality styloid, and part of the distal au- M 39 5-9, 11-13 Anatiferocystis papillata (Bassler) UCM 46036, carapace, plastron, and right lateral views of a theca with distorted and displaced proximal aulacophore 5-7 [Note the spine base for the single articulating spine in figure 46037, right lateral view of a theca, x4.5 6.], x4.5 Locality N UCM UCM 46039, carapace and incomplete plastron views of a theca with an incomplete proximal aulacophore attached, 9, 13 x4.5 Locality N UCM II, 12 46038, carapace and plastron views of a theca [Note the axial displacements of the plates of the plastron.], X 5.0, Locality N 10 new genus and species 6408 (holotype), slightly distorted carapace with an incomplete distal aulacophore attached [Note the anal-pyramid series.], x 4.5, Locality O at the distal end of the theca exposed by erosion of the overlying Kopficystis kirkfieldi, USNM 1 platelets CM 35 Bulletins of American Paleontology, Volume Plate 100 '•^Vi » t'j % ^>.#iMiii^ ^^^^^^^HV^' 11 10 12 13 ,v Plate Bulletins of American Paleontology, Volume 100 11 10 i North American Mitrate Stylophorans: Parsley Explanation of Plate 53 Page Figure 1-11 Anatiferocystis papillata (Bassler) 1, 39 UCM 46037, carapace and disturbed plastron of a theca with remnants of the proximal aulacophore attached, x4.5 Locality N 4, UCM UCM 46040, carapace of a theca, x 5, Locality N 46041, carapace and plastron of a distally-incomplete theca [The incomplete distorted proximal aulacophore is attached.], x4.5 Locality N 6, USNM 401450, well-preserved carapace and plastron of a theca [Note the protruded subanal plate and spine base.], x4.5 Locality N 8, 11 10 USNM 401451, carapace and UCM UCM plastron of a theca [Note the well-defined spine base at end of plastron.], x5.0 Locality N 46042, proximal view of a theca with attached distorted proximal aulacophore, x6.0 Locality N 46043, theca with articulated spine lying across it [Note the stereom structure of the spine.], x5.0, Locahty N Bulletin 336 54 INDEX Note: Page numbers are in light face; plate numbers are in bold face type; the page numbers on which principal discussions occur are in itaUcs; B indicates the foldout inside the back cover aboral, discussed barrandei Anatiferocystis Africa Barrandeocarpus XJha^s, 1979 North South and Gill in Allanicytidium Caster and Gill in Ubaghs, Bassler (1915) 23,32,36,37 16 Bassler (1932) 5,25,30,32,41,49,50 30,32 16 Bassler (1935) Bassler (1938) 19 5,16.17,19 Bassler (1943) 30,38,39 Ubaghs, 1968a Allanicytidiinae 6,7 17,19 africanus, Placocystis Allanicytidiidae Caster Bassler (1950) 968a 10,11,16,17,19,20 Amecystis Ulrich and Kjrk, 1921 AMNH U [American Museum of Natural History, New York, NY, 18,22,46,48 S A.] AmygdalothecaUtta^s, 1969 10,14,15 Anatiferocystis ChauMeX, 1941 15,i5,40,41 38 barrandei Chau\£\ 1941 papillaia {Bass\eT, 1943) 38,i9,40,52,53 7,8 spwosa Ubae):\s 1979 Anomalocystida Caster, 1952 Anomalocystis Hall Ammalocystites HaW, 1858 balanoides (Meek) corwu/iis Hall, 1859 dispariHs HaW, 1859 Anomalocyslites 36 23,32 Anomalocystitida Caster, 1952 5,16,75,19,20,35 Anomalocystitidae Bassler, 1938 5,16,17,79,20,23 Anomalocystitinae Sassier, 1938 5,16,17,79,23 Arnold A Bather (1899) Bather (1900) 14,19,23,32,36,37 Bather (1913) 10 BelemnocystitesM\\\eT and G\a\ey 1894 Benbolt Formation 39 19 Blastozoa Spnnkle 1973 5-7,15,34,38,40 Haj near Zahofany Branson and Peck (1940) Museum British Broadhead T Bromide Formation, Pooleville Bull Fork 39 12 (Natural History), London, England, U K Calcichordata Jefferies, Member 42 1967 14 Carlson and Fisher (1981) Caster (1952) (1 16 39,42 6,8,18 Cannon Limestone Caster 18,39 Formation capensis Placocyslella 6,7 Bohemia 19 18 Asia 19 20 37 Anomocystis Hall Antedon FreminviWe, 1811 1952 5,9,10,18,21,23,32,34-36,37 5,20,26 Meek, 1872 , disparilis (Hall) Billings (1854) 32 (Aleleocystites'?) Basslerocystis Caller 32 Billings (1858) 5,18,20,22,35,46-48,8 balanoides Meek 1872 39 and Moodey (1943) 18 19 huxleyi (Billings) Bassler 5,39 9,10,13,17,79,20,22,23,32,35,36 1,2,3 38 15-17,34,35 30,41 16 8,12 5,14,18-20,23,24,28,30-34,36-38 954a) 13 16,19,20 Aspidocarpus \J\3a^%, 1979 15 Caster (1954b) Atelecystis Billings 36 37 Caster (1968) 6,8,11,12,20 Caster (1983) 6,10,12,16,19,20 37 Caster, Dalve, huxleyi (Billings) Ateleocystis huxleyi (Billings) 9,10,13,17-23,32,34,35,i6,37 Ateleocystites^i\\m%.s, 1858 balanoides (Meek) 32 gutlenbergensis V.o\a\a and ioWie, 1982 /i«.v/e,V'' cf /I 26,36,37 5,36,i7 Bilhngs, 1858 /lu-v/ey; Billings, Ateleocystites lAndsXrom 1858 non 36 Billings Atelocystites HaeckeX, 1896 36 Australia New i7,52 5,6,16,19 South Wales Caster, K and Pope (1955) 30 E Casrwcww 16 va// Ubaghs and Robison, 1985 Catheys Limestone Formation 30,41 Ceralocyslts Jaeke\ 1901 12 Chauvel (1937) Chauvel (1941) Chauvel (1981) 38 15,38,40 Chinianocarpos Vbag^s, 1961 15,41 Chordata 8,18 Tasmania 16,19 Cincta Jaekel, 1918 Victoria 16,19 Climacograptus spiniferus Zone 36 F Coonigan Formation 16 Australocystinae Caster, 1954b Australocystis CasXer, 16,17,19 1954b 16,17 Collier, Comuta BalanocystitesBanaxiAe, 1887 15,38,40,41 balanoides 32 Anomalocystites 23,32 Anomalocystites (Ateleocystites?) balanoides (cf.), Barrande (1887) Enoploura 1,2,3 5,18,20,22,35,46-48,8 Costeltaria beds 30 Cothurnocystis Baxher, 1913 10 Crustacea 32 Enoploura 5,33,49 5,18,i2,33,34,36,49 Placocystis 18,32 Aleleocyslites Enoploura cornutus .Anomalocystites 10,12,29 5,6 ii,50,51 9,12,15,19,25,38.40 Ctenocystoidea Robison and Sprinkle, 1969 Cystitidae 6,7 37 North American Mitrate Stylophorans: Parsley 39 Czechoslovakia 55 19,20 Hall (1861) 13 Hall and Clarke (1892) Dalve, E R 18 Deak, W Dehm Dehm (1932) 13,30,35 (1934) 6,30 34 denticulatus Willmanocystis Derstler K 7,18,36 19 Diamphidiocystida Kolata and Guensburg, 1979 Diamphidiocystidae Kolata and Guensburg, 1979 Diamphidlocystis Kolata and Guensburg, 1979 6,7 6,7 6,7 37 37 Anomalocystites 19,22,41 and Price (1975) Derstler, 22,41 huxleyi 19,22,41 Derstler(1979) Haude(1980) Helderberg, Lower Homalozoa Whitehouse, 1941 Homoiostelea Gill and Caster, I960 Homostelea Gill and Caster 1960 19 18,19 Atelecystis 37 Ateleocystis 5,i6,37 Ateleocystites huxleyi (cf.), Ateleocystites i7,52 disparilis, 5,20,26 Anornatocystiles 20 Basslerocyslis distal, discussed 10 dorsal, discussed Idaho 8,9 Enopleura'? papillala Bass\er, 1943 S A road cut south of Liberty [Locality H] in 16 24 Indiana St Echinodeimata Elkhom Formation Enopleura Bassler, 1932 punctata Bassler, 1932 Geological Survey, Urbana, IL, U Illinois State 34,42,50 Marys 34 Weisberg, railroad cut 13, lower Suman 42 W., 32 inferior, 30 39 insertus, Kierocystis IVi' 'h, SE 'U, Sec 8, T N., R 34,42,50 quadrangle [Locality P] 10 discussed /owacv^ni 5,22,23,49 Thomas and Ladd, 1926 Enoploura Wetherby, 1879 5,8-10,13,16,17,20-22,25,25,27,30-33,36,37 balanoides (Meek, \S12) cf E balanoides [Meek, \S72) 5, 18,i2,33, 34,36,49 ii,50,51 5,6 Jaekel (1901) 12,19,23,25 Jaekel (1918) 6,15,18,36,37 5,8,18 Jefreries(1967) Crustacea (Haeckel 1896) 5.33,49 Jefferies (1968a) meeki easier, 1952 popei Caster, 1952 5,33,51 Jefferies (1968b) 14,38 5,9,11, 12,24,25,27,28, 31,32,ii,34,51 Jefferies (1984) 8,23,34 34,50 5,i(^,3 1-33,49-51 Jefferies (1986) punctata Bassler, \932 4,5,6 51 sp 5,32 wetherbyi easier, 1952 Enoploura"! Wetherby Enoplourinae Caster, 1952 Jefferies 14 and Uwis (1978) and Prokop (1972) 14,23,37 15,16,10 , 16 Jeffenes, R P S and Banks (1985) Jince Formation Jell Barrett, 12 JoUie (1982) 6,8 5,6,38 32,42 Fairview Formation Fatka and Kordule (1985) 23,26,34 forbesianus, Placocystites 36 Placocystites 6,38 France Montagne Noire Freminville(1811) 14 Frest, Kolata, Jefferies 33,38 5,16,17,20,2i,36 Eumorphocystis Branson and Peck, 1940 Europe (aff.), 6,8 Jefferies (1981) cl E popei easier, \952 forbesianus 5,8 and Mapes (1985) 38 Kentucky Mercer Co., Curdsville [Locality G] Kier, P 13,24 22,42,49 M 16 Kierocystis, n gen 11,17,19,22,23 insertus, n sp 5,22,23,49 Kirk (1911) 14,16,19,26,36 42 Kirkfield Limestone kirkfieldi Kopficystis 15,i« Kirkocystidae Caster, 1952 38 Kirkocystis QassXer, 1950 Galliaecystis Gill 10 \Jha^s, 1969 and Caster (1960) 6,16,18,19,34,36 37,42 Glens Falls Limestone Formation Glossopleura Zone Golf Course Formation, Gene Autry Shale Grand Lake Formation Grant Lake Formation, Corryville Member GSC [Geological Survey of Canada, Ottawa, Ontario, Canada] 5,39 papillata (Bassler 1932) Kolata (1984) Kolata and Guensburg (1979) 38 Kolata and JoUie (1982) Kolata, D R 42 42 de Koninck (1869) Kopficystis, n gen Member kirkfieldi, r\ 5,i5,52 18,19 6,8,9,12,18,19,21-23,26,34,36,37 16,18,36 23,26,35,36 16,17,20,34,i5,36 sp 5,i5,52 18,37 Guensburg, T E Guttenberg Formation 18,30,41 guttenbergensis Ateleocystites 26,36,37 Haeckel (1896) 36 5,18,19,23,32.36,37 Hall (1858) 9,10.13,19.23.32 Hall (1859) 5,18,20.26 Lagynocystida 16,37 Lagynocystis }aeke\, 1918 15,37,38 Landing, E 16 Letna Formation 39 Lexington Limestone Curdsville Limestone Lindstrom (1888) 13 Member 22,42 37 Bulletin 336 56 Oklahoma Localities: A: New B: New York, Litchfield, unknown locality New York, Herkimer Co., Days Comer, York, Dayville, Saltsburg quarries, "Jerusalem Hill" 5,38 Carter Co., Criner Hills, Rock Crossing [Locality N] 11,41 Ad C: 22,47,46 Saltsburg quarries 22,47,46 D: Maryland, Cumberland 22,47,47,48 E: Ohio, Oxford F: Tennessee, 0.5 mi north-northeast of Pulaski 32,47,49 30,47,49-51 G: Kentucky, Mercer Co Curdsville H; Indiana, in road cut south of Liberty 1: Ohio, Clermont Co., Stonelick Creek 22,23,42,49 , J: Ohio, Warren Q: Ohio, Cincinnati R: Quebec, near Hull, S: lower 'h, SE 'A, Sec 8, T 34,42,50 32,33,42.50 Bngham Quarry 36,37,42 36,37,42 Malvinokaffric Lebanon Co., Swatara Gap 36,37,42 Realm 10,16,19-21 22,41 Manlius Formation, Olney Member Martinsburg Formation 36,37,42 19 Maryland 22,47,47,48 Cumberland [Locality D] MCZ [Museum of Comparative Zoology, Harvard University, Cambridge, Meek Meek MA, U 18,36,37,52 S A.] (1872) 5,18,23,31-33,36 (1873) 23,32,36 meeki Enoploura Meyer, D L 5,33 16 19,23,32,36,37 Miller (1889) Miller and Gurley (1894) it rocysli I es Parsley (1981) 13 Parsley (1982) 12,13,39 Parsley (1988) 10,11,13,14 and Caster (1982) Parsley and Mintz (1975) Paul and Smith (1984) Parsley Paul, C R 10,12 12 6,12 C 7,18 Peltocystida Jefferies and Lewis, 1978 5,16,i7 38 Peltocystidae Pellocyslis Thoral 1935 15,37,38,40,41 19 22 Curtin Gap Lebanon Co., Swatara Phihp (1979) 36,37,42 [Locality T] 5,6,8,18,41 Philip (1981) 10,19,20 Phylhcystis Thoral, 1935 9,10,12,16 Placocystella capensis Rennie, 1936 16 Placocystida 18 Placocystidae Caster, 1952 34 34 Placocystinae Caster, 1952 Placncyslis de Koninck 23,33 afncanus Reed 1925 Crustacea Haeckel, 1896 16 18,32 f/ac«-m/to de Koninck 1869 17.23.35,36 forhesianus de Y^onincV., 1869 afT 23,26,34 36 forbesianus 18,19 Placocystitida Placocystitidae Caster, 1952 5,16,17,19,34 Placocystitinae Caster 1952 5.20,i4,35,36 25 Barrande 1887 9,12,26 New York 19 Dayville, Saltsburg quarries, "Jerusalem Hill" [Locality A] 1854 Plei4rocysliles BiWings, Enoploura popei (cf.), Enoploura 5,9,1 1,1 2,24,25,27,28,3 l,32,ii,34,51 popei, i4,50 proximal, discussed punctata 22,47,46 Herkimer Co., Days Comer, Saltsburg quarries 13 11,12 Parsley (1980) 36 Milrocyslella }aeke\ 1901 M Paranacystis Caster, 1954a 10,12,14,18,37,40 Mississippi River Valley Mitrata Jaekel, 1918 iS,39,40,52,53 7,8 Pennsylvania Quebec, near Hull, presumably close to Brigham Quarry T: Pennsylvania, Anatiferocystis 39 35,42,52 13, papillaia 5,39 39,42,52,53 quadrangle 36,37,42 Kirkocystis 42,51 71/2' discussed Ottawa Limestone Hull Beds 34,42,51 Co Suman oral, 35,42,52 Enopleural Avenue 34,42,51 L: Ohio, one mile northeast of Camden, on Seven Mile Creek, 32,33,42,51 parallel to main highway 36,37,42,52 M; New York, Trenton Falls N: Oklahoma, Carter Co., Criner Hills, Rock Crossing N., R W., 38 Ontario, Kirkfield [Locality O] 34,42,50 K: Ohio, Cincinnati, Boudinot O: Ontario, Kirkfield P: Indiana, Weisberg, railroad cut 39,42,52,53 southern [Locality C] 30 Enopleura 4,5,6 5,iO,3 1-33,49-51 near Hull, Brigham Quarry [Locality R] 36,37,42 Enoploura 22,47,46 Litchfield, unknown Trenton Falls [Locality M] New Zealand Notocarpos Philip, 1981 NYSM [New York 22,47,46 locality [Locality B] State 37,42,52 16 presumably close 36 to Brigham Quarry [Locality S] 36,37,42 10,17,19,20 Museum, Albany, NY, U S A.] 16,18,22,46 Ohio 24 Cincinnati [Locality Q] 32,33,42,50 Boudinot Avenue [Locality K] Clermont Co., Stonelick Creek [Locality I] one mile northeast of Camden, on Seven Mile Creek, main highway [Locality Oxford [Locality E| Warren Co [Locality J] Quebec L] Rafinesquina Hall and Clarke, 1892 13 Reed (1925) 16 Regnell (1945) 37 36 Regnell (1960) Rennie (1936) 16 and Prokop, 1972 33,42,51 /?«;fu/ofarpo.s Jefferies 33,42,51 Rhenocystis Xiehm, 1932 parallel to 32,33,42,51 Ridgely Formation Robison and Sprinkle (1969) 5,15-17,19 17,30,35,36 22,41 32,47,49 42,51 Saluda Formation 41 57 North American Mitrate Stylophorans: Parsley 18.19 Schuchert(1904) Shriver Formation 22 Smith (1988) 12 Sonnier, S 18 Signer and Brett (1984) South America Southern Hemisphere Southern Illinois University, Edwardsville, IL, U sp., Enoploura 5,10 S A Sprinkle (1976) 12 Sprinkle (1982) 42 7,18 CA, U S A.] 18,22,46 UCM [University of Cincinnati, Cincinnati, OH, U S A.] 9,11,16,18,22,24.27.32,33,34,39,40,45,48-52 K University of Liege, Liege, Belgium New Orleans New Orleans LA, U University of LIniversity of Tennessee Knoxville TN, U S A 18 16 S A 16 18 TX 18 U S A USNM [U S National Museum of Natural History, Washington, DC, U.S A.] 16,75,22,30,3 1,35,37,39,46-53,8 University of Texas Austin, Utah 34 Stocker, J 5-7,5,9,12,16,18.26 Stylophora Gill and Caster, 1960 10 superior, discussed 24 Tennessee mi north-northeast of Pulaski [Locality F] 30,'#7,49-51 39 eastern Udd vali, 10 Castericystis venlral discussed K(c;onan'ito Gill and Caster, 1960 16,17 Waynesville Formation 34,42 10,17,19.20 Tasinarucytidium Caster, 1983 Thomas and [University of Cahfomia, Berkeley, University of Liverpool, Liverpool, England, U Sprinkle (1973) 0.5 16,18 51 19,23,36 Sprinkle, J UCB G Ulrichand Kirk (1921) Springer (1913) Spnnkle and Robison (1978) Ubaghs 18 5,39 spinosa Analiferocystis Ubaghs (1987) Ubaghs and Robison (1985) (1926) Thoral (1935) Tulane University, New Orleans, LA, U search of the Graduate School] 9,10,12.16,37,38,40 S A., [Committee of Re18 22 West Virginia, Ridgely Wetherby (1879) welherbyi Enoptoura Whitehouse(1941) Whitewater Formation Willmanocystis Kolata and 5.8.10.16.20.21.23.32.33,36 33,38 Jollie, 1982 denticulatus V^XaXa anA ioWis, 1982 Ubaghs(1961) Ubaghs (1968a) Ubaghs (1968b) Ubaghs (1969) Ubaghs (1975) Ubaghs (1979) 5,8,10,12.13.15.41 8-12.14-16,18.19,23,26,31,37,38,39,41 Wilson (1946) Woodward Woodward 34,42 17-19,23 36,37 (1871) 19,23,37 (1880) 19,23,33,36,37 9,10,14,15,37,38,41 6,7,8 5,12,15,16,35,39^1 Zittel (1879) 36 -I I M'2 M'3 M'4 Text-figure figure is 5.— Carapace based on USNM detail of Anomalocystites cornutus This 33661A m'5 Carapace (PI fig I) — Plastron detail of Anonialocysiiles cornulus on USNM 33661A (PI 5) Text-figure figure IS based fig This Text-figure «M/U5 This figure 2720 in m6 — Internal is m5 plastron surface of Anonialocysiiles ror- a composite based on USNM 33661Fand AMNH Extending from M'l to the muscle pit the diagonally -transverse septum Tnangular area devel- (PI 3, figs 4, 13) M4 is oped mostly on M'l-MI is the raised (thickened) oral field The triangular area developed on m'5, m6 mS A and CS is the thickened distal field Between M'4-in'5 and M4-m5 are the muscle pits These pits probably contained muscles thai moved the spines toward the plane and (probably) muscles that were part of the anal sagittal sphincter system Overlayering of plate matenal to form the walls and deepen the muscle pits has obscured the plate sutures Narrow inner marginal grooves run the length of the theca from the muscle pits to the extremities of M2-M'2 Table 1.— Explanations of plate abbreviations used 'MA- CM3^ in this paper ... numbers and volumes may be had on request Volumes 1-23 of Bulletins of American Paleontology have been reprinted by Kraus Reprint Corporation, Route 100, Millwood, New York 10546 USA Volume of Palaeontographica... Jr (to 6/30/91) BULLETINS OF AMERICAN PALEONTOLOGY and PALAEONTOGRAPHICA AMERICANA Peter R Hoover Editor Reviewers for this issue T E Broadhead T E Guensburg D R Kolata A list of titles in both... Palaeontographica Americana has been reprinted by Johnson Fifth Ave., New York, NY 10003 USA Reprint Corporation 1 1 Subscriptions to Bulletins of American Paleontology may be started at any volume or